BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY VOL. ii 1963-1965 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1965 DATES OF PUBLICATION OF THE PARTS No. i . . . . . .25 October 1963 No. 2 . . . . . .2 December 1963 No. 3 . . . . . .11 February 1964 No. 4 . . . . -13 February 1964 No. 5 . . . . .31 December 1963 No. 6 ...... i April 1964 No. 7 . ... . . 2 March 1964 No. 8 ...... 2 July 1964 No. 9 ...... 20 May 1965 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING 2 - DEC 1965 CONTENTS ZOOLOGY VOLUME n PAGE No. i. A revision of the genus Hipposideros. By J. E. HILL I No. 2. A revision of the British mites of the genus Pergamasus Berlese s. lat. (Acari : Mesostigmata) . By S. K. BHATTACHARYYA (Pis. 1-8) 131 No. 3. The Cheilostomatous Polyzoa Neoeuthyris woosteri (MacGillivray) and Reginella doliaris (Maplestone). By ANNA B. HASTINGS (Pis. 1-3) 243 No. 4. The marine Enoplida (Nematoda) : a comparative study of the head. By WILLIAM G. INGLIS 263 No. 5. The swimbladder in African Notopteridae (Pisces) and its bearing on the taxonomy of the family. By P. H. GREENWOOD (Pis. 1-4) 377 No. 6. A revision of the genus A earns L., 1758 (Acaridae, Acarina). By D. A. GRIFFITHS (PI. i) 413 No. 7. A collection of Mesostigmata (Acari) associated with Coleoptera and Hemiptera in Venezuela. By K. H. HYATT 465 No. 8. The genus Steatonyssus Kolenati (Acari : Mesostigmata). By W. M. TILL & G. OWEN EVANS 511 No. 9. Form and function in the evolution of the Vermetidae. By J. E. MORTON 583 Index Volume II 631 U.ll. n.u. ^ 1 OCT A REVISION OF THE GENUS HIPPOSIDEROS J. E. HILL BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. i LONDON: 1963 A REVISION OF THE GENUS HIPPOSIDEROS BY J. E. HILL Department of Zoology British Museum (Natural History) Pp. 1-129 : 41 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. i LONDON: 1963 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. n, No. i of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. Trustees of the British Museum 1963 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued October, 1963 Price Thirty-five Shillings A REVISION OF THE GENUS HIPPOSIDEROS By J. E. HILL SYNOPSIS A taxonomic revision of the Microchiropteran genus Hipposideros is presented in this study which is primarily devoted to an examination of the genus at the specific and higher levels, with marked emphasis on the detailed diagnosis of its numerous species, their relationships, supraspecific groupings and probable phylogeny. INTRODUCTION THIS revision is based on the unequalled collections of the British Museum (Natural History), which contain an extensive series of the genus Hipposideros, complemented by a wealth of type specimens, and is principally concerned with the classification of the genus at the specific and higher levels. Detailed monographic treatment at the subspecific level would entail larger series than are at present available for many subspecies and to avoid uneven treatment of the genus has not been under- taken, although wherever possible some note has been taken of subspecific variation. Attention has been directed explicitly to the determination of the relationships between the species and species groups of this large genus and for this purpose detailed study has been concentrated at the level of the species. The results of this study are presented in the form of diagnoses of the numerous species included within Hipposideros, often based on the original material, in many cases supported by a further series of specimens, and of the species groups into which the genus can be divided. It has been possible to determine relationships between species, especially among those hitherto inadequately described or poorly known and to determine the validity and extent of the supraspecific groupings within the genus. The subsequent study of these groupings and their relationships has suggested a classification of Hipposideros and has given some indication of phylogeny within the genus. All measurements are in millimetres : forearm lengths are given in the form of histograms in the text, while minima, maxima and means (in parentheses) of representative cranial dimensions appear in a terminal table. Genus HIPPOSIDEROS Gray, 1831 Hipposideros Gray, 1831 : 37. Type species by subsequent designation (Sclater (1901 : 116)) Vespertilio speoris Schneider. Hipposiderus Gray, 1834 : 53. Emendation. Phyllorrhina Bonaparte, 1837 : fascicule 21 (sub. Rhinolophus ferrum-equinum) . Type species by subsequent designation (Sclater (1901 : 116)) Rhinolophus diadema Geoffroy. Macronycteris Gray, i866a : 82. Type species by monotypy Rhinolophus gigas Wagner. Gloionycteris Gray, i866a : 82. Type species by monotypy Rhinolophus armiger Hodgson. Rhinophylla Gray, i866a : 82. Type species by monotypy Phyllorrhina labuanensis Tomes. 4 J. E. HILL Preoccupied by Rhinophylla Peters (i865a : 355, 18655 : 520), a genus of Phyllostomatidae. Speorifera Gray, i866a : 82. Type species by monotypy Hipposideros vulgaris Blyth = Rhinolophus larvatus Horsfield. Chrysonycteris Gray, i866a : 82. Type species by monotypy Hipposideros fulvus Gray. Doryrhina Peters, iSyia : 324. Type species by monotypy Phyllorrhina cyclops Temminck. Sideroderma Peters, iSyia : 324. Type species by monotypy Phyllorr hina fuliginosa Temminck. Ptychorhina Peters, iSjia, : 325. Type species by monotypy Rhinolophus caffer Sundevall. Cyclorhina Peters, iSyia : 326. Type species by subsequent designation (Tate (1941 : 354)) Phyllorhina obscura Peters. Thyreorhina Peters, iSyia : 327. Type species by monotypy Phyllorhina coronata Peters. Syndesmotis Peters, iSyia : 329. Type species by monotypy Phyllorrhina megalotis Heuglin. Hipposiderus Blanford, 1888 : 637. Emendation. Taxonomic history of the genus The early generic history of Hipposideros is confused and the availability of the names employed for the genus has been discussed by Blanford (1888 : 637), Palmer (1904 : 327, 535) and Oey & Feen (1958 : 226). The first generic name to be applied exclusively to a horseshoe bat was Rhinolophus Lacepede, 1799, and this name, for type species Vespertilio ferrum-equinum Schreber, 1774, the European greater horseshoe bat, was gradually brought into general use for ah 1 species of horseshoe bat as each was discovered and made known to science. Later, Leach (1816 : 5) introduced Phyllorhina for type species (by monotypy) Vespertilio minutus Montagu, 1808, the lesser horseshoe bat of Great Britain and Ireland, listing a specimen presented to the British Museum by G. Montagu with the citation : Phyllorhina minuta Small Leafnose Torquay, Devon. G. Montagu, Esq. This citation follows that for Rhinolophus ferrum-equinum. The combination Phyllorhina minuta employed by Leach is listed by Miller (1912 : 149) and EUerman & Morrison-Scott (1951 : 115) as a nomen nudum and as a possible synonym of Rhinolophus hipposideros hipposideros (Bechstein), 1800. However, Leach has clearly applied a new generic name to the species described by Montagu, now called Rhinolophus hipposideros minutus. Oey & Feen (1958 : 226) state that Leach employed Phyllorhina as a subgeneric name within Rhinolophus for the lesser horseshoe bat from England, retaining the greater horseshoe bat within the nominate subgenus. Leach, however, used both names in the generic sense, calling the greater horseshoe bat Rhinolophus ferrum-equinum and the lesser horseshoe bat Phyllorhina minuta. As Oey & Feen point out, no subsequent author has arranged Rhinolophus ferrum-equinum and Rhinolophus hipposideros (of which Vespertilio minutus Montagu, 1808 is now considered to be the British subspecies) under different genera or subgenera. Phyllorhina Leach, 1816, must therefore be considered a junior synonym of Rhinolophus Lacepede, 1799. A REVISION OF HIPPOSIDEROS 5 Gray (1831 : 37) first recognized the characteristic distinctions of the hippo- siderine noseleaf , and with a valid diagnosis proposed the generic name Hipposideros for a number of Asiatic species hitherto referred to Rhinolophus. Sclater (1901 : 116) designated Vespertilio speoris Schneider as its type species. In a later work, Gray (1834 : 53) referred to this genus, using the amended spelling Hipposiderus. Peters (i87ia : 312) and Dobson (1876 : 58, 1878 : 127) have quoted this name and reference in their synonymies of the genus and were apparently unaware of the earlier citation as Hipposideros. Later, Gray (1838 : 492) reverted to the original spelling and provided a further brief diagnosis. Hipposideros Gray, 1831, however, did not gain general acceptance and a number of authors, including Peters (i87ia : 312) and Dobson (1876 : 58, 1878 : 127) employed Phyllorrhina Bonaparte, 1831, often rendering the name as Phyllorhina. This name is generally cited from C. L. Bonaparte, 1831, Saggio di una distribuzione metodica degli Animali Verte- brati. Two versions of this work exist in the Library of the British Museum (Natural History). It was published in two sections, the first concerned principally with the homoiothermic animals and the second the poikilothermic animals. The second section includes a further part giving additions and corrections to the first section. The first publication of this work is apparently in an Italian journal, Giornale Arcadico di Scienze, Lettere ed Arti, published quarterly in Rome. The first section appears in the issue for January, February and March, 1831, tome 49, part i, pp. 3-77, the part number referring only to the number of issue for any one year and appearing only on the binding. The second section appears in the issue for October, November and December, 1831, tome 52, part 4, pp. 129-209. In this version of the work the name (spelt Phyllorrhina) appears on p. 15 as follows, 40 referring to its position in the table of genera and 16 to the number of species the genus includes : 40. Rhinolophus, Geoffr. Eur. As. Af. Oc. 16. 1. Rhinolophus, Leach 2. Phyllorrhina, Leach The second version of this work in the Library of the British Museum (Natural History) consists of two separately bound sections, corresponding to the two sections of the work as published in the Giornale Arcadico. The first section is almost identical with that in the Giornale Arcadico, but has a title page, the heading Scienze omitted and has been separately paginated. It is dated 1831 and was printed by Antonio Boulzaler, the printer of the Giornale Arcadico. The entry quoted above appears on p. 16 of this version of the work, and this apparently is the version quoted by Peters (i87ia : 312), Dobson (1876 : 58, 1878 : 127), Blanford (1888 : 637), Palmer (1904 : 535), Sherborn (1929 : 4932) and Oey & Feen (1958 : 226) who without exception cite the name as Phyllorhina. The second section of the work has been similarly treated in the second version, with the addition of a title page, the omission of the heading Scienze and with separate pagination, but is dated 1832, suggesting that this separately bound version, quoted by authors, is but a reprint of the version printed in the Giornale Arcadico. It is not clear from the citation whether Bonaparte in this work employed Phyllorrhina as a subgenus 6 J. E. HILL of Rhinolophus in the sense of its generic usage by Leach (1816 : 5) or in the sense of Hipposideros as proposed by Gray (1831 : 37). In any case, the absence of diagnosis or of type species invalidates its proposal as a new name, as recognized by Palmer (1904 : 535), Sherborn (1929 : 4932) and Oey & Feen (1958 : 226). Bonaparte (1837 : fascicule 21) under Rhinolophus fermm-equinum, revived Phyllorrhina as a subgeneric name for the first section of Rhinolophus as listed by Temminck (1835 : I ~ 2 4)> corresponding to the genus Phyllorhina of subsequent authors. Bonaparte gives a brief diagnosis and Sclater (1901 : 116) has designated Rhinolophus diadema Geoffrey as its type species. It was subsequently raised to generic rank by Peters (1852 : 31) who cited it, however, from Bonaparte (1831). Blanford (1888 : 637) rejected Phyllorrhina Bonaparte, 1837 on the grounds that Hipposideros Gray, 1831 had priority and that the name, which he rendered as Phyllorhina, had been originally proposed by Leach for a species of Rhinolophus and could not therefore be used for another genus. Palmer (1904 : 535) and Sherborn (1929 : 4932) reject it on ground of homonymy with Phyllorhina Leach, 1816, although both correctly citing the name as Phyllorrhina. Oey & Feen (1958 : 226) reject the name since at that time the alteration in spelling was considered insufficient under the International Code of Zoological Nomenclature to revive an unavailable generic name. However, Article 56 (a) of the International Code of Zoological Nomenclature, 1961 states explicitly that two genus-group names are not to be considered homonyms even if the difference between them is due only to one letter, although it is recommended (Appendix D.3) that the proposal of such names should be avoided. Under these circumstances, therefore, Phyllorrhina Bonaparte, 1837, cannot be held to be a homonym of Phyllorhina Leach, 1816, and must be regarded as available within Hipposideros if subgeneric division of that genus is required. Earlier students of the genus or of parts of it include Temminck (1835), Gray (1838, i866a), Peters (i87ia), Dobson (1876, 1878), Andersen (1905, igoGa, 1906!}, 1918), Tate (1941) and Aellen (1952, I956a, I956b). Gray (1831) originally asso- ciated seven Asiatic species with the genus Hipposideros. Temminck (1835) reviewed the genus Rhinolophus in some detail, but made no formal recognition of the separation of that genus introduced by Gray. This author instead separated Rhinolophus into two sections, the first of these corresponding to Hipposideros of Gray (1831) or to Phyllorhina of later authors such as Peters (i87ia) and Dobson (1876, 1878), adding a number of species to those attributed to Hipposideros by Gray and including an African species for the first time. Gray (1838) gave a brief review of Hipposideros and added a number of Asiatic species to the genus as listed by Temminck. Later, Gray (i866a) proposed six new generic names for species hitherto referred to Hipposideros. Peters (i87ia) reduced these to subgenera and proposed six further subgeneric names. This work, the first critical study of the genus, thus greatly divided it but at the same time grouped under numerous sub- generic names the many species and their synonyms that had accumulated since Temminck and Gray first studied the genus as a whole. Gray (i866a) and Peters (i87ia) wete principally concerned with the diagnosis of the species groups into A REVISION OF HIPPOSIDEROS 7 which Hipposideros is divisible, and although no great use has been made by other authors of subgenera within the genus, their subgenera in a number of cases have been recognized as species groups by most subsequent authors. Dobson (1876, 1878) placed the names proposed by Gray in the synonymy of Phyllorhina (= Hipposideros} and did not mention the subgeneric groupings of Peters, but at the specific level greatly amplified the revisionary work begun by Peters and provided detailed descriptions of the ears and noseleaves of individual species. Andersen (1905, I9o6a, I9o6b) began the first modern detailed revisionary work on the genus with studies of the diadema, armiger, commersoni and coffer groups, and in Andersen (1918) Oldfield Thomas published on his behalf a compilation of diagnostic characters presented in the form of keys to the species and subspecies of some of the groups of Hipposideros, including brief diagnoses of a number of new forms. This work was probably an extract from a detailed revision upon which Andersen was working at the time of his disappearance and which was never completed. Tate (1941) in a comprehensive review of much of the genus greatly amplified the work of Gray, Peters and Andersen in the definition and diagnosis of its major groups, retaining for the most part the divisions originally proposed by Gray and Peters and allocating to them the species and subspecies more recently proposed. Tate, however, was primarily concerned with the Asiatic species of the genus and made no detailed survey of its African representatives, touching only briefly on their relationships to their Asiatic congeners. His work is nevertheless the only extensive survey of the genus since it was studied by Peters (1871 a) and Dobson (1876, 1878) and has provided an excellent basis for its further revision. Although subsequent authors, notably Aellen (1952, i956a, i956b), have made valuable contributions to our knowledge of Hipposideros, especially to that of its African members, no student since Peters (i87ia) and Dobson (1876, 1878) has attempted revisionary study of the genus as a whole, and it now stands badly in need of synthetic study. Morphological criteria in the genus The external features of the genus Hipposideros have been described by Dobson (1878 : 127) and the principal features of its skull by Miller (1907 : in), while Tate (1941 : 357) has reviewed the major diagnostic criteria of importance within the genus. As might be expected of so large a genus, it displays a wide range of variation both externally and cranially. The ears, exceptionally united at the base by a low band of integument, vary in outline from comparatively short, broad and rounded or bluntly pointed to long, narrow and with an acute point. Their anterior or inner edge is generally convex, their posterior or outer edge straight or with a shallow concavity or emargination just behind the tip. Some species exhibit a well-defined internal fold at the antitragal lobe, in others absent or represented by a thickening of the membrane of the ear at this point. The external surface of the ear is usually naked except at its base but in some species can be covered with body fur for one half to two thirds the length of the ear. The nasal foliations are complex and consist basically of three sections, in this paper described as the anterior, 8 J. E. HILL intermediate and posterior noseleaves. The noseleaf in many species is flanked by one or more lateral supplementary leaflets, of which in some cases one may extend anteriorly beneath the anterior leaf on to the upper lip, sometimes forming a complete supplementary leaflet encircling the muzzle beneath the anterior leaf. The anterior noseleaf, extending over the upper lip, is the horseshoe of earlier authors, and is rarely much modified but occasionally displays a narrow median emargination. The nostrils open in paired depressions in the centre of this leaf, the narial openings separated by an internarial septum which is usually narrow or only slightly inflated but which may be bulbous, inflated or specialized to form a disc-like structure between the nostrils. Lateral narial lappets project from the outer walls of the narial depressions and together with the internarial septum may become modified to form deep pockets in which lie the narial apertures. The intermediate part of the noseleaf lies immediately behind the narial depressions, forming their posterior boundary, and is a cushion-like structure extending transversely across the entire noseleaf. It may be variously modified by the development of raised median and lateral eminences or ridges, or of a median club-like structure. Its posterior margin forms the base of the posterior leaf, an erect structure with a smooth or sometimes slightly lobulated upper edge, usually convex in outline but on occasion slightly triangular or specialized by the development of a median projection. The anterior face of the posterior leaf is more or less concave, the concavity smooth or divided by one or more vertical septa, enclosing small cells or pockets. The posterior face is usually smooth but in some species is modified by the development of a transverse supplementary structure with a serrated upper edge. The nasal foliations exhibit an exceptionally wide range of variation within the genus from small, comparatively simple structures showing little or no evidence of specialization to large, greatly modified structures completely covering the entire muzzle. A frontal sac, usually less developed in female specimens than in male, with its opening behind the posterior noseleaf, is to be found in a number of species, while a few species are noted for the development, especially in males, of transverse fleshy lobate prominences on each side of the opening of this sac. The skull demonstrates a similarly wide range of variation, especially in the rostral, palatal and sphenoidal regions. Its basic outline varies from elongate and narrow, the zygomatic width less than or equal to the mastoid width, to short and comparatively broad with the zygomatic width exceeding the mastoid width of the skull. The braincase is generally elongate and never globose, and in some species is somewhat inflated. Sagittal and lambdoid crests are usually present and in the larger species are often greatly developed. The interorbital region is usually markedly constricted but exceptionally is broader with no sharp constriction between the braincase and the rostrum. The supraorbital ridges in some species are low and poorly defined and in others are well defined and prominent : they may partially enclose a frontal depression, itself sometimes absent. The rostrum, rounded in outline in some species, is in others more markedly pentagonal. It exhibits paired, inflated rostral eminences anterior to the anteorbital region, separated from each other by a shallow groove. The rostrum is expanded laterally to a greater or lesser A REVISION OF HIPPOSIDEROS g degree, and in some species is markedly flattened and more greatly ossified. Con- siderable variation is displayed in the form of the premaxillae, which basically form a projecting structure with a V-shaped or U-shaped junction with the maxillae. Considered together, they vary from a narrow oblong structure not greatly expanded posteriorly at its junction with the maxillae to a wide structure almost filling the anterior palatal emargination with a wide, sometimes fan-shaped union with the maxillae. Their lateral edges may be deeply notched so that with the maxillae they form the walls of the anterior palatal foramina. In some species, delicate anterior enclosing processes are developed to form the anterior walls of these foramina : these processes in other species completely enclose the foramina which are thus contained within the premaxillae. The anterior palatal foramina in some species are small and rounded and in others are large and oval, elongate or slit-like. The palate is short and broad, the palation usually more or less U-shaped or square, with or without a median emargination or post-palatal spicule. The mesopterygoid fossa is wide and the pterygoids vary considerably in relative length between species. The sphenoidal bridge, flanked by rounded or elongate lateral apertures, usually partially conceals them but in some species is very markedly constricted. A sphenoidal depression is usually present. The cochleae typically are approxi- mately equal in width to their distance apart but exceptionally may be greatly enlarged so that their width is equal to six or eight times this distance. The upper incisors are usually bilobed : the outer lobe, however, is present in varying degrees of obsolescence and in some species is virtually obsolete. The upper canines sometimes have low anterior or posterior cusps. The anterior upper premolar (pm 2 ) is small or minute, often extruded outwards from the toothrow so that the canine and the second upper premolar (pm 4 ) are in contact or nearly so : rarely the anterior upper premolar (pm 2 ) is absent. The posterior cusp of the third upper molar is usually obsolescent or obsolete and its third commissure undeveloped, with the W-pattern of the tooth incomplete : exceptionally the third cusp may be more or less unreduced and the commissure present with the W-pattern of the tooth virtually complete. The crown area of the outer lower incisors in some species is less than or equal to the crown area of the inner teeth : in other species it is greater than the crown area of the inner teeth. The anterior lower premolar (pm 2 ), sometimes almost equal in size to the second lower premolar (pm 4 ), is more usually reduced, sometimes to one quarter or one third the size of the second tooth. Supraspecific groupings within the genus Tate (1941 : 355) has rightly pointed out that the characters of the numerous species currently included within Hipposideros seldom combine to indicate clearly evident evolutionary trends. As this author says, such evidence in this genus is perplexing and often contradictory, and the morphological criteria in Hipposideros apparently represent tendencies latent in perhaps all of its species, active in some and quiescent in others. Similarly, such criteria combine but rarely to form an aggregate of features defining natural groups of species within the genus. Despite this, Peters (iSyia) was able to discern twelve subgenera in Hipposideros (albeit io J. E. HILL with fewer species under consideration than are currently included in the genus) and Tate (1941) was able to reduce these only to eleven major species groups, while Aellen (1954) has added a further group to those denned by Tate. A review of the morphological features of these groups indicates that considerable similarity exists between several of them, and that some at least may be separated only with difficulty from each other. It is clearly evident that the great degree of subdivision advocated by Peters is wholly unjustified and that subgeneric division at this level is unwarranted. It is possible to discern three primary divisions in Hipposideros. One, in general a more primitive section, appears to represent the basic group of the genus from which the other, more specialized sections have been derived. It has itself developed a number of specialized forms as well as retaining species exhibiting comparatively simple and unspecialized features. Members of the groups allocated to this section of the genus are generally of small size, with broad, usually rounded ears, often modified by the presence of an internal fold or thickening at the antitragal lobe, while in most the noseleaves are comparatively simple. Lateral supplementary leaflets may be absent : one leaflet may be incipient or present and in some two lateral leaflets are to be found. The skull is more or less elongate and narrow, with an inflated braincase, the zygomatic width less than or not greatly exceeding the mastoid width. The upper incisors are usually weak and lack much of their outer lobe, while the crown dimensions of the outer lower incisors only exceptionally greatly exceed those of the inner lower incisors. The megalotis, bicolor, calcaratus and galeritus groups of Tate (1941) and the curtus group of Aellen (1954) fall into this division. Of these, the megalotis group, containing only the African species megalotis, is perhaps the most primitive. Despite the uniquely conjoined ears of H. megalotis and its large outer incisors, which in crown area greatly exceed the inner incisors, such characters as its small, simple noseleaf lacking lateral leaflets, and its elongate skull with inflated braincase and weak upper incisors suggest that it is not far removed from the basal stem of Hipposideros. The bicolor and calcaratus groups of Tate (1941) are for the most part little more specialized, their species usually with simple noseleaves which either lack lateral leaflets or have one leaflet only, sometimes in incipiency. They have an elongate, narrow skull similar in outline to that of H. megalotis. The galeritus group of Tate (1941) represents a more specialized derivative of bicolor, its species with two lateral leaflets and a more specialized, shorter, wider skull. The bicolor and galeritus groups as understood by Tate (1941) include a number of isolated species, usually monotypic and in some cases displaying complex developments of the noseleaves. These, although in some instances representing independent but relatively minor lines of modification, in others are highly specialized offshoots of the bicolor and galeritus types, or of the stem connecting them. In many respects such species connect the two groups, while the calcaratus group appears to be a derivative of the bicolor type. In the present work, the bicolor, calcaratus and galeritus groups as they are defined by Tate (1941), together with the curtus group of Aellen (1954), are united to form a single group, for which the earliest name is bicolor. A REVISION OF HIPPOSIDEROS II The second primary division of the genus contains only the highly specialized cyclops and muscinus groups of Tate (1941), here united to form a group for which cyclops is the prior name, restricted to Papua, northern Australia and West Africa. This group exhibits modifications of the noseleaves so peculiar that despite the curious distributional pattern of its members they must be considered to share a common if remote origin. It shares some features with the bicolor group, notably the small size of some of its members, but the species allocated to the cyclops group have longer, narrower ears, sharply triangular, with little or no antitragal modifica- tion. Their noseleaves have two lateral leaflets and are uniquely distinguished in the genus by the second lateral leaflet, which forms an integral part of the posterior leaf and in Australasian species extends anteriorly beneath the anterior leaf, over the upper lip. The noseleaves are further specialized by the development of median tubercles or club-like processes from the intermediate and posterior noseleaves. The skull is less elongate and comparatively wider than in the megalotis and bicolor groups : the upper incisors are weak and the outer lower incisors are very slightly larger in crown area than the inner lower incisors. The least specialized members of the group share some degree of affinity with the bicolor group and it seems likely that the origins of the cyclops group lie remotely with the bicolor group of species. A third division of Hipposideros is represented by the pratti, armiger, speoris, diadema and commersoni groups of Tate (1941). Their members are characterized principally by their larger size, their smaller, triangular ears, which usually lack any antitragal modification, by their comparatively simple noseleaves, which have two or more commonly three or four lateral leaflets, and by their comparatively shorter, broader skulls, which have wider, more expanded zygomata. In the species allocated to these groups the upper incisors are stout and retain much of the outer lobe, while the crown area of the outer lower incisors is greater than that of the inner lower incisors, sometimes greatly so. Monotypic species are rare among these groups, and there are no species with the exotic modifications such as are to be found in the first and second divisions. There are, however, some indications of specialization in the presence of greatly developed transverse supplementary lappets behind the posterior leaf in the pratti group and in the slightly trilobate posterior leaf of the armiger group. The commersoni group of Tate (1941) appears to be the African representa- tive of the Austro-Malayan diadema group as understood by that author, and in the present work these have been united to form a single group for which the prior name is diadema. The primary divisions of the genus indicate three general but distinct evolutionary trends within Hipposideros. One, exemplified by the megalotis and bicolor groups, is towards small or medium size and development and proliferation of the noseleaves, combined with a corresponding increase in the size of the ears and only exceptionally with extensive cranial modification, the skull generally elongate and not greatly widened. The aberrant cyclops group represents a second trend sufficiently differen- tiated as to justify its separation from that shown by megalotis and bicolor, with greatly developed noseleaves and much modification of the ears and auditory region of the skull. This group displays a number of unique features, especially in the 12 J. E. HILL combination of small size, large ears and complex noseleaves with a broadened skull in its Australasian species and in the combination of large size, large ears and complex noseleaf with a very wide skull in its West African species. A third trend, exempli- fied by the pratti, armiger, speoris and diadema groups, is towards greater size, without extensive increase in the size and complexity of the ears and noseleaf, but with corresponding broadening of the skull. Each of these divisions include subsidiary lines of development, and while this view is perhaps an over-simplification of major evolutionary trends in Hipposideros it is clear that the genus cannot be divided readily into more primitive and more advanced groups of species. It can only be stated in general terms that these divisions represent three more or less parallel lines of development within the genus. However, the megalotis-bicolor section includes a number of species which are not greatly specialized, and is therefore to be regarded as the more primitive. It has at the same time developed species which in many of their features are as highly developed as those included in the cyclops section or in the section containing the pratti, armiger, speoris and diadema groups. The three primary groups express to some extent at least the major lines of development that have occurred within Hipposideros : the divisions between them, however, are not clearly defined and there is manifestly no justification for the extreme subdivision of the genus practised by Peters (1871 a) in his recognition of twelve subgenera of Hipposideros. The genus appears instead to consist of an aggregation of loosely defined groups of species, each exhibiting varying combinations of characters, some highly specialized. Tate (1941) recognized this and divided the genus into eleven species groups but did not study its African species in detail. In the present work these have been related to Asiatic species groups and through the combination of some of the groups of Tate the varying levels of development within Hipposideros have been expressed by the division of the genus into seven species groups, one divided into two subgroups. The groups are listed in the follow- ing summary, with their included species, excluding doriae Peters, probably identical with sabanus and which therefore has been excluded from the following discussion. megalotis group megalotis Heuglin bicolor group bicolor subgroup galeritus subgroup bicolor Temminck pygmaeus Waterhouse ater Templeton galeritus Cantor fulvus Gray breviceps Tate cineraceus Blyth curtus Allen nequam Andersen fuliginosus Temminck calcaratus Dobson coffer Sundevall cupidus Andersen beatus Andersen coronatus Peters coxi Shelford ridleyi Robinson & Kloss papua Thomas & Doria jonesi Hayman dyacorum Thomas A REVISION OF HIPPOSIDEROS 13 bicolor group bicolor subgroup (contd.} sabanus Thomas obscurus Peters marisae Aellen cyclops group cyclops Temminck camerunensis Eisentraut muscinus Thomas & Doria wollastoni Thomas semoni Matschie stenotis Thomas pratti group pratti Thomas lylei Thomas armiger group armiger Hodgson turpis Bangs speoris group abac Allen larvatus Horsfield speoris Schneider diadema group lankadiva Kelaart schistaceus Andersen diadema Geoffroy dinops Andersen inexpectatus Laurie & Hill commersoni Geoffroy Forty-three species are recognized, of which it has been possible to examine all but coronatus and marisae. The bicolor group as here understood includes the bicolor, galeritus and calcaratus groups of Tate (1941) and is a complex of related species not readily separable into different groups. The cyclops group listed by Tate (1941) has been incorporated into the muscinus group as recognized by that author to form a cyclops group, while the commersoni group of Tate (1941) is closely related to the diadema group and has been united with it. Relationships between the groups are briefly summarized in the form of a diagram (Fig. i). The genus is remarkable for its high content of monotypic species, with no fewer than twenty-seven of its forty-three species not divided into subspecies. Of the remainder, seven are divided into two subspecies and one into three, while only eight species have more than three subspecies. The distribution of subspecies in the species groups of Hipposideros is shown in Table i. The megalotis, bicolor and cyclops groups contain by far the majority of monotypic species and while some of J. E. HILL FIG. i. Group relationships in Hipposideros 15 these are clearly closely related, others are separated from each other by very distinctive external and cranial features. Rather than representing comparatively recent developments within the genus, from the wide divergence of their isolating characters and their usually restricted distribution, these species appear to be relicts of a rather remote phase of radiation within Hipposideros. The three species groups themselves support this contention, their considerable dissimilarities suggesting their separation at an early stage in the evolution of the genus. The megalotis group is monotypic and of very restricted distribution, while the bicolor group contains a wide diversity of loosely allied species, few of them successful and wide- spread, and many apparently representing independent lines of development. These groups are widely separated from the cyclops group, also of restricted distribu- tion and so distinctively specialized as to demand a long period of modification. The bicolor and cyclops groups have both Asiatic and African representatives, whose wide separation from each other in a number of basic features indicates that the partition of each parent group into Asiatic and African sections now no longer continuous was evidently a remote event. African representatives of the bicolor group are rather less distantly removed from their Asiatic congeners than are the African cyclops and camerunensis from the Australasian muscinus and its allies and the bicolor group includes a few species demonstrating to a limited extent the connection between its Asiatic and African species. However, the evidence of this connection is tenuous and the development of the group in the Asiatic and African regions apparently represents two independent, parallel lines of modification from a common but remote origin, often displaying considerable similarity and conver- gence, and with a number of independent offshoots in each region. CO CO w oj CO ... 0) .CD g CD g 'o 'CD i CD ^ 11^ X. PH -(-> ft* 3 CO w w w *M O *O -H MH -H *+H -*-* O d a? "S d W H CD t-i JH & J2 M If T3 '^ ^ Qj 'CD S T3 ^ co S'S a g ^2 co a ^ ^ ST^J *H '71 ""* S Sn 8 CD a o 3 ^ 'S J3 ^ r^ ^ ^ 2 o f-\ CO fc a K> "-! 3 (^ 'O CO ^ r 9 co ^ ^ 4- CO megalotis group I i - - - bicolor ,, 23 16 3 - 4 cyclops 6 6 - - - pratti ,, 2 2 - - armiger ,, 2 - i I speoris ,, 3 I i - i diadema ,, 6 I 2 3 Totals 43 27 7 I 8 TABLE i. Distribution of species and subspecies among the species groups of Hipposideros. 16 J. E. HILL The pratti, armiger, speoris and diadema groups contain fewer isolated, monotypic species and represent two allied lines of modification developed independently but more or less parallel to the development of the megalotis, bicolor and cyclops groups. One, exemplified by the wholly Asiatic pratti and armiger groups, is towards specialization of the noseleaves and rostral region. The pratti group, with two species, both monotypic, is of very restricted distribution and is evidently related rather distantly to the more widespread armiger group. The other trend includes the speoris and diadema groups, and is towards great size and its corresponding modifications of the skull, both groups having representatives in the Asiatic and African regions. In the speoris group, less specialized than diadema, the sole African species is less closely related to its Asiatic congeners than the African representative of the diadema group is to the Asiatic species of the group, and the speoris and diadema groups apparently represent a rather more recent phase of radiation than do the pratti and armiger groups. These four groups differ widely from the megalotis, bicolor and cyclops groups, and it is evident that their separation occurred at a very early stage in the evolution of the genus. The seven species groups into which the genus is divided in the present paper may be keyed : 1 Ears united at the base by a low frontal band . . . megalotis group (p. 17) Ears not united ............ 2 2 Ears long and narrow, pointed (Figs. 20, 22-25) : cochleae greatly expanded, their width equal to at least four times their distance apart . cyclops group (p. 72) Ears short, broad, rounded or triangular : cochleae not expanded, their width approxi- mately equal to their distance apart ........ 3 3 Ears rounded or broadly triangular, bluntly pointed : upper incisors weak, the outer lobe obsolescent or obsolete ; crown area of outer lower incisors equal to or only slightly greater than that of the inner lower incisors . . bicolor group (p. 18) Ears triangular, pointed : upper incisors strong, usually retaining much of the outer lobe ; crown area of the outer lower incisors much greater than that of the inner lower incisors ............ 4 4 Noseleaf with two lateral supplementary leaflets : frontal depression well-defined ; maxillae elongated ........ pratti group (p. 87) Noseleaf with three or four lateral supplementary leaflets : frontal depression lacking or shallow ; maxillae not elongated ........ 5 5 Noseleaf with upper edge of posterior leaf slightly lobate, the posterior leaf narrower than the anterior leaf : rostral eminences not inflated and rostrum flattened ; a distinct discontinuity between roofs of narial and mesopterygoid canals armiger group (p. 91) Noseleaf with upper edge of posterior leaf forming an arc of a circle, the posterior leaf equal in width to the anterior leaf : rostral eminences moderately inflated and rostrum elevated ; roofs of narial and mesopterygoid canals not sharply discontinuous ............ 6 6 Ears with a small projection at the antitragal fold : sphenoidal bridge narrow, not concealing lateral apertures ; pterygoid wings undeveloped speoris group (p. 94) Ears without antitragal modification : sphenoidal bridge wide, partially concealing lateral apertures ; pterygoid wings expanded . . diadema group (p. 103) A REVISION OF HIPPOSIDEROS 17 HIPPOSIDEROS MEGALOTIS group This group contains only the isolated African species Hipposideros megalotis (Heuglin), which may be distinguished at once from all other species of the genus by the low band of integument uniting the ears at their base. Peters (iSyia : 329) proposed subgeneric recognition for megalotis under the name Syndesmotis, largely on account of its conjoined ears, but Dobson (1878 : 151) ignored this separation and concluded from an examination of the type specimen that the species belonged to that section of Hipposideros typified by H. bicolor and its associated species. Senna (1905 : 275) provided a detailed account of two topotypical specimens of megalotis and raised Syndesmotis of Peters to generic rank. His diagnosis of Syndesmotis was the first comprehensive definition to appear and was based on the conjoined ears, notched, sinuous upper edge of the posterior noseleaf, low cranium, undeveloped sagittal crest, high occipital, elevated frontal, narrow basioccipital and the absence of the anterior upper premolars (pm 2 - 2 ) in megalotis, characters which he considered to render the species genetically distinct from Hipposideros. However, apart from the first, these features are to be found in one or other of the remaining species groups of Hipposideros, and no later author has employed Syndesmotis in the subgeneric or generic sense. Tate (1941 : 357) first listed the species as H. (Syndesmotis) megalotis and postulated a relationship to the Megader- matidae but later (1941 : 359) listed megalotis as the sole species of the megalotis group. Hayman (1954 : 285) reviewed earlier accounts of the species, which hitherto had been known only from the type locality, Keren, in Eritrea, and extended its range to the Kenya highlands. Later (1960 : 61) the same author recorded the species from Ethiopia. There is little to justify the separation of megalotis from Hipposideros. Despite its uniquely specialized ears, and its expanded bullae, which are reminiscent of the cyclops group, megalotis is clearly a primitive species of the genus, allied in many respects to the less specialized species of the bicolor group. Such characters as its small, simple noseleaf lacking lateral leaflets, elongate skull with inflated braincase and narrow rostrum and its weak upper incisors display its affinities with H. bicolor and its allies. At the same time, an evidently long period of modification has led to the development of a greatly specialized auditory region, and to specialization of the dentition in the loss of the anterior upper premolars (pm 2 - 2 ) and considerable increase in the dimensions of the outer lower incisors, which greatly exceed the inner lower incisors in crown area. These features, together with its uniquely joined ears, adequately justify the position of megalotis as the sole member of an individual group of Hipposideros. Hipposideros megalotis (Heuglin) Phyllorrhina megalotis Heuglin, 1862 : 4, 8, Keren, Eritrea. The ears of Hipposideros megalotis are large, rounded, their posterior margins without a concavity behind the tip, and have a small internal fold at the antitragal lobe : they are haired for a little less than one half of their length. They are joined zoo 11, i. B i8 J. E. HILL at the base by a low band of integument. The noseleaf is small and simple, and lacks lateral leaflets : the anterior leaf is unemarginated, the intermediate part of the leaf uninflated, with a low median eminence, while the posterior leaf is moderate, supported by three poorly defined ridges and with its upper edge slightly sinuous, more or less semicircular. The skull is small and elongate, with an inflated braincase and narrow rostrum. The sagittal crest is low and there are no supraorbital ridges. The frontal region is slightly elevated and lacks a frontal depression : the rostral eminences are moderately inflated. The zygomata are moderate, with a low jugal projection : the zygomatic width is less than the mastoid width. The anteorbital foramina are large and rounded, closed by a narrow bar of bone. The junction of the premaxillae with the maxillae is U-shaped and the incisive foramina are rounded and not enclosed within the premaxillae, their antero-posterior walls formed by the maxillae. The palation is U-shaped, with a median emargination, and the vomer projects slightly into the mesopterygoid fossa. The sphenoidal bridge is moderate and flanked by large, rounded lateral apertures. There is a shallow sphenoidal depression : the basioccipital is narrow and the cochleae large, their width equal to approximately three times their distance apart. The upper incisors are widely spaced and weakly bilobed, the outer lobe a little smaller than the inner lobe. The upper canines are slender, with a well-developed posterior cusp. The anterior upper premolar (pm 2 ) is absent and the posterior ridge of the third upper molar obsolescent. The outer lower incisors are much larger in crown area than the inner pair and the anterior lower premolar (pm 2 ) is one half the height and three quarters or more the length of the second lower premolar (pm 4 ). APPROXIMATE DISTRIBUTION : Eritrea ; Ethiopia ; Kenya. 34 36 38 H. MEGALOTIS 34 36 38 FIG. 2. Length of forearm in Hipposideros megalotis HIPPOSIDEROS BICOLOR group The bicolor group as here understood contains the bicolor, calcaratus and galeritus groups of Tate (1941) together with a number of species not studied or allocated to group by that author. The ears of members of the bicolor group are large, broad, rounded or more or less triangular in outline, bluntly pointed, with an internal fold or a thickening of the membrane of the ear at the antitragal lobe. Noseleaves within the group vary from the relatively unspecialized and simple structures of bicolor and its immediate allies to the complex, greatly developed, sometimes bizarre foliations of such species as jonesi or coxi. The number of lateral supple- mentary leaflets varies from none or incipiently one in the more primitive species A REVISION OF HIPPOSIDEROS 19 of the group to one or more usually two in those more specialized, while a somewhat anomalous species allocated to the bicolor group, has an incipient, very poorly developed third leaflet. Species allocated to this group have elongate, narrow skulls, with moderately inflated braincases and narrow rostra and an un- specialized auditory region, the cochleae never more than a little wider than their distance apart. Their upper incisors are weak, the outer lobe obsolescent or obsolete, while the crown dimensions of the outer lower incisors are less than or only slightly greater than those of the inner lower incisors. There is no good reason to divide the bicolor group into separate species groups : the bicolor and calcaratus groups of Tate (1941) clearly represent an aggregation of more primitive species while the galeritus group and its derivatives as understood by that author consists of more specialized species whose connection with bicolor and its immediate allies in some cases can still be traced. The group is of remote origin and is the most wide-ranging of the genus, distributed in one form or another from Africa to the New Hebrides and northern Australia, although it is almost equalled in this respect by the diadema group, which has a similar distributional pattern but extends eastward only to the Solomon Islands and does not enter Australia. It is by far the largest group within the genus and while it includes a number of unspecialized species, the group at the same time includes species which have developed this basic pattern into highly specialized elaborations. The species allocated to the bicolor group may be divided into two loosely defined subgroups, corresponding approximately to the bicolor and galeritus groups of Tate (1941). The bicolor subgroup includes the calcaratus group of Tate (1941), which has few characters to distinguish it from bicolor and its allies, and is predomi- nantly Indo-Australian in distribution, with two species extending its distribution to Africa. Its members are in general less specialized than those of the galeritus subgroup and usually have broader, more rounded ears, which in most cases have an internal fold at the antitragal lobe and are haired for one half or less of their length. Lateral supplementary leaflets as a rule are absent or are exceptionally represented by a single small leaflet. The skull is elongate and narrow, with an inflated braincase, the zygomatic width rarely exceeding the mastoid width. Although the subgroup includes some of the least specialized species of the genus, it contains also such more advanced derivatives as dyacorum, sabanus and jonesi exhibiting a considerable degree of specialization. The galeritus subgroup is represented in the Asiatic and Australasian regions by galeritus and its associated species and in the Ethiopian region by coffer and its allies. Its members are characterized by their usually more or less triangular ears, which often display a concavity in their posterior margins just behind the tip of the ear. The antitragal fold is less prominent and the ears are usually haired for about two thirds of their length. The noseleaves are in general more specialized and have two or exceptionally three lateral supplementary leaflets. Members of this subgroup have shorter, broader skulls than the members of the bicolor subgroup, with the zygomatic width exceeding the mastoid width. As in the bicolor subgroup, some species of the galeritus subgroup exhibit great elaboration of its basic pattern. 20 J. E. HILL No definitive line of separation can be found between the subgroups, and they are linked by species exhibiting their respective characteristics in differing combinations. Hipposideros sabanus and H. dyacorum both lack lateral leaflets and have the comparatively simple noseleaf of the bicolor subgroup yet have the shortened, rather broadened skull characteristic of the galeritus subgroup. Hipposideros obscurus has likewise no lateral leaflets but has somewhat triangular ears and a short, broad skull. Hipposideros marisae has ears and noseleaf similar to those characteris- tic of the galeritus subgroup, but has one lateral supplementary leaflet and a some- what elongate skull. Hipposideros curtus has two lateral leaflets and cranially resembles the galeritus type, but has the ears haired for only one half of their length. Hipposideros fuliginosus and H. pygmaeus have two lateral leaflets but have the sparsely haired ears and elongate skulls of the bicolor type, while H. coxi has two lateral leaflets and ears haired for two thirds of their length yet has an elongate, unbroadened skull. Such species serve to demonstrate the close affinity of the members of the bicolor group and its essential unity. This treatment of the group greatly extends the views of Tate (1941 : 358, 366) who recognized that the African coffer and its allies should be associated with his Asiatic galeritus group, but did not relate these species and their associates to those he included in the bicolor group as he defined it. The allocation of the African species of the bicolor group as here understood contrasts sharply with the views of Aellen (1952 : 62), who divided them widely by the recognition of a coffer group for coffer and its associated species curtus, fuliginosus and beatus, and later (1954 : 479, 480) recognized additionally a jonesi group for the sole species jonesi and a curtus group for the species curtus and marisae. Hipposideros coffer and its allies, together with curtus, are closely related to galeritus, and in the present work are referred to the galeritus subgroup, while jonesi and marisae have affinities with the bicolor subgroup, to which they are referred. Possible relationships within the group are summarized in the form of a diagram (Fig. 3). Morphological characters in the group rarely combine to indicate definite trends, and the relationships of the numerous species comprising the bicolor group are not easily discerned and are difficult of definition. The considerable degree of elaboration of the noseleaves displayed by some species tends further to obscure their basic pattern. General tendencies discernible within the group are for the ears to become less rounded and more triangular, and for their posterior edges to develop a concavity just behind the tip. At the same time, an increase in the area of body fur extending over the outer surface of the ears can be seen, and the internal fold tends to become less prominent. Lateral supplementary leaflets are absent in the more primitive species of the group, more advanced species having one or more frequently two leaflets, while one species, papua, possesses a third leaflet, incipient and poorly developed. Cranially, the elongate, rather narrow outline of the skull in the primitive members of the group tends to become shorter and broader in those more advanced. These general trends, however, are obscured to a large extent by considerable overlap between a number of species, indicated in fig. 3 by enclosure between two parallel dotted lines. Such species exhibit varying combina- A REVISION OF HIPPOSIDEROS 21 FIG. 3. Possible relationships in the Hipposideros bicolor group 22 J. E. HILL tions of primitive and advanced morphological characters and must be regarded as transitional between the simplest members of the group and those most specialized, although some such as coxi display a high degree of individual specialization. A number of less general trends exist within the group and are confined in some cases to one or two species. Such a trend is found in ridleyi and jonesi, large species of the bicolor subgroup which have however developed the internarial septum to form a concave circular disc between and partially covering the nostrils : jonesi is more developed in this respect and additionally has a deeply pocketed posterior leaf reminiscent of that of coxi. Similarly, calcaratus and cupidus have developed a broad interorbital region and their zygomata are widened to an extent considerably exceeding the mastoid width, although in other respects both are very simple species with broad ears which possess an antitragal fold and with a simple noseleaf, lacking lateral leaflets. Hipposideros coxi has a highly complex noseleaf with two lateral leaflets, the anterior leaflet extending forwards beneath the anterior leaf to the median line and with the posterior leaf elaborated into a complex, deeply pocketed structure. Its skull, however, is comparatively elongate and narrow and basically is of the bicolor type. A trend towards galeritus is characterized by the retention of a skull architecturally rather like that of the bicolor type, although broadened, with long pterygoids and a short sphenoidal bridge. It can be traced through dyacorum, a simple species lacking lateral leaflets and with a comparatively elongate skull, obscurus, a species with a specialized noseleaf, no lateral leaflets and shortened skull and pygmaeus, similar to obscurus, but with a more specialized noseleaf and two lateral leaflets. The trend towards coffer is characterized by a shortening of the pterygoids and consequent extension of the sphenoidal bridge, and in some species by the development of a transverse, serrated structure from the rear face of the posterior leaf. It includes sabanus, a simple species lacking lateral leaflets but with a transverse supplementary structure behind the rear leaf and a short, broad skull with short pterygoids and fuliginosus, with two lateral leaflets, a trace of a supplementary structure behind the posterior leaf yet with an elongate skull with long pterygoids. This trend must include also marisae, with broad ears, one small leaflet and elongate skull, and curtus, with similar ears, two lateral leaflets and shorter, broadened skull with short pterygoids, two species further specialized by the considerable inflation of the internarial septum. Hipposideros papua, apparently derived from the stem leading to galeritus and caffer, is a highly specialized species unique in the bicolor group by the possession of three lateral supplementary leaflets, the third small and undeveloped. The anterior leaflet extends upwards towards the base of the posterior leaf : the second or central leaflet extends anteriorly beneath the anterior leaf but does not reach the median line. The ears are acutely pointed but lack a concavity in their posterior margins. The skull is short and broad, with a very wide rostrum and long pterygoids. The bicolor group contains a high proportion of monotypic species, with only ater, bicolor, galeritus and caffer of widespread distribution and divisible into a number of clearly defined subspecies while fulvus and cineraceus are each divisible into two subspecies but are much less widely distributed. The remaining species are A REVISION OF HIPPOSIDEROS 23 monotypic and while this may in part be due to the inadequacy of collections, there seems little doubt that this feature, with their sharply distinct morphological separa- tion from each other in many cases and their restricted distribution, is an indication that some at least are relict forms that have survived the competition of the more successful species either by considerable specialization or in restricted habitats. Predominantly Indo-Australian in distribution, in Africa the group has rarely given rise to such highly specialized forms as Pygmaeus, coxi and papua, and apart from jonesi forms a more closely related aggregation of species than those of the Indo- Australian region. However, the African representatives of the group have been isolated for a considerable period as is demonstrated by the great development of the noseleaf of jonesi when compared with that of ridleyi and by the structural differences in the posterior noseleaf and skull of coffer and its allies with those of galeritus and its associated species. Lack of diversity might be expected on a large land mass such as Africa : the island habitats of Indo- Australia have evidently encouraged diversification and have enabled species such as dyacorum, sabanus, obscums, Pygmaeus, coxi and papua to survive. Hipposideros bicolor (Temminck), 1834, appears to be the first identifiable name in the group and is therefore adopted as the group name. Lesueur & Petit, in Peron, Voyage de Decouvertes aux Terres Australes, Atlas, 1807, pi. 35 (volume not dated, date taken from Sherborn (1925 : 1661) : discussed by Oey & Feen (1958 : 230)) figured three bats from Timor under the name Rhinolophe Crume*nifere (Rhinolophus crumeniferus N.). The status of this name has been discussed by Tate (1941 : 367 (footnote), 382), Laurie & Hill (1954 : 56) and Oey & Feen (loc. cit.). It is usually ascribed to P6ron, the author of the text of the Voyage de Decouvertes aux Terres Australes : however Rhinolophus crumeniferus is based solely on plate 35 of the Atlas, the title page of which clearly attributes this part of the work to Lesueur & Petit. Tate (1941 : 382) considered the plate to depict bats of the genus Hipposideros " probably related to galeritus and cervinus " while Laurie & Hill (loc. cit.) concluded that the bats represented were most nearly allied to cervinus (here considered to be a race of H. galeritus), an opinion supported by Oey & Feen (loc. cit.). The plate by Lesueur & Petit portrays bats with broad, triangular ears, which have a concavity in their posterior margins and with two lateral supplementary leaflets, thus referable to the bicolor group. The ears, however, are not noticeably covered with body fur. The anterior noseleaf is simple and the internarial septum is not inflated while the posterior leaf is high, supported by three well-defined septa not quite reaching the upper edge of the leaf, which is semicircular in outline. The leaf is largely unpigmented and there is a large frontal sac. It seems likely, therefore, that the three bats depicted by Lesueur & Petit represent H. galeritus cervinus or a closely related form. Tate (1941 : 387) postulates affinity with H. papua but the second leaflet shown in the plate by Lesueur & Petit does not extend forward beneath the anterior part of the horseshoe and these authors portray no incipient third leaflet. Pending discovery of the type or the collection of topotypes, the name must remain incertae sedis. 24 J. E. HILL The species here allocated to the bicolor group (with the exception of coronatus, which from its description is evidently near to calcaratus, and doriae, probably synonymous with sabanus) may be keyed : 1 Lateral supplementary leaflets none or one ....... 2 Lateral supplementary leaflets two or three . . . . . . . 14 2 Anterior upper premolar (pm 2 ) obsolescent or obsolete, if present minute, extruded from toothrow, with second upper premolar (pm 4 ) and canine in contact, anterior lower premolar (pm 2 ) one quarter the length and one third or less the height of the second lower premolar (pm 4 ) . . . . . . . . 3 Anterior upper premolar (pm 2 ) present, never minute, anterior lower premolar (pm 2 ) one third or more the length and height of second lower premolar (pm 4 ) . 4 3 Anterior leaf without a median emargination : posterior leaf supported by a well- defined median septum and weaker lateral septa : pterygoids long, sphenoidal bridge wide ......... dyacorum (p. 43) Anterior leaf with well-defined median emargination : posterior leaf lacking supporting septa : pterygoids short, sphenoidal bridge narrow . sabanus (p. 44) 4 Internarial septum expanded to form a more or less disc-like structure between the nostrils : one lateral supplementary leaflet, sometimes rudimentary ... 5 Internarial septum not greatly expanded or modified, more or less parallel sided : no lateral supplementary leaflets ......... 7 5 Internarial septum expanded to form a concave sub-circular disc ... 6 Internarial septum expanded to form an ellipsoidal structure . marisae (p. 48) 6 Posterior leaf low, rounded above : lateral supplementary leaflet incipient, barely visible : palation U-shaped, without a post-palatal spicule . . ridleyi (p. 39) Posterior leaf high, sub-triangular above : lateral supplementary leaflet well developed, extending anteriorly beneath anterior leaf to median line : palation square, with a small post-palatal spicule ..... jonesi (p. 40) 7 Interorbital region wide, not sharply constricted, its width nearly equal to that of the rostrum ............. 8 Interorbital region decidedly constricted, its width considerably less than that of the rostrum ............ 9 8 Sphenoidal bridge well developed, broad, partially concealing small, elongate lateral apertures : well developed sphenoidal depression . . calcaratus (p. 37) Sphenoidal bridge narrow, not concealing large, rounded lateral apertures : poorly developed sphenoidal depression . . . . . . . cupidus (p. 38) 9 Anterior leaf without a median emargination : posterior leaf with three supporting septa : no glandular ridge on muzzle beneath margin of anterior leaf . . 10 Anterior leaf with small median emargination : posterior leaf without supporting septa : a low glandular ridge on muzzle beneath margin of anterior leaf obscurus (p. 47) 10 Internarial septum thickened and bulbous : anterior half of zygomata slender . n Internarial septum uninflated : anterior half of zygomata massive . . . 12 11 Superior projection of zygomata lacking or poorly developed : anterior upper premolar (pm 2 ) not extruded from toothrow .... cineraceus (p. 35) A low superior zygomatic projection : anterior upper premolar (pm 2 ) extruded or partially extruded from toothrow . . . . . . . atev (p. 30) 12 Posterior projecting portion of vomer blade-like . . . . . . 13 Posterior projecting portion of vomer thickened .... bicolor (p. 25) 13 Anterior lower premolar (pm 2 ) much reduced, one third the length of second lower premolar (pm 4 ) .......... fulvus (p. 33) Anterior lower premolar (pm 2 ) less reduced, one half the length of second lower premolar (pm 4 ) . . . . . , , . . nequam (p. 36) A REVISION OF HIPPOSIDEROS 25 14 Anterior lateral supplementary leaflet extending anteriorly beneath anterior leaf to the median line ........... 15 Anterior lateral supplementary leaflet not extending anteriorly beneath anterior leaf to the median line .......... 16 15 Noseleaf not excessively specialized : intermediate leaf without a median eminence : posterior leaf supported by three septa of equal width, not deeply pocketed pygmaeus (p. 49) Noseleaf greatly specialized : intermediate leaf with prominent median eminence : posterior leaf supported by narrow median septum and two broad lateral septa, deeply pocketed .......... coxi (p. 68) 1 6 Second lateral supplementary leaflet not extending anteriorly beneath the anterior leaf : no trace of a third leaflet ...... . . . 17 Second lateral supplementary leaflet extending anteriorly beneath the anterior leaf : an incipient third leaflet ....... papua (p. 70) 17 Posterior leaf having a transverse supplementary structure with a serrated upper edge developed from its posterior face . . . . . . . . 18 Posterior leaf without a transverse supplementary structure developed from its posterior face or with such a structure low, undeveloped and lacking a serrated upper edge ............ 19 1 8 Anterior upper premolar (pm 2 ) small, slightly extruded from toothrow, or com- pressed between canine and second upper premolar (pm 4 ) . . caffer (p. 62) Anterior upper premolar (pm 2 ) minute, extruded from toothrow, canine and second upper premolar (pm 4 ) in contact or nearly so . . . . . beatus (p. 66) 19 Posterior leaf supported by three septa ........ 20 Posterior leaf without supporting septa, paired low lateral ridges sometimes present fuliginosus (p. 61) 20 Internarial septum not expanded : pterygoids long, sphenoidal bridge wide, partially concealing lateral apertures . . . . . . . .21 Internarial septum expanded to form a slightly disc-like structure : pterygoids short, sphenoidal bridge narrow, not concealing lateral apertures . curtus (p. 60) 21 Ears haired for one half of their length : tips of upper incisors strongly convergent breviceps (p. 58) Ears haired for two thirds of their length : tips of upper incisors not strongly convergent .......... galeritus (p. 52) Hipposideros bicolor The ears are large and rounded, their anterior margins strongly convex and their posterior margins lacking any concavity just behind the tip. There is a distinct antitragal fold. The noseleaf is simple, of moderate size, and lacks lateral supple- mentary leaflets. The internarial septum is more or less triangular, broad at the base, narrowed between the nostrils, very slightly inflated and separated from the lateral parts of the anterior leaf by deep grooves. The lateral parts of the anterior leaf adjacent to the nostrils are not expanded and do not partially conceal the narial openings. The intermediate part of the leaf is unspecialized and the posterior leaf is supported by three septa. The skull is elongate and slender, with a com- paratively broad braincase, narrowed interorbital region and slightly inflated rostral eminences. There is a low sagittal crest. The zygomata are massive, with or without a low superior jugal projection, and the anteorbital foramina are rather elongats, closed by a narrow bar of bone. The premaxillae are narrow and taken 26 J. E. HILL together make a wedge-shaped junction with the maxillae. Their anterior enclosing processes are delicate and do not enclose the rounded anterior palatal foramina. The palation is shallowly V-shaped and the mesopterygoid fossa is wide, with a projecting, thickened vomer. The pterygoids are long and the sphenoidal bridge wide, partially concealing large, elongate lateral apertures. There is a shallow oval sphenoidal depression and the cochleae are of moderate size, their width a little greater than their distance apart. The anterior upper premolar (pm 2 ) is very small and is extruded from the toothrow, while the posterior ridge of the third upper molar is reduced to approximately one half of the length of the anterior ridge. The crown area of the outer lower incisors is equal to that of the inner lower incisors or is very slightly greater, while the anterior lower premolar (pm 2 ) is one half or more the length of the second lower premolar (pm 4 ) and two thirds to three quarters its height. Andersen (1918 : 379) provided the first study of the group of species allied to Hipposideros bicolor, recognizing among others a small species, H. cineraceus, a species of medium size, which he called H. bicolor and two larger species, H. pomona and H. gentilis (here considered to be conspecific) in southeastern Asia. Tate (1941 : 360), however, in the course of revisionary work on the genus, has designated a lectotype for Rhinolophus bicolor Temminck, 1834, and has left its exact application in some doubt. The original description by Temminck, 1834, Tijdschr. Natuur. Gesch. 1 (i) : 19, pi. i, fig. 3 listed no specimens, but Temminck stated that the species was to be found in Java, Amboina and Timor and said that it had been received in considerable numbers at the Netherlands Museum. Later, Temminck, 1835, Monogr. Mamm. 2 : 18 (which Tate apparently thought to be the original description) provided a further, more detailed description, basing it on the exami- nation of ten females and four males, and giving as the provenance of the species the islands of Java, Amboina and Timor. His series is clearly composite, the speci- mens from Amboina, which Temminck said to be smaller than those from Java being referable to the form subsequently described by Peters (iSyia : 323) as Phyllorhina amboinensis. Temminck gives measurements of an adult from Java with forearm " i pouce 8 lignes " (approximately 43 mm.), perhaps a representative of H. bicolor in the sense of Andersen (1918 : 380), a species of medium size widely distributed in the Indo- Australian region. Jentink (1887 : 272, 1888 : 168) has listed among other specimens in the collections of the Rijksmuseum van Natuurlijke Historic, Leiden, a large part of the series probably forming the basis of the description of Rhinolophus bicolor by Temminck. Tate (1941 : 361) has examined these specimens and has designated as lectotype specimen " d " of Jentink (1888 : 168), collected by Van Hasselt (from the collections of Kuhl and Van Hasselt according to Jentink) on the Cote d'Anjer, in the extreme northwest of Java, whence Tate restricts the type locality. This selection introduces a change of name into the group as originally defined by Andersen (1918 : 379) since from the detailed notes on the lectotype provided by Tate, it is evidently a member of the species called Hipposideros gentilis by Andersen (1918 : 380), considered by Ellerman & Morrison-Scott (1951 : 127) and in the present study to be conspecific A REVISION OF HIPPOSIDEROS 27 with H. pomona, also proposed by Andersen in this work. Rhinolophus bicolor Temminck is by far the earliest name in this section of the genus and as a result must replace H. pomona as used by Andersen (1918 : 380) and some subsequent authors. The earliest name identifiable with certainty for the species of medium size called H. bicolor by Andersen (1918 : 380) appears to be Hipposideros ater Templeton, 1848. APPROXIMATE DISTRIBUTION : India east to Hainan and the Philippine Islands ; Malay Peninsula, Sumatra, Java and adjacent islands. Hipposideros bicolor bicolor (Temminck) Rhinolophus bicolor Temminck, 1834 : 19, pi. i, fig. 3 ; 1835 : 18 (further description). Lectotype designated and type locality restricted to Anjer coast, northwestern Java by Tate (1941 : 361). Hipposideros javanicus Sody, 1937 : 2I 5- Babakan, Kroja, Tjilatjap, central Java. No Javan specimens are available for study : from Tate (1941 : 361) the lectotype has an elongate skull with slight nasal eminences and a rather pronounced posterior interparietal swelling. The anterior upper premolar (pm 2 ) is slightly extruded from the toothrow, while the anterior lower premolar (pm 2 ) is three quarters the height of the second lower premolar (pm 4 ). Tate suggested that Hipposideros javanicus Sody, 1937 might be synonymous with H. b. bicolor as represented by the lectotype in Leiden. The measurements of the type specimen of H. javanicus as quoted by Sody agree closely with those of the lectotype of H. b. bicolor as recorded by Tate. DISTRIBUTION : Java ; Banka Island. Hipposideros bicolor pomona Andersen Hipposideros pomona Andersen, 1918 : 380. Haleri, north Coorg, India. The zygomata have a low jugal projection and the anterior upper premolar (pm 2 ) is very small while the anterior lower premolar (pm 2 ) is one half the length and two thirds the height of the second lower premolar (pm 4 ). DISTRIBUTION : Southern India. Hipposideros bicolor gentilis Andersen Hipposideros gentilis Andersen, 1918 : 380. Thayetmyo, Burma. Cranially similar to H. b. pomona but the zygomata lack a definite jugal projection. The anterior lower premolar (pm 2 ) is slightly more than one half the length and two thirds the height of the second lower premolar (pm 4 ). DISTRIBUTION : Northern India ; Assam ; Sikkim ; Burma. 28 J. E. HILL Hipposideros bicolor sinensis Andersen Hipposideros gentilis sinensis Andersen, 1918 : 380. Foochow, Fukien, China. The anterior lower premolar (pm 2 ) is three quarters or more the length of the second lower premolar (pm 4 ), sometimes almost equal to it in length, and is two thirds its height. Cranially, the subspecies otherwise resembles H. b. gentilis. Osgood (1932 : 221) suggested that H. b. sinensis may be a synonym of H. b. gentilis, while Bourret (ig42b : n) considered that H. b. sinensis was not a valid subspecies. DISTRIBUTION : Southern China ; Hainan ; recorded from Hong Kong by Romer (1960 : 2) ; Siam (part) ; Indochina (recorded as H. gentilis by Osgood (1932 : 220) and Bourret (ig42b : n ; 1944 : 6)). Hipposideros bicolor atrox Andersen Hipposideros gentilis atrox Andersen, 1918 : 380. Semangko Gap, Selangor, Federation of Malaya, 2,800 feet. There is a low jugal projection on the zygomata and the sphenoidal pits are very elightly wider than in the foregoing subspecies. The anterior lower premolar (pm 2 ) is one half or less the length and one half the height of the second lower premolar (pm 4 ). Davis (1961 : 90) gives a description, with measurements, of a series of H . b. atrox from the Federation of Malaya. DISTRIBUTION : Malay Peninsula ; Terutau Island ; Tioman Island ; Sumatra. Hipposideros bicolor major Andersen Hipposideros gentilis major Andersen, 1918 : 380. Bua-Bua, Engano Island, off west coast of Sumatra. Cranially exactly like H. b. atrox with the anterior lower premolar (pm 2 ) one half the length and height of the second lower premolar (pm 4 ). Although Andersen in the original description stated that H. b. major was larger than H. b. atrox, the two subspecies seem likely to prove synonymous. Hipposideros bicolor erigens Lawrence Hipposideros erigens Lawrence, 1939 : 56. Lower slopes of Mount Halcon, northern side, near Calapan, Mindoro, Philippine Islands. There are no specimens of this form in the collections of the British Museum (Natural History). Its large size, large ears, noseleaf, bullae and teeth suggest alliance with H. bicolor. Hipposideros bicolor macrobullatus Tate Hipposideros bicolor macrobullatus Tate, 1941 : 357. Talassa, near Maros, south Celebes, 300 metres. There are no specimens of this form in the collections of the British Museum (Natural History). Its measurements and characters as given by Tate agree closely with those of H. bicolor. A REVISION OF HIPPOSIDEROS 36 38 4O 42 44 46 48 H. B. POMONA H. B. GENTILIS H. B. SINENSIS H. B. ATROX H. B. MAJOR H. B. ERIGENS H. B. MACROBULLATUS (FROM TATE (1941:357)) H. B. BICOUOR 36 38 40 42 44 46, 46 FIG. 4. Length of forearm in Hipposideros bicolor 30 J. E. HILL Hipposideros ater The ears and noseleaf are very much like those of H . bicolor but the internarial septum is swollen and inflated, slightly bulbous, narrow at its upper end and separated from the lateral parts of the anterior leaf by deep grooves. The narial margins of the anterior part of the leaf are expanded and partially conceal the nostrils. Cranially, there is considerable similarity to H. bicolor but the zygomata are slender anteriorly and the vomer projects slightly into the mesopterygoid fossa and is slightly thickened. The anterior upper premolar (pm 2 ) is usually extruded from the toothrow and the posterior ridge of the third upper molar is one half or less the length of the anterior ridge. The outer lower incisors are very slightly larger in crown area than the inner lower incisors. The anterior lower premolar (pm 2 ) varies in length from one half or less of the length of the second lower premolar (pm 4 ) in western subspecies to a length almost equal to that of the second lower premolar in eastern subspecies, while its height is one half to two thirds that of the second lower premolar. Formerly known as H . bicolor, this species must now be called H. ater since the designation of a lectotype for Rhinolophus bicolor Temminck by Tate (1941 : 361) transfers that name to the species formerly called H. pomona or H. gentilis. DISTRIBUTION : India to the Philippine Islands, Papua and northern Australia. Hipposideros ater ater Templeton Hipposideros ater Templeton, 1848 : 252. Colombo, Ceylon. Hipposideros atratus Kelaart, i85oa : 208. Substitute for ater. The zygomata have a well developed jugal projection and the posterior ridge of the third upper molar is greatly reduced and obsolescent. The anterior lower premolar (pm 2 ) is less than one half the length of the second lower premolar (pm 4 ) and its height is one half or less than that of the second lower premolar. There appears to be no difference between specimens from Peninsular India (called Hipposideros (?) bicolor fulvus by Ellerman & Morrison-Scott (1951 : 127)) and those from Ceylon. DISTRIBUTION : Ceylon ; India. Hipposideros ater nicobarulae Miller Hipposideros nicobarulae Miller, 1902 : 781. Little Nicobar Island, Bay of Bengal. Similar to H. a. ater but slightly larger, with the anterior lower premolar (pm 2 ) one half the length of the second lower premolar (pm 4 ) and one half or less its height. Hipposideros ater saevus Andersen Hipposideros albanensis saevus Andersen, 1918 : 380. Kei Islands. Hipposideros gentilis toala Shamel, 1940 : 352. Toeare, Celebes. Very similar to H. a. ater but a little larger, with the jugal projection of the zygomata very low or lacking. The anterior lower premolar (pm 2 ) varies from slightly more than one half the length to three quarters the length of the second lower premolar (pm 4 ), and is one half or slightly more its height. A REVISION OF HIPPOSIDEROS 31 Andersen (1918 : 380) employed Rhinolophus bicolor Temminck, 1834, for a medium sized (forearm 37-42 mm.) species of bat of his bicolor group, giving as its provenance Sumatra and Java. In this he was followed by a number of subse- quent authors including Chasen (1940 : 44), Ellerman & Morrison-Scott (1951 : 126) and Laurie & Hill (1954 : 54). However, Tate (1941 : 361) has designated as lectotype of Rhinolophus bicolor Temminck a specimen from that part of the original series still extant in the Rijksmuseum van Natuurlijke Historic, Leiden. It is obvious from the notes and measurements of the lectotype as quoted by Tate that this specimen is not representative of the species of Hipposideros hitherto called bicolor. It is a large bat in comparison with the species formerly called bicolor, with a forearm of 47 mm., zygomatic width 9-1 mm., greatest mastoid width 9-4 mm., least inter- orbital width 3-0 mm. and c-m 3 6-5 mm. Tate also points out that Hipposideros javanicus Sody, 1937, from Java, is very probably synonymous with the species represented by this lectotype. These factors taken in conjunction indicate that the lectotype designated by Tate belongs to the species called gentilis by Andersen (1918 : 380) and subsequent authors, and Rhinolophus bicolor Temminck, 1834, must there- fore be transferred to this species. This leaves the bats from Java and its environs, hitherto called H. bicolor bicolor, without a name, and in the ordinary course of events a new subspecific term would be required. However, there seems little to distinguish these bats from H. ater saevus : specimens from Java, Sumatra and the Malay Peninsula average very slightly smaller than those from more easterly localities and are a little paler while the anterior lower premolar (pm 2 ) is slightly more reduced, but these points scarcely seem to warrant subspecific recognition. Shamel (1940 : 352) described Hipposideros gentilis toala, a Celebesian bat which he considered related to the mainland species gentilis ( = bicolor in the present sense) . However, Tate (1941 : 361, 390) thought toala a probable synonym of H. a. saevus : there seems in fact to be little size difference between toala and saevus (measurements of the type specimen of toala as quoted by Tate (1941 : 361) conflict with those given by Shamel in the original description) and in the present study they are considered to be synonymous. DISTRIBUTION : Mergui Archipelago ; Tenasserim ; Peninsular Siam ; Condor Island ; Federation of Malaya ; Teratau Island ; Tioman Island ; Sumatra; Java; Bali ; Celebes ; Peleng Island ; Kei Island ; Buru ; Ceram ; possibly also Sanghir and Talaud Islands. Tate (1941 : 362) states that a specimen in the American Museum of Natural History from Halmahera is identical with one from Java in the Museum of Comparative Zoology, which from the measurements quoted by Tate is apparently an example of H. ater. Hipposideros ater antricola (Peters) Phyllorhina antricola Peters, 1861 : 709. Paracali, Luzon, Philippine Islands. Hipposideros wrighti Taylor, 1934 : 2 37- Baguio, Benguet (near Headwaters gold mine), Luzon, Philippine Islands. A specimen from Balabac Island (B.M. 94.7.2.51) is referred to this subspecies. It has the anterior lower premolar (pm 2 ) considerably reduced, one half the length 32 J. E. HILL and height of the second lower premolar (pm 4 ). Lawrence (1939 : 55) summarizes the taxonomic history of Hipposideros wrighti Taylor and concludes that it may be in fact a re-description of H . a. antricola. DISTRIBUTION : Philippine Islands : Luzon ; Marinduque ; Mindoro ; Mindanao (Sanborn (1952 : 104)) ; Palawan (Sanborn (1952 : 104)) ; Balabac. 32 34 36 38 4O 42 44 H. A. ATER H. A. NICOBARULAE H. A. SAEVUS H. A. ANTRICOLA H. A. ARUENSIS H. A. GILBERT! 32 34 36 38 4O 42 44 FIG. 5. Length of forearm in Hipposideros ater A REVISION OF HIPPOSIDEROS 33 Hipposideros ater aruensis Gray Hipposideros aruensis Gray, 1858 : 107. Aru Islands. Hipposideros albanensis Gray, i866b : 220. Port Albany, northwest Queensland. The zygomata bear a small jugal projection while the anterior lower premolar (pm 2 ) is rather larger than in H. a. saevus and is equal or almost equal in length to the second lower premolar (pm 4 ) and is two thirds its height. The type specimen is the only available example of Hipposideros albanensis Gray. Its skull is frag- mentary and only the left-hand maxillary toothrow and the mandible remain. The dentition is identical with that of H . a. aruensis. DISTRIBUTION : Aru Islands ; southern and northwestern New Guinea ; northern Queensland. Hipposideros ater amboinensis (Peters) Phyllorhina amboinensis Peters, iSyia : 323. Amboina. There is no example of this subspecies in the collections of the British Museum (Natural History). It is very probably synonymous with H. a. aruensis (see Tate (1941 : 380)). Hipposideros ater gilberti Johnson Hipposideros bicolor gilberti Johnson, 1959 : 183. Oenpelli, East Alligator River, Northern Territory, Australia (12 21' S., 133 04' E.). The collections of the British Museum (Natural History) contain four topotypes (B.M. 23.5.14.9-12) of this subspecies. They agree closely with the description and measurements given by Johnson : there seems little to distinguish them from H. a. aruensis except their slightly paler colour. Hipposideros fulvus The ears are very large and rounded, longer than the head, the upper third of their posterior margins very slightly flattened.* 'The noseleaf closely resembles that of H. bicolor and has the internarial septum narrow, uninflated, broadened at its base and narrowed between the nostrils. The narial margins of the anterior leaf are not expanded, and the nostrils are clearly visible. The skull is elongate and comparatively slender, with a low to moderate sagittal crest and uninflated rostral eminences. The zygomata are broad, with a moderate jugal projection. The premaxillae are like those of H. bicolor but make a more rounded, less wedge- shaped junction with the maxillae. The palation is more or less V-shaped, with long pterygoids, wide mesopterygoid fossa and a very thin, blade-like projecting vomer. The cochleae are of moderate size, their width a little greater than their distance apart. There is an ovate sphenoidal depression. The anterior upper premolar (pm 2 ) is minute, extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact or nearly so. The third upper molar is reduced, zoo 11, i. c 34 J. E. HILL its posterior ridge one quarter or less the length of the anterior ridge. The outer lower incisors are slightly larger in crown area than the inner lower incisors. The anterior lower premolar (pm 2 ) is much reduced and is from one quarter to one third the length and one quarter to one half the height of the second lower premolar (pm 4 ). Hipposideros fulvus may be distinguished from H. bicolor and H. ater by its larger ears : from H. bicolor by its blade-like vomer and much reduced anterior lower premolar and from H. ater by its uninflated internarial septum and broadened zygomata. DISTRIBUTION : India east to Tonkin and Annam ; Lower Siam. Hipposideros fulvus fulvus Gray Hipposideros fulvus Gray, 1838 : 492. Dharwar, India. Hipposideros murinus Gray, 1838 : 492. Dharwar, India. Rhinolophus murinus Elliot, 1839 : 99. Dharwar, India. (Re-description of Hipposideros murinus Gray, 1838, perhaps based on the same original material.) Rhinolophus fulgens Elliot, 1839 : 99. Dharwar, India. (Re-description of Hipposideros fulvus Gray, 1838, perhaps based on the same original material.) Phyllorhina aurita Tomes, i85gb : 76. India. Hipposideros fulvus fulvus occurs in two colour phases : one chestnut brown above and below, the other with the dorsal surface dark brown, the hairs with paler bases and with the ventral surface rather paler than the back. The two colour phases furnish the basis of the names Hipposideros fulvus and Hipposideros murinus proposed by Gray and likewise of Rhinolophus fulgens and Rhinolophus murinus proposed by Elliot. This fact was recognized by Wroughton (igi2a : 829, igi2b : 1179) who first realized that the descriptions by Elliot most probably related to the bats described by Gray. The type locality of both Hipposideros fulvus Gray and Hipposideros murinus Gray is given by their describer as Madras. This appears to be an error for Dharwar, whence Elliot obtained the original specimens. Brosset (1962 : 613) has studied the biology of H. fulvus and gives measurements and notes on its colour variation. Tomes in describing Phyllorhina aurita failed to compare it in detail with any Hipposideros of the bicolor type hitherto described from India. His type specimen agrees closely with H . fulvus. DISTRIBUTION : Ceylon ; Peninsular India (on west coast north to Bombay) ; Bengal, Bhutan Duars ; Sikkim ; Assam ; Burma ; Tenasserim ; Lower Siam ; Tonkin ; Annam. Hipposideros fulvus pallidus Andersen Hipposideros fulvus pallidus Andersen, 1918 : 381. Junagadh, Kathiawar, India. This subspecies differs from H. f. fulvus only in its paler back and underparts, the ventral surface being creamy, faintly tinged with grey and lacking all trace of brown. DISTRIBUTION : Kathiawar ; Gwalior ; Bihar and Orissa ; Sind ; Cutch ; Rajputana ; Baluchistan. A REVISION OF HIPPOSIDEROS 38 4O 42 44 35 H. F. FULVUS H. F.PALLIDUS 38 40 42 44 FIG. 6. Length of forearm in Hipposideros fulvus Hipposideros cineraceus The ears are large and rounded and are similar to those of H. fulvus. The noseleaf is similar to that of H. bicolor, the internarial septum more or less parallel-sided, inflated and bulbous, with the narial margins of the anterior leaf expanded to partially conceal the nostrils. The skull is comparatively small, with an inflated braincase, low sagittal crest, narrow interorbital region and slightly inflated rostral eminences. The zygomata are narrow and delicate and lack a jugal projection. The premaxillae resemble those of H. bicolor and make a wedge-shaped junction with the maxillae. The palation is V-shaped and the mesopterygoid fossa wide, with slightly flared pterygoids. The vomer projects into the mesopterygoid fossa and is thickened posteriorly. The anterior upper premolar (pm 2 ) is small, compressed between the canine and the second upper premolar (pm 4 ) but not markedly extruded from the toothrow. The posterior ridge of the third upper molar is one third to one half the length of the anterior ridge. The outer lower incisors are a little larger in crown area than the inner lower incisors. The anterior lower premolar (pm 2 ) is reduced to a little more than one half the length and height of the second lower premolar (pm 4 ). The small size of H. cineraceus readily distinguishes it from its associated species, except perhaps from H. ater : it may be distinguished from this species by its slender, slightly smaller skull, with delicate zygomata which lack a jugal projection, and its 3 6 J. E. HILL less reduced, unextruded anterior upper premolar. It occurs in two colour phases, the one brownish, the other a brighter, redder phase. DISTRIBUTION : Northern India west to the Punjab ; Assam ; Burma ; Siam ; Tonkin ; Malay Peninsula ; Riau Archipelago ; Anamba Islands ; Borneo. Hipposideros cineraceus cineraceus Blyth Hipposideros cineraceus Blyth, 1853 : 410. Near Find Dadan Khan, Salt Range, Punjab. DISTRIBUTION : as above, except perhaps for the foothills of the Himalayas. Hipposideros cineraceus micropus (Peters) Phyllorhina micropus Peters, 1872 : 256. Dehra Dun, near Simla, northwestern India. The type specimen is the only available example of H. c. micropus. Its skull is smaller in some respects than the skull of H. c. cineraceus and it may represent a northern race. 32 H. C. CINERACEUS H. C. MICROPUS 32 34 36 FIG. 7. Length of forearm in Hipposideros cineraceus Hipposideros nequatn Andersen Hipposideros nequam Andersen, 1918 : 381. Klang, Selangor, Federation of Malaya. The type specimen appears to be the only known example of H. nequam. The original description is very brief and no further diagnostic or comparative notes have appeared, although Tate (1941 : 386) noted its relationship to H. bicolor and pointed out (p. 362) that its greatly reduced anterior lower premolar (pm 2 ) separated it from H. bicolor with which the length of its forearm would otherwise place it. The ears are large and rounded, similar to those of H. fulvus, while the noseleaf is A REVISION OF HIPPOSIDEROS 37 closely similar to that of H. bicolor, but is larger. The internarial septum is un- inflated and the narial margins of the anterior leaf are not expanded. The skull of the type specimen is badly damaged and only part of the rostrum with both upper toothrows and the anterior part of the mandible remain. The rostral eminences are slightly inflated and the anteorbital foramen is large and rather elongate The premaxillae are short and broad, with delicate enclosing processes which do not encircle the anterior palatal foramina. They make a wedge-shaped junction with the maxillae. The palation is shallowly V-shaped and the mesopterygoid fossa is wide, with a projecting, blade-like vomer. The upper canines have a low posterior cusp, and the anterior upper premolar (pm 2 ) is small, not extruded from the toothrow, compressed between the canine and the second upper premolar (pm 4 ), which has a small anterior cusp. The posterior ridge of the third upper molar is reduced to one half the length of the anterior ridge. The outer lower incisors are very slightly larger in crown area than the inner lower incisors, while the anterior lower premolar (pm 2 ) is reduced to one half the length and height of the second lower pre- molar (pm 4 ). Cranially, H. nequam resembles H. bicolor atrox and is approximately the same size, but it differs from H. bicolor in its slightly more inflated rostral eminences, shorter, broader premaxillae, blade-like vomer and more greatly reduced anterior lower premolar (pm 2 ) . Although the large ears, blade-like vomer and greatly reduced anterior lower premolar (pm 2 ) suggest a possible relation to H. fulvus, H. nequam is larger, with a broader rostrum, shorter, much broader premaxillae, which make a wedge-shaped and not rounded junction with the maxillae, wider mesopterygoid fossa and more massive dentition, the canine with a low posterior cusp and the second upper premolar (pm 4 ) with a small anterior cusp. Hipposideros calcaratus (Dobson) Phyllorhina calcarata Dobson, 1877 : 122. Duke of York Island. The ears of H. calcaratus are broad and more or less triangular, their posterior margins with a slight concavity just behind the tip. There is a distinct internal fold. The noseleaf is simple, of medium size, without lateral supplementary leaflets and in general similar to that of H. bicolor, with the internarial septum uninflated and the narial margins of the anterior leaf no 1 |(jSxpanded. The posterior leaf is simple and unwidened, with an ill-defined medfan supporting septum, the lateral septa weak or absent. A frontal sac is present in both sexes. The skull is un- specialized, very little less elongate than that of H. bicolor, with an elongated, uninflated braincase, low sagittal crest, unconstricted interorbital region and slightly inflated rostral eminences. The zygomata are broad, with a well developed jugal projection, the zygomatic width greater than the mastoid width. The pre- maxillae are elongate and narrow, with delicate anterior enclosing processes which do not encircle the elliptical anterior palatal foramina. They make a U-shaped junction with the maxillae and the palation is shallowly V-shaped, with a wide mesopterygoid fossa and blade-like projecting vomer. The pterygoids are long and 38 J. E. HILL wide and the sphenoidal bridge is wide, partially concealing elongate lateral apertures between pterygoids and alisphenoids. There is a small oval or ovate sphenoidal depression and the width of the cochleae is approximately equal to their distance apart. The mandible is massive, with a substantial coronoid process and a heavy, knob-like angular process. The upper incisors are simple with their outer lobes almost obsolete. The upper canines are massive, with a well developed posterior cusp extending one third or more the length of the tooth. The anterior upper premolar (pm 2 ) is small, compressed between the canine and the second upper premolar (pm 4 ), sometimes partially extruded. The third upper molar has its posterior ridge one third or less the length of the anterior ridge. The outer lower incisors are equal in crown area to the inner pair. The anterior lower premolar (pm 2 ) is only slightly reduced, two thirds or more the length and three quarters the height of the second lower premolar (pm 4 ). Tate (1941 : 358, 362) allocated H. calcaratus (Dobson) and H. cupidus Andersen to a calcaratus group distinct from his bicolor and galeritus groups. There seems to be no justification for this comparatively wide separation : H. calcaratus and H. cupidus are not greatly removed from H . bicolor and its allies and closely resemble them in the basic features of the ears, noseleaf and skull. They differ from H, bicolor and its associated species chiefly in their more triangular, less rounded ears, more simplified noseleaves and in having the interorbital region of the skull unconstricted. DISTRIBUTION : New Guinea ; Bismarck Archipelago : Duke of York Island ; Solomon Islands : Russell ; New Georgia ; Nissan ; Rennell. Hipposideros cupidus Andersen Hipposideros cupidus Andersen, 1918 : 383. Eaga, Papua. The original diagnosis is very brief : Tate (1941 : 362, 381, 382) and Hill (1956 : 77, 78) give supplementary notes. The ears and noseleaf are as in H. calcaratus. In its cranial characters H. cupidus is closely similar to H. calcaratus, but the zygomata have only a low jugal projection and the pterygoids and sphenoidal bridge are narrow, the sphenoidal bridge not partially concealing wide, rounded lateral aper- tures. The sphenoidal depression is very poorly developed. The dentition is closely similar to that of H. calcaratus but the canines are slender while retaining the high posterior cusp. Hipposideros cupidus is in general very similar to H. calcaratus and the two species are sympatric for part of their range. Tate (1941 : 362) studied series of both and formulated a key for their separation. Hill (1956 : 77, 78) on the basis of a re-examination of the type specimens commented on the diagnostic characters used by Tate and pointed out that criteria of size do not appear to be valid in the Solomon Islands and that the height of the posterior canine cusp is not a reliable diagnostic character. DISTRIBUTION : New Guinea ; Japen Island ; Bismarck Archipelago : Duke of York Island ; Tabar Islands ; Solomon Islands : New Georgia ; Banika. A REVISION OF HIPPOSIDEROS 39 Hipposideros coronatus (Peters) : 327. Mainit, Surigao, northeastern Mindanao, Philippine Phyllorhina coronata Peters, Islands. There is no example of this species in the collections of the British Museum (Natural History). Its size, lack of lateral supplementary leaflets and unspecialized posterior leaf suggests affinity with H. calcaratus. 44 46 48 50 52 54 44 46 46 50 52 54 H. NEQUAM H. CALCARATUS H. COPIOUS H. CORONATUS (FROM PETERS (1 871 o: 328)) FIG. 8. Length of forearm in Hipposideros nequam, H. calcaratus, H. cupidus and H. coronatus Hipposideros ridleyi Robinson & Kloss Hipposideros ridleyi Robinson & Kloss, 1911 : 241. Botanic Gardens, Singapore Island. Gibson-Hill (1949 : 191) believed the type specimen of H. ridleyi to be lost. However, in the course of the present study, it was found among a collection of bats sent many years ago to Andersen from the Federated Malay States Museum for study at the British Museum (Natural History). No further specimens appear to have been recorded. The ears are very large and broad, sub-triangular and bluntly pointed, their anterior margins convex, their posterior margins straight for the upper third and not concave. There is a well-developed fold at the antitragal lobe and the ears are haired for their basal quarter. The noseleaf is large, almost completely covering the muzzle and lacks lateral supplementary leaflets. A slight longitudinal swelling behind and parallel to the outer margin of the anterior leaf however, may represent an incipient leaflet. The anterior leaf is broad, with its narial margins slightly expanded and partially concealing the nostrils. The inter- narial septum is greatly expanded to form a concave circular disc between and anterior to the nostrils which however does not obscure the narial openings. The 40 J. E. HILL narial flaps or lappets are considerably developed to form a pocket encircling the nostrils, but do not rise above the level of the horseshoe. The intermediate part of the leaf is cushion-like, with a low median eminence. The posterior leaf is high, its upper edge semicircular, its lower half supported by three prominent septa of equal depth enclosing four deep pockets, its upper half smooth. The rear walls of the central pockets form a low projecting structure on the posterior face of the leaf. A frontal sac is present in the male type specimen. The skull is comparatively large and elongate, with broad braincase, low sagittal crest, slight supraorbital ridges, narrow interorbital region, a very shallow frontal depression and slightly inflated rostral eminences. The zygomata are slender with a well developed jugal projection. The anteorbital foramen is elongate, closed by a moderate bar. The premaxillae are short and broad anteriorly, with short anterior enclosing processes. They are narrowed posteriorly to make a wedge- shaped junction with the maxillae and do not enclose the anterior palatal foramina. The palation is U-shaped, with a wide mesopterygoid fossa and a thin, blade-like projecting vomer. The pterygoids are long and wide with a wide sphenoidal bridge, partially concealing elongate lateral apertures. There is an ovate sphenoidal depression and the width of the cochleae is a little greater than their distance apart. The upper incisors are weakly bilobed, their tips strongly convergent and almost touching. The upper canines are slender, with well developed cingula. The anterior upper premolar (pm 2 ) is small, compressed between the canine and the second upper premolar (pm 4 ) and the posterior ridge of the third upper molar is one half the length of the anterior ridge. The outer lower incisors are very slightly greater in crown area than the inner lower incisors and the anterior lower premolar (pm 2 ) is nearly as long as the second lower premolar (pm 4 ) but only one half its height. It is evident that H. ridleyi is closely related to H. bicolor and its immediately associated species, resembling them in having ears with a well developed internal fold, its lack of lateral supplementary leaflets and its narrow, elongate skull with broad brain case and zygomatic width less than the mastoid width. The curious specialization of the internarial septum appears to be a further development of the condition in H. ater and H. cineraceus, in which the internarial septum, although broadened and bulbous, has not developed an internarial disc or pad, and which tends towards H. jonesi, an African species exhibiting a yet more greatly developed internarial disc. This contention is supported by the parallel appearance in H. ridleyi of narial pockets and a deeply pocketed posterior leaf, structures found more greatly developed in H. jonesi. Hipposideros jonesi Hayman [Figure 9] Hipposideros jonesi Hayman, 1947 : 71. Makeni, Sierra Leone, West Africa. The ears are very large, broad, sub-triangular, with an acute point, their anterior margins slightly convex, their posterior margins very slightly so, with a faint concavity just behind the tip. There is a distinct internal fold at the antitragal A REVISION OF HIPPOSIDEROS 41 lobe and the ears are haired for their basal third. The noseleaf is a greatly specialized structure with one well-developed lateral supplementary leaflet extending from the base of the intermediate part of the leaf anteriorly beneath the anterior leaf to the median line, with a deep emargination above the centre of the lip. The anterior leaf is broad, covering almost the entire width of the muzzle, with a faint anterior emargination. The internarial septum is greatly expanded into a large concave, more or less circular disc between and just anterior to the nostrils, which it partially conceals. The narial flaps or lappets form pockets enclosing the nostrils and rise above the level of the anterior leaf. The intermediate part of the leaf is elevated but is otherwise unspecialized. The posterior leaf is high, its outline sub-triangular, with a blunt, rounded point, its lower half supported by a shallow median septum and two deeper lateral septa, forming three deep pockets, the central pocket larger than the lateral pockets and divided by the lower median septum. The posterior walls of the pockets do not form a projecting structure on the posterior face of the leaf and the upper half of the leaf is smooth. The frontal sac is very poorly de- veloped in male specimens and lacking in female examples. FIG. 9. Hipposideros jonesi $ (Type B.M. 47.629) (x3) The skull is of medium size and is elongate with a broad braincase, low sagittal crest, barely definable supraorbital ridges, narrowed interorbital region and rostrum, with a shallow frontal depression and well-inflated rostral eminences. The rostrum is elongate, the premaxillae projecting beyond the canines. The zygomata are 42 J. E. HILL slender, with a low jugal process, and the zygomatic width is considerably less than the mastoid width. The anteorbital foramen is rounded and closed by a narrow bar. The premaxillae are broad anteriorly with delicate anterior enclosing processes : they are narrowed posteriorly and do not enclose the large rounded anterior palatal foramina, making a V-shaped junction with the maxillae. The palation is square, with a post-palatal spicule. The mesopterygoid fossa is wide, with a blade-like, projecting vomer. The pterygoids are long and flaring, partially concealing large lateral apertures, although the sphenoidal bridge is narrow. There is a shallow sphenoidal depression. The cochleae are large, their width equal to two times or a little more their distance apart, with a rather narrow basioccipital. The upper incisors lack the outer lobe, and although convergent their tips are widely separated. The upper canines have weak anterior and posterior cusps. The anterior upper premolar (pm 2 ) is of moderate size, slightly extruded and compressed between the canine and the second upper premolar (pm 4 ). The posterior ridge of the third upper molar is one half or slightly less the length of the anterior ridge. The crown area of the outer lower incisors is slightly greater than that of the inner lower incisors and the anterior lower premolar (pm 2 ) is equal approximately to one half the length and height of the second lower premolar (pm 4 ). Hayman (1947 : 73) pointed out that H. jonesi stood apart from any of the groups defined and keyed by Tate (1941), noting that while in size, the form of the ears and in the mastoid width exceeding the zygomatic width it approached the bicolor group of species as understood by Tate, in other features such as the well-developed single supplementary leaflet and the absence of a frontal sac it differed widely from this and the other groups of the genus. Later, Aellen (1954 : 480) keyed and listed H. jonesi as the sole species of a jonesi group. However, there seems little doubt that H. jonesi should be allocated to the group of species typified by H. bicolor and its allies. As in these species, the ears have a distinct internal fold and are haired for one third of their length. The skull has the elongate outline typical of H. bicolor and its associated species, with a similarly broad braincase. The nearest ally to H. jonesi appears to be H. ridleyi, a Malaysian species exhibiting similar specialization of the noseleaf, although as might be expected from the wide geogra- phical separation of the two species, there are very considerable differences between them. Both have large, sub-triangular ears with a well-defined internal fold : their noseleaves are wide, covering the muzzle almost entirely : in each the internarial septum is expanded to form a disc-like structure and circumnarial pockets have been developed while both have a high posterior leaf, its lower half deeply pocketed and its upper half smooth. Hipposideros jonesi differs markedly from H. ridleyi in the possession of a well-developed lateral supplementary leaflet which in H. ridleyi is merely incipient : the internarial disc in H. jonesi is larger and the posterior leaf more developed, with its upper edge triangular in outline and not rounded as in H. ridleyi. Cranially, the two species are closely similar and have skulls essentially resembling those of H. bicolor and its immediate allies : H. jonesi has a smaller, more slender skull than H. ridleyi with a square and not U-shaped palation and larger cochleae, with narrowed basioccipital and sphenoidal bridge. Their dentition A REVISION OF HIPPOSIDEROS 43 differs principally in that the upper incisors of H.jonesi are placed at the outer margins of the premaxillae and although convergent have their tips separated : those of H. ridleyi are more medially sited and have their tips strongly convergent and almost touching. Hipposideros jonesi is the most specialized species of the more primitive bicolor section of the bicolor group, retaining the more or less rounded ears, elongate rostrum and narrowed zygomata of the bicolor type and yet with a greatly specialized noseleaf associated with some specialization of the auditory region of the skull. Together with the similar but less specialized species H. ridleyi it constitutes an offshoot of the bicolor type leading apparently to no further specialization. Its differences from the Malaysian H. ridleyi are of a considerable order of magnitude and indicate remote separation of the parental stems. DISTRIBUTION : West Africa : Sierra Leone ; Guinea (for notes on specimens from Guinea see Eisentraut & Knorr (1957 : 333)). 44 46 48 50 H. JONESI H. RIDLEYI 44 46 48 SO FIG. 10. Length of forearm in Hipposideros jonesi and H. ridleyi Hipposideros dyacorum Thomas Hipposideros dyacorum Thomas, 1902 : 271. Mount Mulu, Baram, Sarawak, Borneo. The ears are of moderate size, broad at the base, sub-triangular, their anterior margins convex, their posterior margins with a slight concavity just behind the tip but otherwise convex. There is a well-defined internal fold at the antitragal lobe and the ears are sparsely haired for about one half of their length. The noseleaf is small, narrow and simple, and lacks lateral supplementary leaflets. It is very like the noseleaf of H. bicolor, with the internarial septum slightly inflated, triangular, broad at its base, narrowed between the nostrils and with the narial margins of the anterior leaf slightly expanded but not concealing the nostrils. The intermediate part of the leaf is cushion-like, with a low, slightly inflated median eminence. The posterior leaf is thin, its upper edge semicircular, and is supported by a well-defined median septum and two less prominent lateral septa. There is a small frontal sac in the female type specimen. 44 J- E. HILL The skull is short and the braincase strongly inflated with a low sagittal crest. The interorbital region is constricted and the supraorbital ridges, although low, are sharply defined and prominent. There is a shallow frontal depression and the rostrum is slightly broadened, with moderately inflated rostral eminences. The zygomata are slender with a moderate jugal projection, and the anteorbital foramen is elongate, closed by a very slender bar. The premaxillae are entirely fused, their junction with the maxillae shallowly V-shaped, the maxillae with an abrupt emar- gination at the apex of the V. The enclosing processes of the premaxillae are delicate and hook-shaped, but do not enclose the rounded anterior palatal foramina. The palation is U-shaped, with an abrupt median emargination. The vomer does not project into the mesopterygoid fossa, which is not greatly widened. The pterygoids are long, the pterygoid wings flaring, together with the slightly constricted sphenoidal bridge partially concealing elongate lateral apertures. There is a shallow sphenoidal depression. The width of the cochleae is equal to their distance apart. The upper incisors are very weakly bilobed, and the upper canines have weak anterior and posterior cusps. The anterior upper premolar (pm 2 ) is minute, extruded, with the canine and the second upper premolar (pm 4 ) in contact. The posterior ridge of the third upper molar is obsolete. The crown area of the outer lower incisors is very slightly greater than that of the inner lower incisors. The anterior lower premolar (pm 2 ) is much reduced, to one quarter the length and height of the second lower premolar (pm 4 ). Measurements of specimens of H. dyacorum from southwestern Borneo are given by Lyon (1911 : 129, 130) while a description, with measurements, of a series from North Borneo is provided by Davis (1962 : 39). Hipposideros dyacorum, forms a link between H. bicolor and H. galeritus, and although independently slightly specialized in some respects displays some characters indicative of the aggregations of species to which they belong. The ears and nose- leaf of H. dyacorum correspond closely to those of H. bicolor and its allies, while its skull is shortened like that of H . galeritus and its associated species, and has the long pterygoids common to both aggregations of species. The dentition of H. dyacorum is more advanced than that of H . bicolor and its allies and in the reduc- tion of the premolars tends towards H. galeritus. DISTRIBUTION : Borneo. Hipposideros sab anus Thomas Hipposideros sabanus Thomas, 1898 : 243. La was, North Borneo. The ears are large and broad, rounded and not acutely pointed, their posterior margins evenly convex. They are thickened at the antitragal lobe but lack a definite internal fold, and are haired for the basal third of their length. The noseleaf is small and comparatively simple, lacking lateral supplementary leaflets. The anterior leaf is narrow, with a well-defined median emargination. The internarial septum is swollen and bulbous, especially posteriorly, but does not conceal the narial apertures, which are flanked by small narial lappets. The intermediate part A REVISION OF HIPPOSIDEROS 45 of the leaf is cushion-like but not greatly inflated. The posterior leaf is high, without supporting septa and has a semicircular upper edge. It is specialized by the development from its posterior face of a transverse supplementary structure, with a slightly serrate upper edge below that of the posterior leaf. There is a small frontal sac in the female type specimen. The skull is small, with a broad braincase and low sagittal crest. The interorbital region is moderately constricted while the supraorbital ridges are not sharply defined and are very weak. There is no frontal depression, the rostrum in profile exhibiting a slight convexity. The rostrum is short and not broadened, the rostral eminences only slightly inflated. The zygomata are slender with a high jugal process, their combined width greater than the mastoid width. The anteorbital foramen is elongate, closed by a very slender bar. The premaxillae are short and broad, their junction with the maxillae V-shaped, while the anterior palatal foramina are rounded and enclosed within the premaxillae by a narrow bar. The palation is only slightly rounded, almost square, without a median emargination. The mesopterygoid fossa is wide, with a slightly projecting vomer, while the pterygoids are short and the sphenoidal bridge narrow, exposing large rounded lateral apertures. There is a shallow sphenoidal depression and the cochleae in width slightly exceed their distance apart. The upper incisors are not bilobed, the outer lobe obsolete, while the upper canines have a weak anterior cusp and a trace of the posterior cusp. The anterior upper premolar (pm 2 ) is obsolete, the second upper premolar (pm 4 ) with a low anterior cusp in contact with the canine. The posterior ridge of the third upper molar is reduced to one third the length of the anterior ridge. The crown area of the outer lower incisors is very slightly greater than that of the inner lower incisors, while the anterior lower premolar (pm 2 ) is much reduced, one quarter the length and height of the second lower premolar (pm 4 ) and is slightly extruded from the toothrow. The collections of the British Museum (Natural History) contain a specimen from Sumatra (B.M. 27.5.9.3) which hitherto has been referred to H. doriae (Peters). However, its ears agree exactly with those of H . sabanus, and although the noseleaf has been partially destroyed, sufficient remains intact to show that the posterior leaf lacks supporting septa and has the low transverse serrated supplementary structure typical of H. sabanus developed from its posterior face. Cranially, this specimen agrees very closely with H. sabanus but has the sphenoidal depression a little more sharply defined and the upper incisors weakly bilobed, the outer lobe obsolescent. In view of these considerations it is referred to H. sabanus, which Chasen (1940 : 481) had already recorded from Sumatra. I am unable to agree with Tate (1941 : 366, 383, 388) that H. dyacorum and H. sabanus are closely related or even allied. At most they appear to share a common origin among the more primitive bats of the bicolor subgroup to which they both display affinity, but otherwise they appear to represent two differing and widely divergent lines of development. Although H. dyacorum has more or less triangular ears with a slight posterior concavity, its ears nevertheless have a well-defined internal fold and its noseleaf is simple, lacking lateral supplementary leaflets. 46 J. E. HILL Hipposideros sabanus has the broad, rounded ears characteristic of the bicolor subgroup, with convex posterior edges but without an evident internal fold. Its noseleaf, although rather small, without lateral supplementary leaflets and basically of the bicolor type, is more specialized, with the posterior leaf lacking supporting septa and with a transverse supplementary structure on its posterior face. The two species differ sharply in their cranial characters. Hipposideros dyacorum has a longer skull than H. sabanus, and has a wider rostrum with the rostral eminences more inflated. The palate in H. dyacorum is very much longer and the palation U-shaped and not square : its pterygoids are long and the sphenoidal bridge is wide, in contrast to H. sabanus, which has short pterygoids and a narrow sphenoidal bridge. The anterior lacerated foramina of H. dyacorum are elongate and narrow, while those of H. sabanus are rounded and wide. These differences in cranial architecture indicate considerable separation, and the two species do not appear to be closely related. Hipposideros dyacorum appears to be more closely related to H. bicolor and its immediate allies than does H. sabanus. Its ears retain the internal fold : its noseleaf is largely unspecialized and the skull is slightly elongate, the palate not extensively shortened. Its long pterygoids and wide sphenoidal bridge suggest relationship with H . galeritus and its associated species and it is clearly derived from the line connecting them to the less specialized species associated with H. bicolor. Hippo- sideros sabanus, although with ears of the bicolor type, has a more advanced noseleaf and a shorter, less elongate skull. Its much shorter palate indicates closer relation- ship to the galeritus subgroup than is evident in H. dyacorum : the features of its posterior leaf, which lacks supporting septa and which has a low posterior transverse structure, together with the short pterygoids and narrow sphenoidal bridge suggest that it has been derived from the stem leading to H. caffer and its allies. DISTRIBUTION : Borneo ; Sumatra. Hipposideros doriae (Peters) Phyllorhina doriae Peters, iSyia : 326. Sarawak, Borneo. The collections of the British Museum (Natural History) contain no specimen referable to H. doriae and it has not been possible to examine the type specimen, described by Peters from the collections of the Marquis J. Doria and apparently deposited in the collections of the Museo Civico di Storia Naturale at Genoa. Although Peters described only the external features of H . doriae there seems little doubt from his description that it belongs to that section of the genus typified by H. bicolor and is very similar, if not identical to H. sabanus. The ears lack a distinct tip and have their anterior and posterior margins equally convex for their terminal third. The noseleaf lacks supplementary lateral leaflets and the posterior leaf has a smooth anterior face, without supporting septa. There is a small but distinct frontal sac. Further notes on the type specimen were obtained by Oldfield Thomas from R. Gestro of the Museo Civico di Storia Naturale and are preserved in the archives of the British Museum (Natural History) for 1902. They have also been 47 recorded by Thomas as a marginal note to the account of H. doriae (p. 146) in a copy of Dobson, 1878, Catalogue of the Chiroptera in the collection of the British Museum, now in the library of the British Museum (Natural History). The features of the type specimen noted by Gestro are that the anterior leaf has no median emargination : that the anterior upper premolar (pm 2 ) is absent and that the anterior lower premolar (pm 2 ) is small, a diagram drawn by Gestro suggesting that it is less than one half of the size of the second lower premolar (pm 4 ). Thomas adds in his marginal note that H. sabanus is probably equal to H. doriae : the close agreement of noseleaf and dentition suggests that H. doriae is related to H. sabanus rather than to H. bicolor as was suggested by Dobson (1878 : 147) and Tate (1941 : 383). The two species are clearly very similar and seem likely to be at least conspecific, in which case it should be noted that doriae is the prior name by many years. Hipposideros obscurus (Peters) Phyllorhina obscura Peters, 1861 : 709. Paracali, Luzon, Philippine Islands. The following notes are based on a small series (B.M. 77.10.6.14-18) in the collec- tions of the British Museum (Natural History), from Dinagat Island, Philippine Islands, consisting of an adult male and female, together with three young males. The ears are sharply triangular, broad at the base, their anterior margins markedly convex and their posterior margins with a concavity just behind the acute point. There is no internal fold but the ear membrane is thickened at the antitragal lobe and there is a small antitragal projection. The ears are haired for one half of their length. The noseleaf is of moderate size and does not entirely cover the muzzle : it lacks lateral supplementary leaflets but has a distinct raised glandular ridge directly beneath the edge of the anterior leaf, extending laterally under the margins of the anterior leaf and anteriorly beneath this leaf to the median line. The anterior leaf in the adult male has a small but distinct median emargination which in the adult female is very small : it is present but very small in two of the young male specimens and absent in the third. The internarial septum is not inflated and is broadly triangular, narrow between the nostrils, which lie at the base of a deep depression : the narial lappets are moderately developed and the narial margins of the anterior leaf are slightly expanded. The intermediate part of the leaf is cushion-like, slightly inflated, with a low median ridge or eminence. The posterior leaf is high, its upper edge semicircular and lacks supporting septa, with a low transverse ridge on its posterior face. There is a well-developed frontal sac in the male, represented by a depression in the female specimen. The skull is short and comparatively broad, with a broad, inflated braincase, low sagittal crest and a moderately constricted interorbital region. There is no frontal depression and the rostrum is broad and rounded, with slightly inflated rostral eminences. The zygomata are slender, their combined width a little greater than the mastoid width. The anteorbital foramen is large and elongate, closed by a narrow bar. The junction of the premaxillae with the maxillae is shallowly V-shaped and the palate is short : the palation is U-shaped with a small post-palatal spicule. 48 J. E. HILL The mesopterygoid fossa is wide, the vomer not projecting and the pterygoids long, together with the moderate sphenoidal bridge partially concealing wide lateral apertures. There is a small sphenoidal depression and the width of the cochleae is equal to their distance apart. The anterior upper premolar (pm 2 ) is small and is extruded into the angle between the canine and the second upper premolar (pm 4 ), which however are not in contact, while the posterior ridge of the third upper molar is obsolescent. The crown area of the outer lower incisors is slightly greater than that of the inner pair. The anterior lower premolar (pm 2 ) is three quarters the length and height of the second lower premolar (pm 4 ). Hipposideros obscurus is the last species of the bicolor type and like H. dyacorum, apparently links the more primitive of these to galeritus and its allies, although it has no apparent close connection with H . dyacorum and in some respects inclines towards H. sabanus. Its ears are very like those of H. dyacorum but lack the internal fold while its noseleaf, although of bicolor type, has deeply depressed nostrils suggestive of H. pygmaeus or H. coxi, and a posterior leaf lacking septa and with a posterior transverse structure reminiscent of H. sabanus. The short, broad skull has the long pterygoids characteristic of H. galeritus and its associates as distinct from H. caffer and its allies but the dentition, retaining the anterior upper premolar, is less advanced than in H. dyacorum and H. sabanus. DISTRIBUTION : Philippine Islands : Luzon ; Dinagat ; Mindanao (Sanborn (1952 : 104)). Hipposideros marisae Aellen Hipposideros marisae Aellen, 1954 : 474- n g- I - White Panther Rock, Duekoue, Ivory Coast. No example of H. marisae has been examined, and the following notes have been compiled from the original description. The ears are large, broad and bluntly pointed, with a concavity behind the tip. There is no antitragal fold. The nose- leaf is small with one very small lateral supplementary leaflet. The anterior leaf has apparently no median emargination and the internarial septum is inflated, slightly disc-like between the nostrils. The intermediate part of the leaf is slightly inflated while the posterior leaf is high, its upper edge semicircular, and is supported by three septa enclosing deep pockets. A frontal sac is present in the male type specimen. The skull is said to be in general similar to that of H. bicolor and H. curtus, with a weak sagittal crest, narrow rostrum, comparatively wide zygomata, their combined width greater than the mastoid width, and small bullae. The dentition is apparently similar to that of H . curtus : the upper incisors lack the outer lobe and the anterior upper premolar (pm 2 ) is small and slightly extruded while the anterior lower pre- molar (pm 2 ) is one half the height of the second lower premolar (pm 4 ). Aellen (1954 : 474 et seq.) has considered the taxonomic status of H. marisae and its relative H. curtus in some detail, and concluded that neither are closely related to H. caffer and its immediate allies but to the bicolor group (the bicolor subgroup as here understood). Aellen further proposed that H. marisae and H. curtus should be considered to represent a group within Hipposideros, the curtus A REVISION OF HIPPOSIDEROS 49 group, characterized by large emarginated ears lacking an internal fold, one or two lateral supplementary leaflets, swollen claviform internarial septum, the presence of a frontal sac in males, sometimes absent in females, the fourth metacarpal longer than the third and fifth and by a small, slightly extruded anterior upper premolar (pm 2 ). However, there seems no justification for this action. Hipposideros curtus, with ears and noseleaf similar to those of the bicolor subgroup yet with two lateral supplementary leaflets and skull tending towards the galeritus subgroup, particularly towards H. caffer, is clearly derived from the Ucolor-caffer stem. Similarly, another such derivative, H. fuliginosus, which Aellen places in a caffer group, has ears and noseleaf closely similar to those of H. caffer, yet has a skull exhibiting a number of the features of the bicolor subgroup. From the original description, H. marisae presents yet a further combination of characters, having slightly specialized ears lacking an internal fold but otherwise like those of the bicolor subgroup and in the possession of a simple noseleaf similar to that of H. bicolor, with one small, rudimentary lateral supplementary leaflet. Its skull appears to be similar to the skulls of the members of the bicolor subgroup, rather elongate and narrow, with the zygomatic width only barely exceeding the mastoid width. It too represents the bicolor-caffer stem, and is a less advanced species than either H. curtus or H. fuliginosus. Hipposideros marisae, H. curtus and H. fuliginosus form connecting links between H. bicolor and its associated species and the more specialized H. caffer. As is not unusual in the genus, specialization of the various species has reached different levels : H . marisae and H. curtus have ears and noseleaves rather of the pattern of H. bicolor, modified by the presence of one lateral supplementary leaflet in H. marisae and the presence of two such leaflets in H . curtus, while H. marisae has a skull of the bicolor type and H. curtus a skull tending towards H. caffer. Hippo- sideros fuliginosus has coffer-like ears and noseleaf associated with a skull similar to that of the members of the bicolor subgroup. The characters of its ears and nose- leaf ally it more closely to H. caffer than to H. marisae and H . curtus and it is less closely related to these species than they are to each other. Both are characterized by a swollen, claviform internarial septum which tends to form a disc-like structure between the nostrils, a specialization found in far more greatly developed form in H. ridleyi and H. jonesi, two species forming a development of H. bicolor and its allies. For this reason, H. marisae and H. curtus are considered to form an indepen- dent offshoot of the bicolor-caffer stem. There seems little point in constructing groups for intermediate species, based largely on the characters that they share in common with the species they link together, and the curtus group of Aellen (1954 : 479) in the present work is considered to form a part of the bicolor group as here understood. Hipposideros pygmaeus (Waterhouse) Rhinolophus pygmaeus Waterhouse, 1843 : 67. Philippine Islands. The type specimen is the only available example of H. Pygmaeus. It is a very small bat with the ears sub-triangular, very broad at the base, pointed, with the zoo 11, i. D 5 o J. E. HILL upper portion of their anterior margins very slightly concave and their posterior margins concave behind the tip. The ears are slightly thickened at the antitragal lobe and are haired for one half of their length. The noseleaf is large, entirely covering the muzzle, and has two lateral supplementary leaflets, the anterior leaflet extending from the base of the intermediate part of the leaf forward beneath the anterior leaf to the median line but with a median emargination, the posterior leaflet well developed and extending slightly anteriorly. The internarial septum is very small and reduced, insignificant between the nostrils : the narial lappets are greatly developed, especially laterally, and rise above the level of the horseshoe. They are elongated and almost completely encircle the nostrils, which lie at the base of the small pockets so formed. The intermediate part of the leaf is well inflated but lacks eminences. The posterior leaf is high, thin, its upper edge semicircular, and is supported by three well-defined septa which enclose four small but moderately deep pockets. There is no frontal sac in the female type specimen : according to Tate (1941 : 369) it is present in males. A small anterior portion of the skull is all that remains. The sagittal crest, represented by a remnant, is evidently low and the interorbital region is constricted, but not sharply so, with no evident supraorbital ridges. There is no frontal depression. The rostrum is slightly shortened, high and rounded, the rostral eminences well inflated with a slight median depression between them. The anteorbital foramen is large and rounded, closed by a very narrow bar. The upper incisors are weakly bilobed, the outer lobe obsolescent. The upper canines have a moderate anterior cusp and a stronger posterior cusp extending for about one third of the length of the tooth. The anterior upper premolar (pm 2 ) is small but is barely out of alignment in the toothrow, separating the canine and the second upper premolar (pm 4 ), which has a distinct anterior cusp. The posterior ridge of the third upper molar is one half of the length of the anterior ridge. Tate (1941 : 367, 369, 388) considered H. pygmaeus to be an isolated species derived from the line leading to H. galeritus and its allies. This view appears to be correct, and despite its extended anterior lateral supplementary leaflets and inflated rostral eminences which suggest relationship to H. coxi, its affinities lie more closely with H. galeritus and its associated species. Although the anterior lateral supplemen- tary leaflets in H . pygmaeus extend anteriorly beneath the anterior leaf to the median line as in H . coxi, the two species differ sharply in that the leaflets in H. pygmaeus are not continuous over the upper lip as in H. coxi, but are divided to the base by a sharp median emargination. This condition suggests H. galeritus, in which the leaflets, although extending forward beneath the anterior leaf, do not reach the median line. The narial part of the noseleaf in H. pygmaeus is slightly more specialized than that of H. obscurus, also a derivative of the galeritus stem. The posterior leaf is divided by blade-like septa as in H. galeritus, and its margins at the base display none of the extraordinary complexity of H. coxi. The skull is not excessively shortened and the dentition not greatly specialized, providing a link with the bicolor type of skull. Vertical inflation of the rostral eminences appears to be a correlation of increasing complexity of the narial region of the noseleaf. Hippo- A REVISION OF HIPPOSIDEROS 51 sideros obscurus, in which the narial apertures are situated at the base of a deep depression but are not noticeably pocketed, has slightly inflated rostral eminences. In H. pygmaeus the narial region is slightly more specialized : the narial lappets are elongated to form small pockets almost completely encircling the nostrils and the rostral eminences are well inflated. Hipposideros coxi has the narial apertures completely concealed within a pocket formed by the narial lappets and the inter- narial septum : correspondingly the rostral eminences are much inflated and appear as two separate raised structures. On the balance of characters, therefore, H. Pygmaeus is associated with H. galeritus and its allies. DISTRIBUTION : Philippine Islands : Luzon ; Negros (Sanborn (1952 : 104)). 34 36 38 40 42 44 46 48 50 52 - 34 36 38 40 42 44 46 48 50 52 FIG. ii. Length of forearm in Hipposideros dyacorum, H. sabanus, H. obscurus, H. marisae, H. breviceps, H. curtus, H. pygmaeus, H. coxi and H. papua H. DYACORUM H. SABANUS H. OBSCURUS H. MARISAE H. BREVICEPS H. CURTUS H. PYGMAEUS H. COXI H. PAPUA 52 J. E. HILL Hipposideros galeritus The ears are broad, triangular and acutely pointed, their anterior margins convex, their posterior margins concave behind the tip but otherwise convex, without a marked antitragal fold but thickened at the antitragal lobe, with a small antitragal projection. They are haired for three quarters of their length. The noseleaf is small and comparatively simple, with two well-developed lateral supplementary leaflets, which project beyond the lateral margins of the anterior leaf. The anterior lateral supplementary leaflet extends from the base of the posterior leaf anteriorly beneath the anterior leaf on to the upper lip but does not reach the median line. This condition is similar to that found in H. pygmaeus, in which the anterior leaflets on each side do in fact extend anteriorly to the median line, but are separated by a deep emargination above the centre of the upper lip. The posterior lateral supple- mentary leaflet is broad but is shorter than the anterior leaflet, only just extending on to the upper lip. The anterior leaf is not emarginated. The internarial septum is more or less parallel-sided and is not inflated : the narial lappets are well developed and project slightly above the level of the anterior leaf. Although lying in a depres- sion, the nostrils are not enclosed by narial pockets. The intermediate part of the leaf is slightly inflated and cushion-like but has no swollen prominences. The posterior leaf is thin, its upper edge semicircular with no trace of lobulation, and is supported by three well-defined septa enclosing four small pockets. There is a frontal sac in males, represented in female specimens by a depression containing a tuft of hairs. The skull is short and broad, with an inflated, almost globose braincase and a low to moderate sagittal crest. The interorbital region is constricted, with sharply defined supraorbital ridges. There is a shallow frontal depression and the rostral eminences are moderately inflated, separated by a shallow groove. The zygomata are slender, with a low to moderate jugal projection, the zygomatic width exceeding the mastoid width. The postorbital processes are incipient, giving the rostrum from above a slightly pentagonal aspect. The anteorbital foramen is elongate and closed by a narrow bar. The premaxillae are fused for their entire length, their junction with the maxillae shallowly V-shaped. Their anterior enclosing processes are slender and do not entirely enclose the more or less oval anterior palatal foramina. The palate is short and broad, the palation U-shaped with usually a small median post-palatal spicule. The mesopterygoid fossa is wide, the vomer not projecting or projecting only slightly, the pterygoids elongated, with the wide sphenoidal bridge partially concealing elongate lateral apertures. There is a well-defined sphenoidal depression, while the width of the cochleae is a little greater than their distance apart. The upper incisors are not bilobed or are only very weakly bilobed, with only a trace of the external lobe. The upper canines have a moderate anterior cusp and a well developed posterior cusp, both low. The anterior upper premolar (pm 2 ) is usually much reduced, minute, extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is reduced to one third of the length of the anterior ridge. The crown area of the outer lower incisors is about the same or very slightly greater A REVISION OF HIPPOSIDEROS 53 than that of the inner pair. The anterior lower premolar (pm 2 ) is usually reduced to one half or less of the length and height of the second lower premolar (pm 4 ) . DISTRIBUTION : India ; Malay Peninsula east to Solomon Islands and New Hebrides ; northern Australia. Hipposideros galeritus galeritus Cantor Hipposideros galeritus Cantor, 1846 : 183. Penang. Tate (1941 : 367) has thrown considerable doubt on the authenticity of the skull of the type specimen of Hipposideros galeritus Cantor and has suggested that the skin and skull may be mismatched. The type specimen is now B.M. 79.11.21.85 in the collections of the British Museum (Natural History), a skin and skull, labelled as a male. In the register of the mammal collections in the British Museum (Natural History) for 1879, the specimen is listed as a skin only, with no mention of a skull. The skin is preserved in the dry state and fortunately is in good condition. The ears are broad at the base, triangular and sharply pointed, their posterior margins with a concavity just behind the tip. The noseleaf has two lateral supplementary leaflets and the posterior leaf is thin, supported by three septa. The skull associated with this skin is small, rather elongate, with a low sagittal crest, constricted inter- orbital region, no supraorbital ridges and moderately inflated rostral eminences. The zygomata are slender, with a well-developed superior projection, the zygomatic width (by extrapolation) less than the mastoid width. The anteorbital foramen is slightly elongate, closed by a narrow bar. The junction of the premaxillae with the maxillae is acutely V-shaped. The palation is U-shaped, with the vomer projecting very slightly into the mesopterygoid fossa. The sphenoidal bridge is moderate, not concealing the lateral apertures. There is a shallow sphenoidal depression. The upper incisors are weakly bilobed and the upper canines are slender, with a low posterior cusp. The anterior upper premolar (pm 2 ) is minute, extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact. The posterior ridge of the third upper molar is one half the length of the anterior ridge. The crown area of the outer lower incisors is slightly greater than that of the inner pair and the anterior lower premolar (pm 2 ) is two thirds the length and one half the height of the second lower premolar (pm 4 ). It is clearly the skull of a member of the bicolor subgroup and agrees closely with H. ater. It is therefore necessary to adopt the suggestion by Tate that Hipposideros galeritus Cantor be restricted to the skin of the type specimen, disregarding the skull hitherto associated with that skin. The characters of H. g. galeritus are much as in the species diagnosis. The upper incisors are not bilobed, the outer lobe being obsolete. The anterior upper premolar (pm 2 ) is very small, extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact. The anterior lower premolar (pm 2 ) is one third to one half the length and height of the second lower premolar (pm 4 ). Davis (1961 : 90) gives a description, with measurements, of a series of H. g. galeritus from the Federation of Malaya. DISTRIBUTION : Malay Peninsula ; Riau Archipelago ; Banka Island ; South Natuna Islands. 54 J. E. HILL Hipposideros galeritus brachyotus (Dobson) Phyllorhina brachyota Dobson, 18745 : 237. Central India. The characters of this subspecies are largely those of the nominate subspecies, but the post-palatal spicule is usually very small or absent, and the dentition is less advanced. The anterior upper premolar (pm 2 ), although small, is not always completely extruded from the toothrow but is compressed tightly between the outer margins of the canine and the second upper premolar (pm 4 ), while in some specimens it is extruded from the toothrow with the canine and the second upper premolar in contact. The posterior ridge of the third upper molar is one half the length of the anterior ridge. The anterior lower premolar (pm 2 ) is less reduced than in H. g. galeritus, and is nearly as long as the second lower premolar (pm 4 ) and two thirds its height. Brosset (1962 : 618) has studied the biology of H. galeritus in India, and gives measurements and notes on its colour variation. DISTRIBUTION : Ceylon ; India : Mysore ; Bombay ; Bengal. Hipposideros galeritus schneideri Thomas Hipposideros schneidersi Thomas (misprint), 19045 : 722. Upper Langkat, Sumatra. The posterior leaf is supported by a prominent median septum, the lateral septa not greatly developed and represented by low ridges. The post palatal spicule is absent. The upper canines have a low anterior cusp, their posterior cusp insignifi- cant or low. The anterior upper premolar (pm 2 ) is minute or lacking (it is absent in one side of the jaw of the type specimen, vestigial in the other side) the canine and the second upper premolar (pm 4 ) in contact. The anterior lower premolar (pm 2 ) is much reduced and is one third to one half the length and height of the second lower premolar (pm 4 ). Tate (1941 : 367) suggests that H. g. schneideri is a derived species of H. galeritus. There seems no reason, however, to separate it from this species with which it shares a majority of characters, differing principally in the greater reduction of the anterior premolars. DISTRIBUTION : Sumatra ; Engano Island ; Sipora Island ; Mentawi Islands : North Pagi. Hipposideros galeritus labuanensis (Tomes) Phyllorrhina labuanensis Tomes, i85Qa : 537. Labuan Island. Hipposideros insolens Lyon, 1911 : 129. Upper Pasir River, southeastern Borneo. This subspecies is very similar to H. g. galeritus. The post-palatal spicule may be present or absent and there is a well-defined sphenoidal depression. The upper incisors are not bilobed or only slightly so, and the upper canines have well defined anterior and posterior cusps. The anterior upper premolar (pm 2 ) is small, extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is one third to one half the length of the anterior ridge. The anterior lower premolar (pm 2 ) is reduced to A REVISION OF HIPPOSIDEROS 42 44 46 48 SO 52 54 55 H. G. BRACHYOTUS H. G. GALERITUS H.G. SCHNEIDERI 44 46 48 5O 52 54 FIG. 12. Length of forearm in Hipposideros galeritus one half or a little less the length and height of the second lower premolar (pm 4 ) to two thirds its length and height. Davis (1962 : 39) provides a description, with measurements, of a series of H . g. labuanensis from North Borneo. Lyon originally described H. insolens as a species distinct from H. galeritus, recording both from the same localities in Borneo. According to Lyon the dis- tinguishing characters of H. insolens were its longer forearm and tibia when compared with H. galeritus. Chasen (1940 : 46) listed H. insolens as a subspecies of H. longicauda, here considered to be a subspecies of H. galeritus. Tate (1941 : 368) retained H. insolens as an apparently distinct species and from an examination of the original series described by Lyon stated that in comparison with H. g. labuanensis 5 6 J. E. HILL the jugal prominence of the zygoma in H. insolens rose more abruptly, the ante- orbital foramen was almost pore-like and not elongate as in H. g. labuanensis while the palate was a little longer (extending a little behind the second upper molar) and lacked a spine. He also noted that the toothrow was shorter, with the W- pattern of the third upper molar less reduced and that the anterior lower premolar (pm 2 ) was more elongate than wide. No topotypical specimens of H. insolens are available but the collections of the British Museum (Natural History) contain an adequate series of Hipposideros galeritus from Borneo and this series suggests that H. insolens as understood by Lyon and Tate may well refer to large individuals of H. g. labuanensis. The jugal prominence in a long series of H. g. labuanensis, although usually moderate in its development is sometimes found to be not greatly developed, rising only gently from the zygoma while the anteorbital foramen, usually elongate, is on occasion rounded and more pore-like. The rear of the palate varies from a line joining the posterior faces of the second upper molars (m z - z ) to a line joining the centres of the third upper molars (m 3 - 3 ), while the post- palatal spicule is sometimes absent or incipient. The length of the posterior ridge of the third upper molar varies from one third to one half the length of the anterior ridge, while the anterior lower premolar (pm 2 ) varies from less than one half the length and height of the second lower premolar (pm 4 ) to two thirds its length and height. DISTRIBUTION : Borneo ; Labuan Island ; Philippine Islands : Mindanao (Sanborn (1952 : 104) as H. g. galeritus : see also Laurie & Hill (1954 : 55)). Hipposideros galeritus longicauda (Peters) Phyllorhina longicauda Peters, 1861 : 708. Java. There are no specimens of this subspecies in the collections of the British Museum (Natural History). Sody (1930 : 270) describes four specimens in some detail. DISTRIBUTION : Java. Hipposideros galeritus celebensis Sody Hipposideros celebensis Sody, 1936 : 47. Mampoe Cave, 20 kilometres north of Watoe Pone, south Celebes. There are no specimens oi this subspecies in the collections of the British Museum (Natural History). A detailed description (as H. g. galeritus) is given by Sody (1930 : 268) of specimens from Celebes subsequently described by the same author as H. celebensis. DISTRIBUTION : Celebes. Hipposideros galeritus batchianus Matschie Hipposideros batchianus Matschie, 1901 : 273. Batchian Island, Molucca Islands. There is no example of this subspecies in the collections of the British Museum (Natural History). A REVISION OF HIPPOSIDEROS 57 Hipposideros galeritus cervinus (Gould) Rhinolophus cervinus Gould, 1863 : pi. 34, letterpress. " Caves on Albany Island " (label on skin of type specimen). Cape York, northern Queensland, Australia. The characters of this subspecies are largely those of the nominate subspecies, the anterior upper premolar (pm 2 ) reduced, extruded from the toothrow, the canine 42 46 H. G. LABUANENSIS H. G. CELEBENSIS (FROM SODY (1930:268")) H. G. CERVINUS 44 46 43 SO 52 54 FIG. 13. Length of forearm in Hipposideros galeritus 58 J. E. HILL and the second upper premolar (pm 4 ) in contact or nearly so and the anterior lower premolar (pm 2 ) usually one half or less the length and height of the second lower premolar (pm 4 ) . Specimens from the Aru Islands have the anterior upper premolar (pm 2 ) much reduced, sometimes minute while in those from the Kei Islands this tooth is minute, almost invisible. Specimens from both groups of islands have the anterior lower premolar (pm 2 ) usually less than one half the length and height of the second lower premolar (pm 4 ). Those from Japen Island have a small but clearly visible anterior upper premolar (pm 2 ) and have the anterior lower premolar (pm 2 ) one half the length and height of the second lower premolar (pm 4 ). In specimens from New Guinea the anterior upper premolar (pm 2 ) is rather larger and more prominent while the anterior lower premolar (pm 2 ) is one half the length and height or slightly more of the second lower premolar (pm 4 ). Examples from New Ireland, the Solomon Islands, the New Hebrides and Cape York exactly resemble those from New Guinea but a single specimen from Kiriwina Island has the anterior lower premolar (pm 2 ) larger, equal nearly to two thirds the length and height of the second lower premolar (pm 4 ). There is some size variation over the range, specimens from the Aru and Kei Islands, Japen Island, Cape York and the Solomon Islands having the forearm averaging slightly shorter than those from New Guinea and the New Hebrides. DISTRIBUTION : New Guinea ; Waigeo Island ; Japen (=Jobi) Island ; Kei Islands ; Aru Islands ; northern Australia ; Bismarck Archipelago : New Ireland ; Trobriand Islands : Kiriwina ; Solomon Islands : Bougainville ; Guadalcanar ; Fauro ; Russell ; Rennell ; New Hebrides : Espiritu Santu ; Efate ; Santa Cruz Islands : Fenvaloa ; Vanikoro. Hipposideros crumeniferus (Lesueur & Petit) Rhinolophus crumeniferus Lesueur & Petit, 1807 : pi. 35. Timor Island. The status of this early name has already been discussed. Despite a careful search by the authorities of the Museum National d'Histoire Naturelle in Paris, no trace can be found of the type specimen. Hipposideros breviceps Tate Hipposideros breviceps Tate, 1941 : 358. North Pagi Island, Mentawei Islands, off west coast of Sumatra. Through the courtesy of the authorities of the American Museum of Natural History, New York, I have been able to examine two of the paratypes of H. breviceps Tate. The ears are broad, triangular, their anterior margins convex and their posterior margins with a concavity behind the acute point but otherwise convex. No definite internal fold can be discerned in the dry skins, but the membrane of the ears is thickened at the antitragal lobe. The ears are haired for one half or slightly more of their length. The noseleaf is small, with two lateral supplementary leaflets projecting beyond the margins of the anterior leaf. The anterior lateral A REVISION OF HIPPOSIDEROS 59 supplementary leaflet extends from the base of the intermediate part of the leaf anteriorly beneath the anterior leaf on to the upper lip, but does not reach the median line. The posterior leaflet does not extend forward in the same manner but termi- nates anteriorly at a point just on the upper lip. The anterior leaf has no median emargination, and the internarial septum is undeveloped. The narial lappets are well developed and project above the level of the anterior leaf. The nostrils lie in deep depressions but are not pocketed, while the intermediate part of the leaf is slightly inflated and cushion-like. The posterior leaf is high, its upper edge semi- circular, and is supported by the median septum and two lateral septa. The skull is very short and broad, with an inflated, broadened braincase and low sagittal crest. The interorbital region is moderately constricted and the supra- orbital ridges are poorly denned. There is a shallow frontal depression. The rostrum is rounded and not markedly pentagonal in outline, with the rostral emi- nences much inflated and individually swollen, separated by a groove. The zygomata are slender, with a high, well developed jugal projection, the zygomatic width exceeding the mastoid width. The anteorbital foramen is elongate, closed by a narrow bar. The premaxillae are short and do not project beyond the canines, making a shallowly V-shaped junction with the maxillae. The palate is short, the palation shallowly U-shaped, level with a line joining the posterior faces of the third upper molars (m 3 ~ 3 ), without a post-palatal spicule. The mesopterygoid fossa is wide, the vomer not projecting, and the expanded pterygoids are long, together with the wide sphenoidal bridge almost concealing elongate lateral aper- tures. There is a shallow sphenoidal depression and the cochleae are a little wider than their distance apart. The upper incisors are weakly bilobed, the outer lobe obsolescent and are strongly convergent, their tips almost touching. The upper canines have a small posterior cusp. The anterior upper premolar (pm 2 ) is minute and completely extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is one third the length of the anterior ridge. The crown area of the outer lower incisors is very slightly greater than the crown area of the inner lower incisors, while the anterior lower premolar (pm 2 ) is very small, one quarter or little more the length and height of the second lower premolar (pm 4 ) . Externally, H. breviceps is very similar to H. galerilus, but has the ears slightly less extensively haired. The skull closely resembles that of H. galeritus but is relatively shorter, with more inflated rostral eminences. In the shortening of the skull it tends towards H. dyacorum and this feature, together with the reduction of the anterior premolars, no doubt led Tate (1941 : 358) to consider H. breviceps as a half-way stage between an insolens-like form and dyacorum. However, there appears to be no close relation between H. breviceps and H. dyacorum. The ears of H. dyacorum have a well-defined internal fold, absent in H. breviceps, while the noseleaves of the two species differ sharply in the absence of lateral supplementary leaflets in H. dyacorum and the presence of two well-developed leaflets in H. breviceps. The skull of H. breviceps, although similar in outline to that of H. dyacorum, is less shortened, and has a much less constricted interorbital region. 60 J. E. HILL There is a shallow frontal depression, absent in H. dyacorum, and there are no marked supraorbital ridges like the prominent, sharply defined ridges of that species. Hipposideros breviceps has the palate less shortened than H. dyacorum and its palation lacks a median emargination : it has a wider mesopterygoid fossa and wider, more spreading pterygoids, with an unconstricted sphenoidal bridge. The affinities of H. breviceps clearly lie with H . galeritus, from which it differs in a number of cranial features, such as its relatively shorter skull and less well-defined supraorbital ridges, slightly more inflated rostral eminences, shorter premaxillae, shorter palate and strongly convergent upper incisors. It is approached most nearly by H. g. schneideri, which closely resembles H. breviceps and has a similarly shortened skull, but which is larger and has rather more prominent supraorbital ridges, less inflated rostral eminences and longer premaxillae. The dentition of H. g. schneideri is almost identical with that of H. breviceps, with the anterior upper premolar (pm 2 ) minute or absent and the anterior lower premolar (pm 2 ) much reduced. The upper incisors of H. g. schneideri, however, are much less strongly convergent at their tips than in H. breviceps, in which the tips almost touch, and the anterior lower premolar (pm 2 ) is slightly less reduced. The majority of its characters indubitably ally H. breviceps closely to H. galeritus, with which it is apparently sympatric in the Mentawei Islands (Tate (1941 : 368, 391) records H. g. schneideri from North Pagi Island) and it is considered to be a recently derived species, close to H. galeritus and having much the same relationship to that species as H. beatus has to H. coffer in the Ethiopian region. DISTRIBUTION : Mentawei Islands : North Pagi Island. Hipposideros curtus G. M. Allen Hipposideros curtus G. M. Allen, 1921 : 194. Sakbayeme, Cameroons. Hipposideros sandersoni Sanderson, 1937 : 2 9> 2 9^- Near Mamfe, Nigeria. The ears are very large and broad, rather rounded and bluntly pointed, with a sharp concavity behind the tip. There is a small antitragal fold and they are haired for one half their length. The noseleaf is broad, with two small lateral supplementary leaflets. The anterior leaf has a small median emargination. The internarial septum is swollen and inflated, especially posteriorly between the nostrils, where it forms a slightly disc-like structure, the disc not however concealing the nostrils. The intermediate part of the leaf is cushion-like, with a low median eminence. The posterior leaf is high, its upper edge semicircular, and is supported by three septa enclosing deep pockets. A frontal sac is present in both sexes. The skull is small, short and broad, with a low sagittal crest. The interorbital region is moderately constricted, with ill-defined supraorbital ridges. The frontal region is more or less pentagonal as in H. galeritus, with a shallow frontal depression and well-inflated rostral eminences separated by a shallow groove. The zygomata are moderate with a low jugal projection and the anteorbital foramen is large and rounded, closed by a narrow bar. The premaxillae make a V-shaped junction with the maxillae and the palate is short, with a square palation. The mesopterygoid A REVISION OF HIPPOSIDEROS 61 fossa is wide with a slightly projecting vomer, while the pterygoids are short, with a narrow, sharply constricted sphenoidal bridge exposing slightly elongate lateral apertures. There is a very shallow sphenoidal depression and the width of the cochleae is equal to their distance apart or is very slightly greater. The upper incisors are weakly bilobed and the upper canines shallowly grooved anteriorly, with low anterior and posterior cusps. The anterior upper premolar (pm 2 ) is small, slightly extruded but compressed between the canine and the second upper premolar (pm 4 ). The posterior ridge of the third upper molar is much reduced and obsolescent. The crown area of the outer lower incisors is a little greater than that of the inner lower incisors, while the anterior lower premolar (pm 2 ) is considerably reduced, to one half the length and one third the height of the second lower premolar (pm 4 ). Hipposideros curtus, like H. fuliginosus, is a species evidently derived from the bicolor -caff er line and presents features which ally it to both the bicolor and galeritus sections of the bicolor group. Its external characters, especially the form of the ears, demonstrate its affinity to the bicolor subgroup but its short, broad skull tends towards the galeritus subgroup while H. curtus has also the short pterygoids and narrow sphenoidal bridge characteristic of H . caffer and its immediate allies. DISTRIBUTION : West Africa : Cameroons ; Nigeria. Hipposideros fuliginosus (Temminck) Phyllorrhina fuliginosa Temminck, 1853 : 77. " La cote de Guine ". The ears are large and broad, sharply triangular and acutely pointed, with a slight trace of an internal fold. Their posterior margins are slightly concave behind the tip, and they are haired for one half of their length. The noseleaf is comparatively broad, with two lateral supplementary leaflets, the anterior leaflet extending anteriorly for a short distance below the anterior leaf. The anterior leaf is simple with the internarial septum uninflated and with well-developed narial lappets. The intermediate part of the leaf is inflated, with a low median eminence. The posterior leaf is high, its upper edge semicircular and unlobulated. It lacks supporting septa but has a very low transverse structure, its upper edge not serrated, developed from its posterior face. There is no trace of a frontal sac in two male specimens (see also Hayman (1946 : 772)). The skull is elongate, with a low to moderate sagittal crest. The interorbital region is long and relatively unconstricted, with barely denned supraorbital ridges. There is no frontal depression and the rostral eminences are moderately inflated and separated by a shallow trough. The zygomata are slender with a moderate jugal projection and the anteorbital foramen is elongate and closed by a narrow bar. The anterior palatal foramina are elongate and not enclosed by the premaxillae, which make a V-shaped junction with the maxillae. The palation is square with a shallow median emargination. The mesopterygoid fossa is wide with long, moder- ately expanded pterygoids which partially conceal elongate lateral apertures. The sphenoidal bridge is moderately constricted and there is a moderate sphenoidal depression. The cochleae are rather less in width than their distance apart. The 62 J. E. HILL upper incisors bear only a trace of the outer lobe and the upper canines have a shallow groove on their anterior faces, with weak anterior and posterior cusps. The anterior upper premolar (pm 2 ) is much reduced, extruded, with the canine and the second upper premolar (pm 4 ) almost in contact. The posterior ridge of the third upper molar is one third the length of the anterior ridge. The crown area of the outer lower incisors slightly exceeds that of the inner lower incisors, and the anterior lower premolar (pm 2 ) is one half the length and height of the second lower premolar (pm 4 ). There is little doubt that H. fuliginosus is a derivative of the line leading from H. bicolor and its allies to the more specialized H. coffer, possessing certain features common to the bicolor subgroup and others common to the galeritus subgroup, to which H. caffer is allocated. It has broad, sharply triangular ears, two lateral supplementary leaflets and the posterior leaf, which lacks supporting septa, has a low transverse structure developed from its posterior face. These features ally it to H. caffer : however, its skull is elongate with the long interorbital region characteristic of the members of the bicolor subgroup. DISTRIBUTION : West Africa : Liberia ; Togo ; Ghana ; Nigeria ; Cameroons ; Gaboon ; Congo. 58 6O 62 64 H. FULIGINOSUS 58 6O 62 64 FIG. 14. Length of forearm in Hipposideros fuliginosus Hipposideros caffer The ears are large, broad, triangular and acutely pointed. Their posterior margins are concave just behind the tip. There is a slight thickening at the antitragal fold and the ears are haired for two thirds to three quarters of their length. The nose- leaf is comparatively simple with two lateral supplementary leaflets, the anterior leaflet extending anteriorly beneath the anterior leaf, sometimes reaching or almost reaching the median line. The anterior leaf is simple and has no median emargina- tion. The internarial septum is slightly inflated. The intermediate part of the leaf is cushion-like, with a low to moderate median eminence, sometimes with low lateral eminences. The posterior leaf is thin, its upper edge semicircular, unlobulated and lacks supporting septa. Its posterior face bears a transverse supplementary structure with a serrated upper edge below that of the true posterior leaf. A frontal sac is present in both sexes but is very small in females. The skull is short and broad, closely similar in outline to the skull of H. galeritus. The braincase is broad with a low sagittal crest and the interorbital region is not A REVISION OF HIPPOSIDEROS 63 greatly constricted, the supraorbital ridges low but sharply defined. There is no frontal depression and the rostral eminences are moderately inflated. The zygomata have a high jugal process, set far back, and the anteorbital foramen is elongate, closed by a narrow bar. The premaxillae make a U-shaped or shallowly V-shaped junction with the maxillae and the anterior palatal foramina are slightly elongate, not enclosed within the premaxillae but almost closed posteriorly by the delicate anterior enclosing processes. The palation is nearly square with a small median notch while the mesopterygoid fossa is wide, the vomer not projecting and the ptery goids short, with a narrow, sharply constricted sphenoidal bridge exposing wide lateral apertures. There is a shallow sphenoidal depression. The cochleae are moderate, their width equal to or a little greater than their distance apart. The upper incisors are weakly bilobed, their outer lobes obsolescent. The upper canines have their anterior faces flattened or with a shallow, ill-defined groove and have a moderate anterior cusp and a slightly higher posterior cusp. The anterior upper premolar (pm 2 ) is small or very small, slightly extruded, compressed between the canine and the second upper premolar (pm 4 ), which usually are not in contact, sometimes more fully extruded with these teeth in contact or nearly so. The posterior ridge of the third upper molar is not greatly reduced, equal to one half the length of the anterior ridge or more. The crown area of the outer lower incisors is very slightly greater than that of the inner pair. The anterior lower premolar (pm 2 ) is not excessively reduced, one half or more the length and height of the second lower premolar (pm 4 ). Hipposideros caffer is the principal representative of the galeritus subgroup in the Ethiopian region, and like H. galeritus, is the dominant and most widely distri- buted species of the subgroup in its region. Despite its superficial resemblance to H. galeritus, it differs widely from this and its associated species in the structure of its posterior noseleaf and in the characters of the post-palatal region. These differences suggest remote origin in the bicolor group although there seems little doubt that H. galeritus and H. coffer have developed from a common if remote source. Both are similarly specialized in a number of features, notably in the extent of body fur on the ears and in the number of lateral supplementary leaflets : both have a characteristic short broad skull very different from the elongate narrow skull of the bicolor type yet both can be linked to the bicolor subgroup by a series of species exhibiting a variety of features on the one hand found in bicolor and galeritus and on the other in bicolor and caffer. A number of subspecies have been proposed to divide H. caffer, mainly on the basis of size, and there is considerable dimensional overlap between them. No attempt has been made in the present work to establish the validity of the named forms of H. caffer but there seems little doubt from the very large series from most parts of its range now preserved in the collections of the British Museum (Natural History) that the variation in size of the species is largely clinal. Smaller subspecies (H. c. caffer, H. c. tephrus) are found in the Yemen, in the eastern part of Africa, in North Africa and on the west coast south to Sierra Leone and the Gold Coast. A larger subspecies (H. c. ruber) replaces H. c. caffer in the west of Kenya, Uganda 64 J. E. HILL and Tanganyika, extending westwards through the Congo, to be itself replaced in west Africa by another large subspecies (H. c. guineensis) and in Angola and south- west Africa by yet another large subspecies (H. c. angolensis). The distributional limits of the subspecies are difficult of definition, and series from some localities cannot be allocated with certainty to subspecies. DISTRIBUTION : Yemen ; the greater part of Africa excluding the Sahara desert and the extreme south. Hipposideros caffer coffer (Sundevall) Rhinolophus caffer Sundevall, 1846 : 118. Near Port Natal. Phyllorr hina gracilis Peters, 1852 : 36, pi. 7, figs. 1-4, pi. 13, figs. 14-15. Tette, Lower Zambezi, Mozambique. Phyllorrhina bicornis Heuglin, 1861 : 4, 7. Keren, Eritrea. DISTRIBUTION : Yemen ; Eritrea ; Somaliland ; Sudan ; Ethiopia ; Kenya (part) ; Tanganyika (part) ; Southern Rhodesia ; Northern Rhodesia (part) ; Nyasaland ; Natal ; Transvaal ; Zanzibar ; Pemba Island; Congo (part). Hipposideros caffer tephrus Cabrera Hipposideros tephrus Cabrera, 1906 : 358. Mogador, Morocco. Hipposideros braima Monard, 1939 : 73, fig. 5. Bagingara, Portuguese Guinea (see Aellen (i956b : 26)). Hipposideros braima is further described and illustrated by Veiga-Ferreira (1949 : 193). DISTRIBUTION : North Africa : Morocco ; Senegal (part) ; Nigeria (part) ; Sierra Leone (part) ; Ghana (part) ; Senegambia. Hipposideros caffer ruber (Noack) Phyllorhina rubra Noack, 1893 : 586, pi. 18, figs. 14-15. " Lugerrunjere Fluss ", Tanganyika. Hipposideros caffer centralis Andersen, I9o6b : 275, 277. Entebbe, Uganda. DISTRIBUTION : Tanganyika (part) ; Kenya (part) ; Uganda ; Congo (part) ; North Rhodesia (part) ; Angola (part) (Sanborn (1950 : 58)) ; recorded from Guinea by Aellen (iQ56a : 889). Hipposideros caffer angolensis (Seabra) Phyllorhina angolensis Seabra, 1898 : 256. Rio Coroca, Angola. DISTRIBUTION : Angola ; southwest Africa ; Gaboon (part) ; recorded from Ghana by Booth (1956 : 137). Hipposideros caffer guineensis Andersen Hipposideros caffer guineensis Andersen, igo6b : 275, 278. Como River, 70 miles from Gaboon. DISTRIBUTION : Gaboon (part) ; Cameroons ; Nigeria (part) ; Ghana (part) ; Gambia ; Sierra Leone (part) ; Liberia ; San Thom6 Island ; Principe Island ; Fernando Po ; Spanish Guinea ; Senegal (part) (Aellen (i956b : 26)). A REVISION OF HIPPOSIDEROS 42 44 46 48 SO 52 54 56 H. C. CAFFER H. C.TEPHRUS H. C 42 44 46 48 SO 52 54 56 ZOO 11, 1. E FIG. 15. Length of forearm in Hipposideros caffer 66 J. E. HILL Hipposideros caffer niapu J. A. Allen Hipposideros caffer niapu J. A. Allen, 1917 : 431. Niapu, northeastern Congo. DISTRIBUTION : Congo (part). Hipposideros beatus The ears are large, broad, triangular and pointed. Their posterior margins are concave just behind the tip. There is a slight thickening at the antitragal fold and they are haired for two thirds of their length. The noseleaf exactly resembles H. C.ANGOLENSIS H C.GUINEENSIS H.C. NIAPU H. BEATUS 42 44 46 48 SO 52 54 56 58 FIG. 1 6. Length of forearm in Hipposideros caffer and H. beatus A REVISION OF HIPPOSIDEROS 67 that of H. coffer. The skull is short and broad and very similar to that of H. coffer. There is a low sagittal crest and the interorbital region is only slightly constricted, with low but sharply denned supraorbital ridges. There is a very shallow frontal depression and the rostral eminences are inflated and separated by a shallow groove. The zygomata are slender, with a high jugal process, and the anteorbital foramen is rounded, closed by a narrow bar. The premaxillae make a V-shaped junction with the maxillae and the palation is square with a median notch. The mesoptery- goid fossa is a little narrower than in H. coffer, the pterygoids short with a narrow, sharply constricted sphenoidal bridge exposing wide lateral apertures. There is a shallow sphenoidal depression and the width of the cochleae is equal to their width apart or is slightly greater. The upper incisors are weakly bilobed and the upper canines have a weak anterior cusp and a low but well developed posterior cusp. The anterior upper premolar (pm 2 ) is minute, extruded, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is one third or less the length of the anterior ridge. The crown area of the outer lower incisors is slightly greater than the crown area of the inner pair. The anterior lower premolar (pm 2 ) is a little more than one half the length and height of the second lower premolar (pm 4 ). Hipposideros beatus differs only very slightly from H. coffer, and may be dis- tinguished from that species chiefly by its smaller size and the greater degree of reduction of its anterior lower premolar (pm 2 ) . The two species provide an interest- ing parallel in the Ethiopian region to the Asiatic species H. galeritus and H. breviceps, the latter differing only very slightly from the former. Hipposideros beatus is slightly more advanced than H. coffer in the details of its dentition and is regarded as a very close derivative of H. coffer. DISTRIBUTION : West Africa from the Congo to Sierra Leone. Hipposideros beatus beatus Andersen Hipposideros beatus Andersen, igo6b : 279. 15 miles from Benito River, West Africa. Hipposideros nanus J. A. Allen, 1917 : 434. Faradje, Uele District, eastern Congo. Hayman (1936 : 919) thought that H. nanus Allen probably referred to the dark phase of H. beatus. Verschuren (1957 : 371) considered it to be a distinct species, distinguished chiefly by its colour and by the insertion of the wing membrane on the tibia and not on the metatarsals as in H. beatus. These features are variable in a small series of H . beatus and for the present H. nanus is provisionally retained in the synonymy of this species. DISTRIBUTION : Congo (part) ; Cameroons ; Nigeria ; Ghana ; Liberia ; Sierra Leone. Hipposideros beatus maximus Verschuren Hipposideros beatus maximus Verschuren, 1957 : 3 62 > 3 6 5- Pidigala-Nord, Pare National de la Garamba, Congo. 68 J. E. HILL Hipposideros coxi Shelford (Figure 17) Hipposideros coxi Shelford, 1901 : 113. Mount Penrisen, Sarawak, Borneo, 4,200 feet. A species of medium size with very large, sub-triangular ears, broad at the base, bluntly pointed, with their anterior margins slightly concave in the upper portion, their posterior margins strongly so for the upper third, thereafter convex. The antitragal lobe is well developed, with a prominent internal fold, and the ears are haired for two thirds of their length. The noseleaf is large and forms a structure of extraordinary complexity, covering the entire muzzle, with two lateral supplementary leaflets. The anterior leaflet on each side extends anteriorly beneath the anterior leaf to become continuous over the upper lip, with no median emargination. At its upper or posterior end this leaflet terminates on each side in a small wart-like protuberance or papilla, at about the level of the eye, pierced by a minute pore. A small, sharply triangular pointed leaflet extends upwards from this papilla behind the lateral margin of the anterior leaf. This extension of the anterior supplementary leaflet at its upper end joins a further cutaneous leaflet extending from the rear of the intermediate part of the leaf, to which it is attached, downwards to a point just above the eye, forming a structure of the shape of an inverted Y. The lateral margin of the posterior leaf is joined to the upper surface of this cutaneous leaflet : the junction of the triangular extension of the supplementary leaflet is on its lower surface at a point nearer to the eye, these leaflets enclosing deep pockets behind the intermediate part of the leaf. Cutaneous outgrowths composed either of the anterior lateral supplementary leaflet or of extensions therefrom thus extend completely beneath the anterior leaf and beneath much of the margins of the inter- mediate part of the leaf, forming deep pockets beneath the latter. The posterior lateral supplementary leaflet is broad and extends from a point level with the wart terminating the anterior leaflet only for a short distance on to the muzzle, just reaching the upper lip. This observation is contrary to that of Shelford, who stated in the original description that the lower (or posterior) leaflet consisted of two separate parts : it is clear from the type specimen, however, that Shelford mistook the cutaneous leaflet extending downwards from the base of the posterior leaf to be a part of the posterior leaflet, but it is apparently a structure quite distinct from either of the lateral supplementary leaflets. The anterior leaf is very large, lacking a median emargination, completely covering the muzzle and with its margin pro- jecting beyond the upper lip and concealing the anterior lateral supplementary leaflets, which however project beyond the lateral margins of the anterior leaf. The internarial septum is slightly bulbous anteriorly, dividing two deep pockets anterior to the nostrils. The narial region is greatly specialized, the narial lappets greatly developed and with their bases expanded towards the median line to join the internarial septum, forming deep pouches at the base of which the nostrils are situated. These pouches have slightly crescentic, elongate openings and the narial lappets project above the level of the anterior leaf. The intermediate part of the leaf is inflated with a prominent median ridge, sparsely haired at its upper end : A REVISION OF HIPPOSIDEROS 69 laterally, there are two low eminences on each side of this ridge, just beneath the upper margin of the intermediate part of the leaf. The lateral margins of the inter- mediate part of the leaf are extended laterally to form a triangular lappet on each side extending over the bases of the margins of the anterior leaf and over the bases of the lateral supporting septa of the posterior leaf. The posterior leaf is very high, thick, its upper edge rounded and is supported by three septa. The central or median septum is narrow and blade-like, with a large, deep cell flanking it on each side. The lateral septa are very broad, their upper ends continuous with the face of the posterior leaf, the cells external to these septa narrow but very deep. The upper third of the posterior leaf extends above these septa and in the type specimen (in alcohol) is folded back upon itself. Laterally, the margins of the posterior leaf extend downwards and are joined to cutaneous leaflets attached on each side to the rear of the lateral lappets of the intermediate part of the leaf. At their upper ends these leaflets form the floor of the deep lateral cells of the posterior leaf and the roof of deep pockets beneath the intermediate part of the leaf. FIG. 17. Hipposideros coxi $ (Type B.M. 1.6.23.1) (x3) The skull is elongate and comparatively narrow, with an elongate braincase and low sagittal crest. The interorbital region is not sharply constricted and there are no supraorbital ridges. There is no frontal depression and the rostrum is elongate and high, with the rostral eminences much inflated, separated by a shallow depression. The zygomata are slender with a low jugal projection and the zygomatic width is less than the mastoid width. The anteorbital foramen is small and rounded, separated from the orbit by a massive bar of bone enclosing a long anteorbital canal. 7 o J. E. HILL The premaxillae of the type specimen are missing. The palate is not markedly shortened, the palation shallowly V-shaped with a small post-palatal spicule. The mesopterygoid fossa is wide, the vomer not projecting, with long, expanded pterygoids and wide sphenoidal bridge almost concealing narrow, elongate lateral apertures. There is a shallow sphenoidal depression and the width of the cochleae slightly exceeds their distance apart. The upper canines have low anterior and posterior cusps and the anterior upper premolar (pm 2 ) is very small, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) almost in contact. The second anterior upper premolar (pm 4 ) has a well-developed anterior cusp. The posterior ridge of the third upper molar is one half the length of the anterior ridge. The crown area of the outer lower incisors slightly exceeds that of the inner pair and the anterior lower premolar (pm 2 ) is slightly over one half the length of the second lower premolar (pm 4 ) and is one half of its height. Hipposideros coxi is very sharply removed from the other species of the bicolor group by the great specialization of its noseleaf , which in the degree of its complexity is approached by no other species of the group. Certain features, however, ally it to the bicolor group. Its ears are of the bicolor type, with a well-developed internal fold. The anterior lateral supplementary leaflet extends beneath the anterior leaf to reach the median line : a similar condition is found in H. jonesi and H. pygmaeus, which have the leaflets divided above the centre of the upper lip by a deep emargina- tion. There is no such emargination in H. coxi, and the anterior lateral supplemen- tary leaflets form an uninterrupted cutaneous frill beneath the anterior leaf. Narial pockets surrounding the nostrils, greatly developed in H. coxi, are less pronounced in H. pygmaeus and are developed to some extent in H. obscurus. The elongate skull and narrow zygomata of H. coxi demonstrate its affinity to the bicolor subgroup, but its complex noseleaf and long anteorbital canal, enclosed by a massive bar of bone, clearly remove it from any close relation to the members of this subgroup or of the galeritus subgroup. Its great degree of specialization suggests that H. coxi must be considered an isolated species of remote origin and it is probably derived from the basal stem of the galeritus subgroup. Hipposideros papua (Thomas & Doria) (Figure 18) Phyllorhina papua Thomas & Doria, 1886 : 204. Korido, Misori Island, Geelvinck Bay, Netherlands New Guinea. (?) Phyllorhina cervina var. misorensis Peters, 1906: pi. 5L, figs. 4, 4a, 4b. The ears are long, broad and triangular, acutely pointed, their posterior margins forming a smooth convex curve. There is a small, slightly thickened antitragal fold and the ears are apparently haired for one third or a little more, perhaps one half of their length (the syntype and only available specimen has the skin in bad condition). The noseleaf is broad, covering the entire muzzle, with three lateral sup- plementary leaflets, the first normal, extending upwards to the base of the posterior leaf just anterior to the eye, the central or second leaflet extending from a point below and just anterior to the eye anteriorly beneath the anterior leaf on to the A REVISION OF HIPPOSIDEROS 71 upper lip but not reaching the median line and the third leaflet very small, rudi- mentary and incipient. The anterior leaf is large, extending over the upper lip, but has no median emargination. The internarial septum is uninflated and the narial lappets well developed, with the internarial septum forming shallow narial pockets. The intermediate part of the leaf is simple with a low median ridge. The posterior leaf is high, its upper edge semicircular and thickened and is supported by three moderate septa enclosing shallow pockets. There is a small depression but no frontal sac in the female type specimen. FIG. 18. Hipposideros papua ? (Syntype B.M. 86.11.3.9) (x3) The skull is only slightly elongate, with a long, broadened braincase and moderate sagittal crest. The interorbital region is moderately constricted with barely defined supraorbital ridges. There is a moderate frontal depression, and the rostrum is slightly broadened, with well-inflated rostral eminences separated by a shallow groove. The zygomata are slender, with a moderate jugal process, and the ante- orbital foramen is large, elongate and closed by a narrow bar. The premaxillae make an acutely V-shaped junction with the maxillae and the anterior palatal foramina are elongate and not totally enclosed. The palate is not markedly shortened, the palation U-shaped with a small post-palatal spicule. The mesoptery- goid fossa is moderate and not greatly widened, the vomer not projecting and the pterygoids long with a wide sphenoidal bridge, almost completely concealing elongate lateral apertures. There is a moderate sphenoidal depression and the width of the cochleae is slightly greater than their distance apart. The upper incisors are not bilobed and the upper canines have low anterior and posterior cusps. The anterior upper premolar (pm 2 ) is small and is only slightly out of alignment in the toothrow. 72 J. E. HILL The posterior ridge of the third upper molar is obsolescent. The crown area of the outer lower incisors is equal to that of the inner pair and the anterior lower premolar (pm 2 ) is one half the length and height of the second lower premolar (pm 4 ). Tate (1941 : 369, 387) was uncertain of the taxonomic position of H. papua and regarded it as a perplexing species. Its ears and cranial characters suggest affinity with the bicolor group, but H. papua has a specialized noseleaf which indicates no close connection with any of the members of that group. Some affinity to the cyclops group is suggested by the extension of the second lateral leaflet beneath the anterior leaf, a feature otherwise peculiar to cyclops and its allies, but H . papua differs from these in its broad ears, the presence of an incipient third leaflet, un- enlarged bullae and convergent toothrows. A more tenuous affinity to the speoris group may be indicated by the presence of a rudimentary third leaflet. For the present H. papua must be regarded as a species of remote origin within the bicolor group, associated probably with the line leading to the muscinus group and less certainly with that leading to the speoris group. HIPPOSIDEROS CYCLOPS group The cyclops group as here understood includes the cyclops and muscinus groups of Tate (1941), both characterized by the great development of the ears and auditory region. The group includes six species, H. cyclops, H. camerunensis , H. muscinus, H. wollastoni, H. semoni and H. stenotis, all demonstrating a high degree of modifica- tion and specialization. The ears are long, narrow and acutely pointed, lacking any antitragal modification and not extensively haired. The noseleaves are much specialized, with two lateral supplementary leaflets. The anterior lateral supple- mentary leaflet is small, the posterior leaflet larger, extending posteriorly to the base of the posterior leaf, of which it forms a continuous part and in the majority of species extending anteriorly beneath the anterior leaf. The anterior leaf has no median emargination and the internarial septum is uninflated or only slightly thickened. The intermediate part of the leaf is flat or slightly cushion-like, specialized by the development to a greater or lesser degree of a median tubercle. The posterior leaf is moderate, with three supporting septa, its upper edge more or less semicircular, thickened, usually with a median club-like process (H. cyclops, H. camerunensis, H. muscinus, H. semoni, H. stenotis} or with a transverse supple- mentary structure developed from the posterior face of the leaf (H. wollastoni). The frontal sac is usually absent. The skull is short and broad, the braincase wide and almost globose, with a low sagittal crest. The interorbital region is much constricted, with low supraorbital ridges. The rostrum is greatly broadened, with well-inflated rostral eminences. The. zygomata are slender or moderate, with a low jugal process, and the anteorbital foramen is rounded, closed by a narrow bar, or pore-like. The palate is short and the pterygoids are long, together with the wide sphenoidal bridge almost concealing elongate lateral apertures. There is a shallow sphenoidal depression and the cochleae are greatly developed, their width equal to four or more times their distance apart. The outer lobe of the upper incisors is obsolescent or absent and the upper canines A REVISION OF HIPPOSIDEROS 73 in most cases lack cusps but have well-developed cingula. The anterior upper premolar (pm 2 ) is usually much reduced and is absent in one species. The posterior ridge of the third upper molar is obsolescent or well developed, in some species almost complete with the W-pattern of the tooth virtually complete. The anterior lower premolar (pm 2 ) is much reduced. The group is one of restricted distribution, with two species in the Ethiopian region and the remainder in New Guinea and northern Australia. As might be expected from this markedly discontinuous range, the group can be divided readily into two sharply denned sections, corresponding to the cy clops and muscinus groups of Tate (1941). Despite this sharp division, the two sections share features in common that make their independent origin unlikely. Their specialized ears and auditory region are unknown elsewhere in Hipposideros, as is the rearward extension of the posterior lateral supplementary leaflet to form a continuous part of the posterior leaf. The noseleaves are further specialized by the development of club-like projections, of which Tate (1941 : 379) observes that they are " structures so peculiar, specialized and seemingly functionless that they are unlikely to have arisen independently ". The two sections share further the same typically shortened, broadened pattern of skull, unwidened at the mastoid region and with wide pterygoids and greatly enlarged cochleae. The Ethiopian species H. cyclops and H. camerunensis are clearly the most primi- tive members of the group and may be readily distinguished from the Australasian species H. muscinus, H. wollastoni, H. semoni and H. stenotis by their much larger size. Their noseleaves differ markedly from those of the Australasian species in that the posterior lateral supplementary leaflet does not extend anteriorly beneath the anterior leaf, while the skulls of the Ethiopian species are more elongate than those of H. muscinus and its allies. The interorbital region is less sharply constricted in the Ethiopian species, and the rostral swellings, although inflated, are less swollen and more ossified. The premaxillae are wide and completely enclose narrow, elongate anterior palatal foramina, making a broad junction with the maxillae extending over almost the entire width of the palatal emargination in contrast to the narrow premaxillae of the Australasian species, which do not enclose the large oval anterior palatal foramina and which make a narrow junction with the maxillae, extending over only one third of the width of the palatal emargination. The cochleae are large but are less developed than those of the Australasian species, and are equal in width to approximately four times their distance apart. Hipposideros muscinus and its allies, confined to New Guinea and northern Australia, are much smaller and more specialized than H. cyclops and its close relative H. camerunensis. The noseleaves of the Australasian species are charac- terized by a number of specializations, among which the forward extension of the posterior lateral supplementary leaflet beneath the anterior leaf is the most notice- able. These species have the skull much shortened, especially in the rostral region, with a broad braincase and a constricted interorbital region. As in H. cyclops and H. camerunensis, the inflated braincase is not abruptly widened at the level of 74 J. E. HILL the mastoids. The interorbital region is short and much less elongate than in H. cyclops and H. camemnensis, while the rostrum, although broadened, is rounded and less angular. As in H. cyclops and H. camerunensis , the Australasian species have long pterygoids and a wide sphenoidal bridge, but the cochleae are more greatly expanded, their width in the majority of species equal to six or more times their distance apart. These differences suggest comparatively remote separation of the parental stocks of the two sections : H. cyclops and H. camerunensis have retained to some extent the elongate outline of the skull typical of the bicolor group, particu- larly in the elongate, broadened braincase, although the rostrum and palate are shortened, while H . muscinus and its allies are rather more specialized and represent a further trend of modification within the group. Although H. muscinus, H. wollastoni, H. semoni and H. stenotis are closely related, it is possible to discern a weak division among them. Hipposideros muscinus is the least specialized : the tubercles on its intermediate and posterior noseleaves are not greatly developed, it lacks a frontal depression, its anteorbital foramen is elongate, closed by a narrow bar, there is a shallow sphenoidal depression traversed by a low median ridge and its cochleae are the least expanded, in width equal to approximately four to five times their distance apart. The anterior upper premolar (pm 2 ) is not greatly extruded and the posterior ridge of the third upper molar is small, less than one third the length of the anterior ridge, with the W-pattern of the tooth incomplete. Hipposideros wollastoni, despite the divergence of its posterior noseleaf, is cranially very similar to H. muscinus. However, its rostral eminences are rather more swollen, its anteorbital foramen is rounded, closed by a narrow bar, the sphenoidal bridge is less constricted, the sphenoidal depression not traversed by a median ridge and the cochleae more expanded, their width equal to six times their distance apart. The anterior upper premolar (pm 2 ) is small, extruded and the posterior ridge of the third upper molar is less reduced, one third or a little more the length of the anterior ridge, with the W-pattern of the tooth less incomplete. Hipposideros semoni and H . stenotis are more closely related to each other than to H. muscinus or H. wollastoni, although their affinities as demonstrated by the nose- leaves clearly lie nearer to H . muscinus than to H. wollastoni. In both H. semoni and H. stenotis the tubercles on the intermediate and posterior noseleaves are developed into club-like processes similar to those of H. cyclops. Cranially, both have a frontal depression and the anteorbital foramen is small and rounded, closed in H. semoni by a moderate bar and in H. stenotis by a narrow bar. In both the sphenoidal bridge is slightly more constricted than in H. muscinus or H. wollastoni. Neither H. semoni nor H. stenotis has a sphenoidal depression and in both the cochleae are greatly expanded, their width equal to eight times their distance apart. In H. semoni the anterior upper premolar (pm 2 ) is small and extruded from the tooth- row : it is absent in H. stenotis while in both the posterior ridge of the third upper molar is not greatly reduced and is equal nearly in length to the anterior ridge, the W-pattern of the tooth nearly complete. In view of these considerations I am unable to agree with Tate (1941 : 379) that there can be no doubt that H. muscinus, H. semoni and H . stenotis are conspecific : although the available material A REVISION OF HIPPOSIDEROS 75 is limited, it is evident that H. muscinus is widely separated from H. semoni and H. stenotis, although these by comparison are closely related, as Tate (p. 389) recognized. Tate (1941 : 378, 379) has provided an account of the muscinus group as he understood it, evidently based on series from various localities in Papua which he considered to represent H. muscinus (pp. 379, 386, 392). Furthermore, he notes (p. 386) that " there is no doubt that muscinus is very closely related to semoni and stenotis " and after reviewing some of the features of the ears and noseleaf of H. muscinus as given by Thomas & Doria in the original description states that the skull, which they did not describe, as represented by the syntype in the British Museum (Natural History) " shows the characteristics of semoni and agrees with my topotypical series of muscinus : the parallel toothrows, the greatly enlarged rostrum, the closely approximated cochleae, etc.". A comparison of his detailed account (pp. 378, 379) with the syntype of H. muscinus in the British Museum (Natural History) suggested that the specimens that Tate used as the basis of his study and that he referred to H. muscinus in fact did not represent that species but should be referred to H. semoni. Through the courtesy of the authorities of the American Museum of Natural History I have been able to examine a part of this series of specimens, including three of those from the Fly River, Papua, thought topotypical of H. muscinus by Tate. Comparison of this selection of the specimens studied by Tate with the syntype of H. muscinus and with examples of H. semoni in the collections of the British Museum (Natural History) shows that without doubt they represent the latter species. A specimen preserved in alcohol, A.M.N.H. 108684, a ma l e from the bank of the Fly River, opposite Sturt Island, Papua, has the noseleaf as described by Tate, its intermediate and posterior leaves with club-shaped median protuberances. These can be readily discerned in four other dry skins from Papua, A.M.N.H. 105341, a female from the same locality ; A.M.N.H. 105057, a male from five miles below Palmer Junction, Upper Fly River ; A.M.N.H. 108500, a young female from Baruari, Astrolabe Range, Central Division and A.M.N.H. 108504, a male from Sogeri, Central Division. These protuberances in the syntype of H. muscinus are not developed into club-shaped structures as in H. semoni and H . stenotis but take the form of tubercles. The skulls of the four dry specimens are available for study and agree exactly with the account by Tate. All have a small frontal depression, absent in H. muscinus but present in H. semoni. The anteorbital foramen is small, rounded and separated from the orbit by a moderate bar as in H. semoni : in H. muscinus this aperture is not rounded but is elongate and is separated from the orbit by a narrow bar. There is a low median sphenoidal ridge and no sphenoidal depression as in H. semoni : a shallow sphenoidal depression is present in H. muscinus. The cochleae are very large, their width equal to six or eight times their distance apart as in H . semoni : those of H . muscinus are less expanded with their width equal to four or five times their distance apart. The angular process of the mandible is elongate, connected to the articular process by a thin web of bone. This condition is very evident in H, semoni, which in this respect exactly resembles the series 7 6 J. E. HILL studied by Tate as represented by these specimens : it is much less pronounced in H. muscinus. The anterior upper premolar (pm 2 ) is minute, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact : this is the condition exhibited by H. semoni and not by H . muscinus in which the anterior upper premolar is larger and separates the canine and the second upper premolar. The posterior ridge of the third upper molar is nearly as long as the anterior ridge, the W-pattern of the tooth almost complete, as in H. semoni and H. stenotis : in H. muscinus the posterior ridge is less than one third the length of the anterior ridge and the W-pattern of the tooth is incomplete. These specimens must be referred to H. semoni and agree exactly with A.M.N.H. 154710, a female from Upper Nesbit River, Cape York, Queensland (labelled Hipposideros muscinus semoni), with B.M. 99.10.24.1, a female, also from Queensland and with B.M. 50.1154-1155, a male and female from the Buntibasa district, Kratke Mountains, Northeast New Guinea. The collections of the British Museum (Natural History) also contain a specimen from Avera, Aroa River, Papua, B.M. 4.4.11.4, with only the rostrum and mandible remaining of the skull, which must be referred to this species. Tate, who thus studied H. semoni in the mistaken belief that he had before him specimens of H. muscinus, thus came to the logical conclusion that H. muscinus and H. semoni were conspecific. Furthermore, he wrote (1941 : 379) " Dahl and Collett, both of whom wrote in 1897, evidently had North Australian material, which they alluded to as muscinus". However, at that time neither H. semoni nor H. stenotis had been described, and Tate was apparently unaware that one of the specimens collected by Dahl at the Mary River, Northern Territory, Australia and referred by Dahl (1897 : 191) and Collett (1897 : 320) to H. muscinus was received by the British Museum (Natural History) in exchange with the then Christiania Museum in 1897 and subsequently became the type specimen of H. stenotis. For the present, H. semoni and H. stenotis are treated as distinct species, although their differences are those of degree rather than of structure. Hipposideros stenotis, however, is a much smaller bat than H. semoni, and has the protuberances of the intermediate leaf and especially the posterior leaf less greatly developed. The frontal depression is more greatly excavated than in H. semoni and its rostral eminences are less inflated, while its anteorbital foramen is larger in relation to the size of its skull. The teeth of H . stenotis are smaller and less massive than those of H. semoni. The anterior upper premolar (pm 2 ) in H. stenotis is absent : in H. semoni this tooth is absent in the type (Thomas (i9i3b : 206)) but is present in both sides of the jaw of all the specimens of H. semoni examined during the preparation of these notes. Relationships within the group are summarized in Figure 19. The muscinus group evidently originates from within the bicolor group or from the bicolor-speoris stem and some of its features are incipiently displayed by H. papua, a species remotely allied to H. bicolor and its allies. The muscinus group is isolated within Hipposideros by the highly modified characters of its ears, noseleaves and auditory region, which are specialized to an extent otherwise nowhere approached in this A REVISION OF HIPPOSIDEROS 77 large genus. Its extremely discontinuous distribution and the complexity of its characters suggest a relict group of remote origin, a view supported by the profound differences between its Ethiopian representatives and its remaining representatives in the Australasian region. voltattoni As muscinus but posterior leaf with transverse supplementary structure, no posterior tubercle: anteorbital foramen rounded, cochleae greatly expanded. semoni, stenotis As muscinus but foliar tubercles club-shaped: with a frontal depres- sion, anteorbital foramen rounded, no sphenoidal depression, cochleae greatly expanded, m 3 unreduced. cyclops, camerunensis Posterior lateral supple- mentary leaflet not extend- ing beneath anterior leaf, foliar tubercles prominent, club-shaped: skull elongate, anterior palatal foramina elongate, enclosed, no sphenoidal depression, cochleae not greatly ex- panded, m 1 reduced. Posterior lateral supplementary leaf- let extending beneath anterior leaf, foliar tubercles not club-shaped: skull shortened, no frontal depres- sion, anteorbital foramen elongate, anterior palatal foramina rounded, not enclosed, shallow sphenoidal depression, cochleae not greatly expanded, m 3 reduced. FIG. 19. Possible relationships in the Hipposideros muscinus group The species of the cyclops group may be keyed : Posterior lateral supplementary leaflet not extending anteriorly beneath the anterior leaf : premaxillae wide, enclosing the anterior palatal foramina, making a broad junction with the maxillae .......... 2 Posterior lateral supplementary leaflet extending anteriorly beneath the anterior leaf : premaxillae narrow, not enclosing the anterior palatal foramina, making a narrow, spatulate junction with the maxillae ....... 3 Smaller, length of forearm less than 74-0 mm. : anteorbital foramen relatively large cyclops (p. 78) Larger, length of forearm exceeding 74-0 mm. : anteorbital foramen relatively small camerunensis (p. 80) Median process of intermediate leaf not greatly developed or club-shaped : frontal depression absent, shallow sphenoidal depression present, posterior ridge of m 8 reduced, W-pattern of tooth incomplete 4 78 J. E. HILL Median process of intermediate leaf well developed, club-shaped : frontal depression present, sphenoidal depression absent, posterior ridge of m 3 little reduced, W- pattern of tooth almost complete ......... 5 4 Posterior leaf with a median process and without a transverse supplementary structure developed from its posterior face : width of cochleae four to five times their distance apart ......... muscinus (p. 81) Posterior leaf without a median process and with a transverse supplementary structure developed from its posterior face : width of cochleae six or more times their distance apart ......... wollastoni (p. 83) 5 Median process of posterior leaf well developed : frontal depression shallow, rostral eminences greatly inflated, anteorbital foramen relatively small . . semoni (p. 84) Median process of posterior leaf not greatly developed : frontal depression deep, rostral eminences moderately inflated, anteorbital foramen relatively large . stenotis (p. 86) Hipposideros cyclops (Temminck) (Figure 20) Phyllorrhina cyclops Temminck, 1853 : 75. River Boutry, Ghana. Rhinolophus micaceus De Winton, 1897 : 524. Como River, 75 miles from Gaboon. Hipposideros langi J. A. Allen, 1917 : 434. Avakubi, eastern Congo. The ears are exceptionally long and narrow, their posterior margins with a con- cavity behind the tip and without antitragal modification. The noseleaf is large, with two lateral supplementary leaflets. The anterior leaflet is short but wide : the posterior leaflet extends upwards to join the base of the posterior leaf but does not extend anteriorly beneath the anterior leaf. The anterior leaf has no median emargination, the internarial septum is not greatly inflated and the narial lappets are slightly expanded. The intermediate part of the leaf is flat but is specialized by the development of a median club-like process. The posterior leaf is moderately high, its upper edge more or less semicircular, thickened but not lobulated, and is supported by three septa enclosing deep pockets. It is specialized by the develop- ment of a median club-like process from the upper part of its anterior face. A frontal sac is present in both sexes. The skull is large and elongate, with an elongate braincase and low sagittal crest. The interorbital region is rather constricted but not greatly shortened, and the supraorbital ridges are well defined. There is a moderate frontal depression and the rostral eminences are moderately inflated, separated by a shallow groove. The zygomata are comparatively massive, with a low jugal projection. The anteorbital foramen is large and rounded, closed by a moderate bar of bone. The premaxillae are broad, together almost filling the anterior palatal emargination, and wholly enclose the anterior palatal foramina. They are very wide posteriorly and make a broad, shallowly V-shaped junction with the maxillae. The palate is short, with a square palation. The mesopterygoid fossa is moderate, with the vomer projecting very slightly, and the pterygoids are long, with the sphenoidal bridge only very slightly constricted, almost completely concealing narrow, elongate lateral apertures. There is a shallow sphenoidal depression and the cochleae are large, their width equal to four times their distance apart. The upper incisors are A REVISION OF HIPPOSIDEROS FIG. 20. Hipposideros cyclops $ (B.M. 30.11.11.151) (x3) very weakly bilobed, and the upper canines lack cusps but have prominent cingula. The anterior upper premolar (pm 2 ) is not greatly reduced but is extruded from the toothrow with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is one third the length of the anterior ridge. The crown area of the outer lower incisors is equal to or slightly exceeds the crown area of the inner pair and the anterior lower premolar (pm 2 ) is much reduced, one quarter the length and one half the height of the second lower premolar (pm 4 ). Hayman (1935) gives an account of H. cyclops. DISTRIBUTION : Portuguese Guinea (Veiga Ferreira (1949 : 195)) ; Sierra Leone ; Liberia ; Ghana ; Nigeria ; Cameroons ; Congo ; Kenya ; Uganda. So J. E. HILL Hipposideros camerunensis Eisentraut Hipposideros camerunensis Eisentraut, 1956 : 526. Buea, Cameroons. The ears and noseleaf of H. camerunensis closely resemble those of H. cydops but the ears of H. camerunensis are rather larger and the noseleaf is broader than in this species. A frontal sac is present in both sexes. The skull is similar to that of H. cyclops and is long and rather elongate, with a broadened braincase and low sagittal crest. The interorbital region is sharply constricted with well-defined supraorbital ridges. There is a moderate frontal depression and the rostrum is expanded and broad, with moderately inflated rostral eminences separated by a shallow groove. The zygomata are massive, with a moderate jugal projection. The anteorbital foramen is comparatively small, closed by a narrow bar of bone. The premaxillae are broad, like those of H. cyclops, together almost filling the anterior palatal emargination, and wholly enclose the elongate anterior palatal foramina. As in H. cyclops, they are wide posteriorly, and make a broad, shallowly V-shaped junction with the maxillae. The palate is short and the palation almost square. The mesopterygoid fossa is moderate, the vomer projecting very slightly, and the pterygoids are long with the sphenoidal bridge only slightly constricted and almost concealing elongate lateral apertures. There is a shallow sphenoidal depression and the cochleae are large as in H. cyclops, their width equal to four times their distance apart. The dentition almost exactly resembles that of H. cyclops. The upper incisors are weakly bilobed, the upper canines lack cusps and the anterior upper premolar (pm 2 ) is not greatly reduced, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is one third the length of the anterior ridge. The crown area of the outer lower incisors slightly exceeds the crown area of the inner pair and the anterior lower premolar (pm 2 ) is one quarter the length and one half or slightly less the height of the second lower premolar (pm 4 ). Hipposideros camerunensis is closely related to H. cyclops and differs from this species in its larger size, broader skull with broader braincase and rostrum and H. C. CYCLOPS H. CAMERUNENSIS 56 56 6O 62 64 66 66 7O 72 74 76 FIG. 21. Length of forearm in Hipposideros cyclops and H. camerunensis A REVISION OF HIPPOSIDEROS 81 smaller anteorbital foramen. The two species are the most primitive of the cyclops group, with their skulls elongate and not markedly shortened as in H. muscinus and its allies. Nevertheless, they display the specializations of the group to a considerable degree, notably in the elongation of their ears, the development of foliar prominences and the enlargement of their cochleae. Eisentraut (1963 : 87) discusses the status of H. camemnensis and records further specimens from Buea, Cameroons. DISTRIBUTION : Cameroons ; Congo (first record : specimen from Shabunda, eastern Congo, in collection of British Museum (Natural History)). Hipposideros muscinus (Thomas & Doria) (Figure 22) Phyllorrhina muscina Thomas & Doria, 1886 : 201, i fig. Fly River, Papua. The ears are very long and comparatively narrow, although not excessively so, with their posterior margins slightly concave behind the tip. They lack any anti- tragal modification and are largely naked. The noseleaf is large, with two lateral supplementary leaflets. The anterior leaflet is short and broad, projecting widely beyond the lateral margin of the anterior leaf. The posterior leaflet is broad and extends posteriorly to join the posterior leaf and anteriorly beneath the anterior leaf on to the upper lip. The anterior leaf has no median emargination and the internarial septum is slightly swollen, especially anteriorly. The narial lappets are well developed, but the nostrils are only slightly pocketed. The intermediate part of the leaf is very slightly inflated with a small median tubercle-like process. The posterior leaf is high, its upper edge flattened, thickened and with a small median FIG. 22. Hipposideros muscinus $ (Syntype B.M, 86.11.3.10) (x3) ZOO 11, 1. F 82 J. E. HILL projecting tubercle. The posterior leaf is supported by three septa enclosing shallow pockets. The skull is small, short and broad, with a low sagittal crest. The interorbital region is short and sharply constricted, the supraorbital ridges well denned. There is no frontal depression and the rostral eminences are moderately inflated, separated by a shallow groove. The zygomata are slender, with a low jugal projection and the anteorbital foramen is small, elongate and closed by a narrow bar. The junction of the premaxillae with the palate is U-shaped : the palate is very short with a U-shaped palation. The mesopterygoid fossa is wide, the pterygoids long and wide and the sphenoidal bridge barely constricted, partially concealing elongate lateral apertures. There is a shallow sphenoidal depression bisected by a low median ridge. The cochleae are large, their width equal to four or five times their distance apart. The upper canines are slender, without cusps, and the anterior upper pre- molar (pm 2 ) is not excessively reduced and is only slightly out of alignment in the toothrow, separating the canine and the second upper premolar (pm 4 ) . The posterior ridge of the third upper molar is obsolescent, its length less than one quarter the length of the anterior ridge. The crown area of the outer lower incisors is slightly greater than the crown area of the inner pair and the anterior lower premolar (pm 2 ) is one third the length and height of the second lower premolar (pm 4 ). Although the only specimen available for study is the syntype, there is no reason to consider that H. muscinus is conspecific with either H. semoni or H. stenotis. Hipposideros muscinus has ears shorter and broader than in these species, and has the projecting processes of the intermediate and posterior noseleaves less developed. Cranially, it differs quite markedly from either, with more sharply defined supra- orbital ridges and a wider rostrum, the rostral eminences less inflated, the rostrum from its lateral aspect horizontal above, lacking the marked concavity over the interorbital region separating the rostral eminences and braincase in H. semoni and H. stenotis and lacking the frontal depression found in these species. In H. muscinus the anteorbital foramen is small and elongate, in contrast to the rounded foramina of H. semoni and H. stenotis : the sphenoidal bridge of H. muscinus is wider than in these species and a sphenoidal depression, absent in both, is present in H. muscinus. Its cochleae are less expanded than those of H. semoni or H. stenotis and the angular and articular processes of the mandible are not markedly connected by a thin web of bone as in these species. The third upper molar of H. muscinus has its posterior ridge greatly reduced and almost obsolete, contrary to the condition evident in H. semoni and H. stenotis, which have the posterior ridge of this tooth nearly equal in length to the anterior ridge and almost entire. These considerations indicate that while H. muscinus without doubt is allied to H. semoni and H. stenotis, it can by no means be considered conspecific with either of these species. It is less specialized than either H. semoni or H. stenotis, and cranially is similar to H. wollastoni. This species, however, differs from H. muscinus in the development of a transverse supplementary structure from the posterior face of the posterior leaf, in more inflated rostral eminences (similar in fact to those of H. semoni and H. stenotis} and in a greater degree of expansion of the cochleae. A REVISION OF HIPPOSIDEROS Hipposideros wollastoni Thomas : 205. Camp 3, Utakwa River, southwestern Hipposideros wollastoni Thomas, Netherlands New Guinea. The ears are long and comparatively narrow, pointed, their posterior margins slightly concave behind the tip. They lack antitragal modification and are largely naked. The noseleaf is large with two lateral supplementary leaflets exactly similar to those of H. muscinus. The anterior leaf resembles that of H. muscinus but the internarial septum is slightly more inflated and the nostrils are slightly pocketed. The intermediate leaf is not inflated but has a large projecting median eminence. The upper edge of the intermediate part of the leaf is elevated laterally, these elevations forming a base for and largely concealing the lateral supporting septa of the posterior leaf. The posterior leaf has its upper edge semicircular, unthickened and lacking a median projection, and is supported by three septa enclosing shallow pockets. A transverse supplementary structure with a serrate upper edge is developed from its posterior face, and is equal in height almost to the posterior leaf. This structure is joined longitudinally to the posterior leaf by two external and two intermediate septa, enclosing three small pockets. A wart-like eminence arises on each side of the muzzle just behind the posterior leaf. There is a small depression but no frontal sac in the female type specimen. FIG. 23. Hipposideros wollastoni $ (Type B.M. 13.6.18.4) (x3) The skull in general outline is very like that of H. muscinus, with a moderate sagittal crest and a constricted interorbital region. The supraorbital ridges are evident but not sharply defined, especially behind the rostral eminences. There is 84 J. E. HILL no frontal depression and the rostral eminences are considerably inflated, separated by a deep groove, and give the rostrum from its lateral aspect a marked elevation. The zygomata are slender, with a moderate jugal projection while the anteorbital foramen is small, rather rounded and closed by a narrow bar. The premaxillae are narrow, spatulate posteriorly, and make a narrow junction with the maxillae. They do not enclose the rounded anterior palatal foramina, the anterior walls of which are formed by the delicate anterior enclosing processes of the premaxillae. The palate is short and the palation U-shaped with a small post-palatal spicule. The mesopterygoid fossa is wide and the pterygoids are long, with an almost uncon- stricted sphenoidal bridge, partially concealing elongate lateral apertures. There is a small sphenoidal depression, lacking any median ridge. The cochleae are greatly expanded, their width equal to six times their distance apart. The angular and articular processes of the mandible are not markedly connected by a web of bone. The upper incisors are not bilobed and the upper canines lack cusps, while the anterior upper premolar (pm 2 ) is small and extruded, the canine and the second upper premolar (pm 4 ) almost in contact. The posterior ridge of the third upper molar is equal in length to one third of the length of the anterior ridge, the W-pattern of the tooth incomplete. The crown area of the outer lower incisors is slightly greater than that of the inner pair, and the anterior lower premolar (pm 2 ) is a little under one half the length and height of the second lower premolar (pm 4 ). Despite its specialized noseleaf , H. wollastoni is more closely related to H . muscinus than to H. semoni or H . stenotis. Its ears, which are more acutely pointed than those of H. muscinus but shorter and broader than those of H. semoni or H. stenotis, its more advanced noseleaf, broader, more inflated rostrum and more greatly expanded cochleae show H. wollastoni to be a more specialized bat than H. muscinus. Hipposideros semoni Matschie (Figure 24) Hipposideros semoni Matschie, 1903 : 774 (Heft 6 : 132). Cooktown, northern Queensland. The ears are very long and narrow, with an acute, narrow point. Their posterior margins are markedly concave just behind the tip, they lack any antitragal modifica- tion and are haired for one third of their length. The noseleaf is a highly developed structure covering much of the muzzle, and has two lateral supplementary leaflets. The anterior leaflet is short and broad, projecting widely beyond the lateral margin of the anterior leaf as in H. muscinus and H. wollastoni. As in these species, the posterior leaflet extends posteriorly to become continuous with the posterior leaf and anteriorly beneath the anterior leaf. The anterior leaf has no median emargination and the internarial septum is slightly inflated with the narial lappets well developed, the nostrils not pocketed. The intermediate part of the leaf is flat, its upper edge elevated laterally and bears a very well-developed club-shaped projection. The posterior leaf is high, supported by three septa, its upper edge semicircular, with a well-developed median club-shaped projection. A REVISION OF HIPPOSIDEROS FIG. 24. Hipposideros semoni $ (B.M. 99.10.24.1) (x3) The skull is short and broad, with a broad braincase and prominent sagittal crest. The interorbital region is moderately constricted with prominent supraorbital ridges which are less developed behind the rostral eminences. There is a shallow frontal depression and the rostral eminences are greatly inflated, to give the rostrum from its lateral aspect a much elevated appearance, deeply concave above the interorbital region. The zygomata are moderate, with a low jugal process and the anteorbital foramen is small and round, separated by a narrow bar. The premaxillae are much as in H . wollastoni, narrow, spatulate posteriorly and making a narrow junction with the maxillae. They do not enclose the rounded anterior palatal foramina. The palate is short, the palation square with a post-palatal spicule. The mesopterygoid fossa is wide while the pterygoids are long and the sphenoidal bridge moderately constricted, exposing elongate lateral apertures. There is no sphenoidal depression, but the sphenoidal region is traversed by a low longitudinal ridge. The cochleae are enormous, their width equal to eight times their distance apart, almost touching, separated by a very narrow basioccipital. The angular and articular processes of the mandible are united by a thin web of bone. The upper incisors are weakly bilobed, the upper canines without cusps and the anterior upper premolar (pm 2 ) very small, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact : Thomas (igi^b : 206) notes that this 86 J. E. HILL tooth is absent in the type specimen. The posterior ridge of the third upper molar is not greatly reduced, its length equal to two thirds the length of the anterior ridge and with the W-pattern of the tooth almost complete. The crown area of the outer lower incisors is slightly greater than that of the inner pair and the anterior lower premolar (pm 2 ) is less than one quarter the length and height of the second lower premolar (pm 4 ). Hipposideros semoni is clearly quite distinct from H. muscinus, differing from this species in a number of salient features, among which the greater development of the ears and noseleaf, rostrum and cochleae are the most important. It is patently one of the most, if not the most specialized species of the cyclops group and together with the closely related H. stenotis represents the culmination of a line of specializa- tion of which H. papua is perhaps an offshoot and H. muscinus a later development. DISTRIBUTION : Northern Australia ; Papua ; Northeast New Guinea. Hipposideros stenotis Thomas (Figure 25) Hipposideros stenotis Thomas, igi3b : 206. Mary River, Northern Territory, Australia. Externally, H. stenotis is closely similar to H. semoni but is considerably smaller. The ears are exactly as described for that species, while the noseleaf almost exactly resembles that of H. semoni except that the median projection of the posterior leaf is less prominent. The skull is short and broad, with moderate sagittal crest and supraorbital ridges like those of H. semoni. There is a deep frontal depression and the rostral eminences are moderately inflated, sufficiently that the rostrum from its lateral aspect appears elevated. The zygomata are slender, with a low jugal projection and the anteorbital foramen is rounded, closed by a narrow bar. The premaxillae are similar to those of H . wollastoni and H. semoni, spatulate posteriorly, making a narrow junction with the maxillae and not enclosing the rounded anterior palatal foramina. The palate is short and the palation square, with a small post- palatal spicule. The mesopterygoid fossa is moderate, the pterygoids long and the sphenoidal bridge moderately constricted exposing wide, elongated lateral apertures. There is no sphenoidal depression and the sphenoidal region is traversed by a low longitudinal ridge. The cochleae are enormous, their width equal to eight times their distance apart, almost touching, separated by a very narrow basioccipital. The angular and articular processes of the mandible are connected by a web of bone. The upper incisors are weak and not bilobed : the type specimen is unusual in that a second right hand upper incisor is present. It is a peg-like tooth, longer but more slender than the first upper incisor. The upper canines lack cusps and the anterior upper premolar (pm 2 ) is absent, with the canine and the second upper premolar (pm 4 ) in contact. The posterior ridge of the third upper molar is not greatly reduced and is two thirds the length of the anterior ridge, with the W-pattern of the tooth almost complete. The crown area of the outer lower incisors is a little greater than that of the inner pair and the anterior lower premolar (pm 2 ) is much reduced, one quarter the length and height of the second lower premolar (pm 4 ) . A REVISION OF HIPPOSIDEROS FIG. 25. Hipposideros stenotis $ (Type B.M. 97.4.12.7) (x3) Although the close structural similarity of H. stenotis to H. semoni suggests that the two may be conspecific, there is considerable disparity in size and H. stenotis has the protuberances of its intermediate and posterior noseleaves less developed. Cranially, its frontal depression is deeper than that of H. semoni and its rostral eminences less inflated : its anteorbital foramen is larger and the molars much less massive. Only the type specimen of H. stenotis is available for study and for the present it is retained as a distinct species. Tate (1941 : 389) records a series of H. stenotis from Papua in the collections of the American Museum of Natural History. Despite careful search by the authorities of that Museum, no trace of this series has been found. HIPPOSIDEROS PRATTI group This group contains only two species, H. pratti and H. lylei, both Asiatic in distribution. The ears are large and broad, bluntly pointed with their posterior margins slightly concave behind the tip. They lack antitragal modification and are haired for one third of their length. The noseleaf has two lateral supplementary leaflets and the anterior leaf a median emargination. The intermediate part of the leaf is expanded, with a median eminence and the posterior leaf is high in the centre, triangular, supported by a prominent median ridge flanked by two much weaker ridges. The group is noteworthy for the development of transverse fleshy lobate prominences on each side of the opening of the frontal sac to form a structure resembling a supplementary posterior noseleaf traversing the muzzle behind the true posterior leaf. The degree of development of this structure varies considerably, but is always at its greatest in male specimens. 88 J. E. HILL 42 44 46 48 H. MUSCINUS H. WOLLASTONI H. SEMONI H. STENOTIS 46 48 42 44 FIG. 26. Length of forearm in Hipposideros muscinus, H. wollastoni, H. semoni and H. stenotis The skull is of moderate to large size, with a moderate sagittal crest. The inter- orbital region is short, with well-defined supraorbital ridges and there is a shallow frontal depression. The rostrum is low and broad with moderately inflated rostral eminences. The zygomata are slender and have a moderate jugal projection, while the anteorbital foramen is rounded and closed by a narrow bar. The pre- maxillae make a fan-shaped junction with the maxillae and totally enclose the rounded anterior palatal foramina. The palate is short and wide, the pterygoids com- paratively short and the sphenoidal bridge not markedly constricted. There is a well-defined sphenoidal depression and the cochleae are small, their width equal to or less than their distance apart. The outer lobe of the upper incisors is only slightly smaller than the inner lobe, and the anterior upper premolar (pm 2 ) is reduced, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is almost obsolete. The crown area of the outer lower incisors is greater than that of the inner pair, and the anterior lower premolar is one half the length and height of the second lower premolar (pm 4 ). Although Ellerman & Morrison-Scott (1951 : 129) considered H. pratti and H. lylei to be conspecific, the two evidently are widely separated by differences in the structure of the noseleaf and cranial architecture which in no way support the view that they are conspecific. Their taxonomic position within the genus is not easy to determine. The presence of two lateral supplementary leaflets and their short, broadened skulls recalls the more specialized species of the bicolor group while the development of a supplementary lobate structure behind the posterior leaf in both forms and the presence of an abrupt discontinuity between the roofs of the meso- pterygoid and narial canals in H. pratti suggest an affinity with the armiger group. A REVISION OF HIPPOSIDEROS 89 The depressed rostrum and the presence of a frontal depression in the pratti group, however, render close alliance with the armiger group unlikely. In the present work, the pratti group is regarded as linking bats of the bicolor group to the more specialized bats of the armiger group, displaying no affinities towards either the speoris or diadema groups. The two species of the pratti group may be keyed : i Lateral margins of anterior leaf not continuous with those of posterior leaf : rostrum lacking lateral pits, an abrupt step-like discontinuity between roofs of narial and mesopterygoid canals, vomer projecting posteriorly beyond palate . pratti (p. 89) Lateral margins of anterior and posterior leaves continuous : rostrum with lateral pits, roofs of narial and mesopterygoid canals merging smoothly, vomer not projecting posteriorly beyond palate ...... lylei (p. 90) Hipposideros pratti Thomas Hipposiderus (sic) pratti Thomas, 1891 : 527. Kiatingfu, Szechuan, China. The ears and noseleaf agree closely with the group description. The anterior leaf has a single median emargination and its lateral margins are not continuous with those of the posterior leaf. The internarial septum is uninflated and the narial lappets moderately developed. The intermediate part of the leaf is inflated with a prominent median eminence. Its upper edge is elevated laterally to form a large transverse structure across the noseleaf anterior to the lower part of the posterior leaf. The posterior leaf is more or less triangular in outline and is supported by a well-defined median septum, flanked laterally by weaker ridges, barely enclosing small cells. Two large lobate projections are developed transversely behind the posterior leaf, one each side of the opening of the frontal sac, to form an enlarged supplementary transverse structure, the projections separated by a deep median notch at the aperture of the sac. A small tuft of bristle-like hairs projects through this notch. This transverse structure is more greatly developed in old male animals than in young male or female examples. A frontal sac is present in both sexes. The skull is large and broad, with a well-developed sagittal crest. The interorbital region is markedly constricted and there are prominent supraorbital ridges. There is a shallow frontal depression and the rostrum is broad, its upper surface below the level of that of the braincase, bluntly angular, with a pentagonal outline and with the rostral eminences not greatly inflated. The zygomata and the anteorbital foramen are as described for the group as are the premaxillae, which make a U-shaped junction with the maxillae. The palation is U-shaped and the mesopterygoid fossa wide with the posterior part of the narial canal deeply excavated, an abrupt step-like discontinuity separating its roof from the roof of the mesopterygoid fossa. The vomer is thin and blade-like, projecting posteriorly beyond the edge of the palate. The pterygoids are moderate and the sphenoidal bridge constricted, exposing wide, elongate apertures. The sphenoidal depression, cochleae and dentition are as described for the group. Osgood (1932 : 222) gives detailed illustrated notes on specimens from Tonkin. go J. E. HILL Pohle (1943 : 323) considers a specimen in the Berlin Museum to be the holotype of Phyllorhina swinhoii Peters (in Swinhoe, Proc. zool. Soc. Lond. 1870 (1871) : 616) (more correctly swinhoei) and has suggested that H. pratti is synonymous with the species described by Peters. However, Peters (i87ia : 317) considered H. swinhoei to be a synonym of H. armiger (Hodgson) and was followed in this opinion by Dobson (1878 : 135) and Andersen (i9o6a : 37) while Tate (1941 : 371, 389) who had available a photograph " of the type skull ", a topotype from Amoy and other specimens from China, considered H. swinhoei to be a subspecies of H. armiger. There are three specimens from the original series collected by Swinhoe in the collections of the British Museum (Natural History). All have three lateral supplementary leaflets and cannot therefore be regarded as H. pratti. In the present work H. swinhoei (Peters) is regarded as a synonym of H. armiger armiger (Hodgson). DISTRIBUTION : Southwestern China ; Indochina : Tonkin. Hipposideros lylei Thomas Hipposideros lylei Thomas, igisa : 88. Chiendao Cave, 50 miles north of Chiengmai, Siam, 350 metres. The ears are broad and triangular, slightly less acutely pointed than in H . pratti, their posterior margins shallowly concave behind the tip and slightly thickened at the antitragal lobe. The noseleaf in its essentials resembles that of H. pratti, with two small lateral supplementary leaflets, but is slightly more specialized. The anterior leaf projects forward over the upper lip and has a deep median emargi- nation, the edge of the leaf lateral to this emargination shallowly emarginated on each side to form two small projecting lappets. Its lateral margins are continuous with those of the posterior leaf but are separated from them at the level of the intermediate part of the leaf by a shallow notch on each side. The internarial septum is not greatly inflated and the narial lappets are well developed. The inter- mediate part of the leaf is similar to that of H. pratti and has a well-developed median eminence. Its lateral margins are embellished on each side with a small outwardly projecting lappet at the level of the notches in the lateral margins of the anterior and posterior leaves. The posterior leaf is triangular in outline and is supported by a median septum and two weaker lateral ridges. There is a greatly developed trans- verse supplementary structure behind the posterior leaf as described for H . pratti. The skull is of moderate to large size, with a low sagittal crest. The interorbital region is markedly constricted and the interorbital ridges are sharply denned. There is a well-developed frontal depression and the rostrum is broad, low as in H. pratti, with the rostral eminences slightly inflated. There is a lateral depression on each side of the rostrum immediately above the anteorbital foramen : these depressions are a little deeper than the frontal depression. The zygomata are moderate with a moderate jugal process and the anteorbital foramen, premaxillae and palate are as described for the group, the premaxillae making a broad, U-shaped junction with the maxillae and the palation more acute, almost V-shaped. The A REVISION OF HIPPOSIDEROS 91 mesopterygoid fossa is wide, and the roof of the narial canal, although discontinuous with the roof of the mesopterygoid canal, merges into it smoothly, without an abrupt, step-like discontinuity. The vomer does not project posteriorly beyond the edge of the palate. The pterygoids are short and wide, the sphenoidal bridge less con- stricted than in H. pratti, partially concealing elongate lateral apertures. The sphenoidal depression, cochleae and dentition are as described for the group. Hipposideros lylei differs from H. pratti in the deeper median emargination of its anterior leaf, flanked by two shallow lateral emarginations absent in H . pratti : in the continuity of the lateral margins of the anterior and posterior noseleaves, which are separated only by lateral marginal notches and in the presence of small lateral lappets at the margins of the intermediate part of the leaf. It has a deeper frontal depression : lateral rostral pits, present in H. lylei, are absent in H. pratti and the zygomata of H. lylei are more massive than those of H. pratti. In H. lylei the roofs of the narial and mesopterygoid canals merge smoothly, without the abrupt, step-like discontinuity characteristic of H. pratti and the sphenoidal bridge of H . lylei is wider than that of H. pratti, partially concealing the lateral apertures. DISTRIBUTION : Burma ; North Shan States ; Siam ; Federation of Malaya. 74 76 H. PRATTI H. LYLEI 76 78 8O 82 84 86 88 FIG. 27. Length of forearm in Hipposideros pratti and H. lylei HIPPOSIDEROS ARMIGER group The ears are large, broad and acutely pointed, their posterior margins concave behind the tip. They are slightly thickened at the antitragal lobe and are haired for one third of their length. The noseleaf is broad, with four supplementary leaflets, the fourth small, sometimes rudimentary. The anterior leaf has no distinct median notch, the internarial septum is not inflated and the narial lappets are not greatly 92 J. E. HILL developed. The intermediate part of the leaf is slightly inflated with a prominent median eminence. The posterior leaf is high and narrow, not as wide as the anterior leaf, its upper edge flattened, thickened, becoming trilobate and is supported by a prominent median septum and two less evident lateral septa. Prominent fleshy elevations arise behind the posterior leaf on each side above the eyes to form a transverse supplementary structure, found at its greatest development in old male specimens, less developed in young male and female examples. There is a frontal sac in male specimens : in female examples it is small or is represented by a depression. The skull is of moderate to large size with well-developed sagittal crest, short, constricted interorbital region and sharply defined supraorbital ridges. There is no frontal depression and the rostral eminences are uninflated. The naso-frontal region is decidedly pentagonal in outline and the rostrum is elevated posteriorly, in profile forming a nearly horizontal surface on a level with the upper surface of the braincase. The zygomata are moderate to strong, with a moderate to well- developed jugal projection. The anteorbital foramen is large and rounded, closed by a very narrow bar. The premaxillae make a fan-shaped j unction with the maxillae and do not enclose the elongate anterior palatal foramina. The palation is U-shaped and the mesopterygoid fossa wide, with the roof of the narial canal separated from the roof of the mesopterygoid fossa by a sharp, step-like discontinuity, the vomer projecting beyond the edge of the palate. The pterygoid wings are expanded and the sphenoidal bridge is wide, almost concealing elongate lateral apertures. There is a moderate sphenoidal depression and the cochleae are small, their width less than their distance apart. The upper incisors are bilobed with the outer lobe approximately equal to the inner lobe while the upper canines have a low anterior cusp. The anterior upper premolar (pm 2 ) is small, extruded from the toothrow with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is obsolescent. The crown area of the outer lower incisors is greater than that of the inner pair and the anterior lower premolar (pm 2 ) is one third to one half the length and height of the second lower premolar (pm 4 ). The armiger group contains two very closely related species, H. armiger and H . turpis, essentially identical in their structural characteristics and separable only by the much smaller size of H. turpis as compared with H . armiger, a situation not unparalleled elsewhere in Hipposideros. The group is a specialized one not closely related to the other group of large bats of the genus Hipposideros, the diadema group, with which it was compared by Andersen (i9o6a : 35). Its affinities lie with the pratti group with which the armiger group shares a number of specializations in the development of lobate transverse structures behind the posterior noseleaf, broad premaxillae joined to the maxillae by a wide, fan-shaped junction, a sharp discontinuity between the roofs of the narial and mesopterygoid canals (least developed in H. lylei, more so in H. pratti, H. armiger and H. turpis} and small cochleae. The armiger group, although having the transverse lobes behind the noseleaf less developed than in the pratti group, seems nevertheless slightly more specialized with a shorter, less elongate skull, no frontal depression, a high rostrum A REVISION OF HIPPOSIDEROS 93 level with the upper surface of the braincase and with the canine-bearing parts of the maxillae not prolonged anteriorly. The armiger group seems to have developed from the speoris-diadema stem, to which it is linked by the pratti group. Hipposideros armiger Size large, otherwise characters as in the group diagnosis. DISTRIBUTION : Northern India to Formosa and the Malay Peninsula. Hipposideros armiger armiger (Hodgson) Rhinolophus armiger Hodgson, 1835 : 699. Nepal. Phyllorhina swinhoii Peters, 18710 : 616. Amoy, Fukien. Hipposideros armiger debilis Andersen, igo6a : 37. Province Wellesley, Federation of Malaya. The status of Phyllorhina swinhoii Peters has been discussed in connection with H. pratti. The diagnosis of H. a. debilis Andersen rests entirely on a slightly narrower interorbital region and shorter mandibular toothrow in Malayan specimens when compared with H. a. armiger. Long series in the collections of the British Museum (Natural History) do not support this contention. DISTRIBUTION : Northern India ; Nepal ; Assam ; Burma ; southern China ; Hong Kong (Romer (1960 : 2)) ; Indochina : Tonkin (Osgood (1932 : 222)), Bourret (ig42b : n)) ; Federation of Malaya ; Langkawi Island. Hipposideros armiger terasensis Kishida Hipposideros armiger terasensis Kishida, 1924 : 42. Formosa. According to Tate (1941 : 390) H. a. terasensis is likely to prove indistinguishable from H. a. armiger. H. A. ARMIGER FIG. 28. Length of forearm in Hipposideros armiger 94 J. E. HILL Hipposideros armiger tranninhensis Bourret Hipposideros tranninhensis Bourret, iQ42a : 20. Plaine des Jarres, Tran-Ninh, Indochina. From the description (repeated in part by Bourret (19425 : 12)) this form appears to be a subspecies of H. armiger, distinguished from H. a. armiger by its more trilobate posterior noseleaf. Hipposideros turpis The characters of this species are exactly as in H. armiger but it is, however, of much smaller size. The two species of the armiger group provide a close parallel to the situation found in some other groups of Hipposideros whereby two closely similar species differing chiefly in size are found to be sympatric. Such is the case with H. galeritus and H. breviceps ; H. caffer and H. beatus ; H. cyclops and H. camerunensis ; H. lankadiva and H. schistaceus and also H. diadema and H. dinops. In all of these, however, there is a small degree of structural differentiation between each pair of species : H. armiger and H. turpis are exceptional in that they differ only in size. Hipposideros turpis turpis Bangs Hipposideros turpis Bangs, 1901 : 561. Ishigaki, South Liukiu Islands. Hipposideros turpis pendleburyi Chasen Hipposideros pendleburyi Chasen, 1936 : 133. Near the foot of Khao Ram, Nakon Sri Tamarat, Peninsular Siam. H.T. PENDLEBURYI 62 64 66 68 7O 72 74 76 78 SO FIG. 29. Length of forearm in Hipposideros turpis HIPPOSIDEROS SPEORIS group The ears are large and comparatively broad, triangular in outline, their posterior margins concave behind the tip. Either there is a small process at the antitragal lobe or the membrane of the ear is thickened at this point. The noseleaf is simple, with three lateral supplementary leaflets. The skull is of medium size, comparatively short and with a moderate sagittal crest. The interorbital region is short and constricted and the supraorbital ridges are well developed. There is no definite frontal depression and the rostrum is low, with moderately inflated rostral eminences. A REVISION OF HIPPOSIDEROS 95 The zygomata are slender with a moderate jugal projection and the anteorbital foramen is rounded and closed by a narrow bar. The premaxillae make a wedge- FIG. 30. Possible relationships in the Hipposideros speoris group 96 J. E. HILL shaped or slightly U-shaped junction with the maxillae and partially or wholly enclose the anterior palatal foramina. The palate is short and wide with a V-shaped or U-shaped palation. The pterygoid wings are not greatly developed and the sphenoidal bridge is not widened, exposing elongate lateral apertures. There is a well-developed sphenoidal depression and the width of the cochleae is equal to their distance apart. The outer lobe of the upper incisors is obsolescent or absent and the upper canines have no definite cusps, although barely defined low anterior and posterior cusps are sometimes present. The anterior upper premolar (pm 2 ) is small and extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is obsolete or nearly obsolete. The outer lower incisors are greater in crown area than the inner pair and the anterior lower premolar (pm 2 ) is one half to two thirds the length and height of the second lower premolar (pm 4 ) . As presently defined, the speoris group includes three species, H. abac, H. larvatus and H. speoris. It is distributed in both the Ethiopian and Asiatic regions, with H. abac representing the group in Africa while H. larvatus and H. speoris are exclusively Asiatic in distribution. As might be expected, H. abae is rather sharply divided from the Asiatic species to much the same degree as the Ethiopian representa- tives of the bicolor group are from their Asiatic relatives, although its affinities lie nearer to the predominantly Indian species H. speoris than to the more Malaysian H. larvatus. The group is not one of extensive specialization in the manner of some of the members of the bicolor or cyclops groups and in fact the ears of its members retain some degree of antitragal modification and their nasal foliations are comparatively simple, apart from the development of a third lateral supplemen- tary leaflet. The structure of the ears and noseleaves in the speoris group indicates quite clearly its affinity to the bicolor group, and similar indications are evident in its cranial architecture, although specialization has proceeded further. In the speoris group the anterior palatal foramina in H. larvatus are wholly enclosed within the premaxillae and in the other species partially so ; the upper canine cusps are virtually obsolete and the crown area of the outer lower incisors is greater than the crown area of the inner pair. The speoris group apparently represents an offshoot of the line leading to the diadema group and has no close relation to the pratti and armiger groups, which represent a different line of modification characterized by the marked development of noseleaves, rostral region and mesopterygoidal region. The species of the speoris group may be keyed : 1 Anterior leaf without median emargination : prominent supraorbital ridges, rostrum flattened, vomer not projecting beyond palate, upper canines with shallow antero- internal groove and low posterior cusp . . . . . . . 2 Anterior leaf with definite median emargination: supraorbital ridges barely developed, rostrum rounded, vomer projecting beyond palate, upper canines not grooved and lacking a posterior cusp ....... larvatus (p. 98) 2 Posterior leaf with supporting septa : posterior palatal foramina undeveloped, palation U-shaped, sphenoidal bridge unconstricted .... speoris (p. 101) Posterior leaf lacking supporting septa : posterior palatal foramina prominent, palation nearly square, sphenoidal bridge constricted . . . abae (p. 97) A REVISION OF HIPPOSIDEROS 97 Hipposideros abae J. A. Allen Hipposideros abae ]. A. Allen, 1917 : 432. Aba, Uele district, Congo. The ears are very large, triangular and sharply pointed, with their posterior margins markedly concave behind the tip. There is no definite process at the antitragal lobe, but the ear membrane is thickened at this point. The anterior leaf has no median emargination and the internarial septum is not inflated, while the narial lappets are moderately developed. The intermediate part of the leaf is not greatly inflated and has a low median eminence flanked by weaker lateral eminences. The posterior leaf is thin and lacks supporting septa, with its upper edge semicircular and not lobate. The skull is slightly more elongate than the skull of H. speoris, with a low to moderate sagittal crest. The interorbital region is elongate and constricted with prominent supraorbital ridges. There is a very shallow frontal depression and the rostrum is narrow, with slightly inflated rostral eminences. The zygomata are slender anteriorly, with a high jugal process, and the anteorbital foramen is large and elongate, closed by a narrow bar. The pre- maxillae make a slightly wedge-shaped junction with the palate and partially or wholly enclose the elongate anterior palatal foramina. The posterior palatal fora- mina, small and insignificant in H. larvatus and H. speoris, are well developed and the palation is almost square. The mesopterygoid fossa is wide, with the vomer not projecting beyond the posterior edge of the palate and with expanded pterygoids. The sphenoidal bridge is narrow and sharply constricted, exposing large elongate lateral apertures. There is a well-developed sphenoidal depression and the cochleae are equal in width to their distance apart or are a little smaller. The upper incisors are large, weakly bilobed and situated near the outer corners of the premaxillae, while the upper canines have their anterior faces shallowly grooved and have a weak anterior cusp and a larger posterior cusp extending for one quarter of the height of the tooth. The anterior upper premolar (pm 2 ) is small, extruded, com- pressed between the canine and the second upper premolar (pm 4 ) with these teeth nearly in contact, while the posterior ridge of the third upper molar is one quarter or less the length of the anterior ridge. The crown area of the outer lower incisors is only slightly greater than that of the inner pair, and the anterior lower premolar (pm 2 ) is one half or slightly less the length and height of the second lower premolar (pm 4 ). Hippjsideros abae is divided sharply from H. larvatus and H. speoris by both external and cranial characters. It is a larger species, with the posterior leaf lacking supporting septa. Its interorbital region is more elongated than in the Asiatic species, its rostrum narrower with the anterior part of the maxillae bearing the canines more elongated, and it has a much larger anteorbital foramen. The pre- maxillae are broader and its prominent posterior palatal foramina, nearly square palation and constricted sphenoidal bridge are in direct contrast to the insignificant foramina, U-shaped palations and unconstricted sphenoidal bridges of H. larvatus and H. speoris. Such differences indicate comparatively remote divergence from the Asiatic members of the group and although in some respects such as its elongate skull and narrow, elongate rostrum H. abae is more primitive than H. larvatus or zoo 11. i. c 9 8 J. E. HILL H. speoris, it has the grooved upper canines of H. speoris and at the same time has developed independent specializations in its unsupported posterior leaf, prominent posterior palatal foramina and constricted sphenoidal region. The position of H. abae within the group is difficult to determine in view of these considerations. Its elongate skull and narrow rostrum suggest some affinity with the bicolor group and H. abae evidently forms a link between that group and the speoris group, but it is otherwise as specialized as H. larvatus and H. speoris in some features and in others perhaps more so. The degree of affinity of H. abae to its Asiatic relatives is similar to that of H. jonesi to its related Asiatic species in the bicolor group or to that of H. cyclops and H. camerunensis to the related Australian and Papuan species of the cyclops group. Hipposideros abae, however, is rather less divergent from its associated Asiatic species than H . jonesi, H. cyclops and H . camerunensis are from their closest congeners in the Asiatic and Australasian regions, perhaps reflecting a slightly less remote dichotomy in the speoris group. Its degree of divergence from its Asiatic relatives is paralleled by H. caffer and its associated species, which have a similar affinity to the Asiatic H. galeritus and its allies, while the African H. commersoni diverges less from the related Asiatic H. diadema than H. abae does from H. larvatus and H. speoris. DISTRIBUTION : Portuguese Guinea (Veiga Ferreira (1949 : 192)) ; Guinea (Aellen (i956a : 888), Eisentraut & Knorr (1957 : 331)) ; Sierra Leone ; Ghana ; Nigeria ; Congo ; Uganda (first record : specimen from Metu, West Madi County, West Nile District, in collection of British Museum (Natural History)). 56 60 62 64 66 H. ABAE J_ J 66 58 6O 62 64 FiG.3i. Length of forearm in Hipposideros abae Hipposideros larvatus The ears are as described for the group, with a small process at the antitragal fold. The anterior leaf has a small but distinct median emargination, the inter- narial septum is uninflated and the narial lappets are well developed, the nostrils slightly pocketed. The intermediate part of the leaf is slightly expanded, with a moderate median eminence and weaker lateral eminences. The posterior leaf is moderate, supported by three well-defined septa, its upper edge semicircular but slightly flattened and thickened, displaying a tendency to become lobate. There is A REVISION OF HIPPOSIDEROS 99 a frontal sac in both sexes, in females reduced sometimes to a depression containing a tuft of hairs. The skull is short and massive, with a well-developed sagittal crest and low supraorbital ridges. There is a shallow frontal depression and the rostral eminences are well inflated, separated by a shallow groove, the rostrum rounded. The anteorbital foramen is slightly elongate. The premaxillae make an acutely V-shaped junction with the maxillae and wholly enclose the slightly elongate anterior palatal foramina. The palation is U-shaped, with the vomer projecting beyond the posterior edge of the palate and the sphenoidal bridge only very slightly constricted, nearly parallel-sided. The sphenoidal depression is well developed. The upper incisors are weakly bilobed, the upper canines with a low anterior cusp but no posterior cusp and with their antero-internal faces not grooved. The anterior upper premolar (pm 2 ) is slightly extruded and compressed tightly between the canine and the second upper premolar (pm 4 ), while the posterior ridge of the third upper molar is one quarter the length of the anterior ridge. The anterior lower premolar (pm 2 ) is one third to one half the length of the second lower premolar (pm 4 ). Although H. larvatus is very similar to H. speoris, the two species are separated by a number of differences indicating comparatively recent divergence. Of the two, H. larvatus is perhaps slightly the more primitive, with a prominent median emargination in the anterior leaf, longer interorbital region with less prominent supraorbital ridges, longer, rounded rostrum and ungrooved canines, although these have lost the posterior cusp. The two species are more closely related to each other than either is to the African species H. abae, which displays some affinities to H. speoris but clearly is widely separated from the two Asiatic species. Hipposideros larvatus exhibits a cline in size in southeast Asia : H . I. grandis from Burma and H. I. alongensis from Indochina are the largest subspecies, while H. 1. barbensis from St. Barbe Island and H. I. larvatus from Java are the smallest. Specimens from Siam approach H. I. grandis, while those from Sumatra and its adjacent islands are nearer in size to H. 1. larvatus : H. I. neglectus from Borneo and the Malay Peninsula is intermediate between these extremes. DISTRIBUTION : Burma to Indochina and the Malay Peninsula ; Java ; Borneo ; Sumatra and adjacent islands. Hipposideros larvatus larvatus (Horsfield) Rhinolophus larvatus Horsfield, 1823 : No. 6, pi. Java. Rhinolophus vulgaris Horsfield, 1823 : No. 6, pi. Java. Rhinolophus deformis Horsfield, 1823 : No. 6, pi. Java. Rhinolophus insignis Horsfield, 1823 : No. 6, pi. Java. DISTRIBUTION : Java. Hipposideros larvatus sumbae Oei Hipposideros larvatus sumbae Oei, 1960 : 28. Eastern part of Sumba Island, Lesser Sunda Islands. Lectotype designated by Bree (1961 : 122). ioo J. E. HILL Hipposideros larvatus barbensis Miller Hipposideros barbensis Miller, 1900 : 233. St. Barbe Island, South China Sea. DISTRIBUTION : St. Barbe Island ; Johore Archipelago : Aor Island (Hill (1960 : 28)). Hipposideros larvatus neglectus Sody Hipposideros larvatus neglectus Sody, 1936 : 46. Roema Manoeal, south foot of Mount Kenepai, central Indonesian Borneo. Miller (1942 : 116) refers a specimen from Nias Island to H. I. neglectus but suggests that it may prove separable from both H. 1. larvatus and H. I. neglectus : Hill (1960 : 28) provisionally refers specimens from Butang Island and Simalur Island to H. I. neglectus. 58 H. L. LARVATUS H. L. BARBENSIS H. L. NEGLECTUS H. L. GRANDIS H. L. LEPTOPHYLUIS 54 56 56 6O 62 64 66 68 FIG. 32. Length of forearm in Hipposideros larvatus A REVISION OF H1PPOSIDEROS 101 DISTRIBUTION : Borneo ; Karimata Island ; South Natima Islands : Sirhassen Island ; Sumatra ; Nias Island ; Simalur Island ; Butang Island ; Malay Peninsula ; Tioman Island. Hipposideros larvatus grandis G. M. Allen Hipposideros larvatus grandis G. M. Allen, 1936 : 345. Akanti, Upper Chindwin, Burma, 500 feet. Shamel (1942 : 322) compared H. 1. grandis and H. I. neglectus. DISTRIBUTION : Burma ; Siam (Shamel (1942 : 322)) ; Indochina (part). Hipposideros larvatus atongensis Bourret Hipposideros larvatus alongensis Bourret, 19423. : 27. Bay d'Along, Indochina. External measurements of a small series of specimens are given by Bourret (19420 : 10). Hipposideros larvatus poutensis J. A. Allen Hipposideros poutensis J. A. Allen, 1906 : 483. Pouten, Hainan. Hipposideros larvatus leptophyllus (Dobson) Phyllorhina leptophylla Dobson, i8j<\a : 234. Khasi Hills, Assam. Hipposideros speoris The ears are as described for the group, pointed, their posterior margins slightly concave behind the tip and have a small projecting process at the antitragal lobe. The anterior leaf has only a faint trace of a median emargination, the internarial septum is slightly inflated and the narial lappets are well developed, the nostrils slightly pocketed. The intermediate part of the leaf is slightly expanded, with inflated median and lateral eminences. The posterior leaf is supported by three septa and its upper edge is semicircular and not especially thickened. There is a frontal sac in male specimens : it is absent or represented by a tuft of hair in female examples. The skull is similar to the skull of H. larvatus, with a low to moderate sagittal crest and low but definite and more prominent supraorbital ridges. There is a shallow frontal depression and the rostral eminences are well inflated, the rostrum flattened dorsally. The premaxillae make an acutely angled wedge-shaped junction with the maxillae and partially or wholly enclose the rather elongate anterior palatal foramina. The palation is U-shaped and the vomer does not project beyond the edge of the palate. The sphenoidal bridge is moderate and only slightly constricted and there is a shallow but well-defined sphenoidal depression. The upper incisors are not bilobed or only very weakly so and the upper canines have low anterior and posterior cusps and a shallow groove on their antero-internal faces. The 102 J. E. HILL anterior upper premolar (pm 2 ) is slightly extruded, compressed tightly between the canine and the second upper premolar (pm 4 ), while the posterior ridge of the third upper molar is one third the length of the anterior ridge. The anterior lower premolar (pm 2 ) is one half the length and height of the second lower premolar (pm 4 ). Brosset (1962 : 608) provides a study of the biology of H. speoris in India, with measurements and notes on its colour variation. DISTRIBUTION : India ; Ceylon. Hipposideros speoris speoris (Schneider) Vespertilio speoris Schneider, 1800 : pi. 5Qb. Tate (1941 : 377) has suggested restriction of the type locality to Tranquebar, India : it is briefly reviewed by Oey & Feen (1958 : 231). Rhinolophus marsupialis Desmarest, 1820 : 126. Rhinolophus dukhunensis Sykes, 1831 : 99. Deccan, India. Hipposideros apiculatus Gray, 1838 : 492. Madras, India. Hipposideros templetonii Kelaart, i85oa : 208. Ceylon. Hipposideros aurens Kelaart, 1853 : 18. Ceylon. Hipposideros blythi Kelaart, 1853 : 20. Ceylon. 46 48 H. S. SPEORIS H. S. PULCHELLUS 46 48 5O 52 54 FIG. 33. Length of forearm in Hipposideros speoris A REVISION OF HIPPOSIDEROS 103 Oey & Feen (1958 : 227) discuss the date of description of Vespertilio speoris Schneider and reprint the original text. They also study (p. 232) the external differences between H. speoris and H. larvatus in considerable detail. DISTRIBUTION : India (part) ; Ceylon. Hipposideros speoris pulchellus Andersen Hipposideros speoris pulchellus Andersen, 1918 : 383. Vijayanagar, Bellary, India. DISTRIBUTION : India (part). HIPPOSIDEROS DIADEMA group The ears are triangular, acutely pointed, their posterior margins concave behind the tip, with no antitragal modifications. The noseleaf is comparatively simple, with three or four lateral supplementary leaflets. The anterior leaf has no median emargination and the internarial septum is not inflated. The intermediate part of the leaf is expanded and the posterior leaf is high, supported by a median septum and two weaker lateral septa. A frontal sac may be present or absent. The skull is large, with moderate or strongly developed cranial crests. A frontal depression is present in the majority of species. The rostral eminences are moderately inflated, the rostrum broad and high, its rounded upper surface level with the roof of the braincase. The zygomata are massive, with a prominent jugal process and the anteorbital foramen is large, elongate and closed by a narrow bar. The premaxillae make a wedge-shaped junction with the maxillae and partially or wholly enclose the anterior palatal foramina. The palate is short, the vomer not projecting or projecting only very slightly beyond its posterior edge. The mesopterygoid fossa is wide, the pterygoid wings expanded, terminating in delicate processes. The sphenoidal bridge is wide, partially concealing elongate lateral apertures. The sphenoidal depression is moderately developed and the cochleae are large, their width as great or greater than their distance apart. The mandible is massive, with a deep symphysis and a high coronoid process : the angular process is sometimes flexed outwards. The upper incisors are bilobed, the upper canines with or without a posterior cusp. The anterior upper premolar (pm 2 ) is small, partially or wholly extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is much reduced and obsolescent. The crown area of the outer lower incisors is greater than that of the inner pair and the anterior lower premolar (pm 2 ) is from one third the length and height to one half or two thirds the length and height of the second lower premolar (pm 4 ). As defined in the present paper, the diadema group has been extended to include the commersoni group as listed by Tate (1941 : 358), including solely the African species H. commersoni. There appears no good reason for the wide separation of this species from the diadema group since it shares with H. diadema and its associated species a number of features indicative of a common origin. Like H. diadema and its allies, it is characterized by its acute, triangular ears, simple noseleaf with IO4 J. E. HILL three or four lateral supplementary leaflets and massive skull with greatly developed cranial crests. As might be expected, the African H. commersoni is rather sharply FIG. 34. Possible relationships in the HipposiJeros diadema group A REVISION OF H1PPOSIDEROS 105 separated from H. diadema, which with its allies forms a comparatively closely related group of wholly Asiatic and Australasian species. In general, these are more primitive and less modified than H. commersoni , with broader ears, less de- veloped postorbital processes, the naso-frontal region rounded from its dorsal aspect and not pentagonal, and with the upper canines not grooved anteriorly. Hipposideros lankadiva and H. schistaceus are the least specialized species in the group : H. diadema and more particularly its immediate derivatives H. dinops and H. inexpectatus are more modified, while H. commersoni is the most advanced species of the group, with narrow ears, projecting postorbital processes, more pentagonal naso-frontal region, grooved canines and with the anterior premolars (pm-f) more reduced. Possible relationships of the species of the diadema group are summarized in Figure 34. The species of the diadema group may be keyed : i Ears broad at base : postorbital processes not projecting, naso-frontal region rounded, upper incisors in contact or nearly so, upper canines not grooved anteriorly, lacking posterior cusps ......... 2 Ears narrow at base : postorbital processes projecting, naso-frontal region pentagonal, upper incisors widely separated, upper canines grooved anteriorly, with high posterior cusp ......... commersoni (.p 115) z Frontal region convex or flattened, no frontal depression ; palation acute, V-shaped ; sphenoidal depression well developed ........ 3 Frontal region concave, with a shallow frontal depression ; palation rounded, U-shaped ; sphenoidal depression shallow . . . . . . . 4 3 Skull larger, condylocanine length exceeding 26-5 mm., length of maxillary toothrow (c-m 3 ) exceeding 12-0 mm. ....... lankadiva (p. 105) Skull smaller, condylocanine length less than 25-5 mm., length of maxillary toothrow (c-m 3 ) less than u-8 mm. ....... schistaceus (p. 107) 4 Smaller, length of forearm less than 98-0 mm., condylocanine length less than 33-0 mm. ; intermediate part of noseleaf with prominent median ridge, sometimes form- ing a projection ............ 5 Larger, length of forearm exceeding 100 mm., condylocanine length exceeding 34-0 mm. ; intermediate part of noseleaf swollen but without a prominent median ridge or projection ........ inexpectatus (p. 113) 5 Smaller, length of forearm less than 92-0 mm., condylocanine length less than 31-0 mm., length of maxillary toothrow (c-m 3 ) less than 13-8 mm. . . diadema (p. 108) Larger, length of forearm exceeding 92-0 mm., condylocanine length exceeding 32-0 mm., length of maxillary toothrow (c-m 3 ) exceeding 14-0 mm. dinops (p. 112) Hipposideros lankadiva The ears are large and acutely pointed, with their posterior margins slightly concave behind the tip. The noseleaf has usually four lateral supplementary leaflets, the fourth much reduced and rudimentary, sometimes absent. The anterior leaf has no median emargination and the internarial septum is not inflated, while the narial lappets are well developed. The intermediate part of the leaf is expanded, its central part inflated and swollen but not forming a distinct median ridge, flanked laterally by lesser eminences. The posterior leaf is high and broad, its upper margin semicircular, with a swollen median projection, flanked by narrow notches, and is supported by three well-defined septa enclosing small cells. The skull is large and io6 J. E. HILL heavily built, with well-developed cranial crests. There is no frontal depression and the frontal region immediately anterior to the sagittal crest is convex or flattened. The zygomata are strong, with a high jugal process, while the anteorbital foramen is large, elongate and closed by a narrow bar. The premaxillae make a U-shaped junction with the maxillae and enclose or nearly enclose the rounded anterior palatal foramina. The palation is V-shaped, the mesopterygoid fossa slightly narrowed with the vomer projecting slightly beyond the posterior edge of the palate. The sphenoidal bridge is wide, almost completely concealing elongate lateral aper- tures and the sphenoidal depression is well developed. The width of the cochleae is a little less than their distance apart. The mandible is massive, with a high coronoid process and the angular process flexed strongly outwards. The upper incisors are bilobed, the outer lobe a little larger than the inner lobe, and the upper canines are not grooved anteriorly : they have low anterior cusps but lack posterior cusps. The anterior upper premolar (pm 2 ) is small, extruded or partially extruded from the toothrow. The posterior ridge of the third upper molar is obsolescent. The crown area of the outer lower incisors is considerably greater than that of the inner pair, and the anterior lower premolar (pm 2 ) is one third the length and height of the second lower premolar (pm 4 ). Hipposideros lankadiva is closely related to H. diadema but is perhaps slightly less specialized. Externally, it is very similar to H. diadema, but the intermediate part of the noseleaf has no definite median ridge and the median projection of the edge of the posterior leaf is more swollen and prominent. Cranially, it differs rather sharply in its convex or flattened frontal region, which lacks the frontal depression of H. diadema, in its V-shaped and not U-shaped palation and in its well-developed sphenoidal depression. With the very closely related H. schistaceus it represents H. diadema in the Indian subcontinent. Brosset (1962 : 621) has studied the biology of H. lankadiva in India and gives measurements and notes on its colour variation : this author suggests that in India the species is perhaps not subspecifically separable. DISTRIBUTION : Ceylon ; Peninsular India. Hipposideros lankadiva lankadiva Kelaart Hipposideros lankadiva Kelaart, 18505 : 216, Kandy, Ceylon. DISTRIBUTION : Ceylon. Hipposideros lankadiva Indus Andersen Hipposideros indus Andersen, 1918 : 382. Gersoppa, Kanara, India. Hipposideros lankadiva mixtus Andersen Hipposideros indus mixtus Andersen, 1918 : 382. Kolar, eastern Mysore, India. The available material of this and the other subspecies of H. lankadiva is ve^ limited, but there seems little difference between H. I. indus and H. I. mixtus and they are likely to prove synonymous. A REVISION OF HIPPOSIDEROS Hipposideros lankadiva unitus Andersen 107 Hipposideros indus unitus Andersen, 1918 : 382. Mundra, Saugor, Central Provinces, India, i, 600 feet. ao 92 H. L. LANKADIVA H. L.INDUS H.L. MIXTUS H.L. UNITUS 78 90 92 FIG. 35. Length of forearm in Hipposideros lankadiva Hipposideros schistaceus Andersen Hipposideros schistaceus Andersen, 1918 : 382. Vijayanagar, Bellary, India. Andersen gave only a brief diagnosis of this species. Its ears and noseleaf are exactly as in H. lankadiva but its coloration is paler and less brown and the skull, although resembling that of H. lankadiva very closely, is smaller, less massive, lower and flatter : the cranial crests are less developed and the cochleae are com- paratively wider, their width equal to their distance apart. The remaining features of the skull, and its dentition, exactly resemble H. lankadiva. Hipposideros schistaceus is evidently very closely related to H. lankadiva, differing from this species chiefly in its smaller, less massive skull and relatively larger bullae. The 72 74 78 ao 82 84 - H. SCHISTACEUS 72 74 76 78 8O 82 84 FIG. 36. Length of forearm in Hipposideros schistaceus io8 J. E. HILL available material, both of H. schistaceus and of the smaller subspecies of H. lankadiva, is too limited to determine the extent of size variation in either H. schistaceus or H. lankadiva, and for the present their exact relationship must remain uncertain. Hipposideros diadema The ears are of moderate size, broad at the base and acutely pointed, with their posterior margins concave behind the tip. The noseleaf is well developed, with three or four lateral supplementary leaflets, the fourth small and sometimes rudi- mentary. The anterior leaf has no median emargination and the internarial septum is not inflated. The narial lappets are well developed and the nostrils are slightly pocketed. The intermediate part of the leaf is expanded with a prominent median ridge forming a median projection, flanked laterally by two much smaller projections. The posterior leaf is high, thick and fleshy, its upper edge semicircular, with a small median projection, and is supported by a median septum and two weak lateral septa. There is no frontal sac. The skull is large, its characters mainly those of the group. The cranial crests are moderately developed, the postorbital processes rounded and there is a shallow frontal depression. The rostral eminences are moderately inflated, the naso-frontal region from its dorsal aspect rounded and not pentagonal. The premaxillae make a wedge-shaped or slightly rounded junction with the maxillae and partially or wholly enclose the slightly elongate anterior palatal foramina. The palation is U-shaped or slightly V-shaped with the vomer projecting only very slightly beyond the posterior edge of the palate. The pterygoids and sphenoidal bridge are wide, almost completely concealing the elongate lateral apertures. There is a shallow sphenoidal depression and the cochleae are moderate, their width as great or nearly as great as their distance apart. The upper incisors are bilobed, the outer lobe usually a little larger than the inner lobe and are closely approximated with their tips convergent. The upper canines are not grooved anteriorly, and have no posterior cusps. The anterior upper premolar (pm 2 ) is small, partially or wholly extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is obsolescent or obsolete. The crown area of the outer lower incisors is considerably greater than that of the inner pair, while the anterior lower premolar (pm 2 ) is one half to two thirds the length and height of the second lower premolar (pm 4 ). Hipposideros diadema has been divided into numerous subspecies over its extensive range, which includes almost the entire Indo-Australian region. As Tate (1941 : 373 et seq.) recognized, two broad groups of subspecies can be discerned among them. One group, composed of larger subspecies, is confined to the Asiatic mainland, Sumatra, Borneo, Java and their adjacent islands : the other, composed of smaller subspecies, is distributed from Celebes and the Philippine Islands eastwards to the Solomon Islands and northern Australia. There is some overlap between the two groups and larger subspecies are found in the Moluccas on the islands of Ceram and Batchian. Although in general the larger subspecies occur in the western parts A REVISION OF HIPPOSIDEROS 109 of the range of H. diadema and the smaller towards the eastern limits of its distribu- tion, there appears to be little evidence of a clinal pattern of size variation. DISTRIBUTION : Burma, Nicobar Islands and Philippine Islands east to the Solomon Islands and northern Australia. Hipposideros diadema diadema (Geoffrey) Rhinolophus diadema Geoffrey, 1813 : 263, pis. 5, 6. Timor Island. Rhinolophus nobilis Horsfield, 1823 : No. 6, pi. Java. DISTRIBUTION : Timor Island ; Sumbawa Island ; Java. Hipposideros diadema masoni (Dobson) Phyllorhina masoni Dobson, 1872 : 338. Moulmein, Burma. Hipposideros diadema vicarius Andersen, 1905 : 499. Sarawak. According to Andersen (1905 : 500) Phyllorhina masoni Dobson is of uncertain application but later (1907 : 6) he recognized it as a subspecies of H. diadema and gave brief diagnostic characters to separate this from the nominate subspecies. There appear to be no grounds for the separation of specimens from the Asiatic mainland from those of Borneo and Sumatra when large series are examined. Osgood (1932 : 221) gives notes on specimens from Annam and Shamel (1942 : 322) compares specimens from Siam with those from Java. DISTRIBUTION : Burma ; Indochina : Annam (Osgood (1932 : 221)) ; Tonkin ; Siam (Shamel (1942 : 322)) ; Tenasserim ; Malay Peninsula ; Borneo ; Sumatra ; Nias Island (Miller (1942 : 117)). Hipposideros diadema nicobarensis (Dobson) Phyllorhina nicobarensis Dobson, 1871 : 262. Nicobar Islands, Bay of Bengal. Hipposideros diadema enganus Andersen Hipposideros diadema enganus Andersen, 1907 : 8. Engano Island. Sody (1940 : 394) considers H. d. enganus to be a synonym of H. d. masoni. Hipposideros diadema natunensis Chasen Hipposideros diadema natunensis Chasen, 1940 : 43. Bunguran Island, North Natuna Islands. Hipposideros diadema griseus (Meyen) Rhinolophus griseus Meyen, 1833 : 608, pi. 46. San Matheo Cave, Montalban, near Manila, Luzon, Philippine Islands. Hipposideros diadema anderseni Taylor, 1934 ' 246. Novaliches, Rizal Province, Luzon. Lawrence (1939 : 53) commented on the status of H. d. anderseni. DISTRIBUTION : Philippine Islands : Mindoro ; Cebu ; Mindanao ; Catanduanes ; Leyte ; Luzon ; Guimaras. J. E. HILL FIG. 37. Length of forearm in Hipposideros diadema A REVISION OF HIPPOSIDEROS in Hipposideros diadema speculator Andersen Hipposideros diadema speculator Andersen, 1918 : 381. Kalao Island, Flores Sea, south of Celebes. DISTRIBUTION : Celebes ; Kalao Island. Hipposideros diadema ceramensis Laurie & Hill Hipposideros diadema ceramensis Laurie & Hill, 1954 : 5&- Teleoti Bay, south Ceram Island, Molucca Islands. DISTRIBUTION : Molucca Islands : Ceram ; Buru. Hipposideros diadema euotis Andersen Hipposideros euotis Andersen, 1905 : 502. Batchian Island, Molucca Islands. Hipposideros diadema pullatus Andersen Hipposideros diadema pullatus Andersen, 1905 : 498. Haveri, Papua, 700 metres. DISTRIBUTION : New Guinea. Hipposideros diadema custos Andersen Hipposideros diadema custos Andersen, 1918 : 381. Ara, Kei Island. Hipposideros diadema mirandus Thomas Hipposideros demissus mirandus Thomas, 1914 : 437. Manus Island, Admiralty Islands. Hipposideros diadema trobrius Troughton Hipposideros diadema trobrius Troughton, 1937 : 2 ?6- Kiriwina Island, Trobriand Islands. Hipposideros diadema oceanitis Andersen Hipposideros diadema oceanitis Andersen, 1905 : 497. Guadalcanar, Solomon Islands. DISTRIBUTION : Solomon Islands : Guadalcanar ; Fauro ; Vella Lavella ; Ysabel ; Bougainville. Hipposideros diadema demissus Andersen Hipposideros demissus Andersen, 1909 : 268. Yanuta, San Christoval Island, east Solomon Islands. Hipposideros diadema reginae Troughton Hipposideros diadema reginae Troughton, 1937 : 2 75- Bloomfield River, Cooktown area, Queensland, Australia. ii2 J. E. HILL 64 66 66 TO 72 74 76 78 8O 92 H. D PULLATUS H. D. GUSTOS H. D. MiRANDUS H. D. OCEANITIS H. 0. DEMISSUS FIG. 38. Length of forearm in Hipposideros diadema Hipposideros dinops The ears are large, broad at the base and acutely pointed, their posterior margins concave just behind the tip. The noseleaf is large, with three lateral supplementary leaflets, the third small. The anterior leaf is broad and has no median emargination while the internarial septum is not inflated, the narial lappets are well developed and the nostrils slightly pocketed. The intermediate part of the leaf is expanded, with a prominent projecting median eminence, flanked laterally by much lesser swellings. The posterior leaf is thick and fleshy, its upper edge semicircular with a small, incipient median projection, and is supported by a median septum and two weak lateral septa. There is no frontal sac in the female type specimen. The skull is large, with well-developed lambdoid and sagittal crests. The postorbital processes are rounded and project very slightly, and there is a shallow frontal depression. The rostral eminences are moderately inflated and the rostrum is broadened, the naso-frontal region from its dorsal aspect very slightly less rounded and more angular than in H. diadema. The zygomata are moderate, with a prominent jugal process, and the anteorbital foramen is large, elongate and closed by a narrow bar. The premaxillae make a V-shaped contact with the maxillae and do not enclose or only just enclose the elongate anterior palatal foramina. The palation is U-shaped, the vomer projecting only very slightly beyond the posterior edge of the palate. The mesopterygoid fossa is wide and the pterygoids expanded, with the wide sphenoidal bridge almost completely concealing elongate lateral apertures. There is a shallow sphenoidal depression and the cochleae are a little wider than their distance apart. The mandible is massive, with a high coronoid process and with the massive angular process flexed slightly outwards. The upper incisors are strong and almost in contact, weakly bilobed, while the upper canines are massive and lack anterior or posterior cusps. The anterior upper premolar (pm 2 ) is small and A REVISION OF HIPPOSIDEROS 113 extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact, while the posterior ridge of the third upper molar is almost obsolete. The crown area of the outer lower incisors is considerably greater than that of the inner pair and the anterior lower premolar (pm 2 ) is one half the length and height of the second lower premolar (pm 4 ). Hipposideros dinops differs principally from H. diadema in its greater size, more greatly developed intermediate noseleaf, more angular naso-frontal region, more expanded zygomata and larger teeth. It is closely related to H . diadema and although evidently presenting some of the features of H. diadema in more exaggerated form, its angular, slightly pentagonal naso- frontal region, expanded zygomata and massive mandible are clearly correlations of the trend towards great size. In this respect, H. dinops links H . diadema to the giant species H. inexpectatus and displays to some extent the specializations of this species and of the more remotely related large African species H. commersoni. DISTRIBUTION : Celebes ; Peleng Island ; Solomon Islands. Hipposideros dinops dinops Andersen Hipposideros dinops Andersen, 1905 : 502. Rubiana Island, Solomon Islands. DISTRIBUTION : Solomon Islands : Rubiana ; Bougainville. Hipposideros dinops pelingensis Shamel Hipposideros pelingensis Shamel, 1940 : 353. Peleng Island, east of Celebes. Hipposideros pelingensis Shamel is unrepresented in the collections of the British Museum (Natural History) but there seems very little doubt from its brief descrip- tion that it is only subspecifically related to H. dinops. The tibia of H. d. pelingensis is shorter than that of H. d. dinops and Shamel says of the upper incisors that they " are small and bicusped, their inner cusps larger than the outer ones ". In the type specimen of H. d. dinops the outer lobes of the upper incisors are slightly larger than the inner lobes. Tate (1941 : 374) referred a series from south Celebes to H . d. pelingensis and noted that in size they were almost as large as H. d. dinops from Rubiana Island. Apparently with reference to this series, Tate stated that the premaxillae do not wholly enclose the anterior palatal foramina : in the type specimen of H. d. dinops these apertures are just enclosed by the anterior enclosing processes of the premaxillae. DISTRIBUTION ; Peleng Island ; Celebes (Tate (1941 :_374, 38% 391)). Hipposideros inexpectatus Laurie & Hill Hipposideros inexpectatus Laurie & Hill, 1954 : 60. Posso (=Poso), north Celebes. A very large species, with ears similar to those of H. diadema, very large and broad at the base, acutely pointed and with their posterior margins concave just behind the tip. The noseleaf is very large and has four lateral supplementary leaflets : the third leaflet is reduced and the fourth minute. The anterior leaf is zoo 11, i. H n 4 J. E. HILL broad and has no median emargination. The internarial septum is not inflated and the narial lappets are well developed. The intermediate part of the leaf is greatly expanded with its central part much swollen and inflated but with only a very low, indefinite median ridge which does not extend to the upper margin of the intermediate part of the leaf : the swollen central part is flanked laterally by lesser eminences. The posterior leaf is thick and fleshy, its upper edge semicircular with a very small, incipient median projection and is supported by a narrow median septum and rather broader, less definite lateral septa. The skull is very large, with greatly developed lambdoid crests and an enormously developed, crescentic sagittal crest, merging into prominent supraorbital ridges. The postorbital processes project slightly and there is a shallow frontal depression. The rostral eminences are moderately inflated and the rostrum is broad, the naso-frontal region from its dorsal aspect slightly pentagonal, less rounded and more angular than in H. diadema or H. dinops. The zygomata are massive, greatly expanded and have a prominent jugal process, while the anteorbital foramen is very large and elongate, closed by a long, narrow bar. The premaxillae make a shallowly V-shaped junction with the maxillae and just enclose the slightly elongate anterior palatal foramina. The palation is U-shaped, the vomer projecting very slightly beyond the posterior edge of the palate, with wide mesopterygoid fossa, expanded pterygoids and large, wide sphenoidal bridge almost completely concealing elongate lateral apertures. There is a shallow sphenoidal depression and the width of the cochleae is a little greater than their distance apart. The mandible is massive, with a deep symphysis, high coronoid process and a thick, substantial angular process flexed strongly outwards. The upper incisors are large, their tips almost in contact and are weakly bilobed while the upper canines are massive and lack anterior or posterior cusps. The anterior upper premolar (pm 2 ) is small and extruded from the toothrow, the canine and the second upper premolar (pm 4 ) in contact. The posterior ridge of the third upper molar is obsolescent. The crown area of the outer lower incisors is considerably greater than that of the inner pair, and the anterior lower premolar (pm 2 ) is one half the length and height of the second lower premolar (pm 4 ). 94 96 98 !OO 1O2 H. D. DINOPS H. INEXPECTATUS 94 96 98 IOO 1O2 FIG. 39. Length of forearm in Hipposideros dinops and H. inexpectatus A REVISION OF HIPPOSIDEROS 115 Hipposideros inexpectatus is evidently closely related to H. dinops, but it differs from that species in its larger size, in its slightly less specialized intermediate nose- leaf, which has no definite median ridge or projection as in H. dinops and in the greater development of those cranial specializations associated with great size. The skull of H. inexpectatus is larger than that of H. dinops, with more greatly developed cranial crests, a more angular, slightly pentagonal naso-frontal region and a much more massive mandible, and in these features approaches the large African species H. commersoni. Its broad ears, closely approximated upper incisors and the absence of a groove in the anterior faces of the upper canines ally it indubi- tably to H. diadema and H . inexpectatus is evidently an extreme of the trend towards great size of which H. dinops is a part. Hipposideros commersoni A very large species with the ears narrow at the base, triangular, rounded at the tips and with their posterior margins concave just behind the tip. The noseleaf is large and not greatly specialized, with three or four lateral supplementary leaflets, the fourth small, sometimes rudimentary and rarely but on occasion absent. The anterior leaf is broad and has no median emargination and the internarial septum is not inflated, while the narial lappets are well developed and the nostrils are slightly pocketed. The intermediate part of the leaf is expanded, with a low, indefinite median ridge or eminence and lacking lateral inflations. The posterior leaf is of moderate height, its upper edge flattened, without a median thickening or projection, and is supported by a median septum and two very weak lateral septa. A frontal sac is present in both sexes, its opening more or less longitudinal. The skull is large, with well-developed lambdoid crests and a greatly developed sagittal crest merging into sharply defined supraorbital ridges. The postorbital processes project slightly and the rostral eminences are moderately inflated. The rostrum is broad and the naso-frontal region from its dorsal aspect is distinctly pentagonal in outline. The zygomata are strong with a prominent jugal process, and the anteorbital foramen is large, elongate and closed by a narrow bar. The premaxillae make a V-shaped junction with the maxillae and wholly enclose the slightly elongate anterior palatal foramina. The palation is U-shaped, rather square, the vomer not projecting beyond the posterior edge of the palate. The mesopterygoid fossa is moderate and the pterygoids slightly constricted posteriorly, while the sphenoidal bridge is moderate, not excessively constricted, partially concealing elongate lateral apertures. There is a well-developed sphenoidal depres- sion and the width of the cochleae is a little greater than their distance apart. The mandible is massive, with a deep symphysis, high coronoid process and a substantial angular process, flexed strongly outwards. The upper incisors are very weakly bilobed and set widely apart, at the outer margins of the premaxillae, while the upper canines have their anterior faces shallowly grooved and have low posterior cusps. The anterior upper premolar is very small, extruded from the toothrow, with the canine and the second upper premolar (pm 4 ) in contact or nearly so. The posterior ridge of the third upper molar is obsolete or almost obsolete. The crown n6 J. E. HILL area of the outer lower incisors is a little greater than that of the inner pair, while the anterior lower premolar (pm 2 ) is one half or less the length and height of the second lower premolar (pm 4 ). Hipposideros commersoni stands rather sharply apart from the Asiatic species of the group by virtue of its narrower ears, pentagonal naso-frontal region, widely separated upper incisors and grooved upper canines, which possess low posterior cusps. Although in some respects H. commersoni is approached by the large Australasian species H. dinops and H. inexpectatus, these are basically species of the diadema type which appear to have developed specializations similar to those of H. commersoni as correlations of their large size, and H. commersoni must be regarded as a species of rather remote origin in the diadema group. DISTRIBUTION : Madagascar ; eastern Africa south to Nyasaland and Northern Rhodesia ; Southwest Africa and Angola north to Gambia ; Congo. Hipposideros commersoni commersoni (Geoffroy) Rhinolophus commersoni Geoffroy, 1813 : 263, pi. 5. Fort Dauphin, Madagascar. DISTRIBUTION : Madagascar. 78 80 82 84 88 9O 92 H. C. COMMERSONI H. C. THOMENSIS 78 8O 82 84 86 88 9O 92 FIG. 40. Length of forearm in Hipposideros commersoni Hipposideros commersoni marungensis (Noack) Phyllorhina commersoni var. marungensis Noack, 1887 : 272, pi. 10, figs. 31-33. Qua-Mpala, Marungu, western Tanganyika. Hipposideros commersoni mostellum Thomas, 1904^ : 385. Tana River, Kenya. DISTRIBUTION : Kenya ; Tanganyika ; Zanzibar ; Nyasaland ; Northern Rhodesia ; Southwest Africa. Hipposideros commersoni gigas (Wagner) Rhinolophus gigas Wagner, 1845 : 148. Benguela, Angola. Phyllorrhina vittata Peters, 1852 : 32, pi. 6, figs. 1-3, pi. 13, figs. 7-13. Ibo Island, Cap Delgado group, 12 20' S. (in part, male co-type only : lectotype selected by Andersen (igoGa : 45)). (?) Hipposideros gigas gambiensis Andersen, igo6a : 42. Gambia. Hipposideros gigas viegasi Monard, 1939 : 70. Madina Boe, Portuguese Guinea. A REVISION OF HIPPOSIDEROS 117 FIG. 41. Length of forearm in Hipposideros commersoni n8 J. E. HILL Hipposideros gigas gambiensis was separated from H. c. gigas by Andersen on the grounds of slightly wider noseleaves. It seems unlikely to be a valid subspecies. DISTRIBUTION : Angola ; Congo (part) ; Tanganyika (part) ; Cameroons ; Nigeria ; Ghana ; Guinea ; Portuguese Guinea ; Gambia. Hipposideros commersoni thomensis (Bocage) Phyllorhina commersoni var. thomensis Bocage, 1891 : 88 and 1905 : 67. San Thome Island. Hipposideros commersoni niangarae J. A. Allen Hipposideros gigas niangarae J. A. Allen, 1917 : 438, pi. 51, fig. i. Niangara, Uele district, Congo. SUMMARY The genus Hipposideros presents a wide range of morphological variation, and the characters of its numerous species rarely combine to indicate clearly defined natural groups or evolutionary trends. This revision, based chiefly on the features of the ears, noseleaf and skull, discusses the morphological criteria within the genus and its supraspecific groupings, recognizing seven species groups containing a total of forty- three species. These groups and their included species are described in some detail, with keys, and their probable relationships have been defined and discussed. These studies have suggested a less diffuse classification of Hipposideros than those proposed by earlier revisers, and indicate that despite the wide morphological dissimilarity of many of its species, the genus includes but three major evolutionary trends. ACKNOWLEDGEMENTS My thanks are due to Mr. H. M. van Deusen, Curator of the Archbold Collections of the American Museum of Natural History, who arranged the loan of specimens ; to Dr. G. B. Corbet, for his valuable suggestions on the presentation of data ; to Mr. R. W. Hayman, whose knowledge of Africa and of African bats has been readily available and to Miss Annette Bown, who prepared the illustrations and diagrams used in this paper. A REVISION OF HIPPOSIDEROS 119 (giu-o) Avojq^oo^ AJBJIIXBUI jo q^Suaq 00 MMUO uo ON t^ 00 -* * ^ oo N Tt - 'uol>.O "o-J O "3* O ^ uo uo'SoUOo^uo'uouo 1 1 o M i, o 6 1 M^'JOOO^ON^M^,^ w i> t^i> w ^2 ro Tf uo uo O uouououo uo uo O *O ^t" suouipads jo jgquin^f M-MTJ-I^M MO fOOO M CO MM M M M O M M M N \T> fOTj-M UO 00 UO Tfr- O\. f^ ON t^. ON M O> O* 1 ON M OO O MOOM o ob ON ob obobooob TJ- ob 00 M ob ro 1 co ob M 9 ^ ,030 ob M ^=0 22,o CO suguipgds jo agquinjvj ro MO M t>- MM M O M M M o oo M ri W ,.0^ OMZ O C^ Tf ro O ^~ ON Tt-ONf^0 ONMuoOOO ^ OOorO O OOroOOo O^^ON0^ uo000 ob ob ob > ob f>. M M CO ob & o fr ON ob 1 vO i 'ro' STOUHKKbjoMqumu M MTJ-MOO MM M f^M M MM M M CO M M N M ON t^ 00 IT> 0. IX O M CO ^ cc siftsiiiil&Jltisi? 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CO CO M CO T^ M ^" O O MM in M M >-, H i H 1[^.SU9| 9UTU'BOO{XpUO3 H 3 i M - . c**^ - " r^** * * o ' * r*** " ^ r*** co " * f^l w M CO M "^" CO ^^ C^ ^ C 1 ! 04 ^* 1 M 1 'sO ' vO ' O ^ t*^* ' ' t*^* O^ 1 fsl ^ M ^^ M '^ CO ^^ C 1 ! ^ '^ C^l o ^~^" ^o N *~ "" *o N ^^' o ^~^ i *o > ^ r^ r^* *"""'' ^s| M M CO N 01 M M O "^1" ON O A in ' OO ' OO ON 00 J. ^*O ' ON M M o | ^^ | ^ | ^9 ( x'; 4 N M M M MM su9iup9ds jo J9qmn^[ CO CO CO ro vO N ro M M CO M OO MMCO MM M J a i H Q - ^ 3 ^ 5 ^ pP ^ Jg .g S ^N lO "^ "^ ^ H "S * "fe * a 'S s 1- | ' to to * ;< -^ to o to g ' S to J I ' -S 1 55 g S ^ 2 c-2 ^ $ ? ^ 1 s c S -^ t ^ ^3 S 'toStio t^vj e S 8 * 2 e "*"* ^i Qi ^> ^i ^ ^i ^i 2 * ^ ^' r ^^ -si e _ e . . to ^ ^ '^S'^S'TS ^^S Bi -aj- -^ as &j&3 A REVISION OF HIPPOSIDEROS 123 iO HI ^ O\ O O O oo to CM ^""^ i i r* O HI - ^ CM --~r* Tt~ - * M -~~-* M rOMOpOr^M7tH|NCM|_ir^|_|9 w inM9 Q ._. ro hdMuOw^MM^M *O i_f O HH ON i CO ^ ^ H) M O O ' H) CO ^ O 00 M N CO M 9 9 op 9 T+- TJ- ro ' Tt* O OO t ro I iO ' VO ' M M h- 1 O M MM ^0 M ^ M "^ M 00 TJ- ro Tj- TJ- N M O MNCO NMMfO >O >O ro '*- T)-Ot>-N O 1o ro CO f) Co~ ^ O t^ ^ 'I-T ^ O* ^ ST ^ > o -^ . ^ . ^^ . H T*" ON "ON OO^ N ^ T*" ^ o ^^ ONOO k ob, ^ P M 6 " ' ob ob t~> M M to N H M rj- N |p VN 00 M 1 O oo ^ I M N M VP S to ^ g 1 I s S s -2 K) o ^ g ^y .. ^ ^ "xs ts "xs ^s *e ts o s ^^s 124 J- E. HILL REFERENCES AELLEN, V. 1952. Contribution a 1'etude des chiropteres du Cameroun. Mem. Soc. Neuchatel Sci. Nat. 8 : 1-121, 26 figs., map. - 1954. Description d'un nouvel Hipposideros (Chiroptera) de la Cote d'lvoire. Rev. suisse Zool. 61 : 473-483, 2 figs. - 19563. Speleologica africana. Chiropteres des grottes de Guinee. Bull. Inst. franc. Afr. Noire ISA : 884-894, i tab. - i956b. Le Pare National du Niokolo-Koba (premier fascicule). II. Chiropteres. Mem. Inst. franc. Afr. Noire No. 48 : 23-34, 5 tabs. ALLEN, G. M. 1921. A new horseshoe bat from West Africa. Rev. zool. afr. 9 : 193-196. - 1936. Two new races of Indian bats. Rec. Indian Mus. 38 : 343-346. ALLEN, J. A. 1906. Mammals from the island of Hainan, China. Butt. Amer. Mus. nat. Hist. 22 : 463-490, i pi. - 1917. Part i. 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(ACARI: MESOSTIGMATA) S. K. BHATTACHARYYA BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. 2 LONDON: 1963 A REVISION OF THE BRITISH MITES OF THE GENUS PERGAMASUS BERLESE S.LAT. (ACARI: MESOSTIGMATA) BY S. K. BHATTACHARYYA Pp. 131-242 ; Plates 1-8 ; 313 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. 2 LONDON: 1963 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. n, No. 2 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. Trustees of the British Museum (Natural History) 1963 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued December, 1963 Price Forty-five shillings A REVISION OF THE BRITISH MITES OF THE GENUS PERGAMASUS BERLESE S.LAT. (ACARI: ME SO STIGMATA) By S. K. BHATTACHARYYA CONTENTS Page 1. INTRODUCTION .......... 134 2. METHODS ........... 135 3. EXTERNAL MORPHOLOGY ........ 135 4. CLASSIFICATION .......... 145 KEY TO THE SPECIES OF BRITISH MITES OF THE GENUS Pergamasus BERLESE s. LAT. ......... 148 Pergamasus (Pergamasus} crassipes (L.) Berlese . . . 151 longicornis Berlese .... 155 septentrionalis (Oudemans) . . 159 quisquiliarum (Canestrini) . . . 163 mirabilis Willmann . . . . 166 hamatus (Koch) . . . . 169 hortensis sp. nov. . . . . 173 Pergamasus (Paragamasus) robustus (Oudemans) . . . 174 alpestris Berlese . . . . 178 alstoni sp. nov. . . . . 181 armatus Halbert . . . . 183 cambriensis sp. nov. . . . 186 cassiteridum sp. nov. . . . 188 celticus sp. nov. .... 192 diversus Halbert .... 192 femoratus sp. nov. .... 197 integer sp. nov. .... 197 lapponicus Tragardh . . . 200 leruthi Cooreman .... 204 londonensis sp. nov. .... 207 longisetosus sp. nov. . . . 207 minimus sp. nov. . . . . 211 misellus Berlese . . . . 211 nathistmus sp. nov. . . . . 214 parrunciger sp. nov. . . . 216 rothamstedensis sp. nov. . . . 219 runciger (Berlese) .... 222 schweizeri sp. nov. .... 225 suecicus (Tragardh) . . . 228 teutonicus Willmann . . . 231 truncus Schweizer . . . . 234 wasmanni (Oudemans) . . . 236 RECORDS OF OTHER BRITISH SPECIES OF Pergamasus BERL. s. LAT. 239 5. SUMMARY ........... 240 6. ACKNOWLEDGEMENTS ......... 240 7. REFERENCES .......... 241 134 s - K - BHATTACHARYYA INTRODUCTION MITES of the family Parasitidae, in its present concept, were included in the genus Gamasus Latr. until Berlese (1892) subdivided the genus into three subgenera, Gamasus s. str., Eugamasus Berl. and Hologamasus Berl. This division was based primarily on the form of the sclerotization of the idiosoma in the adult female and the nature of the gnathosomal corniculi in the male. In 1904, Berlese proposed three other subgenera, namely, Trachygamasus (type Gamasus pusillus Berl.), Pergamasus (type Acarus crassipes Linn.) and Amblygamasus (type Gamasus tiberinus Can.) although these were not defined until Berlese (1906) published his " Monografia del Genere Gamasus ". The genus Trachygamasus was characterized by the absence of metasternal shields in the female (actually the metasternals are completely fused with each other and the sternal shields) while the two other sub- genera were essentially based on the armature of the second pair of legs in the male. Berlese's concepts of the classification of the 'Gamasus' -complex have been subject to little revision subsequently although each of his subgenera has, at one time or another, been given generic rank. Hull (1918) for example, recognized Berlese's groupings but gave generic rank to Ologamasus, Pergamasus and Gamasus, whilst he considered Amblygamasus to be a subgenus of Pergamasus, and Eugamasus a subgenus of Gamasus. This author also proposed a further division of Pergamasus and erected the subgenera Paragamasus and Plesiogamasus. Since Berlese's monograph four genera have also been added to the family, namely, Oocarpais Berlese, 1916, Parasitellus Willmann, 1939, Gamasodes Oudemans, 1939 and Pergamasellus Evans, 1957. Certain nomenclature changes have also been necessary, such as Parasitus Latr. having priority over Gamasus Latr. and Holoparasitus Oudemans 1936 being proposed as a new name for Ologamasus Berlese, 1892 non Berlese, 1888. The contributions to the knowledge of the Parasitidae within the past forty years have largely been in the form of descriptions of new species with little or no attempt at critical revisions at generic level. European workers have been particularly prominent in the field, new species having been described, amongst others, by Oudemans (1902, 1904, 1912, and 1926), Voigts and Oudemans (1904), Bonnet (1911), Hull (1916 and 1918), Halbert (1915), Tragardh (1910 and 1936), Vitzthum (1926), Willmann (1932, 1938, 1939, 1940, 1949, 1951, 1953, 1954 and 1956), Sellnick (1929, 1940), Pax & Willmann (1937), Leitner (1946 & I946a), Cooreman (1943, 1951), Schmolzer (1953), Halaskova (1959) and Schweizer (1961). Schweizer's key to the Parasitidae of Switzerland, published posthumously in 1961, has been the only attempt at a complete regional revision of the family since Hull (1918). This work presents a revision of the British mites of the genus Pergamasus Berl. s. lat. Members of this genus are amongst the most important acarine predators in Palaearctic soils although they appear to be replaced by the Rhodacaridae in tropical soils (Evans, in litt.}. The confusion concerning the identity of the species of this relatively large genus limits the scope of ecological work on the role played by these mites in the arthropod community of the soil. Although Turk (1953) lists twenty- REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 135 four species and varieties of Pergamasus (including Amblygamasus, Paragamasus and Plesiogamasus) for the British Isles only a small percentage of these can be identified with certainty. It is intended that this work will remedy, in part, this confusing situation and also, by the detailed study of the external morphology of the species, provide new criteria for their classification. The type material is deposited in the Collections of the British Museum (Natural History) . METHODS This revision is largely based on identified and unidentified specimens of Pergamasus in the Collections of the British Museum (Natural History). This material was supplemented by Berlese-Tullgren extractions of litter and other organic debris made by the author. The specimens were prepared for microscopical study by clearing them in 60% lactic acid. All the figures have been made from temporary preparations in lactic acid. Structures requiring detailed study, for example leg II of the male, chelicerae, pedipalps and the genital shield of the female, were dissected and prepared separately. Specimens were orientated in cavity slides for studying structures situated laterally, for example the peritreme and peritrematal shield. The three species cultured under laboratory conditions were confined in small petri dishes containing a charcoal and Plaster of Paris base of the type used by Bhattacharyya (1962). They fed on various species of Collembola. EXTERNAL MORPHOLOGY Chelicerae The chelicerae are three-segmented with the basal segment short, the second segment long and terminating in the fixed digit, and the third segment represented by the movable digit. Both digits are invariably dentate. The movable digit in the male and females of the Paragamasus-group of species bears one, two or three well- defined teeth (Text-figs. 185, 228) whilst in the Pergamasus-group the dentition is more variable. The pilus dentilis is simple in all species examined. The shape and number of teeth on the fixed digit of both groups shows considerable interspecific variation; the number of teeth ranging from one to several. A simple (Text-fig. 256), comb-like or spatulate (Text-fig. 106) dorsal seta is present as well as two lyriform fissures. The fixed digit in certain males is truncated distally, for example in Pergamasus runciger Berlese (Text-fig. 262) and P. diversus Halbert (Text-fig. 166). The spermadactyl is simple and fused with the movable digit distally (Text-fig. 206) . The processes of the arthrodial membrane at the base of the movable digit may be simple or branched (Text-figs. 247 & 185) ; branched processes being rare in the females. The ontogenetic development of the dentition of the digits of the chelicerae in Pergamasus robustus (Oudems.) is shown in Text-figures I, 2 and 3. It will be noted that in P. robustus the dentition of the movable digit is constant throughout the 136 S. K. BHATTACHARYYA FIGS. 1-5. Figs. 1-3, Chelicerae of the larva (fig. i), protonymph (fig. 2), and deutonymph (fig. 3) of Pergamasus (Paragamasus) robustus (Oudemans). Figs. 4 and 5. Gnathosoma of the larva (fig. 4) in Pergamasus (Pergamasus) septentrionalis (Oudemans) and (fig. 5) in Pergamasus (Paragamasus) robustus (Oudemans). REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 137 post-embryonic developmental stages. In P. septentrionalis (Oudems.), on the other hand, there is a progressive addition of teeth to the digit up to the deutonymph. The dorsal seta is constant in form throughout ontogeny. FIGS. 6-10. Figs. 6-8 : Gnathosoma of the protonymph (fig. 6) of Pergamasus (Para- gamasus) robustus (Oudemans), protonymph (fig. 7) of Pergamasus (Pergamasus) septentrionalis (Oudemans) and deutonymph (fig. 8) of Pergamasus (Paragamasus) robustus. Figs. 9-10 : Tectum of the protonymph (fig. 9) and deutonymph (fig. 10) of P. (P.) robustus. 138 S. K. BHATTACHARYYA Gnathosomal base, hypostome, tectum and pedipalps The gnathosomal base and hypostome are well developed. The chaetotaxy of the venter of the hypostome comprises three setae in the larva, one of which has hyper- trophied to form horn-like corniculi. The other two setae (gs.i and gs.2) are simple or slightly pilose (Text-figs. 4 & 5). In the protonymph two pairs of setae are added, namely gs.3 on the venter of the hypostome and gs.4 on the gnathosomal base (Text-figs. 6 & 7). This chaetotaxy is retained in the deutonymph (Text-fig. 8) and adult (Evans, 1957). Setae gs.2 and gs.3 normally form a more or less transverse row between gs.i and gs.4 ; the exception occurs in some males, where gs.3 lies anterior to gs.2. The corniculi in females and immature stages are sessile but in males they are borne on stalk-like projections of the hypostome which bear the hypostomal setae (Text-fig. 102). The hypostomal processes are strongly developed and show interspecific differences. Their external margins are invariably fringed with setae-like processes. The deutosternum in the adults bears eight to fifteen transverse rows of denticles. There are no deutosternal denticles in the larva. With few exceptions the number of rows of denticles is constant in the protonymph, deutonymph and adult of a given species. The tectum in the adults is usually produced into three distinct prongs although this number may be increased to five or more in the neotrichous species. The form of the tectum is often markedly dissimilar in different post-embryonic stages of a species (Text-figs. 9 & 10). The pedipalp has five free segments excluding the apotele which is represented by a three-tined seta-like structure on the inner basal angle of the tarsus. The chaetotaxy of the palp is an important taxonomic criterion both as regards the number of setae and their form. The palpal chaetotactic formulae are as follows (Text-figs. 11-13) : ~ Larva (0-4-5-1 2-1 i-apotele) . Protonymph (1-4-5-12-1 5-apotele) . Deutonymph and adult (2-5-6-14-1 5-apotele). In certain males the palp trochanter has one or two ventral protuberances ; the larger protuberance always bears a seta. Forms with two protuberances have the proximal one considerably smaller than the distal (Text-fig. 195). The seta on the internal (anterior) face of the femur is always comb-like whereas the two setae on the internal face of the genu may be spatulate or comb-like (Text-figs. 227 & 64). The other setae of these two segments may be simple or pilose. Idiosoma DORSUM The dorsum of the larva has a well-defined podonotal shield bearing the normal nine pairs of setae (ii, i2, 13, i4, i5, zi, Z2, si and 55). The unsclerotized cuticle posterior to the podonotal shield also carries nine setae comprising J2-J5, 57, 82-85 (Text-fig. 14). In the protonymph, there are fifteen setae (including 57) in the podonotal region of which eleven setae are situated on the podonotal shield. The REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 139 FIGS. 11-15. Fig 3 - II - I 3 : Pedipalp of the larva (fig. n), protonymph (fig. 12), and deutonymph (fig. 13) of Pergamasus (Paragamasus) robustus (Oudemans). Figs. 14 and 15 : Dorsum of the larva (fig. 14) and protonymph (fig. 15) of Pergamasus (Para- gamasus) robustus. setae added to this region of the dorsum in the protonymph are : si, s6, 14, r5 and ry (Text-fig. 15). Fourteen pairs of setae occur on the opisthonotal region of which four or five pairs may be situated on a small, weakly sclerotized opisthonotal shield (Text-fig. 15). Setae Ji, Zi-Z4 and Ri are added at this stage. Mesonotal scutellae may be present or absent. I have observed three types of chaetotaxy of the podonotal region in deutonymphs depending on the number of setae in the r series. In P. robustus twenty-one setae 140 S. K. BHATTACHARYYA FIGS. 16-18. Dorsum of the deutonymph of Pergamasus (Paragamasus) robushis (Oudemans) (fig. 16), Pergamasus (Pergamasus) crassipes (L.) Berlese (fig. 17) and Pergamasus (Pergamasus) septentrionalis (Oudemans) (fig. 18). REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 141 are present in this region with the r series (n-ry) complete (Text-fig. 16). P. crassipes, on the other hand, has nineteen setae, r2 and r3 being absent (Text-fig. 17) and P. septentrionalis only seventeen setae, 12, r3, r5 and ij being absent (Text-fig. 18). There are probably more than these three types I have observed, particularly amongst the species showing neotrichy of the podonotal region in the adult, for example Pergamasus mirabilis Willmann and Pergamasus hamatus (Koch), but I have not seen deutonymphs of these species. The deutonymphal opisthonotal shield is invariably well-defined and in the Paragamasus-group bears twelve or thirteen pairs of setae. The thirteen setae present on the opisthonotal shield of P. robustus, for example, are Ji-j5, Zi-Z/j., 81-83 and Ri (Text-fig. 16). Neotrichy affects the unsclerotized cuticle of the opisthonotal region and the opisthonotal shield of members of the Pergamasus-group (Text-figs. 17 & 18) to such an extent that it is not possible to distinguish the primary setae. The dorsal chaetotaxy of the deutonymph of the Paragamasus group is retained by the adult and there is a fusion of the podonotal and opisthonotal elements to form an entire dorsal shield (Text-fig. 103). In the males of some species the line of fusion of these two sclerites is discernible (Text-fig. 96). The fusion of the deutonymphal podonotal and opisthonotal shield also occurs in the neotrichous Pergamasus-gToup and setae appear to be added to the dorsum at the deutonymphal ecdysis. The neotrichy is usually restricted to the opisthonotal region although in P. hamatus and P. mirabilis the podonotal region is also involved (Text-figs. 88 &76). The surface of the dorsal shield or shields is usually regionally reticulated and provided with numerous pore-like structures. The dorsal setae are usually simple although some pilose setae occur in immature and adult stages. VENTER Larva : The tritosternum in the larva has a rectangular base, longer than wide and a pair of pilose laciniae (Text-fig. 19). The sternal region bears three pairs of simple or pilose setae but there is no definite indication of a sternal shield. All the species have a well-defined anal shield with the normal complement of setae, namely, a pair of paranals and a post-anal seta. These setae are invariably long. Each anal valve is provided with an euanal seta. The remainder of the opisthogastric region carries four pairs of setae distributed as in the Text-figure 19. There are no stigmata or peritr ernes. Protonymph : The tritosternum is essentially the same as in the larva. The inter-coxal region bears four pairs of setae of which the anterior three pairs, homo- logous with the three sternal setae in the larva, are situated on a sternal shield provided with two pairs of lyriform fissures (Text-fig. 20). The remaining pair of setae situated between coxae IV is homologous with the genital setae of the adult. The anal shield bears the three setae present in the larva but these are considerably shorter in length. There are no euanal setae. The chaetotaxy of the remainder of the opisthogastric region shows an increase of one pair of setae which is located lateral to the anal shield. A pair of stigmata and short peritremes are present. I 4 2 S. K. BHATTACHARYYA FIGS. 19-22. Venter of the larva (fig. 19) of Pergamasus (Paragamasus) robustus (Oudemans), protonymph (fig. 20) of PeYgamasus (Pergamasus) crassipes (L.) Berlese, deutonymph (fig. 21) of P. (P.) robustus and deutonymph (fig. 22) of P. (P.) crassipes. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 143 Deutonymph : The tritosternum is similar to that of the protonymph but is invariably flanked by one or more pairs of prae-endopodal shields. The inter-coxal region carries five pairs of setae ; those additional to the protonymphal complement being setae IV (the metasternals) . Sternal setae I-IV are situated on a well- sclerotized sternal shield (Text-fig. 21). The anal shield is essentially the same as in the protonymph although in the Pergamasus-group some of the opisthogastric setae may be located on the shield. In the Paraga masus-group, unsclerotized cuticle of the opisthogastric region bears eight, nine or ten pairs of setae (Text-fig. 21), but in the Pergamasus-group neotrichy occurs particularly in the posterior region of the opisthogaster (Text-fig. 22). Both groups have well developed stigmata, peritremes and peritrematal shields. The length of the peritreme varies interspecifically. Small sclerites are often present on the opisthogaster. Female : The tritosternum resembles that of the immature stages although the pilosity of the laciniae may be indistinct in some species. The prae-endopodal shields are well-defined in all the species examined but there is considerable inter- specific variation in their shape, size and number. All species have a sternal shield bearing three pairs of setae and two pairs of lyriform fissures. The sternal shield is fused with the exopodal shields between coxae I and II and with the endopodal shields in the region of coxae II and III. Its posterior margin is often deeply incised medially and its surface may be variously ornamentated with a network of lines or punctures. The metasternal setae are situated on large discrete metasternal shields which are fused with the endopodal elements in the region of coxae IV and flank the triangular genital shield. Rarely, the metasternal shields are weakly fused along the median line. They bear a pair of lyriform fissures. The genital shield is large and with the exception of Paragamasus suecicus (Tragardh) (Plate 7!) is broadly triangular in outline. It bears a pair of genital setae. The anterior portion of the shield is strengthened by more heavily sclerotized areas whose outlines are of taxonomic importance (Plate yb and Plate 8d). The genital shield overlies the genital orifice and a pair of sclerites (paragynials) referred to as " nymphae " by Berlese (1906). The wall of the vagina is variously sclerotized, the sclerotization being in the form of endogynial processes (Text-fig. 70, 118). Some species also have a pair of horn-like structures extending beneath the meta- sternal shields. The opisthogastric shield is large and is fused with the podal elements in the region of coxae IV. The anus is subterminal and paired paranal and the post-anal setae are always present. In the Paragamasus-group (Text-figs. 264, 217) the opisthogastric shield bears eight, nine or ten (Text-figs. 213, 123, 117) pairs of setae excluding those associated with the anus, whereas the number is markedly increased in the neotrichous species of the Pergamasus-group (Text-figs. 36 & 57). The stigmata and peritremes are conspicuous ; the length of the peritreme shows interspecific variation. The peritrematal shield shows varying degrees of fusion with the dorsal shield and the opisthogastric shield. In Pergamasus crassipes (Linn.), P. longicornis (Berl.), P. septentrionalis (Oudems.) and P. quisquiliarum (Can.), the shield is completely fused with the dorsal shield (Text-fig. 23). In I 4 4 S. K. BHATTACHARYYA 25 26 FIGS. 23-26. Peritrematal shields of females. Fig. 23 Pergamasus (Pergamasus) longicornis Berlese, fig. 24 Pergamasus (Pergamasus) hamatus (Koch), fig. 25 Pergamasus (Paragamasus) diver sus Halbert and fig. 26 Pergamasus (Paragamasus} alstoni sp. nov. P. hamatus (Koch) and P. hortensis sp. nov. the extent of fusion with the dorsal shield is less ; the shield being free in its posterior half (Text-fig. 24) . In the Paragamasus- group the shield is free (Text-fig. 25) or fused with the opisthogastric shield posteriorly (Text-fig. 26). The unsclerotized cuticle of the opisthogastric region is provided with a variable number of setae. Male : The dorsal, lateral and ventral surfaces of the idiosoma in the male are completely sclerotized except at the junction of the exo- and endopodal shields, between coxae II and III, and III and IV. The genital orifice is situated immediately anterior to the sternal shield and its ventral wall is strongly sclerotized (Text-fig. 27) . The orifice is protected by a genital lamina whose anterior margin may be produced into one or two processes. The lamina is connected by a pair of elongate apodemes to strong retractor muscles and it is assumed that the lamina is capable of being withdrawn into the orifice. The base of the tritosternum, covered by the lamina, is strongly reduced but the pilose laciniae resemble those of the female. Prae- endopodal shields are invariably present. The chaetotaxy of the venter of the male resembles that of the females of the same species. Sternal seta 5 (the genital) may be situated near (P. diversus, P. alpestris) or on (P. lapponicus, P. teutonicus) scale- like tubercles (Text-figs. 163, 193). The peritrematal shields (including the stigmata and peritremes) are entirely fused with the dorsal shield and the exopodals. A post- stigmatic seta may or may not be present. REVISION OF THE GENUS PERGA MASUS BERLESE S. LAT. 145 27 28 FIGS. 27-28. Pergamasus (Paragamasus) lapponicus Tragardh, male. Fig. 27 internal view of the sterniti-genital region. Fig. 28 tarsus and prae-tarsus of leg I. Legs The legs are six segmented (excluding the ambulacrum) in all post-embryonic developmental stages. Lyriform fissures occur in the basal half of the femora and tarsi, and result in false divisions of these segments. All the segments of the legs have a well-defined chaetotaxy which has been studied in detail by Evans (1963) whose terminology I have adopted. The chaetotaxy of each instar shows little or no intra or interspecific variation in setal complement although the relative lengths and form of the setae may vary. Leg II in the male is specialised for grasping the female during mating which, as in the majority of the Mesostigmata, is an indirect method involving a spermatophore. Leg II is often enlarged (crassate) and two setae (v 2 and v 3 ) on the femur and one on the genu (av x ) and the tibia (av x ) hypertrophy and form spur-like structures. The form of the spurs on these segments is of taxo- nomic importance. A dorsal seta on the basal half of tarsus IV is invariably long and erect. The ambulacrum comprises a lobed pulvillus and two claws (Text-fig. 28). CLASSIFICATION The Paragamasus-group is at present, divided into four subgenera. According to Hull (1918) these subgenera may be characterized as follows: i ' Body of both sexes piriform, rostrum very prominent, cuticle polished and smooth, Femur II of male crassate but spurless. Colour ruddy brown '........ Amblygamasus Berlese. 146 S. K. BHATTACHARYYA 2 ' Body of both sexes piriform, cuticle conspicuously reticulate and rather rough. Femur II crassate with a strong falcate spur ; patella with a prominent apical spur or branch projecting inwards and forwards. Colour ruddy brown, rather dull ' .... Pergamasus s. str. 3 ' Body oblong, more or less parallel sided in both sexes. Ped II as in (2) but without the patellar spur. Colour pale yellowish brown ' Paragamasus Hull. 4 ' Body of male narrow, oblong, of female rather piriform. Ped II of the male only slightly crassate all spurs more or less cylindrical. Colour very pale ' Plesiogamasus Hull. The validity of Hull's subdivision of the genus must be assessed on the basis of the characters associated with leg II of the male since those based on the shape of the idiosoma and its colour are less reliable ; being subjected to considerable intra- specific variation. His reference to the armature of the patella is confusing. Assuming that his " patella " refers to the genual segment, then the prominent apical spur on leg II of P. crassipes (the type of the subgenus Pergamasus) is not located on that segment but on the tibia whereas in P. alpestris, which Hull includes in Pergamasus s. str., the characteristic L-shaped process is present on the genu. Thus, Hull's use of the armature of the patella in separating Pergamasus and Paragamasus has no foundation. Further, the statements that members of Paragamasus have no patellar spur is also inaccurate, the only British species of the genus without a spur on genu II are P. quisquiliarum and P. diversus and Hull considered that the latter should probably be placed in Pergamasus s. str. The type species of Paragamasus Hull (type Pergamasus robustus (Oudemans)) and Plesiogamasus Hull (type Pergamasus hamatus (Koch) sensu Berlese, 1905) were designated by Turk & Turk (1952). Within the limited number of species (thirty-two) I have examined the groupings formed on the basis of various combinations of external morphological characters indicate an intrageneric rather than an intergeneric relationship between the groups. The emphasis laid on the characters associated with leg II in the male in the classifica- tion at subgeneric level appears to be unwarranted. The nature of the armature of leg II does not appear to be correlated with any other distinctive morphological character and the degree of development of spurs (hypertrophied setae) and sclerotized protuberances show every gradation in form, from the weakly developed nodular spurs of P. nathistmus sp. nov. to the massive spurs and protuberances of P. robustus and P. crassipes. The only character I have found which will give a reliable practical division of the genus is the chaetotaxy of the dorsum and the opisthogastric region of the idiosoma. Two groups are readily definable ; one in which the chaetotaxy displays neotrichy and the other in which neotrichy is not apparent. This character has the advantage over characters restricted to the male or female in being determined at the deuto- nymphal stage and of applying to both sexes. I have, provisionally, included the neotrichous species in the subgenus Pergamasus Berl. s. str. and the non-neotrichous species in Paragamasus Hull. These subgenera have the following characteristics : REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 147 Family PARASITIDAE Oudemans Genus PERGAMASUS Berlese, 1904 s. lat. Type : Pergamasus crassipes (Linn.) Berlese, 1906. Subgenus Pergamasus s. str. (including Amblygamasus Berl. and Plesiogamasus Hull). The chaetotaxy of the dorsum and of the opisthogastric region of the idiosoma displays neotrichy in all the members of this subgenus. With the exception of P. (P.) hamatus, neotrichy of the dorsal chaetotaxy is restricted to the opisthosomal region. Two distinct groups of species may be recognized on the basis of female characteristics, namely : the crassipes-group (P. (P.) crassipes and P. (P.) longi- cornis ; P. (P.) septentrionalis and P. (P.) quisquiliarum) in which the peritrematal shield is entirely fused with the dorsal shield in the region of coxae II-IV and the tectum is essentially five-pronged ; and the hamatus-group (P. (P.) hamatus and P. (P.) hortensis) in which the peritrematal shield is free posteriorly and the tectum is essentially three-pronged. The female of P. (P.) mirabilis is not known. Members of the crass^s-group are large (up to i,383|x), strongly sclerotized species ; the males of which may be readily distinguished by the armature of leg II and the form of the internal setae of the genu of the pedipalp and of the dorsal seta of the chelicera. The form of the sclerotized structures of the endogynium is characteristic for the females. The seven British species which I have assigned to this subgenus are keyed below. Subgenus Paragamasus Hull, 1918 (including Leptogamasus Tra.ga.rdh) Type : Pergamasus robustus (Oudemans, 1902) The majority of the British species of Pergamasus fall into this subgenus. There are never more than thirty-five pairs of setae on the opisthonotal region of the dorsal shield and the setae of the opisthogastric shield in the female number eight, nine or ten (excluding the three setae associated with the anus). The species range in length from 400-1, 200(x. The dorsal shield in the male often shows an incomplete fusion of the podonotal and opisthonotal elements ; their juncture being indicated by a transverse suture. The tectum is basically three-pronged throughout. The separation of the males has been made largely on the armature of leg II, the form and chaetotaxy of the trochanter of the pedipalp, the shape and dentition of the chelicera and the chaetotaxy of the dorsum. The females show greater morphological homogeneity and although it has been possible to divide them into groups on the basis of such characters as the fusion of the peritrematal shield with the opistho- gastric shield, the number of setae on the opisthogastric shield and the chaetotaxy of the dorsal shield, specific diagnosis in a number of cases depends on the form of the genital shield and the endogynial processes. The characters associated with the genital shield and endogynium are so difficult to define for key purposes that for identification purposes it will be necessary to remove the genital shield and compare its structure with the illustrations. 148 S. K. BHATTACHARYYA KEY TO ADULTS OF THE BRITISH MITES OF THE GENUS Pergamasus BERL. s. LAT. MALES 1 Opisthonotal region with a dense covering of setae, more than 45 pairs (Text-fig. 82) (Subgenus Pergamasus) 2 Opisthonotal region with a moderate covering of setae, not more than 35 pairs (Text-fig. 152) . . . . . . , (Subgenus Paragamasus) 7 2 Chaetotaxy of the podonotal region displaying neotrichy, more than 45 pairs of setae present (Text-fig. 82) ; femur II with two spurs (Plate 2c) ; dorsal shield with a distinct transverse suture indicating the juncture of the podonotal and Opisthonotal shields . . . .P. (Pergamasus) hamatus (Koch) (p. 169) - Chaetotaxy of the podonotal region not displaying neotrichy, 25-27 pairs of setae present (Text-fig. 76) ; femur II with at the most a single spur (Plate la) . . 3 3 Dorsum with a transverse suture (Text-fig. 76) ; coxa II with processes (Plate 2a) ; tibia II unarmed (Plate 2b) ; prae-endopodal shields almost meeting in the mid- line ; genital lamina weakly developed . P. (Pergamasus) mirabilis Willmann (p. 166) - Dorsum without a transverse suture (Text-fig. 50) ; coxa II unarmed ; tibia II armed with one or more spur-like processes ; prae-endopodal shields widely separated ............. 4 4 Internal setae of the genu of the pedipalp spatulate, entire ; tibia of leg II with a large ventro-lateral process (Plate ib) ; dorsal seta of the chelicera simple . 5 - Internal setae of the genu of the pedipalp denticulate distally (Text-fig. 64) ; tibia II without such large process (Plate xe) ; dorsal seta of the chelicera denticulate distally (Text-fig. 65) 6 5 Ventro-lateral process of tibia II hook-like distally (Plate ib) P. (Pergamasus) longicornis Berlese (p. 155) - Ventro-lateral process of tibia II rounded distally (Plate la) P. (Pergamasus) crassipes (L) Berlese (p. 151) 6 Genu II strongly spurred (Plate ic) ; femur II without spur ; genital lamina produced into a median spine (Text-fig. 51) P. (Pergamasus) septentrionalis (Oudemans) (p. 159) Genu II unarmed (Plate id) ; femur II with a truncated spur ; genital lamina may be medially incised (Text-fig. 62) P. (Pergamasus) quisquiliarum (Canestrini) (p. 163) 7 Dorsum entire, without transverse suture (Text-fig. 235) ..... 8 Dorsum with a transverse suture (Text-fig. 239) ...... 13 8 Coxa and trochanter of leg II tuberculated (Plate 20 & i) ; movable digit of the chelicera bidentate (Text-fig. 113), fixed digit truncate distally ; with scale-like elevations near sternal setae V (Text-fig, no) P. (Paragamasus) alpestris Berlese (p. 178) - Coxa and trochanter of leg II not tuberculated ; movable digit of the chelicera uni- or bidentate, fixed digit rounded distally (Text-fig. 251) without scale-like elevations near sternal setae V . . . . . . . ; 9 9 Dorsal shield greater than 700^ in length . . . . . . ... 10 Dorsal shield less than 500 [A in length . . . . . . . 1 1 10 Dorsal seta RS about twice the length of 84 (Text-fig. 203) : spur on tibia II in the form of a ridge (Plate 46) . . P. (Paragamasus) leruthi Cooreman (p. 204) - Dorsal setae R$ and 84 subequal in length (Text-fig. 181) ; spur on tibia II digitiform (Plate 4b) ..... P. (Paragamasus) integer sp. nov. (p. 197) 1 1 Fixed digits of the chelicera tapering distally, not hooked ; movable digit unidentate (Text-fig. 282) . . . .P. (Paragamasus) suecicus (Tragardh) (p. 228) - Fixed digit of chelicera hooked distally ; movable digit bidentate (Text-fig. 251) . 12 REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 149 12 Spurs on femur II minute, main spur scarcely longer than the accessory spur (Plate 5b) ...... P. (Paragamasus) nathistmus sp. nov. (p. 214) - Spurs on femur II larger, main spur about twice the length of the accessory spur (Plate 5f) . . . .P. (Paragamasus) rothamstedensis sp. nov. (p. 219) 13 Corniculus with triangular hyaline appendage (Text-fig. 167) ; genu II unarmed (Plate 3b) ..... P. (Paragamasus) diversus Halbert (p. 192) Corniculus without such appendage ; genu II armed with a spur . . . 14 14 Trochanter of pedipalp without seta-bearing tubercle (Text-fig. 99) . . 15 Trochanter of pedipalp with a strong seta-bearing proximal tubercle (Text-fig. 195) 16 15 Trochanter II spurred (Plate 2d) . P. (Paragamasus) robustus (Oudemans) (p. 174) - Trochanter II not spurred ; leg II figured (Plate 6d) P. (Paragamasus) truncus Schweizer (p. 234) 16 Sternal seta V arising from a scale-like elevation (Text-fig. 193) . . . . 17 - Sternal seta V not arising from a scale-like elevation . . . . . . 18 17 Trochanter II with a strong spur-like projection (Plate 4c) ; leg II (Plate 4d) ; trochanter IV with process (Text-fig. 197) P. (Paragamasus) lapponicus Tragardh (p. 200) Trochanter II without such projection ; leg II (Plate 6c) ; trochanter IV without process .... P. (Paragamasus) teutonicus Willmann (p. 231) 18 Femur II with a conspicuous tubercle proximal to the lyriform fissure (Plate 4a) ; anterior rows of deutosternal denticles strongly arched (Text-fig. 180) ; leg II figured (Plate 4a) ... P. (Paragamasus) femoratus sp. nov. (p. 197) Femur II without tubercle proximal to lyriform fissure ..... 19 19 Dorsal shield 500^ or less in length ; main spur on femur II considerably shorter than the length of tibia II (Plate 6d) ........ 20 Dorsal shield JOO\L or greater in length ; main spur on femur II about equal in length to or longer than tibia II (Plate 3g) ....... 22 20 Femur II with a strong sclerotized ridge proximal to the main spur (Plate 6e) ; spur on tibia slender . . P. (Paragamasus) wasmanni (Oudemans) (p. 237) Femur II without sclerotized ridge proximal to the main spur (Plate 5a) ; spur on tibia II broad . . . . . . . . . . .21 21 Proximal seta on the trochanter of the pedipalp about equal to the height of the tubercle (Text-fig. 138) ; leg II (Plate 3a) P. (Paragamasus) cambriensis sp. nov. (p. 186) Length of the proximal seta on the trochanter of the pedipalp about twice the height of the tubercle (Text-fig. 227) ; leg II (Plate 5a) P. (Paragamasus) misellus Berlese (p. 211) 22 Accessory spur on femur II broad, truncate (Plate 5c) ..... 23 Accessory spur on femur II slender, tapering distally (Plate 6a) .... 25 23 Main spur on femur II slender without tubercle (Plate 3d) ; trochanter II with sclerotized process distally (Plate 3c) ; leg II (Plate 3d) P. (Paragamasus) cassiteridum sp. nov. (p. 188) Main spur on femur II with distinct tubercle (Plate 5g) ; trochanter II without sclerotized process distally .......... 24 24 Trochanter of pedipalp with a distinct tubercle proximal to the seta-bearing tubercle (Text-fig. 242) ; leg II (Plate 5c) . P. (Paragamasus) parrunciger sp. nov. (p. 216) Trochanter of pedipalp without tubercle proximal to the seta-bearing tubercle (Text-fig. 261) ; leg II (Plate 5g) . P. (Paragamus) runciger (Berlese) (p. 222) 25 Fixed digit of the chelicera markedly broad anterior to the lyriform fissure (Text- fig. 156) ; leg II (Plate 3g) . . .P. (Paragamasus) celticus sp. nov. (p. 192) - Fixed digit of the chelicera relatively slender (Text-fig. 131) ; leg II (Plate 2g) . 26 150 S. K. BHATTACHARYYA 26 Trochanter of pedipalp with a distinct tubercle proximal to the seta-bearing tubercle (Text-fig. 270) ; spurs on leg II as in Plate 6a P. (Paragatnasus) schweizeri sp. nov. (p. 225) - Trochanter of pedipalp without tubercle proximal to the seta-bearing tubercle (Text-fig. 130) ; spurs on leg II as in Plate 2g P. (Paragatnasus) armatus Halbert (p. 183) FEMALES 1 Opisthonotal region of dorsal shield with a moderate covering of setae, never more than 35 pairs (Text-fig. 134) ; peritrematal shield free posteriorly or fused with the opisthogastric shield (Text-figs. 141 & 123) ; opisthogastric shield with a maximum of ten pairs of setae, excluding the three setae associated with the anus ........ (Subgenus Paragamasus) 7 Opisthonotal region of dorsal shield with a dense covering of setae, more than 45 pairs (Text-fig. 55) ; peritrematal shield fused with the dorsal shield along its entire length or free in its posterior half (Text-figs. 47 & 93) ; opisthogastric shield with never less than 14 pairs of setae (Text-fig. 47) (Subgenus Pergamasus) 2 2 Podonotal region of the dorsal shield neotrichous with more than 35 pairs of setae (Text-fig. 88) ; peritrematal shield free in its posterior half (Text-fig. 24) ; tectum three-pronged (Text-fig. 90) . P. (Pergamasus) hamatus (Koch) (p. 169) - Podonotal region of the dorsal shield with never more than 25 pairs of setae (Text- fig- 35) I peritrematal shield fused with the dorsal shield along its entire length or free in its posterior half (Text-figs. 36 & 93) ; tectum basically three- or five- pronged (Text-figs. 95 & 72) ......... 3 3 Tectum basically three-pronged (Text-fig. 95), peritrematal shield free in its posterior half (Text-fig. 93) ; endogynium characteristic (Text-fig. 94) ; dorsal shield i,2oofz in length . . . . .P. (Pergamasus) hortensis sp. nov. (p. 173) Tectum basically five-pronged (Text-fig. 38) ; peritrematal shield fused with the dorsal shield along its entire length ; large species, greater than 1,200(1 in length 4 4 Endogynium with a pair of large horn-like structures (Text-fig. 56) ; no reticulated endogynial sacs ; trochanter IV with a sclerotized protuberance (Text-fig. 60) P. (Pergamasus) septentrionalis (Oudemans) (p. 159) Endogynium without paired horn-like structure but with a pair of reticulated sacs separated by a median process (Text-fig. 70) ; trochanter IV without sclerotized protuberance ............ 5 5 Internal setae of the genu of the pedipalp denticulate distally (Text-fig. 74) ; dorsal seta of the chelicera denticulate distally (Text-fig. 73) ; opisthogastric shield with twenty or more pairs of setae excluding the anals (Text-fig. 68) P. (Pergamasus) quisquiliarum (Can.) (p. 163) Internal setae of the genu of the pedipalp spatulate ; entire dorsal seta of the chelicera simple (Text-fig. 49) ; opisthogastric shield with about 15 pairs of setae (Text-fig. 36) .... . 6 6 Median process of endogynium strongly bifurcate distally (Text-fig. 47) ; genital shield as in Plate yb . . .P. (Pergamasus) longicornis Berl. (p. 155) - Median process of endogynium entire (Text-fig. 37) ; genital shield as in Plate ya P. (Pergamasus) crassipes (L) Berl. (p. 151) 7 Peritrematal shield fused with the opisthogastric shield posteriorly (Text-fig. 284) ; with 9 pairs of opisthogastric setae (Text-fig. 284) ...... Peritrematal shield free posteriorly (Text-fig. 25) ; with 8 or 10 pairs of opisthogastric setae (Text-figs. 254 & 275) . ..... . . . . . . 10 8 Genital shield rounded anteriorly (Plate 7!) P. (Paragamasus) suecicus Tragardh (p. 228) - Genital shield triangular anteriorly (Plate 7g) ...... REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 151 9 Endogynial processes as in Text-fig. 210 P. (Paragamasus) leruthi Cooreman (p. 204) Endogynial processes as in Text-fig. 124 . P. (Paragamasus) alstoni sp. nov. (p. 181) 10 Opisthogastric shield with 8 pairs of setae excluding anals (Text-fig. 254) . . u Opisthogastric shield with 10 pairs of setae excluding anals (Text-fig. 188) . . 12 11 Genital shield as in Plate yk . P. (Paragamasus) rothamstedensis sp. nov. (p. 219) - Genital shield as in Plate 7J . .P. (Paragamasus) londonensis sp. nov. (p. 207) 12 Dorsal setae R5 and 84 subequal in length (Text-fig. 187) ; genital shield and endogynial processes as in Plate 7i and Text-fig. 189 respectively P. (Paragamasus) integer sp. nov. (p. 197) - Dorsal seta R5 considerably longer than 84 (Text-figs. 231 & 302) . . . 13 13 Small species 450-550(0. in length ; peritreme extending to the middle of coxa II ; Genital shield as in Plate 8a . .P. (Paragamasus) misellus Berlese (p. 211) Genital shield as in Plate 8e . P. (Paragamasus) wasmanni (Oudemans) (p. 237) Genital shield as in Plate 8b . . P. (Paragamasus) truncus Schweizer (p. 234) Sterniti-genital region as in Text-fig. 222 P. (Paragamasus) minimus sp. nov. (p. 211) Genital shield as in Plate 8d .P. (Paragamasus) cambriensis sp. nov. (p. 186) - Large species 700-1, 200^. in length ; peritreme usually extending beyond coxa II . 14 14 Large species, dorsal shield 1,100-1,200^ in length . . . . . . 15 - Smaller species, dorsal shield usually 700-800^ in length (except P. (Paragamasus) longisetosus, 975-1,000^) .......... 16 15 Endogynial processes and genital shield as in Text-fig. 104 and Plate 70 respectively ; peritreme extending to the middle of coxa II P. (Paragamasus) robustus (Oudemans) (p. 174) - Endogynial processes and genital shield as in Text-fig. 118 and Plate 7f respectively P. (Paragamasus) alpestris Berlese (p. 178) 16 Dorsal setae conspicuously long and thick (Text-fig. 216) ; species 975-1,000^ in length . . . . .P. (Paragamasus) longisetosus sp. nov. (p. 207) - Dorsal setae relatively shorter and never thickened ; less than 900^ in length . 17 17 Posterior incision of the sternal shield extending to the level of sternal setae II (Text-fig. 170) ; genital shield with an anterior arrow-like sclerotized area (Plate 7h) P. (Paragamasus) diversus Halbert (p. 192) - Posterior incision of the sternal shield never reaching the level of sternal setae II ; genital shield without such sclerotized area a Genital shield as in Plate 8k . .P. (Paragamasus) runciger Berlese(p. 222) b Genital shield as in Plate 8j .P. (Paragamasus) parrunciger sp. nov. (p. 216) c Genital shield as in Plate 8c . . .P. (Paragamasus) armatus Halbert (p. 183) d Genital shield as in Plate 8h & i . P. (Paragamasus) lapponicus Trag. (p. 200) e Genital shield as in Plate 8m & n .P. (Paragamasus) teutonicus Willmann (p. 231) f Genital shield as in Plate 8f & g . .P. (Paragamasus) celticus sp. nov. (p. 192) g Genital shield as in Plate 81 .P. (Paragamasus) schweizeri sp. nov. (p. 225) Pergamasus (Pergamasus) crassipes (L) Berlese Gamasus (Pergamasus) crassipes : Berlese A (1906) Redia 3 : 229 (male only). Gamasus (Pergamasus) crassipes var. longicornis Berlese (1906) Redia 3 : 232 (female only). MALE. The male is ruddy brown in colour. The dorsal shield (1,268-1, 383^ in length x 747-864(0, wide) is pear-shaped, regionally reticulated, strongly sclerotized and without a transverse suture. The opisthosomal region is densely covered with setae (Text-fig. 29). Ventrally, there are two large prae-endopodal shields flanking the genital sclerite. The tritosternum has a pair of well developed pilose laciniae. About half of the 152 S. K. BHATTACHARYYA 31 34 FIGS. 29-34. Pergamasus (Pergamasus) crassipes (L.) Berlese, male. Fig. 29 dorsum of idiosoma. Fig. 30 venter. Fig. 31 tectum. Fig. 32 chelicera. Fig. 33 venter of gnathosoma. Fig. 34 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 153 tritosternum is covered by the genital lamina. The genital lamina is produced anteriorly into a sharply pointed spine. The arrangement of the sternal setae and pores are shown in Text-fig. 30. The stigma is situated ventro-laterally between coxae III and IV ; the peritreme extends to coxa I. The post-stigmatal extension of the peritrematal shield reaches the posterior region of coxa IV. A pair of post-stigmatal setae is present. The tectum is produced anteriorly into five short tooth-like processes (Text-fig. 31). The most distinctive features of the chaetotaxy of the pedipalp are, the comb-like seta on the femur and the spatulate setae on the genu. The palpal trochanter is without a tubercle. The chelicera is strong with a simple dorsal seta. The fixed digit is truncated. Both digits possess rows of teeth (Text-fig. 32). The sperma- dactyl is normal. The corniculus is distinctly stalked. The entire lateral (outer) margins of the hypostomal processes are fringed (Text-fig. 33). The hypostomal setae are pilose. The deutosternal teeth of the ventral groove are arranged in eleven transverse rows. The tarsus (333-360(0.) of leg I is longer than the tibia (254-266(0,). The prae- tarsus is 19-25(0, long. The femur of leg II is crassate with a strong falcate spur. The terminal end of the main process of the tibia is almost rounded and its distal spur is shown in Plate la. The trochanter of leg II is shown in Text-fig. 34. FEMALE. The dorsum of the dark brown, strongly sclerotized idiosoma is neotrichous (Text-fig. 35). The dorsal shield (1,200-1, 351(0, long x 782-917(0, wide) is oval and regionally reticulated. The tritosternum is well developed with a pair of pilose laciniae. There is a pair of prae-endopodal shields almost meeting in the medial line (Text-fig. 36). The metasternal shields are free. The endogynium is furnished with a pair of round faintly reticulated sacs with a short median process arising between them (Text-fig. 37). The terminal end of the process may be smooth or slightly serrated. The genital shield is shown in Plate 7a. The opisthogastric shield usually has fifteen pairs of setae of variable length (excluding the three setae associated with the anus) . Ventrally the stigma is situated a little behind coxa III and the peritreme extends to coxa I. The peritrematal shield is fused with the dorsal shield. The tectum is five-pronged (Text-fig. 38) . The pedipalp is essentially the same as in the male. The dentition of the fixed digit is well shown in Text-fig. 39. Fringes are confined to the proximal portion of the hypostomal processes (Text-fig. 40). The deutosternal denticles are arranged in twelve transverse rows. The gnatho- somal setae are distinctly pilose. Leg I has the tarsus (342-404(0.) longer than the tibia (241-291(0.) ; the prae- tarsus is 25-29(0, long. The trochanter of leg IV is without a process. DISTRIBUTION AND HABITAT. I have examined material from : In litter under Cupressus sp. (Rothamsted Lodge, Rothamsted Experimental Station, Harpenden, Hertfordshire, G. Owen Evans and E. Browning, i8.iii.i957). In leaf mould (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951). Oak and birch leaves (Newton Abbot, S. Devon, R. G. Lewesther, 21. vi. 1951). Litter and humus under pines and holly (summit of Dove Crag, near Ambleside, Westmorland, M. E. Bacchus, 154 S. K. BHATTACHARYYA FIGS. 35-40. Pergamasus (Pergamasus) crassipes (L.) Berlese, female. Fig. 35 dorsum of idiosoma. Fig. 36 venter. Fig. 37 endogynial process. Fig. 38 tectum. Fig. 39 chelicera. Fig. 40 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 155 28.ii. 1954). Sphagnum, etc. (Cock Hill, Yorkshire, J. T. Salmon, 21. vii. 1951). In leaf mould (Epping Forest, Essex, J. T. Salmon, 27. v. 1951). In leaf mould (Saver- nake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). In moss of wall (Tarn Hows, North Lancashire, M. E. Bacchus, 4 . xii . 1954) . In moss (Grassington, Grass Woods, North Riding, Yorkshire, J. T. Salmon, 24^1.1951). Sphagnum (Ascot, Berkshire, A. H. G. Alston, 8 . xii . 1957) . Leaf litter from woods (near Buxton, Derbyshire, J. T. Salmon, 23. vi. 1951). Moss on tree stumps and stones (Forest of Dean, Gloucester- shire, J. T. Salmon, 27. ix. 1951). In beech leaf mould from steep bank (Rickmans- worth, Hertfordshire, M. E. Bacchus, 25.11.1951). In leaf mould (Richmond Park, Surrey, J. T. Salmon, 19. ill. 1951). In leaf mould (Burnham Beeches, Buckingham- shire, J. T. Salmon, 3^1.1951). (Roy. Bot. Gdns., Kew, Surrey, J. L. Gilbert, .iii.igGi). Moss, humus, hollow tree (Swansea University, Swansea, P. N. Lawrence, 26 . iii . 1962). Sphagnum, etc. (Moel Siabod, North Wales, P. N. Lawrence, 22 . x . 1960). In moss (Tarbert, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3 . vii . 1951). In dead leaves (Loch Lomond between Glasgow and Tarbert, Dunbar- ton, J. T. Salmon, 8. vii. 1951). BIOLOGICAL NOTES. Eggs from four females kept under laboratory conditions hatched within one to three days (average 2-16 days) of deposition. The eggs are densely covered with hair-like outgrowths. The larva is sluggish and does not feed. It moults in one to eight days (average 3-16) into an active feeding protonymph. The protonymphal stage lasts nine to thirteen days (average 10-0). The deuto- nymph is also active and feeding lasts seven to thirty-five days (average 24-22). Unlike P. (P.) robustus, males and females were reared from the egg batch laid by each female. There is considerable confusion in the literature concerning the identity of P. (P.) crassipes. Berlese (1906) in his monograph describes and figures two forms, namely, Gamasus (Pergamasus) crassipes (L.) and G. (P.) crassipes var. longicornis Berlese. These two forms are readily distinguishable by the armature of the tibia of leg II in the male. Unfortunately, Berlese connected the wrong female and male ; thus the female of G. (P.) crassipes : Berlese is really the female of G. (P.) crassipes var. longicornis Berlese. I have verified this by examining material in the Berlese Collection at Florence and by rearing P. (P.) crassipes in the laboratory. I consider the two " forms " to be distinct sympatric species which can be distinguished by a number of morphological characters. Owing to the confusion regarding the identity of these species, I have not attempted a full synonymy. Pergamasus (Pergamasus) longicornis Berlese Gamasus (Pergamasus) crassipes var. longicornis Berlese, A. (1906). Redia 3 : 232, fig. Gamasus (Pergamasus) crassipes (L.) Latr. Berlese, A. (1906). Redia 3 : 229, fig. (female only). Pergamasus crassipes (L.) var. longicornis : Tragardh, I. (1912), Arch. Zool. Exper. gen. 8 : 523, figs. ; Tragardh, I. (1931). The Natural History of Juan Fernandez and Easter Islands 3 : 603, figs. ; Turk, F. A. (1945). Ann. Mag. nat. Hist. 12 n : 802, figs. ; Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 59, figs. Gamasus (Pergamasus) crassipes (L.) var. longicornis : Halbert, J. N. (1915). Proc. R. Irish Acad. 31 39 ii : 54, fig. 156 S. K. BHATTACHARYYA 43 FIGS. 41-45. Pergamasus (Pergamasus) longicornis Berlese, male. Fig. 41 dorsum of idiosoma. Fig. 42 venter. Fig. 43 tectum. Fig. 44 chelicera. Fig. 45 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 157 Pergamasus crassipes (L.) Latr. var. longicornis : Womersley, H. (1942). Trans. Roy. Soc. S. Aust. 66 (2) : 146, figs, (male only). Pergamasus crassipes longicornis : Cooreman, J. (1943). Bull. Mus. nat. Bel., 19 63 : 4, fig. Amblygamasus septentrionalis belgicus Cooreman, J. (1943). Bull. Mus. nat. Bel., 19 63 : 4, figs. MALE. The dorsal shield (1,275-1, 284^ long x 714-747(1 wide) is pear-shaped and strongly sclerotized. The opisthonotal shield is densely covered with setae (Text-fig. 41). The tritosternum is provided with a pair of pilose laciniae. The genital lamina is anteriorly produced into a sharp median spine (Text-fig. 42). A pair of prae- endopodal shields is present. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. There is a post-stigmatal prolongation of the peritrematal shield which reaches to coxa IV. A pair of post-stigmatal setae is present. The tectum has five prongs (Text-fig. 43). The most distinctive features of the pedipalp are : the trochanter without any tubercle, the femur with a comb-like seta and the genu with two spatulate setae. The chelicera is robust and shown in Text-fig. 44. The corniculus and hypostomal processes are similar to those in P. crassipes, The tibia, tarsus and prae-tarsus of leg I are 250-266^, 304-342^ and 25-35^ long respectively. Leg II : the femur is crassate with a strong falcate spur and the genu has a tubular spur ; the tibia has three spurs, namely distal, medial and lateral (Plate ib). The median process terminates in a hook-like structure, and this shape is constant in all the specimens I have examined. The distinctive shape of the trochanter of leg IV is figured (Text-fig. 45). FEMALE. The dorsal shield measures i,268-i,3i7[x long x 797-815^ wide and is dark brown to deep yellow in colour. The distribution and relative lengths of the dorsal setae are shown in Text-fig. 46. The tritosternum and prae-endopodal shields are delineated as in Text-fig. 47. The metasternal shields are free. The endogynium has a pair of round sacs with a bifurcate median process. Each sac is strongly reticulated. The genital shield is shown in Plate 7b. The setation of the opisthogastral shield is shown in Text- fig. 47. The stigma is situated between coxae III and IV with the peritreme extending to coxa I. The peritrematal shield is entirely fused with the dorsal shield. The tectum is five-pronged (Text-fig. 48). The pedipalp is essentially similar to that in the male. The chelicera is figured (Text-fig. 49). Leg I has the tibia, tarsus and prae-tarsus 262-266^, 358-363^ and 29-35^ long respectively. The trochanter of leg IV lacks a spur. DISTRIBUTION AND HABITAT. This species has been recorded from Northern Europe (Berlese, 1906), Germany, Norway (Tragardh, 1912), Ireland (Halbert, 1915), Juan Fernandez and Easter Islands (Tragardh, 1931), Australia (Womersley, 1942) and Belgium (Cooreman, 1943). I have examined specimens from litter under Cupressus sp. (Rothamsted Lodge, Rothamsted Experimental Station, Harpenden, Hertfordshire, G. Owen Evans and E. Browning, 18 . iii . 1957). Oak and birch leaves 158 S. K. BHATTACHARYYA 48 FIGS. 46-49. Pergamasus (Pergamasus) longicornis Berlese, female. Fig. 46 dorsum of idiosoma. Fig. 47 venter. Fig. 48 tectum. Fig. 49 chelicera. (Newton Abbot, S. Devon, R. G. Lewesther, 21 .vi. 1951). Litter and humus under pines and holly (Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, io.ix.i956). Moss leaves (Adelaide Hill, Bowness, Cumberland, J. T. Salmon, 29. vi. 1951). Summit of Dove Crag, 2,600 ft. (near Ambleside, Westmorland, M. E. Bacchus, 28.11.1954). Sphagnum, etc. (Cock Hill, Yorkshire, J. T. Salmon, 2i.vii.i95i). In leaf mould (Epping Forest, Essex, J. T. Salmon, 27. v. 1951). In leaf mould (Saver- nake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). In moss of wall (Tarn Hows, N. Lancashire, M. E. Bacchus, 4.xii.i954). Litter under thick evergreen hedge (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 29-iii. 1957). Litter and soil under REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 159 evergreen trees (St. Agnes, Isles of Stilly, Cornwall, K. H. Hyatt, 5.^.1957). (Royal Botanical Gardens, Kew, Surrey, J. L. Gilbert, 5.1^.1961). Moss from crevices in rocks (near Force Forge, Westmorland, M. E. Bacchus, 2.xii.i954). In oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). Moles' (Talpa europaea) nest, near pasture (Godstow, Berkshire, Prof. P. A. Larkin, 4^1.1962). Rotten black leaves at riverside rocks (on the river Brathy, S.W. Ambleside, Lancashire, P. N. Lawrence, 23. ix. 1956). From pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Salt marsh debris, (Llanrhidian, Glamorgan, P. N. Lawrence, 27.^1.1962). Yew and rhododendron humus (Educational Garden, Swansea University, Swansea, P. N. Lawrence, 26.^.1962). In moss (Tarbert, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3 . vii. 1951). In moss and heather (Glen Garry, Perth, J. T. Salmon, 5. vii. 1951). In dead leaves (Loch Lomond, between Glasgow and Tarbert, J. T. Salmon, 8. vii. 1951). Moss on rock (Caherbannagh, Co. Clare, Ireland, Nottingham University Expedition, vii Pergamasus (Pergamasus) septentrionalis (Oudemans) Parasitus septentrionalis Oudemans, A. C. (1902). Tijdschr. Ent. 45 : 39, figs. Gamasus (Amblygamasus) septentrionalis : Berlese, A. (1906). Redia 3 : 188, figs. Gamasus (Amblygamasus} septentrionalis var. norvegicus Berlese, A. (1906). Redia 3 : 190, figs. ; Sellnick, M. (1940). Goteborgs VetenskSamh. Handl. (5) 68 14 : 58, figs. Amblygamasus septentrionalis (Oudemans) 1902. Haarlov, N. (1957). Spolia zool. Mus. Hauniensis 17 : 16, figs. Amblygamasus septentrionalis (Oudemans) norvegicus Berlese 1905. Halaskova, V. (1959). Zvldstni. Otisk Mus. Zprdvy Prazskeho Kraje 4 : 6, figs. MALE. This species is strongly sclerotized and reddish-brown in colour. The dorsal shield (1,383-1,463^ long x 7i4-72i(x wide) is regionally reticulated. Only the opisthonotal shield is densely covered with setae (Text-fig. 50). Some dorsal shield setae appear to be pilose under high magnification. The tritosternum has a pair of pilose laciniae. The genital lamina is produced, anteriorly, into a well developed sharp, median spine (Text-fig. 51). There is a pair of prae-endopodal shields. The sternal setae are sparsely pilose under high magnifica- tion. The pitted reticulation extends from the anterior margin of the sternal shield to a little behind coxae IV. The stigma is situated between coxae III and IV ; and the peritreme extends as far as coxa I. The post-stigmatal extension of the peritrematal shield extends to coxa IV. The tectum is four-pronged (Text-fig. 52). The most distinctive features of the pedipalp are : the femur with one and the genu with two distally denticulated setae. The chelicera is robust and its digit has a row of closely-set teeth. The distinctive shape of the fixed digit is shown in Text-fig. 53 ; the dorsal seta being terminally denticulated. The corniculus is distinctively stalked and the hypostomal processes are shown in Text-fig. 54. The deutosternal denticles are arranged in fourteen or fifteen transverse rows. The hypostomal setae are pilose. i6o S. K. BHATTACHARYYA FIGS. 50-54. Pergamasus (Pergamasus) septentrionalis (Oudemans), male. Fig. 50 dorsum of idiosoma. Fig. 51 venter. Fig. 52 tectum. Fig. 53 chelicera. Fig. 54 gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 161 The tarsus (462-475^) of leg I is considerably longer than the tibia (337-358(0.) ; the prae-tarsus is 41-49^ long. Leg II ; the femur is without a distinct spur, the genu has three processes and two of them are well developed and retrogade (Plate ic). The main tibial process is large, conical and stalked. The trochanter of leg IV is simple. FEMALE. The dorsal shield (1,400-1,567^ long x 847-950^ wide) is strongly sclerotized and tapers anteriorly. The neotrichy is confined to the posterior half of the dorsal shield (Text-fig. 55). A few dorsal setae appear pilose under high magnification. Ventrally, there are two large prae-endopodal shields almost meeting in the median line and forming a cup-shaped structure into which the base of the tritosternum is situated. The sternal setae I-III are pilose. The metasternal shields are free. The endogynium has a pair of blunt horn-like lobes joined together by a common stalk (Text-fig. 56). The genital shield is shown in Plate 7c. A very short median process may be present between the sacs. The pitted reticulation of the ventral shield extends from the anterior margin of the sternal shield to a little behind coxae IV. The distribution of setae on the opisthogastral shield is shown in Text- fig. 57. The peritrematal shield is entirely fused with the dorsal shield. The stigma is situated between coxae III and IV ; the peritreme extends to the level of coxa I. The tectum is shown in Text-fig. 58. The pedipalp is similar to that in the male. The dentition of the chelicera and the form of the spermadactyl process are shown in Text-fig. 59. The dorsal seta of the fixed digit is terminally denticulated. The tarsus (477-5o8(z) of leg I is much longer than the tibia (367-400^) ; the prae- tarsus is 37-49^ long. The trochanter of leg IV is shown in Text-fig. 60. DISTRIBUTION AND HABITAT. This species has already been recorded from Holland (Oudemans, 1902), Germany and Norway (Berlese, 1906), Iceland (Sellnick, 1940), Denmark (Haarlov, 1957) and Czechoslovakia (Halaskova, 1959). I have examined specimens from : leaf mould (Savernake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). Litter under thick evergreen hedge (St. Agnes, Isles of Stilly, Cornwall, K. H. Hyatt, 29.^1.1957). Rotten black leaves at riverside rocks (on the river Brathy, S.W. Ambleside, Lancashire, P. N. Lawrence, 23. ix. 1956). Leaf mould from woods (near Buxton, Derbyshire, J. T. Salmon, 23. vi. 1951). Rotten reeds (Colletts Bridge, nr. Wisbech, Cambridge, P. N. Lawrence). Ash humus (Colletts Bridge, nr. Wisbech, Cambridge, P. N. Lawrence, i6.x.i96o). Rotten wood (Blurridge Farm, Ridge Hill, Combe Martin, Devonshire, M. E. Bacchus, I2.iii.i957). In leaf mould (Richmond Park, Surrey, J. T. Salmon, ig.iii.ig^i). In leaf mould (Leith Hill, Surrey, J. T. Salmon, i6.vi.i95i). Summit of Dove Crag (near Ambleside, Westmorland, M. E. Bacchus, 29.11.1954). Stratton Strawless, near Aylsham, Norwich, Norfolk, J. T. Salmon). Yew and rhododendron (Educa- tional Garden, Swansea University, Swansea, P. N. Lawrence, 26.iii. 1962). Bracken litter (Bishopston Common, Swansea, S. K. Bhattacharyya, 4. i. 1961). In moss and heather (Glen Garry, Meall Doire an Daimh 1900 ft., Perth, J. T. Salmon, 5. vii. 1951 1 62 S. K. BHATTACHARYYA 59 FIGS. 55-60. Pergamasus (Pergamasus) septentrionalis (Oudemans), female. Fig. 55 dorsum of idiosoma. Fig. 56 endogynial process. Fig. 57 venter. Fig. 58 tectum. Fig. 59 chelicera. Fig. 60 trochanter of leg IV. In moss (Tarbert, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3.vii.i95i). Moss on rock under Hazel (Caherbannagh, Co. Clare, Ireland, Nottingham University Expedition, viii.i96i). BIOLOGICAL NOTES. Six females were kept in the laboratory and all of them laid eggs erratically. Ten eggs were laid by one female and the oviposition period extended for thirty-one days. The eggs are relatively large in size and lustrous. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 163 The outer surface of the chorion is densely covered with hair-like outgrowths. The eggs hatched in two to five days and the non-feeding larval stage lasted one to three days (average 2.44 days). The larva is sluggish. There are two feeding nymphal stages, the proto- and deutonymph. High mortality in the culture was observed for the protonymphs. The duration of the protonymphal stage was seven to twenty- two days (average 14.5 days). Two deutonymphs moulted into adult females and the duration of this stage was seven and twenty-one days, respectively. The nymphal and adult stages are very active and cannibalistic. Pergamasus (Pergamasus) quisquiliarum (Can.) Gamasus quisquiliarum Canestrini, G. & R. (1882). Atti. 1st. Veneto, 8 (5) : 920. Gamasus (Pergamasus) quisquiliarum : Berlese, A. (1906). Redia 3 : 223, figs. Pergamasus quisquiliarum : Cooreman, J. (1943). Bull. Mus. roy. Hist. nat. Bel. 19, 63 : 7, fig. ; Halaskova, V. (1959). Zvldstni Otisk Mus. Zprdvy Prazskeho Kraje 4 : 3, figs. ; Schweizer, J. (1961). Denks, schweiz. naturf. Ges. 84 : 41, figs. MALE. This species is deep yellow to reddish-brown in colour and well sclerotized. The dorsal shield (1,157-1,317^ long x 582-6i5[x wide) is without a transverse suture and is regionally reticulated. The opisthonotal shield is densely covered with setae, neotrichous (Text-fig. 61). Some dorsal setae appear to be pilose, under high magnification. The tritosternum is well developed with a pair of pilose laciniae. The genital lamina has a median incision (Text-fig. 62). Two large prae-endopodal shields flank the genital sclerite. The pitted reticulation extends from the anterior margin of the sternal shield to a little behind coxae IV. The stigma is situated between coxae III and IV ; the peritreme anteriorly extends to coxa I and its outer margin is crenate. Posteriorly, the post-stigmatal prolongation of the peritrematal shield extends to coxa IV. The tectum has five prongs (Text-fig. 63). The distinctive features of the pedipalp are shown in Text-fig. 64. The fixed digit of the chelicera is multi-dentate and the teeth are confined to the distal half of the digit. The fixed digit is truncated and the dorsal seta has a denticulated end (Text-fig. 65). The anterior lateral margins of the hypostomal processes are fringed (Text-fig. 66) . The corniculus is stalked and is of distinctive shape. The tibia, tarsus and prae-tarsus of leg I are 254-312^, 388-39250, and 45-49^ long, respectively. Leg II : the femur lacks an accessory spur and the main spur is distally truncated. The genual process is very weakly developed (Plates id & e). The tibia is provided with a vermiform process. The trochanter of leg IV is without any process. FEMALE. The dorsal shield (1,317-1,417^ long x 764-815^ wide) is regionally reticulated. The podonotal shield is non-neotrichous, whilst the opisthonotal shield is densely covered with setae (Text-fig. 67) . The tritosternum has a pair of pilose laciniae. The tritosternal base is situated in a cup-like structure formed by a pair of prae-endopodal shields. The ventral shield 164 S. K. BHATTACHARYYA 64 FIGS. 61-66. Pergamasus (Pergamasus) quisquiliarum (Canestrini), male. Fig. 61 dorsum of idiosoma. Fig. 62 venter. Fig. 63 tectum. Fig. 64 trochanter, femur and genu of pedipalp. Fig. 65 chelicera. Fig. 66 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 165 67 68 70 FIGS. 67-70. Pergamasus (Pergamasus) quisquiliarum (Canestrini), female. Fig. 67 dorsum of idiosoma. Fig. 68 venter. Fig. 69 genital region without epigynial shield. Fig. 70 bifurcated endogynial process. 166 S. K. BHATTACHARYYA is regionally ornamented with a pitted reticulation (Text-fig. 68). The metasternal shields are free and flank the epigynial shield. The genital shield is shown in Plate yd. The endogynial process is complicated and its sacs are somewhat rounded in shape (Text-fig. 69). There is a median, elongated process in between the endogynial sacs, the terminal end of which may be bi- or trifurcated (Text-figs. 70 and 71). The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. The entire peritrematal shield is fused with the dorsal shield. The tectum and dentition of the chelicera are shown in Text-figs. 72 and 73 respectively. The pedipalp is essentially similar to that of the male (Text-fig. 74). Approximately half of the lateral margins of the hypostomal processes are fringed (Text-fig. 75). The deutosternal denticles are arranged in fourteen to sixteen trans- verse rows. The tarsus (470-487^ long) of leg I is considerably longer than the tibia (350- 367^) ; the prae-tarsus is 29-55^ long. The trochanter of leg IV is without a tubercle. DISTRIBUTION AND HABITAT. This species has previously been recorded from Italy (Canestrini & Canestrini, 1882), Norway (Berlese, 1906), Belgium (Cooreman, 1943), Czechoslovakia (Halaskova, 1959), Switzerland (Schweizer, 1961) and South America (Sheals, 1962). I have examined specimens from the following localities : Litter and soil under evergreen trees (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 5.^.1957). In leaf mould (Houghton Green, Rye, Sussex, J. T. Salmon, 19. v. 1951). Bramble- covered ditch (Colletts Bridge, Norfolk, P. N. Lawrence). From soil (between Rockingham and Corby, Northants, B. N. K. Davies, 14.1.1958). Rotten reeds (Colletts Bridge, Norfolk, P. N. Lawrence). Litter under Cupressus sp. (Rothamsted Lodge, Rothamsted Experimental Station, Harpenden, Hertfordshire, G. Owen Evans and E. Browning, i8.iii.i957). Oak and beech litter (Singleton Park, Swansea, S. K. Bhattacharyya, 3.1.1961). Pergamasus (Pergamasus) mirabilis Willmann Pergamasus mirabilis Willmann, C. (1951). SitzBer. Osterr. Akad. Wiss. Abt. i, 160 : 99, figs. Amblygamasus mirabilis : Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 66, figs. MALE. The dorsal shield (77i-838(x long x 400-424^ wide) is reddish in colour and has a transverse suture (Text-fig. 76). Some dorsal setae appear to be pilose under high magnification. The form of the prae-endopodal shields is shown in Text-fig. 77. The trito- sternum has a pair of pilose laciniae but the tritosternal base is completely hidden by the genital lamina. The setation of the opisthogastral shield is figured. The pitted reticulation extends from the anterior margin of the sternal shield to a little behind the coxa of leg IV. The stigma is situated between coxae III and IV ; and the undulating peritreme reaches to coxa I. A pair of post-stigmatal setae is present. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 167 71 75 73 FIGS. 71-75. Pergamasus (Pergamasus) quisquiliarum (Canestrini) , female. Fig. 71 trifurcated endogynial process. Fig. 72 tectum. Fig. 73 chelicera. Fig. trochanter, femur and genu of pedipalp. Fig. 75 venter of gnathosoma, 74 168 S. K. BHATTACHARYYA FIGS. 76-81. Pergamasus (Pergamasus) mimbilis Willmann, male. Fig. 76 dorsum of idiosoma. Fig. 77 venter. Fig. 78 tectum. Fig. 79 trochanter, femur and genu of pedipalp. Fig. 80 chelicera. Fig. 81 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 169 The tectum is three-pronged, the median being longer and broader than the lateral tines (Text-fig. 78). The palpal trochanter, femur and genii are shown in Text-fig. 79. The chelicera is rather elongated with well developed dentition, (Text-fig. 80). A cone-shaped projection is especially discernible above the dorsal seta of the chelicera. The fixed digit is not truncated and has a simple dorsal seta. The teeth of the movable digit are well separated and the spermadactyl is normal. The corniculus is stalked ; and more than the distal half of the hypostomal processes is provided with fringes. The deutosternal denticles in the ventral groove of the gnathosoma are shown in Text-fig. 81. The tibia (162-168(0.) of leg I is remarkably shorter than the tarsus (221-247^) ; the prae-tarsus is only 17-25^ long. The coxa of leg II has well-developed spurs (Plate 2a). The femur of leg II is not crassate (Plate 2b). The genu has a large falcate spur, and the tibia is devoid of processes. The trochanter of leg IV possesses no spur. FEMALE. Unknown. DISTRIBUTION AND HABITAT. This species is previously recorded from Austria (Willmann, 1951) and Switzerland (Schweizer, 1961). I have examined specimens from soil (between Rockingham and Corby, Northants, B. N. K. Davis, 14.1.1958). Pergamasus (Pergamasus) hamatus (C. L. Koch) Berlese, 1906 Gamasus hamatus Koch, C. L. (1839). Deutsch. Crust. Myr. Arachn. fasc. 26 t. 2. Gamasus (Pergamasus} hamatus : Berlese, A. (1906). Redia 3 : 212, figs. Gamasus (Pergamasus) hamatus var. gracilis Berl. (sic) : Berlese, A. (1906). Redia 3 : tab. V, fig. Tab. XIV, figs. Pergamasus hamatus : Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 61, figs. Pergamasus hamatus (Koch) var. longipes Schweizer, J. (1961). Denks. schweiz. natiirf. Ges. 84 : 62, figs. MALE. The dorsal shield (1,157^ long x 58251 wide) is strongly sclerotized and provided with a median transverse suture. This species differs from other neotrichous species in that the neotrichy extends into the podonotal region of the dorsal shield (Text-fig. 82). Some dorsal setae appear pilose under high magnification. Ventrally, the tritosternum has a pair of well-developed pilose laciniae but the tritosternal base is completely hidden by the genital lamina. The distribution of the prae-endopodal shields, and the shape of the genital lamina, is shown in Text- fig. 83. The setation of the opisthogastral shield is figured. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. The peritrematal shield lies along the dorso-ventral margin of the body. The tectum is three-pronged, the median prong being longer than the lateral ones (Text-fig. 84). The dorsal surface of the tectum is faintly reticulated. The pedipalp trochanter is tuberculated (Text-fig. 85) ; the femur and genu are provided with one comb-like and two palmate setae, respectively. The fixed digit of the chelicera has S. K. BHATTACHARYYA FIGS. 82-83. Pergamasus (Pergamasus) hamatus (Koch), male. Fig. 82 dorsum of idiosoma. Fig. 83 venter. a row of closely-set teeth ; the movable digit is bidentate with a normal sperm- adactyl (Text-fig. 86). The terminal ends of both digits are pointed and the arthrodial membranes are simple and branched. The corniculus has a distinct stalk and its peculiar shape is shown in Text-fig. 87. The fringed hypostomal processes are as figured. The deutosternal denticles on the ventral groove of the gnathosoma are arranged in ten transverse rows. The tibia, tarsus and prae-tarsus of leg I are 254^, 333-337^ and 41 \i long respectively. The distinctive feature of leg II is shown in Plate 2c. The femur has a relatively small main falcate spur at the base of which is an elongated accessory spur. The genu and tibia are armed with conical rather than elongate spurs. The trochanter of leg IV is without any prominence. FEMALE. The dorsal shield (1,200-1,268^ long x 665^ wide) tapers posteriorly. The setation of the dorsal shield is similar to that of the male (Text-fig. 88). REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 171 85 84 86 87 FIGS. 84-87. Pergamasus (Pergamasus) hamatus (Koch), male. Fig. 84 tectum. Fig. 85 trochanter of pedipalp. Fig. 86 chelicera. Fig. 87 venter of gnathosoma. The tritosternum is well developed with an elongated basal part and a pair of pilose laciniae. A pair of large prae-endopodal shields is present. The sternal shield is furnished with a median vertical groove which anteriorly reaches to about the level of sternal setae II. The metasternal shields may be fused. The endogynium has a pair of horns. The endogynial processes are relatively simple. The setation of the opisthogastral shield is shown in Text-fig. 89. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. The antero-lateral margin of the peritrematal shield is coalesced with the dorsal shield at the level of coxa II, but the shield is free posteriorly. The tectum is three-pronged (Text-fig. 90). The pedipalp is essentially similar to that in the male (except the trochanter, which lacks a tubercle, Text-fig. 91). The fixed digit is furnished with a row of closely-set teeth : the movable finger is tridentate, the teeth being well separated (Text-fig. 92). Leg I with the tibia, tarsus and prae-tarsus 270^, 35o-367(j, and 37(x long, respectively. I 7 2 S. K. BHATTACHARYYA FIGS. 88-92. Pergamasus (Pergamasus) hamatus (Koch), female. Fig. 88 dorsum of idiosoma. Fig. 89 venter. Fig. 90 tectum. Fig. 91 trochanter of pedipalp. Fig. 92 chelicera. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 173 DISTRIBUTION AND HABITAT. This species has been recorded from Germany (Koch, 1839), Italy (Berlese, 1906) and Switzerland (Schweizer, 1961). I have examined specimens from Brigster, Lancashire, and bulk soil, Ayside, Westmorland (J. Satchell, xi.i953). Pergamasus (Paragamasus) hortensis sp. nov. FEMALE. The dorsal shield, in the compressed specimen, measures about i,200(x long and 683^ wide, and is lightly sclerotized and yellow in colour. The chaetotaxy of the opisthonotal region shows neotrichy (Text-fig. 93). Ventrally, the tritosternum has an elongate basal part and a pair of pilose laciniae. A pair of large prae-endopodal shields is present. The posterior margin of the sternal shield is provided with a short, median vertical fissure. The metasternal shields are free. The endogynial horns and processes are delineated in Text-fig. 94. The setation of the opisthogastral shield is figured. The stigma is situated between FIGS. 93-95. Pergamasus (Pergamasus) hortensis sp. nov., female. Fig- 93 venter. Fig. 94 magnified structure of the sterniti-genital region. Fig. 95 tectum. 174 S. K. BHATTACHARYYA coxae III and IV ; the peritreme extends to coxa I. The peritrematal shield is free posteriorly. The tectum has three prongs (Text-fig. 95) . The chelicerae of the unique specimen have not been examined. The most distinctive features of the pedipalp are, the femur with a comb-like seta and the genu with two spatulate setae. The hypostomal process is similar to that in P. (P.) quisquiliarum. Leg I with tibia, tarsus and prae-tarsus 274-279^, 354(x and 43-45^ long respectively. The trochanter of leg IV is without any tubercle. MALE. Unknown. HABITAT. I have examined a single damaged female from greenhouse soil, Marton, Blackpool, Lancashire, M. Cohen, 1934 (Holotype 1963.2.7.10). Pergamasus (Paragamasus) robustus (Oudemans) Parasitus longulus (C. L. Koch), var. robustus Oudemans, A. C. (1902). Tijdschr. Ent. 45 : 38, figs. Parasitus robustus : Oudemans, A. C. (1905). Abh. Nat. Ver. Bremen 18 : 82. Gamasus (Pergamasus) robustus : Berlese, A. (1906). Redia 3 : 219, figs. Pergamasus robustus : Tragardh, I. (1910). Naturw. Untersuch. Sarekgeb. 4 (4) : 404, figs. MALE. The dorsal shield (i,ioo-i,2OO(ji long x 6i5-665[x wide) has a median transverse suture (Text-fig. 96). The podonotal region bears twenty-one pairs of setae and their distribution is figured. The opisthonotal region has a variable number of setae. The tritosternum has a pair of well-developed ciliated laciniae. The genital lamina is distinctive in shape (Text-fig. 97). The anterior margin of the sternal shield is slightly concave. The setation of the opisthosomal region is figured. The stigma is situated between coxae III and IV ; the peritreme extends to coxa II. A pair of post-stigmatal setae is present. The tectum is rounded in shape and provided with three prongs (Text-fig. 98). The most distinctive features of the chaetotaxy of the pedipalp are, the comb-like seta on the femur and the two spatulate setae on the genu. The palpal trochanter is without any tubercle (Text-fig. 99). The chelicera is robust, the fixed digit being truncated and with a variable number of teeth (Text-figs. 100 & 101). The movable digit is unidentate with a normal spermadactyl. There are ten transverse rows of deutosternal denticles. The entire outer margins of the hypostomal processes are fringed (Text-fig. 102). The corniculus is distinctly stalked. The tibia, tarsus and prae-tarsus of leg I are i83-i88pL, 237-258(1 and 17-25^ long respectively. The distinctive features of leg II are shown in Plate 2d and Plate 3f ; the trochanter being spurred. The trochanter of leg IV is devoid of processes. FEMALE. The dorsal shield (i,ii7(j, long x 598-632^ wide) is regionally reticulated. The anterior half of the dorsal shield is furnished with twenty-one pairs of setae, whilst the number of setae on the posterior half is greatly variable (Text- fig. 103). REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. IOO FIGS. 96-102. Pergamasus (Paragamasus) robustus (Oudemans), male. Fig. 96 dorsum of idiosoma. Fig. 97 venter. Fig. 98 tectum. Fig. 99 trochanter, femur and genu of pedipalp. Fig. 100 chelicera. Fig. 101 variation in the dentition of the fixed digit. Fig. 102 venter of gnathosoma. The tritosternum has a well-developed basal part. There is a granulated region in between a pair of large prae-endopodal shields. The sternal shield bears three pairs of setae and two pairs of pores. Posteriorly the sternal shield has a median narrow incision reaching the level of sternal setae II. The metasternal shields are free. A pair of endogynial horns is present. The endogynial processes are very complicated (Text-fig. 104). The genital shield is shown in Plate ye. The stigma is situated between coxae III and IV ; the extension of the peritreme is as in the male. 1 7 6 S. K. BHATTACHARYYA FIGS. 103-108. Pergamasus (Paragamasus) robustus (Oudemans), female. Fig. 103 dorsura of idiosoma. Fig. 104 venter. Fig. 105 tectum. Fig. 106 chelicera. Fig. 107 venter of gnathosoma. Fig. 108 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 177 The tectum is three-pronged (Text-fig. 105). The pedipalp is essentially similar to that of the male. The dorsal seta of the fixed digit is pilose ; the movable digit of the chelicera is tridentate (Text-fig. 106). The structure of the gnathosoma is shown in Text-fig. 107. The tibia, tarsus and prae-tarsus of leg I are 183-187^, 254-258^ and 17-21(1 long respectively. The trochanter of leg IV may be with or without any processes (Text-fig. 108). DISTRIBUTION AND HABITAT. This species has been recorded from Holland (Oudemans, 1902), Germany, Norway (Berlese, 1906), and Swedish Lapland (Tragardh, 1910). I have examined the following material : Litter and humus under pines and holly (near Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, io.ix.i956). Moss and leaves (Adelaide Hill, Bowness, Cumberland, J. T. Salmon, 29. vi. 1951). Sphagnum, etc. (Cock Hill, Yorkshire, J. T. Salmon, 21 . vii . 1951) . In leaf mould (Epping Forest, Essex, J. T. Salmon, 27. v. 1951). In leaf mould (Savernake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). In leaf mould (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951), (Stratton Strawless, near Aylsham, Norwich, Norfolk, J.T. Salmon, 13 . v . 1951) . In moss (Grassington, Grass Woods, North Riding, Yorkshire, J. T. Salmon, 24.^.1951). Sphagnum (Ascot, Berkshire, A. H. G. Alston, 8.xii.i957). Moss on tree stumps and stones (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951), (Red Bank, Grasmere, Westmorland, M. E. Bacchus, 3.xii.i954). In oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). In leaf mould (Boltons Wood, Boltons Abbey, North Riding, Yorks., J. T. Salmon, 24. vi. 1951). In leaf mould (Burnham Beeches, Buckinghamshire, J. T. Salmon, 3. v. 1951). Sphagnum, etc. (Moel Siabod, North Wales, P. N. Law- rence, 22.x. 1960). Rotten stump (Gwydyr Forest, North Wales, P. N. Lawrence, 22 . x . 1960) . Oak humus (Gwydyr Forest, North Wales, P. N. Lawrence, 22.x. 1960) . In moss (Tarbert, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3. vii. 1951). Floor of scotch fir wood (Aviemore, Inverness, J. T. Salmon, 7. vii. 1951). In moss and heather (Glen Garry, Perth, Meall Doire and Daimh, J. T. Salmon, 5. vii. 1951). BIOLOGICAL NOTES. Two females collected from oak litter were reared under laboratory conditions. Twenty eggs were laid by one female and twenty- two eggs by the other. The female which laid twenty eggs lived for eighty-eight days in culture. The eggs are smooth and considerably smaller in size than those of P. (P.) crassipes or P. (P.) septentrionalis. The eggs hatched in three to five days (average 4-5 days) after deposition and the active feeding larval stage lasted five to seventeen days (average 9-08 days). The duration of the protonymphal and deutonymphal stages was seven to fifteen (average 10-18 days) and fourteen to twenty-three (average 20-0) days, respectively. Both these instars were active and fed voraciously on Collembola. Cannibalism was also noted. The eight deutonymphs reared all moulted to give males. These males resembled the parent female in lacking a tubercle on the trochanter of leg IV. There were no tubercles on the trochanters of the pedipalps in the reared males. i 7 8 S. K. BHATTACHARYYA Pergamasus (Paragamasus) alpestris Berlese Gamasus (Pergamasus) alpestris Berlese, A. (1904). Redia 1 : 236 ; Berlese, A. (1906). Redia 3 : 220, figs. Pergamasus alpestris : Cooreman, J. (1954). Bull. Inst. roy. Sci. nat. Belg, 30, 34 : 5, figs. HalaSkova, V. (1959) Zvldstni Otisk Mus. Zprdvy Prazsktho Kraje 4 : 3, figs. ; Schweizer, J. (1961) Denks. schweiz. naturf. Ges. 84 : 63, figs. MALE. The dorsum is dark brown in colour and sub-globular in shape. The dorsal shield (1,150-1,190^ long x 632-648^ wide) has no transverse suture and the lengths of the dorsal setae are variable (Text-fig. 109). The prae-endopodal shields and the shape of the genital lamina are shown in Text-fig, no. The tritosternum has an extremely reduced base and a pair of pilose laciniae. r I \ l I / *i r V r M- f ' " ' /.\ in 'i V.'-M IO9 FIGS. 109-112. Pergamasus (Paragamasus) alpestris Berlese, male. Fig. 109 dorsum of idiosoma. Fig. no venter. Fig. in tectum. Fig. 112 trochanter, femur and genu of pedipalp. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 179 The venter is very characteristic ; a scaly, elongated structure is situated near the coxa of leg IV. The setation of the opisthogastral shield is figured. The stigma is situated between the coxae III and IV ; and the peritreme extends to coxa I. A pair of relatively long post-stigmatal setae is present. The tectum is three-pronged and situated on a cup-like structure (Text-fig, in). The distinctive features of the pedipalp and chelicerae are shown in Text-figs. 112 and 113 respectively. The corniculus is distinctly stalked ; and the hypostomal process fringed (Text-fig. 114). The deutosternal denticles are arranged in ten transverse rows. The tibia, tarsus and prae-tarsus of leg I are 204-208^, 300-304^ and 2i-25fx long respectively. Leg II, with coxa, trochanter, femur, genu and tibia, is armed with variously shaped spurs (Plate 21). Leg II is remarkable for the presence of a spur on the coxa (Plate 2e). The femur is crassate with a well-developed falcate major spur and accessory spurs. The armature of the trochanter of leg IV is shown in Text-fig. 115. FEMALE. The dorsal shield (1,117-1,132^ long x 732^ wide) is broadly oval and regionally reticulated (Text-fig. 116). Lengths of the dorsal setae are greatly variable. The posterior half of the shield bears about twenty-nine pairs. / FIGS. 113-115. Pergamasus (Paragamasus) alpestris Berlese, male. Fig. 113 chelicera. Fig. 114 venter of gnathosoma. Fig. 115 trochanter of leg IV. i8o S. K. BHATTACHARYYA FIGS. 116-121. Pergamasus (Paragamasus) alpestris Berlese, female. Fig. 116 dorsum of idiosoma. Fig. 117 venter. Fig. 118 endogynial process. Fig. 119 tectum. Fig. 120 chelicera. Fig. 121 venter of gnathosoma. REVISION OF THE GENUS PERGA MA S US BERLESE S. LAT. 181 Ventrally, the tritosternum has an elongated basal part and a pair of well- developed pilose laciniae. There is a pair of large prae-endopodal shields which are coalesced in the median line. The metasternal shields are free (Text-fig. 117). The endogynium has a pair of horns. The endogynial process is complicated (Text- fig. 118). The genital shield is as seen in Plate ji. The opisthogastral shield is furnished with ten pairs of setae (excepting paranal and post-anal setae) of variable length . The stigma is in the normal position with the anterior prolongation of the peritreme to coxa I. The tectum is three-pointed (Text-fig. 119). The chelicera is shown in Text- fig. 120. The proximal half of the hypostomal margins are fringed. The deuto- sternal denticles are arranged in parallel transverse rows (Text-fig. 121). The tarsus (287-2891.1) of leg I is considerably longer than the tibia (187-196^). The prae-tarsus (29-30^) is long. DISTRIBUTION AND HABITAT. This species is previously known from Italy (Berlese, 1904, 1906), Belgium (Cooreman, 1954), Czechoslovakia (Halaskova, 1959), Switzerland (Schweizer, 1961). I have examined material from moss on tree stumps and stones (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951 ; Red Bank, Grasmere, Westmorland, M. E. Bacchus, 3.xii.i954) ; peat (Blackbush, Kew, Surrey, A. H. G. Alston, 18.1.1958) ; moss from crevices in rocks (near Force Forge, Westmorland, M. E. Bacchus, 2.xii.i954) ; leaf mould (Boltons Wood, Boltons Abbey, North Riding, Yorks., J. T. Salmon, 24. vi. 1951) ; in moss of wall (Tarn Howes, North Lancashire, M. E. Bacchus, 4.xii. 1954) and yew and rhododendron humus (Educational Garden, Swansea University, P. N. Lawrence, 26.1^.1962). Pergamasus (Paragamasus) alstoni sp. nov. MALE. Unknown. FEMALE. The dorsal shield (622-648^ long x 350-357^ wide) is sclerotized, regionally reticulated and yellow in colour. The setation of the dorsal shield is shown in Text-fig. 122. The tritosternum has a pair of pilose laciniae. The distribution of the prae- endopodal shields is characteristic (Text-fig. 123). The interscutal membrane between the anterior margin of the sternal shield and the large prae-endopodal shields is striated. The metasternal shields are free. The endogynium has a pair of horns and its process is remarkable in shape and structure (Text-fig. 124). The genital shield is shown in Plate 7g. The opisthogastral shield has nine pairs of setae (excepting three setae associated with the anus). The stigma is situated between coxae III and IV ; and the peritreme extends as far as the level of coxa II. The peritrematal shield is characteristically fused with the exopodal and opisthogastral shields. The posterior extremity of the peritrematal shield curves inwards in a tubular form to meet coxa IV. 182 S. K. BHATTACHARYYA 124 FIGS. 122-126. Pergamasus (Paragamasus] alstoni sp. nov., female. Fig. 122 dorsum of idiosoma. Fig. 123 venter. Fig 124. endogynial process. tectum. Fig. 126 chelicera. Fig. 125 The tectum is three-pronged (Text-fig. 125). The distinctive features of the pedipalp are : femur with a comb-like seta and genu with two spatulate setae. The dentition and shape of the chelicera are shown in Text-fig. 126. The ventral groove of the gnathosoma possesses ten or eleven rows of deutosternal denticles. The tibia, tarsus and prae-tarsus of leg I are 95-102(0., 150-154^ and i2-i7(x long respectively. The trochanter of leg IV is devoid of a tubercle. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 183 HABITAT. I have examined three females from soil in filmy-fern house (Royal Physic Garden, Chelsea, London, A. H. G. Alston, January, 1956) and a single female from yew and rhododendron humus (Educational Garden, Swansea University, Swansea, P. N. Lawrence, 26.1^.1962). I have designated a female from the Royal Physic Garden, Chelsea, London (Coll. A. H. G. Alston, January, 1956) as the Holotype (1963.2.7.16). Pergamasus (Paragamasus) armatus Halbert Gamasus (Pergamasus} runciger Berl. var. armatus Halbert, J. N. (1915). Proc. R. Irish Acad. 31 39 ii : 51, figs. MALE. The dorsal shield (838-898^ long x 433^ wide) is provided with a median transverse suture. The anterior dorsal shield is furnished with twenty-one pairs of setae and the setation of the posterior half is shown in Text-fig. 127. The placement of the prae-endopodal shields and the shape of the genital lamina is shown in Text-fig. 128. The tritosternum has a reduced basal part and a pair of ciliated laciniae. The setation of the opisthogastral shield is shown in the figure. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. A pair of post-stigmatal setae is present. The tectum is shown in Text-fig. 129. The palpal trochanter has a large seta- bearing protuberance (Text-fig. 130) ; the femur and genu are provided with one comb-like and two spatulate setae, respectively. The chelicera is shown in Text- fig. 131. The corniculus is distinctly stalked. Almost the entire lateral margins of the hypostomal processes are fringed. The deutosternal denticles on the ventral groove of the gnathosoma are shown in Text-fig. 132. The tibia, tarsus and prae-tarsus of leg I are I2i-i^0[i, 150-210^ and 17-25^ long respectively. The armature of the femur, genu, tibia and tarsus of leg II is of special interest (Plate 2g). The femur is strongly crassate and has a large falcate main spur and an accessory spur. The trochanter of leg IV is armed with a tubercle (Text-fig. 133). FEMALE. The dorsal shield (882-898^ long x 467-500^ wide) is well sclerotized and not entirely reticulated. The anterior half of the dorsal shield as usual bears twenty-one pairs of setae, and the posterior half is with twenty-nine pairs (Text- fig- 134). Ventrally, the tritosternum has an elongated basal part, and a pair of ciliated laciniae. There are several pairs of prae-endopodal shields some of which are small. The interscutal membrane, between the anterior margin of the sternal shield and the prae-endopodal shields, is striated (Text-fig. 135). The endogynium consists of a pair of horns. The genital shield is shown in Plate 8c. The opisthogastral shield has ten pairs of setae (excepting a post-anal and a pair of paranal setae). The stigma is situated between the coxae III and IV, and the peritreme extends to the level of coxa I. The tectum is three-pronged (Text-fig. 136). The distinctive features of the pedipalp are the femur with a comb-like seta and the genu with spatulate setae. The 184 S. K. BHATTACHARYYA 131 FIGS. 127-133. Pergamasus (Paragamasus) armatus Halbert, male. Fig. 127 dorsum of idiosoma. Fig. 128 venter. Fig. 129 tectum. Fig. 130 trochanter of pedipalp. Fig. 131 chelicera. Fig. 132 venter of gnathosoma. Fig. 133 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 185 FIGS. 134-137. Pergamasus (Paragamasus) armatus Halbert, female. Fig. 134 dorsum of idiosoma. Fig. 135 venter. Fig. 136 tectum. Fig. 137 chelicera. proximal half of the hypostomal process is fringed. The chelicera is as figured (Text-fig. 137). The tibia, tarsus and prae-tarsus of leg I are 116-150^, 154-183^ and 8-17^ long respectively. The trochanter of leg IV is without a tubercle. i86 S. K. BHATTACHARYYA DISTRIBUTION AND HABITAT. The female of the species is previously known from Ireland only (Halbert, 1915). I have examined material from : hornbeam humus (Bookham Common, Surrey, Mrs. R. E. Teagle, xi. 1956). In oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). Algae and rushes (Colletts Bridge, Norfolk, P. N. Lawrence, i6.x.i96o). In beech leaf mould from steep bank (Rickmansworth, Herts., M. E. Bacchus, 25. ii. 1951). Beech leaf mould (Amersham, Buckingham- shire, M. E. Bacchus, 13^.1951). Inside bark of fallen elm tree (Harefield, Middlesex, A. H. G. Alston, 22.1.1956). Rotten reeds (Colletts Bridge, Norfolk, P. N. Lawrence). Moss on trees and humus (Hen Pare Wood, Killay nr. River Clyne, nr. Swansea, Wales, P. N. Lawrence, 24.^.1962). Pergamasus (Paragamasus) cambriensis sp. nov. MALE. The dorsal shield measures 490-500^ long x 240-250^ wide and is provided with a transverse suture. The anterior region of the dorsal shield bears twenty-one pairs of setae. The tritosternum has a short basal part with a pair of pilose laciniae. A pair of large prae-endopodal shields is present. The genital lamina resembles that of P. wasmanni Oudemans. The stigma is situated between coxae III and IV ; and the peritreme extends to coxa II. A pair of post-stigmatal setae is present. The tectum is three-pronged, the median tine being shorter than the lateral ones. The palpal trochanter has a seta-bearing tubercle (Text-fig. 138). The chelicera is shown in Text-fig. 139. The corniculus is distinctly stalked, and the entire lateral margins of the hypostomal processes are fringed. (39 138 FIGS. 138-139. Pergamasus (Paragamasus) cambriensis sp. nov., male. Fig. 138 trochanter of pedipalp. Fig. 139 chelicera. Leg I with the tibia, tarsus and prae-tarsus 66-70^, Ii6-i29[x and 8-i2[A long, respectively. The femur, genu and tibia of leg II are armed with spurs (Plate 3a). The genual spur is characteristic in shape and position. The trochanter of leg IV is tuberculated. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 187 FIGS. 140-146. Pergamasus (Paragamasus) cambriensis sp. nov., female. Fig. 140 dorsum of idiosoma. Fig. 141 venter. Fig. 142 magnified structure of the sterniti-genital region. Fig. 143 tectum. Fig. 144 chelicera. Fig. 145 venter of gnathosoma. Fig. 146 trochanter of leg IV. FEMALE. The dorsal shield (515-548^ long x 267-300^ wide) is sclerotized and yellow in colour. The setation of the dorsal shield is shown in Text-fig. 140. The tritosternum has an elongated basal part and a pair of pilose laciniae. A i88 S. K. BHATTACHARYYA narrow elongated strip of prae-endopodal shield is situated in between large prae- endopodal shields (Text-fig. 141). The metasternal shields are free. The genital region is as in Text-fig. 142, and the genital shield in Plate 8d. The opisthogastral shield with ten pairs of setae. Although the peritrematal shield is fused with the dorsal shield at the level of coxa I the peritreme only extends to coxa II. The position of the stigma is the same as in the male. The tectum is three-pronged (Text-fig. 143). The dentition of the chelicera is shown in Text-fig. 144. The deutosternal denticles are arranged in ten transverse rows. Leg I with the tibia, tarsus and prae-tarsus 78-87^1, 125-137^ and 8-i2(x long respectively. The trochanter of leg IV is shown in Text-fig. 146. HABITAT. Two males and three females from hornbeam humus (Bookham Common, Surrey, Mrs. R. E. Teagle, xi.i956), two females from beech leaf mould (Amersham, Buckinghamshire, M. E. Bacchus, 13^.1951). Three females in soil (between Rockingham and Corby, Northants, B. N. K. Davis, 14.1.1958). Six males and thirteen females from litter under thick evergreen hedge (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 29.01.1957). Two females from pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Two females from oak litter (Llew Reservoir, Swansea, S. K. Bhattacharyya, 2.1.1961). I have designated a male and female from Bookham Common, Surrey (Coll. Mrs. R. E. Teagle, xi . 1956) as the Holotype (1963 .2.7.1) and Allotype (1963 .2.7.2) respectively. Pergamasus (Paragamasus) cassiteridum sp. nov. MALE. This species is strongly sclerotized and dark brown in colour. The dorsal shield (86-88[z long x 433-467^ wide) has a transverse suture (Text-fig. 147). The anterior dorsal shield bears twenty-one pairs of setae, whilst the posterior half about twenty-six pairs of setae. The tritosternum has a reduced base with a pair of pilose laciniae. The anterior- most margin of the genital lamina is convex in shape. The distribution of the prae- endopodal shields and the setation of the opisthogastral shield is shown in Text-fig. 148. The stigma is situated between coxae III and IV ; the peritreme anteriorly extends as far as the level of coxa I. A pair of post-stigmatal setae is present. The tectum is three-pronged. The most distinctive features of the pedipalp are, the trochanter with two tubercles (Text-fig. 149), the femur with a comb-like seta and the genu with two spatulate setae. The dentition of the chelicera and the form of the spermadactyl process is shown in Text-fig. 150. There are distinctly stalked corniculi. The entire lateral margins of the hypostomal processes are fringed. The tibia, tarsus and prae-tarsus of leg I are I2i-i29[x, 145-15451 and I7(x long respectively. The trochanter, femur and genu of leg II are armed with variously shaped processes (Plate 3c-e). The femur is crassate. The tibial spur is smaller than in robustus and the distal end has a prominence. The armature of the trochanter of leg IV is delineated in Text-fig. 151. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 189 150 149 FIGS. 147-151. Pergamasus (Paragamasus] cassiteridum sp. nov., male. Fig. 147 dorsum of idiosoma. Fig. 148 venter. Fig. 149 trochanterof pedipalp. Fig. 150 chelicera. Fig. 151 trochanter of leg IV. i go S. K. BHATTACHARYYA FEMALE. Unknown. HABITAT. One male from litter under thick evergreen hedge, St. Agnes, Isles of Stilly, Cornwall, K. H. Hyatt, 29.^.1957, and two males from lichens in crevices, Moel Siabod, North Wales, P. N. Lawrence, 22.x. 1960. I have designated the male from St. Agnes, Isles of Stilly, Cornwall (Coll. K. H. Hyatt, 29.111.1957) as the Holotype (1963.2.7.7). 157 156 FIGS. 152-157 Pergamasus (Paragamasus) celticus sp. nov., male. Fig. 152 dorsum of idiosoma. Fig. 153 venter. Fig. 154 tectum. Fig. 155 trochanter of pedipalp. Fig. 156 chelicera. Fig. 157 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 19! 161 FIGS. 158-161. Pergamasus (Paragamasus) celticus sp. nov., female. Fig. 158 dorsum of idiosoma. Fig. 159 venter. Fig. 160 tectum. Fig. 161 chelicera. 192 S. K. BHATTACHARYYA Pergamasus (Paragamasus) celticus sp. nov. MALE. The dorsal shield measures 732-747^ long and 367^ wide and is provided with a transverse suture (Text-fig. 152). The dorsum bears about forty-nine pairs of setae of which twenty-one pairs are situated on the anterior dorsal shield. Ventrally, the tritosternum has a pair of pilose laciniae. The distribution of the prae-endopodal shields and the shape of the genital lamina are shown in Text- fig. 153. The stigma is situated between coxae II and IV and the peritreme reaches to coxa I. A pair of post-stigmatal setae is present. The tectum is three-pronged (Text-fig. 154). The distinctive features of the pedipalp are, the trochanter has a seta-bearing process (Text-fig. 155), the femur a comb-like seta and the genu two spatulate setae. The characteristic shape of the fixed digit and the dentition of the chelicera are shown in Text-fig. 156. The corniculus is distinctly stalked. The entire lateral margins of the hypostomal processes are fringed. Leg I is with the tibia, tarsus and prae-tarsus ioo-i33[x, 168-175(0. and i6-2ifji long, respectively. The trochanter of leg II lacks a spur but the femur, genu and tibia are armed with variously shaped processes (Plate 3g). The trochanter of leg IV is shown in Text-fig. 157. FEMALE. The dorsal shield (704-732^ long x 357-374^ wide) is well sclerotized and yellow in colour. The podonotal shield invariably bears twenty-one pairs of setae but the opisthonotal shield has about twenty-seven pairs (Text-fig. 158). The tritosternum has an elongated basal part and a pair of pilose laciniae. The form of the prae-endopodal shield is shown in Text-fig. 159. Ventro-medially, the interscutal membrane between the anterior margin of the sternal and prae- endopodal shields is striated and granular. The metasternal shields are free. The endogynium has a pair of horns and its processes are delineated in Text- fig. 159. The epigynial shield is shown in Plate 8f. The opisthonotal shield is with ten pairs of setae. The position of the stigma and extension of the peritreme are similar to those in the male. The tectum is three-pronged (Text-fig. 160). The chelicera is shown in Text- fig. 161. The ventral groove of the gnathosoma has nine or ten rows of denticles. The tibia, tarsus and prae-tarsus of leg I are H2[x, i33-i85[x and io-i2(ji long, respectively. The trochanter of leg IV is without any process. HABITAT. Three males and four females have been obtained from pines and rhododendrons (Parknasilla, Co. Kerry, Ireland, Miss T. Clay, 27.1.1953); Holotype male (1963.2.7.17) and Allotype female (1963.2.7.18). Pergamasus (Paragamasus) diversus Halbert Gamasus (Pergamasus) diversus Halbert, J. N. (1915). Proc. R. Irish Acad. 31 39 ii : 52, figs. MALE. The dorsal shield (831-898^ long x 400-440^ wide) is well sclerotized and provided with a transverse suture (Text-fig. 162). The anterior region of the REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 193 dorsal shield is furnished with twenty-one pairs of setae and is regionally reticulated. The posterior region of the shield bears twenty-five pairs of setae and is entirely reticulated. 165 FIGS. 162-168. Pergamasus (Paragamasus) diver sus Halbert, male. Fig. 162 dorsum of idiosoma. Fig. 163 venter. Fig. 164 tectum. Fig. 165 trochanter, femur and genu of pedipalp. Fig. 166 chelicera. Fig. 167 venter of gnathosoma. Fig. 1 68 trochanter of leg IV. S. K. BHATTACHARYYA FIGS. 169-174. Pergamasus (Paragamasus) diversus Halbert, female. Fig. 169 dorsum of idiosoma. Fig. 170 venter. Fig. 171 endogynial process. Fig. 172 tectum. Fig. 173 chelicera. Fig. 174 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 195 The tritosternum has a short basal part and a pair of pilose laciniae. The genital lamina is medially notched into two lobes which have serrated margins (Text-fig. 163). Between coxae IV are a pair of well sclerotized, scale-like structures. Each scale-like structure is associated with a seta. The stigma is situated between coxae III and IV ; the peritreme extends to coxa II. A pair of post-stigmatal setae is present. The tectum is three-pronged (Text-fig. 164). The palpal trochanter has a seta- bearing process ; the femur and genu are provided with a comb-like and two spatulate setae respectively (Text-fig. 165). The chelicera is shown in Text-fig. 166. The corniculus is stalked and provided with a blade-like hyaline appendage (Text-fig. 167). The hypostomal process and deutosternal denticles are shown in the figure. The tibia, tarsus and prae-tarsus of leg I are 125-158(1,, I9r-2i3(j. and 12-17^ long, respectively. The femur of leg II is crassate with a strong falcate main spur and an accessory spur. The distal end of the femur also bears a tubular process. The genu lacks a process ; whilst the tibia bears a discoidal spur (Plate 3b) . The trochanter of leg IV is shown in Text-fig. 168. FEMALE. The dorsal shield (831-847^ long x 433-450^ wide) is sclerotized. The anterior region of the dorsal shield has twenty-one pairs of setae and the posterior region about twenty-four pairs (Text-fig. 169). The tritosternum has a well developed basal part. The distribution of the prae- endopodal shields is shown in Text-fig. 170. The posterior margin of the sternal shield has a median incision which extends to the level of the second pair of sternal setae. The metasternal shields are free and flank the epigynial shield. A pair of endogynial horns is present. The complicated endogynial processes are figured (Text-fig. 171). The genital shield is shown in Plate 7h. The opisthogastric shield bears ten pairs of setae (excepting three setae associated with the anus) . The position of the stigma and the extension of the peritreme are similar to that in the male. The tectum is three-pronged (Text-fig. 172). The palp is basically similar to that in the male. The dentition of the chelicera is shown in Text-fig. 173. The ventral groove of the gnathosoma has ten transverse rows of denticles (Text-fig. 174). The tibia, tarsus and prae-tarsus of leg I are I2i-i45(z, 170-191(0, and 8-i7(x long, respectively. k DISTRIBUTION AND HABITAT. The male of this species has previously been re- >rded only from Ireland (Halbert, 1915). I have examined specimens from : litter and soil under evergreen trees (St. gnes, Isles of Scilly, Cornwall, K. H. Hyatt, 5.^.1957). In leaf mould (Rye, Houghton Green, Sussex, J. T. Salmon, 19. v. 1951). Rotten wood (Blurridge Farm, Ridge Hill, Combe Martin, Devonshire, M. E. Bacchus, I2.iii.i957). Salix sp. litter (British Museum (Nat. Hist.), London, S. K. Bhattacharyya, 5.1^.1962). Rotten reeds (Colletts Bridge, Norfolk, P. N. Lawrence). Litter under thick evergreen hedge (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 29.^.1957). Pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Moss humus ig6 S. K. BHATTACHARYYA 178 180 FIGS. 175-180. Pergamasus (Paragamasus) femoratus sp. nov., male. Fig. 175 dorsum of idiosoma. Fig. 176 venter. Fig. 177 tectum. Fig. 178 trochanter of pedipalp. Fig. 179 chelicera. Fig. 180 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 197 and humus in small wood (Swansea University, Swansea, P. N. Lawrence, 26 . iii . 1962) . Moss on trees and humus (Hen Pare Wood, Killay, nr. River Clyne, Swansea, P. N. Lawrence, 24.^1.1962). Oak and beech litter (Singleton Park, Swansea, S. K. Bhattacharyya, 3.1.1961). Under pines and rhododendrons (Parknasilla, Co. Kerry, Ireland, Miss T. Clay, 27.1.1953). Pergamasus (Paragamasus) femoratus sp. nov. MALE. The dorsal shield (854^ long x 440 jj. wide) is moderately sclerotized and provided with a transverse suture (Text-fig. 175). The anterior dorsal shield has twenty-one pairs of setae and the posterior dorsal shield about twenty-six pairs. Ventrally, the tritosternal base and the proximal part of the pilose laciniae are covered by the genital lamina. The distribution of the prae-endopodal shields and the setation of the opisthogastral shield are shown in Text-fig. 176. The stigma is situated between coxae III and IV with the extension of the peritreme to coxa I. The tectum is three-pointed (Text-fig. 177). The palpal trochanter is shown in Text-fig. 178. The femur and genu is with one comb-like and two spatulate setae respectively. The characteristic features of the chelicera are shown in Text-fig. 179. The corniculus, hypostomal process and rows of deutosternal denticles are figured (Text-fig. 180). The tibia, tarsus and prae-tarsus of leg I are 145 y., 200-204^ and 41 [j. long, respec- tively. The femur, genu and tibia of leg II are armed with variously shaped processes (Plate 4a). The femur is characterized by the presence of a tubular spur near the proximal end of the segment, thus the name femoratus. The trochanter of leg IV lacks a tubercle. HABITAT. I have examined a single male (Holotype 1963.2.7.12) from mosses on rocks (Ballyporty Loughs, Co. Clare, Ireland, Coll. P. N. Lawrence, I2.vii.i96o). Pergamasus (Paragamasus) integer sp. nov. MALE. This species is reddish-brown in colour, well sclerotized and regionally reticulated. The dorsal shield (747^ long x 389^ wide) has no transverse suture (Text-fig. 181). It usually bears forty-six pairs of setae of which twenty pairs are situated on the anterior half of the dorsal shield. The number of setae varies on the posterior half of the dorsal shield. The tritosternum has a pair of pilose laciniae and its base is hidden by the genital sclerite. The distribution of the prae-endopodal shields is shown in Text-fig. 182. The setation of the opisthogastral shield is as figured. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. A pair of post-stigmatal setae, one on each side, is present. The tectum is as figured with three prongs, of which the median is much longer and broader based (Text-fig. 183). The palpal trochanter is shown in Text-fig. 184 ; a comb-like seta is present on the femur and the genu carries two spatulate setae. The fixed digit of the chelicera has a row of closely-set teeth and a simple 1 98 S. K. BHATTACHARYYA 186 FIGS. 181-186. Pergamasus (Paragamasus) integer sp. nov., male. Fig. 181 dorsum of idiosoma. Fig. 182 venter. Fig. 183 tectum. Fig. 184 trochanter of pedipalp. Fig. 185 chelicera. Fig. 186 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 199 dorsal seta. The movable digit is unidentate and shorter than the fixed digit (Text-fig. 185). The spermadactyl is normal. The corniculus is stalked. The entire lateral margins of the hypostomal processes are fringed. The deutosternal 189 FIGS. 187-191. Pergamasus (Paragamasus) integer sp. nov., female. Fig. 187 dorsum of idiosoma. Fig. 188 venter. Fig. 189 endogynial process. Fig. 190 tectum. Fig. 191 chelicera. 200 S. K. BHATTACHARYYA denticles are arranged in ten transverse rows (Text-fig. 186). The hypostome bears the usual four pairs of setae of which three pairs are distinctly pilose. The tibia, tarsus and prae-tarsus of leg I are 116-121^, 183-18711 and 17(0. long, respectively. The distinctive features of leg II are delineated in Plate 4b. The trochanter of leg IV is as drawn simple. FEMALE. The dorsal shield (889-967^ long x 500-565^ wide) is well sclerotized and its setation shown in Text-fig. 187. Ventrally, the tritosternum has an elongated basal part and a pair of pilose laciniae. A pair of large prae-endopodal shields is present (Text-fig. 188). The metasternal shields are free. The endogynium has a pair of horns. The endogynial process is shown in Text-fig. 189. The genital shield is shown in Plate y\. The opisthogastral shield bears ten pairs of setae (excepting three setae associated with the anus). The stigma lies between coxae III and IV ; and the peritreme extends to coxa I. The peritrematal shield runs for a short distance as a post-stigmatal prolongation. The tectum is three-pronged, the median prong being in advance of the lateral ones (Text-fig. 190). The pedipalp essentially resembles that of the male. The chelicera is as figured (Text-fig. 191). The tibia, tarsus and prae-tarsus of leg I are 125-129^, 208-215^ and i7-2i(j. long respectively. The trochanter of leg IV is without a special prominence. HABITAT. I have examined two males and two females from moss on trees and humus (Hen Pare Wood, Killay nr. River Clyne, nr. Swansea, P. N. Lawrence, 24.^1.1962). One female in leaf mould (Boltons Wood, Boltons Abbey, North Riding, York- shire, J. T. Salmon, 24. vi. 1951). Two females in oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). Three females in rotten wood from Pinus sp. in woodland belt (near St. Catherines Castle, Fowey, Cornwall, Mrs. R. E. Teagle, ii. 1957). Nine females and one male in moss on wall (Tarn Hows, N. Lancashire, M. E. Bacchus, 4.xii.i954). I have designated a male and female from Hen Pare Wood, Killay, nr. River Clyne, nr. Swansea, as the Holotype (1963.2.7.3) and Allotype (1963.2.7.6), respectively. Pergamasus (Paragamasus) lapponicus Tragardh Pergamasus lapponicus Tragardh, I. (1910). Naturw. Untersuch. Sarekgeb. 4 (4) : 408, figs. Cooreman, J. (1943). Bull. Mus. roy. Hist. nat. Belg., 19, 63 : 5, figs. ; Schweizer, J. (1949). Ergebn. wissens. Untersuch. Schweiz. Nationalparkes (n.s.) 2 : 28, figs ; Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 56, figs. MALE. The dorsal shield (797-831^ long x 400-417^ wide) has a transverse suture (Text-fig. 192) and is regionally reticulated. The anterior dorsal shield bears twenty-one pairs of setae and the posterior dorsal shield about twenty-eight pairs. The tritosternum has a pair of pilose laciniae but its base is completely hidden by the genital sclerite. The shape of the genital lamina is shown in Text-fig. 193. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 197 FIGS. 192-197. Pergamasus (Paragamasus) lapponicus Tragardh, male. Fig. 192 dorsum of idiosoma. Fig. 193 venter. Fig. 194 tectum. Fig. 195 trochanter, femur and genu of pedipalp. Fig. 196 chelicera. Fig. 197 trochanter of leg IV. The venter has a remarkable rounded, scaly, seta-bearing structure near coxa IV. The number of setae on the opisthogastric shield is shown in Text-fig. 193. The stigma is situated between the coxae III and IV ; the peritreme extends to coxa I. There is a post-stigmatal seta. 202 S. K. BHATTACHARYYA The tectum is three-pointed (Text-fig. 194). The palpal trochanter has a modified nipple-like setae-bearing process, at the base of which is a small protuberance ; a comb-like seta is present on the femur whilst the genu has two spatulate setae (Text-fig. 195). The distinctive shape of the fixed digit of the chelicera is shown in Text-fig. 196. The movable digit is bidentate, and the spermadactyl is normal. The corniculus is distinctly stalked. The entire lateral (outer) margins of the hypo- stomal processes are fringed. The tarsus (i87-2o8(ji) of leg I is longer than the tibia (i4i-i45(j.) ; the prae-tarsus is i7-2i[A long. The trochanter (Plate 4c) of leg II has a well developed spur ; the femur, genu and tibia are spurred (Plate 4d). The main process of the crassate femur is falcate. The armature of the trochanter of leg IV is figured (Text-fig. 197). FEMALE. The dorsal shield (764-797^ long x 400-467 [x wide) is more or less oval and yellow in colour and regionally reticulated. The anterior shield has only twenty-one pairs of setae and the posterior half bears about twenty-nine pairs (Text-fig. 198). The length of the dorsal setae vary considerably. Ventrally, the tritosternum has a pair of well developed pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 199. The inter- scutal membrane in between the anterior margin of the sternal shield and the prae-endopodal shield is striated. The metasternal shield is free. There is a pair of endogynial horns. The genital shields are shown in Plate 8h and i. The opistho- gastral shield is with ten pairs of setae. The stigma is situated between coxae III and IV ; and the peritreme reaches to the level of coxa I. The tectum is basically three pronged but may be variable (Text-fig. 200). The fixed digit of the chelicera has four teeth but the movable digit is tridentate (Text-fig. 201). The tibia, tarsus and prae-tarsus of leg I are 137-141^, 196-200^ and i6-i8(ji long, respectively. The trochanter of leg IV is figured (Text-fig. 202) . DISTRIBUTION AND HABITAT. This species is known from Swedish Lapland (Tragardh, 1910), Belgium (Cooreman, 1943), Germany and Switzerland (Schweizer, 1949, 1961). I have examined material from leaf mould (Epping Forest, Essex, J. T. Salmon, 27. v. 1951). Sphagnum, etc. (Cock Hill, Yorkshire, J. T. Salmon, 2i.vii.i95i). In marsh debris (Surlingham, Norfolk, J. T. Salmon, 12. v. 1951). In leaf mould (Savernake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). Litter and humus under pines and holly (near Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, 10 . ix . 1956) . In leaf mould (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951). From litter (Stratton Strawless, near Aylsham, Norwich, Norfolk, J. T. Salmon, 13 .v. 1951). In moss of wall (Tarn Hows, North Lancashire, M. E. Bacchus, 4 . xii . 1954) . In moss (Grassington, Grass Woods, North Riding, Yorkshire, J. T. Salmon, 24 . vi . 1951) . In Sphagnum (Virginia Water, Surrey, A. H. G. Alston, 18 . xii . 1951) . Sphagnum (Ascot, Berkshire, A. H. G. Alston, 8. xii. 1957). Moles' (Talpa europaea] nest, near pasture (Godstone, Surrey, Prof. P. A. Larkin, 4.xi.i962. Per C. Elton). Moss from trees and wall (Pont Hill, Cornwall, Mrs. R. E. Teagle, 24 . ii . 1957) . Litter under yew REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 203 FIGS. 198-202. Pergamasus (Paragamasus) lapponicus Tragardh, female. Fig. 198 dorsum of idiosoma. Fig. 199 venter. Fig. 200 tectum. Fig. 201 chelicera. Fig. 202 trochanter of leg IV. 204 S. K. BHATTACHARYYA (summit of Box Hill, Surrey, A. H. G. Alston, 15.1.1956). Hornbeam humus (Bookham Common, Surrey, Mrs. R. E. Teagle, xi . 1956) . Holly humus (near Groom- bridge, Sussex, P. N. Lawrence, 3 . iii . 1957) . In oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). Heath and birch (St. Faith's Common, Horsford, Norwich, Norfolk, J. T. Salmon, 13^.1951). In leaf mould (Leith Hill, Surrey, J. T. Salmon, i6.vi.i95i). In leaf mould (Richmond Park, Surrey, J. T. Salmon, 19. iii. 1951). Moss from crevices in rocks (near Force Forge, Westmorland, M. E. Bacchus, 2.xii.i_954). In leaf mould (Boltons Wood, Boltons Abbey, North Riding, Yorkshire, J. T. Salmon, 24. v. 1951). In leaf mould (Burnham Beeches, Bucks., J. T. Salmon, 3. vi. 1951). Oak humus (Gwydyr Forest, North Wales, P. N. Lawrence, 22 . x . 1961). Moss on boulder (Llandudno, North Wales, P. N. Lawrence, 12 . xi . 1961). Rhododendron humus (Swansea University, Swansea, P. N. Lawrence, 26. iii. 1962). Turf among bracken (Clyne Common, Swansea, P. N. Lawrence, 29.^.1962). Bracken litter (Bishopston Common, Swansea, S. K. Bhattacharyya, 4.1.1961). Floor of scotch fir wood (Aviemore, Inverness, J. T. Salmon, 7.vii.i95i). In moss (Tarbet, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3. vii. 1951). Pergamasus (Paragamasus) leruthi Cooreman Pergamasus leruthi Cooreman, J. (1951). Bull Mus. Hist. nat. Beige. 27, 42 : 5, figs. MALE. The dorsal shield (732(j. long x 382^ wide) is without a median transverse suture and yellow in colour. The distribution of setae on the dorsal shield is shown in Text-fig. 203. The tritosternum is provided with a pair of pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 204. The genital lamina is not produced into a median spine. There is a pair of undulating fissures which extend downward from the anterior margin of the sternal shield (Text-fig. 204) . The stigma is situated between coxae III and IV ; and the peritreme extends to coxa I. The post-stigmatal extension of the peritrematal shield reaches coxa IV. The tectum is three-pronged (Text-fig. 205). The palpal trochanter is without a prominence ; the femur and genu are provided with a comb-like and two spatulate setae, respectively. The terminal end of the fixed digit is slightly blunt and provided with a simple seta ; the movable digit is unidentate, with a spermadactyl process (Text-fig. 206). The corniculus is stalked. The tibia, tarsus and prae-tarsus of leg I are 126-166(1, 187-208(1 and 17-21(1 long, respectively. Leg II (Plate 4e) is remarkable for the peculiar shape of the spurs present on the genu and tibia. The trochanter of leg IV is as shown in Text- fig. 207. FEMALE. The dorsal shield (88251 long x 457(1 wide) is sclerotized. The distri- bution and relative lengths of the dorsal setae are shown in Text-fig. 208. The tritosternum has an elongated basal part and a pair of pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 209. The meta- sternal shields are free. The endogynium has a pair of horns. The endogynial process is very complicated (Text-fig. 210). The opisthogastral shield bears only REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 205 2O6 FIGS. 203-207. Pergamasus (Paragamasus) leruthi Cooreman, male. Fig. 203 dorsum of idiosoma. Fig. 204 venter. Fig. 205 tectum. Fig. 206 chelicera. Fig. 207 trochanter of leg IV. 2O6 S. K. BHATTACHARYYA 2O9 FIGS. 208-211. Pergamasus (Paragamasus) leruthi Cooreman, female. Fig. 208 dorsum of idiosoma. Fig. 209 venter. Fig. 210 magnified structure of the Sterniti-genital region. Fig. 211 chelicera, REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 207 nine pairs of setae (excepting three setae associated with the anus). The stigma is situated between coxae III and IV ; and the extension of the peritreme is not clearly distinguishable from the damaged specimen. The entire peritrematal shield is coalesced with the ventral shield. The post-stigmatal extension of the peritrematal shield reaches to coxa IV. The tectum is three-pronged. The pedipalp is similar to that of the male. The fixed digit of the chelicera is furnished with a row of closely-set teeth ; the movable digit is with four teeth (Text-fig. 211). The tibia, tarsus and prae-tarsus of leg I are 158-166^, 225-229^ and 21^ long, respectively. DISTRIBUTION AND HABITAT. This species has been recorded from Roumania (Cooreman, 1951). I have found only one male from Scotland, in dead leaves (Loch Lomond, between Glasgow and Tarbet, Dunbarton, J. T. Salmon, 8.vii.i95i). I have examined Cooreman's specimens of both sexes. The above description and figures of the female are based on the paratype. Pergamasus (Paragamasus) londonensis sp. nov. MALE. Unknown. FEMALE. The dorsal shield (474-483^ long x 267-283^ wide) is light yellow in colour and not heavily sclerotized. The anterior half of the dorsal shield bears twenty-one pairs of setae and the posterior half about twenty pairs (Text-fig. 212). Ventrally, the tritosternum has a long basal part and a pair of pilose laciniae. The distribution of the prae-endopodal shields in relation to the tritosternum is shown in Text-fig. 213. The metasternal shields are free. The endogynial structure is comparatively simple. The genital shield is shown in Plate 7]. The opisthogastral shield is provided with only eight pairs of setae (excepting the three setae associated with the anus). The stigma is situated between coxa III and IV ; and the peritreme extends to coxa I. The tectum is three-pronged (Text-fig. 214). The trochanter, femur and genu of the pedipalp are similar to those in P. rothamstedensis. The chelicera is shown in Text-fig. 215. The hypostomal processes and deutosternal denticles resemble those of the female of P. rothamstedensis. The tibia, tarsus and prae-tarsus of leg I are 70fx, 1 16-121 (x and 8-i2[x long, respectively. The trochanter of leg IV is devoid of prominences. HABITAT. I have examined four females from Salix sp. litter (British Museum (Nat. Hist.), London, S. K. Bhattacharyya, 5.^.1962) and have designated a female as the Holotype (1963.2.7.11). Pergamasus (Paragamasus) longisetosus sp. nov. MALE. Unknown. FEMALE. The dorsal shield (974-1,000^ long x 632^ wide) is reddish-brown in colour and strongly sclerotized. The dorsal setae are comparatively long (some of 208 S. K. BHATTACHARYYA 212 215 214 FIGS. 212-215. Pergamasus (Paragamasus) londonensis sp. nov., female. Fig. 212 dorsum of idiosoma. Fig. 213 venter. Fig. 214 tectum. Fig. 215 chelicera. them sparsely ciliated) and stout, and their distribution is shown in Text-fig. 216. The podonotal shield has an invariable number of setae (twenty-one pairs) but the number of setae on the opisthonotal shield is about twenty-eight pairs. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 209 Ventrally, the tritosternum has a pair of pilose laciniae. The prae-endopodal shields are present. The posterior margin of the sternal shield has a short median incision. The metasternal shields are free. The endogynium has a pair of horns and its process is comparatively simple. The setation of the opisthogastral shield is shown in Text-fig. 217. The stigma is situated between coxae III and IV ; the peritreme extends to the level of coxa I. The tectum is three-pronged, the median tine being longer than the lateral ones (Text-fig. 218). The most distinctive features of the chaetotaxy of the pedipalp FIGS. 216-219. Pergamasus (Paragamasus) longisetosus sp. nov., female. Fig. 216 dorsum of idiosoma. Fig. 217 venter. Fig. 218 tectum. Fig. 219 chelicera. S. K. BHATTACHARYYA 223 222 x FIGS. 220-223. Pergamasus (Paragamasus) minimus sp. nov., female. Fig. 220 dorsum of idiosoma. Fig. 221 venter. Fig. 222 magnified structure of the sterniti-genital region. Fig. 223 chelicera. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 211 are ; the comb-like seta on the femur and the two spatulate setae on the genu. The dentition of the chelicera is shown in Text-fig. 219. The deutosternal denticles are arranged in ten transverse rows. The tibia, tarsus and prae-tarsus of leg I are 149-154^, 204-225^ and i8(x long respectively. The trochanter of leg IV is without any prominence. HABITAT. One female has been obtained from inside the bark of a fallen elm tree (Harefield, Middlesex, A. H. G. Alston, 22.1.1956), and a single female from the Royal Botanic Gardens, Kew, Surrey, by John L. Gilbert (5.^.1961). I have designated the female from Harefield, Middlesex as the Holotype (1963.2.7.8). Pergamasus (Paragamasus) minimus sp. nov. MALE. Unknown. FEMALE. The dorsal shield (500^ long x 265^ wide) is very weakly sclerotized. The dorsal setae are comparatively short (Text-fig. 220). The tritosternum has an elongated base and a pair of well-developed pilose laciniae. The distribution of the prae-endopodal shields is delineated in Text-fig. 221. The metasternal shields are free. The endogynium has a pair of horn-like structures (Text-fig. 222). The opisthogastric shield has ten pairs of setae (excepting three setae associated with the anus). The stigma is present, between coxae III and IV ; and the peritreme extends as far as the level of coxa I. The tectum is three-pronged. The most distinctive features of the pedipalp are ; the femur is with a comb-like and the genu with a spatulate seta, respectively. The dentition of the chelicera is shown in Text-fig. 223. Approximately half of the lateral margins of the hypostomal processes are fringed. The deutosternum of the ventral groove of the gnathosoma has nine (ten?) transverse rows of denticles. Leg I with the tibia 77^ and tarsus 121^ long, the prae-tarsus is 8(A long. The trochanter of leg IV is tuberculated. HABITAT. I have examined a single female (Holotype 1963 : 2.7.19) from soil and compost (Tettenhall, Stafford, Dr. D. W. Empson, 4.1^.1959). Pergamasus (Paragamasus) misellus Berlese Gamasus (Pergamasus) misellus Berlese, A. (1904). Redia 1 : 237 ; Berlese, A. (1906). Redia 3 : 207, figs. Pergamasus misellus : Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 49, figs. ; Karg, W. (1962). Mitt. Zoo/. Mus. Berlin 38 : 77, figs. MALE. The dorsal shield is 515-548(0, long and 283^ wide and is provided with a transverse suture (Text-fig. 224). The anterior dorsal shield bears twenty-one pairs of setae. The tritosternum, prae-endopodal shields and the shape of the genital lamina is shown in Text-fig. 225. The stigma is situated between coxae III and IV ; and the peritreme extends to coxa II. A pair of post-stigmatal setae is present. S. K. BHATTACHARYYA 23O FIGS. 224-230. Pergamasus (Paragamasus) misellus Berlese, male. Fig. 224 dorsum of idiosoma. Fig. 225 venter. Fig. 226 tectum. Fig. 227 trochanter, femur and genu of pedipalp. Fig. 228 chelicera. Fig. 229 venter of gnathosoma. Fig. 230 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 213 FIGS. 231-234. Pergamasus (Paragamasus) misellus Berlese, female. Fig. 231 dorsum of idiosoma. Fig. 232 venter. Fig. 233 magnified structure of the sterniti-genital region. Fig. 234 chelicera. 2i 4 S. K. BHATTACHARYYA The tectum is three-pronged (Text-fig. 226). The distinctive features of the pedipalp are shown in Text-fig. 227. The chelicera is as in Text-fig. 228. The form of the hypostomal processes and rows of deutosternal denticles are shown in Text-fig. 229. The tarsus of leg I (i29-i33[x) is longer than the tibia (78-83^) ; the prae-tarsus being 8^ long. The femur, genu and tibia are spurred (Plate 5a). A tubercle is present on the trochanter of leg IV (Text-fig. 230) . FEMALE. The dorsal shield measures 565-582^ long and 332-339^ wide. The anterior region of the dorsal shield has twenty-one pairs of setae and the posterior region about twenty-five pairs (Text-fig. 231). Ventrally, the tritosternum has an elongated basal part and a pair of pilose laciniae. The prae-endopodal shields are distributed as in Text-fig. 232. The metasternal shields are free ; and a pair of endogynial horns is present. The enlarged view of the genital region is shown in Text-fig. 233. The genital shield is shown in Plate 8a. There are ten pairs of opisthogastral setae (excepting a pair of paranal and a postanal seta) . The position of the stigma and the extension of the peritreme are similar to the male. The tectum has three prongs. The chelicera is delineated in Text-fig. 234. The tibia, tarsus and prae-tarsus of leg I are 74^, 125^, and 8(j, long respectively. The trochanter of leg IV has a tubercle. DISTRIBUTION AND HABITAT. P. misellus has been previously recorded from Italy (Berlese, 1904, 1906), Switzerland (Schweizer, 1961) and Germany (Karg, 1962). I have examined specimens from litter under Cupressus sp. (Rothamsted Lodge, Rothamsted Experimental Station, Harpenden, Hertfordshire, G. Owen Evans and E. Browning, i8.iii.i957). Salix sp. litter, British Museum (Nat. Hist.), London, S. K. Bhattacharyya, 5.^.1962). Turf among bracken (Clyne Common, Swansea, P. N. Lawrence, 29.^.1962). Humus in small wood (Swansea University, P. N. Lawrence, 26.^.1962). Pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Bracken litter (Bishopston Common, Swansea, S. K. Bhattacharyya, 4.1.1961). Pergamasus (Paragamasus) nathistmus sp. nov. MALE. The dorsal shield (374[x long x 183(0. wide) is weakly sclerotized, regionally reticulated and somewhat oval in shape. The dorsum is devoid of a transverse suture (Text-fig. 235). Ventrally, there are two large prae-endopodal shields flanking the genital sclerite and partly covered by the genital lamina. The genital lamina has essentially the same shape as in P. rothamstedensis sp.n. The basal part of the tritosternum is completely hidden by the genital sclerite. The chaetotaxy of the venter is shown in Text-fig. 236. The stigma is situated between coxae III and IV ; the peritreme extends to coxa I. The post-stigmatal extension of the peritrematal shield is not clearly discernible. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 215 236 238 FIGS. 235-238. Pergamasus (Paragamasus) nathistmus sp. nov., male. Fig. 235 dorsum of idiosoma. Fig. 236 venter. Fig. 237 tectum. Fig. 238 venter of gnathosoma. 216 S. K. BHATTACHARYYA The tectum is three-pronged, the centre prong being longer than the lateral ones (Text-fig. 237). The palp trochanter has no tubercle. The femur has a comb-like seta, and the genu two spatulate setae. Owing to the small size and weakly sclerotized body of the single specimen it did not seem advisable to risk the dissection of the chelicera, so that its dentition is not included in the description. The gnathosoma is shown in Text-fig. 238. The tibia, tarsus and prae-tarsus of leg I are 60-62^1, 95^ and 8-iOfx respectively. Leg II is very weakly spurred (Plate 5b). The femur is not crassate and its main spur is as long as the tibial process. HABITAT. I have examined a single male (Holotype 1963 : 2.7.9) from Salix sp. litter, British Museum (Natural History), London (Coll. S. K. Bhattacharyya, 14.^.1962). Pergamasus (Paragamasus) parrunciger sp. nov. MALE. The dorsal shield (797[x long x 374-38251, wide) is rather weakly sclerotized and provided with a transverse suture (Text-fig. 239). The anterior dorsal shield bears twenty-one pairs of setae and is regionally reticulated. The posterior dorsal shield has about twenty-eight pairs of setae. The tritosternal base is extremely reduced and provided with a pair of pilose laciniae. The distribution of the prae-endopodal shields and the shape of the genital lamina is shown in Text-fig. 240. The setation of the opisthogastral shield is as figured. The stigma is situated between coxae III and IV. The peritreme extends to coxa I. A pair of post-stigmatal setae is present. The tectum is three-pronged (Text-fig. 241). The median tine has a broader base than those of the lateral ones. The trochanter, femur and genu of the pedipalp is shown in Text-fig. 242. The chelicera is robust and its dentition is delineated in Text-fig. 243. Some of the processes of the arthrodial membrane at the base of the movable digit are branched. There is a simple dorsal seta on the fixed digit. The corniculus is distinctly stalked. The lateral margins of the hypostomal processes are fringed. The tibia, tarsus and prae-tarsus of leg I are 103-128^, 158-187^ and i4-i7(x long respectively. The femur, genu and tibia of leg II are spurred (Plate 5c and e). The femur is crassate. The tibial process may vary in form (Plate 5d). The trochanter of leg IV is simple. FEMALE. The dorsal shield (747-789^ long x 440-467^ wide) is broadly oval and sclerotized. The anterior half of the dorsal shield bears twenty-one pairs of setae and the posterior half is with twenty-eight. The length of the setae is variable in the posterior half (Text-fig. 244) . Ventrally, the tritosternum has a pair of pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 245. The interscutal membrane in between the anterior margin of the sternal and prae-endopodal shields is striated. The endogynium has a pair of horns. The genital shield is shown in Plate 8j. The opisthogastral shield bears ten pairs of setae (excepting three setae associated with REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 217 242 FIGS. 239-243. Pergamasus (Paragamasus) parrunciger sp. nov., male. Fig. 239 dorsum of idiosoma. Fig. 240 venter. Fig. 241 tectum. Fig. 242 trochanter, femur and genu of pedipalp. Fig. 243 chelicera. 218 S. K. BHATTACHARYYA the anus). The stigma is situated between coxae III and IV and the peritreme extends to the level of coxae I. The peritrematal shield extends for a short distance behind the stigma. The tectum has three prongs of equal length (Text-fig. 246). The palpal femur and genu are with one comb-like and two spatulate setae, respectively. The fixed 246 FIGS. 244-247. Pergamasus (Paragamasus) parrunciger sp. nov., female. Fig. 244 dorsum of idiosoma. Fig. 245 venter. Fig. 246 tectum. Fig. 247 chelicera. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 219 digit of the chelicera has four well-developed teeth ; the movable finger is tridentate (Text-fig. 247). Approximately the proximal half of the hypostomal processes are fringed. The tibia, tarsus and prae-tarsus of leg I are 112-129^, 175-179^ and 12-14^ long respectively. The trochanter of leg IV is without a tubercle. HABITAT. I have examined five males and nine females from leaf mould (Leith Hill, Surrey, J. T. Salmon, 16. vi. 1951) and have designated a male as the Holotype (1963.2.7.19) and a female as the Allotype (1963.2.7.20). Pergamasus (Paragamasus) rothamstedensis sp. nov. MALE. The dorsal shield (417^ long x 200-250^ wide) is light yellow in colour and oval in shape. The dorsum is devoid of a transverse suture and the posterior half of the dorsal shield bears a variable number of setae. The setae of the dorsal shield are comparatively short (Text-fig. 248). Ventrally, the distribution of the prae-endopodal shields is shown in Text-fig. 249. The tritosternum is with a pair of pilose laciniae but its base is completely hidden by the genital sclerite. The anterior margin of the genital lamina is smooth. The distribution of setae on the opisthogastric shield is figured. The stigma is situated between coxae III and IV ; the peritreme anteriorly extends as far as the level of coxa I. Posteriorly the peritrematal shield extends to coxa IV. The post- stigmatal seta is absent. The tectum is three pointed. The median prong is the longest and is blunt (Text- fig. 250). The fixed digit of the chelicera (Text-fig. 251), is tridentate, and the movable digit is bidentate. The palp trochanter is without a tubercle. The corniculus is distinctly stalked (Text-fig. 252). The lateral margins of the hypostomal processes, excepting the basal part, are fringed. The deutosternum has ten trans- verse rows of denticles. Leg I has the tibia, tarsus and prae-tarsus 62-66^, 95-108(1, and 8-i2[x long, respectively. The femur of leg II (Plate 51), is with a main and an accessory spur ; and the genual spur is less developed than the tibial process. The trochanter of leg IV is simple. FEMALE. The dorsal shield (450-483^ long X 25-275^ wide) is weakly sclerotized and oval in shape (excepting the anterior region) (Text-fig. 253). The anterior half of the dorsal shield has twenty-one pairs of setae and the posterior half about twenty- two pairs. There are three prae-endopodal shields, as shown in Text-fig. 254. The trito- sternum has a pair of pilose laciniae. The metasternal shields are free. The epigynial shield is as shown in Plate 7k. The opisthogastric shield has eight pairs of setae (excepting the three anal setae). The stigma is situated between coxae III and IV ; and the peritreme extends to coxa I. The tectum is three-pronged, the median being the longest (Text-fig. 255). The pedipalp is essentially similar to that in the male. The dentition of the chelicera is S. K. BHATTACHARYYA 251 252 FIGS. 248-252. Pergamasus (Paragamasus) rothamstedensis sp. nov., male. Fig. 248 dorsum of idiosoma. Fig. 249 venter. Fig. 250 tectum. Fig. 251 chelicera. Fig. 252 venter of gnathosoma. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 221 256 257 FIGS. 253-257. Pergamasus (Paragamasus) rothamstedensis sp. nov., female. Fig. 253 dorsum of idiosoma. Fig. 254 venter. Fig. 255 tectum. Fig. 256 chelicera. Fig. 257 venter of gnathosoma. 222 S. K. BHATTACHARYYA shown in Text-fig. 256. About half of the lateral margins of the hypostomal processes are fringed. The arrangement of the transverse rows of denticles is figured (Text-fig. 257). The tibia, tarsus and prae-tarsus of leg I are 66-74(1,, io6-i2i(x and io-i2(x long respectively. The trochanter of leg IV is without any tubercle. HABITAT. I have examined four females from litter under Cupressus sp. (Rothamsted Lodge, Rothamsted Experimental Station, Harpenden, Hertfordshire, G. Owen Evans and E. Browning, i8.iii.ig57), an d two males and four females from Salix sp. litter (British Museum (Natural History), London, S. K. Bhatta- charyya, 5.^1.1962). I have designated a male and female from the British Museum (Nat. Hist.), London (Coll. S. K. Bhattacharyya, 5.^.1962) as the Holotype (1963.2.7.13) and Allotype (1963.2.7.14) respectively. Pergamasus (Paragamasus) runciger (Berlese) Parasitus longulus (C. L. Koch) Oudemans, A. C. (1902). Tijdschr. Ent. 45 : 37, figs. Gamasus (Pergamasus) runciger Berlese, A. (1904). Redia 1 : 263 ; Berlese, A. (1906). Redia 3 : 214, figs. : Halbert, J. N. (1915). Proc. R. Irish Acad. 31 39 ii : 51, figs. Pergamasus runciger : Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 54, figs, (in part). MALE. The dorsal shield (838^ long x 433^ wide) is well sclerotized and provided with a transverse suture. The distribution of the setae on the dorsum is shown in Text-fig. 258. The tritosternum has a pair of pilose laciniae and its base is greatly reduced. The prae-endopodal shield and shape of the genital lamina is shown in Text-fig. 259. The stigma is situated between coxae III and IV ; and the peritreme extends to coxa I. A pair of post-stigmatal setae is present. The tectum is three-pointed (Text-fig. 260). The distinctive features of the pedipalp are, the trochanter with a seta-bearing tubercle (Text-fig. 261), the femur with a comb-like and the genu with two spatulate setae. The chelicera is shown in Text-fig. 262. The tibia, tarsus and prae-tarsus of leg I are Ii2-i2i[j,, 150-154^ and i2(ji long, respectively. The femur, genu and tibia of leg II are spurred (Plate 5g and h). The femur is crassate with a main falcate spur and an accessory spur. The trochanter of leg IV is simple. FEMALE. The dorsal shield (8i5[x long x 450-483^ wide) is oval (excepting the anteriormost region) and bears about forty-eight pairs of setae of which twenty-one pairs are situated on the podonotal shield (Text-fig. 263). Ventrally, the tritosternum has a well-developed basal part and a pair of pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 264. The metasternal shields are free. The endogynium has a pair of horns. The epigynial shield and details of the sterniti-genital region are shown in Plate 8k and Text-fig. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 223 FIGS. 258-262. Pergamasus (Paragamasus) runciger (Berlese), male. Fig. 258 dorsum of idiosoma. Fig. 259 venter. Fig. 260 tectum. Fig. 261 trochanter of pedipalp. Fig. 262 chelicera. 265 respectively. Ten pairs of setae are situated on the opisthogastral shield. The position of the stigma and extension of the peritreme are similar to that in the male. The tectum is three-pronged. The dentition of the chelicera is shown in Text- fig. 266. 224 S. K. BHATTACHARYYA FIGS. 263-266. Pergamasus (Paragamasus) runciger (Berlese), female. Fig. 263 dorsum of idiosoma. Fig. 264 venter. Fig. 265 magnified structure of the sterniti-genital region. Fig. 266 chelicera. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 225 The tibia, tarsus and prae-tarsus of leg I are 121-137^, 170-191^ and 12-17^ long, respectively. The trochanter of leg IV is without any process. DISTRIBUTION AND HABITAT. This species has been recorded from Holland (Oudemans, 1902), Norway (Berlese, 1904, 1906), Ireland (Halbert, 1915) and Switzerland (Schweizer, 1961). I have examined specimens from litter and humus under pines and holly (near Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, io.ix.i956). Moss and leaves (Adelaide Hill, Bowness, Cumberland, J. T. Salmon, 29. vi. 1951). In marsh debris (Surlingham, Norfolk, J. T. Salmon, 12. v. 1951). Sphagnum, etc. (Cock Hill, Yorks, J. T. Salmon, 2i.vii.i95i). In leaf mould (Epping Forest, Essex, J. T. Salmon, 27. v. 1951). In litter (Stratton Strawless, near Aylsham, Norwich, Norfolk, J. T. Salmon) . Hornbeam humus (Bookham Common, Surrey, Mrs. R. E. Teagle, xi . 1956) . In oak and beech leaf mould (Grasmere, Westmorland, J. T. Salmon, 26. vi. 1951). Algae and rushes (Colletts Bridge, Norfolk, P. N. Lawrence, i6.x.i96o). Beech leaf mould (Amersham, Bucks., M. E. Bacchus, 13. v. 1951). Moss from crevices in rocks (near Force Forge, Westmorland, M. E. Bacchus, 2.xii.i95i). Mosses on rotten wood (Killay, near River Clyne, Hen Pare Wood, Swansea, P. N. Lawrence, 24.^1.1962). Yew and rhododendron humus (26.1^.1962) and moss on tree and humus underneath (Killay, near River Clyne, Swansea, P. N. Lawrence, 24.^.1962). Pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Pergamasus (Paragamasus) schweizeri sp. nov. MALE. The dorsal shield (697^ long x 332-350(1, wide) is provided with a transverse suture and tapers posteriorly (Text-fig. 267). The tritosternal base is completely hidden by the genital lamina. The distribution of the prae-endopodal shields and the shape of the genital lamina is shown in Text- fig. 268. The stigma is situated between coxae III and IV ; and the peritreme anteriorly extends to coxa I. A pair of post-stigmatal setae is present. The tectum is three-tined (Text-fig. 269). The palp trochanter has two tubercles, and one of them (the distal) bears a simple seta (Text-fig. 270). The palpal femur and genu are provided with one comb-like and two spatulate setae, respectively. The dentition of the chelicera and the form of the spermadactyl process are shown in Text-fig. 271. There are stalked corniculi. The entire lateral margins of the hypostomal processes are fringed (Text-fig. 272). The tibia, tarsus and prae-tarsus of leg I are io8-i2i[x, 158-170^ and I2(x long, respectively. The crassate femur of leg II is provided with a strong falcate spur and a small accessory spur ; the genu and tibia are armed (Plate 6a). The distinc- tive feature of the trochanter of leg IV is constant in all males of the species examined (Text-fig. 273). FEMALE. The dorsal shield measures 764-797^ long x 367-400^ wide and is provided with about forty-eight pairs of setae (Text-fig. 274). The tritosternum has a pair of pilose laciniae. Ventro-medially, the interscutal membrane between the prae-endopodal shields and the anterior margin of the 226 S. K. BHATTACHARYYA FIGS. 267-273. Pergamasus (Paragamasus) schweizeri sp. nov., male. Fig. 267 dorsum of idiosoma. Fig. 268 venter. Fig. 269 tectum. Fig. 270 trochanter, femur and genu of pedipalp. Fig. 271 chelicera. Fig. 272 venter of gnathosoma. Fig. 273 trochanter of leg IV. REVISION OF THE GENUS PERGA MAS US BERLESE S. LAT. 227 sternal shield is striated (Text-fig. 275). The metasternal shields are free. The endogynium has a pair of horns and its process is comparatively simple. The genital shield is shown in Plate 81. The opisthogastral shield is normal, bearing ten pairs of setae. The stigma and peritreme are similar to those in the male. The tectum is basically three-pronged but shows some variation (Text-fig. 276). The dentition of the chelicera is shown in Text-fig. 277. 274 275 277 FIGS. 274-277. Pergamasus (Paragamasus) schweizeri sp. nov., female. Fig. 274 dorsum of idiosoma. Fig. 275 venter. Fig. 276 tectum. Fig. 277 chelicera. 228 S. K. BHATTACHARYYA The tibia, tarsus and prae-tarsus of leg I are 116-129^, 170-181^ and 12-17^ long, respectively. HABITAT. I have examined the following material : Seven males and four females from litter under thick evergreen hedge (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 29.^.1957). Twenty-two males and twenty females in litter and soil under evergreen trees (St. Agnes, Isles of Scilly, Cornwall, K. H. Hyatt, 5.^.1957). One male and three females in moss (Grassington, Grass Woods, North Riding, Yorkshire, J. T. Salmon, 24. vi. 1951). Two males and two females from moss on tree stumps and stones (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951). One male from Stratton Strawless (near Aylsham, Norwich, Norfolk, J. T. Salmon, 13. v. 195?). Three males and two females from leaf mould (Forest of Dean, Gloucestershire, J. T. Salmon, 27. ix. 1951). One male from litter and humus under pines and holly (near Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, 10 . ix . 1956). Six males and eight females from lichens in crevices (Moel Siabod, North Wales, P. N. Lawrence, 22.x. 1960) . One male in mosses on rotten wood (Killay, near River Clyne, Hen Pare Wood, near Swansea, P. N. Lawrence, 24.iii.i962). One male from rotten stump (Gwydyr Forest, North Wales, P. N. Lawrence, 22.x. 1960). Two males and two females in moss (Tarbet, Crianlarich, Loch Lomond, Dunbarton, J. T. Salmon, 3.vii.i95i). I have designated a male and a female from St. Agnes, Isles of Scilly (Coll. K. H. Hyatt, 29 . iii . 1957) as Holotype (1963 .2.7.4) an d Allotype (1963 .2.7.5) respectively. Pergamasus (Paragamasus) suecicus (Tragardh) Leptogamasus suecicus Tragardh, I. (1936). Ent. Tidskr. 227, figs. Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 10, figs. MALE. The dorsal shield (450-467^ long x 240-249^ wide) is deep yellow, well sclerotized and not entirely reticulated. The dorsum is without a median transverse suture and narrows posteriorly (Text-fig. 278). Ventrally, the tritosternum has a pair of well developed pilose laciniae. The basal part of the tritosternum is completely hidden by the genital sclerite. The prae-endopodal shields and pilose laciniae are also partly covered by the genital lamina. The setation of the opisthogastral shield is shown in Text-fig. 279. The stigma is situated between coxae III and IV ; the peritreme reaches to coxa II. The post-stigmatal prolongation of the peritrematal shield extends to coxa IV. The tectum is three-pronged (Text-fig. 280). The most distinctive features of the pedipalp are, the trochanter without any tubercle, the femur with a comb-like seta and the genu with two spatulate setae (Text-fig. 281). The fixed digit of the chelicera is provided with closely-set teeth of variable size ; a pilus dentilis and a simple dorsal seta are also present (Text-fig. 282). The movable finger is unidentate and the spermadactyl is normal. A number of arthrodial processes, at the base of the movable digit, are branched. REVISION OF THE GENUS PERGA MAS US BERLESE S. LAT. 229 I MM ' i ? u 252 281 FIGS. 278-282. Pergamasus (Paragamasus) suecicus (Tragardh), male. Fig. 278 dorsum of idiosoma. Fig. 279 venter. Fig. 280 tectum. Fig. 281 trochanter, femur and genu of pedipalp. Fig. 282 chelicera. 230 S. K. BHATTACHARYYA The tibia, tarsus and prae-tarsus of leg I are 74-78^, 109-112^ and 8-10^ long, respectively. The femur, genu and tibia of leg II are spurred (Plate 6b). The femur is crassate, with a strong falcate main spur and an accessory spur. The genual spur is smaller than the tibial spur and almost rounded in shape. Trochanter IV is weakly tuberculated. FIGS. 283-287. Pergamasus (Paragamasus) suecicus (Tragardh), female. Fig. 283 dorsum of idiosoma. Fig. 284 venter. Fig. 285 tectum. Fig. 286 chelicera. Fig. 287 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAX. 231 FEMALE. The shape and setation of the dorsal shield (474-490^ long x 249-256^ wide) are shown in Text-fig. 283. The dorsal shield bears about forty-seven pairs of setae of which twenty-one pairs are situated on the anterior half. The number of setae is only variable on the posterior half of the dorsal shield. Ventrally, the tritosternum has an elongated basal part and a pair of pilose laciniae. There are three pairs of prae-endopodal shields, distributed as in Text-fig. 284. The metasternal shields are completely fused. Anteriorly, the epigynial shield is rounded in shape, except the anterior extremity, where the shield terminates in a knob-like structure (Plate 7!). The opisthogastral shield is somewhat reduced and bears nine pairs of setae (excepting three setae associated with the anus). The peritrematal shield is completely coalesced with the opisthogastric shield posteriorly ; the peritreme only reaches to coxa II. The narrow, tubular post- stigmatal prolongation of the peritrematal shield extends to coxa IV. The tectum is three-pronged (Text-fig. 285). The pedipalp is essentially similar to that in the male. The dentition of the fixed digit of the chelicera is figured, and the movable digit has four teeth (Text-fig. 286). The tibia, tarsus and prae-tarsus of leg I are 78-83^, i25[A and 8^ long, respec- tively. The distinctive feature of trochanter IV is shown in Text-fig. 287. DISTRIBUTION AND HABITAT. This species is previously known from Sweden (Tragardh, 1936), Britain (Evans, 1957) and Switzerland (Schweizer, 1961). I have examined specimens from soil (Silwood Park, Berkshire, D. K. Choudhuri, 1958.2.13.12-16). (Corby, Northants, B. N. K. Davis, 1958). Larch litter (Merlewood, Lancashire, D. Macfarlane, v.1955). In moss and humus in hollow tree (Swansea University, Swansea, P. N. Lawrence, 26.^1.1962). Moss on cliff (Gwydyr Forest, North Wales, P. N. Lawrence, 22.x. 1960). Pergamasus (Paragamasus) teutonicus Willmann v Pergamasus lapponicus Tragardh var. teutonica Willmann, C. (1956). Ceskoslov. parasit. 33 : 215, figs. MALE. The dorsal shield is 697-732^ long x 350-417^ wide and has a transverse suture. The anterior shield bears twenty-one pairs of setae and the posterior half about twenty-six pairs (Text-fig. 288). The distribution of the prae-endopodal shields is delineated in Text-fig. 289. The base of the tritosternum and its laciniae are covered by the genital sclerite and lamina. The most distinctive features of the venter are the presence of a rounded setae-bearing scale-like structure near the coxa IV. The setation of the opistho- gastric shield is figured. The stigma is situated between coxae III and IV ; and the peritreme reaches to the level of coxa I. A pair of post-stigmatal setae is present. The tectum is as figured (Text-fig. 290). The palp-trochanter and the dentition of the chelicera are shown in Text-figs. 291 and 292, respectively. The corniculus is stalked. 232 S. K. BHATTACHARYYA FIGS. 288-292. Pergamasus (Paragamasus) teutonicus Willmann, male. Fig. 288 dorsum of idiosoma. Fig. 289 venter. Fig. 290 tectum. Fig. 291 trochanter of pedipalp. Fig. 292 chelicera. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 233 293 294 295 FIGS. 293-296. Pergamasus (Paragamasus) teutonicus Willmann, female. Fig. 293 dorsum of idiosoma. Fig. 294 venter. Fig. 295 tectum. Fig. 296 chelicera. 234 S. K. BHATTACHARYYA The tibia, tarsus and prae-tarsus of leg I are 104-121^, 164-175^ and long, respectively. The trochanter of leg II is without any tubercle. The femur, genu and tibia are armed with spurs (Plate 6c). The trochanter of leg IV may be with a weakly developed tubercle. FEMALE. The dorsal shield (697-747^ long x 389-433^ wide) is oval, and yellow in colour and regionally reticulated. The anterior half of the dorsal shield has twenty-one pairs of setae and the posterior half about twenty-seven pairs (Text-fig. 293). The tritosternum has an elongated base and a pair of pilose laciniae. The distribution of the prae-endopodal shields is shown in Text-fig. 294. The interscutal membrane between the anterior margin of the sternal shield and the prae-endopodal shield is striated. The reticulation of the sternal shield is figured. The metasternal shields are free. The endogynium has a pair of horns. The genital shield is as in Plate 8m and n. The opisthogastral shield, with ten pairs of setae (excepting three setae associated with the anus). The position of the stigma and extension of the peritreme are similar to those in the male. The tectum is three-pronged (Text-fig. 295). The pedipalp is essentially similar to that of the male except for the absence of the armature on the trochanter. The dentition of the chelicera is as figured (Text-fig. 296). The proximal half of the hypostomal margins are fringed. The deutosternal denticles are arranged in nine to ten transverse rows. The tibia, tarsus and prae-tarsus of leg I are 104-125^, 160-183^ and i2-2i[x long, respectively. The trochanter of leg IV is without any tubercle. DISTRIBUTION AND HABITAT. Only the male of this species has previously been recorded from Czechoslovakia (Willmann, 1956). I have examined material from : under bark of fallen elm tree (Harefield, Middlesex, A. H. G. Alston). Beech leaf mould (Amersham, Buckinghamshire, M. E. Bacchus, 13 .v. 1951). In beech leaf mould from steep bank (Rickmansworth, Herts., M. E. Bacchus, 25.^.1951). Soil in aquarium house (Botanic Garden, Cambridge, A. H. G. Alston, 13.^.1957). In pine litter (Clyne Park, Swansea, S. K. Bhattacharyya, 4.1.1961). Moss and humus in hollow tree (Swansea University, P. N. Lawrence, 26.^.1962). Pergamasus (Paragamasus) truncus Schweizer Pergamasus truncus Schweizer, J. (1961). Denks. schweiz. naturf. Ges. 84 : 44, figs. MALE. The dorsal shield (515^ long x 240-250^ wide) has a median transverse suture and is lightly sclerotized. The anterior region of the dorsal shield bears twenty-one pairs of setae. The dorsum has comparatively short setae. The tritosternum has a pair of pilose laciniae. The genital lamina is somewhat similar to that of P. wasmanni. The stigma is situated between coxae III and IV ; the peritreme extends to coxa II. The post-stigmatal seta is present. The tectum is three-tined, the median prong being the shortest (Text-fig. 297). The most distinctive features of the pedipalp are : the trochanter is without a REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 235 298 301 FIGS. 297-301. Pergamasus (Paragamasus) truncus Schweizer. Fig. 297 tectum of male. Fig. 298 chelicera of male. Fig. 299 dorsum of idiosoma in female. Fig. 300 venter of female. Fig. 301 chelicera of female. tubercle, the comb-like seta on the femur and the two spatulate setae on the genu are present. The chelicera is shown in Text-fig. 298. The corniculus is distinctly stalked. The tibia, tarsus and prae-tarsus of leg I are 74-78^, 108-129^ and io-i2[ji long, respectively. The distinctive spurs of leg II are delineated in Plate 6d. The 236 S. K. BHATTACHARYYA trochanter of leg IV is weakly spurred and the process somewhat resembles that in P. misellus. FEMALE. The dorsal shield (532[x long x 267^ wide) is yellow in colour and provided with comparatively short setae (Text-fig. 299) . The anterior region of the dorsal shield bears twenty-one pairs of setae and the posterior region twenty-four pairs. Ventrally, the tritosternum has an elongated basal part and a pair of pilose laciniae. The distribution of prae-endopodal shields is shown in Text-fig. 300. The endogynium has a pair of horns ; the genital shield is shown in Plate 8b. The opisthogastral shield is with ten pairs of setae (excepting three setae associated with the anus). The stigma is situated between coxae III and IV. Although the peritrematal shield anteriorly extends to the level of coxa I, the peritreme only reaches to coxa II. The tectum is three-tined. The pedipalp is essentially the same as in the male. The dentition of the chelicera is shown in Text-fig. 301. The tarsus (n6-i29(x) of leg I is longer than the tibia (74-80^). The prae-tarsus is 6-8[j. long. The trochanter of leg IV is armed. DISTRIBUTION AND HABITAT. This species has been recorded only from Switzer- land (Schweizer, 1961). I have examined specimens from a number of localities in Wales : Turf among bracken (Clyne Common, Swansea, P. N. Lawrence, 29.111.1962). Salt marsh debris (Llanrhidian, Glamorgan, P. N. Lawrence, 27.111.1962). Moorland mosses, Sphagnum by stream (S. Llanrhidian, S. Wales, P. N. Lawrence, 27.111.1962). Mosses on rotten wood (Killay, near River Clyne, Hen Pare Wood, Swansea, P. N. Lawrence, 24.111.1962). Oak litter (Llew Reservoir, Swansea, S. K. Bhattacharyya, 2.1.1961). Pergamasus (Paragamasus) wasmanni (Oudemans) Parasitus wasmanni Oudemans, A. C. (1902). Tijdschr. Ent. 45 : 39, figs. MALE. The dorsal shield (467-483^ long x 240-274^ wide) has a transverse suture (Text-fig. 302). The anterior half of the dorsal shield bears twenty-one pairs of setae. The tritosternum has a reduced basal part and a pair of pilose laciniae. There is a pair of prae-endopodal shields flanking the genital sclerite. The shape of the genital lamina and the setation of the opisthogastric shield are shown in Text-fig. 303. The stigma is situated between coxae III and IV ; and the peritreme extends to coxa II. A pair of post-stigmatal setae is present. The tectum is three-pronged (Text-fig. 304) . The palpal trochanter is tuberculated (Text-fig. 305). The femur and genu are provided with a comb-like and two spatu- late setae, respectively. The chelicera is as shown in Text-fig. 305. The sperma- dactyl is normal. The corniculus is distinctly stalked. Leg I is with tibia and tarsus 66-78(0, and H2-u6[ji long, respectively ; the prae-tarsus is IO[JL long. The diagnostic characters of leg II are shown in Plate 6e. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 237 The tibial spur differs from the genual process by being slightly broader. The trochanter of leg IV is delineated in Text-fig. 307. FEMALE. This sex is weakly sclerotized and light yellow in colour. The dorsal shield (507-522^1 long x 267(0, wide) is regionally reticulated and provided with about forty-six pairs of setae, of which twenty-one pairs of setae are situated on the anterior half of the dorsal shield (Text-fig. 308). The tritosternum is biramous. There are three prae-endopodal shields, the median shield being an elongated strip (Text-fig. 309). The interscutal membrane FIGS. 302-307. Pergamasus (Paragamasus) wasmanni (Oudemans), male. Fig. 302 dorsum of idiosoma. Fig. 303 venter. Fig. 304 tectum. Fig. 305 trochanter, femur and genu of pedipalp. Fig. 306 chelicera. Fig. 307 trochanter of leg IV. 238 S. K. BHATTACHARYYA 3IO 311 FIGS. 308-313. Pergamasus (Paragamasus) wasmanni (Oudemans), female. Fig. 308 dorsum of idiosoma. Fig. 309 venter. Fig. 310 endogynial process. Fig. 311 tectum. Fig. 312 chelicera. Fig. 313 trochanter of leg IV. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 239 between the anterior margin of the sternal shield and the prae-endopodal shields is without striations. The metasternal shields are almost fused in the mid-line. The endogynium has a pair of horns and its process is shown in Text-fig. 310. The genital shield is as in Plate 8e. The setation of the opisthogastral shield is figured. The stigma is situated between coxae III and IV. Anteriorly, the peritrematal shield extends to the level of coxa I, but the peritreme reaches to coxa II. The tectum is three-pointed, the median prong being short and narrow (Text-fig. 311). The pedipalp resembles that of the male but the trochanter lacks tubercles. The chelicera is shown in Text-fig. 312. The lateral margins of the hypostomal processes are fringed. The ventral groove of the gnathosoma is provided with ten transverse rows of deutosternal denticles. The tibia and tarsus of leg I are 70-74^ and I2i-i25(x long, respectively ; the prae-tarsus is 8-iojji long. The trochanter of leg IV is tuberculated (Text-fig. 313) . DISTRIBUTION AND HABITAT. Only the female of this species is previously known from Holland (Oudemans, 1902). I have examined material from litter and humus under pines and holly (near Duddleswell, Ashdown Forest, Sussex, K. H. Hyatt, io.ix.i956). Moss and leaves (Adelaide Hill, Bowness, Cumberland, J. T. Salmon, 29. vi. 1951). In leaf mould (Savernake Forest, Berkshire, J. T. Salmon, 26. ix. 1951). In moss (Grassington, Grass Woods, North Riding of Yorkshire, J. T. Salmon, 24^1.1951). Leaves from Sequoia sempervirens (Virginia Water, Surrey, A. H. G. Alston, i8.xii.i955). (Red Bank, Grasmere, Westmorland, M. E. Bacchus, 3 . xii . 1954) . Oak and rhododendron humus (Penn Ponds, Richmond Park, Surrey, P. N. Lawrence, io.iii.i957). Holly humus (near Groombridge, Sussex, P. N. Lawrence, 3.^.1957). Heath and birch (St. Faith's Common, Horsford, Norwich, Norfolk, J. T. Salmon, 13. v. 1951). In leaf mould (Burnham Beeches, Buckinghamshire, J. T. Salmon, 3.vi.i95i). Rhododendron humus (Swansea University, Swansea, P. N. Lawrence, 26.^.1962). RECORDS OF OTHER BRITISH SPECIES OF Pergamasus Berlese s. lot. According to " A synonymic Catalogue of British Acari " compiled by Turk (1953) the following species also occur in the British Isles. I have not been able to examine authentic British specimens of these species. 1. Amblygamasus dentipes (Koch, 1839). 2. Pergamasus alpestris var. coniger Hull, 1916. 3. Pergamasus processiferus Halbert, 1915. 4. Pergamasus johnstoni Oudemans, 1936. (nom. nov. pro Gamasus marginatus Johnston, 1848 non Gamasus marginatus Latr., 1806). 5. Paragamasus runcatellus (Berlese, 1906). The British records of this species probably refer to P. (Paragamasus} schweizeri sp. nov. 6. Paragamasus decipiens (Berlese, 1904). 7. Paragamasus minor (Berlese, 1892). 2 4 o S. K. BHATTACHARYYA 8. Plesiogamasus paripes (Hull, 1918). 9. Plesiogamasus parvulus (Berlese, 1903). 10. Plesiogamasus parvulus var. dilatatellus (Berlese, 1906). The records of Pergamasus barbarus Berl. and Pergamasus mediocris Berl. given by Turk & Turk (1952) are based on misidentifications ; the species involved being P. (P.) septentrionalis (Oudemans) and P. (P.) longicornis Berlese. SUMMARY This work deals with a revision of the British species of the genus Pergamasus Berlese s. lat. On the basis of the chaetotaxy of the dorsum of the idiosoma the genus is divided into two subgenera, Pergamasus s. str. and Paragamasus Hull. Thirty-two species are described, figured and keyed. The following new species are described : Pergamasus (Pergamasus) hortensis sp. nov. Pergamasus (Paragamasus) alstoni sp. nov cambriensis sp. nov. cassiteridum sp. nov. celticus sp. nov. femoratus sp. nov. integer sp. nov. londonensis sp. nov. longisetosus sp. nov. minimus sp. nov. nathistmus sp. nov. parrunciger sp. nov. rothamstedensis sp. nov. schweizeri sp. nov. The following species are recorded for the first time from Britain : P. (Pergamasus) mirabilis Willm., P. (Paragamasus) leruthi Cooreman, P. (Paragamasus) teutonictis Willm., P. (Paragamastis) truncus Schweizer and P. (Paragamasus) wasmanni (Oudemans) . ACKNOWLEDGEMENTS I am extremely grateful to Mr. T. E. Hughes for his interest in the work, to the Trustees of the British Museum (Natural History) for providing me with the facilities to work on the material of Pergamasus in the collections of the Arachnida Section, and to Dr. G. Owen Evans for his guidance and ready assistance at all times. My thanks are also due to Dr. J. Cooreman (Brussels) for the loan of type material, to Dr. L van der Hammen (Leiden) for material from the Oudemans Collection, and Prof. A. Melis (Florence) for permission to work on the Berlese Collection. Finally, I have pleasure in expressing my gratitude to the Central Research Funds Committee (University of London) for financial assistance. REVISION OF THE GENUS PERGAMASUS BERLESE S. LAT. 2 4 i REFERENCES BERLESE, A. 1888. Acari Austro-Americani. Bull. Soc. ent. Ital., 20, 171-222. - 1892. Acari Myriopoda et Scorpiones hucusque in Italia reperta. Portici et Padova. Order Mesostigmata (Gamasidae) : 1-143, figs. - 1904. Acari Nuovi. Redia, 1 : 235-252. - 1906. Monographia del genere Gamasus Latr. Redia. 3 : 65-304, figs. BHATTACHARYYA, S. K. 1962. Laboratory studies on the feeding habits and life cycles of soil inhabiting mites. Pedobiologia 1 : 291-298, fig. BONNET, A. 1911. Biospeologica. xxi. Description des Gamasides cavernicoles recoltes par A. Vire. Arch. zool. Paris (5) 8 : 381-398, figs. COOREMAN, J. 1943. Note sur la faune des Hautes-Fagnes en Belgique (i). xi Acariens (Parasitiformes) (2) Bull. Mus. roy. Hist. nat. Belg, 19 63 : 1-28, figs. - 1951. Etudes Biospeologiques. xxxiv (i) Acariens de Transylvanie. Bull. Mus. Hist, nat. Belg. 27 42 : 1-15, figs. EVANS, G. O. 1957. An introduction to the British Mesostigmata (Acarina) with keys to families and genera. /. Linn. Soc. Lond. Zool. 43 : 203-259, figs. 1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari : Mesostigmata). Bull. Brit. Mus. (nat. Hist.) Zool. 10 : 275-303. HALASKOVA, V. 1959. Zur Kenntnis der freilebenden Gamasiden der Tschechoslovakeia i. (Acarina, Parasitiformes). Acta Soc. ent. Cech. 56 : 97-108, figs. HALBERT, J. N. 1915. Clare Island Survey, Part 39, ii, Acarinida ii Terrestrial and marine Acarina. Proc. roy. Irish Acad., 31 : 45-136, figs. HULL, J. E. 1916. A November week at Grange-over-Sands in Acari (Terrestrial). Lane. Nat. 8 : 381-386. - 1918. Terrestrial Acari of the Tyne Province. Trans, nat. Hist. Soc. Northumb. (n.s.), 5 : 13-88. LEITNER, E. 1946. Zur Kenntnis der Milbenfauna auf Diingerstatten. Zbl. Gesamt. Geb. Ent. Lienz, 1, 3 : 75-95. - i946a. Zur Kenntnis der Milbenfauna auf Diingerstatten. Ibid. 1, 5-6 : 129-156, figs. OUDEMANS, A. C. 1902. New list of Dutch Acari. Second Part. With remarks on known and descriptions of a new subfamily, new genera and species. Tijdschr. Ent. 45, no. 1/2 : 1-52, figs. - 1904. Acarologische Aanteekeningen X. Ent. Ber. Amst. 1 (16) : 140-141. - 1912. Acarologische Aanteekeningen XXXIX. Ent. Ber. Amst., 3 (63) : 215-217. - 1926. Acarologische Aanteekeningen LXXXII. Ent. Ber. Amst., 7 (150) : 119-126, figs. - 1936. Kritisch Historisch Overzicht der Acarologie. Derde Gedeelte. Band A : 1-430, figs. Leiden. - 1939. Neue Funde auf dem Gebiete der Systematik und der Nomenklatur der Acari. III. Zool. Anz. 125, no. 1/2 : 20-24. PAX, F. & Willmann, C. 1937. Die Wasserfalle des Schneeberggaues und ihre Fauna. Pax, Beitr. Biol. Glatzer Schneeberges, 3 : 267-288. SCHMOLZER, K. 1953- Vorkommen und Verbreitung der Gattung Pergamasus Berl. 1903 in Oesterreich. Zool. Anz. 150 : 289-298, figs. SCHWEIZER, J. 1961. Die Landmilben der Schweiz. (Mittelland, Jura und Alpen). Parasiti- formes Reuter. Denks. schweiz. naturf. Ges. 84 : 1-207, figs. SELLNICK, M. 1929. Eine neue Pergamasus. Art. Bull. Inst. Rech. biol. Perm. 6, 7 : 319-322, figs, (paper in Russian, translation in German pp. 322-326). - 1940. Die Milbenfauna Islands. Goteborgs Vetensk. Samh. Handl. (5) 6B 14 : 1-129, figs. TRAGARDH, I. 1910. Acariden aus dem Sarekgebirge. Naturw. Untersuch. Sarekgeb., 4 : 375-586, figs. - 1936. Leptogamasus, a new genus of Acari from Sweden. Ent. Tidskr., 57 : 227-234, figs, TURK, F. A. 1953. A synonymic catalogue of British Acari. Ann. Mag. nat. Hist., (12). 6 : 1-26, 81-99. 242 S. K. BHATTACHARYYA TURK, F. A. & TURK, S. M. 1952. Studies of Acari yth Series. Records and descriptions of mites new to the British fauna together with short notes on the biology of sundry species. Ibid., (12), 5 : 475-506. WILLMANN, C. 1932. Acari aus sudostalpinen Hohlen. Mitt. Hohlen u. Karstf. Berlin, 4 : 158-161. 1938. Beitrag zur Kenntnis der Acarofauna des Komitates Bars. Ann. hist. nat. Mus. hung. 31 : 144-172, figs. 1939 Die Arthropodenfauna von Madeira .... XIV. Terrestrische Acari (exk. Ixodidae). Ark. Zool. Stockholm, 31A (10) : 1-42, figs. - I939a. Terrestrische Acari der Nord- und Ostseekuste. Abh. Nat. Ver. Bremen, 31 3 : 521-550, figs. 1940. Neue Milben aus Hohlen der Balkanhalbinsel gesammelt von Prof. Dr. K. Absolon, Brunn. Zool. Anz. Leipzig, 130 (9-10) : 209-218. - 1949. Uber eine Milbenausbeute aus dem Naturschutzgebiet ' Verlorenes Wasser ' bei Panten (Kr. Liegnitz). Abb. naturw. Ver. Bremen 32, 2 : 339-348, figs. 1951. Untersuchungen iiber die terrestrische Milbenfauna im pannonischen Klimagebiet Osterreichs. Sitz.-Ber. Osterr. Akas. Wiss. Abt. i. 160 : 91-176, figs. 1953. Neue Milben aus den Ostlichen Alpen. Ibid., abt. i, 162 : 449-519, figs. - 1954. Mahrische Acari hauptsachlich aus dem Gebiete des Mahrischen Karstes. Cesk. parasit. 1 : 213-272, figs. 1956. Milben aus dem Naturschutzgebiet auf dem Spieglitzer (Glatzer) Schneeberg. Cesk. parasit. 3 : 211-273, figs. VITZTHUM, G. H. 1926. Die Acarofauna der Harzfliisse (Der " Acarologischen Beobach- tungen " 12 Reihe) SitzBer. Ges. Naturf. Freunde : 89-110, figs. VOIGTS, H. & OUDEMANS, A. C. 1904. Zur Kenntnis der Milben-Fauna von Bremen Abh. Ver., Bremen 18 : 199-252, figs. PLATE i Armature of leg II in the male of Pergamasus s. lat. Pergamasus (Pergamasus) crassipes (L.) Berlese Pergamasus (Pergamasus) longicornis Berlese Pergamasus (Pergamasus) septentrionalis (Oudemans) Pergamasus (Pergamasus) quisquiliarum (Canestrini) Genu & tibia of Pergamasus (Pergamasiis) quisquiliarum (Canestrini) Bull. B.M. (N.H.) Zoo/, u, 2 PLATE i a PLATE 2 Armature of leg II in the male of Pergamasus s. lat. a. Coxa II of Pergamasus (Pergamasus) mirabilis Willmann b. Pevgamasus (Pergamasus) mirabilis Willmann c. Pergamasus (Pergamasus) hamatus (Koch) d. Pergamasus (Paragamasus) robustus (Oudemans) e. Coxa II of Pergamasus (Paragamasus) alpestris Berlese f. Pergamasus (Paragamasus) alpestris Berlese g. Pergamasus (Paragamasus) armatus Halbert Bull. B.M. (N.H.) Zool. n, 2 PLATE 2 PLATE 3 Armature of leg II in the male of Pergamasus s. lat. a. Pergamasus (Paragamasus) cambriensis sp. nov. b. Pergamasus (Paragamasus) diversus Halbert c. Trochanter II of Pergamasus (Paragamasus) cassiteridum sp. nov. d. Pergamasus (Paragamasus} cassiteridum sp. nov. e. Ventral view of femur of leg II in Pergamasus (Paragamasus) cassiteridum sp. nov. f. Ventral view of femur of leg II in Pergamasus (Paragamasus) robustus (Oudemans) g. Pergamasus (Paragamasus) celticus sp. nov. Bull. B.M. (N.H.) Zoo/, n, 2 PLATE 3 PLATE 4 Armature of leg II in the male of Pergamasus s. lat. Pergamasus (Paragamasus) femoratus sp. nov. Pergamasus (Paragamasus) integer sp. nov. Trochanter II of Pergamasus (Paragamasus} lapponicus Tragardh Pergamasus (Paragamasus} lapponicus Tragardh Pergamasus (Paragamasus} leruthi Cooreman Bull. B.M. (N.H.) Zool. n, 2 PLATE 4 PLATE 5 Armature of leg II in the male of Pergamasus s. lat. a. Pergamasus (Paragamasus) misellus Berlese b. Pergamasus (Paragamasus) nathistmus sp. nov. c. Pergamasus (Paragamasus) parrunciger sp. nov. d. Variation of tibial spur in Pergamasus (Paragamasus) parrunciger sp. nov. e. Ventral view of femur in Pergamasus (Paragamasus) parrunciger sp. nov. f. Pergamasus (Paragamasus) rothamstedensis sp. nov. g. Pergamasus (Paragamasus) runciger (Berlese) h. Ventral view of femur in Pergamasus (Paragamasus) runciger (Berlese) Bull. B.M. (N.H.) Zoo/, n, 2 PLATE 5 PLATE 6 Armature of leg II in male of Pergamasus s. lat. a. Pergamasus (Paragamasus) schweizeri sp. nov. b. Pergamasus (Paragamasus) suecicus (Tragardh) c. Pergamasus (Paragamasus) teutonicus Willmann d. Pergamasus (Paragamasus) truncus Schweizer e. Pergamasus (Paragamasus) wasmanni (Oudemans) Bull. B.M. (N.H.) Zool. n, 2 PLATE 6 PLATE 7 Genital shields of the females of Pergamasus s. lat. a. Pergamasus (Pergamasus) crassipes (L.) Berlese b. Pergamasus (Pergamasus) longicornis Berlese c. Pergamasus (Pergamasus) septentrionalis (Oudemans) d. Pergamasus (Pergamasus) quisquiliarum (Canestrini) e. Pergamasus (Paragamasus) robustus (Oudemans) f. Pergamasus (Paragamasus) alpestris Berlese g. Pergamasus (Paragamasus) alstoni sp. nov. h. Pergamasus (Paragamasus) diversus Halbert i. Pergamasus (Paragamasus) integer sp. nov. j. Pergamasus (Paragamasus) londonensis sp. nov. k. Pergamasus (Paragamasus) rothamstedensis sp. nov. 1. Pergamasus (Paragamasus) suecicus (Tragardh) Bull. B.M. (N.H.) Zoo/, n, 2 PLATE 7 PLATE 8 Genital shields of the females of Pergamasus s. lat. a. Pergamasus (Paragamasus) misellus Berlese b. Pergamasus (Paragamasus) truncus Schweizer c. Pergamasus (Paragamasus) armatus Halbert d. Pergamasus (Paragamasus) cambriensis sp. nov. e. Pergamasus (Paragamasus) wasmanni (Oudemans) f. Pergamasus (Paragamasus) celticus sp. nov. g. Pergamasus (Paragamasus) celticus sp. nov. (internal view) h. Pergamasus (Paragamasus) lapponicus Tragardh i. Pergamasus (Paragamasus) lapponicus Tragardh (internal view) j. Pergamasus (Paragamasus) parrunciger sp. nov. k. Pergamasus (Paragamasus) runciger (Berlese) 1. Pergamasus (Paragamasus) schweizeri sp. nov. m. Pergamasus (Paragamasus) teutonicus Willmann n. Pergamasus (Paragamasus) teutonicus Willmann (internal view) Bull. B.M. (N.H.) Zool. n, 2 PLATE 8 PRINTED IN GREAT BRITAIN BY THOMAS DE LA RUE & COMPANY LIMITED LONDON THE CHEILOSTOMATOUS POLYZOA NEOEUTHYRIS WOOSTERI (MacGILLIVRAY) AND REGINELLA DOLIARIS (MAPLESTONE) ANNA B. HASTINGS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. 3 LONDON: 1964 THE CHEILOSTOMATOUS POLYZOA NEOEUTHYRIS WOOSTERI (MacGILLIVRAY) AND REGINELLA DOLIARIS (MAPLE STONE) BY ANNA B. HASTINGS British Museum (Natural History) Pp. 243-262 ; Plates 1-3 ; 4 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ii No. 3 LONDON: 1964 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. n, No. 3 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. Trustees of the British Museum (Natural History) 1964 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued February, 1964 Price Eleven Shillings THE CHEILOSTOMATOUS POLYZOA NEOEUTHYRIS WOOSTERI (MacGILLIVRAY) AND REGINELLA DOLIARIS (MAPLE STONE) By ANNA B. HASTINGS CONTENTS Page 1. ABSTRACT .......... 245 2. NEOEUTHYRIS Bretnall ........ 245 a. Neoeuthyris woosteri (MacGillivray) ..... 245 3. REDUCED AND VESTIGIAL OVICELLS ...... 250 4. REGINELLA Jullien ......... 252 a. Reginella furcata (Hincks) ....... 253 b. Reginella doliaris (Maplestone) ...... 254 5. COMPARISON OF Reginella doliaris AND Conescharellina . . . 259 6. ACKNOWLEDGMENTS ......... 260 7. REFERENCES .......... 260 8. ADDENDA ........... 262 i. ABSTRACT Neoeuthyris woosteri (MacG.), unlike the other undoubted members of the Euthyrisellidae (in which ovicells are vestigial or absent), has well developed ovicells. Urceolipora, removed from the family by Harmer, has ovicells with some likeness to those of Neoeuthyris. There is evidence of a tendency to reduction of the ovicells of the Cheilostomata, but our knowledge of the factors concerned is still insufficient to frame an explanation. Metracolposa Canu & Bassler is a synonym of Reginella Jullien. Metracolposa mucronata Canu & Bassler falls within the range of variation of Reginella furcata (Hincks) of which it is thus a synonym. Cellepora doliaris Maplestone is a member of the Cribrilinidae. Its zooecia have much in common with those of Reginella, to which it is tentatively referred. The cribriform frontal shield of its erect zooecia faces the periphery of the low conical colony which is built with a profusion of kenozooecia and avicularia, apparently budded from the septula of the zooecia. Though belonging to a different major systematic group, the zoarium of R. doliaris shows parallel features to that of Conescharellina and helps to elucidate the arrangement of the zooecia in the Conescharellinidae. 2. NEOEUTHYRIS Bretnall Neoeuthyris Bretnall, 1921 : 157 ; Hastings, 1960 : 244, 245 ; Opinion 617, 1961 : 363. TYPE-SPECIES : Euthyris woosteri MacGillivray, by monotypy. The status of the generic name Neoeuthyris Bretnall, and of the name of its type- species, was established by Opinion 617 of the International Commission on Zoological Nomenclature. It remains to discuss the material of N. woosteri in the British Museum. 2a. Neoeuthyris woosteri (MacGillivray) Text-figs, i, 2 Euthyris woosteri MacGillivray, 1891 : 77, pi. 9, figs. 2, 2a. Neoeuthyris woosteri Bretnall, 1921 : 157, text-fig, i ; Hastings, 1960 : 244 ; Opinion 617, 1961 : 363. DISTRIBUTION : Cooktown, Queensland, on an alga (MacGillivray ; Bretnall) ; Western Australia, on Metamastophora plana (Gray ; 1938.8.10.1) ; Fremantle, 246 ANNA B. HASTINGS Western Australia, on part of one of four specimens of Metamastophora plana, Harvey's Australian Algae, No. 442 (1948.3.12.1, transferred from Department of Botany). HOLOTYPE : Cooktown, Queensland, divided between National Museum of Victoria, Melbourne, Victoria (MacGillivray's specimen), and Australian Museum, Sydney, N.S.W. (the rest of the specimen from which MacGillivray's lobe was taken, 11.875. Being part of the same specimen this, too, is holotype, not a paratype, c.f. Bretnall, 1921 : 159). REMARKS : As already noted (Hastings, 1960 : 245), 1938.8.10.1 is the type- material of Lichenella brentii Gray (1858), and the algal portion, there chosen as lectotype of Gray's species, is now in the Botanical Department of the British Museum (Nat. Hist.). No intact Polyzoa remain on this lectotype material, though some basal and lateral walls are to be seen. The part including the Polyzoan has been retained in the Zoological Department under the original number. The known colonies of N. woosteri all encrust algae and cause a wrinkling of the surface of the encrusted fronds. MacGillivray described the species from a single lobe from a colony whose form was unknown. Bretnall examined the whole, small specimen from which MacGillivray's lobe was taken (Australian Museum, 11.875), and established the algal nature of the basal layer. The basal surface, as described by MacGillivray, is like the basal algal layer in Gray's specimen. Miss Elizabeth Pope has very kindly examined the specimen in the Australian Museum, in which one fragment is mounted to show the alga, and she has confirmed the presence of ridges similar to those in Gray's specimen. No. 442 of Harvey's Australian Algae in the British Museum consists of four specimens of Metamastophora plana, one of which bears extensive growths of Neoeuthyris. The contrast between the ridged surface of the parts of the fronds bearing the polyzoan, or remains of it, and the smoothness of the colonized parts is striking. It is also interesting to find that a photograph of a specimen which, Mr. Ross tells me, is presumably part of the type-gathering of Metamastophora plana shows exactly similar ridges on the fronds (Foslie, 1929, pi. 25, fig. 5). The colonies of N. woosteri are very fragile, being delicately calcified and covered with a thin epitheca which readily breaks away. My material is all old (Gray's specimen has been in the Museum for over a century) and has been dried and preserved between paper as herbarium material. The specimens are therefore considerably damaged. Further, the epitheca may be obscured by a thin growth of a calcareous alga. Fortunately, enough remains intact to show the essential features of the anatomy and the beauty of the species. In fact the damage is some- times helpful, for zooecia are to be found in which the loss of the epitheca exposes the underlying calcareous parts, others in which the fracture of the calcareous wall ex- poses the floor of the compensation sac, and others again in which the compensation sac has also been destroyed exposing the interior of the zooecium. As in Euthyrisella obtecta (Hincks), the epitheca is stretched above the depressed, calcareous, frontal wall. It is attache