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A Ps i . ‘ a . ie yer af Je rt ’ ‘© on. | ‘ - - ? a | “ F fl a % na z oo | : “ | z | _ - sg F: te a ee a) sv g “Aro *vV 3, Sy 7 e @* s AUG 1974 THE BRITISH CYNIPOIDEA tg? (HYMENOPTERA) DESCRIBED BY P. CAMERON J. QUINLAN BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 1 LONDON : 1974 14 AUG1974 THE BRITISH CYNIPOIDEA (HYMENOPTERA) yy, ss DESCRIBED BY P. CAMERON BY JOHN QUINLAN Pp. I-21 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 1 LONDON : 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, ts issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 31 No. i of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) © Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 1 August, 1974 Price £1.15 THE BRITISH CYNIPOIDEA (HYMENOPTERA) DESCRIBED, BYP: CAMERON By J. QUINLAN CONTENTS SYNOPSIS 3 INTRODUCTION 3 ACKNOWLEDGEMENTS 6 CYNIPIDAE 6 Charipinae 6 Cynipinae 9 EUCOILIDAE Io FIGITIDAE F F : : : : ; : : : 16 Anacharitinae . ‘ : : ; : : ‘ : : 16 Aspicerinae sy Figitinae . , L7 SUMMARY OF THE PRESENT NAMES DISCUSSED 18 SUMMARY OF THE LECTOTYPES AND NEOTYPES DESIGNATED 18 REFERENCES 19 INDEx . 3 ; ; : ; ; : ; ; : . 20 SYNOPSIS The British species of Cynipoidea described by P. Cameron are re-classified. Five new combinations and seven new specific synonyms are established. Twelve lectotypes and five neotypes are designated. Nine specific names are at present nomina dubia. INTRODUCTION CAMERON described 50 British species of Cynipoidea during the period 1875-89. Most of these species have been variously referred to by European workers but some have remained unrecognized. Before further progress could be made with the preparation of keys to the British Cynipoidea in the Handbooks for the Identification of British Insects series it became necessary to study the type-material of Cameron’s species and to correlate it with the original descriptions. The type-specimens of 36 of these species have been located in the British Museum (Natural History) (hereafter abbreviated to BMNH); of the remaining 14 species, neotypes have been designated for five and nine remain nomina dubia. Three Cameron collections are to be found in the BMNH. One bears the registration label ‘1886-3’ and is a collection of galls and gallflies purchased from Cameron and collected mainly in Europe. The second collection bears the regi- stration label ‘Cameron 1896—76’ and was presented to the BMNH by F. D. Godman; the Museum Register states that this collection consisted of Tenthredinoidea and 4 J. QUINLAN Cynipoidea, “618 sps., 141 types, 391 microscope slides, 103 larvae and 242 drawings’. The third collection has the registration label ‘1914-110’. In preparing this paper I have contacted the following British and European Museums in order to ensure, as far as possible, that no type-material of Cameron’s British Cynipoidea has been overlooked: Leicester Museum; Royal Scottish Museum, Edinburgh; University Museum, Oxford; Természettudomanyi Mtzeum, Budapest. Most of these museums have kindly loaned me material that might have contained the Cameron material for which I was looking. During the course of this study I have traced where possible the collections of Thomas Richard Billups, the Rev. Thomas Anself Marshall and Eduard Arthur Fitch, who were contemporaries of Cameron and in a few instances the actual collectors of species described by him. Cameron’s British Cynipoidea are all mounted on rectangular cards. Many of them have been remounted by various people on similar rectangular cards, but in most such cases the original card mount has been left attached to the pin. Most type-material has been found in that collection bearing the accession label ‘Cameron 1896—76’, and in many instances the locality has been written on this label. Cameron’s original specific labels are sometimes attached to the specimens. Some of the rectangular cards on which Cameron mounted his material have a number in the bottom left-hand corner. In a few cases this number has been crossed out and another number has been written in the right-hand corner. The number in the left-hand corner can usually be correlated with the figure number on the plate in Cameron (1890); occasionally this number seems to be the paragraph number of the species description in Cameron’s monograph (1890). Only when the descrip- tion, the locality and other data such as Cameron’s original determination labels are present and agree with that published, has credence been given to this number. Cameron did not indicate which specimen of a series was to be regarded as the type. Unfortunately, prior to my examination of the Cameron material, the specimens had been moved in the collection from species to species and had not been given labels to indicate their original placements. There is usually no way of telling from Cameron’s descriptions how many specimens he had before him when describing a new species; only very rarely did he state the number of specimens upon which he based a description. Where only one type-specimen has been found and there is no evidence that more than one specimen ever existed, I have accepted and labelled that specimen as the holotype, provided that it agrees with the original description and bears the same data as that published. Cameron was not always the collector of the specimens he described and in those cases where another collector is cited in the descriptions this is indicated in the type-data. All nominal species are arranged alphabetically under the genus in which they were described and within their respective subfamilies, together with references to the original description and other relevant papers in which they have been referred to or discussed, in the following sequence. Name; author, date and page reference of the original publication; status and sex of primary type where known; present lectotype designation (if necessary); locality of primary type; type-depository. BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 5 Number and sex of paralectotypes if such exist, with data and depository information as for primary types. Statement on the labelling of the type-material and its condition if damaged. A statement (headed ‘Jdentity.’) on the generic placement and taxonomic validity of the name, accompanied when known by similar data to that above for the names of senior and junior synonyms. Notes on other aspects of the information supplied are given below. Lost types. In those cases where the type-material is lost and there is no evidence on the number of original specimens or their sex, the statement ‘Type(s) (? sex)’ is used to show this lack of information. Locality. Cameron did not always write the locality on his labels. In a number of instances one, two or three letters only were used to indicate the locality; I have given these localities in full. I have also added ‘GREAT Britain’ and the county, although this information was not published in the original descriptions. Collectors. Unless specifically stated to the contrary Cameron was the collector. In those cases where another collector was named and the material has not been found, it is presumed to have been returned to the collector and has had to be regarded as lost. Type-depositories. These are given for the primary types of the synonyms of Cameron’s names and for Cameron’s nominal species. The following abbreviations are used to indicate these depositories. BMNH _ British Museum (Natural History), London. IPK Institut fiir Pflanzenschutzforschung Kleinmachnow, Eberswalde. MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin. NM Naturhistorisches Museum, Vienna. NR Naturhistoriska Riksmuseum, Stockholm. UM University Museum, Oxford. UZI Universitetets Zoologiska Institution, Lund. ZSBS Zoologische Sammlung des Bayerischen Staates, Munich. Taxonomic arrangement. The family and subfamily arrangement is as given by Eady & Quinlan (1963: 7). The number of segments in the antennal club has been important in the identification of many of the species. It is not clear what Cameron meant by the term ‘club’; I have regarded as club segments those antennal segments that bear rhinaria, as defined by Richards (1956). Kleidotoma Westwood (1833 : 494). Cameron first used the name Kleditoma (x888a : 165) when describing the species nigripes. In subsequent papers, including his monograph (1890 : 216) in which he describes and keys species in the genus Kleidotoma Westwood, he consistently spells the generic name as Kleditoma and refers to Kleditoma Westwood (1833 : 494). In interpreting Cameron’s spelling of Kleidotoma it cannot be demonstrated that he intended to correct the original name and, therefore, the name Kleditoma is an ‘incorrect subsequent spelling’ as defined in Article 33(b) of the International Code of Zoological Nomenclature. Neotypes. The neotypes here designated accord with the provisions of Article 75(a) of the current International Code of Zoological Nomenclature, and this paper * 6 J. QUINLAN is considered to be revisionary work in the terms of the Code and of the proposed amendments to Article 75(a) (Bull. zool. Nom. 29(2) : 90, 1972). ACKNOWLEDGEMENTS I would like to thank those colleagues in other museums and institutions who have provided me with types on loan and information on the Cameron collection. For such assistance I warmly thank the following: Dr R. Danielson (Universitetets Zoologiska Institution, Lund); Dr Max Fischer (Naturhistorisches Museum, Vienna) ; Dr M. W. R. de V. Graham (University Museum, Oxford), Dr W. Hellén (Zoological Museum of the University, Helsinki); Dr E. Kénigsmann (Museum fiir Naturkunde der Humboldt-Universitat, Berlin); Dr F. Kiihlhorn (Zoologische Sammlung des Bayerischen Staates, Munich), Dr G. Morge (Institut fiir Pflanzenschutzforschung Kleinmachnow, Eberswalde). CYNIPIDAE CHARIPINAE Allotria ancylocera Cameron, 1886 :85. Holotype g, GREAT Britain: Scotland, West Lothian, Carruber Glen, 14. viii (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Carruber Glen’. On the underside of the label is the name ‘ancylocera’. A further rectangular card mount has the name ‘Carruber’ and ‘ancylocera’ together with the date ‘14/8’. A separate label with the word ‘Holotype’ printed in the centre has been added together with a determination label ‘HOLOTYPE Of Allotria ancylocera Cam. det. J. Quinlan. 1973’. Identity. Syn. n. of Alloxysta victrix (Westwood, 1833 : 495), holotype 9, GREAT BRITAIN (UM, Oxford) [examined]. Hellén (1963 : 16) first established the combination Alloxysta victvix (Westwood) and it is here confirmed. Allotria basimacula Cameron, 1886 :87. LECTOTYPE 9, Great Britain: Scotland, Stirlingshire, Mugdock (BMNH, ex coll. Cameron), here designated. Paralectotype. 1 ¢, same data as the lectotype (BMNH). The lectotype and the paralectotype have the BMNH accession label ‘Cameron 96-76 Mugdock’. Both specimens have the original rectangular card mount with the name ‘Mugdock’ and an unintelligible date written on the underside. The card mount attached to the lectotype has the name ‘basimacula’ written on it. The paralectotype has a separate label in Cameron’s handwriting ‘basimacula’. A further purple-edged circular label with the printed word ‘Lectotype’ in the centre has been added to the lectotype. A similar blue-edged paralectotype label has been added to the -paralectotype. Both specimens have a determination label ‘LECTOTYPE’ and ‘PARALECTOTYPE’ respectively ‘of Allotria basimacula Cam. det. J. Quinlan. 1973’. Identity. Valid species of Alloxysta Férster. Dalla Torre & Kieffer (1910 : 258) first established the combination of Alloxysta basimacula (Cameron) and it is herein confirmed. Allotria caledonica Cameron, 1886 : 88. Holotype 9, Great Britain: Scotland, Stirlingshire, Mugdock, 27. vii (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Mugdock’. On the underside of the label is the name ‘caledonica’. The rectangular card on which the holotype is mounted has the number ‘2’ in the left-hand corner; this is the figure number on the plate in Cameron (1890). On the underside of the card mount is the name ‘caledonica Cam.’ BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 7 and the date ‘27/7’ and the letters ‘Mugd’. A further label ‘caledonica’ in Cameron’s handwriting is attached to the holotype. A red-edged circular label with the word ‘Holotype’ printed in the centre has been added together with a determination label ‘HOLOTYPE of Allotria caledonica Cam. det. J. Quinlan 1973’. The holotype has only 8 segments of the right antenna and to of the left antenna remaining; all the tarsi are damaged and the wings are damaged by the glue used for mounting the specimen. Identity. Valid species of Alloxysta Forster, combination first established by Dalla Torre & Kieffer (1910 : 256) as Alloxysta caledonica (Cameron) and herein confirmed. Allotria collina Cameron, 1889 :57. Type(s) 2, GREAT Britain: Scotland, Stirlingshire, Mugdock (Cameron Coll.) (lost). Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer (1910 : 280) placed collina in Charips Marshall (1870 : 181), attributing the latter name to ‘Haliday in Marshall’ (loc. cit.) as does Ionescu (1969 : 237). It seems most probable from the species description that collina is assignable to Alloxysta Férster but in the absence of type-material the name must remain a nomen dubium. Allotria crassa Cameron, 1889:59. LECTOTYPE 9Q, Great Britain: Scotland, Sutherland, Bonar Bridge (BMNH, ex coll. Cameron), here designated. Paralectotypes. GREAT Britain: 1 Q, Scotland, Argyll, Cladich; 1 9, Scotland, Dumfries-shire, Dumfries (correctly associated) ; 1 9 (misassociated), Scotland, Dumfries-shire, Dumfries. The lectotype and the paralectotypes have the BMNH accession label ‘Cameron 96-76’ together with their respective localities as listed above. The lectotype has the original rectangular card mount on the underside of which is the locality ‘Bonar’. A further handwritten label ‘crassa Cam’ is attached, on the underside of which is the name ‘scipes Thom’. This is deduced as being originally ‘fuscipes’ Thom’ before being used to write the name ‘crassa Cam’ on the other side. A purple-edged circular label with the printed word ‘Lectotype’ has been added together with a determination label ‘LECTOTYPE of Allotria crassa Cam. det. J. Quinlan. 1973’. The two correctly associated paralectotypes both have blue-edged circular paralectotype labels and determination labels ‘Allotria crassa Cam. det. J. Quinlan 1973’. The misassociated paralectotype has a blue-edged circular paralectotype label and a label in Cameron’s handwriting ‘Allotria fuscipes’. A further label ‘PARALECTOTYPE of Allotria crassa Cam. wrongly associated det. J. Quinlan 1973’ is attached. This wrongly associated paralectotype is referable to the genus Phaenoglyphis Forster. Identity. Valid species of Alloxysta Forster, first established by Dalla Torre & Kieffer (1910 : 261) and herein confirmed. Allotria dolichocera Cameron, 1889 :56. LECTOTYPE 9, Great Britain: Scotland, Dumfries-shire, Dumfries (BMNH, ex coll. Cameron). Paralectotypes. GREAT BRITAIN: I @ (misassociated), Scotland, Lanarkshire, Cadder Wilderness (BMNH); 1 Q (misassociated), England, London, Peckham (T. R. Billups) (BMNH). The lectotype herein designated and the paralectotypes have the BMNH accession label ‘Cameron 96-76’ together with their respective localities as listed above. The lectotype has a label in Cameron’s handwriting ‘dolichocera’, on the other side of this label is the name ‘brevis Thoms’ but the top of this name has been cut off. A further rectangular card on which the lectotype was originally mounted has on the underside the locality ‘Dumfries’ and the word ‘brevis’. The two misassociated paralectotypes clearly belong in the genus Alloxysta Forster and do not agree with Cameron’s descripton of dolichocera on the relative lengths of the antennal segments. The paralectotype collected by Billups has both antennae damaged but the critical segments are still present. A purple-edged circular label with the word ‘Lectotype’ has been added, together with a determination label ‘LECTOTYPE of Allotria dolichocera Cam. det. J. Quinlan 1973’. Identity. Valid species of Phaenoglyphis Forster, new combination Phaenoglyphis dolichocera (Cameron) comb. n. here established. 8 J. QUINLAN Allotria maculicollis Cameron, 1886: 87. Type(s) 3, GREAT Britain: Scotland, Kirkcud- brightshire, New Galloway (Cameron Coll.) (lost). Identiy. Unknown, the name remains a nomen dubium. The identity cannot be deduced from Cameron’s (1890 : 252) key placement, in which Allotria macrophadna is compared with maculicollis. The name was applied almost certainly to an Allovysta species. Allotria megaptera Cameron, 1889:54. Holotype ¢, Great Britatn: Scotland, Lanarkshire, Cadder Wilderness (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Cadder Wilderness’. On the underside of this label is the name ‘megaptera’. Two further labels in Cameron’s handwriting are attached to the holotype pin: they read ‘Allotria’ and ‘megaptera Cam’. On the original rectangular card is the number ‘7’; this is the figure number on the plate in Cameron (1890). A further red-edged circular label with the word ‘Holotype’ printed in the centre has been attached together with a determination label ‘HOLOTYPE of Allotria megapterva Cam. det. J. Quinlan 1973’. The left fore wing and both hind wings are missing. Identity. Valid species of Alloxysta Forster, new combination Alloxysta megaptera (Cameron) comb. n. here established. Cameron first listed megaptera in a key to the British Allotria without designating it ‘sp. nov.’ as was usually his custom. The description in the couplet of the key to Allotria satisfies the provisions of Article 11 of the International Code of Zoological Nomeclature. Allotria melanogaster Hartig (1840 : 200), misidentified by Cameron (1886 : 86), was later described as Allotria megaptera Cameron (1889 : 54). Allotria mullensis Cameron, 1883 : 366. Holotype 9, Great Britain: Scotland, Argyll, Mull (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Mull’. On the underside of this label is the name ‘mullensis’. The original card mount has the number ‘1’ in the left- hand corner; this is the figure number on the plate in Cameron (1890). A further: label ‘mullensis Cam’. in Cameron’s handwriting is attached. A red-edged circular label with the word ‘Holotype’ printed in the centre has been added together with a determination label ‘HOLOTYPE of Allotria mullensis Cam. det. J. Quinlan 1973.’ Identity. Valid species of Alloxysta Forster, new combination Alloxysta mullensis (Cameron) comb. n. here established. Allotria perplexa Cameron, 1889 : 58. Type(s) 9, GREAT Britain: Scotland, Sutherland; Inverness-shire, Kingussie; Lanarkshire, Clydesdale; Kirkcudbrightshire, New Galloway; Dumfries-shire, Dumfries (Cameron Coll.) (lost). Identity. Unknown, the name remains a nomen dubium. Hellén (1963 : 12) placed perplexa in the genus Alloxysta Forster, but without seeing type-material. In the absence of type-material the name must remain a nomen dubium. Allotria piceomaculata Cameron, 1883 : 367. Holotype 9, GREAT Britain: Scotland, Dumfries-shire, Dumfries, vi (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Dumfries’, on the underside of this label is the name ‘piceomaculata’. The rectangular card on which the holotype was originally mounted has the number ‘3’; this is the figure number on the plate in Cameron (1890). On the underside of the card mount is the locality ‘Dumfries’ and the name ‘piceomaculata’. The rectangular card mount on which the holotype is mounted also has the number ‘3’ in the left-hand corner. A further label in Cameron’s handwriting ‘piceomaculata Cameron’ is attached to the holotype. A red-edged circular label with the word ‘Holotype’ printed in the centre has been added together with a determination label ‘HOLOTYPE of Allotria piceomaculata Cam. det. J. Quinlan 1973’. The holotype has only one segment remaining of the left antenna. Identity. Valid species of Alloxysta Forster, combination first established by Cameron (1886 : 88) and here confirmed. Allotria pleuralis Cameron, 1879 :113. LECTOTYPE 9, Great Britain: Scotland, Lanarkshire (BMNH, ex coll. Cameron), here designated. Paralectotypes. GREAT BRITAIN: 2 Q, Scotland, Lanarkshire (BMNH, ex coll. Cameron). The lectotype has the BMNH accession label ‘Cameron 96-76’. Both the rectangular BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 9 card on which it is mounted and the original card mount have the number ‘8’ in the bottom left-hand corner. This number is the reference to the figure number on the plate in Cameron (1890). A further purple-edged circular lectotype label has been added together with a determination label ‘LECTOTYPE of Allotria pleuralis Cam. det. J. Quinlan 1973’. The left antenna has the apical segment missing. The two paralectotypes are labelled ‘Cameron 96-76’ and carry the label ‘pleuralis’. Two further specimens have not been located. (Five specimens were referred to in the original description, three from Clyde near Newton and two from Possil Marsh, all taken in July. One paralectotype has the locality Possil Marsh.) Identity. Syn. n. of Xystus testacea Hartig, 1841 : 352, holotype 9, Austria (ZSBS, Munich) and new combination Alloxysta testacea (Hartig) comb. n. here established. Allotria ruficeps Cameron, 1883 : 365. Type(s) 9, GREAT BritaIn: Scotland, Kirkcud- brightshire, New Galloway, vi (Cameron Coll.) (lost). Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer (1910 : 283) proposed a replacement name Charips (Charips) cameroni for Allotria ruficeps Cameron, which was preoccupied by Allotria ruficeps (Zetterstedt, 1883 : 410). Allotria ruficollis Cameron, 1883 : 385. Holotype 9, GREAT Britain: Scotland, Argyll, Mull, vi (BMNH, ex coll. Cameron). : The holotype has the BMNH accession label ‘Cameron 96-76 Mull’. The original rectangular card mount has on the underside the words ‘Mull ruficollis’. A further hand- written label in Cameron’s handwriting ‘ruficollis Cam’. is attached. A red-edged circular label with the word ‘Holotype’ printed in the centre has been added together with a determination label ‘HOLOTYPE of Allotria ruficollis Cam. det. J. Quinlan 1973’. Identity. Syn. n. of Xystus erythrothorvax Hartig, 1840 : 200, holotype g, GERMANY (ZSBS, Munich). Dalla Torre & Kieffer (1910 : 257) first established the combination Alloxysta erythrothorax (Hartig) and it is here confirmed. Allotria salicis Cameron, 1883 : 367. Type(s)? 9, GREAT Britain: Scotland, Lanarkshire, Clydesdale, Kilpatrick Hills (Rev. T. A. Marshall) (lost). Identity. This nominal species clearly belongs in the genus Phaenoglyphis Forster. Cameron (1890 : 237) first established salicis as being in Phaenoglyphis but in the absence of the type(s) or reliably determined material the specific name remains a nomen dubium. Phaenoglyphis forticornis Cameron, 1888): 210. Type(s) 9, GREAT Britain: England, Barnstaple, Lastingham (Rev. T. A. Marshall) (lost). Identity. Unknown, the name remains a nomen dubium. Dalla Torre & Kieffer (1910 : 294) and Ionescu (1969 : 274) placed forticornis in Phaenoglyphis Forster but without seeing type-material; the name must remain a nomen dubium. CYNIPINAE Aulax graminis Cameron, 1875 : 322. LECTOTYPE 9, Great Britain: Scotland, Lanarkshire, Glasgow, near Partick (BMNH, ex coll. Cameron), here designated. Paralectotypes. 3 9, same data as the lectotype (BMNH). The lectotype and the paralectotypes have the BMNH accession label ‘Cameron 96-76’. The lectotype has a label in Cameron’s handwriting ‘graminis Cam’. A further purple-edged circular label with the printed word ‘Lectotype’ has been added, together with a further determination label ‘LEcToTyPE of Aulax graminis Cam. det. J. Quinlan 1973’. A blue- edged circular label with the printed word ‘Paralectotype’ has been added to each paralectotype. Identity. Junior synonym of Aulacidea hieracii (Bouché, 1834: 164), holotype g, GERMANY (DEI, Eberswalde). The synonymy of graminis with hieracii was first established by Cameron (1893 : 50) and recently confirmed by Eady & Quinlan (1963 : 20), who inadvertently marked the synonymy as new. Diastrophus (?) aphidivorus Cameron, 1889 :67. Holotype gj, GREAT BRITAIN: England, ?Barnstaple (Rev. T. A. Marshall) (BMNH, ex coll. Cameron). 10 J. QUINLAN The holotype has the BMNH accession label ‘Cameron 96-76’. The original retangular card mount has on the underside the words ‘Aphis of Nettle N’. On the upperside of the card mount can be seen the lighter areas where the holotype and the aphis referred to in Cameron’s description had been mounted. The number ‘7’, badly faded, can be seen that refers to the plate figure reference in Cameron (1893). A further label ‘aphidivorus’ in Cameron’s handwriting is attached to the pin. A red-edged circular label with the printed word ‘Holotype’ in the centre has been added together with the determination label ‘HOLOTYPE of Diastrophus aphidivorus Cam. det. J. Quinlan 1973’. The holotype has the left hind tibia and tarsus missing. The biological information in Marshall’s handwriting ‘Aphis of Nettle N’ is suspect. The ‘N’ on the label gives rise to doubt on the actual type-locality published by Cameron as ‘Barnstaple’. Morley (1915 : 23), who lists the Rev. T. A Marshall’s localities, gives the letter ‘N’ as Marshall’s abbreviation for Nunton, near Salisbury (Wiltshire). Identity. Syn. n. of Diastrophus rubi (Bouché, 1834 : 163), holotype 2, GERMANY (MNHU, Berlin) [examined]. Dalla Torre & Kieffer (1910 : 49) listed aphidivorus as a species of Trischiza Forster (Figitinae). EUCOILIDAE Eucoila fortinervis Cameron, 1889 : 66. Holotype g, GREAT BRITAIN: England, Gloucester (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Gloster’. The original rectangular card on which it was mounted has the number ‘g’ on the upper surface and the locality ‘Gloster’ on the underside. The number ‘9’ is the reference to the figure number on the plate in Cameron (1890). The holotype has the label ‘fortinervis Cam’ in Cameron’s handwriting attached to it. Although Cameron did not describe the female of fortinervis, pl. 9, fig. 9 in Cameron (1890) of a female suggests that the figure was transposed with that of another species, most probably proxima. A red-edged circular. label with the word ‘Holotype’ printed in the centre has been attached, together with a determination label ‘HOLOTYPE of Eucoila fortinervis Cam. det. J. Quinlan 1973.’ The holotype has segments 14-15 of the right-hand antenna missing. Identity. Valid species of Trybliographa Forster, new combination Trybliographa fortinervis (Cameron) comb. n. here established. Eucoila gracilicornis Cameron, 18884 : 168. LECTOTYPE 9, Great Britain: Scotland, Lanarkshire, Banks of Clyde nr Cambuslang. Paralectotypes. 2 3, same data as the lectotype (BMNH, ex coll. Cameron). The lectotype herein designated has the BMNH accession label ‘Cameron 96-76 S. BK. Clyde’. A rectangular label with ‘gracilicornis CBL’ and the name ‘similis’ crossed out is attached. On the underside of this label is the number ‘4’; this number seems to be the paragraph number in Cameron (1890 : 199), in which the species is described. Another label added at a later stage by G. J. Kerrich has the abbreviation ‘Pseudeuc’ on it. A circular purple-edged label with the printed word ‘Lectotype’ has been added together with a determination label ‘LEctotyPE of Eucoila gracilicornis Cam. det. J. Quinlan 1973’. Both paralectotypes have had blue-edged circular paralectotype labels added to them, together with determination labels. Identity. This valid species was first placed in Trybliographa Forster by Hellén (1960 : 12). The placing of gracilicornis in Trybliogrvapha by Hellén is herein confirmed. Eucoila proxima Cameron, 1889 :67. LECTOTYPE 9, Great Britain: England, Essex, Benfleet (BMNH, ex coll. Cameron), here designated. Paralectotypes. 29, same data as the lectotype except that Benfleet is omitted (BMNH, ex coll. Cameron). The lectotype has the BMNH accesssion label ‘Cameron 96-76 Benfleet’. A further BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON II purple-edged circular label with the printed word ‘Lectotype’ in the centre has been added together with a determination label ‘LEcToTyPE of Eucoila proxima Cam. det. J. Quinlan 1973’. One paralectotype has the additional label ‘proxima Cam’ in Cameron’s handwriting and the number ‘16/83’, together with the number ‘14’ in one corner and the number ‘8’ in the other. The number ‘8’ is the number of the figure on the plate in Cameron (1890). This plate and figure shows a male; Cameron did not describe the male of proxima and this suggests that the figure was transposed with that of fortinervis Cameron. The other paralectotype has a label ‘proxima Cam’. in Cameron’s handwriting and the number ‘15’ on the original card mount. Identity. Syn. n. of Trybliographa glottiana (Cameron 1883 : 368), holotype 2, GREAT Britain (BMNH, ex coll. Cameron) [examined, see p. 14]. Eucoila scotica Cameron, 1889 :65. LECTOTYPE 9, Great Britain: Scotland, Dumfries- shire, Dumfries (BMNH, ex coll. Cameron), here designated. Paralectotypes. GREAT BriTAIn: 1 9, Scotland, Stirlingshire, Mugdock (BMNH, ex coll. Cameron); 1 g, 1 9, Scotland, Dumfries-shire, Dumfries (BMNH, ex coll. Cameron); 1 9, Scotland, Dumfries-shire, Dumfries (BMNH, ex coll. Cameron); 1 9, Scotland, Kirkcud- brightshire, Colvend (BMNH, ex coll. Cameron); 1 9, Scotland, no locality; 1 g, Scotland, West Lothian, Carruber Glen (BMNH, ex coll. Cameron); 2 9, Scotland, Ayrshire, Dalry (BMNH, ex coll. Cameron). The lectotype has the BMNH accession label ‘Cameron 96-76. Dumfries’, together with another rectangular label with the locality and name ‘Dumfries scotica’. A further label in Cameron’s handwriting is attached and reads ‘scotica Cam’. On the reverse side of this label is the abbreviation ‘clia (new sp. ?) 9’. A purple-edged circular label with the printed word ‘Lectotype’ has been added, together with a determination label ‘LECTOTYPE of Eucoila scotica Cam. det. J. Quinlan 1973’. The paralectotypes all have the BMNH accession label ‘Cameron 96—76’ together with their respective locality names and the deter- mination label ‘scotica’. The paralectotype from Colvend has the name labels ‘ciliaris’ and ‘scotica’; on the underside of this label is the number ‘3’, which is the figure number on the plate in Cameron (1890). Identity. Valid species of Trybliographa Forster, new combination Trybliographa scotica (Cameron) comb. n. here established. Kleditoma affinis Cameron, 1889 : 64. Holotype 9, GREAT Britarn: Scotland, Sutherland, Bonar Bridge (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Bonar Bridge’. A further label in Cameron’s handwriting is attached and reads ‘affinis Bonar’. A circular red-edged label with the printed word ‘Holotype’ has been added, together with a determination label ‘HOLOTYPE of Kleidotoma affinis Cam. det. J. Quinlan. 1973.’. Identity. This valid nominal species is clearly assignable to Kleidotoma subgen. Tetrahoptra Forster (1869 : 342), senior synonym of Tetratoma Cameron (1890 : 223). Kleditoma caledonica Cameron, 1888a : 167. Type(s) 9, GREAT Britain: Scotland, Argyll, Claddich, Loch Awe (Cameron) (lost). NEOTYPE Q, Great Britain: England, Buckinghamshire, Greenlands, 31.vii.1936, ‘reared from nest of Heron, No. 2.’ (BMNH), here designated. I have been unable to locate the original female type(s). The neotype here designated has, in addition to the data above, a red-edged circular label with the printed word ‘Neotype’ in the centre, an identification label ‘Kleidotoma caledonica Cam. J. F. P. 1947.’, and the label ‘NEOTYPE of Kleidotoma caledonica Cam. det. J. Quinlan 1973.’. Identity. This valid species is clearly assignable to Kleidotoma subgen. Kleidotoma Westwood. Dalla Torre & Kieffer (1910 : 210), Hellén (1960 : 24) and Ionescu (1969 : 139) have referred to this species and have used the characters that Cameron used in his original description for identifying it. The BMNH collection contains 2 9, identified and labelled as caledonica by J. F. Perkins, which agree with the original description; one of these has been designated neotype. Cameron’s description isolates caledonica from other related species on the antennal club being shorter than the flagellum, the scutellum not projecting at the 12 J. QUINLAN apex, the legs being black, the radial cell elongate and the joints of the antennal club being attenuate. Cameron did not know the male of the species. Dalla Torre & Kieffer (1910 : 211) and Ionescu (1969 : 139) refer in error to both male and female in Cameron’s works (1888a : 166; 1890 : 224). Kleditoma crassiclava Cameron, 1888a:166. Holotype 9, GREAT Britain: Scotland, Sutherland, Bonar Bridge (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘P. Cameron Coll. B.M. 1914-110’. The original rectangular card mount has the name ‘crassiclava’ on the underside; the same card mount has the locality ‘Bonar’ written across the specific name. Both the species name and the locality are written in Cameron’s handwriting. The holotype has been remounted on a card point. A red-edged circular label with the printed word ‘Holotype’ in the centre has been attached, together with a determination label ‘HoLoTYPE of Kleidotoma crassiclava Cam. det. J. Quinlan. 1973.’. Identity. This valid species is assignable to the genus Rhynchacis Forster. The combination with Rhynchacis Forster was first established by Cameron (1890 : 217). Kleditoma elegans Cameron, 1889 :60. LECTOTYPE 9, Great Britain: Scotland, Stirlingshire, Mugdock Wood, nr Glasgow (BMNH, ex coll. Cameron), here designated. Paralectotype. GREAT BRITAIN: I @ (misassociated), Scotland, Lanarkshire, Clober Glen, nr Glasgow (BMNH, ex coll. Cameron). The lectotype has the BMNH accession label ‘96-76 Mugdock Wood’. The specific name ‘elegans Cam.’ in Cameron’s handwriting is attached to the lectotype. The originai card mount has the number ‘g’ in the bottom right-hand corner; this is the figure number to the plate in Cameron (1890). The misassociated paralectotype has a specific label ‘elegans?’ in Cameron’s handwriting; although similar to elegans, it has the antennal segments differently shaped and of different comparative lengths. It is Kleidotoma pentatoma Thomson. Identity. This valid species is assignable to Kleidotoma subgen. Pentakleidota Weld and is the type-species of this subgenus. Kleditoma filicornis Cameron, 1889 : 62. Type(s) 9, Great Britain: England, Devon, Bishopsteignton (Rev. T. A. Marshall) (lost). NEOTYPE Q, Great Britain: England, Norfolk, Holkham, 17.vii.1968 (M. Crisp) (BMNH), here designated. I have been unable to locate the original female type(s). Various authors, Dalla Torre & Kieffer (1910), Hellén (1960) and Ionescu (1969) have all referred to this species in published works and have used the characters that Cameron used in his original description for identifying this species. The neotype here designated is one of 8 females with identical data; a red-edged circular label with the printed word ‘Neotype’ in the centre has been attached, together with a further label ‘NEOTYPE of Kleidotoma filicornis Cam. det. J. Quinlan 1973.’. Identity. This valid species is assignable to Kleidotoma subgen. Kleidotoma Westwood and is separated from related species on the long thin antennae, the less abrupt club and the long narrow radius of the forewing. Dalla Torre & Kieffer (1910) wrongly cite Cameron (1890) as describing the male. Kleditoma gracilicornis Cameron, 1889 : 63. Type(s) 2, GREAT BRITAIN: England, Wiltshire, Nunton (publ. as Munton] (Rev. T. A. Marshall) (lost). Identity. This nominal species clearly belongs in the genus Kleidotoma subgen. Tetvahoptra Forster, but in the absence of the type(s) or reliably determined material the specific name remains a nomen dubium. The male of this species was not described. Kleditoma longicornis Cameron, 1899 : 62. Type(s) 9, GREAT Britain: England, Devon, Barnstaple (Rev. T. A. Marshall) (lost). Identity. This nominal species belongs in the genus Kleidotoma subgen. Kleidotoma Westwood, but in the absence of the type(s) or reliably determined material the specific name remains a nomen dubium. BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 13 Kleditoma longipennis Cameron, 1889:59. Holotype 9, Great Britarn: Scotland, Lanarkshire, Clober Moor, nr Glasgow (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Clober Moor’, on the reverse side of which is the name ‘longipennis’. A further label in Cameron’s handwriting ‘longipennis Cam.’ and a smaller label with the name ‘longipennis’ is attached. The original card on which the holotype was mounted has the data ‘Clober Moor’. The rectangular card on which the holotype is mounted has the number ‘8’ in the bottom left-hand corner with a line crossing the number out; in the right-hand corner of the card mount is the number ‘9’. The latter number is the figure number on the plate in Cameron (1890). Identity. This valid species belongs in the genus Kleidotoma subgen. Pentakleidota Weld. The male of this species was not described. Kleditoma marshalli Cameron, 1889 : 61. Type(s) 2 g, GREAT Britain: England, Devon, Barnstaple (Rev. T. A. Marshall) (lost). Identity. This nominal species clearly belongs in the genus Kleidotoma_ subgen. Kleidotoma Westwood, but in the absence of the type(s) or reliably determined material the specific name remains a nomen dubium. Kleditoma melanopoda Cameron, 1888a:167. Holotype 9, Great Britain: England, London District (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76’; no locality data are attached. The holotype is mounted on a rectangular card in the bottom left-hand corner of which is the number ‘1o’; this number is the figure number on the plate in Cameron (1890). A label ‘melanopoda’ in Cameron’s handwriting is attached. A red-edged circular label with the word ‘Holotype’ printed in the centre has been attached, together with a determination label ‘HOLOTYPE of Kleidotoma melanopoda Cam. det. J. Quinlan. 1973.’. (Although not stated in the original description, Cameron (1890 : 231) indicates that the Rev. T. A. Marshall was the collector of this specimen from the same locality ‘London District’.) Identity. This valid species belongs in the genus Kleidotoma subgen. Arhoptra Kieffer. The male of this species was not described. Kleditoma nigripes Cameron, 18884 : 165. Type(s) 2, GREAT Britain: England, London, Dulwich (T. R. Billups) (lost). NEOTYPE 9, Great Britain: England, Hertfordshire, Harpenden, Rothamsted, 1951 (B. R. Laurence), B.M. 1951-450 (BMNH), here designated. No Cameron material of this species stands in the BMNH collection under this name and all attempts to trace the type(s) in collections that could have contained T. R. Billups material have been fruitless. The neotype here designated is one of four females with the above data in the BMNH collection determined as ‘Rhynhacis nigripes (Cam)’ by G. J. Kerrich, 1951. It has a red-edged circular label with the word ‘Neotype’ printed in the centre, together with a determination label ‘NEOTYPE of Kleidotoma nigripes Cam. det. J. Quinlan. 1973.’ Identity. Valid nominal species of Rhynchacis Forster, combination Rhynchacis nigripes (Cameron) first established by Cameron (1890 : 218) and here confirmed. Kleditoma picipes Cameron, 1886: 92. Holotype 9, Great Britain: Scotland (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76’. No data label for the locality is attached. A label in Cameron’s handwriting is attached and reads ‘picipes’. The holotype is mounted on a rectangular card; in the bottom left-hand corner is the number ‘Io’, which is the figure number on the plate in Cameron (1890). The holotype bears a red-edged circular label with the printed word ‘Holotype’ in the centre, together with a determination label ‘HoLotyPE of Kleidotoma picipes Cam. det. J. Quinlan. 1973.’. Cameron did not give a locality in his original description, although the paper was on the fauna of Scotland with particular reference to Clydesdale. In a later reference to the species, Cameron (1890 : 226) gave Clydesdale as the locality for the species. Identity. This valid species of Kleidotoma Westwood is newly assigned to the subgen. 14 J. QUINLAN Kleidotomidea Rohwer & Fagan (1917 : 369). In placing picipes I have by examination counted as club segments of the antennae those segments having rhinaria. The male of picipes was not described by Cameron. Kleditoma striata Cameron, 1886 : 91. Holotype 3, GREAT Britain: Scotland, Lanarkshire, Lanark, nr Newton on the Clyde (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76. S. Bk. Clyde’. Two determi- nation labels in Cameron’s handwriting are attached, one has the name ‘striata’ and the other has the name ‘Kleditoma striata Cam 3’. The number ‘3’ is repeated again on the rectangular card mount and is the figure number on the plate in Cameron (1890). The holotype bears a red-edged circular label with the printed word ‘Holotype’ in the centre. A further determination label ‘HOLOTYPE of Kleidotoma striata Cam. det. J. Quinlan. 1973.’. has been attached. Identity. This valid species belongs in Kleidotoma subgen. Pentakleidota Weld. Cameron did not describe the male of this species although one male from his collection labelled as striata is in the BMNH collection. Kleditoma striaticollis Cameron, 1888a:167. Holotype 9, Great Britain: Scotland, Kirkcudbrightshire, New Galloway (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Galloway’ together with a further label with the locality ‘Galloway’ and the name ‘striaticollis’ in Cameron’s hand- writing. The holotype is mounted on a rectangular card; in the bottom left-hand corner is the number ‘8’. This number is the paragraph number given to the species in Cameron (1890 : 222). No figure of the species was given by Cameron. A red-edged circular label with the printed word ‘Holotype’ in the centre has been attached to the holotype together with a determination label ‘HoLoTyPE of Kleidotoma striaticollis Cam. det. J. Quinlan. 1973.’ Identity. This valid species belongs in the genus Kleidotoma subgen. Kleidotoma Westwood. The male of stviaticollis was not described by Cameron. Kleditoma truncata Cameron, 1889 : 60. Holotype 9, GREAT Britain: Scotland, Renfrew- shire, Bishopton, 31. vii (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Bishopton’. A further label with the locality ‘Bishopton 31/7’ is attached to the holotype together with a label ‘truncata Cam.’ in Cameron’s handwriting and another label with ‘truncata 2’ is also attached to the holotype. The number ‘2’ on the second name label is the number of the figure on the plate in Cameron (1890). A red-edged circular label with the printed word ‘Holotype’ in the centre has been added, together with a determination label ‘HoLoTYPE of Kleidotoma truncata Cam. det. J. Quinlan. 1973.’. Identity. This valid species of Kleidotoma Westwood is newly assigned to the subgen. Pentakleidota Weld (1952 : 225). Psichacra dalei Cameron 1879: 115. Holotype 9, Great Britain: England, Dorset, Wooton (J. C . Dale) (lost). NEOTYPE 9, Great Britain: England, Monmouth, Monks Wood, 5.x.1878 (BMNH), here designated. I have been unable to locate the original female holotype of this species. Cameron (1890 : 198) gives an additional locality to that in his first description: Worcester (Fletcher). The neotype here designated, in addition to the data above, has the labels ‘Cameron 96-76 Monks Wd.’ and a handwritten label ‘rufula Forst’. The neotype is one of three specimens with identical data in the BMNH and is distinguished from them by a red-edged circular label with the printed word ‘Neotype’ in the centre. A further label ‘NEOTYPE of Psichacva dalei Cam. det. J. Quinlan 1973.’ has been added. Identity. Junior synonym of Psichacra rufula (Forster, 1855 : 257), synonymy first established by Cameron (1890 : 197) and here confirmed. Psichacra glottiana Cameron, 1883 : 368. Holotype 9, Great Britarn: Scotland, Lanark shire, Cambuslang (publ. as Cambusland), along the banks of the Clyde, vii (BMNH, ex coll. Cameron). BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 15 The holotype has the BMNH accession label ‘Cameron 96-76 South Bank of the Clyde’. A further rectangular card on which the holotype was originally mounted has the number ‘7’ in the right-hand corner; this is the number of the figure on the plate in Cameron (1890). On the underside of this label are the initials ‘CBL’ and the specific name ‘glottiana’ in Cameron’s handwriting. A further label in Cameron’s writing ‘glottiana’ is also attached to the holotype. An additional label is attached to indicate the name under which the holotype stood in the BMNH collection; it reads ‘In BM. coll. under Trybliogrvapha mandibularis (Zett)’. A red-edged circular label with the printed word ‘Holotype’ in the centre has been attached, together with a determination label ‘HOLOTYPE of Psichacra glottiana Cam. det. J. Quinlan. 1973’. Identity. Valid species of Trybliogrvapha Forster, new combination Trybliographa glottiana (Cameron) comb. n. here established, and senior synonym of proxima Cameron, 1889 (see p- Io). Psichacra marshalli Cameron, 1883 : 369. Type(s) ¢, GREAT Britain: England, Devon, Barnstaple (Rev. T. A. Marshall) (lost). NEOTYPE 4, Great Britain: England, Surrey, Boxhill (C. G. Champion) (BMNH, ex coll. Cameron), here designated. I have been unable to locate the original male type(s). Dalla Torre & Kieffer (1910) and Hellén (1960) have referred to this species in published works and have used the characters that Cameron used in the original description for identifying it. The identity of marshalli is clear from the neotype here designated. It has a label ‘Box Hill 67 74’, a handwritten label ‘Gronotoma marshalli’, the BMNH accession label ‘Cameron 96-76 Box Hill’, and a red-edged label with the printed word ‘Neotype’ in the centre. A determination label ‘NEOTYPE of Psichacra marshalli Cam. det. J. Quinlan 1973,’ has also been attached to it. A further label attached to the neotype reads ‘New Genus B of Weld 1952, G. J. Kerrich det. 1960’. Identity. This valid species is not assignable to a recognized genus at present. It runs in the keys to genera of Weld (1952 : 108) to his genus ‘B’. It is proposed to publish a description of this new genus in another publication. Psichacra similis Cameron, 1883 : 368. LECTOTYPE 9, Great Britarn: Scotland, Lanarkshire, Cambuslang [publ. as Cambusland along the banks of the Clyde] (BMNH, ex coll. Cameron), here designated. Paralectotype. 1 ¢, same data as the lectotype except that the BMNH accession number is ‘Cameron 86-3 South Bank Clyde’ and a label ‘CBL’. A further label in Cameron’s handwriting ‘Eucoela mandibularis Zett’ is attached. Non-syntypic specimens. In the BMNH collection are five Cameron specimens that agree with the original description but they do not have the lectotype data: 1 g without data; 1 g, Clober; 1 ¢, Gloucester; 1 9, labelled ‘N’; and 1 9, Dalry. Cameron referred only to ‘Cambusland’ as the type-locality. All five specimens have been labelled “These specimens have no type status’. The lectotype, which has been remounted on a rectangular card, has the BMNH accession label ‘Cameron 96-76’. The original card mount is attached to the pin; on the underside of this card mount are the specific names ‘similis’ and ‘mandibularis’ written in ink over the pencilled word ‘Cambusland 7/9?’. On the upper surface of this card in the bottom left-hand corner is the number ‘5’; this is the figure number on the plate in Cameron (1890 : 220) to the species Trybliographa mandibularis (Zetterstedt). A circular purple- edged label with the printed word ‘Lectotype’ in the centre has been attached together with a determination label ‘LEcToTYPE of Psichacra similis Cam. det. J. Quinlan. 1973.’. Identity. Junior synonym of Trybliographa mandibularis (Zetterstedt, 1838 : 410) (Figites), syntypes ¢ 9, SwepEN (NR, Stockholm) [examined]. The synonymy of similis with mandibularis was first established by Cameron (1890 : 200) and is here confirmed after direct comparison of the lectotype and syntypes. Trybliographa crassicornis Cameron, 1889 : 64. Holotype 9, GREAT BRITAIN: Scotland, Lanarkshire, Cambuslang on the Clyde (BMNH, ex coll. Cameron). 16 J. QUINLAN The holotype has the BMNH accession label ‘Cameron 96-76 South Bk Clyde’; a further label in Cameron’s handwriting has the following abbreviations “CBL’ and ‘crassi’. This same rectangular label has the number ‘4’ in the bottom left-hand corner; this is the figure number on the plate in Cameron (1890). A red-edged circular label with the printed word ‘Holotype’ in the centre has been attached to the holotype, together with the determination label ‘HOLOTYPE of Trybliographa crassicornis Cam. det. J. Quinlan. 1973.’. Identity. This valid species clearly belongs in the genus Trybliographa Forster and this combination is herein confirmed. Trybliographa nigricornis Cameron, 1883 : 369. LECTOTYPE 9, Great BriTAIN: Scotland, Lanarkshire, Clober Wood (BMNH, ex coll. Cameron), here designated. Paralectotype. GREAT BRITAIN: I 4, Scotland, Kirkcudbrightshire, Dalry [labelled Daly], (BMNH, ex coll. Cameron). The lectotype has the BMNH accession label ‘Cameron 96-76 Clober Wood’. A further label in Cameron’s handwriting is attached to the lectotype and reads ‘Clober’ and ‘nigricornis Cam’. A circular purple-edged label with the printed word ‘Lectotype’ in the centre has been added, together with the determination label ‘LECTOTYPE of Trybliographa nigricornis Cam. det. J. Quinlan. 1973.’. Identity. Syn. n. of Trybliographa atra (Hartig, 1840: 201) (Cothonaspis), Holotype ©, GERMANY (ZSBS, Munich) [examined]. Trybliographa testaceipes Cameron, 1883 : 370. Holotype 9, GREAT Britain: Scotland, Ayrshire, Dalry (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Dalry’. The original rectangular card mount has the name ‘testaceipes’ on the underside and the number ‘8’ on the upper surface. This number is the figure number on the plate in Cameron (1890). A further label ‘testaceipes’ in Cameron’s handwriting is attached. A red-edged circular label with the printed word ‘Holotype’ has been attached, together with a determination label ‘HOLOTYPE of Trybliogvapha testaceipes Cam. det. J. Quinlan. 1973.’. Identity. Valid species of Trybliographa Forster. FIGITIDAE ANACHARITINAE Aegilips bicolorata Cameron, 1887: 194. Holotype g, GREAT Britain: England [publ. as probably London district], (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76’. On the reverse side of this label is the name ‘bicolorata’. Two further labels with the name ‘bicolorata’ in Cameron’s handwriting are attached to the holotype. A label with the number ‘5’ is attached to the holotype; this is the figure number on the plate in Cameron (1890). The holotype has a label ‘B.M. Type Hym. 7. 39’., together with a determination label ‘B.M. Type Hym. Aegilips bicolovata Cameron. 1887.’ Both forewings are damaged at the apices. Identity. Valid species in the genus Aegilips Walker. Aegilips ruficornis Cameron, 1883 : 372. Holotype 9, GREAT Britain: Scotland, Renfrew- shire, Bishopton (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Bishopton’. On the reverse side of this label is the name ‘ruficornis’. Further labels are attached and read ‘Bishop’ and ‘ruficornis’; both are in Cameron’s handwriting. A label with the number ‘8’ is also attached; this number is the figure number on the plate in Cameron (1890). The holotype bears a BMNH Type Hym. 7.40. label. A further label ‘B. M. Type. Hym. Aegilips ruficornis Cameron. 1883’ is attached to the holotype. Identity. Valid species in the genus Aegilips Walker. BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 17 Aegilips scotica Cameron, 1883 : 372. LECTOTYPE g, Great Britain: Scotland, Inverness-shire, Glenmoriston (BMNH, ex coll. Cameron), here designated. Paralectotype. 1 ¢, same data as the holotype. The lectotype and paralectotype have the BMNH accession label ‘Cameron 96-76’. The lectotype has a label ‘scotica’ in Cameron’s handwriting. A purple-edged circular label with the printed word ‘Lectotype’ has been added, together with a determination label ‘LECTOTYPE of Aegilips scotica Cam. det. J. Quinlan. 1973.’.. The left antenna of the lectotype is damaged. Identity. Syn. n. of Xyalaspis abietina (Thomson, 1861 : 412), holotype 3 [publ. as 9], SWEDEN: Ostergéthland (Prof. Zetterstedt) (UZI, Lund) [examined]. (One other male, not a syntype, stands under abietina in the UZI, Lund.) Aegilips striolata Cameron, 1883 : 373. Holotype g, GREAT BRITAIN: Scotland, Stirlingshire, Mugdock near Glasgow (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76’; on the reverse side of this label is the locality ‘Mugdock’. A further label with the number ‘3’ is attached; this number is the paragraph number on the appropriate page in Cameron (1890 : 181). An additional label in Cameron’s handwriting ‘striolata’ is attached. A red-edged circular label with the printed word ‘Holotype’ has been attached to the holotype, together with a determination label ‘HOLOTYPE of Aegilips striolata Cam. det. J. Quinlan. 1973’. Identity. Valid species in the genus Aegilips Walker. ASPICERINAE Onychia nigripes Cameron, 1879: 112. Holotype 9, Great Britain: England, Suffolk, Norwich (J. B. Bridgman) (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76’; on the reverse side of this label is the name ‘nigripes’. On the reverse side of the original card mount is the number ‘198’ or ‘19S’. A further label has the number ‘6’, which is the figure number on the plate in Cameron (1890). A determination label ‘nigripes’ in Cameron’s handwriting is attached. No locality label is attached but according to the original description Cameron only saw one specimen collected by Bridgman. A red-edged circular label with the printed word ‘Holotype’ in the centre has been attached, together with a determination label ‘HOLOTYPE of Onychia nigripes Cam. det. J. Quinlan. 1973.’. Identity. Junior synonym of Callaspidia defonscolombet Dahlbom, synonymy first established by Cameron (1890 : 177) and here confirmed by comparison of types [defonsco- lombei holotype 3, SWEDEN: Furillen vid Gottland (UZI, Lund)]}. (This species was first described by Dahlbom (1842 : 13) as Callaspidia De Fonscolombei. Dalla Torre & Kieffer (1910 : 64) were the first authors to coalesce the name to defonscolombet, which is correct under the International Code of Zoological Nomenclature.) FIGITINAE Melanips femoralis Cameron, 1883 :371. Holotype 2, GREAT BriTAIN: Scotland, Suther- land, Bonar Bridge, vi (BMNH, ex coll. Cameron). The holotype has the BMNH accession label ‘Cameron 96-76 Bonar Bridge’. A determi- nation label ‘femoralis’ and the data label ‘Bonar’ in Cameron’s handwriting is attached to the holotype. A red-edged circular label with the printed words ‘Type H. T.’, together with a label ‘B. M. Hym. 7.41’ and a determination label ‘B. M. Type Hym. Melanips femoralis Cameron 1883’, are attached to the holotype. Identity. Valid species of Melanips Giraud. 18 J. QUINLAN SUMMARY OF THE PRESENT NAMES DISCUSSED New synonymy Aegilips scotica Cameron syn. n. of Xyalaspis abietina (Thomson). Allotria ancylocera Cameron syn. n. of Alloxysta victrix (Westwood). Allotria pleuralis Cameron syn, n. of Alloxysta testacea (Hartig). Allotria ruficollis Cameron syn n. of Alloxysta erythrothorax (Hartig). Diastrophus aphidivorus Cameron syn. n. of Diastrophus rubi (Bouché). Eucoila proxima Cameron syn. n. of Trybliographa glottiana (Cameron). Trybliographa nigvicornis Cameron syn. n. of Trybliographa atra (Hartig). Confirmed synonymy Aulax graminis Cameron, junior synonym of Aulacidea hieracii (Bouché). Psichacra similis Cameron, junior synonym of Trybliogvapha mandibularis (Zetterstedt). Onychia nigripes Cameron, junior synonym of Callaspidia defonscolombei Dahlbom. New combinations Alloxysta testacea (Hartig) comb. n. Phaenoglyphis dolichocera (Cameron) comb. n. Trybliographa fortinervis (Cameron) comb. n. Trybliographa glottiana (Cameron) comb. n. Trybliogvapha scotica (Cameron) comb. n. Confirmed combinations Alloxysta basimacula (Cameron). Alloxysta caledonica (Cameron). Alloxysta erythrothorax (Hartig). Alloxysta megaptera (Cameron). Alloxysta mullensis (Cameron). Alloxysta piceomaculata (Cameron). Trybliographa gracilicornis (Cameron). Rhynchacis crassiclava (Cameron). Nomina dubia Allotria collina Cameron. Allotria maculicornis Cameron. Allotria perplexa Cameron. Allotria ruficeps Cameron. Allotria salicis Cameron. Phaenoglyphis forticornis Cameron. Kleditoma gracilicornis Cameron. Kleditoma marshalli Cameron. Kleditoma longicornis Cameron. SUMMARY OF LECTOTYPES AND NEOTYPES DESIGNATED Lectotypes Aegilips scotica Cameron. Allotria basimacula Cameron. BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON 19 Allotria crassa Cameron. Allotria dolichocera Cameron. Allotria pleuralis Cameron. Aulax graminis Cameron. Eucoila gracilicornis Cameron. Eucoila proxima Cameron. Eucoila scotica Cameron. Kleditoma elegans Cameron. Psichacra similis Cameron. Trybliographa nigricornis Cameron. Neotypes Kleditoma caledonica Cameron. Kleditoma filicornis Cameron. Kleditoma nigripes Cameron. Psichacra dalei Cameron. Psichacra marshalli Cameron. REFERENCES BoucnkE, P. F. 1834. Naturgeschichte der Insekten pp. i-v, 1-216. Berlin. CaMERON, P. 1875. On some new or little known British Hymenoptera. Proc. nat. Hist. Soc, Glasgow 2 : 304-324. 1879. On some new or little known British Hymenoptera. Trans. ent. Soc. Lond. 12 : 107-119. 1883. Descriptions of sixteen new species of parasitic Cynipidae, chiefly from Scotland. Trans. ent. Soc. Lond. 16 : 365-374. 1886. The Fauna of Scotland with special reference to Clydesdale and the Western District. Hymenoptera, part II. Supplement to Tenthredinidae, pp. 53-95. Glasgow. 1887. Hymenopterological notes. I. On some new or little known British Hymenoptera. Entomologist’s mon. Mag. 23 : 193-195. 1888a. Descriptions of twenty three new species of Hymenoptera. Mem. Proc. Manchr lit. phil. Soc. (4) 1 : 160-163. 1888b. On some new or little known British parasitic Cynipidae. Entomologist’s mon. Mag. 24 : 209-211. 1889. On the British species of Allotrinae, with descriptions of other new species of parasitic Cynipidae. Mem. Proc. Manchr lit. phil. Soc. (4) 2 : 53-69. 1890. Monograph of British phytophagous Hymenoptera 3 (1889): 1-274. London. 1893. Monograph of British phytophagous Hymenoptera 4 (1892): 1-248. London. DaHLBom, G. 1842. Onychia och Callaspidia, tvenne for Skandinaviens Fauna nya Inseht- Slaigten, hévande till Gallaple-Steklarnes naturliga grupp. 1-16, 9 figs Lund. DaLia TorrE, K. W. & KIEFFER, J. J. 1910. Cynipidae. Das Tierreich 24 : 1-891. Berlin. Eapy, R. D. & QuINLAN. J. 1963. Hymenoptera Cynipoidea. Key to families and sub- families, and Cynipinae (including galls). Handbk Ident. Br. Insects 8 (1a) : I-81. Foérster; A. 1869. Ueber die Gallwespen. Verh. zool.-bot. Ges. Wien 19 : 327-370. GrIRAuD, J. 1856. Observations sur quelques espéces d’Hyménoptéres rares ou peu connues, trouvées dans les environs de Vienne. Verh. zool.-bot. Ver. Wien 6 : 179-188. 1860. Enumération des Figitides de l’Autriche. Verh. zool.-bot. Ges. Wien. 10 : 23-176. Hartic, T. 1840. Ueber die Familie der Gallwespen. Z. Ent. (Germar) 2 : 176-209. 1841. Erster Nachtrag zur Naturgeschichte der Gallwespen. Z. Ent. (Germar) 3 : 322-358. 1843. Zweiter Nachtrag zur Naturgeschichte der Gallwespen. Z. Ent. (Germar) 4 : 395- 422. 20 J. QUINLAN HELLEN, W. 1960. Die Eucoilinen Finnlands (Hym: Cyn). Fauna fenn. 9: 1-29. 1963. Die Alloxystinen Finnlands (Hymenoptera, Cynipidae). Fauna fenn. 15 : 1-23. Ionescu, M. A. 1969. Hymenoptera Cynipoidea. Fam. Ibaliidae, subfam. Ibaliinae, Fam. Figitidae—Aspicerinae, Anacharitinae. Figitinae. Fam. Cynipidae—subfam. Eucoilinae, Charipinae. [In Romanian.] Fauna Repub. pop. rom. (Insecta) 9 (6) : 1-285, 57 figs, 22 pls, 8 maps. KerricuH, G. J. & QUINLAN, J. 1960. Studies on Eucoiline Cynipoidea (Hym). Opusc. ent. 25 : 179-196. Kort, G. S. & Hincxs, W. D. 1945. A check list of British insects pp. lix, 483. Stockport. MarsHALL, T. A. 1870. Onsome British Cynipidae. Entomologist’s mon. Mag. 6 : 178-181. Morey, C. 1915. The Rev. T. A. Marshall’s localities. Entomologist 48 : 23-24. RicHarps, O. W. 1956. Hymenoptera, Introduction and Key to Families. Handbk Ident. Br. Insects 6 (1) : 1-38. RoHWER, S. A. & Facan, M. 1917. The type species of the genera of Cynipoidea, or the gall wasps and the parasitic Cynipoids. Proc. U. S. natn. Mus. 53 : 357-380. TuHomson, C. G. 1862. Férsdk till uppstallning och beskrifning af Sveriges Figiter. Ofvers. K. VetenskAkad. Forh. Stockh. 1861 : 395-420. 1877. Ofversigt af Sveriges Cynips-Arter. Opusc. ent. 8 : 778-820. WELD, L. H. 1952. Cynipoidea (Hym.) 1905-1950, pp. 351. Ann Arbor, Michigan. [Privately published. ] WEstwoop, J. O. 1833. Notice of the habits of a Cynipideous insect, parasitic upon the Rose Louse (Aphis rosae), with descriptions of several other parasitic Hymenoptera. Mag. nat. Hist. 6 : 491-497. ZETTERSTEDT, J. W. 1838. Insecta Lapponica 1 : 1-1139. Lipsiae. INDEX Aegilips, 16, 17 forticornis, 9 affinis, 11 fortinervis, 10 Allotria, 6, 7, 8, 9 ancylocera, 6 glottiana, 14 aphidivorus, 9 gracilicornis, Eucoila, 10 Aulax, 9 gracilicornis, Kleditoma, 12 graminis, 9 bicolorata, 16 basimacula, 6 Kleditoma, I1, 12, 13, 14 caledonica, Allotria, 6 caledonica, Kleditoma, I1 longicornis, 12 longipennis, 13 collina, 7 crassa, 7 crassiclava, 12 maculicollis, 18 crassicornis, 15 marshalli, Kleditoma, 13 marshalli, Psichacra, 15 dalei, 14 megaptera, 8 dolichocera, 7 Melanips, 17 melanopoda, 13 elegans, 12 mullensis, 8 Eucoila, 10, 11 nigricornis, 16 femoralis, 17 nigripes, Kleditoma, 13 filicornis, 12 nigripes, Onychia, 17 BRITISH CYNIPOIDEA DESCRIBED BY P. CAMERON Onychia, 17 perplexa, 8 piceomaculata, 8 picipes, 13 pleuralis, 8 proxima, 10 Psichacra, 14, 15 ruficeps, 9 ruficollis, 9 ruficornis, 16 J. QUINLAN Department of Entomology British MusEuM (NATURAL History) CROMWELL ROAD Lonpon SW7 5BD salicis, 9 scotica, Aegilips, 17 scotica, Eucoila, 11 similis, 15 striata, 14 striaticollis, 14 striolata, 17 testaceipes, 16 truncata, 14 Trybliographa, 15, 16 21 - : - oO : 7 ; oe - - — ‘~ _ : p 7 Sy: -_ é _ _ : = ee ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. {3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50. . HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounn, L. A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. 5. . AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. {5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Eviot, J.N. Ananalysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. £12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera Termitidae). Pp. 244: g plates, 661 text-figures. July, 1972. £9.90. . CROSSKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . VON HayeK, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CRossKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS, OLD WOKING, SURREY, ENGLAND A CATALOGUE OF THE TYPE-SPECIMENS OF THE CETONIINAE (COLEOPTERA : SCARABAEIDAE) DESCRIBED BY G. J. ARROW WITH A COMPLETE BIBLIOGRAPHY OF HIS ENTOMOLOGICAL WORKS M. E. BACCHUS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 2 LONDON : 1974 A CATALOGUE OF THE- TYPE-SPECIMENS OF THE CETONIINAE (COLEOPTERA : SCARABAEIDAE) DESCRIBED BY G. J. ARROW WITH A COMPLETE BIBLIOGRAPHY OF HIS ENTOMOLOGICAL WORKS BY MICHAEL EDWARD BACCHUS Pp. 23-44 BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 2 LONDON : 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, ts issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 31 No. 4 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.). © Trustees of the British Museum (Natural History), 1975 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 7 February, 1975 Price £4.80 A GUIDE TO THE GENERA AND: SPEGIES OF PARNASSIINAE (LEPIDOPTERA : PAPILIONIDAE) By P. R, ACKERY CONTENTS Page SYNOPSIS : ° ’ . ° 2 ° . , 5 : 73 INTRODUCTION - : ° . ° ‘ : : - : 73 ACKNOWLEDGEMENTS . , : . ; ; : ; 74 KEY TO THE GENERA OF PARNASSIINAE r : : 74 NOTES ON THE GENERA, WITH KEYS TO THE SPECIES WHERE APPLICABLE . 75 Archon Hiibner . : : : ; : : . : 75 Hypermnestra Ménétriés . : ‘ : : : ; : 75 Parnassius Latreille E ; : : ? : : : : 70 Sericinus Westwood ‘ 3 : ; 4 F A A : 90 Parnalius Rafinesque_ . ; : : ‘ : : ° : go Luehdorfia Criiger . : : ‘ : : : ‘ : : 92 Bhutanitis Atkinson 3 5 : ; ; F ‘ : : 93 REFERENCES . . ‘ ‘ ; 5 : ‘ : : : 94 INDEX . A ‘ é : : : ; ; ; ‘ ; 105 SYNOPSIS Illustrated keys are given to the seven genera and 44 species of Parnassiinae here recognized, together with brief notes on the distribution and larval food plants of each species. For polytypic species a list is given of the subspecies that differ most markedly from the typical form. One generic synonym is newly established. INTRODUCTION THE inconsistency of the wing pattern within many species of the Parnassiinae has inspired the description of numerous subspecies and forms, principally in the works of Fruhstorfer, Bang-Haas, Bryk and Eisner. Thus, although the literature is extensive, it is concerned mainly with intra-specific variation rather than the definition and identification of the recognized species. Bryk (1934; 1935) includes keys to the Parnassiinae in his extensive monograph; however, the lack of availability of this work, together with the great emphasis placed on variation within species throughout the study, renders it unsuitable as a ready means of identification to species. It is hoped that this present work will to some degree fill this apparent gap in the literature available to lepidopterists interested in the Parnassiinae. There has been considerable difference of opinion concerning the higher classifica- tion of this subfamily, the two tribes here recognized, the Parnassiini and Zerynthiini, being regarded by Bryk (1934; 1935) and Ford (1944) as subfamilies, while Clench 74 Pp. KR. ACKERY (1955) and Hemming (1960) have suggested that they be accredited with family status. I have followed Ehrlich (1958) and Munroe (1960) in treating the two groups as tribes which together form the subfamily Parnassiinae. At the generic and specific levels there has been more general agreement, except over the status of the genus Parnassius and its species (see pp. 76-77). Sericinus, Archon and Hypermnestra are at present monotypic, Bhutanitis contains four species and the far-eastern genus Luehdorfia consists of two species. Allancastria, hitherto a valid monotypic genus, is here considered to be a junior synonym of Parnalius, which now includes three species. I have adopted Munroe’s (1960) broad concept of Parnassius, which is regarded here as comprising 32 species. Examples of each genus are illustrated in the Frontispiece, figs 1-7. To include notes on all the described forms is certainly not the purpose of this work. I have, however, listed under the relevant species some of those subspecies that seem to differ most markedly from the more typical forms, together with their localities and points of difference. No attempt has been made to produce a comprehensive list of all the subspecies described. ACKNOWLEDGEMENTS I wish to thank Mr R. I. Vane-Wright for his help in the preparation of this work together with Messrs C. F. Huggins, R. L. Smiles and M. A. Kirby whose helpful suggestions led to many improvements, particularly in the key to Parnassius. I also extend my thanks to Messrs T. G. Howarth, H. K. Clench, H. J. Epstein, O. Kudrna, A. Tsvetejev, P. M. Hammond, R. B. Grubh, J. C. Eisner, Suguru Igarashi and Kazuo Saitoh. To Mr P. York and Mr F. Greenaway I am indebted for the production of the photographs. KEY TO THE GENERA OF PARNASSIINAE I Outer margin of hindwing rounded, without tails . : , (Parnassiini) 2 Tails of various lengths present on the outer margin of hindwing (Zerynthiini) 4 2 (1) Claspers of male narrow (Text-fig. 1) ; forewing with five radial veins (Text-fig. 18) ARCHON Hiibner (p. 75) ~ Claspers of male various, never narrow; forewing with four radial veins . 3 3 (2) Male tarsal claws equal (Text-fig. 29) . . HYPERMNESTRA Ménétriés (p. 75) Male tarsal claws sub-equal (Text-fig. 30) . . PARNASSIUS Latreille (p. 76) 4 (1) Vein M, of hindwing produced to a short tapering or rounded tooth (Text-figs 07;-19) ; . PARNALIUS Rafinesque (p. 90) - Vein M, of hindwing produced to a distinct tail ‘ 5 (4) Uncus short, bifid (Text-fig. 16); hindwing tail on vein M, longer than the discal cell of hindwing; vein R, of oe usually arising from discal cell (Text-fig. 26) : : . SERICINUS Westwood (p. 90) - Uncus elongate, bifid; hindwing tail on vein M; no longer than the discal cell; vein R; normally stalked with R,, 5 . 6 6 (5) Precostal cell wide; tails well developed on veins eae and Cea of hindwing (Text-fig. 25); internal ventral surface of claspers smooth (Text-figs 14, 15) BHUTANITIS Atkinson (p. 93) GENERA AND SPECIES OF PARNASSIINAE 75 - Precostal cell narrow; tails rudimentary on veins Cu,, and Cu,, of hindwing (Text-fig. 23); internal ventral surface of claspers spinose (Text-figs 12, 13) LUEHDORFPFIA Criiger (p. 92) NOTES ON THE GENERA, WITH KEYS TO THE SPECIES WHERE APPLICABLE Tribe PARNASSIINI Parnassinae Swainson, 1840 : 87,90. Type-genus: Parnassius Latreille. ARCHON Hiibner [Doritis Fabricius sensu Hiibner, [1819] : 86.] Archon Hiibner, [1822]:2, 6, 8, 9. Type-species: Papilio thia Hiibner, by subsequent designation (Scudder, 1875 : 117). Dorarchon Rothschild, 1918 : 219. Type-species: Papilio apollinus Herbst, by monotypy. Archon Hiibner; Bryk, 1934 : 19. Archon Hiibner; Munroe, 1960 : Io. The name Doritis is still commonly used for this taxon, although Papilio apollo Linnaeus is clearly the type-species of Dorvitis, not Papilio apollinus Herbst as was erroneously believed. Archon apollinus (Herbst) (Pl. 1, figs 1, 2, Text-figs 1, 18) Papilio apollinus Herbst, 1798 : 156. Papilio thia Hiibner, [1806] : 60. Archon apollinus (Herbst); Bryk, 1922 : 224; 1934 : 23. Archon apollinus (Herbst); Eisner, 1966 : 89. DISTRIBUTION. Rumania. Bulgaria. Turkey. U.S.S.R.: Armenia, Turk- menistan. Greece. Syria. Iraq. Lebanon. Israel. LARVAL FOOD PLANTS. Aristolochia hastata (Higgins & Riley, 1970). A. bodame (Bryk, 1934). A. maurorum (Suguru Igarashi, in litt.). HYPERMNESTRA Ménétriés Ismene Nickerl, 1846: 207. Type-species: Ismene helios Nickerl, by monotypy. [Junior homonym of both Ismene Savigny, 1816, and Ismene Swainson, 1820.] Hypermnestra Ménétriés, 1848 : pl. 6, fig. 1. [Replacement name for Ismene Nickerl.] Hypermnestra Ménétriés; Bryk, 1935 : 4. Hypermnesiva Ménétriés; Munroe, 1960 : Io. Ehrlich (1958) says that there are reports of Parnassius species in which the males have nearly symmetrical claws as in Hypermnestra Ménétriés; however, I have always found the tarsal claws of male Parnassius to be subequal. 76 P. R. ACKERY Hypermnestra helios (Nickerl) (Pl. 1, figs 3, 4, Text-figs 2, 20, 29) Ismene helios Nickerl, 1846 : 208, pl. 3, figs a-g. Hypermnestra helios (Nickerl); Bryk, 1935 : 7. Hypermnestra heltos (Nickerl); Eisner, 1966 : 121 DISTRIBUTION. Iran. Afghanistan. U.S.S.R.: Uzbekistan, Kirghizia. LARVAL FOOD PLANTS. Zygophyllum (Munroe, 1960). Z. turcomanicum (Verity, 1906). Z. atriplicoides (Bryk, 1935). Z. fabago, Z. portulacoides (Eisner, 1968). Z. gontsharovii (Stschetkin, 1963). PARNASSIUS Latreille Parnassius Latreille, 1804 : 185, 199. Type-species: Papilio apollo Linnaeus, by monotypy. Doritis Fabricius, 1807 : 283. Type-species: Papilio apollo Linnaeus, by subsequent designation (Dalman, 1816 : 60). Tadumia Moore, 1902: 116. Type-species: Parnassius acco Gray, by original designation. [Synonymized by Munroe, 1960: I1.] Kailasitus Moore, 1902: 118. Type-species: Parnassius charltonius Gray, by original designation. [Synonymized by Munroe, 1960 : 11.] Koramius Moore, 1902:120. Type-species: Parnassius delphius Eversmann, by original designation. [Synonymized by Munroe, 1960 : 11.] Lingamius Bryk, 1935 : 538-540. Type-species: Parnassius hardwickii Gray, by original designation. [Synonymized by Munroe, 1960 : I11.] Eukovramius Bryk, 1935 : 630, 673-674. Type-species: Parnassius imperator Oberthiir, by monotypy. [Synonymized by Munroe, 1960 : 11.] Bryk (1935) divides Parnassius into five genera on the basis of differences in the wing venation and relative lengths of the foretibial epiphysis. As shown by Munroe (1960) the groups so defined do not agree with those indicated by the male genitalia. Iam adopting Munroe’s broad concept of this genus, but not his suggested division into two subgenera, Parnassius s. str. and Doritis. The latter name cannot have any valid use as it is a junior objective synonym of Parnassius. Elwes (1886) attempted to define the limits of the species, focusing attention on the wide differences in the shape of the sphragis. The comparative morphology of the species is discussed by Hering (1932) who recognizes only six distinct species, this arrangement being followed by Fisher (1950). A revision of the mnemosyne group is presented by Miiller (1973), partly based on new morphological characters derived from the comparative structure and arrangement of the wing scales. Valuable information on this genus is to be found in some of the faunistic studies on butterflies. Kurentsov (1970) illustrates and discusses the Parnassius of the eastern U.S.S.R. and the distinguishing characters exhibited by the three western Palaearctic species are given by Higgins & Riley (1970). Yokohama & Wakabayashi (1968) illustrate the representatives in the Japanese fauna, whilst five species are included by Seok (1939) in his check list of Korean butterflies. The Asian species, nomion Fischer de Waldheim, is doubtfully listed by Wilson (1961) in his identification key of North American Papilionidae, but excluded by Dos Passos (1964). GENERA AND SPECIES OF PARNASSIINAE Wai The studies of the Indian fauna provide useful keys, notably in the works of Evans (1927) and Talbot (1939). A complete key to the 34 species then recognized is given by Bryk (1935). Munroe (1960) notes 37 species, five more than Eisner (1966), but the only species included here are those listed by Eisner in his index to the ‘Parnassiana’ and ‘Parnassius Nova’ series. The apparent uniformity of structure within the species-groups, as defined by Munroe (1960), together with the multiplicity of variations in the wing pattern within most of the species, presents some difficulty in the preparation of a key to the species of Parnassius. The wing pattern is used as the principal means of differentiating between the species since the structural characters derived from the genitalia, wing venation, sphragis and foretibial epiphysis, are generally applicable at the species-group level only. As the following key is based on wing pattern at the species level, it is to be expected that atypical specimens, particularly in the actius-jacquemontii-phoebus subgroup, will not always run out satisfactorily. It must therefore be emphasized that the present work should be used only as a guide. This genus is generally distributed throughout the mountainous and northern areas of the Palaearctic and western Nearctic regions, the inaccessibility of many likely localities probably accounting for the rarity of some species. KEY TO THE SPECIES OF PARNASSIUS LATREILLE I Uncus normally plainly visible, spatulate, lying between the two elongate processes on the tenth tergite (Text-fig. 31); sphragis keeled and with a simple, single, backward-pointing flange (PI. 15, fig. 98) [keel lacking in epaphus Oberthiir]; hindwing upperside lacking submarginal series of blue spots . ; 2 - Uncus sometimes enclosed by eighth abdominal tergite ‘and bifid, never spatulate (Text-fig. 32) [simple and truncate in simo Gray]; sphragis various, never keeled and always lacking a single backward-pointing flange (Pl. 15, figs 99-105); hindwing upperside often with submarginal series of blue centred spots . ‘ : ; : : . F 16 2 (1) Antennae dark, lacking white scales : : ‘ ; : : : : 3 - Antennae with some white scales : b 3 (2) Submarginal band of forewing and hindwi ing broken up into a distinct series : of internervular black spots (Pl. 1, figs 5, 6) . apollonius (Eversmann) (p. 80) ~ Submarginal band of hindwing indistinct or absent 4 (3) Pubescence of body mostly pale beneath; wing veins above exceptionally distinct, with a covering of black scales; forewing above usually lacking red markings (Pl. 2, figs 9, 10) . ’ bremeri Bremer (p. 81) = Body clothed with thick black pubescence beneath; wing veins above without black scales; forewing above usually with red markings (Pl. 1, figs 7, 8) honrathi Staudinger (p. i 5 (2) Scaling of antennae, excluding the club, completely white above . = Scaling of antennae, excluding the club, dark and white above. ; 6 (5) Margins distinctly chequered, being black about the wing veins and white in the internervular areas; upper surface of hindwing often with a distinct red basal spot (Pl. 3, figs 17, 18) . . nomion Fischer de Waldheim (p. 82) = Margins usually white, sometimes indistinctly chequered; upper surface of hindwing seldom with a red basal spot (PI. 3, figs 23, 24) apollo (Linnaeus) (p. 83) (5) (7) (8) Oo 1oIln 10 (9) II (10) 12 (7) 13 (12) Pe. Re ACK ERY: Males. . . ; : : ; ; ; : ‘ : : 8 Females . : ‘ : : : : 12 Eighth abdominal tergite rounded posterio- laterally ; : : : : 9 Eighth abdominal tergite pointed posterio-laterally . , : 10 Marginal and submarginal forewing bands poorly developed; ground colour of upperside creamy white; discal spot in cell Cu,» of forewing upperside absent or rudimentary, seldom distinctly scaled below and never centred with red (Pl. 2, fig. 11) P . phoebus (Fabricius) (p. 81) Marginal and submarginal bands of forewing well developed; ground colour of upperside grey-white, often distinctly dusted with black; discal spot in cell Cu,, of forewing upperside usually present, distinctly scaled below and often centred with red (Pl. 3, fig. 21) : tianschanicus Oberthiir (p. 82) Basal red spot usually present in hindwing upperside; discal spot in cell Cu;p of forewing underside often heavily scaled, sometimes centred with red (Pl. 2, fig. 15) ; . jacquemontii Boisduval (p. 81) Basal red spot normally absent from the hindwing upperside, if present the discal spot in cell Cu, of forewing underside is ssi scaled, not centred with red - - Il Black basal scaling normally quite extensive; margins of forewings usually distinctly chequered, being black about the veins and white in the inter- nervular areas (Pl. 3, fig. 19) . . epaphus Oberthiir (p. 82) Black basal scaling usually less extensive; margins of forewing seldom distinctly chequered (Pl. 2, fig. 13) : : y actius (Eversmann) (p. 81) Sphragis lacking a keel (PI. 3, fig. 20) ; : . epaphus Oberthiir (p. aA Sphragis strongly keeled . ‘ 13 Discal spot in cell Cup of forewing endemade distinctly PANY fea hada! spot never present in hindwing upperside; upperside distinctly dusted with black scales (Pl. 3, fig. 22) 4 : . tianschanicus Oberthiir (p. 82) Discal spot in cell Cu,p of ae dense normally only lightly scaled, if distinctly scaled then red basal spot normally present in poe upperside; upperside not distinctly dusted with black : ; 14 Abdomen usually hairy, almost as much so as the male [Asian species Sey (Ples, Hesi4)) M2 . actius Cate (p. 81) Abdomen not exceptionally hairy [except European phoebus] 3 15 Hindwing upperside normally with a distinct red basal spot [Asian species only] (Pl. 2, fig. 16) : ; . jacquemontii Boisduval (p. 81) Hindwing upperside seldom with a ‘distinct red basal spot [N. American, European & Asian species] (Pl. 2, fig. 12) : phoebus (Fabricius) (p. 81) Foretibial epiphysis short, not reaching the end of the tibia (Text-fig. 27); hindwing upperside without a submarginal series of blue spots (except orleans Oberthiir) % 17 Foretibial epiphysis longer, often reaching the end of the tibia (Text-fig. 28): hindwing upperside usually with a submarginal series of blue spots. . : 24 Hindwing underside lacking red ocelli 2 : : : ° : . 18 Hindwing underside usually with red ocelli : : ° 20 Outer surface of palpi dark, occasionally with a few light yellows per face lacking white hairs; bifid uncus with pane ventral processes (Text-fig. 6, Pl. 4, figs 29, 30) : : glacialis Butler (p. 84) Outer surface of palpi with aie Ag ate with some white hairs; bifid uncus without ventral processes. 19 Forewing upperside usually with two black ae in ee discal call: cieoen of male weakly produced posteriorly, never ending in a long thin point (Pl. 4, figs 25, 26) . - ; : . mmnemosyne (Linnaeus) (p. 83) 22 (20) 23 (22) GENERA AND SPECIES OF PARNASSIINAE Forewing upperside without two black spots in the discal cell; clasper of male strongly produced posteriorly, ending in a long thin point (Pl. 4, figs 27, 28) Face usually with golden yellow pubescence. Pubescence of face black, pale yellow or white Basal black scaling of hindwing upperside sharply Sieresnateu: margins of hindwing underside black; wings of male often yellow (Pl. 4, figs 31, 32, Pl. 5, figs 33, 34) , eversmanni Ménétriés (p. Basal dark scaling of hindwing upperside not sharply divided from the white ground colour; margins of hindwing underside partly white; wings of male never yellow (PI. 5, figs 39, 40) : ; ; clodius Ménétriés (p. Hindwing upperside and underside with a Sueiarernal series of internervular blue-centred spots; margins of forewing upperside usually distinctly chequered, black about the veins and white in the internervular areas (Pl. 6, figs 41, 42) . : : orleans Oberthiir (p. Hindwing upperside lacking blue- centred enue spots; forewing margins never divided into distinct black and white areas : Forewing apex largely hyaline, the submarginal antecheRgalay mye ery usually absent, if present pale and indistinct; veins of hindwing underside pale (Pl. 5, figs 35, 36) . : nordmanni (Nordmann) (p. Forewing above with submarginal internervular markings forming a distinct band in the wing apex; veins of hindwing underside dark (PI. 5, figs 37, 38) ariadne Lederer (p. Vein R, stalked with vein R,_;, not arising from discal cell (Text-fig 24) Vein R, arising from discal cell (Text-figs 21, 22) : z 3 Hindwing upperside usually with large red postdiscal spots . Red postdiscal spots of hindwing upperside small or absent . Hindwing blue-black internervular submarginal spots absent from ail nye sphragis straight (Pl. 9, figs 67, 68, Pl. 15, fig. 105) inopinatus Kotzsch (p. Hindwing blue-black internervular submarginal spot present in cell R;; sphragis coiled (Pl. 9, figs 65, 66, Pl. 15, fig. 104) . charltonius Gray (p. Hindwing upperside without red markings but with an orange postdiscal band, wide in female, narrow in male (Pl. 9, figs 71, 72) . autocrator Avinoff (p. Hindwing upperside with red as and lacking an orange postdiscal band (Pl. 9, figs 69, 70) : ‘ loxias Piingeler (p. Vein R, anastomosing with vein Re (Text- no on Vein Re not anastomosing with vein R, (Text-fig. 22) Hindwing upperside usually with inoee blue-centred cabinarainal ‘spots: sphragis large, bilobate (Pl. 8, figs 63, 64, Pl. 15, fig. 103) imperator Oberthiir (p. Hindwing upperside seldom with large blue-centred submarginal spots; sete never bilobate Uncus short, strongly bifid; aphiage formally iatsiay. flattened: foone a complete ring about the abdomen (PI. 7, figs 51, 52, 53, 54) acco Gray (p. Uncus simple or weakly bifid; sphragis never laterally flattened or forming a complete ring about the abdomen Uncus short, simple (Text-fig. 8); sphragis rudimentary: hindwing underside with few postdiscal spots (Pl. 10, figs 73, 74) . 3 simo Gray (p. Uncus elongate, weakly bifid (Text-fig. 3); sphragis well formed, distinct; hindwing underside with a eae row of postdiscal spots (Pl. 10, figs 75, 76). . : tenedius Eversmann (p. Clasper broad; cenit process half 6 ere thirds the length of the clasper (Text-figs 4, 5); sphragis simple, usually laterally flattened (Pl. 15, fig. 100) . stubbendorfii Ménétriés (p. 79 84) Zr Zz 84) 85) 89) 88) 86) 89) RR) 80 Bo Ro ACKERY. - Clasper narrow, distally pointed; internal process one-third the length of the clasper (Text-fig. 7); sphragis bilobate (Pl. 15, figs 101, 102). 33 (32) Veins of hindwing underside bordered with white scales which cover the veins (Pl. 6, figs 45, 46) szechenyii Frivaldsky (p. 86) Wing veins clearly visible, not covered by white scales ; . 34 34 (33) Male with a distinct row of bristles anterior to the uncus; clasper produced dorsally to a distinct angle (Text-fig. 4); submarginal blue spots of hindwing upperside usually centred with white (Pl. 6, figs 43, 44) hardwickii Gray (p. 86) Male without a distinct row of bristles anterior to the uncus; clasper large, rounded; submarginal spots of hindwing upperside, when present, seldom centred with white (Pl. 6, figs 47, 48) . cephalus Grum-Grshimailo (p. 86) 35 (32) Sclerotized area of male eighth abdominal tergite produced posteriorly to give two finger-like projections; sphragis produced backwards, forming two lateral points (Pl. 15, fig. 101); basal black scaling of hindwing upperside usually very extensive, often surrounding the red ocelli of hindwing above (Pl. 8, figs 61, 62) acdestis Grum-Grshimailo (p. 87) Sclerotized area of male eighth abdominal tergite not strongly produced, never with finger-like projections ; sphragis produced backwards to form two ventral lobes (Pl. 15, fig. 102); basal black scaling of hindwing upperside usually rather less extensive 36 (35) Submarginal spots of hindwing upperside aoaaliy distinct and centred with blue (PI. 8, figs 57, 58, 59, 60) : ; ; delphius (Eversmann) (p. 87) Submarginal spots of hindwing upperside, when present, paler and not centred with blue (Pl. 7, figs 55, 56) patricius Niepelt (p. 87) 35 36 THE APOLLO-Grovup Parnassius apollonius (Eversmann) (Pl. 2; tigs5,.0) Doritis apollonius Eversmann, 1847 : 71, pl. 3, figs 1, 2. Parnassius apollonius (Eversmann); Bryk, 1935 : 176. Parnassius apollonius (Eversmann); Eisner, 1966 : 89. DISTRIBUTION. U.S.S.R.: Uzbekistan, Tadzhikistan, Kirghizia. China: western Sinkiang. LARVAL FOOD PLANTS. Salsola (Elwes, 1886). Scabiosa (Stichel,1907b). Radiola semenovt (A. Tsvetajev, pers. com.) Parnassius honrathi Staudinger (Pl. x, figs 7, 8) Parnassius honrathi Staudinger, 1882 : 161, pl. 1, figs 4, 5a, pl. 2, fig. 5. Parnassius honvathi Staudinger & Bang-Haas; Bryk, 1935 : 185. Parnassius honvathi Staudinger & Bang-Haas; Eisner, 1966 : 122. DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan. GENERA AND SPECIES OF PARNASSIINAE 81 Parnassius bremeri Bremer (Pl. 2, figs 9, 10) Parnassius bremeri Bremer, 1864 : 6 (Felder in litt.). Parnassius bremeri Bremer; Felder & Felder, 1865 : 133. Parnassius bremeri Bremer; Bryk, 1935 : 190. Parnassius bremeri Bremer; Eisner, 1966 : 94. DISTRIBUTION. China: Heilunkiang, Shansi, Hopei. U.S.S.R.: Russia (Chita, Khabarovsk, Sakhalin, Kamchatka). Korea. LARVAL FOOD PLANTS. Various species of Sedum (Stichel, 19070). S. aizon, S. ischida, S. ussuriensis, S. quadriflorum (Kurentsov, 1970). Parnassius phoebus (Fabricius) (Pl. 2, figs 11, 12, Text-fig. 31) Papilio phoebus Fabricius, 1793 : 181. Parnassius phoebus (Fabricius); Bryk, 1935 : 206. Parnassius phoebus (Fabricius); Eisner, 1966 : 157. DISTRIBUTION. Europe: Italy, Austria and Switzerland. [Maritime Alps and eastward to Styria and Grossglockner, rare in north, occasional in Allgauer Alps (Higgins & Riley, 1970).] U.S.A.: Alaska, Washington, Idaho, Montana, Dakota, Wyoming, California, Nevada, Utah, Colorado, New Mexico. Canada: British Columbia, Alberta. China: Sinkiang. U.S.S.R.: Russia (Irkutsk, Amur), Kazakhstan. Mongolia. LARVAL FOOD PLANTS. Saxifraga aizioides, Sempervivum montanum (Higgins & Riley, 1970). Sedum stenopetalum, Sempervivum and Saxifraga (Wilson, 1961). Sedum telephium, S. fabria, S. album, S. roseum and Sempervivum tectorum (Bryk, 1935). Saxifraga calycina, S. nivalis (Kurentsov, 1970). Carex filifolia, Gayophytum diffusum, Phlox douglasii, Sedum debile, S. obtusatum, S. wrightit (Tietz, 1972). Sedum lanceolatum (=stenopetalum) (Scott, 1973). Parnassius actius (Eversmann) (Pl. 2, figs 13, 14) Doritis actius Eversmann, 1843 : 540, pl. 9, figs 2a, b. Parnassius actius (Eversmann); Bryk, 1935 : 249. Parnassius actius (Eversmann); Eisner, 1966 : 82. DisTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan, Kirghizia, Kazakhstan. China: Sinkiang, Kansu. Pakistan. Kashmir. Parnassius jacquemontii Boisduval (Pl. 2, figs 15, 16) Parnassius jacquemontii Boisduval, 1836 : 400. Parnassius jacquemontii Boisduval; Bryk, 1935 : 257- 82 Po RACK ERY, Parnassius jacquemontii Boisduvai; Eisner, 1966 : 128. Parnassius jacquemontii Boisduval; Ackery, 1973 : 6. DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan, Uzbekistan. China: Sinkiang, Kansu, Szechwan. N.India. Pakistan. Tibet. Variable species. Red basal spot sometimes absent. Superficially many forms resemble epaphus Oberthiir, but the sphragis of the female always bears a keel. Subsp. mercurius Grum-Grshimailo [Tibet, Amdo], subsp. jupiterius Bang-Haas {Kansu, Pullow mont, Minschan], subsp. tatungi Bryk & Eisner [Kansu, Richthofen Mts, Nanschan Mts]. Wing margins distinctly chequered. Subsp. thibetanus Leech [Szechwan, How-Kow, Ta-tsien-lou]. Dusted with black scales. Parnassius epaphus Oberthiir (Pl. 3, figs 19, 20) Parnassius epaphus Oberthiir, 1879 : 23. Parnassius epaphus Oberthiir; Bryk, 1935 : 270. Parnassius epaphus Oberthiir; Eisner, 1966 : 106. Parnassius epaphus Oberthiir; Ackery, 1973 : 5. DistrisuTion. Afghanistan. Pakistan. Kashmir. N. India. Nepal. Sikkim. Tibet. China: Sinkiang, Szechwan, Kansu, Tsinghai. Variable species. Red basal spot sometimes present causing many specimens to resemble jacquemontii Boisduval, but sphragis always without a keel. Parnassius tianschanicus Oberthtir (Pisettigs 21,22) Parnassius corybas var. tianschanicus Oberthiir, 1879 : 108. Parnassius tianschanicus Oberthiir; Bryk, 1935 : 288. Parnassius tianschanicus Oberthiir; Eisner, 1966 : 184. DISTRIBUTION. U.S.S.R.: Uzbekistan, Tadzhikistan, Kirghizia. Afghanistan. Pakistan. Kashmir. China: Sinkiang. Parnassius nomion Fischer de Waldheim (PL-3; figs 17, 18) Parnassius nomion Fischer de Waldheim, 1823 : 242, pl. 6, figs 3, 4. Papilio apollo var. nomion; Geyer, [1838] : pl. 207, fig. 1029. Parnassius nomion Hiibner; Bryk, 1935 : 300. Parnassius nomion Hiibner; Eisner, 1966 : 149. DISTRIBUTION. Mongolia. U.S.S.R.: Russia (Irkutsk, Buryat, Amur, Khabarovsk, Altay). China: Kansu, Tsinghai, Shensi, Heilungkiang, Liaoning. Korea. U.S.A.: Alaska? GENERA AND SPECIES OF PARNASSIINAE 83 LARVAL FOOD PLANTS: Yellow-flowered Sedum (Elwes, 1886). S. album (Bryk, 1935). Red basal spot sometimes absent causing specimens to resemble afollo Linnaeus. Subsp. epaphoides Bryk & Eisner [Kansu, Richthofen Mts.]. Much smaller than the typical form. Parnassius apollo (Linnaeus) (Pl. 3, figs 23, 24, Pl. 15, fig. 98) Papilio apollo Linnaeus, 1758 : 465. Parnassius apollo (Linnaeus); Latreille, 1804 : 199. Parnassius apollo (Linnaeus); Bryk, 1935 : 325. Parnassius apollo (Linnaeus) ; Eisner, 1966 : 89. DISTRIBUTION. Sweden. Finland. Poland. Germany. France. Spain. Switzerland. Austria. Czechoslovakia. Hungary. Rumania. Italy. Bulgaria. Greece. Albania. Turkey. Syria. U.S.S.R.: Latvia, Lithuania, Ukraine, Armenia, Caucasus, Russia (Orel, Ural Mountains, Omsk, Altay, Tuva). China: Sinkiang. Mongolia. LARVAL FOOD PLANTS: Stonecrop; Sedum, especially S. album, S. telephium, S. purpurascens, and Sempervivum (Higgins & Riley, 1970). Sedum acre (Rebel & Rogenhofer, 1893). S. annuum, S. villosum and S. roseum (Eisner, 1958). S. maximum (Holik, 1937). Subsp. geminus Stichel (Switzerland: Grindelwald). Faintly yellowish, red spots smaller. Subsp. bartholomaeus Stichel [Germany: Ké6nigsee]. Small, male heavily marked above. Subsp. rhodopensis Markovic [Bulgaria: Rila Planina, Rhodopi Planina]. Very large. Subsp. nevadensis Oberthiir [Spain: Sierra Nevada]. Male above with yellow ocelli. Subsp. sicilae Oberthiir [Sicily]. Very small (Higgins & Riley, 1970). THE MNEMOSY NE-Grovup Parnassius mnemosyne (Linnaeus) (Pl. 4, figs 25, 26, Pl. 15, fig. 99, Text-fig. 32) Papilio mnemosyne Linnaeus, 1758 : 465. Parnassius mnemosyne (Linnaeus); Bryk, 1935 : 19. Parnassius mnemosyne (Linnaeus) ; Eisner, 1966 : 142. DISTRIBUTION. Finland. Sweden. Denmark (Baltic Islands). Germany. France. Spain. Switzerland. Italy. Austria. Poland. Czechoslovakia. Hungary. Yugoslavia. Rumania. Bulgaria. Albania. Greece. Turkey. Syria. Lebanon. Iraq. Iran. Afghanistan. U.S.S.R.: Estonia, Latvia, Lithuania, Ukraine, Uzbekistan, Tadzhikistan, Kirghizia, Russia (Mordov, Ural Mountains). 84 P. RR. ACKERY LARVAL FOOD PLANTS. Corydalis (Higgins & Riley, 1970). C. cava and C. hellert (Bryk, 1935). C. solida (Rebel & Rogenhofer, 1893). C. parnanica (Kolar, 1937). C. intermedia (Eisner, 1958). Parnassius stubbendorfii Ménétriés (Pl. 4, figs 27, 28) Parnassius stubbendorfii Ménétriés, 1849 : 273, pl. 6, fig. 2. Parnassius stubbendorfi Ménétriés; Bryk, 1935 : 107. Parnassius stubbendorfi Ménétriés; Eisner, 1966 : 179. DISTRIBUTION. U.S.S.R.: Russia (Altay, Tuva, Buryat, Chita, Amur, Khabarovsk, Primorye, Sakhalin, Kurile Islands). Mongolia. Tibet. Korea. China: Heilungkiang, Kansu, Szechwan, Tsinghai. Japan: Hokkaido. LARVAL FOOD PLANTS. Corydalis ambigua, C. gigantaea (Kurentsov, 1970). Aristolochia debilis (Lee, 1958). Parnassius glacialis Butler (Pl. 4, figs 29, 30, Text-fig. 6) Parnassius glacialis Butler, 1866 : 50. Parnassius stubbendorfi glacialis Butler; Bryk, 1935 : 128. Parnassius glacialis Butler; Eisner, 1966 : 117. Parnassius glacialis Butler; Ackery, 1973 : 7. DISTRIBUTION. Japan: Hokkaido, Honshu, Shikoku. Korea. China: Hupeh, Shangtung, Kiangsu, Anhwei, Chekiang. LARVAL FOOD PLANTS. Corydalis incisa, C. ambigua, C. decumbens and C. remota (Suguru Igarashi, in litt). Aristolochia debilis (Lee, 1958). Parnassius eversmanni Ménétriés (Pl. 4, figs 31, 32, Pl. 5, figs 33, 34) Parnassius eversmanni Ménétriés, 1855 : 73, pl. 1, fig. 2. Parnassius eversmanni Ménétriés; Hemming, 1934 : 199. Parnassius eversmanni Ménétriés; Bryk, 1935 : 133. Parnassius eversmanni Ménétriés; Eisner, 1966 : 108. DISTRIBUTION. U.S.S.R.: Russia (Irkutsk, Buryat, Primorye, Khabarovsk, Tuva, Chita, Yakut, Magadan, Kamchatka, Yevrey). Mongolia. Japan: Hokkaido. U.S.A.: Alaska. LARVAL FOOD PLANTS. Corydalis gigantea (Fumariaceae) (Wilson, 10961). Dicentra peregrina (Suguru Igarashi, in litt.). Subsp. felderi Bremer [U.S.S.R.: Russia (Amur, Khabarovsk, Yevrey)]. GENERA AND SPECIES OF PARNASSIINAE 85 Yellow pigment absent in male. Postdiscal and costal spots often black. Superficially similar to glacialis Butler. Parnassius nordmanni (Nordmann) (Pl. 5, figs 35, 36) Doritis nordmanni Nordmann, 1851 : 423, pl. 13, figs 1-3. Parnassius novdmanni (Nordmann); Hemming, 1934 : 198. Parnassius nordmanni Ménétriés; Bryk, 1935 : 146. Parnassius nordmanni Ménétriés; Eisner, 1966 : 149. DISTRIBUTION. U.S.S.R.: Caucasus. Parnassius ariadne Lederer (Pl. 5, figs 37, 38) Doritis clarius Eversmann, 1843 : 539, pl. 9, figs 1A, B, C (nec clarius Hiibner, [1806] : 61, nota 6). Parnassius ariadne Lederer, 1853 : 354. Parnassius aviadne Lederer; Hemming, 1934 : 198. Parnassius clarius (Eversmann) ; Bryk, 1935 : 151. Parnassius clarius (Eversmann); Eisner, 1966 : 98. DISTRIBUTION. U.S.S.R.: Russia (Altay), Tadzhikistan. Western Mongolia. Parnassius clodius Ménétriés (Pl. 5, figs 39, 40) Parnassius clodius Ménétriés, 1855 : 7. Parnassius clodius Ménétriés; Bryk, 1935 : 156. Parnassius clodius Ménétriés; Eisner, 1966 : 98. DISTRIBUTION. U.S.A.: Alaska, Washington, Idaho, Montana, Wyoming, Oregon, Nevada. Canada: British Columbia. LARVAL FOOD PLANTS. Viola, Sedum (Stonecrop), Vaccinium?, Rubus? (Wilson, 1961). Saxifraga sp., Vitis californica (Tietz, 1972). Parnassius orleans Oberthiir (Pl. 6, figs 41, 42, Text-fig. 27) Parnassius orleans Oberthiir, 1890 : 1. Parnassius orleans Oberthiir, 1891 : 8, 18, pl. 1, fig. 2. Parnassius orleans Oberthiir; Bryk, 1935 : 163. Parnassius orleans Oberthiir; Eisner, 1966 : 154. DISTRIBUTION. Tibet. Mongolia. China: Sinkiang, Tsinghai, Kansu, Shensi, Szechwan, Yunnan. 86 Pe ReoeACKERY THE HARDWICKII-GrRovup Parnassius hardwickii Gray (Pl. 6, figs 43, 44, Text-fig. 4) Parnassius hardwickii Gray, 1831 : 32. Lingamius hardwickei (Gray); Bryk, 1935 : 541. Lingamius hardwickei (Gray); Eisner, 1966 : 120. DISTRIBUTION. Kashmir. N. India. Nepal. Sikkim. Bhutan. LARVAL FOOD PLANTS. Various species of Saxifrage (Moore, 1902). Superficially similar to orleans Oberthiir but normally distinguishable by the white-centred, blue, submarginal series of spots on the hindwing upperside. THE SZECHENYII-Group Parnassius szechenyii Frivaldszky (Pl. 6, figs 45, 46, Pl. 15, fig. 100, Text-fig. 5) Parnassius szechenyii Frivaldszky, 1886 : 39, pl. 4, figs 1, Ia. Kovamius szechenyit (Frivaldszky) ; Bryk, 1935 : 550. Koramius szechenyi (Frivaldszky) ; Eisner, 1966 : 181. DISTRIBUTION. Tibet. China: Tsinghai, Kansu, Szechwan, Yunnan. Parnassius cephalus Grum-Grshimailo (Pl. 6, figs 47, 48, Pl. 7, figs 49, 50, Text-fig. 22, 30) Parnassius cephalus Grum-Grshimailo, 1891 : 446. Koramius cephalus (Grum-Grshimailo) ; Bryk, 1935 : 558. Koramius cephalus (Grum-Grshimailo) ; Eisner, 1966 : 97. DISTRIBUTION. Tibet. China: Kansu, Szechwan, Tsinghai. Kashmir. Subsp. maharaja Avinoff [Kashmir: Ladakh Range]. Markings generally reduced. Submarginal spots above faint, not centred with blue. Postdiscal and costal spots absent from hindwing upperside. This subspecies (PI. 7, figs 49, 50), treated by both Bryk (1935) and Munroe (1960) as a species, differs from typical cephalus in having veins R, and R, consistently anastomosing, causing it to key out as P. acco Gray. THE ACCO-GROUP Parnassius acco Gray (El) NES SE, 52, 55054) Parnassius acco Gray, 1853 : 76, pl. 12, figs 5, 6. Tadumia acco (Gray); Bryk, 1935 : 631. Tadumia acco (Gray); Eisner, 1966 : 82. GENERA AND SPECIES OF PARNASSIINAE 87 DISTRIBUTION. Kashmir. Tibet. Sikkim. Subsp. liliput Bryk [Tibet: Everest District], subsp. hunningtoni Avinoff (Tibet: Dochar, Tuna, Tsangpo Valley, Dzara, Kyetrak, Chumbi Valley. Sikkim: Gangtok]. Smaller than the typical form, red markings absent. THE DEL PHIUS-Group Parnassius patricius Niepelt (Pl. 7, figs 55, 56) Parnassius patricius Niepelt, 1911 : 274. Koramius patricius (Niepelt); Bryk, 1935 : 568. Koramius patricius (Niepelt); Eisner, 1966 : 155. DISTRIBUTION. U.S.S.R.: Kirghizia. Parnassius acdestis Grum-Grshimailo (PI. 8, figs 61, 62, Pl. 15); fig. rox, Text-fig. 7) Parnassius delphius var. acdestis Grum-Grshimailo, 1891 : 446. Kovramius acdestis (Grum-Grshimailo); Bryk, 1935 : 572. Koramius acdestis (Grum-Grshimailo) ; Eisner, 1966 : 82. DISTRIBUTION. U.S.S.R.: Kirghizia. Kashmir. Tibet. Sikkim. Bhutan. China: Sinkiang, Szechwan. The arrangement of the forewing radial veins is more variable in this species than in any other. Although veins R, and R, are usually separate, in many specimens they do appear to touch and in some cases quite definitely anastomose. Subsp. lucifer Bryk [Sikkim: Gyamtshona]. Postdiscal and costal spots black in hindwing above. Subsp. lux Eisner [Tibet: Jung-jung Khola]. Basal black scaling in hindwing above far less extensive than in the typical form. Postdiscal and costal spots large. Parnassius delphius (Eversmann) (Pl. 8, figs 57, 58, 59, 60, Pl. 15, fig. 102) Doritis delphius Eversmann, 1843 : 541, pl. 7, figs 1a, b. Parnassius delphius (Eversmann) ; Elwes, 1886 : 39. Koramius delphius (Eversmann); Bryk, 1935 : 583. Koramius delphius (Eversmann) ; Eisner, 1966 : 102. DistripuTion. Afghanistan. U.S.S.R.: Tadzhikistan, Kirghizia, Uzbekistan. Pakistan. Kashmir. N. India. Tibet. China: Sinkiang, Tsinghai. Highly variable species. Hindwing discal spots often without red scales. 88 Pik. ACKERY, Subsp. pulchra Eisner | Kirghizia: Kungey Alatau Mountains]. Wings exception- ally dark, semi-transparent. Although treated by Bryk (1935) and Munroe (1960) as a distinct species, Parnassius stoliczkanus is here regarded, in accordance with Eisner (1966), as a subspecies of Parnassius delphius (Eversmann). The following subspecies are those that Bryk and Munroe would have included in Parnassius stoliczkanus Felder & Felder. Subsp. atkinsoni Moore [Kashmir: Pir Pinjal, Sind Valley, Burzil Pass. India: Himachal Pradesh, Kulu], subsp. beate Eisner [Kashmir: Karakoram, Potu-la Pass, Chalsi, Leh], subsp. chitralica Verity [Pakistan: Chitral], subsp. florenciae Tytler [Tibet: Phupes Hundes, Tibu, Churmurti], subsp. gracilis Bryk & Eisner [India: Himachal Pradesh, Kangra, Rohtang Pass], subsp. imitator Bryk & Eisner [U.S.S.R.: Tadzhikistan, Pamirs, Beik Pass], subsp. kumaonensis Riley [India. Uttar Pradesh, Kumaon, Shillung], subsp. nicevillei Avinoff [Kashmir: Pir Pinjal, Burzil Pass, SariSungur Pass, Sapta La], subsp. parangensis Eisner (India: Himachal Pradesh, Parang Pass, Bara Lacha Pass. Kashmir: Tagalang Pass, Lingti, Ladahk], subsp. rileyi Tytler [Kashmir: Rupal Valley, Astor], subsp. spitiensis Bang-Haas [Tibet: Spiti, Tum-Tum-Thang, Churmurti], subsp. stoliczkanus Felder & Felder (Kashmir: Ladak, Rupshu, Sapta La], subsp. tenuis Bryk & Eisner [Kashmir: Gya-Ladahk, Tagalang Pass], subsp. tytlerianus Bryk & Eisner [Kashmir: Chitral, Bangol Pass], subsp. zanskarica Bang-Haas [Kashmir: Nira, Zanskar Mts.], subsp. zogilaica Tytler [Kashmir: Zogila]. Generally smaller than the typical delphius. Red costal spot of hindwing upperside usually absent. THE TIM PE RA TOR-Grovup Parnassius imperator Oberthiir (Pl. 8, figs 63, 64, Pl. 15, fig. 103, Text-fig. 21) Parnassius imperatoy Oberthiir, 1883 : 77. Tadumia imperator (Oberthiir); Bryk, 1935 : 675. Eukoramius imperator (Oberthiir) ; Eisner, 1966 : 123. DISTRIBUTION. Tibet. China: Tsinghai, Kansu, Szechwan, Yunnan. LARVAL FOOD PLANT. Corydalis (Verity, 1907). THE CHA RLTONIUS-GrRovup Parnassius charltonius Gray (Pl. 9, figs 65, 66, Pl. 15, fig. 104, Text-fig. 24) Parnassius charltonius Gray, 1853 : 77, pl. 12, fig. 7. Koramius charltonius (Gray); Bryk, 1935 : 694. Koramius charitonius (Gray); Eisner, 1966 : 97. DISTRIBUTION. Afghanistan. U.S.S.R.: Kirghizia, Tadzhikistan. Pakistan. Kashmir. N. India. Tibet. LARVAL FOOD PLANT. Corydalis gortschakovi (A. Tsvetajev, pers. com.) GENERA AND SPECIES OF PARNASSIINAE 89 Parnassius inopinatus Kotzsch (Pl. 9, figs 67, 68, Pl. 15, fig. 105) Parnassius inopinatus Kotzsch, 1940 : 17. Kailasius inopinatus (Kotzsch); Eisner, 1966 : 123. DISTRIBUTION. Afghanistan: Firus-Kuhi Range, Koh-i-Baba Range. Parnassius loxias Piingeler (Pl. 9, figs 69, 70) Parnassius loxias Piingeler, 1901 : 178, pl. 1, figs 5, 6. Koramius loxias (Piingeler); Bryk, 1935 : 717. Eukoramius loxias (Piingeler); Eisner, 1966 : 132. DISTRIBUTION. U.S.S.R.: Kirghizia. China: Sinkiang. Parnassius autocrator Avinoff (Pl. 9, figs 71, 72, Text-fig. 28) Parnassius charltonius autocrator Avinoff, 1913 : 16, pl. 2, fig. 2. Kovramius charltonius autocratoy (Avinoff); Bryk, 1935 : 716. Eukoramius autocratoy (Avinoff); Eisner, 1966 : 91. DISTRIBUTION. Afghanistan. U.S.S.R.: Tadzhikistan. LARVAL FOOD PLANT. Corydalis adiantifolia (Wyatt & Omoto, 1963). THe TENE DIUS-GrRovup Parnassius tenedius Eversmann (Pl. 10, figs 75, 76, Text-fig. 3) Parnassius tenedius Eversmann, 1851 : 621. Tadumia tenedius (Eversmann); Bryk, 1935 : 647. Tadumia tenedius (Eversmann); Eisner, 1966 : 181. DISTRIBUTION. Mongolia. U.S.S.R.: Russia (Yakut, Tuva, Chita). China: Inner Mongolia. LARVAL FOOD PLANT. Corydalis sp. (bracteata?)( A. Tsvetajev, pers. com.). THe SITMO-GrRouP Parnassius simo Gray (Pl. 10, figs 73, 74, Text-fig. 8) Parnassius simo Gray, 1853 : 76. Tadumia simo (Gray); Bryk, 1935 : 654. Tadumia simo (Gray); Eisner, 1966 ; 178. BF go Pp. Rk. ACKER Y DISTRIBUTION. U.S.S.R.: Kirghizia, Tadzhikistan. Kashmir. N. India. Mongolia. Tibet. China: Sinkiang, Kansu. Tribe ZERYNTHIINI Zerynthianae Grote, 1899: 17. Type-genus: Zerynthia Ochsenheimer. SERICINUS Westwood Sericinus Westwood, 1851 :173. Type-species: Papilio telamon Donovan, by original designation. Sericinus Westwood; Bryk, 1934 : 77. Sericinus Westwood; Munroe, 1960 : 13. Sericinus montela Gray (Pl. xo, figs 77, 78, Text-figs 16, 26) Papilio telamon Donovan, 1798 : pl. 27, fig. 1. [Junior homonym of Papilio telamon Linnaeus, 1758 : 486.] Sericinus montela Gray, 1853 : 78, pl. 13, figs 1, 2. Sericinus telamon (Donovan); Bryk, 1934 : 80. Sericinus telamon montela Gray; Bryk, 1934 : 89. Sericinus montela Gray; Eisner, 1966 : 142. Sericinus telamon (Donovan); Eisner, 1966 : 181. Sericinus montela Gray; Hemming, 1967 : 409. DISTRIBUTION. China: Heilungkiang, Kirin, Liaoning, Hopei, Shangtung, Anhwei, Kiangsu, Hunan, Hupeh, Kiangsi, Kansu. Korea. LARVAL FOOD PLANTS. Aristolochia (Leech, 1893 : 488). A. contorta (Kurentsov, 1970). PARNALIUS Rafinesque Thais Fabricius, 1807 : 283. Type-species: Papilio hypsipyle Fabricius, bymonotypy. [Junior homonym of Thais Réding, 1789.] Parnalius Rafinesque, 1815 : 128. [Replacement name for Thais Fabricius.] Zerynthia Ochsenheimer, 1816 : 29. [Replacement name for Thais Fabricius.] Eugraphis Billberg, 1820: 75. Type-species: Papilio hypsipyle Fabricius, by monotypy. Parnalius Rafinesque; Sherborn, 1929 : 4765. Zerynthia Ochsenheimer; Sherborn, 1932 : 7041. Allancastria Bryk, 1934 : 19, 61-62. Type-species: Thats cerisy Godart, by original designation. Syn. n. Zerynthia Ochsenheimer; Bryk, 1934 : 31. Parnalius Rafinesque; Neave, 1940a : 614. Zerynthia Ochsenheimer; Neave, 19406 : 689. Allancastria Bryk; Munroe, 1960 : Io. Zerynthia Ochsenheimer; Munroe, 1960 : 13. Zerynthia Ochsenheimer; Hemming, 1967 : 464. Parnalius Rafinesque; Cowan, 1970: 11. [Zerynthia Ochsenheimer cited as synonym.] GENERA AND SPECIES OF PARNASSIINAE gI According to Cowan (1970) Rafinesque introduced the name Parnalius for Thais Fabricius, which was invalid as a junior homonym. The name is available and valid, and is a senior objective synonym of Zerynthia Ochsenheimer, 1816. It has been correctly listed by both Sherborn (1929) and Neave (1940). The genus Allancastria Bryk is here treated as a synonym of Parnalius Rafinesque. The differences in venation, as figured by Bryk (1934), do not appear to be consistent although the genitalia are certainly distinct. If Allancastria Bryk is to be recognized as a valid genus it would seem to me that there is equal justification for raising the status of the species groups of Parnassius to genera. In order to maintain con- sistency in approach I am regarding Allancastria Bryk and Parnalius Rafinesque as being subjectively synonymous. KEY TO THE SPECIES OF PARNALIUS RAFINESQUE I Cell 2A of forewing underside uniformly scaled, without distinct red or black discal spot; uncus long, bifid; clasper narrow (Text-fig. 11, Pl. 10, figs 79, 80) cerisy (Godart) (p. 91) - Cell 2A of forewing underside with a distinct red or black discal spot; uncus short, bifid; clasper broad (Text-figs 9, 10) : 2 2 (1) Forewing upperside usually with distinct red spots in the discal cell and with a vitreous spot near the wing apex; uncus in dorsal aspect narrowing towards the base; clasper distinctly produced dorso-posteriorly (Text-fig. 10, Pl. 11, figs 83, 84) ; ; rumina (Linnaeus) (p. 92) Forewing upperside usually with black spots in the discal cell, seldom red, and vitreous spot absent from the wing apex; uncus in dorsal aspect narrowing towards the apex; clasper slightly produced dorso-posteriorly (Text-fig. 9, Pl. 11, figs 81, 82) : - . polyxena (Denis & Schiffermiiller) (p. 91) Parnalius cerisy (Godart) comb. n. (Pl. ro, figs 79, 80, Text-figs 11, 17) Thais cerisy Godart, [1824] : 812. Zerynthia cerisyi (Godart); Stichel, 19074 : 82. Allancastria cerisyi (Godart); Bryk, 1934 : 63. Allancastria cerisy (Godart) ; Cowan, 1970 : 17, 41. DISTRIBUTION. Cyprus. Crete. Greece. Yugoslavia. Bulgaria. Rumania. Albania. Turkey. U.S.S.R.: Armenia. Iran. Iraq. Syria. Israel. Lebanon. LARVAL FOOD PLANTS: Aristolochia clematis and A. hastata (Bryk, 1934). A. maurorum (Suguru Igarashi, in litt.). Parnalius polyxena (Denis & Schiffermiller) comb. n. (Pl. 11, figs 81, 82, Text-figs 9, 19) Papilio hypermnestra Scopoli, 1763 : 149, pl. [17], fig. 425. [Junior homonym of Papilio hypermnestra Linnaeus, 1763 : 40.] Papilio polyxena Denis & Schiffermiiller, 1775 : 162 92 Po RACK ERY: Papilio hypsipyle Fabricius, 1777 : 265. Zerynthia hypermnestra (Scopoli); Bryk, 1934 : 34. Zerynthia hypermnestra (Scopoli); Eisner, 1966 : 123. Zerynthia polyxena (Denis & Schiffermiiller); Hemming, 1967 : 436. Hemming (1967) gives a detailed explanation of the nomenclatorial history of this species and how folyxena was finally established as the valid name. DisTRIBUTION. S. France. Austria. Italy. Sicily. Yugoslavia. Hungary. Rumania. Albania. Greece. Czechoslovakia. LARVAL FOOD PLANTS. Aristolochia pistolochia. A. rotunda and A. clematis (Higgins & Riley, 1970). A. sicula (Bryk, 1934). Parnalius rumina (Linnaeus) comb. n. (Pl. 11, figs 83, 84, Text-fig. 10) Papilio rumina Linnaeus, 1758 : 480. Thais maturna Butler, 1870 : 232. Zerynthia rumina (Linnaeus); Bryk, 1934 : 50. DISTRIBUTION. S. France. Spain. Portugal. Algeria. Morocco. Tunisia. LARVAL FOOD PLANTS. Various kinds of Avistolochia (Higgins & Riley, 1970). A. pistolochia and A. fontanesi (Bryk, 1934). LUEHDORFIA Criiger Luehdorfia Criiger, 1878 : 128. Type-species: Luehdorfia eximia Criiger, by monotypy. Luehdorfia Criiger; Bryk, 1934 : 99. Luehdorfia Criiger; Munroe, 1960 : 13. KEY TO THE SPECIES OF LUEHDORFIA CRUGER 1 Claspers of male covered with thick golden brown pubescence; ventral spinose setae, placed internally on claspers, elongate (Text-fig. 12); sphragis of female keeled (Pl. 11, figs 85, 86) . : . puziloi (Erschoff) (p. 92) — Claspers of male covered with thick black pubescence; ventral setae, placed internally on claspers, shorter (Text-fig. 13); Ppa of female without keel (Pl. 11, figs 87, 88, 89,90) . : : : F . japonica Leech (p. 93) Luehdorfia puziloi (Erschoff) (Pl. 11, figs 85, 86, Text-fig. 12) Thais puziloi Erschoff, 1872 : 315. Luehdorfia eximia Criiger, 1878 : 128. Luehdorfia puziloit (Erschoff); Bryk, 1934 : 102 DISTRIBUTION. U.S.S.R.: Russia (Primorye). Korea. Japan: Hokkaido, Honshu. LARVAL FOOD PLANTS. Asarum (Graeser, 1888). A. sieboldi (Kurentsov, 1970). GENERA AND SPECIES OF PARNASSIINAE 93 Luehdorfia japonica Leech (Pl. 11, figs 87, 88, 89, 90, Text-figs 13, 23) Luehdorfia japonica Leech, 1889 : 25, pl. 1, figs 1, rb, 1c. Luehdorfia japonica japonica Leech; Bryk, 1934 : 102. DISTRIBUTION. Japan: Honshu. Taiwan. China: Liaoning, Kirin, Hupeh, Anhwei, Kiangsu, Kiangsi. LARVAL FOOD PLANTS. Asarum nipponicum, A. tamaense, A. blumet, A. caulescens and A. sieboldi (Suguru Igarashi, in litt.). Subsp. chinensis Leech (China: Hupeh, Anhwei, Kiangsu, Kiangsi). Hindwing upperside with red submarginal band. Internervular marginal spots of hindwing yellow. The latter taxon, variously treated by authors, was regarded by Bryk (1934) as a subspecies of puzilot Erschoff. The male genitalia, however, show a close resemblance to those of japonica Leech both in the shape of the claspers and in the length of the setae placed internally thereon. These characters, together with the unkeeled sphragis of the female, seem to indicate that this taxon was correctly placed by Rothschild (1918) as a subspecies of japonica Leech, and it is here so treated. BHUTANITIS Atkinson Armandia Blanchard, 1871 : 809, nota 3. Type-species: Armandia thaidina Blanchard, by monotypy. [Homonym of Armandia Filippi, 1862.] Bhutanitis Atkinson, 1873 : 570. Type-species: Bhutanitis hdderdalii Atkinson, by monotypy. Bhutanitis Atkinson; Bryk, 1934 : 113. Bhutanitis Atkinson; Munroe, 1960 : 13. KEY TO THE SPECIES OF BHUTANITIS ATKINSON I Hindwing upperside with a series of orange marginal internervular markings . 2 - Hindwing upperside without orange marginal internervular markings, being yellow or grey in these areas 2 (1) Vein M, of hindwing produced to broad spatulate tail; ‘clasper of male bluntly produced posteriorly and bearing a tuft of thick black pubescence (Text-fig. 14, Pl. 12, figs 92, 93) : . thaidina (Blanchard) (p. 93) - Vein M, of hindwing produced to a narrow tail; clasper of male pointed and bearing sparse pubescence only (Text-fig. 15, Pl. 13, figs 94, 95) lidderdalii Boisduval (p. 94) 3 (1) Vein Cu, of hindwing produced to a round lobe; pale bands of wings broad, resembling Luehdorfia; female bearing a sphragis (Pl. 12, fig. 91) mansfieldi (Riley) (p. 94) - Vein Cu,, of hindwing produced to a distinct tail; pale bands of wings narrow; female without sphragis (Pl. 14, figs 96, 97) , . ludlowi Gabriel (p. 94) Bhutanitis thaidina (Blanchard) (Pl. 12, figs 92, 93, Text-figs 14, 25) Armandia thaidina Blanchard, 1871 : 809. Bhutanitis thaidina (Blanchard) ; Bryk, 1934 : 116. 94 P. R. ACKERY DISTRIBUTION. China: Shensi, Szechwan, Yunnan. LARVAL FOOD PLANT. Aristolochia sp. (Bryk, 1934). Bhutanitis lidderdalii Atkinson (Pl. 13, figs 94, 95, Text-fig. 15) Bhutanitis lidderdalii Atkinson, 1873 : 570, pl. 50. Bhutanitis lidderdalii Atkinson; Bryk, 1934 : 118. DISTRIBUTION. Bhutan. Sikkim. N. India: Assam, Nagaland, Manipur. N. Burma. China: Szechwan, Yunnan. ; Bhutanitis ludlowi Gabriel (Pl. 14, figs 96, 97) Bhutanitis ludlowi Gabriel, 1942 : 189. DISTRIBUTION. Bhutan: Trashiyangsi Valley. As far as I am aware, the type-series of /udlowi Gabriel is unique, no other representatives of this species being known to me. Bhutanitis mansfieldi (Riley) (Pl 2. te. Or) Armandia mansfieldi Riley, 1939a : 207, pl. 4. Bhutanitis mansfieldi (Riley); Riley, 1939) : 267. DISTRIBUTION. China: Yunnan. This species, known to me from the female holotype only, bears a curious resemblance to Luehdorfia Criiger in both pattern and wing shape; furthermore it is the only Bhutanitis species in which the female bears a sphragis. When more material becomes available, examination of the male genitalia may show whether it has been correctly placed here. REFERENCES AckERY, P. R. 1973. A list of the type-specimens of Parnassius (Lepidoptera: Papilionidae) in the British Museum (Natural History). Bull. Br. Mus. nat. Hist. (Ent.) 29 : 1-35, 1 pl. ATKINSON, W. S. 1873. Description of a new genus and species of Papilionidae from the south-eastern Himalayas. Proc. zool. Soc. Lond. 1873 : 570-572, I pl. AVINOFF, A. 1913. Quelques formes nouvelles du genre Parnassius Latr. Trudy russk. ent. Obshch. 40 (5) : 1-21, 1 pl. BILLBERG, G. J. 1820. Enumeratio insectorum in museo Gust. Joh. Billberg. ii + 138 pp. Stockholm. BLANCHARD, E. 1871. Remarques sur la faune de la principauté thibétaine du Moupin. C. r. hebd. Séanc. Acad. Sci., Paris 72 : 807-813. GENERA AND SPECIES OF PARNASSIINAE 95 Boispuval, J.B. A.D. 1836. Spécies général des Lépidoptéves 1: vii + 690 pp., 24 pls. Paris. BrREMER, O. 1864. Lepidopteren Ost-Sibiriens insbesondere des Amur-Landes. Mém. Acad. imp. Sci. St. Peterbs. 1-103, 8 pls. Bryk, F. 1922. Zur Prasizierung Puppenhabitus einiger Parnassiiden. Ent. Tidskr. 42 : 224-226. 1934. Lepidoptera, Baroniidae, Teinopalpidae, Parnassiidae. Pars I. Tierreich 64 : xxiii + 131 pp., 87 figs. 1935. Lepidoptera, Parnasstidae. pars II. (Subfam. Parnassiinae). Tierreich 65 : li + 790 pp., 698 figs. Butter, A. G. 1866. A list of the diurnal lepidoptera recently collected by Mr Whitely in Hakodadi (North Japan). J. Linn. Soc. 9 : 50-51. 1870. Catalogue of diurnal lepidoptera described by Fabricius in the collection of the British Museum. 303 pp., 2 pls. London. CiencH, H. K. 1955. Revised classification of the butterfly family Lycaenidae and its allies, Ann. Carneg. Mus. 33 : 261-274. Cowan, C. F. 1970. Annotationes Rhopalocerologicae 1970. 70 pp. Berkhamsted. CRAMER, P. 1775. De Uitlandsche Kapellen voorkomende in de drie Waereld-Deelen Asia, Africa en America. 1 : 1-132, 84 pls. Amsterdam & Utrecht. CRUGER, C. 1878. Ueber Schmetterlinge von Wladiwostok. Verh. Ver. naturw. Unterh. Hamb. 3 : 128-133. Daman, J. W. 1816. Forsok till systematik Uppstallning af Sveriges Fjarillar. K. svenska VetenskAkad. Handl. 37 : 48-101. Denis, J. N. C. M. & ScHIFFERMULLER, I. 1775. Ankiindung eines systematischen Werkes von den Schmetterlingen der Wienergegend.: 323 pp. Vienna. Donovan, E. 1798. An epitome of the natural history of the insects of China: 50 pls. London. EuRLICH, P. R. 1958. The comparative morphology, phylogeny and higher classification of the butterflies (Lepidoptera: Papilionoidea). Kans. Univ. Sci. Bull. 39 : 305-370. EISNER, C. 1958. Parnassiana Nova XVII. Varia. Zodl. Meded., Leiden 36 : 1-3. 1966. Parnassiidae-Typen in der Sammlung J.C. Eisner. Zodl. Verh., Leiden 81 : 1-190, 84 pls. 1968. Parnassiana Nova XLIII. Nachtragliche Betrachtungen zu der Revision der Subfamilie Parnassiinae (Fortsetzung 16). Zodl. Meded., Leiden 43 : 9-17, 2 pls. EtweEs,H.J. 1886. On butterflies of the genus Parnassius. Proc. zool. Soc. Lond. 1886 : 6-53, 4 pls. Erscuorr, N. 1872. Diagnoses de quelques espéces nouvelles de Lépidoptéres appartenant a la faune de la Russie Asiatique. Trudy russk. ent. Obshch. 8 : 315-318. Evans, W. H. 1927. Identification of Indian butterflies. xviii + 302 pp., 32 pls. Madras. EVERSMANN, E. 1843. Quaedam lepidoptorum species novae, in montibus Uralensibus et Altaicus habitantes, nunc descritae et depicta. Bull. Soc. Nat. Moscou 16 : 535-555, 4 Pls. —— 1847. Lepidoptera quaedam nova Rossiae et Sibiriae indigena descripsit et delineavit. Bull. Soc. Nat. Moscou 20 (2) : 66-83, 6 pls. 1851. Description de quelques nouvelles espéces de lépidoptéres de la Russie. Bull. Soc. Nat. Moscou 24 (1) : 610-644. Fasricius, J. C. 1777. Genera Insectorum: xii + 310 pp. Chilonii. 1793. Entomologia Systematica 3 (1) : vi + 488 pp. Hafniae. —— 1807. Die neueste Gattungs-Eintheilung der Schmetterlinge aus den Linneischen Gattung Papilio und Sphinx. Magazin Insektenk. (Illiger) 6 : 277-295. FELDER, C. & FELDER, R. 1865. Reise dey Osterreichischen Fregatte Novara; Lepidoptera, Rhopalocera (published 1865-1867), vi + 548 pp., 140 pls. Vienna. FIisHER, C. 1950. Parnassius Latreille mnemosyne Linne. Bull. Soc. ent. Mulhouse 1950 54-58, 63-66, 69-73, 2 pls. FISCHER DE WALDHEIM, G. 1823. Entomographie de la Russie; Lepidoptera: 242-264, 6 pls. Moscow. 96 Ps R. ACKERY. Forp, E. B. 1944. Studies on the chemistry of pigments in the lepidoptera, with reference to their bearing on systematics. 4: The classification of the Papilionidae. Trans. R. ent. Soc. Lond. 94 : 201-223. FRIVALDSZKY, J. 1886. Lepidoptera Nova et Varietates in Expeditione ad Oras Asie Orientalis Comitis Belae Szechenyi, a Domonins Gustava Kreitner et Ludovico Loczy Collecta et a Joanne Frivaldszky Descripta. Termezetr. Fiiz. 10 : 39-40, 1 pl. GABRIEL, A. G. 1942. A new species of Bhutanitis (Lep. Papilionidae). Entomologist 75 : 189. GEYER, C, [1838] Jn Hiibner J., Sammlung Europdischer Schmetterlinge (Papiliones), (pub- lished [September 1837-1838]) pls 204-207. Augsburg, GopartT, J. P. [1824]. In Latreille & Godart, Encyclopédie Méthodique 9 (Supplément) : 809-828. Paris. GRAESER, L. 1888. Beitrage zur Kenntniss der Lepidopteren-Fauna des Amurlandes. Berl. ent. Z. 32 : 33-152. Gray, G. R. 1831. Descriptions of eight new species of Indian butterflies, (Papilio, Lin.) from the collection of General Hardwicke. Zoological Miscellany : 86 pp., 4 pls. London. 1853. Catalogue of the lepidopterous insects in the British Museum. 1: 84 pp., 1 fig., 12 pls. London. Grote, A. R. 1899. Specializations of the lepidopterous wing; Parnassi-Papilionidae. I. Proc. Am. phil. Soc. 38 : 7-21. GRUM-GRSHIMAILO, G. J. 1891. Lepidoptera nova in Asie Centrali novissime lecta et descripta a Gr. Grum-Grshimailo. Trudy russk. ent. Obshch. 25 : 445-465. HeEmMING, F. 1934. Revisional notes on certain species of Rhopalocera (Lepidoptera). Stylops 3 : 193-200. 1960. Annotationes Lepidopterologicae 2 : 41-47. London. 1967. The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bull. Br. Mus. nat. Hist. (Ent.), Suppl. 9 : 509 pp. Hersst, J. F. W. 1798. In Jablonsky & Herbst, Natursystem aller bekannten in und auslandischen Insekten Schmetterlinge 9 : iv + 206 pp., 30 pls. Berlin. Herinc, M. 1932. Morphologische Untersuchungen in der Gattung Parnassius (Lepidopt.) als Beitrag zu einer Kritik am Begriff der Unterart. Mitt. zool. Mus. Berl. 18 : 273-317, 150 figs. Hiaorns, L. G. & Ritty,N.D. 1970. A field guide to the butterflies of Europe 380 pp., 60 pls. London. Hoiikx, O. 1937. Ein Fadenwurm als Schmarotzer bei Parn. apollo L. Parnassiana 5 : 17-18. HUsBnER, J. [1806]. Sammlung Europdischer Schmetterlinge, Der Ziefey (Papiliones). (published [1805-1806]). iv + iv + 74 pp., pls. 115-124. Augsburg. [1819]. Verzeichniss bekannter Schmettlinge. (6) : 81-96. Augsburg. [1822]. Systematische-alphabetische Verzeichniss. vi-+ 81 pp. Augsburg. Korar, H. 1937. Parnassius mnemosyne L. der Mont Olympe. Lambillionea 37 : 98-99, I pl. Kortzscu, H. 1940. Parnassius inopinatus, eine iiberreschende neue Art. Ent. Z., Frankf. a. M. 54: 17-21. Kurentsov, A. I. 1970. The butterflies of the far east U.S.S.R. (identification key) [In Russian]. 164 pp., 14 pls. Leningrad. LATREILLE, P. A. 1804. Tableau méthodique des Insectes. Nouveau Dictionaire d’ Histoire naturelle 24 : 129-200. Paris. LEDERER, J. 1853. Lepidopterologisches aus Sibirien. Verh. zool.-bot. Ver. Wien. 3 : 351-386, 7 pis. Lee, C. L. 1958. Butterflies [In Chinese]. Dept. Ent., Academia Sinica (Peking), Publ. no. 4. 198 pp., 4 pls., 198 figs. Leecu, J. H. 1889. Description of a new Luehdorfia from Japan. Entomologist 22 : 25-26. 1893. Butterflies from China, Japan and Korea. Part Il; Lycaenidae Papilionidae and Hesperidae (published 1893-1894). 267-680, 17 pls. London. GENERA AND SPECIES OF PARNASSIINAE 97 LinnaEus, C. 1758. Systema Naturae ed. 10 : 824 pp. Holmiae. 1763. Centuria Insectorum, quam praesidae Car. von Linne, proposuit Boas Johansson. Amoenttates Academica 6 : 384-415. MENETRIES, M. 1849. Description des insectes recuillis par feu M. Lehmann. Mém. Acad. Imp. Sci. St. Petersb. (6) 8 : 217-328. 1855. Enumeratio Corporum Animalium Musei Imperialis Scientarum Petropolitanae. Classis Insectorum, Ordo Lepidopterorum. ParsI. Lepidoptera Diurna. xvi+97+[3] pp., 6 pls. St. Petersburg. Moore, F. 1902. Lepidoptera Indica 5 Rhopalocera; Nymphalidae, Riodinidae, Papilionidae. (published 1901-1903). vii + 248 pp., 85 pls. London. MULLER, A. 1973. Die mnemosyne-Gruppe der Gattung Parnassius Latreille unter Beriick- sichtigung neuer Schuppenmerkmale ihrer Arten. (Lep. Parnassiidae). Dt. ent. Z. 20 : 211-276, 5 pls. MunroE, E. 1960. The classification of the Papilionidae. Can. Ent., Suppl. 17 : 1-51. NEAVE, S.A. 1940a. A list of the names of genera and subgenera in zoology from the tenth edition of Linnaeus 1758 to the end of 1935. M-P. Nomencl. zool. 3 : 1065 pp. 1940b. A list of the names of genera and subgenera in zoology from the tenth edition of Linnaeus 1758 to the end of 1935. Q-Zand supplement. Nomencl. zool. 4 : 758 pp. NICKERL, F. A. 1846. Beschreibung einer neuen Gattung Art als Beitrag zur europdischen Lepidoptern-Fauna. Stettin. ent. Zig 7 : 207-209, I pl. NIEPELT, W. 1911. Neue palaearktische Macrolepidopteren. Int. ent. Z. 5 : 274-275. NORDMANN, A. 1851. Die im Gebiete der Fauna Taurico-Caucasica. Beobachteten Schmetterlinge. Bull. Soc. Nat. Moscou 24 (2) : 395-428. OBERTHUR, C. 1879. Catalogue raisonné des Papilionidae de la collection de Ch. Oberthiir. Etudes d’Entomologie 4 : ix + xvii, 19-115, 4 pls. Rennes. 1883. Séance du 27 Juin 1883. Bull. Soc. ent. Fr. (6) 3 : 73-79. 1890. Description d’une espéce nouvelle de lépidoptére appartenant au genre Parnassius. 3 pp. Rennes. 1891. Faunes entomologiques. Descriptions d’insectes nouveaux ou peu connus. Etudes d’Entomologie 14 : x + 19 pp., 3 pls. Rennes. OCHSENHEIMER, F. 1816. Die Schmetterlinge von Europa 4: 212 pp. Leipzig. Passos, C. F. pos 1964. A synonymic list of the Nearctic Rhopalocera. Lepidopterists’ Society Memoirs No. 1, vi + 145 pp. New Haven, Connecticut. PUNGELER,R. 1901. Neue Macrolepidopteren aus Centralasien. Dt. ent. Z. Iris 14 : 177-191 RAFINESQUE-SCHMALTZ, C.S. 1815. Analyse de la Nature 224 pp. Palermo. REBEL, H. & RoGENHOFER, A. 1893. Zur Kenntnis des Genus Parnassius Latr. in Oesterreich- Ungarn. Jber. wien. ent. Ver. 1892 : 51-70, 1 pl. RitEy, N. D. 1939a. A new species of Armandia (Lep. Papilionidae). Entomologist 72 : 207-208, 1 pl. 1939b. Arvmandia mansfieldi (Nota). Entomologist 72 : 267. ROTHSCHILD, W. 1918. Catalogue of the Parnassius in the Tring Museum. WNovit. zool. 25 : 218-262. Scopout, J. 1763. Entomologia Carniolica, Ordo III, Lepidoptera : 142-258, pls 16-36. Vindobonae, Scott, J. A. 1973. Population biology and adult behavior of the circumpolar butterfly, Parnassius phoebus F. (Papilionidae). Entomologica scand. 4 : 161-168. Scupper, S. H. 1875. Historical sketch of the generic names proposed for butterflies: A contribution to systematic nomenclature. Proc. Am. Acad. Arts Sct. : 91-293. SEItz, A. 1906. Luehdorfia in Die Gross-Schmetterlinge der Evde. Palaearktischen Fauna 2:3 25; SEOK,D.M. 1939. A synonymic list of the butterflies of Korea. xxxi + 391 pp.,2pls. Seoul. SHERBORN, C.D. 1929. Index Animalium 19 : 4691-4930. London. —— 1932. Index Animalium 28 : 6807-7056. London. 98 P. R. ACKERY SHirozu, T. & Hara, A. 1960. Early stages of Japanese butterflies in colour 1: 142 pp., 60 pls. Osaka. STAUDINGER, O. 1882. Jn Staudinger, O. & Bang-Haas, O., Ueber einige neue Parnassius- und andere Tagfalter-Arten Central Asies. Berl. ent. Z. 26 : 161-177. STICHEL, H. 1907a. Lepidopterologische Miscellen. III. Zum MHeimatsnachweis von Zerynthia cerisyi God. Ent. Z., Frankf.a. M. 21 : 82-83. 1970b. Parnassius, In Seitz, A., Die Gross-Schmetterlinge der Erde. Palaearktischen Fauna 1 : 19-36, 7 pls. STSCHETKIN, Y.L. 1963. The Lepidoptera of moderate altitudes in the mountains of southern Tadzhikistan, [In Russian.] Trudy Inst. Zool. Parazit., Stalinabad 24 : 21-73. Swarnson, W. 1840. In Swainson, W. & Shuchard, W. E. On the history and natural avrangement of insects. iv + 406 pp. London. TatBot, G. 1939. The fauna of British India, including Ceylon and Burma. Butterflies I : xxix + 591 pp., 3 pls. London. Tietz, H.M. 1972. An index to the described life histories, early stages and hosts of the macro- lepidoptera of the continental United States and Canada. 2 vols. vi-+ 1041 pp. Sarasota, Florida. VERITY, R. 1906. Rhopalocera Palaearctica. 37-76, 8 pls. Florence. 1907. Rhopaloceva Palaearctica. 77-124,12 pls. Florence. WEstwoop, J.O. 1851. On the Papilio telamon of Donovan, with descriptions of two other eastern butterflies. Tvans. ent Soc. Lond. 1 : 173-176. Witson, K.H. 1961. In Ehrlich, P. R. & Ehrlich, A. H., How to know the butterflies. viii + 262 pp., 525 figs. Dubuque, Iowa. Wyatt, C. & Omoto, K. 1963. Auf der Jagd nach Parnassius autocrator Avin. Z. wien. ent. Ges. 48 : 163-170. YoKouHAMA, M. & WAKABAYASHI, M. 1968. Coloured illustrations of the butterflies of Japan 178 pp., 74 pls. Osaka. 4 Fics 1-8. Male genitalia, left clasper removed. 1, Avchon apollinus amasinus (Staudinger & Rebel). 2, Hypermnestva helios maxima Grum-Grshimailo. 3, Parnassius tenedius tenedius Eversmann. 4, P. havdwickii hardwickii Gray. 5, P. szechenyii szechenyii Frivaldsky. 6, P. glacialis glacialis Butler. 7, P. acdestis lathonius Bryk. 8, P. simo simonius Staudinger. ’ 8 Fics 9-16. Male genitalia, left clasper removed. 9, Parnalius polyxena polyxena (Denis & Schiffermiiller). 10, P. rwmina australis (Esper). 11, P. cerisy speciosa (Stichel). 12, Luehdorfia puziloi puziloi (Erschoff). 13, L.japonicajaponica Leech. 14, Bhutanitis thaidina thaidina (Blanchard). 15, B. lidderdalii lidderdalii Atkinson. 16, Sericinus montela magnus Fruhstorfer. GENERA AND SPECIES OF PARNASSIINAE 101 Fics 17-20. Venation of right fore- and hindwings. 17, Parnalius cerisy speciosa (Stichel). 18, Archon apollinus amasina (Staudinger & Rebel). 19, Parnalius polyxena polyxena (Denis & Schiffermiiller). 20, Hypermnestra helios maxima Grum-Grshimailo. 102 P. R. ACKERY Fics 21-24. Venation of right fore- and hindwings. 21, Parnassius imperator imperator Oberthiir. 22, P. cephalus ares Bryk & Eisner [forewing only]. 23, Luehdorfia japonica japonica Leech. 24, Parnassius charltonius deckerti Verity [forewing only]. GENERA AND SPECIES OF PARNASSIINAE 103 Fics 25-26. Venation of right fore and hindwings. 25, Bhutanitis thaidina thaidina (Blanchard). 26, Sericinus montela magnus Fruhstorfer. 104 P, R. ACKERY Pa Fics 27-32. 27-28, forelegs, showing foretibial epiphysis of (27) Parnassius orleans Oberthiir, (28) P. autocrator Avinoff. 29-30. Male tarsal claws of forelegs of (29) Hypermnestva helios (Nickerl), (30) Parnassius cephalus Grum-Grshimailo. 31-32. Dorsal view of uncus of (31) P. phoebus (Fabricius), (32) P. mnemosyne (Linnaeus). GENERA AND SPECIES OF PARNASSIINAE 105 INDEX Synonyms are in tfalics. acco, 86 acdestis, 87 actius, 81 Allancastria, 90 apollinus, 75 apollo, 83 apollonius, 80 Archon, 75 ariadne, 85 Armandia, 93 autocrator, 89 Bhutanitis, 93 bremeri, 81 cephalus, 86 cerisy, 91 charltonius, 88 clarvius, 85 clodius, 85 delphius, 87 Dorarchon, 75 Doritis, 75, 76 epaphus, 82 Eugraphis, 90 Eukoramius, 76 eversmanni, 84 eximia, 92 glacialis, 84 hardwickii, 86 helios, 76 honrathi, 80 Hypermnestra, 75 hypermnestra, 91 hypsipyle, 92 imperator, 88 inopinatus, 89 Ismene, 75 P. R. ACKERY Department of Entomology British Museum (NaTurRAL History) CROMWELL Roap Lonpon SW7 5BD jacquemontii, 81 japonica, 93 Kailasius, 76 Kovramius, 76 lidderdalii, 94 Lingamius, 76 loxias, 89 ludlowi, 94 Luehdorfia, 92 mansfieldi, 94 maturna, 92 mnemosyne, 83 montela, go nomion, 82 nordmanni, 85 orleans, 85 Parnalius, 90 Parnassius, 76 patricius, 87 phoebus, 81 polyxena, 9I puziloi, 92 rumina, 92 Sericinus, 90 simo, 89 stubbendorfii, 84 szechenyii, 86 Tadumia, 76 telemon, 90 tenedius, 89 thaidina, 93 Thais, 90 tianschanicus, 82 thia, 75 Zerynthia, 90 EXGs Fic. EG. Fic. FIG: Fic. Fic. Fic. I ay PLATE 1 Upper- and undersides (natural size) Archon Hiibner apollinus apollinus (Herbst) ¢ (Turkey: Taurus Mts) apollinus apollinus (Herbst) 9 (Turkey: Taurus Mts) Hypermnestra Ménétriés helios maxima Grum-Grshimailo ¢ (U.S.S.R.: Uzbekistan, Buchara) helios maxima Grum-Grshimailo 9 (U.S.S.R.: Uzbekistan, Buchara) Parnassius Latreille apollonius narynus Fruhstorfer g (U.S.S.R.: Kirghizia, Fort Naryn) apollonius narynus Fruhstorfer 9 (U.S.S.R.: Kirghizia, Fort Naryn) honrathi honrathi Staudinger § (U.S.S.R.: Tadzhikistan, Hasret Sultan mont) honrathi honrathi Staudinger 9 (U.S.S.R.: Tadzhikistan, Hasret Sultan mont) PEATE: Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 Fic. EGG. Fic. HIG: Fic. FIG: Fic. HG: PEATE 2 Upper- and undersides (natural size) Parnassius Latreille bremert bremeri Bremer ¢ (China-U.S.S.R.: Amur) bremeri bremeri Bremer Q (China-U.S.S.R.: Amur) phoebus sacerdos Vorbrodt ¢ (Switzerland: St Moritz) phoebus sacerdos Vorbrodt 2 (Switzerland: San Bernhardino) actius actius (Eversmann) ¢ (China-U.S.S.R.: Borokhoro Mts) actius actius (Eversmann) 9 (China-U.S.S.R.: Tien Shan) jacquemontit baroghila Tytler g (Pakistan: Chitral) jacquemontii baroghila Tytler 9 (Pakistan: Chitral) Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 BEAM Bec BiG: RiG: Fic. lacey FIG. FIGs Fic. Fic. A 18. 19% 20: 21, 22. 22: aA PLATE 3 Upper- and undersides (natural size) Parnassius Latreille nomion dis Bryk & Eisner ¢ (U.S.S.R.: Irkutsk) nomion dis Bryk & Eisner 9 (U.S.S.R.: Irkutsk) epaphus sikkimensis Elwes ¢ (S. Tibet: Mago) epaphus sikkimensis Elwes @ (S. Tibet) tianschanicus tianschanicus Oberthiir g§ (U.S.S.R.: Turkestan) tianschanicus tianschanicus Oberthiir 9 (U.S.S.R.: Turkestan) apollo apollo (Linnaeus) ¢ (Sweden) apollo apollo (Linnaeus) 2 (Sweden) PLATE 3 Bull, Br. Mus. nat. Hist. (Ent.) 31, 4 Fic. nies PIG. InTG. BIG. Fic. RIG: Fic. 25. 20. 27 25. 29. 30. Bit 225 PEATE 4 Upper- and undersides (natural size) Parnassius Latreille mnemosyne similis Bryk & Eisner 3 (Hungary) mnemosyne similis Bryk & Eisner 2 (Hungary) stubbendorfii stubbendorfi Ménétriés g (U.S.S.R.: Russia, Buryat, Sajan) stubbendorfii stubbendorfii Ménétriés 9 (U.S.S.R.: Russia, Buryat, Sajan) glacialis glacialis Butler 3 (Japan: Hokkaido, Sapporo) glacialis glacialis Butler 9 (Japan: Hokkaido, Yesso) eversmanni felderi Bremer ¢ (U.S.S.R.: Russia, Yevrey, Pompejefka) eversmanni feldevi Bremer 9 (U.S.S.R.: Russia, Yevrey, Radde) PLATE 4 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 Fic. Hie. Fic. Fic. BiG: HIG. BiG: 1gitex PLATE 5 Upper- and undersides (natural size) Parnassius Latreille eversmanni eversmanni Ménétriés ¢ (U.S.S.R.: Russia, Buryat, Sajan) eversmanni eversmanni Ménétriés 9 (U.S.S.R.: Russia, Buryat, Sajan) novdmanni novdmanni (Nordmann) ¢ (Caucasus) novdmanni nordmanni (Nordmann) 9 (Caucasus, Adshara Mts) aviadne aviadne Lederer ¢ (U.S.S.R.-Mongolia: Altai) ariadne aviadne Lederer 9 (U.S.S.R.: Turkestan) clodius clodius Ménétriés g (U.S.A.: West Washington) clodius clodius Ménétriés 9 (U.S.A.: Oregon) PLATE 5 Bull, Br. Mus. nat. Hist. (Ent.) 31, 4 Fie. Fic. Fic. BIG. lex Fie. luces Fic. PEATE <6 Upper- and undersides (natural size) Parnassius Latreille orleans orleans Oberthiir ¢ (China: Szechwan, How-Kow) orleans orleans Oberthiir 9 (China: Szechwan, How-Kow) hardwickii harvdwicku Gray 3 (N. India: Almora) havdwickii hardwickii Gray 9 (Nepal: Jargeng Khola) szechenyii szechenyii Frivaldsky 3 (China: Tsinghai, Kuku-Nor) szechenyii szechenyti Frivaldsky 92 (Tibet: Amdo) cephalus aves Bryk & Eisner ¢ (China: Kansu) cephalus aves Bryk & Eisner 9 (China: Kansu) PLATE 6 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 BiG: Fie. FIG. Fic. ie: BIG: BIG? BiG: PLATE 7 Upper- and undersides (natural size) Parnassius Latreille cephalus maharaja Avinoff ¢ (Kashmir: Ladakh) cephalus maharaja Avinoff 9 (Kashmir: Ladakh) acco tagalangi Bang-Haas ¢ (Kashmir: Ladakh) acco tagalangi Bang-Haas 9 (Kashmir: Ladakh) acco hunnyngtoni Avinoff 3 (Tibet: Kyetrak) acco hunnyngtoni Avinoff 2 (Tibet: Chumbi Valley) patricius kardakoffi Bryk & Eisner 3 (U.S.S.R.: Kirghizia, Kungei-Ala-Tau) patricius kardakoffi Bryk & Eisner 9 (U.S.S.R.: Kirghizia, Kungei-Ala-Tau) PLATE 7 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 Fic. Fic. ics Hie: BiG: Fic. TG: Fic. PLATE 8 Upper- and undersides (natural size) Parnassius Latreille delphius albulus Honrath ¢ (China: Sinkiang, Tien Shan) delphius albulus Honrath @ (China: Sinkiang, Tien Shan) delphius beate Eisner § (Kashmir: Potu-la Pass) delphius beate Eisner 2 (Kashmir: Chalsi) acdestis acdestis Grum-Grshimailo ¢ (Tibet: Amdo) acdestis acdestis Grum-Grshimailo 2 (Tibet: Amdo) imperator imperator Oberthiir ¢ (China: Szechwan, Ta-tsien-lou) imperator imperator Oberthiir 9 (China: Szechwan, Ta-tsien-lou) PLATE 8 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 IG: IgiKes ines Ric: TG; TG: Fie. BIG. 65. 60. 67. 68. 69. 70. 715 2s PLATE 9 Upper- and undersides (natural size) Parnassius Latreille charltonius sevenissimus (Bryk) ¢ (Kashmir: Kutie Pass) charltonius serenissimus (Bryk) 9 (Kashmir; Dugi Pass & Reyling) inopinatus inopinatus Kotzsch ¢ (Afghanistan: Marak) inopinatus inopinatus Kotzsch 9 (Afghanistan: Marak) loxias loxias Piingeler ¢ (China: Sinkiang, Aksu) loxias loxias Piingeler 9 (China: Sinkiang, Tien Shan) autocratoy autocratoy Avinoff ¢ (Afghanistan: Hindu Kush) autocrator autocratoy Avinoff 2 (Afghanistan: Hindu Kush) PLATE 9 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 Fic. Fic. Fic. Fic. BIG: Fic. InIG: Fic. 79. 80. PLATE tro Upper- and undersides (natural size) Parnassius Latreille simo simonius Staudinger ¢ (U.S.S.R.: Turkestan) simo simonius Staudinger 9 (U.S.S.R.: Kirghizia, Transalai) tenedius tenedius Eversmann ¢ (U.S.S.R.: Sajan Mts, Arasagun-gol) tenedius tenedius Eversmann 9 (U.S.S.R.: Sajan Mts, Arasagun-gol) Sericinus Westwood montela shantungensis Hering g§ (China: Shantung) montela montela Gray 2 (China: Hopei, Peking) Parnalius Rafinesque cerisy speciosa (Stichel) 3 (Lebanon: Beirut) cerisy speciosa (Stichel) 9 (Lebanon: Beirut) PLATE tro Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 PARE Sr Upper- and undersides (natural size) Parnalius Rafinesque Fic. 81. polyxena polyxena (Denis & Schiffermiiller) J (Yugoslavia: Kragouyevatz) Fic. 82. polyxena polyxena (Denis & Schiffermiiller) 9 (Hungary) Fic. 83. vumina rumina (Linnaeus) ¢ (Spain) Fic. 84. vumina rumina (Linnaeus) 9 (Spain) Luehdorfia Criiger Fic. 85. puziloi puziloi (Erschoff) ¢ (U.S.S.R.-China: Amur) Fic. 86. puziloi puziloi (Erschoff) 9 (U.S.S.R.-China: Amur) Fic. 87. japonica japonica Leech § (Japan: Honshu, Oyama) Fic. 88. japonica japonica Leech 9 (Japan: Honshu, Oyama) Fic. 89. japonica chinensis Leech ¢ (China: Kiangsu, Nanking) Fic. 90. japonica chinensis Leech 9 (China: Hupeh, Chang-Yang) PLATE 11 Bull, Br. Mus. nat. Hist. (Ent.) 31, 4 PAT a2 Upper- and undersides (natural size) Bhutanitis Atkinson Fic. 91. mansfieldi (Riley) 2 holotype (China: Yunnan) Fic. 92. thaidina thaidina (Blanchard) ¢ (China: Yunnan, Tsekou) Fic. 93. thaidina thaidina (Blanchard) 2 (China: Szechwan, Ta-tsien-lou) PICATE er Bull, Br. Mus. nat. Hist. (Ent.) 31, 4 DAA: 13 Upper- and undersides (natural size) Bhutanitis Atkinson Fic. 94. lidderdalii lidderdalii Atkinson ¢ (India: Naga Hills, Kohima) Fic. 95. liddevdalii lidderdalii Atkinson 9 (Bhutan) PLATE 13 Bull. Br. Mus. nat. Hist. (Ent.) 31, 4 BIG. Ines HIG. Bic. res Fic. ites Fic. 98. 99. 100. IOT. EO? 103. 104. 105. PLATE 15 Lateral view of sphragis Parnassius Latreille apollo (Linnaeus) mnemosyne (Linnaeus) szechenyii Frivaldsky acdestis Grum-Grshimailo delphius (Eversmann) amperatoy Oberthiir charltonius Gray inopinatus Kotzsch Bull, Br. Mus. nat. Hist. (Ent.) 31, 4 PEATE srs ENTOMOLOGY SUPPLEMENTS Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . Sanps, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxaDA, T.. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures,g maps. February, 1967. £3.50. HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8. . Mounp, L.A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. {5. . AriF1, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. CrosskEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. £12. Sanps, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. CrosskEy, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species, Pp. 167: 109 text-figures. February, 1973. £6.50. . voN Hayex, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CrossKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THR GRESHAM PRESS OLD WOKING SURREY BNGLAND THE NESOTHRIPS COMPLEX OF SPORE-FEEDING THYSANOPTERA (PHLAEOTHRIPIDAE: IDOLOTHRIPINAE) L. A. MOUND BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 5 LONDON : 1974 > Rik “ fone . faa | als [ hae Cas kT Ay CAS TT) Vig ite lad Lon - 7 Mb og tha) oa ea 1 t MA | > ee L ; ay ‘ Any et Pe al : ate ee Sl ask Bee de THE NESOTHRIPS COMPLEX ny oS OF SPORE-FEEDING THYSANOPTERA (PHLAEOTHRIPIDAE : IDOLOTHRIPINAE) BY LAURENCE ALFRED MOUND Pp 107-188; 74 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 5 LONDON : 1974 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), tnstituted im 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 veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 31 No. 5 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Pertodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) © Trustees of the British Museum (Natural History), 1974 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 12 December, 1974 Price £4-80 THE NESOTHRIPS COMPLEX OF SPORE-FEEDING THYSANOPTERA (PHLAEOTHRIPIDAE : IDOLOTHRIPINAE) By L. A. MOUND CONTENES Page SYNOPSIS ‘ : : . é : : : 7 A ‘ 109 INTRODUCTION e ; : : : : : a : 2 109 RECOGNITION OF GENERA . a F ; : ° ; ‘ - IIo DISTRIBUTION OF GENERA . j F : : : ‘ : : IIo SPECIATION IN LITTER THRIPS. ‘ : ; . : , : IIl THRIPS DISPERSION BY MAN ‘ : ; , 5 : : , LEE ACKNOWLEDGEMENTS . ‘ ‘ : F A : P : II2 ABBREVIATIONS : A é : ‘ ; , : TI2 CHECK-LIST OF THE SPECIES DISCUSSED IN THIS PAPER . : : , 113 KEY TO THE GENERA OF THE Nesothrips COMPLEX ; : : ; 115 GENERA AND SPECIES DISCUSSED ALPHABETICALLY i : ; , 117 SPECIES REMOVED FROM THE Nesothrips COMPLEX. ‘ ; ; : 181 REFERENCES . ; ; 5 : 5; ‘ ; p : 182 INDEX . : : ; 3 , F é ; : E : 186 SYNOPSIS Difficulties in the definition of genera among the Cryptothripini are correlated with the biology of these spore-feeding thrips which live in leaf litter and on freshly dead wood. Neso- thrips is divided into five genera, each with a restricted geographical distribution, but the structural characteristics on which these genera are based are weak and overlapping. Keys are provided to twelve genera and one hundred and one species, and three species are transferred to the subfamily Phlaeothripinae. Two new genera and seven new species are described and sixteen new synonymies are established. These synonymies suggest that some species have been transported between various tropical countries by human commerce. INTRODUCTION THE GENUS Nesothrips has been interpreted by recent authors in different ways. Stannard (1957) used the name for a large group of species with a wide range of morphological characteristics and placed eight generic names insynonymy. Priesner (1961), however, retained a large element of the traditional classification and recog- nised five genera in the group when proposing a classification of the suborder Tubuli- fera. Unfortunately neither of these authors was able to examine the full range of forms in the Nesothrips-complex because the collections of J. D. Hood and R. S. IIo L. A. MOUND Bagnall were not readily available at the time. In the present study one hundred species are recognized in the complex and through the courtesy of many colleagues the author has been able to study type-material or authentic specimens in all except two instances. As a result a more traditional view has been adopted using relatively small genera. The generic classification of the whole tribe Cryptothripini is in need of revision, but the present study is concerned solely with those species which have been associated with Nesothrips or the closely related genus Rhaebothrips. RECOGNITION OF GENERA In the Cryptothripini the individual genera intergrade in a confusing pattern and it was for this reason that Stannard (1957) used Nesothrips in a broad sense. However, this has several disadvantages; it obscures the fact that certain species of Nesothrips are morphologically more similar to species in other genera than they are to other species of Nesothrips; it conceals the fact that closely related groups of species tend to occur nearer to each other geographically than do less related species; it conceals the structural diversity of the Nesothrips group. If the genus is used in Stannard’s sense then Rhaebothrips and Acallurothrips must be included. Moreover some species of Dichaetothrips and Scotothrips then become difficult to distinguish from the enlarged genus. In this way the concept of a genus approaches Priesner’s (1961) concept of a tribe or subtribe, and the name Nesothrips no longer calls to mind any concise set of structural, biological or geographical attributes. In order that genera reflect evolutionary relationships between groups of species it is sometimes necessary to admit that there are intermediate species. The use of subgenera may be helpful at times in these circumstances, but this would not be practical in the Cryptothripini in view of the complexity of the relationships between the various forms. The origin of these complex relationships probably lies in the lack of host or habitat specificity in many of these fungal-spore-feeding thrips (Mound & O’Neill, 1974 and also below). DISTRIBUTION OF GENERA The generic analysis in this paper has been facilitated partly by establishing the synonymy of several tropical tramp species, and partly by transferring to Neosmerin- thothrips several South American species described in Gastrothrips. The result is not completely satisfactory, but in that it establishes a correlation between structural similarity and geographical distribution this system probably reflects evolutionary relationships. The genus Nesothrips sensu stricto is based on a species from Hawaii and is used here for a group of species from the Pacific, Australia and New Zealand. This is itself a difficult group of species which are not all closely related, and some of which are very closely related to Rhaebothrips species from the Pacific and New Zealand. Carientothrips is another related genus but this has its centre of diversity in Australia rather than the Pacific Islands. In contrast Bolothrips is a holarctic genus usually associated with grass tussocks, but with several species in South Africa. Gastrothrips is a Neotropical genus with a few species entering North NESOTHRIPS COMPLEX III America and one species-group widespread (?introduced) in the old world. The remaining large genus, Neosmerinthothrips, is more confusing with species apparently native to various parts of the tropics. The characters on which these genera are separated are summarised in the key below, although few of the genera can be distinguished on single characters. SPECIATION IN LITTER THRIPS The species of the subfamily Idolothripinae (= Megathripinae), to which the Nesothrips complex belongs, all feed on fungal spores, judging from the gut contents of the specimens which have been studied. They usually live in leaf litter or on freshly dead wood and they are found mainly in the tropics. This habitat is wide- spread and readily available across large parts of the earth’s surface, and there is no evidence that the fungi on which the thrips feed are restricted in their distribution. This has important effects on the biology of the thrips, their dispersive activity and resultant speciation, as well as the ease with which they are transported by man. Although the habitat is widespread the nutritional status of the fungi at any one site varies with the state of decay of the substratum. This fluctuation in suitability of the habitat has probably stimulated the evolution of winged and wingless morphs in many species, as well as the production of major and minor (oedymerous and gynaecoid) individuals. It is assumed that this structural diversity within individual species is controlled largely by the environment and that the flexibility of the morphogenetic process is itself inherited. The evolution of such a group, within which each species is structurally diverse, can be expected to produce a complicated series of forms difficult to classify into genera. Moreover the lack of host specificity and the irregular production of winged individuals is likely to facilitate gene flow between populations of incipient species. This could have caused the complicated patterns of variation between populations that are found apparently irregularly in Carientothrips mjobergi (Mound, 1974) and Biconothrips species (Mound, 19720) or in clines as in Bolothrips species (q.v.) and Allothrips megacephalus (Mound, 19722). THRIPS DISPERSION BY MAN Several species of fungus-feeding thrips from leaf litter and dead wood appear to have been distributed around the world by human commerce. Some of these have been referred to elsewhere (Mound, 1970 : 87, Nesothrips propinquus and Hoplandro- thrips flavipes), and the establishment of new synonymy in the present study has revealed further examples e.g. Scotothrips claripennis, Neosmerinthothrips collaris and Rhaebothrips lativentris. Such species have probably been transported in sailing ships: in dry grasses used as fodder and bedding; in seed pods and coconut husks; in baskets or tubs of plants; on dead wood, or in ballast. Different species seem to be associated with different shipping routes. Thus N. propinguus is found along the route from Europe to New Zealand via South Africa. N. collaris is found in West Africa and the West Indies, the two ends of one of the major slave 112 L. A. MOUND trading routes, and this lends support to the suggestion made in this paper that Neosmerinthothrips hilaris from West Africa and N. diversicolor from Brazil may be variants of one species. A similar situation is found in the Phlaeothripine genus Rhinoceps. ‘The first species in this genus, R. jansei Faure, 1949, was described from south-east Africa, the second species, R. cornutus zur Strassen, 1972, from Colombia, and there is a third (?) species in the BMNH collection from Ghana. However, in view of the very small differences between these ‘species’ it seems more likely that they are merely local variants which have been distributed by man. A distribution involving south-east Africa and the West Indies is also found in Scotothrips claripennis and Neosmerinthothrips collaris, and again this relationship calls to mind the slave trading pattern of the seventeenth and eighteenth centuries. It would not have been possible to transport more than a million men and women in conditions of appalling squalor without transporting fungus-feeding and detritus- living insects. The fact that S. claripennis is also found in India may reflect even earlier patterns of human trading. Ships have crossed the Indian ocean for several thousand years between such distant points as China and East Africa, and this could well explain much of the present-day distribution of such species as Nesothrips brevicollis. ACKNOWLEDGEMENTS This study would not have been possible without the active support of many colleagues. Dr Jacot-Guillarmod of Grahamstown, South Africa and the late Professor Priesner of Linz, Austria not only loaned many valuable specimens from their collections, but also gave the author the benefit of their experience with the Crypto- thripini in the initial stages of the project. Kellie O’Neill of Washington, Professor Ananthakrishnan of Madras and Dr zur Strassen of Frankfurt also loaned numerous specimens. To all of these the author is deeply grateful. Figures 1-58 in this paper were all drawn to the same magnification by my colleague Mrs Jenny Palmer, and figures 59~74 were all drawn by the author to a different magnification. ABBREVIATIONS The following abbreviations are used for depositories. AMG Albany Museum, Grahamstown, South Africa ANIC Australian National Insect Collection, C.S.I.R.O., Canberra BMNH British Museum (Natural History) BPBM Bernice P. Bishop Museum, Hawaii CAS California Academy of Sciences, San Francisco DSIR Department of Scientific & Industrial Research, Auckland, New Zealand FDA Florida Department of Agriculture, Gainsville, U.S.A. HP Dr H. Priesner collection, Linz, Austria INHS _ Illinois Natural History Survey, Urbana, U.S.A. NR Naturhistoriska Riksmuseum, Stockholm Pretoria Plant Protection Research Institute, Pretoria, South Africa NESOTHRIPS COMPLEX 113 NZFI New Zealand Forest Research Institute, Rotorua QM Queensland Museum, Brisbane, Australia SMF Senckenberg Museum, Frankfurt ™ Természettudomanyi Mizeum, Budapest TNA Professor Ananthakrishnan collection, Loyola College, Madras, India USNM_ United States National Museum (National Museum of Natural History), Washington D.C. The following abbreviations are used in the descriptions of species for the major setae of the pronotum. epim epimeral setae posteroangular setae am anteromarginal setae aa anteroangular setae pa ml midlateral setae CHECK-LIST OF THE SPECIES DISCUSSED IN THIS PAPER Family PHLAEOTHRIPIDAE Uzel Phloeothripidae Uzel, 1895 Subfamily IDOLOTHRIPINAE Bagnall Idolothripidae Bagnall, 1908 Megathripidae Bagnall, 1914 Megathripinae Bagnall; Karny, 1919 CARIENTOTHRIPS Moulton, 1944 stat.n. acti Mound, 1974 badius (Hood, 1918) comb. n. apterus (Girault, 1928) BOLOTHRIPS Priesner, 1926 gen. rev. Bolothrips (Botanothrips) Hood, 1939 bicolor (Heeger, 1852) bicolor brevicornis Priesner, 1928 andrei Watson, 1933 cinctus Faure, 1943 cingulatus (Karny, 1916) dentipes (Reuter, 1880) bagnalli (Karny, 1916) dentis Faure, 1954 embotyi Faure, 1943 gilvipes Hood, 1914 litoreus Hood, 1939 syn. n. icarus (Uzel, 1895) tcarus pallipes (Uzel, 1895) insularis (Bagnall, 1914) icarus tuberculatus (Priesner, 1922) brachyurus (Bagnall, 1927) arenarius Priesner, 1950 Syn. n. italicus sp. n. pratensis Hood, 1939 rachiphilus Cott, 1956 schafferi (Thomasson & Post, 1966) comb. n. varius Hartwig, 1948 biformis (Moulton,1939) comb. n. capricornis (Mound, 1974) comb. n. casuarinae Mound, 1974 denticulatus (De Santis, 1963) comb. n. Jijiensis Moulton, 1944 grayi sp. n. japonicus (Bagnall, 1921) comb. n. loisthus Mound, 1974 magnetis Mound, 1974 miskoi Mound, 1974 miobergi (Karny, 1920) australicus (Priesner, 1928) incisus (Girault, 1927) fiavitibia (Moulton, 1968) pedicillus Mound, 1974 pictilis Mound, 1974 reedi Mound, 1974 semirufus (Girault, 1928) vesper Mound, 1974 DICERATOTHRIPS Bagnall, 1908 validipennis (Hood, 1938) comb, n. 114 DICHAETOTHRIPS Hood, 1914 gloveri (Ramakrishna Ayyar & Marga- bandhu, 1939) comb. rev. GASTROTHRIPS Hood, t912 gen. rev. Probolothrips Moulton, 1941 abditus Hood, 1935 brasiliensis (Moulton, 1938) hambletoni (Moulton, 1941) acuticornis (Hood, 1925) cybele (Girault, 1927) noumeae Bianchi, 1945 alticola Hood, 1942 anolis Morgan, 1925 proteus Hood, 1933 callipus Hood, 1935 corvus Priesner, 1933 capitalis Hood, 1935 syn. n. falcatus Ananthakrishnan, 1968 comb. n. fulvicauda Hood, 1937 fulviceps Hood, 1937 fumipennis Hood, 1952 intonsus Hood, 1941 mandiocae (Moulton, 1941) oeceticola De Santis, 1943 mongolicus Pelikan, 1965 comb. n. monticola Hood, 1942 procerus Hood, 1956 ruficauda Hood, 1912 stygicus Hood, 1935 subulatus (Hartwig, 1948) comb. n. texanus Hood, 1912 NEOSMERINTHOTHRIPS Schmutz, 1913 gen. rev. Coenurothrips Bagnall, 1921 Galactothrips Moulton, 1933 syn. n. affinis (Bagnall, 1921) comb. n. annulipes (Hood, 1950) comb. n. milleforme (De Santis, 1963) Syn. n. brevicollis (Bagnall, 1921) comb. n., stat. rev. collaris (Bagnall, 1917) comb. n. fuscicauda (Morgan, 1925) syn. n. marshalli (Priesner, 1934) Syn. n. dominicanus (Hood, 1925) Syn. n. diversicolor (Moulton, 1933) comb. n. Sijiensis (Moulton, 1944) comb. n. fructuum Schmutz, 1913 ceylonicus (Karny, 1925) Syn. n. hilaris (Priesner, 1937) comb. n. hoodi (Faure, 1954) comb. n. inquilinus Ananthakrishnan, 1960 nigrisetis (Hood, 1935) comb. n. parvidens (Hood, 1935) comb. n. paulistarum (Hood, 1950) comb. n. L. A. MOUND picticornis (Hood, 1936) comb. n. plaumanni (Hood, 1950) comb. n. robustus (Ananthakrishnan, 1964) comb. n. variipes (Hood, 1950) comb. n. xylebori Priesner, 1935 NESIDIOTHRIPS gen. n. alius (Ananthakrishnan, 1970) comb. n. validus (Bagnall, 1921) comb. n. NESOTHRIPS Kirkaldy, 1907 Oedemothrips Bagnall, 1910 aoristus Mound, 1974 artocarpi (Moulton, 1942) brevicollis (Bagnall, 1914) minor (Bagnall, 1921) syn. n. formosensis (Priesner, 1935) Syn. n. formosensis karnyi (Priesner, 1935) syn. n. carverae Mound, 1974 fodinae sp. n. hemidiscus Mound, 1974 malaccae sp. n. melinus Mound, 1974 niger (Moulton & Steinweden, 1932) oahuensis Kirkaldy, 1907 laticeps (Bagnall, 1910) propinquus (Bagnall, 1916) dimidiatus (Hood, 1918) propinquus breviceps (Bagnall, 1924) propinquus obscuricornis (Bagnall, 1924) cestosa (Karny, 1920) oleriae (Moulton, 1949) similis (Hartwig, 1948) Syn. n. rhizophorae (Girault, 1927) semiflavus (Moulton, 1939) yanchepi Mound, 1974 PHACOTHOHRIPS gen. n. ocelloides (Hood, 1950) comb. n. RHAEBOTHRIPS Karny, 1913 doulli sp. n. eastopi sp. n. lativentris Karny, 1913 claripennis (Hood, 1919) seychellensis (Bagnall, 1921) syn. n. difficilis (Bagnall, 1921) syn. n. ipomoeae (Ishida, 1932) magnus (Moulton, 1928) yuasat (Moulton, 1928) fuscus Moulton, 1942 australiensis (Moulton, 1968) leveri sp. n. major Bagnall, 1928 nigrisetis Sakimura, 1972 zondagi sp. n. NESOTHRIPS COMPLEX 115 SCOTOTHAHRIPS Priesner, 1939 diversus (Ananthakrishnan, 1972) claripennis (Moulton, 1934) comb. n. syn. n. trinidadensis (Hood, 1935) Syn. n. firmus (Hood, 1952) comb. n. indicus (Ananthakrishnan, 1968) SYNCEROTHRIPS Hood, 1935 gen. rev. syn. n. harti Hood, 1935 Species removed from Nesothrips complex Subfamily PHLAEOTHRIPINAE Uzel, 1895 Adelothrips lativerticis (Post, 1961) Adelothrips speciosissimus (Karny, 1920) comb. n. Liothrips debilis (Hood, 1936) comb. n. KEY TO THE GENERA OF THE NESOTHRIPS COMPLEX 1 Antennal segment IV with two sense cones (Text-fig. 59). : : . : 2 — Antennal segment IV with three or more sense cones (Text-figs 61-74) 2 Antennal segment VIII less than twice as long as wide, broadly joined to VII; head not projecting in front of eyes; interocellar setae twice as long as distance between two ocelli; female with a fore tarsal tooth [South Africa] Bolothrips dentis (p. 120) — Antennal segment VIII slender, twice as long as wide; head strongly projecting in front of eyes; no elongate setae near the ocelli (Text-fig.1) ; female without a fore tarsal tooth : 3 3 Antennal segment III with to sense cones; prothorax and all legs eoltaws i in both sexes, rest of body brown [Europe and eastern North America] Bolothrips bicolor (p. 118) — Antennal segment III with one sense cone; prothorax brown in 9, legs brown with yellow markings on the femora, ¢ with pronotum and pterothorax yellow with brown shadings laterally [Italy] . : : . Bolothrips italicus (p. 122) 4 Antennal segment IV with three sense cones (Text-figs 64, 6 Slo : : 5 — Antennal segment IV with four or rarely five sense cones (Text-figs 6a) : ‘i 5 Eyes usually longer ventrally than dorsally, if eyes not prolonged ventrally then metanotal setae weak and pelta broadly rounded; tube always black with straight sides (Text-fig. 13); e. and @ usually apterous, 2 rarely macropterous [mainly Holarctic] . : : : BOLOTHRIPS (p. 117) — Eyes rarely prolonged on ventral surface of head: one pair of metanotal setae frequently stout; pelta usually triangular and never broadly rounded; tube variable, frequently yellow, or with apex sharply constricted (Text-figs 14-19) ; 2 usually macropterous, rarely micropterous or apterous; g macropterous or apterous [mainly Neotropical] : : 6 6 Antennal segment VIII distinct from VII, uswally long and slender, rarely less than 0-75 times aslongas VII . ; : . GASTROTHRIPS (p. 134) — Antennal segment VIII broadly fused to VIL, the suture incomplete dorsally SYNCEROTHRIPS (p. 181) 7 Head with an ommatidia-like papilla on cheek, situated midway between posterior margin of eye and posterior margin of head (Text-fig. 47); tube margins strongly convex, maximum diameter about five times apical diameter (Text-fig. 51); fore wing without duplicated cilia [Brazil] . . . . PHACOTHRIPS (p. 170) — Head with normal cheeks, rarely with one isolated ommatidium close behind com- pound eye; tube frequently with straight sides, if convex then maximum diameter 116 Io p At 12 13 14 L. A. MOUND less than three times apical diameter; fore wing when present ies lacking duplicated cilia . : 8 Head with a pair of stout setae anterolateral to the fore ocellus but with no other long setae near the ocelli (rarely with these setae shorter than one side of ocellar triangle) (Text-fig. 9); antennal segment III more than 1-3 times as long as IV, segments VII and VIII with a complete suture but forming a single unit with an almost smooth outline (Text-fig. 66); pronotum broad and weakly reticulate, with anterior and median ridge scarcely thickened even in major males, and mid- lateral setae usually small; fore femora of males frequently with a series of stout spines on inner margin [Neotropics] A ; ; DICERATOTHRIPS (p. 133) Head with a pair of stout setae between or behind the ocelli, or with no long setae near the ocelli; antennal segment III less than 1-2 times as long as IV, segment VIII usually smaller at base than VII at apex; pronotum frequently with a stout median thickening, anterior margin often thickened in males, and midlateral setae rarely small; fore femur of § without spine-like setae ; 9 Head with a pair of long setae arising within the ocellar triangle, midway between anterior and posterior ocelli (Text-fig. 46); fore tarsus of Q withastout tooth . NESIDIOTHRIPS (p. 156) Head with a pair of long setae arising between or behind the posterior ocelli, or ocellar setae all short; fore tarsus of 2 frequently without a tooth . , 10 Fore tarsal tooth well developed in 9; head always longer than wide, eyes never prolonged on ventral surface of head; ocellar setae arising behind posterior ocelli; cheeks frequently with several stout setae; tube slender with straight sides, evenly narrowing from base to apex : II Fore tarsal tooth usually absent in 9; if present then eyes prolonged on ventral surface of head, or ocellar setae arising between posterior ocelli, or head as wide as long and constricted to base without cheek setae, or tube heavy with stout setal tubercles or even with convex sides : . 12 Postocellar setae longer than one side of ocellar triangle; “cheeks with fine setae (Text-fig. 8) : ; , DICHAETOTHRIPS (p. 134) Postocellar setae shorter than | one se of ocellar triangle; cheeks frequently with stout setae . g : . SCOTOTHRIPS (p. 177) Maxillary stylets retracted into ‘head ; as far as postocular setae, parallel in middle of head and about one third of head width apart (Text-fig. 2); pelta of macropterae usually with very slender lateral wings [Pacific, Australia, Falkland Is] CARIENTOTHRIPS (p. 125) Maxillary stylets wide apart, arranged in a V-shape and usually low in head (Text- figs 40-42); pelta of macropterae without slender lateral wings : 13 Tube heavy with sides convex (Text-figs 35-37) and head with no elongate ocellar setae; or fore tarsal tooth present in 9, and tube heavy with one or more pairs of lateral setal bases, ocellar setae rarely elongate [Oriental, Ethiopian and Neo- tropical regions] ; . NEOSMERINTHOTHOHRIPS (p. 148) Tube with straight sides, ‘sometimes slightly constricted at apex (Text-figs 44, 56); fore tarsal tooth rarely present in 9; postocellar setae usually longer than diameter of one ocellus, frequently arising between the posterior ocelli [Pacific and Australian regions, introduced elsewhere] : . 14 Postocellar setae close together arising between the posterior ocelli (Text-figs 52- 54); tube rarely less than three times as long as maximum width; 9 rarely with a fore tarsal tooth ; i . RHAEBOTHRIPS (p. 171) Postocellar setae frequently arising behind posterior ocelli, if arising between the ocelli than the tube is shorter, less than 2°5 times as long as maximum width; ? never with a fore tarsal tooth A ; ; : ; ; ‘ ‘ ; 15 Head more than 1-3 times as long as wide . CARIENTOTHRIPS (part) (p. 125) Head less than 1-2 times as long as wide . : , ‘ NESOTHRIPS (p. 158) NESOTHRIPS COMPLEX 117 GENERA AND SPECIES DISCUSSED ALPHABETICALLY BOLOTHRIPS Priesner gen. rev. Bolothrips Priesner, 1926: 90. Type-species: Phloeothrips bicolor Heeger, by original designa- tion Bolothrips (Botanothrips) Hood, 1939 : 605-606. Type-species: Bolothrips pratensis Hood, by original designation. The genus Bolothrips is used here for a group of fourteen Holarctic and Ethiopian species which live at the base of grass tussocks and only rarely produce macropterae. The more common species appear to exist as irregular clines, for example the gilvipes, litoreus, rhachiphilus complex in North America, and the insularis, icarus, varius complex in Europe and Africa. The interpretation of these clines nomenclatorially must be rather arbitrary. Most species of Bolothrips have three sense cones on the fourth antennal segment, although the ventral sense cone is missing in the type-species bicolor as well as in dentis Faure. Gastrothrips species also have three sense cones on the fourth antennal segment although these are usually rather stouter than in Bolothrips species. More- over most species of Gastrothrips have a pair of stout metanotal setae, a triangular pelta, and well developed wings. Botanothrips has been used for a group of species in which the eyes are not prolonged ventrally, but this is not accepted here in view of the variation in insularis and varius. One species, dentis Faure, is retained in Bolothrips for the present although it combines the characters of several genera. GENERIC DEFINITION. Medium sized, black or bicoloured, rarely pale species of Cryptothri- pini. Head usually longer than wide, projecting in front of eyes, with eyes prolonged or at least narrowed ventrally; maxillary stylets V-shaped and wide apart. Antennae eight segmen- ted, VIII more than 0-5 times as long as VII; segment III with two (rarely one) sense cone, IV with three (rarely two) sense cones. Pronotum with epimeral sutures complete; ¢ with fore tarsal tooth and frequently with fore tibial apical tubercle; 2 without tooth or tubercle. Usually apterous, fore wing with duplicated cilia in bicolor and icarus. Metanotal median setae small and slender. Pelta usually broadly rounded; tube with straight sides. KEY TO THE SPECIES OF BOLOTHRIPS I Body colour light ee brown, abdominal segments VIII-X lightest [South Africa] J : .*embyoti (p. 120) - Body colour dark brown, thorax sometimes yellow: abdoniinal segments VIII-IX darker than rest of body ; k : ‘ ; 2 2 Antennal segment IV with two sense cones, s, the ventral one “missing : : : 3 — Antennal segment IV with three sense cones... 5 3 Fore tarsal tooth present in 9; interocellar setae longer than distance between ‘ocelli; tube paler than abdomen, constricted at apex [South Africa] . . dentis (p. 120) — Fore tarsal tooth not present in 2; interocellar setae not elongate; tube with sides straight : , 4 4 Prothorax and all legs yellow i in n both sexes, rest of body brown; ‘antennal segment III with two sense cones [Europe, eastern North America] ‘ . bicolor (p. 118) - Prothorax brown in 9, legs brown with yellow markings; $ with prothorax and pterothorax yellow but shaded brown laterally; antennal segment III with one sense cone; Italy : ; : : : ; ; ; . italicus (p. 122) 118 Lv Aa MOUND 5 Ventral length of eyes more than 1-6 times the dorsal length . ; — Ventral length of eyes less than 1-3 times the dorsal length : : p 7 II 6 Pronotum, pterothorax and abdominal segment I yellow. é : ‘ . 7 — Pronotum sometimes reddish yellow, but pterothorax brown 8 7 Abdominal segment II yellow; antennal segments II and IV largely brown [Europe] cingulatus (p. 119) — Abdominal segment II brown; antennal ces II brownish yellow, IV and V largely yellow [South Africa] : . . Ccinctus (p. 119) 8 Tube relatively long, at least 2-3 times as long as maximum width: antennal segment III largely yellow ‘ 9 — Tube relatively short, not more than 2 2°I times as ; long ; as maximum width; antennal segment III brown with pedicel light . : ste) 9 Legs uniform dark yellow; antennal segments III, IV and basal eae thirds of Vv yellow [western North America] . : . schafferi (p. 123) — Legs largely brown; antennal segments IV and V brown, TI yellow with brown shading at apex [Europe and North America] : : . dentipes (p. 119) 10 Head of apterae with reduced ocelli; pronotum reddish yellow [south- eastern North America] . : . gilvipes (p. 120) — Head of apterae without ocelli: -proniotuni black {Calitornial . *rhachiphilus (p. 123) 11 Body bicoloured, head and thorax much paler than abdomen [Florida, Texas] pratensis (p. 122) — Head and thorax as brown as abdomen . - 12 12 Dorsal setae on tergite IX less than 0-8 times as plone. as ihe [South Africa] varius (p. 123) — Dorsal setae on tergite IX more than o-9 times as ous as tube [Mediterranean to Latvia] ; 13 13 Antennal segment Iv ass at bases dorsal setae on . tergite IX generally shorter than tube : , . icarus (p. 121) — Antennal segment IV brown: dorsal setae on n tergite IX generally a little longer than tube . : : : ; : : : , : . .énsularis (p. 121) * Not studied Bolothrips bicolor (Heeger) Phloeothrips bicolor Heeger, 1852 : 477-478. Syntypes ?sex, AuSTRIA (? lost). Bolothrips bicolor (Heeger) Priesner, 1926 : go. Bolothrips bicolor f. brevicornis Priesner, 1928a : 687. Holotype 9, Huncary (HP) [not examined]. Oedaleothrips andrei Watson, 1933 : 49-50. Syntypes 4 9, 4 g, U.S.A.: Iowa (FDA) [not examined]. [Synonymized by Stannard, 1957 : 106.] This species is widespread in central and south-eastern Europe, and according to Stannard (1957) it is distributed through the northern part of North America. The present author has not studied specimens from west of North Dakota or Utah. One macroptera is listed below from Minnesota, and Priesner (1928 : 687) refers to macropterae from Austria found in July and August. The most closely related species are dentipes and italicus, whereas cingulatus has much shorter antennal segments with three sense cones on the fourth segment. SPECIMENS STUDIED. Huncary: Simontornia, 1 9, 1 g, 5.v.1924 (F. Pillich). CZECHOSLOVAKIA: I 9 NESOTHRIPS COMPLEX 119 ex Uzel Collection. FRANCE: Villefranche sur Saone, 3 3, 2 9, ix—x.1927 (O. John); Hyéres Plage, 1 g, ix.1927 (R. S. Bagnall) (BMNH). U.S.A.: Illinois, Grundy County, 1 J, x.1970 (L. A. Mound); Minnesota, Mac- Gregor, 7 9, 3 g apterae, 1 2 macroptera, iv.1936 (Ff. Andre); M., Hay Creek, 2 9, I g, xi.1936 (G. Decker); Iowa, Ottumwa, 27 J, 22 9, in Moss, 1.1933; I., Boone, 792,76, from Poa, vii—viii.1934 (Ff. Andre); North Dakota, Northwood, 1 9, iv.1g61 (R. I. Post); Utah, Blacksmith Fork Canyon, I 9, iv.1971 (G. F. Knowlton) (BMNH). Bolothrips cinctus Faure Bolothrips (Bolothrips) cinctus Faure, 1943 : 86-87. Syntypes 2 9, SoutH Arrica: Transvaal (Pretoria) [not examined]. This species has a shorter tube than any other species of Bolothrips, and it can be distinguished moreover by its colour pattern. The two syntype apterae were collected from Themeda triandra near Lydenburg, Transvaal. SPECIMEN STUDIED. SouTH AFrica: Transvaal, Ermelo, 1 9 aptera on Hyparrhenia, 21.v.1949 (J. C. Faure) (HP). Bolothrips cingulatus (Karny) Cryptothrips cingulatus Karny, 1916 : 92. Syntypes ? sex, AUSTRIA (HP) [not examined]. Bolothrips cingulatus (Karny) Priesner, 1926 : go. Priesner (1964 : 143) refers to this as a south-eastern European species and states that the macropterae have only the first and second abdominal segments light. The antennae of cingulatus are shorter than those of bicolor and dentipes. SPECIMENS STUDIED. HunGary: Simontornyia, I 3, 3 9 apterae, vi & ix. 1924 (F. Puillich). FRANCE: Rhone, Curis, 3 g, 19 apterae, vill. 1927 (O. John); Hyéres Plage, 1 2 aptera, ix. 1927 (R. S. Bagnall) (BMNH). Bolothrips dentipes (Reuter) Phloeothrips dentipes Reuter, 1880 : 12-14. 2 Syntypes, ? sex, FINLAND (? lost). Bolothrips dentipes (Reuter) Priesner, 1928a : 689-692. Cryptothrips bagnalli Karny, 1916 : 94. Syntypes, ? sex, SARDINIA (? lost). Cryptothrips dentipes var. bagnalli Karny; Priesner, 1925 : 154. This species is widespread in Europe between Ireland, Poland and Hungary. There are numerous specimens in the BMNH collections from southern England, and also one female without data but bearing a determination label ‘Reuter det. 3.84. Stannard (1968) records the species from Illinois, and one specimen has been studied from Massachusetts. 120 L. A. MOUND Bolothrips dentis Faure Bolothrips dentis Faure, 1954b : 155-159. Holotype g, TRANsvaat (Pretoria) [not examined]. This species was described from 14 males and 26 females, all apterae, collected at Mariepskop in Transvaal, mainly on dead branches covered in lichen. The combination of morphological characters is most unusual. The head is truncate at the anterior and bears a pair of stout interocellar setae, but the eyes are prolonged ventrally. The fore tarsus of the female bears a tooth but the fourth antennal segment has two sense cones. The tube is pale in colour and constricted at the apex. This one species thus shows characteristics from each of the genera of the Nesothrips complex. It is not a true Bolothrips species but is retained in this genus until the South African fauna is better known. SPECIMENS STUDIED. TRANSVAAL: Mariepskop, 2 9 paratypes on dead branches of Syzygium cordatum, iv. 1951 (J. C. Faure) (HP, AMG). Bolothrips embotyi Faure Bolothrips (Bolothrips) embotyi Faure, 1943 : 87-89. Syntypes 5 9, 7 ¢, SoutH Arrica: Pondo- land (Pretoria) [not examined]. No specimens of this species have been studied. Judging from the description it is a true Bolothribs with the eyes prolonged ventrally, the head projecting in front of the eyes, the fourth antennal segment with three sense cones, and the female without a fore tarsal tooth. Bolothrips gilvipes (Hood) Cryptothrips gilvibes Hood, 1914: 169-170. Holotype 9, U.S.A.: Maryland (USNM) [not examined]. Bolothrips (Bolothrips) litoreus Hood, 1939 : 609-612. Holotype 9, U.S.A.: Texas (USNM) [not examined]. Syn. n. This species appears to show a gradient in size and colour with the darkest speci- mens in the southern and western part of its range. The specimens from Florida listed below have the thorax varying in colour from yellow to light brown even at one site. In view of this variation there seems to be little point in distinguishing the larger and darker specimens as litoreus. The Californian species rhachiphilus is possibly another element in this cline. SPECIMENS STUDIED. — U.S.A.: Alabama, Mobile, r g on dead grass (det. gilvibes by J. D. Hood), 17.xii.1938 (J. D. Hood); Florida: Cedar Keys, 17 9, r 3, on salt marsh grass, I2.vii.1939 (P. Oman); F., Lake Wauberg, 1 9, iv. 1938; F., Hudson, 1 ¢ on grass, vii. 1939; F., Deland, 1 9, 1 g on grass, vii. 1939 (BMNH, USNM). NESOTHRIPS COMPLEX 121 Bolothrips icarus (Uzel) Cryptothrips Icarus Uzel, 1895 : 232-233. Syntypes g 9, CzEcHosLovakia: Bohemia (? lost). Cryptothrips icarus var. pallipes Uzel, 1895 : 233. Bolothrips icarus (Uzel) Priesner, 1928a : 692-695. All of the specimens listed below have the legs more or less yellow with a variable amount of brown shading. The species is widespread from southern Europe to Latvia and is very closely related to the Mediterranean insularis and the African varius, but has paler legs and the eyes usually less prolonged ventrally. Three macropterous females have been studied from France. SPECIMENS STUDIED. FRANCE: Villefranche sur Saone, 3 g, I 2, I Q macroptera, vii. 1927; Rhéne, Curis, 1 g, 2 9 macropterae, vill. 1927 (O. John); La Trinite sur Mer, 1 4, viii. 1968 (V. F. Eastop). Austria: Innsbruck, I 3, ix. 1929 (R. S. Bagnall). YUGOSLAVIA: Subotica, I g, vi. 1963. CZECHOSLOVAKIA: Ceje, I 9, vi. 1950 (J. Pelikan); Bzendei, 3 3, v. 1964 (V. F. Eastop). HuncGary: Simontornia, 3 9, I g, vi. 1924 (Pillich). U.S.S.R.: Latvia, 4 J, 1 9, vi. 1928; 2 J, 3 9, viii. 1929 (O. John) (BMNH). Bolothrips insularis (Bagnall) Cryptothrips insularis Bagnall, 1914 : 295. Holotype 9, CANARY IsLANDs (BMNH) [examined]. Cryptothrips icarus var. tuberculatus Priesner, 1922: 105. Holotype 9, YuGostavia (HP) [examined]. [Synonymized by Mound, 1968 : 141.] Cryptothrips brachyurus Bagnall, 1927 : 573-574. Lectotype g, FRancE (BMNH) [examined]. [Synonymized by Mound, 1968 : 141.] Bolothrips arenarius Priesner, 1950 : 36-37. Syntypes gf 9, Eaypr & Syria (HP) [examined]. Syn. n. Priesner (1928 : 693) suggested that this species (under the name tuberculatus) might be a geographical race of tcarus. Whereas icarus is found in continental Europe, imsularis appears to be restricted to the Mediterranean area between the Canary Islands and Syria. Both species are variable, but they can apparently be distinguished by the characters given in the key above. Specimens from the eastern part of the range of insularis, Cyprus and Egypt, generally have the tube shorter and stouter than specimens from the western part of the range, Spain and the Canary Islands. The specimens from Cyprus appear to be intermediate be- tween insularis and typical arenarius from Egypt. SPECIMENS STUDIED. Holotype 9 of insularis, CANARY IsLANDs (T. V. Wollaston) (BMNH). Lectotype 6 with 1 3, r 2 paralectotypes of brachyurus, FRANCE: Tamaris, iii. 1927 (R. S. Bagnall) (BMNH). Holotype 9, allotype ¢ [sic] of twberculatus, YUGOSLAVIA: Dalmatia, Ragusa, from grass, 31.x.1918 (HP). EcyptT: 40 km east of Meadi, r 3, 1 2 paratypes of arenarius, on Retama, 9.iv.1934; Ismailia, I 9, 23.iv.1935; Bingash, ¢ allotype, 17.i.1935 (HP). CANARY IsLANDs: Lancerota, I 2 on grass, iii. 1963, I g, ii. 1968 (zur Strassen). SPAIN: Chiclana, 122 L. A. MOUND 29. FRANCE: St Cyr sur Mer, 1 9, 2 g, ix. 1927 (R. S. Bagnall). Cyprus: Akrotiri Bay, 8 3, 42 on Juncus, vii. 1945 (Mavroumoustakis) (BMNH). Bolothrips italicus sp. n. Q aptera. Colour brown, head and posterior abdominal segments darker than pterothorax; femora variable in colour, mid and hind tibiae and tarsi dark brown, fore tarsi and inner surface of fore tibiae lighter; antennal segment III yellow, IV light brown, V slightly paler at extreme base; terminal setae of abdomen light brown. Structure of body and antennae very similar to dentipes and bicolor, but head more slender, pterothorax distinctly alatiform, metanotum not sculptured; antennal segment III with one sense cone, IV with two sense cones. MEASUREMENTS (holotype 9 in wm). Body length 2900. Head, length 330; median width 230; postocular setae 100. Pronotum, length 175; width across fore coxae 320. Metathorax total width 315. Tergite IX setae B, 170. Tube, length 210; basal width roo. Antennal segments length IIIJ—VIII length, 97; 97; 97; 80; 65; 45. ¢ aptera. Thorax paler than 9, pronotum yellow but shaded brown laterally, pterothorax and abdominal segment I almost clear yellow; head and posterior abdominal segments dark brown; all femora extensively yellow on inner surface, mid and hind tibiae and tarsi brown; antennal segments IV and V paler than in. Similar in structure to 9 but fore tarsus with a slender tooth and fore tibia with an apical tubercle. MEASUREMENTS (paratype gj in wm). Body length 1900 (body contracted). Head length 290. Tube, length 160; basal width go. SPECIMENS STUDIED. Holotype 9, ITALy (taken in quarantine at Detroit, U.S.A.): Stipa tenacissima, 27.x1.1961 (J. Fergusson 4087) (USNM). Paratypes. 5 2, 6g collected with the holotype (USNM, BMNH). CoMMENTS. Unfortunately this species is known only from rather poorly preserved specimens taken in quarantine. It is described here because of its interesting relationship to bicolor and dentipes, and because it is the only member of the genus with one sense cone on the third antennal segment. The females are very similar to dentipes but have a shorter tube, and some of the specimens have the pterothorax and femora pale in colour. Bolothrips pratensis Hood Bolothrips (Botanothrips) pratensis Hood, 1939 : 606-609. Holotype 9, U.S.A.: Texas (USNM) [not examined]. Despite the fact that the eyes are not prolonged on the ventral surface of the head, the present author does not agree with Hood that pratensis is closely related to the European and African icarus-group of species. The tube is particularly stout and the mesothoracic spiracles project as in gilvipes, and it is possible that pratensis is derived from forms with ventrally prolonged eyes. Most of the specimens listed below were collected on dead wood, in contrast to the type-series which came from grasses as is usual in Bolothrips. NESOTHRIPS COMPLEX 123 SPECIMENS STUDIED. U.S.A.; Texas, Palacios (type-locality), 1 9, 1 g paratypes, 28.iii. & 23.iv.1939 (J. D. Hood); Florida, Upper Matecumbe Key, 5 g, 10 2 from dead branches, xii. 1958 (Andre); F., Grassy Key, 1 g, 1 2 on dead branches, xii. 1958 (Andre); F., Cedar Keys, 1 g, I 9, vii. 1939 (P. Oman) (BMNH). Bolothrips rhachiphilus Cott Bolothrips (Bolothrips) rhachiphilus Cott, 1956 : 181-182. Holotype 9, U.S.A.: California (not known). Type-material of this species has not been deposited in the California Academy of Sciences (Arnaud & Lee, 1973) as was stated in the original description. The species appears to be related to the dark forms of gilvipes. Bolothrips schafferi (Thomasson & Post) comb. n. Nesothrips schafferi Thomasson & Post, 1966 : 31-32. Holotype 9, U.S.A.: North Dakota (INHS) [not examined]. This species is closely related to dentipes in the form of the antennae, head and tube, but differs in the colour of the legs and antennae. It is known only from western North America. SPECIMENS STUDIED. U.S.A.: North Dakota, Benson County, Wood Lake, 3 9 paratypes on Spartina pectinata, 27.viii.1964 (Thomasson & McBride) (BMNH & USNM); Oregon, 2 9, I g, 15.iv.1937 (?F. Andre) (BMNH). Bolothrips varius Hartwig Bolothrips varius Hartwig, 1948 : 110-112. Holotype 9, SoutH Arrica: Transvaal (Pretoria) {not examined]. The original description of this species noted that the ventral length of the eyes varied from ‘about equal to the dorsal length to about 1-4 times the dorsal length’. The eyes of inusularis from the Mediterranean region also vary considerably, and these two species are closely related. Hartwig lists material of varius from Cape Province, Basutoland, Natal and Transvaal. The specimen listed below from Kenya has a slender head as in varius and insularis, but the legs and antennal segments are pale as in icarus. It is possible that further collecting will produce intermediates between these named forms. SPECIMENS STUDIED. SoutH Arrica: Transvaal, Pretoria, 1 ¢ paratype on Panicum maximum, 13.11.1946 (E. K. Hartwig [data as for holotype]; 3 9, 1 g on P. maximum, 8.v.1948 (E. K. Hartwig) (BMNH). Kenya: 1 92 on Pinus, vii.1963 (BMNH). 124 L. A. MOUND 4 Fics 1-7. 1, Bolothrips italicus. 2, Carientothrips fijiensis. 3, C. mjobergi. 4, C. grayi. 5-7. Pelta (first abdominal tergite): 5, B.italicus. 6, C. fijiensis. 7, C. grayt. NESOTHRIPS COMPLEX 125 CARIENTOTHRIPS Moulton stat. n. Bolothrips (Carientothrips) Moulton, 1944 : 306. Type-species: Bolothrips (Carientothrips) jijiensis Moulton, by monotypy. This genus was placed as a synonym of Nesothrips by Stannard (1957) but is used here for a group of seventeen species from Australia and the Pacific, with one possibly unrelated form from South America. The macropterae of the type-species fijiensis are similar to a new species grays described below from New Guinea, and also to the macropterae of the mjobergi complex from Australia. The micropterae of fijiensis are very similar to the more slender forms loisthus and magnetis from Australia and New Zealand, but the micropterae of mjobergi are much shorter and broader and at first sight look very different. These are related to other species in Australia with short broad heads, miskoi and pedicillus. Within mjobergi there are forms with the eyes prolonged ventrally or with the eyes equal in size on the dorsal and ventral surfaces, moreover the legs also vary from yellow to brown, and there is no clear evidence that these different forms represent distinct species. The pelta of the macropterae is frequently characteristic with very slender lateral wings, but the pelta of micropterae and apterae is reduced and such specimens are very similar to Nesothrips species. These two genera intergrade in Australia, but Carientothrips is used for a group of species with the stylets relatively close together in the head. Two species with the stylets wide apart are also included from Australia, capricornis which seems to be related to semirufus, and badius which is not closely related to any other species. GENERIC DEFINITION. Medium sized, dark or pale species of Cryptothripini. Head longer than wide, or wider than long; eyes normal or reduced, sometimes longer on ventral than on dorsal surface; maxillary stylets usually parallel in middle of head and about one third of head width apart; ocellar setae usually well developed. Antennae eight-segmented, two sense cones on segment III, four sense cones on IV; segment VIII usually elongate. Fore tarsal tooth absent in 9 (except denticulatus), present ing. Pronotum transverse, oedymerous individuals rare; epimeral sutures usually complete. Wings frequently absent. Pelta frequently with slender lateral wings; tube with straight sides. KEY TO THE SPECIES OF CARIENTOTHRIPS 1 Antennal segment III with a shelf-like ring near the base (Text-fig. 60) : ; 2 Pedicel of antennal segment IIT simple 2 Thorax and legs yellow, rest of body and head brown; antennal segments ‘TI-IV largely yellow [Australia] . : . pictilis (p. 132) — Head light brown, thorax medium brown, abdomen dark, legs yellowish brown; antennal segments II-IV largely brown [Australia] ; : . acti (p. 126) 3 Metanotum with four to six pairs of setae medially in apterae, two pairs in macropterae ; epimeral sutures incomplete : : . . . 4 — Metanotum with one pair of median setae; epimeral sutures complete , . 6 4 Antennal segment III 2-5 times as long as wide, pedical stout bearing several transverse ridges (Text-fig 62) [Japan] : eae (p. 130) — Antennal segment III either longer or shorter, pedicel slender : 5 5 Antennal segment III 2-1 times as long as wide [Australia]. . ‘ reedi (p. 132) — Antennal segment III 2-8 times as long as wide [Australia] ‘ . vesper (p. 133) 6 Head broad, less than 1-2 times as long as wide ‘ : : > ° : 7 Head slender, more than 1-3 times as long as wide. ; : : . : 12 126 L. A. MOUND Body, legs and antennae dark brown except pedicel of III which is paler; pelta of macropterae and apterae rounded; eyes not extended on ventral surface of head [Australia] . z . casuarinae (p. At least antennal segments III_-v with yellow pedicels; frequently with legs, several antennal segments and anterior half of body pale Female with very small fore tarsal tooth at base of pretarsus; ocellar setae less than 20 wm long; antennal segment III stout but ey constricted at pedicel [Argen- tina and Falkland Islands] . , ‘ . denticulatus (p. Female without a fore tarsal tooth; ocellar setae more than 30 wm; antennal segment III tapering to base Setae B, on tergite IX about o- °5 times as long a as B, [Society Islands) biformis (p. Setae B, on tergite IX as long as or longer than B, [Australia] Racnual segments VI and VII broadly rounded with a sharply slender yellow pedicel : ; : pedicillus (p. Pedicel of antennal segments VI and VII less slender, VI frequently yellow in basal half . : Apical third of tube sharply yellow in contrast to black basal two- thirds: femora largely yellow. . tmiskoi (p. Apical third of tube paler than base but not sharply yellow: femora brown but tibiae yellow or brown ; : -mjobergi (p. Body and appendages uniformly dark brown except for pale base of antennal segment III; eyes reduced but rounded and protruding [Australia] : -. badius (p. Head and pronotum sometimes paler than abdomen, antennae or legs with extensive pale areas; eyes large, if reduced then not round and protruding Antennal segment IV shorter than segment III [Australia] . : na (. Antennal segment IV longer than segment III Setae B, and B, on tergite IX less than 0-5 times as long a as the tube, with apices clearly expanded; head and thorax ies abdomen brown, antennal segments I and II yellow [Australia] . . semirufus (p. Setae B, and B, on tergite IX more than 0'5 times as ; Jong: as tube, apices acute or blunt; head and thorax usually as brown as abdomen, if paler then antennal segment II brown Antennal segments III and IV almost clear yellow, also basal half of V; ‘segment IV about 3-5 times as long as wide; body colour brown, legs yellow [Australia] magnetis (p. Antennal segments III and IV variously shaded with brown; if pale then segment IV less than 3-0 times as long as wide; body and femora usually dark Eyes prolonged on ventral surface of head with one large isolated facet; dorsal surface of head and pronotum conspicuously reticulate (Text-fig. 4); antennal segments IV and V with elongate slender pedicels [New Guinea] , grayi (p. Eyes not longer ventrally than dorsally (Text-fig. 2); dorsal surface of head weakly sculptured; antennal segments IV and V with short pedicels (Text-fig. 61) Eyes reduced to about five facets ventrally; antennal segment III yellow in basal half, sometimes brown in apical half [Australia & New Zealand] . loisthus (p. Eyes with at least ten facets on ventral surface; antennal segment III pale at base and apex, but with a transverse brown band at basal third [Fiji] . fijiensis (p. Carientothrips acti Mound (Text-fig. 60) Carientothrips acti Mound, 1974 : 25-26. Holotype 9, Austratia (ANIC) [examined]. 131) 129) 17 130) 128) The acti group of species includes japonicus (Bagnall), pictilis Mound, reedi NESOTHRIPS COMPLEX 27 Mound and vesper Mound. Both acti and pictilis are unusual in having the terminal setae on the tube flattened at the apex instead of acute, and in having the pedicel of antennal segment three flared into a ring. However, japonicus has the pedicel of segment three slightly thickened, and this probably represents an intermediate condition. These species, apart from fictilis, have the epimeral sutures incomplete, and the metanotum bears more than one pair of median setae. SPECIMENS STUDIED. Holotype 9, AUSTRALIA: Australian Capital Territory, Canberra, Black Mt, on dead leafy Eucalyptus branches, 3.i.1961 (E. M. Reed) (ANIC). Paratypes from New South Wales, A.C.T. and northern Victoria from leaf litter and grass tussocks are listed in detail in Mound (1974). Carientothrips badius (Hood) comb. n. Cryptothrips badius Hood, 1918 : 143-144. Holotype 9, AUSTRALIA: Queensland (USNM) [examined]. Elaphrothrips apterus Girault, 1928 (42):2. Holotype g, AUSTRALIA: Queensland (QM) [examined]. [Synonymized by Mound, 1974 : 23]. Bolothrips badius (Hood) Hartwig, 1948 : 117. This species was treated in the genus Bolothrips by Mound (1974), but is now removed from that genus because the fourth antennal segment bears four sense cones. However, it is an unusual species with a relatively long head and small rounded eyes which is not closely related to any other known species. C. badius is widespread in Australia in grass tussocks, and a detailed list of specimens from Queensland, New South Wales, and Western Australia is given in Mound (1974). Carientothrips biformis (Moulton) comb. n. Bolothrips biformis Moulton, 1939 : 146-147. Holotype 9, Tanit1 (BPBM) [not examined]. This species is removed from Bolothrips because it has four sense cones on the fourth antennal segment. It is not closely related to the other species of Cariento- thrips but has the maxillary stylets closer together than the species of Nesothrips. SPECIMEN STUDIED. SocrETy IsLranps: Rapa, Mangaoa Pk, 1000-1200 ft, 25.vii.1934 (E. C. Zimmermann), I Q microptera paratype (BMNH). Carientothrips capricornis (Mound) comb. n. Bolothrips capricornis Mound, 1974 : 23-24. Holotype 9, AUSTRALIA: Queensland (ANIC) [examined]. This species was described in Bolothrips because of its resemblance to the American species B. pratensis. However, the fourth antennal segment bears four sense 128 L. A. MOUND cones unlike any species of Bolothrips, and capricornis is probably most closely related to Carientothrips semirufus. SPECIMENS STUDIED. Holotype 2 microptera, AUSTRALIA: Queensland, Townsville, 2 miles west of Saunders Beach, in leaf litter, 21.vii.1968 (L. A. Mound) (ANIC). Paratype 9 collected with the holotype (BMNH). Carientothrips casuarinae Mound Carientothrips casuarinae Mound, 1974 : 26-29. Holotype 9, AUSTRALIA: New South Wales (ANIC) [examined]. This species, which is known from macropterae as well as apterae, is dark brown but is otherwise similar to semirufus. It has been taken on dead Casuarina branches at several sites in eastern Australia. SPECIMENS STUDIED. Holotype 9 aptera, AUSTRALIA: New South Wales, Normanhurst (15 miles north of Sydney), on dead Casuarina foliage, 1.i1.1959 (E. M. Reed) (ANIC). Paratypes g and 2 macropterae from New South Wales listed in detail in Mound (1974). Carientothrips denticulatus (De Santis) comb. n. Nesothrips (Bolothrips) denticulatus De Santis, 1963b : 66. Holotype 9, ARGENTINA: Tierra del Fuego (Museos de La Plata) [not examined]. This species is removed from Bolothrips because it has four sense cones on the fourth antennal segment. It is not close to Nesothrips species because the stylets are parallel in the middle of the head, but it seems unlikely to be closely related to the other species of Carientothrips. Some of the specimens listed below were identified by Prof. Luis de Santis as denticulatus. The fore tarsal tooth of the female is less than I0 pm long, although that of the male is about 30 ym long. SPECIMENS STUDIED. EAST FALKLAND ISLANDs: Kidney Is., 55 9, 24 ¢ in Poa flabellatum tussocks, 29.x1.1967 (E. Holdgate) (BMNH). Carientothrips fijiensis (Moulton) comb. n. Bolothrips (Carientothrips) fijiensis Moulton, 1944 : 306-307. Holotype 9, Fiji (BPBM) {not examined]. ; The original description refers to an apterous paratype female, although this specimen is actually micropterous with wing lobes 160 wm long. The eyes of the microptera are reduced and flattened externally as in loisthus, although the eyes _ of the macropterae are large and rounded as in grayi described below. The Australian species mjobergi is similar to the macropterae of fijtensis although with a NESOTHRIPS COMPLEX 129 shorter head, but the apterae and micropterae of mjobergi are much shorter and broader. The pelta of fijiensis is similar to that of grayi, and the male has a small fore tarsal tooth. SPECIMENS STUDIED. Fiji: Vitilevu, Nandarivatu, 2 9, r ¢ macropterae, I 2 microptera (all paratypes), g.ix.1938, I Q macroptera paratype I.ix.1938; Mt Korobamba, 1 2 macroptera paratype I.viii.1938 (E. C. Zimmermann) (CAS). Carientothrips grayi sp. n. (Text-figs 4, 7, 63) Q macroptera. Colour dark brown with red internal pigment; tarsi light brown; antennal segments IV and V yellow in basal third, III with yellow pedicel and yellowish brown in apical half otherwise dark brown; fore wings deeply shaded in third and fourth fifth, but clear in apical fifth, basal two-fifths clear with dark longitudinal stripe; major setae almost hyaline, bases of setae at apex of tube dark. Head longer than wide, projecting in front of large compound eyes, cheeks rounded to base with about four small setae; dorsal surface strongly sculptured; postocellar setae well behind ocelli; maxillary stylets retracted to postocular setae; eyes with one large ventral facet dis- placed posterior to normal hind margin of eye. Antennae with segment III much shorter than IV; segment VII with a distinct pedicel, VIII slender and slightly constricted at base. Prono- tum similarly sculptured to head, with weak median thickening; epimeral sutures complete, setae with blunt apices. Metanotum reticulate, median setae slender, 30 um long. Fore wing slightly constricted medially; subbasal setae short, duplicated cilia variable. Fore tarsus with no tooth. Pelta reticulate with pronounced lateral wings; setae on tergite IX softly rounded at apex; tube slender with straight sides bearing fine setae. MEASUREMENTS (holotype 9 in wm). Body length 3250. Head, length 350; median width 260; postocular setae 65; ocellar setae 35. Pronotum, length 195; median width 320; major setae -am 30, aa ?30, ml ?50, epim I00, pa 70. Fore wing, length 1400; distal width 100; subbasal setae 40, 45, 65; number of duplicated cilia 4-7 (3-6 in one paratype). Tergite VII setae, B, 70; B, 100. Tergite IX setae, B, 160; B, 190; B,; 155. Tube, length 290; basal and apical width 90, 45; terminal setae 175. Antennal segments length, 55; 65; 95; 125; 120; 77; 58; 35; semse cones on III 75. SPECIMENS STUDIED. Holotype 9, NEw GuINEA: Wau, under bark of Avaucaria cunninghamiana, 22.1x.1970 (B. Gray) (BMNH). Paratypes, NEw GuINEA: Wau; I 9, 9.i.1966 (J. Sedlacek); 1 9, 6.x.1966 (G. Samuelson) (BPBM). CoMMENTs. This species is very similar to the macropterae of jijiensis, although larger and with the eyes prolonged ventrally. In the BPBM collection there is a female from north-western New Guinea, Lake Wissel, with almost uniformly brown antennae and no duplicated cilia on the fore wings, but otherwise very similar to grayi. This may represent a further species. The micropterae of Jijiensis are more slender and very similar to loisthus, although a direct comparison of loisthus and grayi would not suggest any close relationship between these two species. 130 L. A. MOUND Carientothrips japonicus (Bagnall) comb. n. (Text-fig. 62) Cryptothrips japonicus Bagnall, 1921 : 355-356. Holotype 9, JAPAN (BMNH) [examined]. Mystrothrips japonicus (Bagnall) Mound, 1968 : 138. The unique holotype of this species is rather damaged but is difficult to distinguish from the eastern Australian species veedi Mound. The third antennal segment is thicker at the base, suggesting the origin of the condition found in acti Mound. The apices of the terminal setae on the tube are flattened in acti and pictilis Mound, but are acute as in most other thrips in japonicus, reedi and vesper. SPECIMEN STUDIED. Holotype 2, JAPAN: Kobe, 23.vill.1g16 (Lewis) (BMNH). Carientothrips loisthus Mound Carientothrips loisthus Mound, 1974 : 29-30. Holotype 9, Austratta: South Australia (ANIC) [examined]. This species was based on a single female aptera collected in South Australia, and at that time the holotype was regarded as a distinct species from specimens collected in New Zealand. However there is more difference between the specimens listed below from various localities within New Zealand than there is between some of these specimens and the Australian holotype. For this reason they are all regarded as one species. Large specimens from both North and South Island, New Zealand have the femora brown, whereas the females from Waiwera have the femora almost as yellow as the tibiae. Moreover these small specimens have the head and thorax much paler than the abdomen. The large specimens have very long pronotal setae with acute apices, whereas the small specimens have the epimeral and midlateral setae bluntly rounded at the apex. The holotype is completely apterous but has three ocelli, whereas the New Zealand specimens have a small lobe (15 wm) attached to the lateral corners of the mesonotum, and the lateral ocelli are scarcely indicated. The males are smaller than the females with almost clear yellow legs and a large fore tarsal tooth. MEASUREMENTS (large New Zealand 2 in wm). Body length 2700. Head, length 310; median width 225; ocellar setae 85; postocular setae 135; mid-dorsal setae 90-120. Pronotum, length 175; median width 290; major setae — am 80, aa 100, ml 120, epim 130, pa 135. Tube length 195. SPECIMENS STUDIED. Holotype 9, AUSTRALIA: South Australia, Adelaide, Waterfall Gulley, grass by stream, 14.1.1968 (L. A. Mound) (ANIC). NEw ZEALAND: South Island, Greymouth, 1 9 on Juncus, 12.ix.1972 (V. F. Eastop); North Island, Levin, 2 9, ix. 1966, I gf, v. 1966, I 9, iv. 1967 (Manson & Pritchard); N.I., near Rotorua, 2 9, ix. 1972 (V. F. Eastop); N.I., Waiwera, 4 9, NESOTHRIPS COMPLEX 131 I g at base of grasses, viii. 1968 (L. A. Mound); N.I., Helensville, 1 3, viii. 1968 (L. A. Mound) (BMNH). Carientothrips magnetis Mound Carientothrips magnetis Mound, 1974 : 30-31. Holotype 9, AusTRALIA: Queensland (ANIC) [examined]. This species has the eyes well developed ventrally, but is intermediate between loisthus and the acti-group of species in the form of the head and epimeral sutures. SPECIMEN STUDIED. Holotype 9, AUSTRALIA: Queensland, 10 miles north of Townsville, on grasses, 20.vii.1968 (L. A. Mound) (ANIC). Carientothrips miskoi Mound Carientothrips miskoi Mound, 1974 : 31. Holotype 9, AusTRALIA: New South Wales (ANIC) [examined]. This species is known only from three females which were collected on two separate occasions at the same site. It differs from mjobergi mainly in having the tube brightly bicoloured. SPECIMENS STUDIED. Holotype 2 aptera, AUSTRALIA: New South Wales, Cabbage Tree Creek, 7 miles north of Nelligen, near Bateman’s Bay, in leaf litter, 2.ii.1969 (S. Misko) (ANIC). Carientothrips mjobergi (Karny) Cryptothrips mjobergi Karny, 1920 : 42. Holotype 9, AUSTRALIA: Queensland (NR) [examined]. Cryptothrips (?) australicus Priesner, 1928b : 649-651. Holotype 9, AUSTRALIA: Queensland (HP) [examined]. [Synonymized by Mound, 1974 : 31.] Cryptothrips incisus Girault, 1927 (37): 1. Syntypes 2 9, AUSTRALIA: Queensland (QM) [examined]. [Synonymized by Mound, 1974 : 31.] Bolothrips (Bolothrips) flavitibia Moulton, 1968 : 117-118. Holotype 9, AUSTRALIA: Queens- land (CAS) [examined]. [Synonymized by Mound, 1974 : 31.] This species is widespread in Australia and highly variable. The eyes are frequently prolonged ventrally, although this is not true of all populations. The tibiae are usually brown but sometimes yellow, and similarly the colour of the fore wings and antennae apparently varies. The number of duplicated cilia on the fore wing has been found to vary from two to eight even within a single population. The pelta of the macropterae is very similar to that of macropterae of fijiensis and grayi, but the distinction between a central body and lateral wings is less evident in the pelta of micropterae and apterae. Numerous specimens of this species, or species complex, have been studied from all the States of Australia except Tasmania and the Northern Territories, mainly from Eucalyptus dead leaves and branches. 132 L. A. MOUND Carientothrips pedicillus Mound Carientothrips pedicillus Mound, 1974 : 32-33. Holotype 9, Austratia (ANIC) [examined]. This species is known only from the holotype. It is very similar to mjobergi but has the antennal segments six and seven sharply constricted to a narrow pedicel. SPECIMEN STUDIED. Holotype 2 aptera, AUSTRALIA: Australian Capital Territory, Mt Gingera, in leaf litter, 13.iv.1967 (E. B. Britton) (ANIC). Carientothrips pictilis Mound Carientothrips pictilis Mound, 1974 : 33-34. Holotype 2 Australia: New South Wales (ANIC) [examined]. The unique holotype of this species is bicoloured as in some species of Bolothrips. However, the fourth antennal segment has four sense cones and the maxillary stylets are parallel in the middle of the head. SPECIMEN STUDIED. Holotype 9, AUSTRALIA: New South Wales, Parkes, from Eucalyptus blakelyi, 7.Vili.1959 (E. M. Reed) (ANIC). Carientothrips reedi Mound Carientothrips reedi Mound, 1974 : 34-35. Holotype 9, AustraL1a (ANIC) [examined]. This species was described from apterae and one macroptera collected in New South Wales and Australian Capital Territory, as well as one macroptera from southern Queensland. This may be a widespread species because the unique holotype of japonicus Bagnall only differs in the detailed structure of the antennae. These two species are related to acti Mound in the form of the head and thorax. SPECIMENS STUDIED. Holotype 9 aptera, AUSTRALIA: Australian Capital Territory, Canberra, Black Mt, in grass tussock, 11.xii.1960 (E. M. Reed) (ANIC). Paratypes from New South Wales, A.C.T., and Queensland are listed in Mound (1974). Carientothrips semirufus (Girault) Elaphrothrips semirufus Girault, 1928(42) : 4. Holotype 9, AustTratia: Victoria (QM) [examined]. Carientothrips semirufus (Girault) Mound, 1974 : 35. This apterous species is very similar to the bicoloured specimens of Jotsthus from New Zealand, but has short expanded major setae and the mesonotum is closely transversely striate. NESOTHRIPS COMPLEX 133 SPECIMENS STUDIED. Holotype 9, AUSTRALIA: Victoria, Melton, in tussocks, v. 1928 (F. E. Wilson) (QM). AUSTRALIA: New South Wales, 13 9, 10 ¢ all collected south-east from the Blue Mountains and detailed in Mound (1974) (BMNH, ANIC). Carientothrips vesper Mound Carientothrips vesper Mound, 1974 : 35-36. Holotype 9, AUSTRALIA: Western Australia (ANIC) [examined]. The unique holotype of vesper is very similar to the apterae of reed: but with longer antennae and shorter expanded setae on tergite nine. SPECIMEN STUDIED. Holotype 2, AUSTRALIA: Western Australia, Manjimup, in grass clump, 28.ix.1967 (L. A. Mound) (ANIC). DICERATOTHORIPS Bagnall Diceratothrips Bagnall, 1908 : 193. Type-species: D. bicornis Bagnall, by monotypy. This genus has been confused with Dichaetothrips (Mound, 1968 : 78), but is best restricted to a group of Neotropical species. Most of these species have a pair of elongate anteocellar setae, although in some of the smaller species such as harts Hood, delicatulus Hood, and setigenis Hood these setae are reduced. It is to this group of species that Gastrothrips validipennis Hood is related. Large species of Diceratothrips have a series of stout spines on the inner surface on the fore femora, at least in the males. The antennae are frequently distinctive, segment three is relatively long, the sense cones on segments three and four unusually short, and segments seven and eight frequently form a single elongate unit. The pronotum is also unusual with no thickening at the anterior margin and very little medially even in large individuals, moreover the anterior third is reticulate and the anterior setae short. Diceratothrips validipennis (Hood) comb. n (Text-figs 9, 66) Gastrothrips validipennis Hood, 1938 : 403-406. Holotype 9, U.S.A.: Florida (USNM) [not examined]. This is a small species of Diceratothrips with the anteocellar setae exceptionally reduced. SPECIMEN STUDIED. U.S.A.: Florida, Homestead, 1 paratype 3, on dead branches, 28.xii.1937 (J. C. Bradley) (AMG). 134 L. A. MOUND DICHAETOTHAHRIPS Hood Dichaetothrips Hood, 1914 : 164-165. Type-species: D. brevicollis Hood, by monotypy. The synonymy of this genus and its relationship to Scotothrips have been discussed recently by Mound (1974). The Indian species referred to below can be distinguished from Nesothrips by the presence of a large fore tarsal tooth in the female as well as the more elongate tube. It is returned to the genus Dichaetothrips on account of the elongate postocellar setae. Dichaetothrips gloveri (Ramakrishna & Margabandhu) comb. rev. (Text-fig. 8) Neosmerinthothrips gloveri Ramakrishna & Margabandhu, 1939 : 31-32. LECTOTYPE 9 Inp1a (TNA), here designated [examined]. Dichaetothrips gloveri (Ramakrishna & Margabandhu) Shumsher, 1947 : 204-205. Nesothrips gloveri (Ramakrishna & Margabandhu) Ananthakrishnan, 1960 : 32-33. The original description of this species does not designate a holotype, although Shumsher (1947) refers to a holotype in the Zoological Survey of India. The specimen here designated as lectotype bears the original data and is the only member of the type-series still available for study. The postocellar setae of the lectotype are 55 um long, rather longer than the distance between the anterior and posterior ocelli. The fore tarsal tooth is slender and almost as long as the width of the fore tarsus. Unfortunately the specimen is crushed, but in the opinion of the present author further collecting is likely to show that this individual is a small specimen of the species known as Dichaetothrips usttatus Ananthakrishnan & Jagadish, 1970 (= Dichaetothrips indicus Ananthakrish- nan, 1961). SPECIMEN STUDIED. Lectotype 9, INDIA, Numkum, Ranchi, on Zizyphus jujuba, 1933 (TV R450) (TNA). GASTROTHRIPS Hood gen. rev. Gastrothrips Hood, 1912 : 156. Type-species: G. ruficauda Hood, by original designation. Probolothrips Moulton, 1941 : 319. Type-species: P. hambletoni, by monotypy; treated here as a synonym of abditus. [Synonymized by Hood, 1952 : 163.] The genus Gastrothrips is used here for a group of nineteen species, mainly from the Neotropics, which have only three sense cones on the fourth antennal segment. The tube is variable in shape, either with straight sides or constricted apically, and frequently much paler than the rest of the abdomen. In contrast the Neo- tropical species related to nigrisetis Hood have four sense cones on the fourth antennal segment (with the exception of paulistarum), and the tube is black, frequently with convex sides. These species are treated here under Neosmerinthothrips. The Gastrothrips species which have a straight-sided tube comprise a single NESOTHRIPS COMPLEX 135 Fics 8-12. 8, Dichaetothrips gloveri. 9, Diceratothrips validipennis. 10-12. Gastrothrips anolis: 10, large male. 11, female. 12, pelta of female. 136 L. A. MOUND species from India, falcatus Ananthakrishnan, and a complicated species-group with named forms distributed throughout the tropics, the acuticornis-group. All the other Gastrothrips species are known only from the Neotropics with a few species extending into temperate latitudes. A major problem with species recognition is the few species that are known from a series of specimens at more than one site. Moreover several species are known from only one morph, aptera, microptera or macroptera, and are therefore difficult to compare with each other. Cryptothrips citriceps Priesner, 1921 may also belong in this genus but has not been studied. GENERIC DEFINITION. Medium sized, usually dark species of Cryptothripini. Head usually rectangular, longer than wide and not projecting in front of eyes; eyes large and usually equally developed on dorsal and ventral surfaces; head rarely wider than long, rarely with cheeks bearing stout setae. Antennae eight-segmented, VIII usually elongate and slender, rarely short and broad; segment IV with three sense cones, III with one or twosense cones. Pronotum transverse, sometimes enlarged in major males, anterior setae usually small, epimeral sutures frequently not complete. Fore tarsal tooth present in g, present or absent in 9. Wings present or absent, duplicated cilia present or absent. Mesonotal anterior angles frequently projecting in major males; metanotal median setae usually well developed. Pelta usually well defined, triangular, with small lateral wings. Tube with straight sides evenly narrowing to apex, or with apex sharply constricted; never with sides strongly convex and not constricted at apex; frequently yellow or paler than rest of abdomen. Males commonly oedymerous. KEY TO THE SPECIES OF GASTROTHRIPS 1 Tube with straight sides, not sculptured nor sharply constricted close to terminal setae (Text-fig. 16); tube dark at least in basal half. A ' 2 — Apex of tube sharply constricted in front of terminal setae, tube often sculptured (Text-figs 17-19); basal half of tube frequently yellow or paler than tergite IX . 3 2 Antennal segment VIII broad at base, scarcely 0-5 times as long as VII (Text-fig. 64) ; pelta with a small central body and slender lateral wings; fore wings without duplicated cilia . : ‘ - falcatus (p. 143) - Antennal segment VIII slender, more > than O°7 times ¢ as long as VII; pelta without slender lateral wings; fore wings with duplicated cilia acuticornis-group (see acuticornis (p. 139), mongolicus (p. 145), procerus (p. 147), & subulatus (p. 148)) 3 Basal half of tube yellow or reddish brown, paler than abdominal segment IX ; 4 — Tube black or dark brown throughout, not paler than tergite IX ‘ : 9 4 Antennal segments I-III largely yellow, paler than the head; tube bright yellow ; 5 — Antennal segments I-III brown, as dark as head; tube reddish atl to reddish brown : 7 5 Pelta small and semicircular, occupying less than 0°5 ‘of anterior margin of tergite II (Text-fig. 12); ag eae ae 6 with pair of elongate tubercles on posterior margin of pronotum : . anolis (p. 142) — Pelta broadly triangular, occupying at least two- thirds of anterior margin of tergite II (Text-fig. 20) . : 6 6 Antennal segments I and II yellow i in sharp contrast to brown segments IV-VIII: head of ? with cheeks evenly narrowing to base (Text-fig. 25); 9 with a fore tarsal tooth ; ‘ . intonsus (p. 145) -— Antennal segments grade ‘evenly from yellow iE to brown VIII, even VII has a pale pedicel; head of 2 with cheeks asia except for slight constriction near base; 9 without a fore tarsal tooth . . fulvicauda (p. 143) 7 Head slightly wider than long (210 um 195 pm); mid and hind femora brown but with a yellow area on distal posterior margin . . . . ruficauda (p. 147) NESOTHRIPS COMPLEX 137 14 Fics 13-22. 13-19. Tube (last abdominal segment): 13, Bolothrips italicus. 14, Gastrothrips intonsus. 15, G. mandiocae. 16, G. falcatus. 17, G. callipus. 18, G. abditus. 19, G. ruficauda. 20-22. Pelta: 20, G. intonsus. 21, G. mandiocae. 22, G. ruficauda. 138 L. A. MOUND — Head longer than wide; mid and hind femora uniformly brown . 8 8 Head less than 1-15 times as long as wide; tube more than o-g times as long as s head corvus (p. 142) — Head more than 1-25 times as long as wide; tube less than 0-75 times as long as head alticola (p. 140) g Legs and antennal segments I-VI largely yellow; tube 0-7 times as long as head; apterous with metanotal setae fine and less than 15 ymlong . fulviceps (p. 144) — Legs and antennal segments largely dark; tube 0-8 times as long as head or longer; macropterous or micropterous with metanotal setae more than 30 wm long and usually stout : i é : . ; : : E : Io 10 Fore wing with duplicated cilia : : : : : : ; II — Fore wing without duplicated cilia, or micropterous : 13 11 Female without a fore tarsal tooth; postocellar setae less than the diameter of one ocellus i ; . abditus (p. 138) — Female with a fore tarsal tooth: - postocellar setae asually at least 40 pm long, ela twice as long as the diameter of one ocellus_ . . 12 12 Antennal segment III brown with yellow pedicel; sides of tube weakly convex, curving from base to apex; pronotal anteromarginal setae less than half as long as anteroangulars; median part of pelta narrowly triangular : . stygicus (p. 147) — Antennal segment III largely yellow, shaded brown at apex; sides of tube parallel medially or even weakly constricted in anterior third; pronotal anteromarginal setae as long as anteroangulars; median part of pelta broadly triangular mandiocae (p. 145) 13 Fore tarsal tooth of 2 acute, almost as long as width of fore tarsus . texanus (p. 148) — Female without a fore tarsal tooth . : 14 14 Femora brown with yellow area distally on inner margin; sides of tube weakly convex, curving from base to apex . ‘ . callipus (p. 142) — Femora uniformly brown without a clear yellow area; tube parallel to weakly constricted medially, or evenly tapering with sides not rounded : 15 15 Micropterous; tube tapering from base to near apex . ‘ _monticola (p. 146) — Macropterous; tube parallel or weakly constricted medially . fumipennis (p. 144) Gastrothrips abditus Hood (Text-fig. 28) Gastrothrips abditus Hood, 1935 : 177-182. Holotype 9, PANAMA: Barro Colorado (USNM) [not examined]. Hoplothrips brasiliensis Moulton, 1938 : 378-379. Holotype 9, Brazit (CAS) [not examined]. [Synonymized by Hood, 1952 : 163.] Probolothrips hambletoni Moulton, 1941 : 320-321. Holotype 9, Braziz (CAS) [not examined]. [Synonymized by Hood, 1952 : 163.] Hood described this species from six female and one male macropterae collected from burrows in dead branches. None of these have been studied by the present author, but specimens from the type-series of Moulton’s two species have been seen. Apparently abditus is widespread in eastern South America, and it is closely related to mandiocae and texanus. There are two undescribed species in the USNM from Mexico which are also related to abditus. One of these is macropterous with a heavy tube and dark brown third antennal segment. The other species includes the only hemimacropterous Gastrothripbs known to the author, and these may NESOTHRIPS COMPLEX 139 represent abditus despite the fact that the wing lobes are shaded. The macropterae of abditus have the fore wings pale or only weakly shaded. SPECIMENS STUDIED. BRAZIL: Minas Gerais, on dead Manioc stems, 3 9, I g, 2I.vil.1933, 2 g, 10.1V.1933 (Hambleton) (BMNH) [type data of brasiliensis and hambletoni]. PARAGUAY: Caaguazu, I ¢ swept, 1.11.1964 (Ff. Andre). TRINIDAD: St Augustine, 1 J in empty Psychidae case, 1.xi.1970 (L. A. Mound) (BMNH). Gastrothrips acuticornis (Hood) Cryptothrips acuticornis Hood, 1925: 65. Holotype 9, West Inpizs: St Croix (USNM) [examined]. Cryptothrips cybele Girault, 1927(38) :1. Holotype J, AUSTRALIA; Queensland (QM) [examined}. [Synonymized by Mound, 1974 : 53.] Gastrothrips noumeae Bianchi, 1945 : 251-254. Holotype 9, New CaLEeponia (BPBM) [not examined]. [Synonymized by Mound, 1974 : 53.] Nesothrips acuticornis (Hood); Ananthakrishnan, 1969 : 181~182. This species has a remarkable distribution and, moreover, it is difficult to distin- guish from three other described species: mongolicus from Mongolia, procerus from Brazil and subulatus from Transvaal. Hood described acuticornis from several specimens collected on St Croix and Barbados, and in the description refers to small pale males. Ananthakrishnan (1969) recorded both sexes from southern India, and Mound (1974) compared Indian specimens with the West Indian holotype and also specimens from New Caledonia and Australia. The Australian specimens have the eighth antennal segment slightly shorter than the seventh in contrast to the other material studied, and moreover the third antennal segment of most . of the Australian specimens is paler than that of either the holotype or the Indian . material. However, one female from Kuala Lumpur in Malaya has the third antennal . segment dark except for the pedicel as in the holotype, but the eighth segment is short as in the Australian specimens. For these reasons cybele and noumeae are regarded as the same species as acuticornis, despite the fact that the males which have been studied from Australia and New Caledonia are macropterous and brown like the females, whereas the males from India and the West Indies are yellow and micropterous. Large males from India have the posterior margin of the meta- notum produced into a dentate fringe which is excavate medially. The females of acuticornis are macropterous with short postocellar setae, and the head of each of the specimens which have been studied is about I-I times as long as the tube. However, there are two micropterous females from Basutoland in the Albany Museum, Grahamstown which may represent this species but one has both postocellar setae 30 »m long and the other has these setae 12 ym and 30 pm long. The single male collected with these specimens is also micropterous and slightly oedymerous but has a simple posterior border to the metanotum unlike the small yellow males from India. Whilst considering the variation in acuticornis it is necessary to refer to three other similar species. The South American species procerus has a relatively short Cc 140 L. A. MOUND tube, the head being 1-5 times as long as the tube in all three of the available speci- mens. The holotype and one male from southern Brazil have a more slender head than a female from Paraguay, but all three specimens have the third antennal segment light brown with a yellow pedicel, and none of them have elongate posto- cellar setae. The South African species subulatus has the head about 1-4 times as long as the tube and the postocellar setae less than 15 ym long, but the third antennal segment is yellow and the fourth variable yellowish brown. Finally, mongolicus has the head 1-2 times as long as the tube, the third antennal segment is yellow in contrast to the dark brown fourth segment, but the postocellar setae are 50 wm long. These three species are known from too few specimens to assess the signifi- cance of these differences at present. The form of the pelta differs between them but this may be a function of body size. It is possible that all three are only local populations of acuticornis. SPECIMENS STUDIED. Holotype 2 macroptera of acuticornis, WEST INDIEs: St Croix, swept from grass, g.ii.1915 (C. B. Williams 553) (USNM 71992). Holotype 3 macroptera of cybele, AUSTRALIA: Queensland, Gympie, in forest, 29.vi.1924 (A. A. Girault). AUSTRALIA: Queensland, Laidley, 1 3, 1 @ in forest, 31.v.1923 (A. A. Girault); Q., Grandchester, 1 2 (labelled as 3) in forest, I.viii.1g24 (A. A. Girault) (QM); Q., Moura, 2 9 on Rhodes grass, 10.x.1967 (Page & Rigby); Q., 100 miles north of Rockhampton, I ¢ in grass, 18.vii.1968 (L. A. Mound) (BMNH). NEw CALEDONIA: near Noumea, 2 9, I g paratypes of noumeae, beaten from grass, 24.ix.1940 (FF. X. Williams) (BMNH, BPBM, HP). _ West Mataysia: Kuala Lumpur, 1 9 from live tree, 26.xii.1969 (F. Andre) (BMNH). Inop1a: Madras, 4 gf on Barteria sp. twigs, 18.vii.1969, 3 2 on dry Jasmine twig, 25.vii.1966, 1 9 from grass, 23.iii.1968; Palghat, 1 9, 1 J on dry twig, 12.v.1970; Kumili, 1 9 on dry twig, 22.vii.1969; Ronningtonn, 1 9 on dry twig, 8.11.1969 (T. N. Ananthakrishnan) (BMNH). Gastrothrips alticola Hood Gastrothrips alticola Hood, 1942 : 570-573. Holotype 9, Peru (USNM) [not examined]. This species was described from thirty-five apterous females and ten apterous males. Hood states in the description ‘basal three fourths of tube blackish brown’, whereas the paratypes which have been studied here have the basal three-fourths of the tube paler than tergite nine. This has made comparison with other species particularly difficult. In the key to species above alticola will run to monticola if the tube is considered to be black. However, monticola is known only from micropterae and alticola only from apterae. However, both forms were collected in the Department of Huanuco, Peru, at 3000 metres. Moreover, in the BMNH collections there is a single macropterous female which was collected within 100 miles of Huanuco. This female has the basal three fourths of the tube dark reddish brown and paler than tergite nine, the fore wing bears six duplicated cilia, and the NESOTHRIPS COMPLEX 141 fore tarsus does not bear a tooth. This specimen probably represents alticola, and monticola may also be the same species. SPECIMENS STUDIED. PERU: Dept. Huanuco, Shishmay, 2 2 paratypes from shrub with dry branches, at 3000 m, 18.ix.1937 (fF. Woytkowski); Panao, 1 2, 1 g from dead branches and leaves at 3000 m, 10.ix.1937 (F. Woythkowski) (AMG). Fics 23-28. 23-26. Gastrothrips intonsus : 23, fore tarsus of small male. 24, small male. 25, female. 26, fore tarsus of female. 27, G. ruficauda. 28, G. abditus. 142 L. A. MOUND Gastrothrips anolis Morgan (Text-figs 10-12) Gastrothrips anolis Morgan, 1925 : 7-8. Holotype 9, PuErto Rico (USNM) [not examined]. Gastrothrips proteus Hood, 1933 : 417-419. Holotype 9, Panama: Barro Colorado (USNM) [not examined]. [Synonymized by Hood, 1935 : 186.] Morgan described this species from one female, and Hood described proteus from one macropterous and seventeen apterous females with seven apterous males. The tube is bright yellow as in fulvicauda, intonsus and ruficauda, and the head of the major males has spiny cheeks as in intonsus. However, the females and the smallest males have a quadrate head similar to several other species of Gastrothrips. The major males bear two elongate tubercles on the posterior margin of the pronotum which overhang the metanotum, and Hood (1935: 185) has published drawings of the variation of this species. SPECIMENS STUDIED. PANAMA: Barro Colorado Is., Gatun Lake, paratypes of proteus; 1 9, 1 3 on dead branches, 27.vii.I933, I 9 on dead grass and leaves, I.vii.1933, I ¢ on grass, 14.vil1.1933 (J. D. Hood) (AMG). CAyMAN ISLANDS: 2 9, 2 @ apterae in Nightingale’s nest, 16.vii.1962 (A. Ventura) (BMNH). Gastrothrips callipus Hood (Text-fig. 17) Gastrothrips callipus Hood, 1935 : 182-186. Holotype9, U.S.A., Texas (USNM) [not examined]. Hood described this species from three female and one male macropterae from Texas, but the species is apparently widespread in the south-eastern part of North America. The femora are bicoloured as in the North American species ruficauda and the Central American species anolis; however, the tube is black with weakly convex sides. The male listed below from Mexico is identified as callipus with some doubt because it is micropterous with the basal three antennal segments yellowish brown. SPECIMENS STUDIED. U.S.A.: Texas, Victoria, paratype 2 on Chenopodium, iv. 1908 (J. D. Mitchell) (USNM); T., Sinton, r 2 on dead branches, 20.xii.1967; T., San Antonio, I 2 on dead branches, 23.x1i.1967 (F. Andre); Oregon, Ashland, 1 9 in berlese funnel, 6.xii.1937 (BMNH); South Carolina, Charleston, 3 2 on Solidago, 15.vi.1943 (W. S. Fields); Florida, Hialiah, 1 2 on grass and weeds, 15.iv.1969 (C. E. Stegmaier). BAHAMA Is.: West Palm Beach, I 9, 13.v.1966 (W. E. Browning. MExIco: in quarantine at Nogales, 1 J on tomato fruit, 15.11.1941 (USNM). Gastrothrips corvus Priesner Gastrothrips (?) corvus Priesner, 1933 : 55-57. Holotype 9, Mexico (HP) [examined]. NESOTHRIPS COMPLEX 143 Gastrothrips capitalis Hood, 1935 : 174-177. Holotype 9, U.S.A.: Texas (USNM) [examined]. Syn. n. The two unique holotypes of corvus and capitalis have been compared with each other and capitalis is here regarded as a small macroptera whereas the corvus holotype is a large microptera. The other differences noted by Hood follow from this morph difference, but it should be noted that in both specimens the basal two-thirds of the tube is dark reddish brown in contrast to the black colour of segment nine. The most closely related species appears to be monticola from Peru which has a dark tube. SPECIMENS STUDIED. Holotype 9 microptera of corvus, MExIco: Stadt, im Zimmer, 20.ii.1924 (A. Dampf) (HP). Holotype 2 macroptera of capitalis, U.S.A.: Texas, Brownsville, Palm Jungle, in beatings, 20.xi.1g1z (C. A. Hart) (USNM 71086). Gastrothrips falcatus (Ananthakrishnan) comb. n. (Text-figs 16, 64) Nesothrips falcatus Ananthakrishnan, 1968 : 969-971. Syntypes 11 9, 5 g, INp1a (TNA) [not examined]. The males of this species show extreme allometry. Major males bear several very stout recurved setae on the posterior margin of the fore femora and fore coxae, and the metanotum bears a median process projecting backwards over the pelta. These features are not developed in minor males. Allometry is also known in four other species of Gastrothrips, acuticornis, alticola, anolis and monticola, although different parts of the body are affected in each species. In most species of Thysan- optera which show allometry the major males are much larger than the minor males, but in anolis and apparently falcatus the differences in body size are not very great. Unfortunately the measurements published by Ananthakrishnan (1970; 1971) are given as ranges for each character and hence cannot be used to establish correlations. G. falcatus appears to be related to the acuticornis-group in the form of the head and tube, but differs from other members of the genus in having antennal segment eight short and broad. There is one specimen in the BMNH collections from Valparai in southern India which is very similar to falcatus in colour and the form of the wings and antennae, but has the tube constricted at the apex, the pelta broadly triangular and the postocellar setae close together. SPECIMENS STUDIED. Inp1A: from dead twigs; Madras, 1 9, ii. 1968, 3 9, xi. 1968, 2 9, vii. 1969, I , xii. 1970; Devikula, 1 ¢, ix. 1970; Goa, 2 9, x. 1968 (BMNH). Gastrothrips fulvicauda Hood Gastrothrips fulvicauda Hood, 1937 : 277-280. Holotype 9, PERu (USNM) [not examined]. This species was based on three apterous females, and unlike ruficauda the head 144 L. A. MOUND is rectangular without any ocelli (Hood, 1937: fig. 4b). The tube is yellowish orange and the distal half of the femora considerably paler than the tibiae. The most closely related species seems to be alticola which is also based on apterae from Peru, but these have the first two antennal segments brown and the tube more darkly coloured. SPECIMENS STUDIED. PERU: Cajamarca Dept., near Celendin, from bush containing dry branches, I paratype 9 aptera, I.vi.1936 (F. Woytkowskt) (USNM). Gastrothrips fulviceps Hood Gastrothrips fulviceps Hood, 1937 : 274-277. Holotype 9, PEru (USNM) [not examined]. This species was described from seven apterous females from Celendin, Peru, and the holotype was collected from a bush containing dry branches and moss. No other member of Gastrothrips has a similar colour combination of dark tube but yellowish legs, antennal segments and head. In colour and general shape fulviceps is very similar to Nesothrips propinquus (Bagnall), but that species has a more transverse head, four sense cones on the fourth antennal segment, a shorter and broader eighth antennal segment, and a differently shaped pelta without a strong subbasal line of sculpture. The head of fulviceps is as broad as long, inter- mediate in shape between that of ruficauda and the majority of Gastrothrips species. SPECIMENS STUDIED. Peru: Cajamarca Dept., Celendin, 1 9 paratype on Alansoa acutifolia flowers, 2.vi.1936 (F. Woytkowski) (AMG). GUATEMALA: Guatemala City, I 9 on Brunella vulgaris, 7.v.1945 (E. J. Hambleton) (USNM). Mexico: in quarantine at Brownsville, Texas, I 2 on rose, 6.x.1939; in quarantine at San Francisco, California, 1 2 on Sedum, 20.iv.1938 (USNM). Gastrothrips fumipennis Hood Gastrothrips fumipennis Hood, 1952 : 163. Holotype 9, Brazit (USNM) [not examined]. Hood described this species from three female and seven male macropterae. According to the description large males bear a pair of lateral processes on the mesothorax as in abditus, falcatus and acuticornis. Despite the absence of duplicated wing cilia fumipennis is very closely related to abditus, although it is rather darker with shaded wings and brown tarsi and third antennal segment. SPECIMENS STUDIED. BraziL: Santa Catarina, 2 g paratypes on dry branches, 16.v.1949 (F. Plaumann) (AMG). NESOTHRIPS COMPLEX 145 Gastrothrips intonsus Hood (Text-figs 23-26) Gastrothrips intonsus Hood, 1941a : 180-183. Holotype 9 microptera, PEru (USNM) [not examined]. f The unique holotype of this species was related by Hood to picticornis on account of the shape of the head. However, picticornis is treated here in the genus Neo- smerinthothrips because of the shape of the tube. Moreover the males of intonsus, particularly large specimens, have the head with convex spiny cheeks as in anolis. These major males, however, do not have the posterior margin of the pronotum produced into tubercles. The females have the head broadest just behind the eyes and strongly tapering to the base, and the pelta is broader than in the male and similar to fulvicauda and alticola. The bright yellow tube is similar to anolis. SPECIMENS STUDIED. PERU: Piedras Grandes, at 3000 m, I major ¢ (det J. D. Hood) in flowers, 2.xi.1937 (F. Woytkowski) (USNM); Huacapistana, 1 2 on dry leaves, 12.11.1940 (F. Woytkow- ski); San Felix, 1 minor ¢ from bush with dry leaves, 25.iii.19g40 (Ff. Woytkowsht) (BMNH). Gastrothrips mandiocae (Moulton) (Text-fig. 21) Dichaetothrips mandiocae Moulton, 1941 : 321-322. Holotype 9, Braziz (CAS) [examined]. Gastrothrips oeceticola De Santis, 1943 : 92-96. Holotype 9, ARGENTINA (Museos de la Plata) {not examined]. [Synonymized by Hood, 1956 : 99.] Hood (1950 : 22-25) redescribed and figured oeceticola from southern Brazil, and later synonymized it with mandiocae after a comparison of the type-specimens. The postocellar setae are frequently more than 100 pm long, but the female listed below from Paraguay has one of these setae less than 20 ym long. Because of this variation the female listed below from Ghana is identified provisionally as mandiocae despite the fact that the postocellar setae are short. The most closely related species are abditus and texanus. SPECIMENS STUDIED. BraziIL: Minas Gerais, 1 9, I ¢ in dead Manioc stems, 15.ix.1933 (Hambleton) [type data of mandiocae]; Vicosa, 3 9, 2 g on dead branches, 8.vii & 20.ix.1964 (F. Andre) (BMNH). Paracuay: Caaguazu, I Q swept, 2.ii.1964 (F. Andre) (BMNH). Guana: Accra, Legon, 1 9 from Tapinanthus on Cacao, 1969 (P. Room) (BMNH). Gastrothrips mongolicus (Pelikan) comb. n. Nesothrips mongolicus Pelikan, 1965 : 231-233. Holotype 9, Moncorta (TM) [not examined]. This species was described from two females and is discussed under acuticornis. 46 L. A. MOUND Contrary to the original description the ventral setae on the head are not particu- Jarly long in comparison to other members of the acuticornis group. SPECIMEN STUDIED. Monco ta: Koschoo Zaidam, in Milvus nest at 1490 m, I paratype 9, I.vii.1964 (Kaszab) (J. Pelikan collection). Gastrothrips monticola Hood Gastrothrips monticola Hood, 1942 : 573-576. Holotype 2, PERu (USNM) [not examined]. Fics 29-30. Gastrothrips intonsus, large male: 29, fore tarsus. 30, head, pronotum and mesonotum with short wings. NESOTHRIPS COMPLEX 147 This species was described from two female and one male micropterae, and its possible synonymy with alticola is discussed under that species. SPECIMEN STUDIED. Peru: Huanuco Dept., Piedras Grandes, 1 2 paratype from a bush at 3000 m, 13.x1.1937 (Ff. Woytkowski) (AMG). Gastrothrips procerus Hood Gastrothrips procerus Hood, 1956 : 99-100. Holotype 9, Brazit (USNM) [examined]. This species was described from one female macroptera, and is discussed under acuticornts. SPECIMENS STUDIED. Holotype 9, BraziL: Santa Catarina, on dead branches, v. 1949 (Ff. Plaumann) (USNM 71989). Brazit: Parana, Ponta Grossa, I ¢ in grass tufts, 14.x.1972 (V. F. Eastop). PARAGUAY: Caaguazu, I 9 sweeping, 2.11.1964 (f°. Andre) (BMNH). Gastrothrips ruficauda Hood (Text-figs 19, 22, 65) Gastrothrips ruficauda Hood, 1912 : 156-157. Syntypes 3 9, U.S.A.: Illinois (USNM) [not examined]. Stannard (1957 : 104-106) records this species from eastern North America. The males have a pointed process in front of the mesothoracic spiracle as in several other species of Gastrothrips. The legs are bicoloured as in callipus, and the tube is bright yellow as in anolis, fulvicauda and intonsus. This is the type-species of Gastrothrips but the head of the female is relatively small with more rounded cheeks than other members of the genus. The only specimen studied was an apterous female with reduced wing-retaining setae but with well developed ocelli. SPECIMEN STUDIED. U.S.A.: Virginia, Rosslyn, 1 9 from dead willow branch, 1.xi.1914 (C. B. Williams & J. D. Hood) (BMNH). Gastrothrips stygicus Hood Gastrothrips stygicus Hood, 1935 : 186-191. Holotype 9, Panama: Barro Colorado (USNM) [not examined]. Hood described this species from five female and one male macropterae. The tube is stouter than in abditus or mandiocae with weakly convex margins; however, these three species, together with texanus, are closely related. The fore tarsal tooth of stygicus is very short and broad, and the postocellar setae are intermediate in length between those of abditus and mandiocae. 148 L. A. MOUND SPECIMEN STUDIED. PANAMA: Porto Bello, 1 9 paratype on dead vegetation, 10.vii.1933 (J. D. Hood) (AMG). Gastrothrips subulatus (Hartwig) comb. n. Bolothrips subulatus Hartwig, 1948 : 113-115. Holotype 9, Sout Arrica: Transvaal (Pretoria) [not examined]. This species, which was described from 45 9 and 5 ¢ collected in Pretoria and one female collected in Lourenco Marques, is discussed under acuticornis. The original description refers to macropterous females and ‘dealate’ females and males. How- ever, the paratypes listed below are micropterae with well developed ocelli and wing retaining setae. SPECIMENS STUDIED. SouTH AFRICA: Transvaal, Pretoria, paratypes; 4 9, I g on Bignonia tweediana, 1.1.1946 (E. K. Hartwig) [same data as holotype]; 1 2 on B. tweediana, 16.ii.1941; I 9 on bean leaves, 6.1.1940 (E. K. Hartwig) (BMNH, AMG, HP). Gastrothrips texanus Hood Gastrothrips texanus Hood, 1912 : 157-159. Holotype 9, U.S.A.: Texas (USNM) [examined]. This species is known only from the unique holotype macroptera and one damaged female from the same locality. The pelta is similar to abditus, but the fore tarsus bears a tooth and the fore wings do not have any duplicated cilia. Despite this unusual combination of characters the species is closely related to abditus and mandtocae. SPECIMENS STUDIED. Holotype 9, U.S.A.: Texas, Brownsville, on Acacia farnesiana, 29.vi.1908 (C. A. Hart) (USNM 71990). I 9, 17.11.1938 (Anderson) (USNM). NEOSMERINTHOTHOHRIPS Schmutz gen. rev. Neosmerinthothrips Schmutz 1913 : 1051. Type-species: N. fructuum Schmutz, by monotypy. Coenurothrips Bagnall, 1921a@:271. Type-species: C. brevicollis Bagnall, by original designation. Galactothrips Moulton, 1933: 404. Type-species: G. diversicolor Moulton, by monotypy. Syn. n. Stannard (1957) placed this genus in synonymy with Nesothrifs on the grounds that fructwum was closely related to those species of the Nesothrips complex ‘formerly called Gastrothrips’. Unfortunately, Stannard did not indicate in what sense he was using the name Gastrothrips, and the present author has removed from Gastro- thrips to Neosmerinthothrips those species which have four sense cones on the fourth antennal segment together with paulistarum Hood. The genus Gastrothrips is limited in this paper to a group of species of the Nesothrips complex with only three NESOTHRIPS COMPLEX 149 sense cones on the fourth antennal segment. Asa result Neosmerinthothrips includes eighteen named species which are found in the Oriental, Ethiopian and Neotropical regions. The Oriental and Ethiopian species are more or less closely related, although it is interesting that the fore tarsal tooth is either present or absent. One species from Brazil, diversicolor, is probably introduced; it is so similar to the West African hilaris. Similarly the West Indian and African species collaris is very closely related to the Indian species, robustus. However, the annulipes group of species, including picticornis, plaumanni, paulistarum and variipes, is almost certainly native to Brazil. This is the group of species, together with two species from Panama and one from Fiji, which are here removed from Gastrothrips. The genus Neosmerinthothrips cannot be defined on any single character, but it includes those members of the Nesothrips complex which have a heavy tube with somewhat convex margins and the setae on tergite nine relatively elongate. GENERIC DEFINITION. Medium-sized black or dark brown species of Cryptothripini. Head variable, longer than wide, or slightly wider than long, stylets V-shaped and wide apart; post- ocellar setae small (except nigrisetis), eyes not prolonged ventrally. Antennae eight-segmented ; segment VIII either broad or narrowed at base, but never elongate and sharply constricted; segment ITI with two sense cones, IV with four sense cones (three in paulistarum). Pronotum transverse, epimeral sutures complete; oedymerous forms rare; fore tarsal tooth present in ¢ but large, small or absent in9. Metanotal median setae usually small and fine; fore wing with duplicated cilia. Pelta broad with lateral wings, lateral setae of tergites VI, VII and IX usually elongate, tube heavy with convex margins or at least with small setal bearing tubercles laterally, rarely almost straight with longitudal ridges in basal third. KEY TO THE SPECIES OF NEOSMERINTHOTHRIPS (excluding inqutlinus, q.v.) Antennal segment I yellow, if shaded with brown then distinctly paler than the head Antennal segment I dark brown, as dark as the head . ; ‘ : Fore tarsal tooth absent in 9 Fore tarsal tooth present in? . Head at least 1-2 times as long as maximum » width (Text-fig. 3 3)% head largely yellow Head less than 0-95 times as long as maximum width (Text-fig. 32) ; head dark brown Pronotal anteroangular and midlateral setae 15 zm long [West Africa]. hilaris (p. 153 Pronotal anteroangular and midlateral setae more than 35 ym long [Brazil] diversicolor (p. 152) Antennal segments I-III largely yellow; tube more than 1-8 times as long as maxi- mum width [India and Ceylon] ; d . fructuum (p. 152) — Antennal segment II brown in basal half, Ill Beowat with a yellow pedicel; tube about 1-5 times as long as wide [Brazil] F ; picticornis (p. 155) 6 Antennal segments IV and V yellow, not darker than Ill and IV [South Africa] hoodi (p. 153) — Antennal segments IV and V dark brown in contrast to yellow-brown of segment ITI 7 Fore tarsal tooth of 9 less than half as long as width of foretarsus; tube more than 1-7 times as long as maximum width; large species, antennal segment III about 80 wm long [Seychelles] ; A brevicollis (p. 151) — Fore tarsal tooth of 2 about as long as 3 width of fore tarsus: tube less than 1-6 times as long as maximum width (Text-fig. 36); small species, antennal segment III less than 60 wm long [Java] ; . ‘ : : ‘ . . xylebori (p. 155) nek AW Oh — 1 | w | vn Iw on 150 L. A. MOUND 8 Fore tarsal tooth absent in@? . z : ‘ : : 9 — Fore tarsal tooth present in ? but sometimes very small : : . : : Pe 9 Antennal segment III brown with pedicel yellow : 3 . affinis (p. 150) — Antennal segment III yellow, shaded light brown in distal half : ‘ 10 10 Fore wing shaded only close to bases of subbasal setae; fore femora largely yellowish brown, darkly shaded along external margin near base [India] . . robustus (p. 155) — Fore wing shaded completely across base; fore femora largely dark brown, paler near apex [Africa and West Indies] ‘ . : ; . collaris (p. 151) Ir Fore wings uniformly shaded light or dark brown ; - : 12 — Fore wings not shaded, or shaded at base with median area ‘much paler P 13 12 Fore wing dark brown; major setae in tergite IX exceptionally dark and stout with pale slender apices, B, longer than tube; pronotal anteromarginal and antero- angular setae scarcely ee than posterior discal setae [Brazil] plaumanni (p. 155) — Fore wing light brown; major setae on tergite IX not exceptionally dark and stout, B, about half as long as tube; pronotal anteromarginal and anteroangular setae twice as long as posterior discal setae [Fiji] . 4 ‘ , . fijiensis (p. 152) 13 Antennal segment IV with three sense cones [Brazil] ; . paulistarum (p. 154) — Antennal segment IV with four sense cones é 14 14 Fore tarsal tooth well developed, length more than o- “5 of fore tarsal width: head broadest across posterior part of eyes [Brazil] , 15 — Fore tarsal tooth small, length less than 0-25 of tarsal width; head broadest across cheeks [Panama] 16 15 Second antennal segment brown at base, Callow at apex; “femora brown, pale at extreme apices; tube about 2:1 times as long as maximum width annulipes (p. 150) — Second antennal segment largely yellow; femora yellow in distal third ; tube less than 2-o times as long as wide (Text-fig. 37) . : 5 . variipes (p. 155) 16 Postocellar setae about 2.0 times as long as diameter of one ocellus; apices of femora yellow; major setae on abdomen black . : ‘ .nigrisetis (p. 154) — Postocellar setae about 0.5 times as long as diameter of one ocellus; femora dark brown; major setae on abdomen pale. : ‘ ‘ ; parvidens (p. 154) Neosmerinthothrips affinis (Bagnall) comb. n. Coenurothrips affinis Bagnall, 1921b : 361-362. Holotype 9, CeyLon (BMNH) [examined]. This species is known from a single crushed specimen which is similar to collaris and rvobustus in structure but with the third antennal segment largely dark brown. The fore femur is dark brown along the external margin and yellow along the internal margin. Unlike brevicollis the fore tarsus has no tooth. SPECIMEN STUDIED. Holotype 9, CEYLON: among cotton from Hettipold, Ex. Gd., 20.vi.1913 (A. Rutherford) (BMNH). Neosmerinthothrips annulipes (Hood) comb. n. Gastrothrips annulipes Hood, 1950 : 13-16. Holotype 9, Brazir (USNM) [not examined]. Nesothrips (Gastrothrips) milleforme De Santis, 1963a : 12-14. Holotype 9, ARGENTINA (Museos de la Plata) [examined]. Syn. n. The head of this species is similar to that of the Brazilian species variipes and picticornis, but the tube is more slender with nearly straight sides and slightly NESOTHRIPS COMPLEX 151 constricted near the apex. The other species described from Brazil, plaumanni, has the head wider across the cheeks than across the eyes but is otherwise similar, and these four species are all closely related. The holotype of milleforme has the distal parts of the major abdominal setae pale not dark, but cannot otherwise be distinguished satisfactorily from the paratype of annulipes listed below. SPECIMENS STUDIED. Holotype 2 of milleforme, ARGENTINA: Tezanosa Pinto (Prov. Entre Rios), 1957 (Rosillo) (Museos de la Plata). BRAZIL: Rio de Janeiro, Jacarepagua, I 2 paratype of annulipes, on dead branches, ai.vi.1948 (J. D. Hood) (USNM). Neosmerinthothrips brevicollis (Bagnall) comb. nov., stat. rev. Coenurothrips brevicollis Bagnall, 1921a : 271-272. Syntypes g, 9, SEYCHELLES (BMNH) [examined]. This species was synonymized in error with Coenurothrips validus Bagnall in Mound (1968 : 142), and placed in the genus Nesothrips sensu lato. However, validus, which is known from a single badly damaged specimen, has a pair of stout interocellar setae and is here placed in the genus Nesidiothribs. N. brevicollis is very similar to collaris and robustus although rather smaller. However, it differs not only in having paler basal antennal segments but in having a small slender fore tarsal tooth in the female. In contrast to hoodi these three species all have yellowish brown setae in tergite nine. SPECIMENS STUDIED. Syntypes. SEYCHELLES: Mahe, 1 9, Cascade Estate, top of Mt Serbert, x. rg08— i. 1909 (H. Scott 104); 1 J, 1908-09 (3. No. 71); 2 9, 1908-09 (15 No. 71) (BMNH). Neosmerinthothrips collaris (Bagnall) comb. n. Cryptothrips collaris Bagnall, 1917 : 26-27. Lectotype 9, St. Vincent (BMNH) [examined]. Gastrothrips fuscicauda Morgan, 1925 : 6-7. Holotype 3, PuERTo Rico (USNM) [examined]. Syn. n. Bolothrips marshalli Priesner, 1934 : 58-60. LECTOTYPE 9, Sierra Leone (BMNH), here designated [examined]. Syn. n. Gastrothrips dominicanus Hood, 1935: 170-174. Holotype g, Dominica (USNM) [not examined]. Syn.n. Bagnall described collaris from two females and Hood described dominicanus from two males but there appear to be no significant differences between the speci- mens. The damaged unique holotype of fuscicauda has been compared with the paratype of dominicanus as well as the other material listed below. More surpris- ingly the African specimens described as marshalli apparently represent the same species together with the specimen from Mozambique, and moreover Indian speci- mens referred to as vobustus Ananthakrishnan can only be distinguished on very 152 L. A. MOUND trivial characters. Thus collaris appears to be a tramp species which has been distributed by man, possibly during the slave trade. SPECIMENS STUDIED. Lectotype 2, with 1 2 paralectotype of collaris, St VINCENT: ?1895 (H. H. Smith) (BMNH). Holotype g of fuscicauda, PUERTO Rico: from stomach of lizard (G. N. Wolcott) (USNM type No. 71987) (labelled ?92). Lectotype 9 of marshalli, SIERRA LEONE: Nijala, inside Cassava seedpod, 4.xi.1932 (E. Hargreaves), with 1 g para- lectotype bearing identical data (BMNH). Dominica: paratype 3 of dominicanus, swept from grass, 13.11.1915 (C. B. Williams 571) (AMG). MozaAmpBiguEe: Lourenco Marques, 2 9, 4 ¢ in dry Cassia pods, vii. 1936 (J. C. Faure) (AMG). Neosmerinthothrips diversicolor (Moulton) comb. n. Galactothrips diversicolor Moulton, 1933 : 404-406. Holotype 9, Braziz (CAS) [not examined]. This is probably only a local and introduced variety of hilaris Priesner from West Africa. The only apparent differences are the slightly longer setae on the pronotum. SPECIMEN STUDIED. BRAZIL: Bahia, on Galactia sp., I paratype 2 (Bondar) (BMNH). Neosmerinthothrips fijiensis (Moulton) comb. n. Gastrothrips fijiensis Moulton, 1944 : 286-287. Holotype 9, F1y1 (BPBM) [examined]. The single specimen from which this species is known has the head rather similar to the annulipes group from Brazil and the tube is also similar to annulipes. How- ever, the abdominal setae are rather short, the dorsal setae on tergite nine being 145 wm and the tube 260 ym. At present fijiensis cannot be placed satisfactorily in any species group. The median part of the pelta is triangular although the lateral wings are well developed. SPECIMEN STUDIED. Holotype 9, Fiy1; Navai Mill, Viti Levu at 2500 ft, 16.ix.1938 (EZ. C. Zimmerman) (BPBM). Neosmerinthothrips fructuum Schmutz (Text-figs 32, 34, 35, 70) Neosmerinthothrips fructuum Schmutz, 1913 : 1052-1053. LECTOTYPE 4g, Cryton (HP), here designated [examined]. Oedemothrips ceylonicus Karny, 1925 : 137-139. Holotype 9, CzEyLton (BMNH) [examined]. Syn. n. NESOTHRIPS COMPLEX 153 The specimen here designated as lectotype was labelled by Prof. H. Priesner ‘Typen material, ex coll. Karny’, although the original description does not indicate if more than one specimen existed. This specimen is very faded and partly crushed, but otherwise cannot be distinguished from specimens labelled as ‘formosensis var. Karni’ by Prof. Ananthakrishnan collected in southern India. The female of the species lacks a fore tarsal tooth. The true formosensis is treated here under Nesothrips, and has a tube with straight not weakly convex margins. SPECIMENS STUDIED. Lectotype ¢ of fructuum, CEYLON: Peradenyia, in Schoten (Uzel 55) (HP). Holo- type 2 of ceylonicus, CEYLON; Peradenyia, in empty galleries in Cassia multijuga with 3 immatures, I.v.1924 (F. T. Jepson) (BMNH). InpIA: Kerala, Kudal, 1 9 in dry twigs, 6.x.1969 (TNA 158); Vyithri, 3 9, 15.vili.1969; Ranni, 2 9, 5.x.1969; Kiruatti, 1 g, 19.ix.1969 (BMNH); Sagar, 1 9 20.x.1966 (HP)... Neosmerinthothrips hilaris (Priesner) comb. n. (Text-fig. 33) Bolothrips hilaris Priesner, 1937 : 624-626. Holotype 3, SrERRA LEONE (BMNH) [examined]. The tube of this species is heavy with slightly convex sides bearing distinct setal bases, as in other species of Neosmerinthothrips. There are four sense cones on the fourth antennal segment, and segment eight is broad at the base. The eyes are smaller on the ventral than on the dorsal surface, the head does not project in front of the eyes and the ocellar setae are minute. The epimeral sutures are sometimes not quite complete, and the setae on tergite nine are elongate. Despite the colour difference hilaris is related to collaris and fructuum in structure, and the Brazilian species diversicolor is probably only a local variant. SPECIMENS STUDIED. Holotype g, with allotype 9, SERRA LEONE: Niala, in twigs of Bauhinia tomen- tosa, 1936 (E. Hargreaves); 1 9 paratype at same locality on dead branch of Ficus (BMNH). GHANA: near Legon, 11 9, 6 g from Tapinanthus banguensis on Cacao branches, 1969 and 1970 (P. Room) (BMNH). Neosmerinthothrips hoodi (Faure) comb. n. Gastrothrips hoodi Faure, 1954a: 9-13. Holotype 9 macroptera dealate, SourH AFRICA (Pretoria) [not examined]. This species is very similar to fructuwm in the shape of the head and tube, and differs mainly in colour and the presence of a pretarsal tooth. Although it was described in Gastrothrips, Faure specifically compared it to fructuwm (as ceylonicus Karny) and moreover listed those characters which are here regarded as excluding species from Gastrothrips sensu stricto. 154 L. A. MOUND SPECIMENS STUDIED. SouTtH Arrica: Zululand, St Richards Bay, 1 2 mac. paratype on dead branches, 17.11.1946; Kluhluwe, 4 9 dealate paratypes on dead branches of Syzygium cordatum, 13.v.1949 (J. C. Faure) (AMG and BMNH). Neosmerinthothrips inquilinus Ananthakrishnan Neosmerinthothrips inquilinus Ananthakrishnan, 1960 : 32-33. Holotype 9, Inpia (?TNA) [not examined]. The holotype female and two male paratypes of inquilinus cannot be found at present in Prof. Ananthakrishnan’s collection (teste T.N.A. in litt.). However, judging from the description this is likely to prove to be a senior synonym of robustus q.v. Neosmerinthothrips nigrisetis (Hood) comb. n. Gastrothrips nigrisetis Hood, 1935 : 161-165. Holotype 9, Panama (USNM) [not examined]. The ocellar setae of nigrisetis are very well developed, about 40 ym long and arising well behind a line joining the posterior ocelli. As a result the head and pronotum look very like a species of Dichaetothrips, but the tube is unusually heavy with small setal tubercles laterally and the abdominal setae are elongate. This species and parvidens are closely related and are probably derived from the annulipes group. They are only distantly related to the other species of Neo- smerinthothrips, although Hood compared them to collaris (as dominicanus) in the original description. SPECIMENS STUDIED. PANAMA: Barro Colorado Island, on dead branches, 3 9, 1 $ paratypes, 30.vii.1933 (J. D. Hood) (AMG). Neosmerinthothrips parvidens (Hood) comb. n. Gastrothrips parvidens Hood, 1935 : 165-168. Holotype 9, Panama (USNM) [not examined]. This species is closely related to migrisetis but has very small ocellar setae, and the tube is more slender than any other species of Neosmerinthothrips. SPECIMENS STUDIED. PANAMA: Barro Colorado Island, on dead branches, I paratype 9, 10.vill.1933 (J. D. Hood) (USNM); Canal Zone, Barro Colorado Is., 1 9 on Swartzia danienensis fls, 10.v.1939 (Zetek) (USNM). Neosmerinthothrips paulistarum (Hood) comb. n. Gastrothrips paulistarum Hood, 1950 : 25-27. Holotype 9, Brazit (USNM) [examined]. Because of the antennal sense cone formula this species will come out to Gastro- NESOTHRIPS COMPLEX 155 thrips in the key above. However, it is based on a single specimen which may be aberrant, and it is closely related to the annulipes group of species. The tube is dark and heavy with slightly convex margins, and the head shape is intermediate between that of annulipes and parvidens. SPECIMEN STUDIED. Holotype 2, BrAziL: Estado de Sado Paulo, Sao Carlos, from dead branches, 13.vi.1948 (J. D. Hood & D. P. de Souza Dias) (USNM). Neosmerinthothrips picticornis (Hood) comb. n. Gastrothrips picticornis Hood, 1936 : 272-275. Holotype 9, Brazir (USNM) [not examined]. The tube of this species is very broad, rather similar to that of xylebori from Java. However, pPicticornis is probably closely related to plaumanni and varitpes which also come from Brazil, but which have a fore tarsal tooth in the female. SPECIMENS STUDIED. Braziv: Rio de Janeiro, 1 2 paratype on dead twigs of Annona squamosa, 10.viii.1934 (D. Mendes) [holotype in USNM bears identical data]; State of Sao Paulo, Itanhaen, I $ on dead branches, 17.vi.1948 (J. D. Hood & J. Lane) (AMG). Neosmerinthothrips plaumanni (Hood) comb. n. Gastrothrips plaumanni Hood, 1950 : 20-22. Holotype 2, Brazit (USNM) [examined]. This is the only South American member of this genus with dark brown fore wings. The abdominal setae are long and dark as in picticornis and varitpes, and the tube is moderately heavy. SPECIMENS STUDIED. Holotype 9, BraziL: Sta Catarina, on dry branches, 5.i.1949 (F. Plaumann) (USNM Type No. 71988). BraziL: Vicosa, 2 2 on dead braches, 20.xi.1964 (F. Andre) (BMNH). Neosmerinthothrips robustus (Ananthakrishnan) comb. n. Nesothrips robustus Ananthakrishnan, 1964 : 102-103. Syntypes g, 9, Inpia (TNA) [not examined). Contrary to the original description the female of this species does not have a fore tarsal tooth (Ananthakrishnan, 1968 : 972). It is placed in the genus Neo- smerinthothrips on account of the heavy tube with weakly convex margins bearing small setal bases, and the elongate setae on tergite nine. It is very similar to the widespread species collaris, being rather smaller and paler, and may eventually prove to be merely a local variety. D 156 L. A. MOUND SPECIMENS STUDIED. Inp1A: Madras, 2 ¢, 2 9; Goa, r g; Adar, r ¢, 1 9; Thenmatai, x 9, all determined by T. N. Ananthakrishnan (BMNH). Neosmerinthothrips variipes (Hood) comb. n. (Text-figs 37, 73) Gastrothrips variipes Hood, 1950 : 16-20. Holotype 2, Brazit (USNM) [not examined]. This species is very close to plaumanni but the fact that it has rather paler wings and more yellowish basal antennal segments and femora may not be significant when more specimens are collected. SPECIMENS STUDIED. BRAZIL: Rio de Janeiro, Jacarepagua, I 2 paratype on dead branches, 22.v.1948 (J. D. Hood & T. Borgmeir); State of Sao Paulo, Sao Carlos, 1 g paratype, 15.vi.1948 (J. D. Hood & D. P. de Souza Diaz) (AMG). Neosmerinthothrips xylebori Priesner (Text-fig. 36) Neosmerinthothrips xylebori Priesner, 1935 : 370. LECTOTYPE 9, Java (SMF), here designa- ted [examined]. @ macroptera. Colour brown, tube blackish; mid and hind femora dark at base and along anterior margin but yellow at apex and posterior margin; fore femora yellow in apical half; antennal segments I and II yellow, III light brown, succeeding segments progressively darker; wings shaded; major setae light brown. MEASUREMENTS (Lectotype 9 in wm). Body length 1550. Head, length 175; maximum width 190. Fore wing, length 700; subbasal setae 50, 50, 58; number of duplicated cilia 6-8. Tergite IX setae, B, 145, B, 130, B, 125. Tube, length 190; basal width 115; apical width 35. Antennal segments length 26; 50; 55; 50; 50; 50; 38; 32. SPECIMENS STUDIED. Lectotype 9, JAvA: Tegallega near Tjibadak, 500-600 m, in tunnel of Xyleborus coffeae (on Cacao?), 30.iii.1925 (Menzel), with r $ paralectotype (SMF); 2 2 para- lectotypes with identical data (HP). ComMMENTS. This species was described in a key, complete with collection data but without reference to the number of individuals. The lectotype selected here is the female labelled as holotype by Prof. Priesner which is mounted on a slide with one larva. Despite the long slender fore tarsal tooth of the female it is similar to fructuum, although the only available male is hemimacropterous and oedymerous. NESIDIOTHRIPS gen. n. Type-species: Nesothrips alius Ananthakrishnan. Medium-sized dark brown species of Cryptothripini with males frequently oedymerous. NESOTHRIPS COMPLEX 157 Head longer than wide or about as wide as long; ocellar setae elongate, arising within the ocellar triangle; postocular setae elongate, major males with two pairs of postocular setae; maxillary stylets wide apart, V-shaped and retracted to postocular setae. Antennae eight- segmented, VIII fairly broad at base; two sense cones on segment III, four sense cones on IV. Pronotum with median thickening, anterior margin thickened in males; fore femora sometimes enlarged, fore tarsal tooth well developed in both sexes. Mesonotal lateral setae small; meta- notal median setae frequently stout; fore wing with duplicated cilia; mid and hind femora each with one stout dorsal seta. Pelta with small lateral wings; sigmoid wing retaining setae well developed on tergites III—-VI, nearly straight on II and VII; lateral setae on tergites VI and VII elongate, on IX shorter than tube; tube heavy, ridged basally and slightly constricted near apex. This genus is erected for two species, one from India and the other from the Seychelles and the Solomon Islands. These species are closely related to Nesothrips and Rhaebothrips species, and the mid and hind femora bear a single stout dorsal seta as in those genera. However, the ocellar setae are elongate and arise within the ocellar triangle, and the fore tarsus bears a large tooth in the female unlike both Nesothrips and Rhaebothrips. KEY TO THE SPECIES OF NEST DIOTHRIPS 1 Setae on tergite IX yellowish brown; midlateral pronotal setae more than 804m long in 9; head slightly longer than tube [India] . ‘ : ? alius (p. 157) — Setae on tergite [IX dark brown; midlateral pronotal setae less than 60 um long in Q; head slightly shorter than tube [Seychelles] : d ; : . validus (p. 157) Nesidiothrips alius (Ananthakrishnan) comb. n. (Text-figs 46, 48, 50, 69) Nesothrips alius Ananthakrishnan, 1970 : 52-55. Holotype 9, Inp1a (TNA) [not examined]. The original description of this species includes illustrations of the dimorphism. The oedymerous males have a second pair of long postocular setae medially on the vertex which are not developed on females or minor males. The median metanotal setae and the pronotal anteroangulars and midlaterals are also elongate in large males. This species is closely related to validus and further collecting may indicate that only one widespread species is involved. SPECIMENS STUDIED. Inp1A: Aryankavu, I 9, I § on dry twiner, 7.x.1969 (I. N. Ananthakrishnan) (TNA) [these data are similar to those published for the type-series — Kerala, Aryankavu, dry twigs, 8.x.1969]. Nesidiothrips validus (Bagnall) comb. n. Coenurothrips validus Bagnall, 1921a : 272-273. Holotype 9, SEYCHELLES (BMNH) [examined]. The holotype of this species is very badly crushed and the interocellar setae are not readily visible. The species C. brevicollis Bagnall is not a synonym of validus contrary to Mound (1968 : 142). The specimens listed below from the Solomon 158 L. A. MOUND Islands are identified provisionally as validus, the males show a similar dimorphism to alius but the terminal abdominal setae are dark brown. SPECIMENS STUDIED. Holotype 2, SEYCHELLES: Silhouette [Mare aux Cochons, ix. 1908] (BMNH). Sotomon IsLaAnDs: Guadalcanal, Mt Austen, 2 3, I @ in litter, 19.xi.1963 (P. Greenslade) (BMNH). NESOTHRIPS Kirkaldy Nesothrips Kirkaldy, 1907 : 103. Type-species: N. oahuensis Kirkaldy, by monotypy. Oedemothrips Bagnall, 1910: 680. Type-species: O. laticeps Bagnall, by monotypy. [Synony- mized by Bianchi, 1944.] Fic. 31. Nesothrips oahuensis, large male. NESOTHRIPS COMPLEX 159 Oedemothrips is a synonym of Nesothrips because the names Jaticeps and oahuensis have been shown to refer to the same species. However, the other genera which were placed as synonyms of Nesothrips by Stannard (1957) are here withdrawn from synonymy for the reasons given in the introduction and key. Unfortunately this still leaves Nesothrips as an unsatisfactory group of fourteen more or less related species from the Pacific, Australia and Indonesia, with two species which have become widespread on trade routes. This group of species is closely related to Rhaebothrips through aoristus and oahuensis, and it is also closely related to Cariento- thrips through melinus and propinquus. Neosmerinthothrips fructuum is sometimes misidentified as Nesothrips brevicollis (or one of its synonyms), but the species of Neosmerinthothrips have a stout heavy tube with weakly convex instead of straight margins. Similarly, small species of Scotothrips and Dichaetothrips have been placed in Nesothrips despite the fact that they have a more elongate tube. In the present interpretation Nesothrips includes both short broad species such as oahuensis and propinquus, as well as more elongate species such as niger and yanchept. The eyes are prolonged on the ventral surface of the head in some species and in propinquus the extent of this prolongation is variable. Duplicated cilia are present on the fore wings of those species in which macropterae are known with the exception of propinquus which does not have duplicated cilia. The syste- matics of this group is not likely to be stabilized without further collecting in the New Guinea and Pacific areas. GENERIC DEFINITION. Small to medium sized dark species of Cryptothripini. Head fre- quently wider than long but sometimes longer than wide, usually only weakly extended in front of eyes; eyes variably prolonged on ventral surface; maxillary stylets wide apart and V-shaped. Antennae eight-segmented, two sense cones on segment III, four sense cones on IV; segment VIII short and broad. Pronotum transverse, sometimes enlarged in major males; epimeral sutures complete. Fore tarsal tooth present in g, absent in9. Wings, when present, usually with duplicated cilia. Metanotal median setae usually small. Pelta with lateral lobes in most species. Tube relatively short, with straight sides. KEY TO THE SPECIES OF NESOTHRIPS rt Antennal segment I yellow or brownish yellow, much paler than VII and VIII and usually paler than the head ‘ ; 2 — Antennal segment I brown, as dark as the head and distal antennal segments , 8 2 Setae B, on tergite IX more than o-8 times as long as the tube; major setae unusually slender; dorsal surface of head with two pairs of elongate postocular setae; mid and hind legs dark brown except for the pale hind ia of femora at extreme apex [Rapa Is.] . i : semiflavus (p. 168) — Setae B, on tergite IX less than Oo P times as long as , tube: major setae stout and dark; head with only one pair of postocular setae; mid and hind legs sometimes yellow, or with apex of femora more or less yellow 3 Head more than 1-1 times as long as wide (Text-fig. 38) [Guam] . artocarpi (p. 161) — Head less than 1-o times as long as wide, usually distinctly wider than long (Text- figs 40-42) : 4 4 Posterior margin of hind femora yellow, anterior margin dark brown, ‘fore femora yellow in distal half; eyes not prolonged on ventral surface of head . Micropterae with anterior margin of head and antennal segments I-V 160 oO loin 5 i i Iz 13 L. A. MOUND yellow or brownish yellow; macropterae and hemimacropterae with head and most of antennal segments IV—V brown [Mascarenes, Indonesia, Japan, Hawaii] (see also rhizophorae) . : : . brevicollis (p. Femora coloured differently from above, or ‘with eyes prolonged on ventral surface of head Tube as long as or a ‘little Jonger than the head: hind femora dark brown but yellow in distal third; antennal segments I and II yellow, III yellow-brown, IV scarcely paler than V—VIII which are dark brown [Malaya, Sumatra] . malaccae (P. Tube 0-8 times as long as head or shorter; hind femora largely yellow . Lateral lobes of pelta separate from the median lobe or with only a slender connec- tion, posterior border of pelta eroded medially and separate from anterior border of tergite II [Australia, New Zealand, South Africa] : . propinquus (p. Lateral lobes of pelta broadly joined to median lobe, posterior border close to anterior border of tergite II (Text-fig. 45) . Body colour dark brown; postocellar setae ‘finely pointed [Fiji] : . fodinae (p. Body colour largely yellow: ocellar setae blunt at apex [Australia] . melinus (p. Ventral length of eyes at least 1-5 times as long as dorsal length [Pacific Is.] Ventral length of eyes less than 1:2 times dorsal length [Australia] ‘ Ocellar setae 60-804 m long, 40-60 wm apart at base os fig. 31); setae By on tergite IX 90-100um [Hawaii] . ; oahuensis (p. Ocellar setae 35 um long, 80m apart at base (Text- fig. 39): setae B, on tergite IX 200 wm [Marquesas Is.] ; 2 niger (p. Pelta semicircular, lateral lobes not developed; head almost transversely oval, broadest across cheeks; pronotum very short, more than three times as wide as long . : ; . hemidiscus (p. Pelta with lateral lobes well developed: ‘head not transversely oval; Po scarcely twice as wide as long : Setae B, on tergite IX 0-6—-0-75 times as long as B, s: head less than 0-9 times as long as wide; mid and hind femora yellow at apex and along posterior margin in distal third (see also brevicollis) . , . rhizophorae (p. Setae B, on tergite IX 1-o-1-2 finies as long as B;; head longer than wide; mid and hind femora brown Setae B, on tergite IX 1-2 times as long as Be but less than. 0°5 times as s long as the tube; major setae including ocellar and metanotal setae ss tanned stout and dark . i : . aoristus (p. Setae B, on tergite IX about as s long as ah and ; more than « o-5 times as long as the tube; major setae slender Antennal segment III brown with pedicel yellow, IV and Wi “dark brown carverae (p. Antennal segment III largely yellow, shaded near apex, IV and V yellow at base yanchepi (p. Nesothrips aoristus Mound Nesothrips aoristus Mound, 1974: 68. Holotype 9, AusTRALIA: South “Australia (ANIC) [examined]. The oedymerous male of this species has enlarged fore femora similar to oahuensis, but the eyes are not prolonged ventrally as in that species. Although the head and fore legs are similar to Rhaebothrips species the tube and antennae are relatively short. NESOTHRIPS COMPLEX 161 SPECIMENS STUDIED. Holotype Q with 2 9, 2 gd paratypes, AUSTRALIA: South Australia, south of Adelaide, on Olearia ramalosa, 4.xii.1967 (L. A. Mound) (ANIC, BMNH). Nesothrips artocarpi (Moulton) (Text-fig. 38) Bolothrips artocarpi Moulton, 1942 : 14-15. Holotype 2, Guam (BPBM) [examined]. Fics 32-37. Neosmerinthothrips species: 32, N. fructuum. 33, N.hilaris. 34, N. fructuum, pelta. 35, N. fructuum, tube. 36, N. xylebori, tube. 37, N. variipes, tube. 162 L. A. MOUND This species, described from two females, is similar to semiflavus in the shape of the head. The maxillary stylets are retracted into the head as far as the posto- cular setae, and the postocellar setae are 30 um long. The metanotum is almost unsculptured around the bases of the median setae, and these setae are slender, 40 wm long and about 45 wm apart. The pelta has well developed lateral wings. The tube is relatively long, 195 um, with straight sides, and the setae on tergite nine are dark brown with pale acute apices. SPECIMEN STUDIED. Holotype 9, GUAM: on Artocarpus communis, 21.v.1936 (Sweezey) (BPBM). Nesothrips brevicollis (Bagnall) (Text-figs 40, 43, 44)| Oedemothrips (?) brevicollis Bagnall, 1914 : 29-30. Holotype 9, JapAN (BMNH) [examined]. Coenurothrips minor Bagnall, 1921a : 287-288. Holotype 9, RopricuEs (BMNH) [examined]. Syn. n. Neosmerinthothrips formosensis Priesner, 1935 : 368-370. LECTOTYPE Q, Tatwan (HP), here designated [examined]. Syn. n. Neosmerinthothrips formosensis var. karnyi Priesner, 1935 : 369-370. LECTOTYPE 9, Java (SMF), here designated [examined]. Syn. n. The name formosensis var. karnyi has been used by Ananthakrishnan in several publications from Madras (e.g. 1973), but most of the specimens bearing that name from southern India are referred to here as Neosmerinthothrips fructuum Schmutz. These specimens have short postocellar setae, the mid and hind femora pale distally, and the tube broad at the base with convex sides. The single specimen listed below from India and bearing Ananthakrishnan’s identification as formosensis karnyt has a pair of setae between the posterior ocelli 35 wm long, the hind femora yellow along the posterior border, and the tube relatively slender with straight sides. Bagnall described this species on a single micropterous female from Japan, and Priesner described formosensis on two micropterous females from Taiwan (the only whole specimen is here designated lectotype). Bagnall’s specimen is slightly larger and crushed but cannot otherwise be distinguished. The variety karnyi and the species minor were both described from macropterous females, which as is indicated in the key to species have a darker coloured head and median antennal segments. The males listed below from Hawaii are hemimacropterous, wing length 500 pm, and have a similar colour pattern to the macropterae. This also applies to the micropterous female from Mauritius listed below, although this specimen has rather long wing lobes, 125 wm. The Australian species rhizophorae is possibly only a local colour variant of brevicollis. In the USNM collections there are three specimens taken in quarentine, apparently from Guam, which have the ocellar setae scarcely 15 xm long and arising just behind a line joining the hind margins of the posterior ocelli. Usually in brevicollis the ocellar setae are about 50 «xm long and just anterior of a line joining the hind margins NESOTHRIPS COMPLEX 163 of the posterior ocelli. Mound (1974) refers to similar variation in the position of the ocellar setae in rhizophorae. SPECIMENS STUDIED. Holotype 9 of brevicollis, JAPAN: Okinawa, Luchu Is., v. 1913 (J. E. A. Lewis) (BMNH). Lectotype 2 of formosensis, TAtwan: Hookotoo, Makoo, 5.vi.1930 (S. Minowa) (HP), with a damaged @ paralectotype on the same slide. Lectotype Q (labelled holotype) of formosensis var. karnyi, JAVA: Tjibodas, 1400m, 1923 (Karny) (SMF). Holotype 2 of minor, RopRIGUES: vii-xi. 1918 (H. J. Snell & H. P. Thomasset) (BMNH). Mauritius: De Pouce Mt, 1 9, I g micropterae on leaves, 2.xi.1939 (kk. Mamet) (HP). ReEvuNIon: St Louis, 1 9 macroptera on tobacco, 14.vii.1951 (J. R. Walliams) (BMNH). Inp1A: Aryankavu, I 2 macroptera on dry twig, 19.vil.1969 (Ananthakrishnan) (BMNH); Pachmarki, 1 9, 20.xi.1966 (Ananthakrishnan) (HP). Hawau: Pupukea, 8 2, 2 ¢ on Leucaena, 16.xii.1969; Barbers Point, 2 9, 6 3, sweeping and on Desmanthus, 16.xii1.1969 (F’. Andre) (BMNH). Nesothrips carverae Mound Nesothrips carveri Mound, 1974: 71. Holotype 9, AusTRALIA: South Australia (ANIC) [examined]. The holotype of this species was collected at Adelaide, but the original description lists specimens from Western Australia, Victoria and the Australian Capital Terri- tory. The nomen nudum ‘Oedemothrips nigricans Bagnall’ in Kelly & Mayne (1934 : 51) refers to this species. The most closely related species is yanchepi with which carverae has been collected in Western Australia near Perth. SPECIMENS STUDIED. AUSTRALIA: South Australia, Adelaide, holotype @ on washing, 15.x.1967 (M. Carver) (ANIC). A complete list of specimens is given in Mound (1974). Nesothrips fodinae sp. n. Q macroptera. Head, thorax and abdomen dark brown, legs brownish yellow with the morphological upper surface of all femora deeply shaded at the base, coxae dark brown, tarsi yellow; antennal segments I-VI yellow, VII and VIII light brown; wings strongly shaded, particularly at base; major setae dark brown. Head broader than long, weakly projecting in front of eyes, postocular setae acute; posto- cellar setae close to inner margin of each posterior ocellus; eyes prolonged on ventral surface. Antennal sensoria rather slender, two sense cones on III, four sense cones on IV; basal pedicel of segment VII broad. Pronotum with a weak median thickening; anterior setae reduced, posterior setae acute; fore femora moderately expanded. Metanotum apparently not sculp- tured medially, median setae slender and acute, 20 zm long. Pelta with broad lateral wings; tergal wing retaining setae reduced on II, sigmoid on III—VII; dorsal and lateral setae on tergite IX short; sides of tube almost straight. 164 L. A. MOUND MEASUREMENTS (holotype 2 in wm). Body length 1850. Head, length 175; maximum width 210; basal width 175; postocular setae 50; postocellar setae 23. Pronotum, length 115; median width 230; major setae —aa, am, ml, 16, epim 58, pa 30-35. Fore wing, length 750; distal width 60; subbasal setae 35, 75, 90; number of duplicated cilia 9. Tergite VII setae, B, 130; B, 95. Tergite IX setae, B, 55; B, 52; B; 70. Tube, length 135; basal width 70; apical width 32; terminal setae 110. Antennal segments length (paratype 9), 27; 50; 67; 58; 55; 52; 36; 27. 6 macroptera. Head and thorax brown, abdomen dark brown; legs yellow with small brown markings particularly at base of femora; otherwise similar to 9. Fore femora strongly swollen, fore tibiae stout, fore tarsi with a stout tooth. Pronotum enlarged, anterior margin and median line strongly thickened ; midlateral setae elongate. Metanotal setae 70m long. MEASUREMENTS (paratype g in wm). Body length 1800. Head, length 170; maximum width 187; postocular setae ?90. Pronotum, length 195; median width 250; major setae — am 20, aa 25, ml 100, epim ?80, pa ?60. Fore wing, length 650; subbasal setae 55, 100, 160; number of duplicated cilia 7. Tergite IX setae, B, 70; B, 60; B; 120. Tube length 135. SPECIMENS STUDIED. Holotype 9, Fiy1: Lau, in [?palm leaf] mines of Promecotheca reicher (Hispidae), li. 1932 (R. W. Paine) (BMNH). Paratypes, 2 9, 4 ¢ collected with the holotype, also several immatures of first, second and fifth instar. CoMMENTS. This species is readily distinguished by the dark body with pale legs and antennae. WN. propinquus is similar in head shape and form of the eyes, but has a different pelta and usually has a paler head and thorax. The males of propinquus are apterous. The males of fodinae are interesting not only because they are macropterous, but also because of the sexual dimorphism in the lengths of the setae on the head, thorax and wings. The postocellar setae vary from 23- 30 um but the series is too short to establish whether these also are subject to sexual dimorphism. Nesothrips hemidiscus Mound Nesothrips hemidiscus Mound, 1974 : 71-72. Holotype 9, AusTRALIA: Queensland (ANIC) [examined]. The D-shaped pelta of this species is unique in the genus. SPECIMENS STUDIED. Holotype 2 with 7 9, 2 ¢ paratypes, AUSTRALIA: Queensland, near Mareeba, on dead Casuarina twigs, 23.vii.1968 (L. A. Mound) (ANIC, BMNH). Nesothrips malaccae sp. n. (Text-fig. 42) Q macroptera. Body and legs dark brown, head and abdomen blackish, all femora yellow in distal third; antennal segments I and II yellow, III light brown with yellow pedicel, IV- VIII dark brown; major setae black; fore wings deeply shaded, particularly at base, with one longitudinal dark stripe. NESOTHRIPS COMPLEX 165 Fics 38-45. Nesothrips species: 38, N. artocarpi. 39, N. niger. 40, N. brevicollis. 41, N. fodinae. 42, N. malaccae. 43, N. brevicollis, pelta. 44, N. brevicollis, tube. 45, N. fodinae, pelta. 166 L. A. MOUND Head broadest behind eyes, cheeks rounded and narrowed to base with two pairs of fine setae; dorsal surface not sculptured; postocellar setae arise behind the posterior ocelli; posto- cular setae almost acute; maxillary stylets wide apart, retracted to postocular setae; eyes not prolonged on ventral surface. Antennae typical of genus, two sense cones on segment III, and four sense cones on IV; VIII not constricted at base. Pronotum transverse, epimeral sutures complete; anteromarginal setae slender and acute, remaining setae stout and bluntly acute. Metanotum with a pair of slender setae medially. Pelta with broad lateral wings; tergal wing retaining setae well developed on II-VII; lateral setae exceptionally long on VII; tube with straight sides. MEASUREMENTS (holotype 2 in wm). Body length 1900. Head, length 210; maximum width 215; basal width 150; postocellar setae ?20; postocular setae 80. Pronotum, length 105; median width 225; major setae — am 35, aa ?20, ml 35, epim 80, pa 50. Fore wing, length 760; distal width 70; subbasal setae 50, 80, 100; number of duplicated cilia 9. Tergite IV setae, B, 100; B, 55. Tergite VII setae B, 175; B, 210. Tergite IX setae, B, 160; By 145; B, 160. Tube, length 210; basal width 95; apical width 42; terminal setae 130. Antennal segments length, 30; 50; 68; 64; 58; 55; 39; 26. SPECIMENS STUDIED. Holotype 2, WEsT MatraysiA: Kuala Lumpur, on dead branch, 29.xi.1969 (R. G. & F. Andre) (BMNH). Paratypes. West Matraysia: I 92 with data similar to holotype except 24.xi1.1969 (BMNH). Sumatra: Fort de Kock, 920 m, xi. 1920 (Jacobson) (SMF) [this paratype bears a manuscript name]. CoMMENTS. This species appears to be related to brevicollis and rhizophorae but has the dorsal setae (B,) on tergite nine elongate, and all the femora dark brown with a sharply yellow apex. The paratype female from Sumatra is larger than the two specimens from Malaya (body length 2400 wm), and was designated holotype of an unpublished name by Dr H. Priesner. Nesothrips melinus Mound Nesothrips melinus Mound,1974:72-73. Holotype gj, AUSTRALIA : Queensland (ANIC) [examined]. This species was described from four males collected in northern Queensland, Australia, although a fifth male from southern New South Wales may represent the same species. These males are largely yellow, and in structure appear to be intermediate between Nesothrips and the Carientothrips species of the mjobergi complex. SPECIMENS STUDIED. Holotype 2 with 3 ¢ paratypes, AUSTRALIA: Queensland, near Ingham, on dead twigs, 21.vii.1968 (L. A. Mound) (ANIC, BMNH). Nesothrips niger (Moulton & Steinweden) (Text-fig 39) Bolothrips nigra Moulton & Steinweden, 1932 : 167-168. Holotype 9, MarguEsas Is. (BPBM) [examined]. NESOTHRIPS COMPLEX 167 This species is known only from the unique holotype which is mounted under a broken cover glass. Because of this its relationships cannot be determined at present. Contrary to the original description the lengths of the tube and B, setae on tergite nine are 260 wm and 210 pm. SPECIMEN STUDIED. Holotype 9, Marquesas Is.: Hiva Oa, Mt Temetiu, 24.vii.19g29 (Adamson & Mumford) (BPBM). Nesothrips oahuensis Kirkaldy (Text-fig. 31) Nesothrips oahuensis Kirkaldy, 1907 : 103. Syntype 9, Hawartan Is.: Oahu (?BPBM) [not examined]. Oedemothrips laticeps Bagnall, 1910 : 680-681. Syntypes g, 9, Hawauan Is.: Oahu, Lanai (BMNH) [examined]. [Synonymized by Bianchi, 1944.] Nesothrips hawaiiensis, lapsus for oahuensis Kirkaldy, Bianchi, 1944 : 31-38. Bianchi studied a male and female specimen from Kirkaldy’s original material and illustrated these. The present author has examined only two female and one oedymerous male syntypes of Jaticeps. The illustration given by Bianchi clearly refers to a rather small male with the fore femora weakly swollen. The male illustrated here has a greatly enlarged fore femur which is L-shaped as in Rhaebo- thrips species. Moreover the head is prolonged in front of the eyes and the ocellar setae are rather close together. The pelta of this specimen is very broad and slender. SPECIMENS STUDIED. I g syntype of laticebs, HAWAIIAN IsLANDs: Oahu, near Honolulu, 2000-3000 ft, vii. 1900 (Perkins 667); 1 2 syntype, HAawattANn IsLanps: Lanai, 2000 ft, i. 1894 (Perkins 91); 1 9 syntype without data (Perkins 489) (BMNH). Nesothrips propinquus (Bagnall) Oedemothrips propingquus Bagnall, 1916 ; 408-409. Holotype 9, AusTRALIA: Victoria (BMNH) [examined]. Cryptothrips dimidiatus Hood, 1918 : 145-146. Holotype 9, AUSTRALIA: Queensland (USNM) [examined]. [Synonymized by Mound, 1968 : 141.] Oedemothrips propinquus var. breviceps Bagnall, 1924 : 634-635. Syntype 9, NEw ZEALAND (BMNH) [examined]. [Synonymized by Mound, 1968 : 141.] Oedemothrips propinquus form obscuricornis Bagnall, 1924 : 635. Bagnalliella cestosa Karny, 1920 : 41. Holotype 9, AUSTRALIA: Queensland (NR) [examined]. [Synonymized by Mound, 1974 : 73.] Neosmerinthothrips oleriae Moulton, 1949 : 492-494. Holotype 9, SourH Arrica (CAS) [not examined]. [Synonymized by Mound, 1968 : 141.] Bolothrips similis Hartwig, 1948 : 103-108. Holotype 9, SoutH Arrica: Pretoria [not examined]. Syn. n. 168 L. A. MOUND In addition to the above synonymy the nomen nudum ‘Oedemothrips australis Bagnall’ in Kelly & Mayne (1934: 51) also refers to this species. The species is widespread and common in Australia and New Zealand, as well as in South Africa, and has been recorded from St Helena and New Caledonia. Material from all these localities is available in the BMNH collections, together with paratypes of oleriae and similis. In the USNM collections there are three females taken in quarantine at Boston apparently from the Azores and Portugal. Moreover in the BMNH collections there is one female apparently collected at Curitiba in southern Brazil, but this is the only Neotropical record and needs further confirmation. Males and most females of propinquus are apterous but female macropterae are collected occasionally. The variation in head shape in Australia includes the form shown by similis, and the ventral prolongation of the eyes is not constant. In a few gynaecoid males the eyes are not elongate ventrally. The colour of the legs and antennae is also variable, and the form obscuricornis has the legs and antennae largely brown. The anterior setae of the pronotum also vary in length from 15 to 60 pm, but there is no evidence from all this variation that more than one species is involved. Populations from New Zealand appear to show the greatest degree of variation. Nesothrips rhizophorae (Girault) Cryptothrips rhizophorae Girault, 1927(38) : 2. Syntype 2 g, AUSTRALIA: Queensland (QM) [examined]. Nesothrips rhizophorae (Girault) Mound, 1974 : 74-75. This species is very similar to bvevicollis (Bagnall) and may be merely a local colour variant with the basal antennal segments brownish instead of yellow. SPECIMENS STUDIED. AUSTRALIA: Queensland, Brisbane, 3 2, I g syntypes from galls on mangrove, 28.vi.1927 (L. Franzen) (QM); Brisbane, Mt Nebo, 1 9 from shrubs, 27.vii.1968 (L. A. Mound) (BMNH). Nesothrips semiflavus (Moulton) Bolothrips semiflavus Moulton, 1939 : 147-148. Holotype 9, Rapa Is. (BPBM) [examined]. This species is known only from the unique holotype. It appears to be similar to artocarpi in having the head slightly longer than wide, but there are two pairs of long setae behind the eyes, the postoculars 110 ym and the mid-dorsals 70 ym long. The postocellar setae are also 70 um long. The median setae on tergite nine are 155 ym and the tube 175 um long. The major setae appear to be paler and more slender than in other species of Nesothrips. SPECIMEN STUDIED. Holotype 9, Rapa IsLanp: Mt Tevaitahu at 700 ft, 8.vii.1935 (BPBM). NESOTHRIPS Fics 46-51. 46, Nesidiothrips alius. 49, P. ocelloides, pelta. 47 50, N. alius, tube. COMPLEX 47, Phacothrips ocelloides. 48, N. alius, pelta. 51, P. ocelloides, tube. 169 170 L. A. MOUND Nesothrips yanchepi Mound Nesothrips yanchepi Mound, 1974: 75. Holotype 9, AusTRALIA: Western Australia (ANIC) [examined]. This species is similar to niger (Moulton & Steinweden) but has the eyes less prolonged ventrally and the setae on tergite nine shorter. It is also closely related to carvert. SPECIMENS STUDIED. Holotype 2 with 1 9, I g paratypes, AUSTRALIA: Western Australia, Yanchep near Perth, from sedges at lakeside, 29.ix.1967 (L. A. Mound) (ANIC, BMNH). PHACOTHRIPS gen. n. Type-species: Gastrothrips ocellovdes Hood. Medium sized, dark brown species of Idolothripinae, Cryptothripini. Head as in Gastro- thrips, not projecting at anterior, eyes large, stylets wide apart, but with one large isolated ommatidium on each cheek midway between posterior margin of head and posterior margin of eye. Antennae with two sense cones on segment III, four sense cones on segment IV; segment VIII more than 0-6 times as long as VII but barely constricted at base. Fore tarsal tooth well developed in both sexes. Median metanotal setae small. Tube very stout, sides strongly convex, maximum width more than o-5 times the total length, and about 5-0 times the minimum width, terminal setae very weak. Macropterous 9 with no duplicated cilia; pelta triangular with slender lateral wings. Micropterous ¢ sometimes oedymerous with enlarged pronotum, pelta broadly triangular, metanotum and tergites with supernumerary setae. This genus is possibly related to the nigrisetis-group of species from South America which is treated here under Neosmerinthothrips. It is distinguished from other Thysanoptera by the ommatidia on the cheeks and the heavy tube which resembles that of some Pygothrips species. Phacothrips ocelloides (Hood) comb. n. (Text-figs 47, 49, 51, 68) Gastrothrips ocelloides Hood, 1950 : 9-12. Holotype 9, Braz (USNM) [not examined]. This species is here removed from Gastrothrips on account of the four sense cones on the fourth antennal segment, as well as the structure of the head and tube. SPECIMENS STUDIED. BRAZIL: Distrito Federal, Jacarepagua, I 9, I ¢ paratype on dead branches, 14.v.1948 (J. D. Hood); Sao Paulo, Itanhaen, 1 9 paratype (sic) on dead branches, 17.vi.1948 (J. D. Hood & J. Lane) (AMG); Vicosa, on dead branches, 2 9, I J, Vii. 1964, 5 9, I g, xi. 1964 (F. Andre) (BMNH). NESOTHRIPS COMPLEX 171 RHAEBOTHOHRIPS Karny Rhaebothrips Karny, 1913 : 128. Type-species: R. lativentris Karny, by monotypy. A revision and diagnosis of this genus was published recently by Sakimura (1971). However, considerably more material is now available and KRhaebothrips is used here in a broader sense. Two species are described below which have a small fore tarsal tooth in the female and one of these, leveri, is unusually small with a short tube, similar to some Nesothrips species. The other species, doulli, is interest- ing because it is closely related to two further new species from New Zealand which do not have a fore tarsal tooth. One of these species is unique in the genus in having the eyes prolonged on the ventral surface of the head as in some Nesothrips species. The distinction between these two genera is tenuous and likely to become more so with further collecting in the Pacific area. Rhaebothrips species always have a stout pair of setae relatively close together between the posterior ocelli. They are usually larger than Nesothrips species with relatively longer antennae and tube. KEY TO THE SPECIES OF RHAEBOTHRIPS 1 Antennal segment III largely yellow or yellowish brown; V paler at base. : 2 — Antennal segment III brown with pedicel pale; V uniformly dark brown ; 4 2 Fore tarsal tooth present in 9; antennal segment III less than 2-0 times as long as wide (Text-fig. 74) [Fiji] 4 F ; leveri (p. 175) — Fore tarsal tooth absent in 9; antennal sepment TI more fan 3°5 times as long as wide . : F : 3 3 Lateral setae in abdominal segments TH_VII light brown t0 sellowiele lativentris (p. 174) — Lateral setae on abdominal segments III-VIII dark brown to black .nigrisetis (p. 175) 4 Setae B, on tergite IX more than 0:8 times as long as tube, sometimes longer than tube [Pacific Is.] : . major (p. 175) Setae B, on tergite IX less than 0-6 oe as long as tube [New Zealand) : : 5 5 Head o-g-1-0 times as long as tube (Text-fig. 58); antennal segment III 2-1-2-2 times as long as wide (Text-fig. 72) d : . zondagi (p. 176) — Head 1-2 times as long as tube, antennal peemeut IIT; 27 nee 5 times as long as wide (Text-figs 67, 71) , 6 6 Fore tarsal tooth present in 2 only micropterae and hemimacropterae sown eyes not prolonged on ventral surface of head (Text-fig. 52) . ; . doulli (p. 171) — Fore tarsal tooth not present in 2 (only macropterae known); eyes 1-4 times as long on ventral surface of head as on dorsal surface (Text-fig. 53). . eastopi (p. 173) Rhaebothrips doulli sp. n. (Text-figs 52, 55, 67) microptera. Colour dark brown with red internal pigment; tarsi and fore tibiae a little paler; antennae dark brown except for yellow pedicel of segment III; fore wing rudiment pale; major setae not very dark, with finely acute apices. Head longer than wide, cheeks rounded, eyes rather small; ocellar setae arising between posterior ocelli, median setae on vertex sometimes elongate; maxillary stylets retracted to postocular setae; antennae similar in structure to eastopi. Pronotum with median thickening 172 L. A. MOUND Nae fel es ae ee oe aes \ | \\I \ -\ i " Fics 52-57. Rhaeobothrips species: 52, R. doulli, male. 53, R. eastopi. 54, R. leveri. 55, R. doulli, fore leg of male. 56, R. eastopi, tube. 57, R. levert, pelta. NESOTHRIPS COMPLEX 173 developed, epimeral sutures complete; metanotum not sculptured, median setae 504m long and 120 wm apart. Fore femur simple, fore tarsus with a small pointed tooth at apex of inner margin about one-third as long as tarsal width. Fore wing lobes with a few cilia. Pelta broad with small lateral wings; tube with straight sides but slightly constricted near apex. MEASUREMENTS (holotype 9 in wm). Body length 2900. Head length 310; median width 260, ocellar setae 100 long, bases 55 apart; post ocular setae 145; mid dorsal setae 85 (35 in Q paratype). Pronotum, length 175; median width 370; major setae —am 50, aa 50, ml 80-85, epim 120, pa 100-120. Fore wing, length 450; subbasal seta 140. Tergite IV setae, B, 175, B, 100. Tergite VII setae, B, 240; B, 280. Tergite IX setae, B, 145, B, 150, Bg 240. Tube, length 260; basal and apical width, 100, 58; terminal setae 195. Antennal segments length, 50, 70, 105; 100; 85; 77; 55; 40. 6 microptera. Colour similar to 9 (2 ¢ paratypes teneral). Pronotum strongly thickened medially and along anterior border; fore femur enlarged L-shaped; fore tarsal tooth curved, almost as long as tarsal width; major setae of head and pronotum longer than in 9. MEASUREMENTS (paratype gj inym). Body length 2400. Head length 290; ocellar setae 100; postocular setae 160, mid-dorsal seta 50. Pronotum, length 225; median width 350; major setae —am 50, aa 65, ml 145, epim 160, pa 145. Fore wing, length 400; subbasal setae 145, 230. Tube length 225. SPECIMENS STUDIED. Holotype 2, NEw ZEALAND: Christchurch, on dead walnut twig, 1951 (K. M. Doull) (BMNH). Paratypes. NEW ZEALAND: I Q, 4 6 collected with holotype (BMNH) (zr ¢ deposited in DSIR); 1 9, 1 g§ on Cytisus bark, Cave [100 miles south-west of Christ- church], 9.vii.1969 (R. Zondag) (NZFI). CoMMENTS. This species is closely related to eastopt described below which unfortunately is known only from a different morph. The third antennal segment of these two species differs from Jativentris and major in being slightly swollen in the basal third. The female paratype from Cave has fore wings 950 wm long and a rather stouter fore tarsal tooth than the micropterae, but the male with which it was collected cannot be distinguished from the other male paratypes. Rhaebothrips eastopi sp. n. (Text-figs 53, 56, 71) Q macroptera. Colour dark brown, fore tarsi yellowish, fore tibiae light brown; antennae brown to dark brown, III slightly lighter with pedical yellow; fore wings very weakly shaded; major setae light brown and finely acute. Head longer than wide; ocellar setae arising between posterior ocelli, maxillary stylets not retracted to postocular setae, eyes prolonged and narrowed on ventral surface; antennal seg- ment III slightly swollen in basal third, VII with a distinct pedicel, two sense cones on III, four on IV. Pronotum smooth, median thickening not developed, epimeral sutures complete; metanotum weakly reticulate medially, setae 65m long. Fore tarsus angulate on inner surface but without a tooth; fore wing narrow (?partially reduced). Pelta with lateral wings extending almost to lateral margins of tergite II; tube with almost straight margins. MEASUREMENTS (holotype 2 in wm). Body length 2800. Head length 320; median width 255; ocellar setae 65 long, bases 62 apart; postocular setae 135. Pronotum, length 195; median width 310; major setae —-am 35, aa 25, ml 50, epim I00—II0; pa 95-100. Fore wing, length 950; distal width 80; subbasal setae 50, 50; number of duplicated cilia 3-4. Tergite E* 174 L. A. MOUND IV setae, B, 145; B, 80. Tergite VII setae, B, 245; B, 225. Tergite IX setae, B, 145; B, 130; B, 225. Tube, length 260; basal and apical width 100, 52; terminal setae 195. Antennal segments length, 50, 65, 100, 115, 100, 70, 56, 45. SPECIMEN STUDIED. Holotype 9, NEw ZEALAND: South Island, 15 miles S.E. of Greymouth, Moana Kotula, from Uncinia and Juncus, 12.1x.1972 (V. F. Eastop) (BMNH). CoMMENTS. The unique holotype is possibly not fully macropterous. The antennae, particularly segment three, are similar to doulli described above, but that species has a distinct fore tarsal tooth in the female and the eyes are not prolonged ventrally. Rhaebothrips lativentris Karny Rhaebothrips lativentris Karny, 1913 : 129-130. Holotype gf, TAIWAN (? lost) [not examined]. Cryptothrips claripennis Hood, 1919: 90. Holotype 9, AUSTRALIA: Queensland (USNM) | [examined]. [Synonymized by Mound, 1974: 91.] Cryptothrips seychellensis Bagnall, 1921 : 274-276. Lectotype g, SEYCHELLES (BMNH) [examined]. Syn. n. Cryptothrips difficilis Bagnall, 1921 : 276. Holotype 9, SEycHELLES (BMNH) [examined]. Syn. n. Machatothrips ipomoeae Ishida, 1932 : 12-14. Holotype 9, Ponape (Hokkaido Univ.) [not examined]. [Synonymized by Kurosawa, 1968 : 60.] Cryptothrips magnus Moulton, 1928a: 299. Holotype 9, Tarwan (CAS) [not examined]. [Synonymized by Sakimura, 1972 : 668.] Gynaikothrips yuasai Moulton, 1928b : 315. Holotype 9, Tarwan (CAS) [not examined] [Synonymized by Sakimura, 1972 : 668] Rhaebothrips fuscus Moulton, 1942: 15-16. Holotype 9, Guam (BPBM) [not examined]. [Synonymized by Sakimura, 1971 : 393.] Bolothrips australiensis Moulton, 1968 : 118-119. Holotype 9, Lorp Howe Istanp (CAS) [examined]. [Synonymized by Mound, 1974: 91.] This remarkable synonymy is a reflection partly of the variation in structure of lativentris, but more particularly of the low standard of preparation of most of the microscope slides on which these nominal species were based. Sakimura (1971) has given an account of the variation of the species, and moreover distinguished a new species, nigrisetis from Fiji and New Guinea. However, in the opinion of the present author, Jativentris is native to — and has its centre of diversity in — the western Pacific. Individuals with black setae which correspond to the descrip- tion of nigrisetis have been studied from the Solomon Islands and the New Hebrides, and also from Fiji where they were collected together with normal Jativentris. It seems unlikely that two species are involved in this pattern of variation although the names are retained separate for the present. There are specimens of lativentris in the BMNH collection from Samoa, Hawaii, Guam, Solomon Islands, New Guinea, Queensland, Malaya, Seychelles, Jamaica, Cayman Islands and Trinidad. Sakimura (1971 ; 1972) also lists southern Japan, Taiwan, Philippines, Java, Mauritius, Puerto Rico. Cuba, Dominican Republic, Panama, Bahama and Florida. NESOTHRIPS COMPLEX 175 Rhaebothrips leveri sp. n. (Text-figs 54, 57, 74) Q macroptera. Colour dark brown, tarsi yellow, fore tibiae with yellow markings; antennal segments I and VI-VIII brown, II yellow at apex, III and IV yellow but light brown distally, V brown with a yellow pedical; fore wing shaded; major setae dark brown. Head longer than wide; ocellar setae arising on line joining posterior margins of posterior ocelli; postocular setae acute; maxillary stylets retracted to postocular setae; antennal seg- ments VI-VIII compact, two sense cones on III, four on IV. Pronotum transverse, median thickening slender; epimeral sutures just complete; epimeral setae bluntly acute. Metanotum not sculptured medially, setae 304m. Fore tarsal tooth acute but less than half as long as tarsal width. Pelta with slender lateral wings; tergal lateral setae not long, blunt at apex; sides of tube straight. MEASUREMENTS (holotype 9 in wm). Body length 1850. Head length 220; width 210; postocellar setae 60; postocular setae 90. Pronotum, length 105; median width 260; major setae —-am 35, aa 35, ml 58, epim 80-85, pa 50-60. Fore wing, length 750; distal width 65; subbasal setae 40, 58, 58; number of duplicated cilia 5-6. Tergite IV setae B, 80; B, 65. Tergite VII setae, B, 135; B, 100. Tergite IX setae, B, 115; B, 125, B, 135. Tube, length 145; width at base 80, at apex 42; terminal setae 135. Antennal segments length, 26, 50, 60; 50; 42; 45; 35; 20. SPECIMEN STUDIED. Holotype 9, Fiji: Viti Levu, Vatuwaga, 12.vii.1g41 (R. A. Lever) (BMNH). CoMMENTS. This species could equally well be placed in Nesothrips. It is here described in Rhaebothrips partly because it is more closely related to lativentris than to N. oahuensis, and partly because the only other related species with a fore tarsal tooth is R. doulli from New Zealand. Rhaebothrips major Bagnall Rhaebothrips major Bagnall 1928 : 75-76. ?Holotype g, Samoa (?lost) [not examined]. The (?unique) holotype male, not female as stated in the original description, should be in Bagnall’s collection at the BMNH but cannot be found (Mound, 1968). Sakimura (1972) has given a redescription of major from specimens collected in Samoa, and there is one macropterous male in the BMNH collections from the Solomon Islands apparently of this species. Rhaebothrips nigrisetis Sakimura Rhaebothrips nigrisetis Sakimura, 1972 : 400-402. Holotype g, F1j1 (BPBM) [not examined.] Sakimura described this species from nearly eighty specimens collected in Fiji and one male from New Guinea. As mentioned above there are specimens appar- ently of nigrisetis in the BMNH collection from New Hebrides, the Solomon Islands and New Guinea, as well as some from Fiji which were collected with Jativentris, and it is doubtful if these two species are really distinct. The Rhaebothrips speci- mens from New Guinea are particularly difficult. Specimens in one series in the 176 L. A. MOUND BMNH collected on various dates at Bulolo in S.E. New Guinea are similar to nigrisetis but with the antennae darker. Another series in the BPBM from N.W. New Guinea (Irian Barat) near Nabire and Lake Wissel have black setae but are much larger than any other Rhaebothrips which have been studied. Rhaebothrips zondagi sp. n. (Text-figs 58, 72) @ macroptera. Colour brown, head and tube darkest; fore tarsi little paler than femora; antennae dark brown, pedicel of segment III yellow; fore wing uniformly shaded dark brown; major setae brown but not very dark, with finely acute apices. Fic. 58. Rhaebothrips zondagi, male. NESOTHRIPS COMPLEX 177 Head longer than wide, ocellar setae between posterior ocelli; maxillary stylets not retracted to postocular setae; antennae relatively short but similar to those of eastopi. Pronotum with weak median thickening; epimeral sutures complete; median area of metanotum not sculptured, setae 50-60.m long and 50um apart. Fore femur simple, fore tarsus with no tooth; fore wing relatively broad. Lateral wings of pelta not extending fully across anterior margin of tergite II; sigmoid setae well developed on II-VII; tube with straight sides. MEASUREMENTS (holotype 2 inyum). Body length 2450. Head, length 260; median width 225; ocellar setae 70, bases 35 apart; postocular setae too. Pronotum, length 160; median width 275; major setae—am 35, aa 20, md 50, epim go, pa 100. Fore wing, length 1100; distal width 115; subbasal setae 65, 100, 115, number of duplicated cilia 12. Tergite IV setae, B, 125, B, 80. Tergite VII setae, B, 220, B, 195. Tergite IX setae, B, 120; B, 145; B; 195. Tube, length 275; basal and apical width 95, 50; terminal setae 160. Antennal segments length, 40, 65, 90, 80, 70, 65, 50, 35. 6 macroptera. Colour and structure similar to 2; frequently oedymerous with fore femora enlarged and L-shaped, and fore tarsus with a stout tooth equal in length to tarsal width; major setae of head and pronotum longer than in 9; fore wing relatively small. MEASUREMENTS (paratype gin wm). Body length 2300. Head, length 260; median width 210; ocellar setae 90; post ocular setae 135; mid dorsal setae 50-65. Pronotum, length 195; median width 320; major setae —am 55, aa 30, ml 115, epim 100, pa 135. Fore wing, length goo; distal width 85; number of duplicated cilia 8; sub-basal setae 115, 160. Tergite IX setae, B, 100; B, 95; B,; 225. Tube length 255. SPECIMENS STUDIED. Holotype 9, NEw ZEALAND: South Island, Hochstetter State Forest 25 miles east of Greymouth, on Phyllocladus alpinus branches, 13.iii.1970 (W. A. Holloway) (DSIR). Paratypes. NEW ZEALAND: 13 9, 8 ¢ collected with the holotype; 2 9 North Island, Mangawiri Basin, Whiriaki (Urewera), 40 miles south-west of Rotorua, on Podocarpus totara twigs, 13.iv.1972 (P. J. Alma) (DSIR, NZFI, BMNH). CoMMENTS. This species is related to the other two species described above from New Zealand in having short setae on tergite nine. The antennae are shorter than any other species of Rhaebothrips, apart from the aberrant Jeveri from Fiji, but the tube is longer than any species of Nesothrips with which it might be confused. SCOTOTHRIPS Priesner Scotothrips Priesner, 1939: 75. Type-species: Adiaphorothrips elepbhas Karny, by original designation. The synonymy of this genus and its relationships to Dichaetothrips have been discussed recently by Mound (1974). Scotothrips species are found mainly in the Pacific, Indonesian and Australian regions, and the Brazilian species firmus listed below is not typical of the group. Scotothrips claripennis (Moulton) comb. n. Dichaetothrips claripennis Moulton, 1934 : 503. Holotype 29, Hawarr (CAS) [not examined]. Gastrothrips trinidadensis Hood, 1935 : 168-170. Holotype 9, Trrnrpap (USNM) [not examined]. Syn.n. 178 L. A. MOUND Nesothrips indicus Ananthakrishnan, 1968 : 967-969. Syntypes g 9, INp1a: Madras (TNA) [not examined]. Syn.n. Nesothrips diversus Ananthakrishnan, 1972 : 434-436. Holotype 9, INp1a: Madras (TNA) [not examined]. Syn.n. This species was based on a single female collected in Honolulu. This specimen has not been studied, but the above synonymy is based on a female from Honolulu determined as claripennis by Bianchi. This species does not belong in Nesothrips ——) Fics 59-66. Right antennae (III-VIII unless stated otherwise): 59, Bolothrips italicus. 60, Carientothrips acti, III. 61, C. fijiensis, III-IV. 62, C. japonicus, III lateral. 63, C. grayi. 64, Gastrothrips falcatus. 65,G.ruficauda. 66, Diceratothrips validipennis. NESOTHRIPS COMPLEX 179 because of the presence of a fore tarsal tooth in the female, nor in Gastrothrips because of the four sense cones on the fourth antennal segment. The elongate straight-sided tube is typical of the Scotothrips/Dichaetothrips complex, and because the postocellar setae are short claripennis is here placed in Scotothrips. The vertex of this species is narrowed in dorsal view but is very deep. This shape is very susceptible to distortion in mounting, and the claripennis and indicus specimens which have been studied have the head apparently rather broad at the base. The specimens described as diversus are here regarded as small pale forms of the larger species. The most constant characteristic in all this material is the presence of a pair of pores on the metanotum anterolateral to the metanotal setae. The colour of antennal segments three and four is variable, although three is usually yellow with a shaded apex and four is usually light brown. The mid and hind tarsi are usually yellow but sometimes light brown. In the BMNH collections there are several specimens of Dichaetothrips usitatus Ananthakrishnan & Jagadish, 1970 (Dichaetothrips indicus Ananthakrishnan, 1961) which had been misidentified as Nesothrips indicus Ananthakrishnan, 1968 despite the presence of very long postocellar setae. One of the male paratypes of diversus is also a misidentification of usitatus. The species claripennis has probably been transported around the tropics by man. SPECIMENS STUDIED. HAWAIIAN IsLANDs: Oahu, Honolulu, Mckinkey High, 1 9 in Monkey Pod, iv. 1946 (F. A. Bianchi) (BMNH). MEXICO: 3 9, 2 9 on Cocos nucifera seed, in quarantine at US (USNM). Banamas: I 2 on Cocos nucifera seed, I 9 on Poinciana regia pod, in U.S. quarantine (USNM). JAMAICA: Caenwood, I 9, vi. 1970 (K. Heinze) (BMNH); 1 9, 2 g on Cocos nucifera fls, r 9 on Delonix regia pod, in U.S. quarantine (USNM); Trinrpap: St Clair, x 9 paratype of tvinidadensis in hole in dead bamboo, 13.i.1917 (C. B. Williams) (AMG). MOZAMBIQUE: Lourengo Marques, I 2 in Cassia pods, vii. 1936 (J. C. Faure) (AMG); SoutH Arrica: Natal, Winterskloof, 2 9, 1 ¢ in Cassia pods, vii. 1942 (F. Casalis) (AMG). Inp1A: Madras, r 9, 2 3, determined as indicus, ii. 1966, I 2 paratype of diversus, xii. 1970 (J. N. Ananthakrishnan) (BMNH). Scotothrips firmus (Hood) comb. n. Gastrothrips firmus Hood, 1952 : 162. LECTOTYPE 9, Brazit (USNM), here designated {not examined]. The head of this species is unusual in Scotothrips as the cheeks are constricted near, but not at, the base, and the eyes are slightly swollen at their posterior margin. The fore tarsus bears a small tooth in the female, the pronotum is thickened medially and at the anterior margin, the median metanotal setae are small and slender, the pelta broadly triangular with the median pores close together, and the tube is long with straight sides. The specimen here designated as lectotype has not been studied 180 L. A. MOUND but is that specimen which was labelled holotype by Hood but not referred to in his text. The lectotype bears identical data to the paratype listed below and has been given the following number: USNM Type Number 71995. SPECIMEN STUDIED. BraziL: Sao Paulo, Itanhaen, I 2 paratype on dead branches, 17.vi.1948 (J. D. Hood & J. Lane) (AMG). Fics 67-74. Right antennae (III-VIII unless stated otherwise): 67, Rhaebothrips doulli. 68, Phacothrips ocelloides. 69, Nesidiothrips alius. 70, Neosmerinthothrips fructuum. 71, Rhaebothrips eastopi, III. 72, R. zondagi. 73, Neosmerinthothrips variipes. 74, Rhaebothrips leveri. NESOTHRIPS COMPLEX 181 SYNCEROTHAHRIPS Hood gen. rev. Syncerothrips Hood, 1935 : 191-192. Type-species: S. harti Hood, by monotypy. Stannard (1957) synonymised this genus with Nesothrips on the grounds that the last two antennal segments of the type-species of Neosmerinthothrips were more or less closely joined. However, Syncerothrips is closely related to the South American species of Gastrothrips. There are three sense cones on the fourth antennal segment and two on the third, the median metanotal setae are very stout, and the pelta triangular. The only species in the genus differs from Gastrothrips species in having the last two antennal segments fused with only a partial suture on the ventral surface. The fore tarsus does not bear a tooth in the female, and the postocellar setae are long. Syncerothrips harti Hood Syncerothrips harti Hood, 1935 : 192-194. Holotype 9, U.S.A.: Texas (USNM) [not examined]. This species was based on a single female, but the specimen listed below was taken at the type-locality and identified by Hood. SPECIMEN STUDIED. U.S.A.: Texas, Brownsville, 1 2 microptera on dead branches, 2.iii.1939 (J. D. Hood) (USNM). SPECIES REMOVED FROM THE NESOTHRIPS COMPLEX The following three species were described in genera related to Nesothrips, but they are not even members of the subfamily Idolothripinae in its strict sense. Species of the genus Adelothrips have moderately broad maxillary stylets and probably feed on fungus spores, but in view of the structure of the male abdomen it is likely that this genus is derived from Hoplothrips-like members of the Phlaeo- thripinae (Mound, 1974). Adelothrips lativerticis (Post) comb. n. Bolothrips lativerticis Post, 1961 : 140-143. Holotype 9, U.S.A.: Oregon (CAS) [examined]. The original illustration of this species shows five sense cones on the third antennal segment and six on the fourth, although the description gives the actual number correctly as three and four. There are two pairs of wing retaining setae on each tergite, unlike any member of the Cryptothripini discussed in this paper. Moreover the males of Jativerticis, like other members of Adelothrips, have setae B, on tergite nine short and stout, and sternites six and seven have glandular areas. This species is a typical member of A delothrips because of the number of antennal sensoria, the fusion of the last two antennal segments, the position of the maxillary stylets close together in the middle of the head, the bell-shaped pelta, and the abdominal structure detailed above. 182 L. A. MOUND SPECIMENS STUDIED. Holotype 9, U.S.A.: Oregon, Hood River County, Herman Creek, in hollow twigs, 31.1.1946 (R. L. Post) (CAS). U.S.A.: Oregon, Benton County, Corvallis, paratype 9, 23.i11.1946 (Leah & Post) (R. L. Post coll. North Dakota); Washington, Puyallup, 3 9, 2 ¢ in dead willow, 20.iv.1937 (B. Baker) (BMNH). Adelothrips speciosissimus (Karny) Nesothrips speciosissimus Karny, 1920: 42. Holotype g, AUSTRALIA: Queensland (NR) [examined]. Adelothrips speciosissimus (Karny) Mound, 1974 : 16. This species may not be native to Australia as the most closely related species appears to be skwarrae Priesner from Mexico. The male has setae B, on tergite nine short, and sternites six and seven have reticulate glandular areas. SPECIMENS STUDIED. Holotype g, AUSTRALIA: Queensland, Cedar Creek, March (Myoberg) (NR). AUSTRALIA: Queensland, Clump Point, Mission Beach, 2 9 on dead branches, 21.vii.1968 (L. A. Mound 764) (BMNH). Liothrips debilis (Hood) comb. n. Neosmerinthothrips (?) debilis Hood, 1936 : 269-271. Holotype g, PANAMA (USNM) [examined]. This species was described from a single male with stout maxillary guides but with slender maxillary stylets. It is a member of the subfamily Phlaeothripinae despite the fact that all three pairs of major setae on tergite nine are elongate, a characteristic shared with several South American species in the genus Ltothrips. The genus is widespread in the tropics and the species feed on the leaves of green plants. SPECIMEN STUDIED. Holotype 3, PANAMA: Frijoles, on Panicum maximum, 23.vii.1933 (P. C. Standley) (USNM). REFERENCES ANANTHAKRISHNAN, T.N. 1960. Neosmerinthothrips inquilinus sp. nov., an inquiline Tubuli- feran (Thysanoptera) from India. Bull. ent. Loyola Coll. 1 : 32-33. 1961. Studies on some Indian Thysanoptera VI. Zool. Anz. 167 : 259-271. 1964. A contribution to our knowledge of the Tubulifera (Thysanoptera) from India. Opusc. ent. Suppl. 25 : 120 pp. NESOTHRIPS COMPLEX 183 1968. Two new species of Nesothrips with further remarks on Nesothrips robustus (Thysan. Tubulifera). Annis Soc. ent. Fr. (N.S.) 4 : 967-973. 1969. Mycophagous Thysanoptera I. Indian Forester 95 : 173-185. 1970. Onsome species of Nesothrips Kirkaldy with notes on the influence of oedymerism and gynaecoidism in species of allied genera of Cryptothripini (Tubulifera: Thysanoptera). J. z00l. Soc. India 22 : 51-62. 1971. Trends in intraspecific sex-limited variations in some mycophagous Tubulifera (Thysanoptera). J. Bombay nat. Hist. Soc. 67 : 481-501. 1972. Mycophagous Thysanoptera IV. Orient. Insects 6 : 425-437. 1973. Mycophagous Tubulifera of India. Occ. Publ. Entomology Res. Unit, Loyola Coll. 2 : I-144. ANANTHAKRISHNAN, T. N. & JaGapisH, A. 1970. The species of Dicevatothrips Bagnall and allied genera from India (Thysanoptera: Megathripinae: Insecta). Ovient. Insects 4 : 265-280. ARNAUD, JR, P. H. & LEE, V. F. 1973. Types of Thysanoptera in the collection of the California Academy of Sciences. Occ, Pap, Calif. Acad, Sci. 105 : 138 pp. BAGNALL, R. S. 1908. On some new genera and species of Thysanoptera. Trans. nat. Hist. Soc. Northumb. 3 : 182-217. 1910. Thysanoptera. Fauna hawaii.3 : 669-701. 1914. Brief descriptions of new Thysanoptera III. Ann. Mag. nat. Hist. (8) 13 : 287-297. 1916. Brief descriptions of new Thysanoptera VIII. Ann. Mag. nat. Hist. (8) 17 : 397— 412. 1917. Ona collection of Thysanoptera from St. Vincent, with descriptions of four new species. J. zool. Res. 2 : 21-27. 1921a. On Thysanoptera from the Seychelles Islands and Rodrigues. Ann. Mag. nat. Hist. (9) 7 : 257-293. 1921b. Brief descriptions of new Thysanoptera. XI. Ann. Mag. nat. Hist. (9) 7 : 355- 368. 1924. Brief descriptions of new Thysanoptera XIV. Ann. Mag. nat. Hist. (9) 14 : 625- 640. 1927. Contributions towards a knowledge of the European Thysanoptera. II. Ann. Mag. nat. Hist. (9) 19 : 564-575. 1928. On some Samoan and Tongan Thysanoptera, with special reference to Ficus gall-causers and their inquilines. Thysanoptera. Insects Samoa 7 : 55-76. Brancul, F. A. 1944. Nesothrips Kirkaldy supersedes Oedemothrips Bagnall. Proc. Hawaii. ent. Soc. 12 : 31-38. 1945. Introduction to the Thysanoptera of New Caledonia. Proc. Hawaii. ent. Soc. 12 : 249-278. Cott, H. E. 1956. Systematics of the suborder Tubulifera (Thysanoptera) in California. Univ. Calif. Publs Ent. 13 : 1-216. De Santis, L. 1943. Especies nuevas y conocidas de la Republica Argentina. Revta Fac. Agron. La Plata 25 : 89-96. —— 1963a. Adiciones a la Fauna Argentina de Tisanopteros III. Notas comn Invest. Cient. Prov. Buenos Aires 1 (6) : 1-14. —— 19636. Tisanopteros de Tierra del Fuego. Revta Soc. ent. argent. 24 : 63-66. Faure, J.C. 1943. Two new species of Bolothrips (Thysanoptera) from South Africa. J. ent. Soc. sth. Afr. 6 : 86-89. —— 1949. Three remarkable new genera of Phlaeothripidae (Thysanoptera) from South Africa. Entomology Mem. Dep. Agric. Un. S. Afr. 2 : 203-217. 1954@. South African Thysanoptera I. J. ent. Soc. sth. Afr. 17 : 9-26. 1954b. South African Thysanoptera II. J. ent. Soc. sth. Afr. 17 : 145-166. Grrau_Lt, A. A. 1927(37). Thysanoptera Nova Australiensis from Queensland. Published privately. pp. [1] Brisbane. 184 1927(38). —— 1928(42). L. A. MOUND Thysanoptera Nova Australiensis II. Published privately. pp. [1-2] Brisbane. Some Insecta and a new all highness (Notes compiled in fear and sorrow). Pub- lished privately. pp. [1-4] Brisbane. Hartwie, E. K. analyses of measurements. Six new species of South African Thysanoptera with statistical J. ent. Soc. sth. Afr. 11 : 83-126. 1948. HEEGER, E. 1852. Beitrage zur Insecten-Fauna Osterreichs. Sber. Akad. Wiss. Wien. 9: 473-490. Hoop, J. D. 1912. Descriptions of new North American Thysanoptera. Proc. ent. Soc. Wash. 14 : 129-160. 1914. Studies in Tubuliferous Thysanoptera. Proc. Biol. Soc. Wash. 27 : 151-172. 1918. New genera and species of Australian Thysanoptera. Mem. Qd Mus. 6 : 121-150. —— 1919. Two new genera and thirteen new species of Australian Thysanoptera. Proc. biol. Soc. Wash. 32 : 75-92. 1925. New Neotropical Thysanoptera collected by C. B. Williams. Psyche, Camb. 32 : 48-69. 1933. New Thysanoptera from Panama. J] N. Y. ent. Soc. 41 : 407-434. 1935. Some new or little known Thysanoptera of the family Phlaeothripidae. Revita Ent., Rio de J. 5 - 159-199. 1936. Studies in Neotropical Thysanoptera I. Revita Ent., Rio de J. 6 : 248-279. 1937. Studies in Neotropical Thysanoptera IV. Revita Ent., Rio de J. 7 : 255-296. 1938. New Thysanoptera from Florida and Carolina. Revta Ent., Rio de J. 8 : 348-420. 1939. New North American Thysanoptera principally from Texas. Revta Ent., Rio de J. 10 : 550-619. 1941. A century of new American Thysanoptera II. Revita Ent., Rio de J. 12 : 139- 243: 1942. Acentury of new American Thysanoptera III. Revta Ent., Rio de J.12 : 547-678. 1950. Brasilian Thysanoptera II Revta Ent., Rio de J. 21 : 1-113. 1952. Brasilian Thysanoptera III. Proc. biol. Soc. Wash. 65 : 141-176. 1956. Brasilian Thysanoptera VI. Revita bras. Ent. 4 : 51-160. IsH1pa, M. 1932. Fauna of the Thysanoptera in Japan. Insecta matsum. 7 : 1-16. Karny, H. 1913. H. Sauter’s Formosa-Ausbeute : Thysanoptera. Supplta ent. 2 : 127-134. 1916. Beitrage zur Kenntnis der Gallen von Java. Z. wiss. InsektBiol. 1916 : 84-94. 1920. Die neuen australischen Thysanopteren der Mjoberg-Ausbeute. Cas. ésl. Spol. ent. 17 : 35-44. 1925. Onsome tropical Thysanoptera. Bull. ent. Res. 16 : 125-142. KELLty, R. & Mayne, R. J. B. Wales. KIRKALDY, G. W. Kurosawa, M. MorGcaun, A. C. from Porto Rico. Moutton, D. 1934. The Australian Thrips. 81 pp. Glebe, New South 1907. Ontwo Hawaiian Thysanoptera. Proc. Hawaii. ent. Soc.1 : 102. 1968. Thysanoptera of Japan. Insecta matsum. Suppl. 4 : I-94. 1925. A new genus, a new subgenus and seven new species of Thysanoptera Fla ent. 9 : 1-9. 1928a. New Thysanoptera from Formosa. Tvans. nat. Hist. Soc. Formosa 18 : 287-328. 1928b. The Thysanoptera of Japan. New species, notes, and a list of all known Japanese species. Annotnes zool. jap. 11 : 287-337. 1933. The Thysanoptera of South America IV. Revta Ent., Rio de J. 3 : 385-419. 1934. New Thysanoptera of the Hawaiian Islands. Proc. Hawaii. ent. Soc. 8 : 499-503. 1938. Thysanoptera from Minas Geraes, Brazil. Revta Ent., Rio de J.9 : 374-382. 1939. Thysanoptera collected by the Mangarevan Expedition. Occ. Pap. Bernice P. Bishop Mus. 15 : 141-148. 1941. Thysanoptera from Minas Gerais (second paper). Revia Ent., Rio de J. 12 : 314- 322. 1942. Thripsof Guam. Bull. Bernice P. Bishop Mus. 172 : 7-16. 1944. Thysanoptera of Fiji. Occ. Pap. Bernice P. Bishop Mus. 17 : 267-311. NESOTHRIPS COMPLEX 185 1949. New Thysanoptera from Africa. Ann. Mag. nat. Hist. (12) 19 : 481-498. 1968. New Thysanoptera from Australia. Proc. Calif. Acad. Sci. 36 : 93-124. Mou ton, D. & STEINWEDEN, J. B. 1932. New Marquesan Thysanoptera. Bull. Bernice P. Bishop Mus. 98 : 165-168. Mounp, L. A. 1968. A review of R. S. Bagnall’s Thysanoptera collections. Bull. Br. Mus. nat. Hist. (Ent.) Suppl. 11 : 1-181. 1970. Thysanoptera from the Solomon Islands. Bull. Br. Mus. nat. Hist. (Ent.) 24: 83-125. 1972a. Polytypic species of spore-feeding Thysanoptera in the genus Allothrips Hood (Phlaeothripidae). J. Aust. ent. Soc. 11 : 23-36. 1972b. Species complexes and the generic classification of leaf-litter thrips of the tribe Urothripini (Phlaeothripidae). Aust. J. Zool. 20 : 83-103. 1974. Spore feeding thrips (Phlaeothripidae) from leaf litter and dead wood in Aus- tralia. Aust. J. Zool. Suppl. Series 27 : 1-106. Mounp, L. A. & O’NEILL, K. 1974. Taxonomy of the Merothripidae, with ecological and phylogenetic considerations (Thysanoptera). J. nat. Hist. [in press] PELIKAN, J. 1965. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolie. 29 Thysanoptera. Annls hist.-nat. Mus. natn, hung. (Zool.) 57 : 229-239. Post, R.L. 1961. Five new Oregon Thysanoptera. Pan-Pacif, ent. 37 : 137-143. PRIESNER, H. 1921. Neue und wenig bekannte Thysanopteren der neotropischen Fauna aus der Sammlung des Berliner Zoologischen Museums. Dé. ent. Z. 1921 : 187-223. 1922. Ueber albanische Thysanopteren. Wéien. ent. Zig 39 : 105-107. 1925. Katalog der europadischen Thysanopteren. Konowia 4 : 141-159. —— 1926. Die Jugendstadien der Malayischen Thysanopteren. Tveubia 8 : 1-259. 1928a. Die Thysanopteren Europas. 755 pp. Wien. —— 1928b. Uber australische Thysanopteren. Sher. Akad. Wiss. Wien Abt. 1. 137: 643- 659. 1933. Neue Thysanopteren aus Mexico gessamelt von Prof. Dr. A. Dampf. Wien. ent. Zig 50 : 49-63. 1934. Drei neue Phloeothripiden (Thysanoptera). Stylops 3 : 58-63. 1935. New or little known oriental Thysanoptera. Philippine J. Sci. 57 : 351-375. 1937. Two new Phlaeothripidae from Sierra Leone. Ann. Mag. nat. Hist. (10) 20 : 624- 629. 1939. Thysanopterologica VIII. Proc. R. ent. Soc. Lond. (B) 8 : 73-78. 1950. Contributions towards a knowledge of the Thysanoptera of Egypt XV. Bull. Soc. Fouad I. Ent. 34 : 25-37. 1961. Das System der Tubulifera (Thysanoptera). Anz. dst. Akad. Wiss. 13 (1960) : 283— 296. RAMAKRISHNA AyyYaAR, T. V. & MARGABANDHU, V. 1939. Notes on new and known Indian Thysanoptera. Rec. Indian Mus. 41 : 21-33. REvTER, E. 1880. Thysanoptera Fennica I. Tubulifera. Bidy. Kann. Finl. Nat. Folk. 40 : 1-26. SakimuRA, K. 1971. A review of the genus Rhaebothrips Karny (Thysanoptera: Phlaeo- thripidae). Pacif. Insects 13 : 391-403. 1972. Synonymies and collection record of Rhaebothrips lativentris Karny (Thysanoptera: Phlaeothripidae). Pacif. Insects 14 : 668. ScuHmutTz, K. 1913. Zur Kenntnis der Thysanopteren Fauna von Ceylon. Sber. Akad. Wiss. Wien Abt. 1. 122 : 991-1089. SHUMSHER, S. 1947. Studies on Indian Thysanoptera II. Rec. Indian Mus. 45 : 201-206. STANNARD, L. J. 1957. The phylogeny and classification of the North American genera of the sub-order Tubulifera (Thysanoptera). Illinois biol. Monogr. 25 : 200 pp. 1968. The Thrips, or Thysanoptera of Illinois. Bull. Ill. nat. Hist. Surv. 29 : 215-552. THomasson, G. L. & Post, R. L. 1966. North Dakota Tubulifera (Thysanoptera). North Dakota Insects Publication 6 : 1-56. 186 L. A. MOUND UzeLt, H. 1895. Monographie der Ordnung Thysanoptera. 472 pp. Konigratz. Watson, J. R. 1933. Two new species of Oedaleothrips with notes on other species. Fla ent. 17 : 48-50. ZUR STRASSEN, R. 1972. Drei neue neotropische Phlaeothripiden-Arten aus dem Regenwald von Kolumbien. Senckenberg. biol. 53 : 269-279. INDEX Synonyms and invalid names are in italics. abditus, 138 Acallurothrips, 110 acti, 126 acuticornis, 139 Adelothrips, 181 affinis, 150 alius, 157 alticola, 140 Allothrips, 111 andrei, 118 annulipes, 150 anolis, 142 aoristus, 160 apterus, 127 arenarius, 121 artocarpi, 161 australicus, 131 australiensis, 174 australis, 168 badius, 127 bagnalli, 119 bicolor, 118 Biconothrips, 111 biformis, 127 Bolothrips, 117 Botanothrips, 117 brachyurus, 121 brasiliensis, 138 breviceps, 167 brevicollis, Neosmerinthothrips, 151 brevicollis, Nesothrips, 162 brevicornis, 118 callipus, 142 capitalis, 143 capricornis, 127 Carientothrips, 125 carverae, 163 casuarinae, 128 cestosa, 167 ceylonicus, 152 cinctus, 119 cingulatus, 119 citriceps, 136 claripennis, Cryptothrips, 174 claripennis, Scotothrips, 177 Coenurothrips, 148 collaris, 151 corvus, 142 cybele, 139 debilis, 182 denticulatus, 128 dentipes, 119 dentis, 120 Diceratothrips, 133 Dichaetothrips, 134 difficilis, 174 dimidiatus, 167 diversicolor, 152 diversus, 178 dominicanus, 151 doulli, 171 eastopi, 173 embyotyi, 120 falcatus, 143 fijiensis, Carientothrips, 128 fijiensis, Neosmerinthothrips, 152 firmus, 179 flavitibia, 131 fodinae, 163 formosensis, 162 fructuum, 152 fulvicauda, 143 fulviceps, 144 fumipennis, 144 fuscicauda, 151 fuscus, 174 Galactothrips, 148 Gastrothrips, 134 gilvipes, 120 gloveri, 134 grayi, 129 hambletoni, 138 harti, 181 hawatiensis, 167 hemidiscus, 164 hilaris, 153 hoodi, 153 Hoplandrothrips, 111 icarus, 121 incisus, 131 indicus, 178 inquilinus, 154 insularis, 121 intonsus, 145 ipomoeae, 174 italicus, 122 japonicus, 130 karnyi, 162 laticeps, 167 lativentris, 174 lativerticis, 181 leveri, 175 Liothrips, 182 litoreus, 120 loisthus, 130 magnetis, 131 magnus, 174 major, 175 malaccae, 164 mandiocae, 145 marshalli, 151 melinus, 166 milleforme, 150 minor, 162 miskoi, 131 mjobergi, 131 mongolicus, 145 monticola, 146 INDEX 187 Neosmerinthothrips, 148 Nesidiothrips, 156 Nesothrips, 158 niger, 166 nigra, 166 nigricans, 163 nigrisetis, Neosmerinthothrips, 154 nigrisetis, Rhaebothrips, 175 noumeae, 139 oahuensis, 167 obscuricornis, 167 ocelloides, 170 oeceticola, 145 Oedemothrips, 158 oleriae, 167 pallipes, 121 parvidens, 154 paulistarum, 154 pedicillus, 132 Phacothrips, 170 picticornis, 155 pictilis, 132 plaumanni, 155 pratensis, 122 Probolothrips, 134 procerus, 147 propinquus, 167 proteus, 142 reedi, 132 rhachiphilus, 123 Rhaebothrips, 171 Rhinoceps, 112 rhizophorae, 168 robustus, 155 ruficauda, 147 schafferi, 123 Scotothrips, 177 semiflavus, 168 semirufus, 132 seychellensis, 174 similis, 167 skwarrae, 182 speciosissimus, 182 stygicus, 147 subulatus, 148 Syncerothrips, 181 188 L.A. MOUND texanus, 148 varius, 123 trinidadensis, 177 vesper, 133 tuberculatus, 121 xylebori, 156 validipennis, 133 yanchepi, 170 validus, 157 yuasat, 174 varlipes, 156 zondagi, 176 L. A. Mounp, B.Sc., D.I.C., D.T.A. Department of Entomology British MusEUM NATURAL HISTORY CROMWELL RoapD Lonpon SW7 5BD i> a ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. Sanps, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures,g maps. February, 1967. £3.50. HemMinc, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. {8. . Mounp, L. A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . ArtF1, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 51 text- figures. December, 1968. £5. . CRoSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Eutot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 68 plates, 15 text-figures. October, 1971. £12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CrosskEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CrosskEy, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND L! A REVIEW OF THE SUBFAMILY ix ys" OXYINAE (ORTHOPTERA : ACRIDOIDEA) RARY & \' D. HOLLIS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 6 LONDON : 1975 rN Mf ‘SS GENERAL A REVIEW OF THE SUBFAMILY OXYINAH 27 JAN i975 (ORTHOPTERA : ACRIDOIDEA) BY. DAVID HOLLIS Ie Pp. 189-234; 69 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 6 LONDON : 1975 BRARY > \4 L! < ~TO,12/ THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), tnstituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially. for each Department. This paper 1s Vol. 31 No. 6 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) © Trustees of the British Museum (Natural History), 1975 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 13 January, 1975 Price £2°75 A REVIEW OF THE SUBFAMILY OXYINAE (ORTHOPTERA : ACRIDOIDEA) By D. HOLLIS CONTENTS Page SYNOPSIS. , . : : = : : : ‘ : : IQI INTRODUCTION : F F : : ; : : ; ‘ 191 ACKNOWLEDGEMENTS ‘ : ; ;: : ; ‘ ; : 192 HISTORICAL. ; : : 193 EVALUATION OF PRESENT CLASSIFICATION AND CHARACTERS USED ; 194 SUBFAMILY OXYINAE : : : : : : y . 198 Key to the genera of Oxyinae : : 5 ; ; 200 Generic definitions and lists of jachaded species : : ; 201 REFERENCES . : 4 : ; : ‘ ; : F ; 229 INDEX . : : ; . : : : : ‘ ; : 232 SYNOPSIS The Old World Acridid subfamily Oxyinae is reviewed and redefined. A key is given to the 2I genera now included in the subfamily. Each genus is defined and a list is given of its included species. Of the 169 described species in the subfamily, 125 are regarded as valid. 125 primary types, including those of the genus Oxya, have been examined and as a result 11 new generic synonyms, two new specific synonyms and 25 new combinations are proposed. One new genus is described; one generic name, one specific name and three combinations ate revived. INTRODUCTION DuRING a recent study on the genus Oxya (Hollis, 1971) it became clear that (i) the genus Oxya did not represent a monophyletic entity, (ii) the derivations of the various species-groups involved in Oxya were obscure, and (iii) the current definition and scope of the subfamily Oxyinae (Dirsh, 1961) needed some form of revision and rearrangement. At that time it was thought practical to publish the results of the study on Oxya separately (Hollis, 1971) considering the genus as a convenient entity but pointing out that some of its species-groups should be transferred to other genera and that study was continuing on the whole subfamily. The results of this study are presented below in review form as it was found impractical, both in terms of time and available information, to make a complete systematic revision of the group. This review is based on a study of the extensive collections of Oxyinae housed in the British Museum (Natural History), London; the Museum fiir Naturkunde der Humboldt-Universitat, Berlin; the Instituto Espafiol de Entomologia, Madrid; 192 D. HOLLIS and the collection of the late C. Willemse. Type and other material was borrowed from institutions in Basle, Eberswalde, Geneva, Leiden, Oxford, Paris and Stockholm. Drawings of the parts of the male phallic complex were made after extraction and maceration in Io per cent. potassium hydroxide solution, and the abbreviations used in these figures are based on those used in Hollis (1971). Drawings of other features were made from dry material. Type-depositories of species included in the subfamily are given in abbreviated form as follows. NM, Basle Naturhistorisches Museum, Basle. MNHU, Berlin Museum fiir Naturkunde der Humboldt-Universitat, Berlin. BMNH British Museum (Natural History), London. IRSNB, Brussels Institut Royal des Sciences Naturelles de Belgique, Brussels. UZM, Copenhagen Universitetets Zoologiske Museum, Copenhagen. DEI, Eberswalde Deutsches Entomologisches Institute, Eberswalde, D.D.R. MHN, Geneva Muséum d’Histoire Naturelle, Geneva. MCSN, Genoa Museo Civico di Storia Naturale, Genoa. RNH, Leiden Rijksmuseum van Natuurlijke Historie, Leiden. ZI, Leningrad Zoological Institute, Academy of Sciences of U.S.S.R., Leningrad. IEE, Madrid Instituto Espafiol de Entomologia, Madrid. UM, Oxford University Museum, Oxford. MNHN, Paris Muséum National d’Histoire Naturelle, Paris. ANS, Philadelphia | Academy of Natural Sciences, Philadelphia. ZIHU, Sapporo Zoological Institute, Hokkaido University, Sapporo. NR, Stockholm Naturhistoriska Rijksmuseum, Stockholm. MRAC, Tervuren Musée Royal de |’Afrique Centrale, Tervuren. ZIUU, Uppsala Zoologiska Institutonen, Uppsala. NM, Vienna Naturhistorisches Museum, Vienna. USNM, Washington U.S. National Museum, Washington. coll. Willemse Dr Fer Willemse, Eygelshoven, Laurastraat 67, Netherlands ACKNOWLEDGEMENTS I would like to express my gratitude to the Centre for Overseas Pest Research for supporting this work and providing funds for visits to museums in Berlin and Madrid. These visits were made most productive and pleasant by the help and hospitality of Dr K. K. Giinther, of the Museum fiir Naturkunde der Humboldt- Universitat, Berlin, and Sefiora Vincente Llorente, of the Instituto Espajfiol de Entomologia, Madrid. I am indebted to the following colleagues for loans of type and other material: Dr C. Baroni-Urbani, Basle; Dr M. Descamps and Dr M. Donskoff, Paris; Dr B. Hauser, Geneva; Dr A. Kaltenbach, Vienna; Dr T. Kronestedt, Stockholm; Dr B. Petersen, Eberswalde, D.D.R; Dr H. Weidner, Hamburg; and Dr Fer Willemse, Eygelshoven, Netherlands. This is an opportune moment for me to thank Dr L. L. Mishchenko, of the Academy of Sciences, Leningrad, for his help in lending type-material of the genus Oxya and to apologize for not acknowledging his valuable help towards the production of my previous paper (Hollis, 1971). REVIEW OF THE OXYINZ 193 HISTORICAL Although first named by Brunner (1893 : 136) as the ‘Oxyae’, the group was recognised earlier by Stal (1878: 45) as ‘part 1 of Division 12’ in his key to Orthopteran genera. I. Bolivar (1918) redefined the group as ‘Section Oxyae’ of the family Acrididae; Willemse (1921) listed the genera he considered to belong to the group Oxyae of the subfamily Cyrtacanthacrinae occurring in the austro- oriental region; Rehn (1957), in his study of the Australasian Acrididae, gave the TABLE I Genera included by various authors in the Oxyinae Genus Stal Brunner’ Bolivar Willemse Rehn Dirsh (1878) (1893) (1918) = (1921) (1957) (1961) Quilta v v v v v Gesonia (now Gesonula) Oxya Caryanda Digentia Bermius Hieroglyphus Tauchiva Racilia Badistica Hieroceryx (now Parahieroglyphus) Chitaura Fer Genditia Carydana Gerista Tauchiridea Maga Cledva (now Cercina) Gerunda Pododula Theomolpus Bermiella ee ee ee ee a Se Ss ee Se << SS Re SS 397- 202 DD. HOLLIS Paratraulia Willemse, 1925a : 355. Type-species: Tvaulia obliquiannulata Brunner von Wattenwy]l, by original designation. [Synonymised by Ramme, 1941 : 114.] Servillenia Ramme, 1941 :121. Type-species: Acridium abbreviatum Audinet-Serville, by monotypy. Syn. n. Diacnosis. Head conical; fastigium of vertex parabolic, wider than long, median longitudinal carinula weakly present; frontal ridge sulcate, complete to clypeus, lateral facial keels complete. Eyes normal or slightly bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process transverse, antero-posteriorly flattened, apex trilobate. Dorsum of pronotum rugulose or subrugulose, slightly flattened, median carina weak, lateral carinae absent, crossed by three transverse sulci; relative width of mesosternal interspace variable. Tegmina and hind wings fully developed or shortened to brachypterous condition; parallel transverse stridulatory veinlets present, sometimes weakly, in 1st radial or medial area. Lower genicular lobe of hind femur spined; hind tibia very weakly expanded apically, with acute upper margins; external apical spine of hind tibia absent. In male 1oth abdominal tergite with a weak furcula on postero-dorsal margin; epiphallus with narrow, divided bridge and well developed ancorae. Female ovipositor valves long, evenly toothed. DISTRIBUTION. China, W. Malaysia, W. Indonesia. SPECIES INCLUDED Tauchira abbreviata (Audinet-Serville) comb. n. Acridium abbreviatum Audinet-Serville, 1839 : 678. Holotype 3, Java (lost). Neotype 4, Java (MNHM, Berlin), designated by Ramme (1941 : 123) [examined]. T. polychroa (Stal) Oxya polychroa Stal, 1875 : 32... Holotype 9, MaLaya (Malacca) (NR, Stockholm). Paratraulia bifasciata Willemse, 19254 : 357, fig. Holotype J, Matava (BMNH) [examined]. [Synonymized by Willemse, 1930 : 190 (footnote). ] T. buae I. Bolivar Tauchiva buae I. Bolivar, 1898 :92. Syntypes g and 9, Sumatra (MCSN, Genoa; IEE, Madrid) [g' examined]. T. obliquiannulata (Brunner von Wattenwy]) Traulia obliquiannulata Brunner von Wattenwyl, 1898: 249. Holotype 9, SUMATRA (? depository). T. rufotibialis (Willemse) Paratraulia rufotibialis Willemse, 1925a : 358. Holotype 9, StIncaporE (BMNH) [examined]. Paratraulia elegantula Willemse, 1930 : 190, fig. 98. Syntypes g and 9, Sumatra (RNH, Leiden). [Synonymised by Willemse, 1956 : 202]. T. karnyi (Willemse) Paratraulia karnyi Willemse, 1925a : 359. Holotype g, Matava (BMNH) [examined]. T. grandiceps (Willemse) Paratraulia grandiceps Willemse, 1928: 9. Holotype 2, SumaTRA (RNH, Leiden). T. gressitti Tinkham Tauchira gressitti Tinkham, 1940 : 304, pl. Io, fig. 4, pl. 13, figs 2, 2a. Holotype g, CHINA (Hainan) (? depository). Discussion. The neotype male of 7. abbreviata differs from other species of Traulia in that the dorsum of the pronotum is slightly less rugulose and flattened REVIEW OF THE OXYINZ 203 and the parallel stridulatory veinlets of the tegmen are weaker but the latter is probably due to the shortening of the tegmen. These slight differences are not considered significant enough to warrant separate generic status for the species and Servillenia is synonymised with Tauchira. 28 29 Fics 20-30. Oxyinae genera, male epiphalli. 20, Thanmoia olivacea (Willemse); 21, T. maculata (Willemse); 22, T. gustavi Ramme; 23, Fer coeruleipennis Willemse; 24, Tauchiva polychroa (Stal); 25, T. abbreviatum (Audinet-Serville) ; 26, Chitaura mirabilis (Carl); 27, C. lucida (Krauss); 28, C. indica Uvarov; 29, C. maculata (Willemse) ; 30, Oxytauchiva aurora (Brunner von Wattenwy)). 204 D.. HOLLIS Oxytauchira Ramme, 1941 (Lext-fig. 30) Oxytauchiva Ramme, 1941 :117. Type-species: Tauchiva gracilis Willemse, by monotypy. Diacnosis. Head conical; fastigium of vertex, from above, parabolic, about as wide as long, median longitudinal carinula weakly present; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes normal. Antenna about as long as combined lengths of head and pronotum. Prosternal process transverse, antero-posteriorly flattened, trilobate apically. Dorsum of pronotum weakly flattened, median carina weak, lateral carinae absent, shallowly crossed by three transverse sulci; mesosternal interspace longer than wide. Tegmen and hind wing fully developed, no trace of transverse parallel stridulatory veinlets on tegmen. Lower genicular lobe of hind femur spined; hind tibia hardly expanded apically but with acute upper margins, external apical spine of hind tibia present. In male 1oth abdominal tergite with a furcula on postero-dorsal margin; epiphallus with narrow divided bridge and developed ancorae. Female ovipositor valves long, slender, unevenly spined. DISTRIBUTION. Burma, Sulawesi. SPECIES INCLUDED Oxytauchira gracilia (Willemse) Tauchira gracilis Willemse, 1931a : 242, fig. 99. Syntypes 2 9, SuLawesi (Central) (NM, Basle) [1 2 examined]. O. aurora (Brunner von Wattenwyl) comb. n. Racilia aurora Brunner von Wattenwyl, 1893 : 155, pl. 5, fig. 53. Holotype J, Burma (MCSN, Genoa). Discussion. The type of O. aurora was unavailable for study and the combination suggested here is based on a study of material in the BMNH collections thought to be this species. O. aurora is clearly not congeneric with other species of Raczlia and the genus is returned to the Catantopinae and is thought to be related to Gerunda. Chitaura I. Bolivar, 1918 (Text-figs 6, 26-29) Chitaura I. Bolivar, 1918 : 32. Type-species: Chitaurva brachyptera I. Bolivar, designated by Willemse, 1931a : 244. Tauracris Willemse, 1931a : 248. Type-species: Tauracris flavolineata Willemse, by monotypy. [Synonymised by Ramme, 1941 : 135.] Baliacris Willemse, 1938 : 116. Type-species: Baliacris maculata Willemse, by monotypy. Syn. n. Chitaurella Ramme, 1941 : 136. Type-species: Tauchira lucida Krauss, by original designation. Syn. n. Diacnosis. Head conical; fastigium of vertex, from above, wider than long, pentagonal or rounded triangular, median longitudinal carinula weakly present; frontal ridge sulcate, REVIEW OF THE OXYIN 205 complete to clypeus; lateral facial keels complete. Eyes normal; antenna about as long as or slightly longer than combined lengths of head and pronotum. Prosternal process antero- posteriorly flattened, trilobate apically. Dorsum of pronotum subcylindrical, median carina weak, lateral carinae absent, shallowly crossed by three transverse sulci; mesosternal interspace as long as or slightly longer than wide. Tegmina and hind wings reduced to micropterous condition, scale-like, tegmen extending to posterior margin of first abdominal sclerite. Lower genicular lobe of hind femur spined; hind tibia slightly expanded apically with acute upper margins; external apical spine of hind tibia small but usually present. In male roth abdominal tergite with furcula on hind margin; epiphallus with divided bridge and clearly defined ancorae. Female ovipositor valves long, slender, serrated. DISTRIBUTION. S. India, W. Indonesia, Sulawesi, Maluku. SPECIES INCLUDED Chitaura lucida (Krauss) comb, rev. Tauchiva lucida Krauss, 1903 : 760, pl. 67, fig. 7. Syntypes ¢ and 9, Java (? Jena Museum). C. mirabilis (Carl) Tauchiva mirabilis Carl, 1916 : 472. Syntypes ¢ and 9, SuLAweEsr (Central) (MHN, Geneva) [examined]. C. samanga (Carl) Tauchiva samanga Carl, 1916: 474. Holotype g, SuLawesi (Central) (MHN, Geneva) [examined]. C. vidua (Carl) Tauchiva vidua Carl, 1916 : 475. Holotype 9, SuLAwesi (Central) (MHN, Geneva) [examined]. C. brachyptera I. Bolivar Chitaura brachyptera 1. Bolivar, 1918 : 33. Holotype 9, SuLtawesr (North) (IEE, Madrid) [examined]. C. indica Uvarov Chitaura (?) indica Uvarov, 1929 : 553. Holotype g, Inp1A (South) (BMNH) [examined]. C. flavolineata (Willemse) comb. n. Tauracris flavolineata Willemse, 1931a : 248, figs 102, 103. Holotype 9, SuLawesr (North) (NM, Basle) [examined]. C. maculata (Willemse) comb. n. Baliacris maculata Willemse, 1938 : 117. Holotype g, Baxi I. (coll. Willemse) [examined]. C. striata Willemse Chitaura striata Willemse, 1938 : 117. Holotype 9, Kry Is. (RNH, Leiden). C. elegans Ramme Chitauva elegans Ramme, 1941 :128. Holotype g, SULAWEsI (South) (MNHU, Berlin) [examined]. C. atrata Ramme Chitaura atrata Ramme, 1941 :129. Holotype g, SuLawes1 (Central) (MNHU, Berlin) [examined]. C. mengkoka Ramme Chitaura mengkoka Ramme, 1941 : 131. Holotype g, SuLtaweEsi (South) (MNHU, Berlin) [examined]. C. ochracea Ramme Chitaura ochracea Ramme, 1941 : 132. Holotype g, SuLaweEs1 (South) (MNHU, Berlin) [examined]. ** 206 D: HOLLIS C. moluccensis Ramme Chitaura moluccensis Ramme, 1941 : 133. Holotype gj, Matuxu (Amboina) (NM, Vienna). C. poecila Ramme Chitaura poecila Ramme, 1941: 131. Holotype g, SuLtawesi (North) (MNHU, Berlin) [examined]. C. bivittata Ramme Chitauva bivittata Ramme, 1941 : 133. Holotype 9, Matuxu (Ceram) (MNHU, Berlin) [examined]. Discussion. Ramme (1941 : 135) first suggested Tawracris may be synonymous with Chitaura and after examining Willemse’s type of 7. flavolineata, a female dried from spirit, this synonymy is confirmed. The type male of Baliacris maculata lacks the external apical spine on both hind tibiae but this character appears to be very unstable in Chitaura, often being present on one hind leg and absent on the other. In general appearance this species is very similar to C. lucida and, on the evidence from external morphology and the structure of the male phallic complex, is placed in Chitaura. C. lucida differs slightly from the bulk of the other species of Chitaura in that the dorsum of the pronotum is less rounded and the transverse sulci are slightly deeper, but these features do not warrant separate generic status and it is recombined with Chitaura. Lucretilis Stal, 1878 (Text-fig. 55) Lucretilis Stal, 1878 : 41, 85. Type-species: Lucretilis taeniata Stal, by monotypy. Diacnosis. Head conical; fastigium forming a stepped join with vertex, pentagonal from above, wider than long, median longitudinal carinula very weak or absent; frontal ridge sulcate but present only down to median ocellus; lateral facial keels absent or if present then disguised by rugulosity of facial integument. Eyes bulbous. Antenna very much longer than combined lengths of head and pronotum. Prosternal process transverse in apical half, trapezoid in anterior view. Dorsum of pronotum rugulose and coarsely pitted, rounded, shallowly crossed by three transverse sulci, median carina very weak, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings somewhat reduced, extending to about half length of abdomen, former without trace of transverse parallel stridulatory veinlets. Hind femur tuberculate, lower genicular lobe spined; hind tibia very weakly expanded in apical half, upper margins acute apically; external apical spine of hind tibia present. In male toth abdominal tergite with or without furcula on postero-dorsal margin; epiphallus with narrow divided bridge, ancorae and only an outer pair of lophi. Female ovipositor valves long, slender, completely smooth. DISTRIBUTION. Malaysia, W. Indonesia. SPECIES INCLUDED Lucretilis taeniata Stal Lucretilis taeniata Stal, 1878 : 85. Holotype g, Sumatra (NM, Vienna). L. antennata I. Bolivar Lucretilis antennata I, Bolivar, 1898 : 88. Holotype gj, Sumatra (IEE, Madrid) [examined]. REVIEW OF THE OXYINZ 207 L. bolivari Miller Lucretilis bolivari Miller, 1934 : 531, fig. 1, pl. 13, fig. 3. Holotype 9, Maravya (BMNH) [examined]. L. uvarovi Miller Lucretilis uvarovi Miller, 1935 : 695, fig. 5, pl. 14, fig. 5. Holotype g, Matava (BMNH) [examined]. L. maculata Willemse Lucretilis maculata Willemse, 1936 : 202. Holotype 3g, BorNrEo (Central) (RNH, Leiden). L. splendens Willemse Lucretilis splendens Willemse, 1938 : 118, fig. 1. Holotype 3, Bornro (North) (UM, Oxford) [topotypic material examined]. L. dohrni Ramme Lucretilis dohrni Ramme, 1941 : 98, pl. 13, fig. 8. Holotype 9, Sumatra (MNHU, Berlin) [examined]. L. jucunda Miller Lucretilis jucunda Miller, 1953 : 142, fig. 1. Holotype g, Matava (BMNH) [examined]. Ochlandriphaga Henry, 1933 (Text-figs 5, 58) Ochlandriphaga Henry, 1933 : 187. Type-species: Ochlandriphaga xanthelytrana Henry, by monotypy. Diacnosis. Head conical; fastigium of vertex forming stepped join with vertex, from above pentangular, without median longitudinal carinula; frontal ridge weakly sulcate, complete to clypeus; lateral facial keels weak and incomplete. Eyes bulbous. Antenna much longer than combined lengths of head and pronotum. Prosternal process antero-posteriorly flattened, with weakly trilobate apex. Dorsum of pronotum rounded, coarsely pitted, moderately deeply crossed by three transverse sulci, median carina very weak, lateral carinae absent; mesosternal interspace slightly longer than wide. Tegmina and hind wings reduced, scale-like, former not extending beyond middle of 3rd abdominal tergite. Lower genicular lobe of hind femur spined; hind tibia hardly expanded apically but dorsal margins acute in apical third; external apical spine of hind tibia present. In male 1oth abdominal tergite without furcula; epiphallus with narrow, divided bridge, without ancorae and only with outer pair of lophi. Female ovipositor valves long, slender, evenly toothed. DISTRIBUTION. Ceylon. SPECIES INCLUDED Ochlandriphaga xanthelytrana Henry Ochlandriphaga xanthelytrana Henry, 1933 : 187, pl. 34, figs 1-5. Holotype gj, CeyLon (BMNH) [examined]. Thanmoia Ramme, 1941 (Text-figs 9, 20-22) Thanmoia Ramme, 1941 : 119. Type-species: Thanmoia gustavi Ramme, by monotypy. Annamacris Willemse, 1956 : 23. Type-species: Annamacris olivacea Willemse, by monotypy. Syn. n. : Oxyacris Willemse, 1956 : 23. Type-species: Oxyacris maculata Willemse, by monotypy. Syn. n. 208 D2 HOLLIS Diacnosis. Head conical; fastigium of vertex, from above, rounded pentagonal, wider than long, median longitudinal carinula absent; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes normal. Antenna longer than head and pronotum combined. Prosternal process slender, conical, with subacute or rounded apex. Dorsum of pronotum from subrugulose to smooth, cylindrical or slightly flattened, median carina weak, moderately deeply or shallowly crossed by three transverse sulci, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings fully developed or somewhat shortened; no trace of transverse parallel stridulatory veinlets on tegmen. Lower genicular lobe of hind femur spined; hind tibia expanded in apical half with acute upper margins; external apical spine of hind tibia absent, or very rarely present. In male 1oth abdominal tergite with weak furcula on hind margin; epiphallus with narrow divided bridge and ancorae (Text- fig. 20). Female ovipositor valves long, slender, with serrate apices (Text-fig. 9), obviously highly specialized. DISTRIBUTION. Vietnam. SPECIES INCLUDED Thanmoia gustavi Ramme Thanmoia gustavi Ramme, 1941 : 120, pl. 16, fig. 5. Holotype 9, NortH VietNAM (MNHU Berlin) [examined]. T. olivacea (Willemse) comb. n. Annamacris olivacea Willemse, 1957 : 480. Holotype g, NortH ViETNAM (MNHN, Paris) [examined]. T. maculata (Willemse) comb. n. Oxyacris maculata Willemse, 1957 : 481, pl. 13, fig. 5. Holotype g, SoutH ViIETNAmM (coll. Willemse) [examined]. Discussion. Superficially these three species appear quite different; 7. gustavi is a relatively large, robust subrugulose species with fully developed temina and hind wings; 7. maculata is smaller, more delicate, with smooth integument and is brachypterous; 7. olivacea is rather intermediate. The male genitalia are very similar in all three species, the epiphallus has a narrow, divided bridge, ancorae, boot-shaped outer lophi and simple inner lophi. I do not consider it necessary to retain these three species in monotypic genera and the above synonymy is made. Apart from the type-specimens there is a female paratype of T. gustavai in NM, Vienna and a male and two females of the same species from the type-locality in IEE, Madrid. Fer I. Bolivar, 1918 (Text-fig. 23) Fer I. Bolivar, 1918 :17. Type-species: Fer coeruleipennis I. Bolivar, by monotypy. Diacnosis. Head conical; fastigium of vertex, from above, rounded triangular, about as wide as or slightly wider than long, median longitudinal carinula weakly present; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes slightly bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process long, weakly flattened REVIEW OF THE OXYIN# 209 posteriorly, slightly expanded subapically and with subacute apex. Dorsum of pronotum rugulose, slightly flattened, shallowly crossed by three transverse sulci, median carina present but weak, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings fully developed, former with transverse parallel stridulatory veinlets present in 1st radial area. Lower genicular lobe of hind femur spined; hind tibia very weakly expanded in apical half, with acute upper margins apically; external apical spine of hind tibia present but very weak. In male roth abdominal tergite with a furcula on postero-dorsal margin; epiphallus with divided bridge, developed ancorae and two pairs of lophi. Female ovipositor valves slender, evenly toothed. DISTRIBUTION. Vietnam. SPECIES INCLUDED Fer coeruleipennis I. Bolivar Fer coeruleipennis 1. Bolivar, 1918: 19. Syntypes, 1 g and 1 9, NortH ViEtNAM (IEE, Madrid) [examined]. Gesonula Uvarov, 1940 (Text-figs 8, 44) Gesonia Stal, 1878 : 46. Type-species: Acridium (Oxya) punctifrons Stal, by monotypy. [Homonym of Gesonia Walker, 1858 : 75.] Gesonula Uvarov, 1940 : 174. [Replacement name for Gesonza Stal.} DiaGnosis. Head conical; fastigium, from above, parabolic, slightly longer than the basal width, without median longitudinal carinula; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes normal. Antenna slightly longer than combined length of head and pronotum. Prosternal tubercle simple, conical with rounded apex. Dorsum of pronotum flattened, shallowly crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace about as wide as long. Tegmina and hind wings fully developed, former with some transverse parallel stridulatory veinlets in 1st radial area. Lower genicular lobe of hind femur spined; hind tibia expanded in apical half, with acute upper margins; external apical spine of hind tibia present. In male postero-dorsal margin of 1oth abdominal tergite with furcula fully developed, weak or absent; epiphallus with narrow divided bridge, well developed ancorae and complex lophi. Female ovipositor valves very short with short lateral and longer, curved apical spines. DISTRIBUTION. Pakistan, eastwards to N. Australia, Solomon Is. SPECIES INCLUDED Gesonula punctifrons (Stal) Acridium (Oxya) punctifrons Stal, 1861 : 336. Holotype g, Cuina (NR, Stockholm). Heteracris tenuis Walker, 1870 :668. Syntypes g¢ and 9, Cuina (BMNH) [examined]. [Synonymised by Kirby, 1910 : 393.] Racilia okinawensis Matsumura, 1910 : 3, pl. 4, fig 7. Syntypes J and 9, Taiwan and JAPAN (Okinawa) (coll. Matsumura). [Synonymised by Tinkham, 1935 : 491.] G. mundata (Walker) Heteracris mundata Walker, 1870 : 672. Syntypes gf and 9, AustraLia (BMNH) [examined]. Gesonia recticercus Sjéstedt, 1920: 21. Syntypes ¢ and 9, Austraria (NR, Stockholm; 210 DD: HOLLES MNHU, Berlin) [examined]. [Synonymized by Sjéstedt, 1935 : 71.] For subspecies of this species see Rehn, 1952. G. rehni Akbar & Balock Gesonula vehni Akbar & Balock, 1970 : 12, pl. 1, figs 1 A-D, pl. 2, figs 1 A-I. Holotype J, PAKISTAN (University of Sind, Janshoro). Hygracris Uvarov, 1921 (Text-figs 12, 43) Hygvacris Uvarov, 1921 : 505. Type-species: Hygvacris palustris Uvarov, by monotypy. Diacnosis. Head conical; fastigium of vertex, from above, parabolic, wider than long, median longitudinal carinula weakly present; frontal ridge sulcate, almost complete to clypeus; lateral facial keels complete. Eyes normal. Antenna slightly longer than combined length of head and pronotum. Prosternal process moderately simple, narrow at base, expanded in apical third and bent backwards, with rounded or subacute apex. Dorsum of pronotum flattened, shallowly crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace longer than wide. Both upper and lower genicular lobes of hind femur spined; hind tibia strongly expanded in apical half, with acute upper margins; external apical spine of hind tibia absent. In male roth abdominal tergite without a furcula; epiphallus with divided bridge, well developed ancorae and two pairs of lophi. Female ovipositor valves long, slender, evenly toothed. DISTRIBUTION. India. SPECIES INCLUDED Hygracris palustris Uvarov Hygvacris palustris Uvarov, 1921 : 506. Holotype g, INp1a (Bengal) (BMNH) [examined]. H. malabaricus Willemse Hygracris malabaricus Willemse, 1962 : 52, fig. 8. Holotype 9, Inp1a (South-west) (MNHN, Paris). Discussion. Superficially species of this genus most closely resemble those of Oxya but are easily distinguished by the acutely pointed upper genicular lobes of the hind femur. The presence of ancorae and bilobed lophi in the male epiphallus suggest a close relationship with the genera Thanmoia, Fer and perhaps Gesonula. Quilta Stal, 1860 (Text-figs 3, 4, II, 59) Acridium (Quilta) Stal, 1861 : 337. Type-species: Acridium (Quilta) mitratum Stal, by monotypy. Quilta Stal; Stal, 1873 : 8o. Diacnosis. Head elongate conical; fastigium of vertex, from above, steeply parabolic, longer than wide, without median longitudinal carinula; frontal ridge sulcate, complete almost to clypeus; lateral facial keels complete. Eyes somewhat flattened. Antenna short, not REVIEW OF THE OXYIN# 211 longer than combined lengths of head and pronotum. Prosternal process simple, subconical with slightly bulbous and backward pointing apex. Dorsum of pronotum flattened; median longitudinal carina very weak, barest traces of lateral carinae present, shallowly crossed by three transverse sulci; mesosternal interspace narrow, longer than wide; metasternal lobes con- tiguous. Tegmina and hind wings fully developed, former without trace of parallel transverse stridulatory veinlets. Both lower and upper genicular lobes of hind femur spined; hind tibia expanded strongly in apical half, with acute upper margins; external apical spine of hind tibia present. In male roth abdominal tergite with a weak furcula on postero-dorsal margin; epiphallus with narrow divided bridge, without ancorae, and with only outer pair of lophi present. Female ovipositor valves long, upper valves smooth, lower valves weakly spined. DISTRIBUTION. Thailand, W. Indonesia. SPECIES INCLUDED Quilta mitrata (Stal) Acridium (Quilta) mitratum Stal, 1861 : 337. Syntypes gj and 9, KEELING Is. (MNHU, Berlin; NR, Stockholm) [examined]. Q. oryzae Uvarov Quilta oryzae Uvarov, 1925 : 159, figs 3-5. Holotype 9, THAILAND (BMNH) [examined]. Q. deschauenseei Rehn Quilta deschauenseei Rehn, 1957 : 6. Holotype 9, THAILAND (ANS, Philadelphia). Stolzia Willemse, 1930 (Text-figs 31-37) Stolzia Willemse, 1930: 117. Type-species: Stolzia rubromaculata Willemse, by monotypy. Incolacris Willemse, 1932 :150. Type-species: Incolacris rubritarsi Willemse, by original designation (Willemse, 1932 : 151). Syn. n. Butonacris Willemse, 1933 :6. Type-species: Butonacris fasciata Willemse, by monotypy. Syn. n. Diacnosis. Head conical; fastigium of vertex, from above, rounded pentagonal, about as wide as long, median longitudinal carinula absent; frontal ridge sulcate, tending to evanesce below median ocellus; lateral facial keels complete. Eyes bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process short, pyramidal, apex subacute. Dorsum of pronotum rugulose, slightly flattened at least in metazona, median carina week, lateral carinae absent, moderately deeply crossed by three transverse sulci; mesosternal interspace wider than long. Lower genicular lobe of hind femur spined; hind tibia hardly expanded apically; external apical spine of hind tibia present. In male roth abdominal tergite with furcula on postero-dorsal margin; supra-anal plate tuberculate; epiphallus asymmetrical, left side larger and with more strongly developed lophi, with divided bridge and developed ancorae. Female ovipositor valves long and slender with apices somewhat spatulate and evenly spined. DISTRIBUTION. China, W. Indonesia, Philippine Is. 212 Dp. TLOLELES Fics 31-37. Stolzia species. Male and female genitalia. S. borneensis (Willemse), 31, male epiphallus; S. aberrans (Willemse), 32, male epiphallus; S. javana Ramme, 33, male epiphallus; S. trifasciata (Willemse) 34, male epiphallus; S. fasciata (Willemse), 35, male epiphallus; S. rubritarsi (Willemse), 36, female ovipositor, ventral view; 37, same, lateral view. REVIEW OF THE OXYINZ 213 Oxyinae genera, male epiphalli. 38, Caryanda diminuta (Walker). 39, C. paravicina (Willemse); 40, C. modesta (Giglio-Tos); 41, C. cylindrica (Ramme); 42, C. palawana (Ramme); 43, Hygracris palustris Uvarov; 44, Gesonula punctifrons (Stal); 45, Oxyina bidentata (Willemse); 46, O. javana (Willemse); 47, O. sinobidentata (Hollis). Fics 38-47. 214 DD; HOLLIS Fics 48-55. Oxyinae genera, male epiphalli. 48, Badistica fascipes Chopard; 49, B. bellula Karsch; 50, B. margarita Jago; 51, B. simpsoni Ramme; 52, B. ornata I. Bolivar; 53, B. lauta Karsch; 54, Gerista dimidiata I. Bolivar; 55, Lucretilis jucunda Miller. REVIEW OF THE OXYINZ 215 Fics 56-62. Oxyinae genera, male epiphalli. 56, Cercina obtusa Stal; 57, Bumacris flavomaculata Willemse; 58, Ochlandriphaga xanthelytrana Miller; 59, Quilta oryzae Uvarov; 60, Cylindrotiltus versicolor Ramme; 61, Digentia rufogeniculata (I. Bolivar) ; 62, D. punctatissima (Stal). 216 D. HOLLIS SPECIES INCLUDED Stolzia rubromaculata Willemse Stolzia rubromaculata Willemse, 1930 : 118, fig. 58. Holotype 3, SumMatTRa (coll. Willemse) [examined]. S. rubritarsi (Willemse) comb. n. Incolacris rubritarsi Willemse, 1932 :151. Holotype 9, PHILIPPINE Is. (coll. Willemse) [examined]. S. trifasciata (Willemse) comb. n. — Incolacris trifasciata Willemse, 1932 :152. Holotype ¢, PHILipPinEe Is. (coll. Willemse) [examined]. S. fasciata (Willemse) comb. n. Butonacris fasciata Willemse, 1933 : 118, fig. 4. Holotype g, SuLAweEs1 (Buton I.) (NM, Vienna) [examined]. Incolacris atrifrons Willemse, 1956 : 186. Holotype 9, SuLawEs1 (Buton I.) (coll. Willemse) [examined]. Syn. n. S. borneensis (Willemse) comb. n. Butonacris borneensis Willemse, 1938 : 83, fig. 1. Holotype g, Borneo (Central) (RNH, Leiden) [1 g paratype examined]. S. aberrans (Willemse) comb. n. Butonacris aberrans Willemse, 1938 : 84, fig. 2. Holotype 4, Bornro (Central) (coll. Willemse) [examined]. S. nigromaculata (Willemse) comb. n. Butonacris nigromaculata Willemse, 1938 : 120, fig. 5. Holotype g, Borneo (North) (UM, Oxford) [could not trace]. S. flavomaculata (Willemse) comb, n. Incolacris flavomaculata Willemse, 1939: 75, 89, fig. 3. Holotype 9, Martaya (Malacca) (coll. Willemse) [examined]. S. hainanensis (Tinkham) comb. n. Incolacris hainanensis Tinkham, 1940 : 302, pl. 11, fig. 2. Holotype g, Cuina (Hainan) (? depository). S.javana Ramme Stolzia javana Ramme, 1941 : 101, pl. 13, fig. 5. Holotype 9, Java (MNHU, Berlin) [examined]. Discussion. Willemse separated Stolzia from Butonacris and Incolacris because of the incomplete frontal ridge of S. rubromaculata, and Incolacris species were separated on the basis of the sculpturing of the metazona. Neither of these characters is considered here to be of any significance at the generic level and the three genera are synomymized. By far the most remarkable feature of this genus is the asymmetrical development of the male epiphallus. As far as I am aware this has not occurred in any other group of Acridoidea although it was noted, to a relatively minor extent, in Oxya fuscovittata (Marschall) (Hollis, 1971). Also the latter species has weakly developed tubercles on the male supra-anal plate, another feature more strongly developed in Stolzia species. REVIEW OF THE OXYINZ 217 Caryanda Stal, 1878 (Text-figs 38-42) Caryanda Stal, 1878 : 47. Type-species: Acridium (Oxya) spurium Stal, by monotypy. Dibastica Giglio-Tos, 1907:9. Type-species: Dibastica modesta Giglio-Tos, by monotypy. Syn. n. Austenia Ramme, 1929 : 331. Type-species: Austenia cylindrica Ramme, by monotypy. [Homonym of Austenia Nevill, 1878 :16.} Syn. n. Austeniella Ramme, 1931 : 934. [Replacement name for Austenia Ramme]. Syn. n. DiaGcnosis. Head conical; fastigium of vertex, from above, pentagonal, wider than long, without median longitudinal carinula; frontal ridge sulcate. Eyes normal. Antenna as long as combined lengths of head and pronotum. Prosternal process conical with subacute apex. Dorsum of pronotum weakly flattened, median carina weak, lateral carinae absent, weakly crossed by three transverse sulci; mesosternal interspace slightly longer than wide, Tegmina and hind wings normally reduced to micropterous condition, some species brachypterous and one species is rarely macropterous. Lower genicular lobe of hind femur spined; hind tibia hardly expanded apically, upper margins acute, external apical spine present. In male 1oth abdominal tergite with or rarely without furcula on hind margin; epiphallus with divided bridge and clearly developed ancorae. Female ovipositor valves long, slender, evenly toothed. DISTRIBUTION. C. Africa, India, Burma, China, Indo-China, W. Indonesia, Philippine Is. SPECIES INCLUDED Caryanda spuria (Stal) Acridium (Oxya) spurium Stal, 1861 : 336. Holotype g, Java (NR, Stockholm) [examined]. C. diminuta (Walker) comb. n. Oxya diminuta Walker, 1871 : 64. Holotype 9, Cu1na (Yunan) (BMNH) [examined]. Oxya rufipes Brunner von Wattenwyl, 1893 : 153. Lectotype g, Campopia (NM, Vienna), designated by Hollis (1971 : 336) [examined]. [Synonymized by Willemse, 1921 : 42]. C. sanguineoannulata Brunner von Wattenwyl Caryanda sanguineoannulata Brunner von Wattenwyl, 1893 : 154, pl. 5, fig. 52. Holotype 36, Burma (MCSN, Genoa). C. modesta (Giglio-Tos) comb. n. Dibastica modesta Giglio-Tos, 1907 : 10. Holotype 9, ZAIRE (IRSNB, Brussels). Dibastica major I. Bolivar, 1908 : 106. Holotype 9, ZAIRE (IRSNB, Brussels). [Synonymised by Descamps & Donskoff, 1964 : 1204]. Dibastica viridis I. Bolivar, 1908 : 106. Holotype 9, ZAIRE (IRSNB, Brussels). [Synonymised by Ramme, 1929 : 328}. Dibastica elegans 1. Bolivar, 1911 : 302. Holotype g, ZAIRE (MRAC, Tervuren). [Synonymised by Descamps & Donskoff, 1968 : 1204]. C. cachara (Kirby) Traulia cachava Kirby, 1914 : 245, fig. 132. Holotype 9, Inp1a (Assam) (BMNH) [examined]. C. elegans I. Bolivar* Caryanda elegans I. Bolivar, 1918 : 20. Syntypes g¢ Burma, 2 2 NortH VIETNAM (IEE, Madrid) [examined, almost certainly a mixed series]. * The homonomy created_here can be resolved when Dr. Fer Willemse publishes his revision of the genus. 218 D. HOLLIS C. pumila Willemse Caryanda pumila Willemse, 1924 : 158. Holotype g, SuMATRA (RNH, Leiden). C. paravicina (Willemse) comb. n. Oxya paravicina Willemse, 1925) : 55, figs 62, 63. Holotype g, INnp1A (North-east) (NM, Vienna) [examined]. C. cylindrica (Ramme) comb. n. Austenia cylindrica Ramme, 1929 : 332, pl. 7, figs 10, 11; figs 46a, 47a, 48a. Holotype g, SIERRA LEONE (BMNH) [examined]. C. methiola Chang Caryanda methiola Chang, 1939 : 43, pl. I, figs 5, 6, 9, pl. 3, fig. 4. Holotype 9, Cuina (Szechwan) (USNM Washington). C. sinensis Chang Caryanda sinensis Chang, 1939: 45, pl. 1, figs 3, 4, 7, pl. 3, fig. 2. Holotype 9, CHina (Szechwan) (USNM, Washington). C. omeiensis Chang Caryanda omeiensis Chang, 1939 : 47, pl. 1, fig. 8. Holotype 9, Cuina (Szechwan) (USNM, Washington). C. pieli Chang Caryanda pieli Chang, 1939 : 48, pl. 2, figs 1-4, pl. 3, figs 1, 7. Holotype 9, Curna (Chekiang) (? depository). C. palawana (Ramme) comb. n. Chitaurella palawana Ramme, 1941: 137. Holotype 4, PHILIPPINE Is. (Palawan) (NM, Vienna) [examined]. C. olivacea Willemse Caryanda olivacea Willemse, 1956:168. Holotype g, NortH ViEtTNAM (NM, Vienna) [examined]. SPECIES DOUBTFULLY INCLUDED C. pulchra Brancsik Caryanda pulchva Brancsik, 1897 : 74, pl. 3, figs 17a, b. Holotype 3, NEw GuINgEa (probably lost). C. flavomaculata I. Bolivar Caryanda flavomaculata I. Bolivar, 1918: 21. Holotype g, NEw Guinea (IEE, Madrid) [examined]. Discussion. There seems to be no valid reason on the basis of external morphology or in the structure of the male phallic complex why Austeniella and Dibastica should remain separate from Caryanda and they are therefore synonymised. As mentioned in my earlier paper on Oxya (Hollis, 1971) that genus was considered to be a somewhat heterogeneous assemblage of species and some rearrangement would be necessary. O. paravicina and O. diminuta cleary differ from other species of Oxya (sensu stricto) in that a male furcula is present and the epiphallus bears ancorae. These two species are considered here more naturally grouped in Caryanda. Chitaurella palawana is clearly not congeneric with C. lucida Krauss, having a simple, conical prosternal process. It seems to be more closely allied to the Caryanda REVIEW OF THE OXYINZE 219 species and is placed in that genus. The holotype male has the external apical spine of the hind tibia present only on the right side, but another male from Palawan, deposited in MNHN, Paris, has the spine on both tibae. The identity of Caryanda pulchra will remain obscure until topotypic material becomes available for study. Caryanda flavomaculata differs from the rest of the genus in having deep transverse sulci on the pronotum, the male furcula is absent, the epiphallus lacks ancorae and is very similar to the type found in Oxya. In view of these differences it is likely that C. flavomaculata should be placed in another genus but, as it is only known from a discoloured male dried from alcohol, there is insufficient information available and the current combination is maintained. Gerista I. Bolivar, 1905 (Text-fig. 54) Gerista I. Bolivar, 1905 : 233. Type-species: Gerista dimidiata I. Bolivar, by monotypy. Diacnosis. Head conical; fastigium, from above, rounded pentagonal, wider than long, median longitudinal carinula absent; frontal ridge sulcate, evanescing below median ocellus; lateral facial keels complete. Eyes bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process conical with subacute apex. Dorsum of pronotum slightly flattened, shallowly crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace about as long as wide. Tegmina and hind wings reduced, former extending only half-way along abdomen and without trace of transverse parallel stridulatory veinlets. Lower genicular lobe of hind femur spined; hind tibia expanded in apical part, with acute upper margins apically; external apical spine of hind tibia absent. In male toth abdominal tergite without a furcula; epiphallus with divided bridge, ancorae and only an outer pair of lophi. Female ovipositor valves long, slender, evenly toothed. DISTRIBUTION. Nigeria, Fernando Po. SPECIES INCLUDED Gerista dimidiata |. Bolivar Gerista dimidiata I. Bolivar, 1905 : 234. Holotype gj, FERNANDO Po (IEE, Madrid) [examined], Discussion. Gerista is the only African genus in the Oxyinae lacking the external apical spines of the hind tibia. However, this feature is not regarded as very significant here and, on the evidence from the form of the male epiphallus, Gerista appears most closely related to the /auta species-group of Badistica. The female of this species was described by Jago (1962) from the Benin forest region of Nigeria but it may be that Jago’s material represents a mainland species distinct from that found on Fernando Po. 220 D. HOLLIS Badistica Karsch, 1891 (Text-figs 48-53) Badistica Karsch, 1891 : 194. Type-species: Badistica bellula Karsch, by monotypy. Dracnosis. Head conical; fastigium, from above, pentagonal, wider than long, median longitudinal carinula absent; frontal ridge sulcate, complete to clypeus or evanescing below median ocellus; lateral facial keels complete. Eyes moderately bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process conical with subacute apex. Dorsum of pronotum rounded or slightly flattened, shallowly or more deeply crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings very reduced, scale-like. Lower genicular lobe of hind femur spined; hind tibia hardly expanded apically, upper margins hardly angular; external apical spine of hind tibia present. In male toth abdominal tergite with or without furcula on postero-dorsal margin; epiphallus with partially or completely divided bridge, ancorae more or less developed. Female ovipositor valves long, slender, smooth, finely serrate in apical quarter or evenly toothed in apical half. DISTRIBUTION. W. Africa. SPECIES INCLUDED Badistica bellula Karsch Badistica bellula Karsch, 1891 : 195, figs. Syntypes g and 9, CamMERouN (MNHU, Berlin) [examined]. B. lauta Karsh Badistica lauta Karsch, 1896 : 282, figs 16, 16a. Lectotype g, Toco (MNHU, Berlin), designated by Jago (1966 : 340) [examined]. B. ornata I. Bolivar Badistica ornata 1. Bolivar, 1905 : 229. Holotype 3, GHana (MNHU, Berlin) [examined]. B. simpsoni Ramme Badistica simpsoni Ramme, 1929 : 336, figs 52b, 53b. Holotype 3, SIERRA LEONE (BMNH) [examined]. : B. fascipes Chopard Badistica fascipes Chopard, 1958 : 128, fig. 1. Holotype 9, Guinea: Mt. Nimba (MNHN, Paris). B. margarita Jago Badistica margarita Jago, 1966 : 331, figs 1A, 1B,2A. Holotype g, Guana (BMNH) [examined]. Discussion. Jago (1966) suggests bellula and simpsoni form one species-group and lauta, fascipes and margarita another, with ornata rather intermediate. B. bellula and simpsoni are distinct from the other species of the genus in that the male epiphallus is incompletely divided (Text-figs 49, 51), a feature altogether rare in the whole subfamily. Furthermore, in these two species the pronotum is flatter and less ‘waisted’. Whether these differences are due to divergence within the genus or convergence of two lines of evolution remains uncertain. Oxya Audinet-Serville, 1831 (Text-figs 7, 10, 13-19) Oxya Audinet-Serville, 1831 : 286. Type-species: Orya hyla Audinet-Serville, by monotypy. Zulua Ramme, 1929 : 327. Type-species: Zulua glabra Ramme, by monotypy. Syn. n. REVIEW OF THE OXYINZ 22% Diacnosis. Head conical; fastigium, from above, rounded pentagonal, wider than long, with weak or without median longitudinal carinula; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes normal. Antenna longer or shorter than combined lengths of head and pronotum. Prosternal process conical with subacute or rounded apex. Dorsum of pronotum flattened or slightly rounded, weakly crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings fully developed or reduced to brachypterous condition, former always touching in mid dorsal line when folded and without trace of transverse parallel stridulatory veinlets. Lower genicular lobe of hind femur spined; hind tibia expanded in apical half and with acute upper margins. In male roth abdominal tergite without furcula; epiphallus with divided bridge, without ancorae and with two pairs of lophi. Female ovipositor valves long, slender, unevenly spined or evenly serrate. DISTRIBUTION. Ethiopian, S.E. Palaearctic, Oriental, Austro-Oriental, Australasian regions. SPECIES INCLUDED (For synonymy and subspecies of these species see Hollis, 1971) Oxya velox (Fabricius) Gryllus velox Fabricius, 1787 : 239. Lectotype 9, Cuina (UZM, Copenhagen), designated by Hollis (1971 : 297) [examined]. O. chinensis (Thunberg) Gryllus chinensis Thunberg, 1815 : 253. Holotype 9, Curna (ZIUU, Uppsala) [examined). O. japonica (Thunberg) Gryllus japonicus Thunberg, 1824 : 429. Holotype 2, JAPAN (ZIUU, Uppsala) [examined]. O. hyla Audinet-Serville Oxya hyla Audinet-Serville, 1831 : 287. Neotype g, SENEGAL (MNHN, Paris), designated by Hollis (1971 : 282) [examined]. O. fuscovittata (Marschall) Gryllus fuscovittatus Marschall, 1836 : 211, pl. 18, fig. 3. Holotype 9, no data [probably N.W. India] (NM, Vienna) [examined]. O. nitidula (Walker) Acridium nitidulum Walker, 1870 : 631. Holotype 9, Inp1A (South) (BMNH) [examined]. O. cyanoptera Stal comb. rev. Oxya cyanoptera Stal, 1873 : 83. Holotype 9, SIERRA LEONE (NR, Stockhom) [examined]. O. cyanipes (Karny) comb. n. Catantops cyanipes Karny, 1907 : 330, pl. 2, figs 11-15. Holotype g, UGanpa (NM, Vienna) [examined]. O. yezoensis Shiraki Oxya yezoensis Shiraki, 1910 : 43. Neotype g, JAPAN (ZIHU, Sapporo), designated by Hollis (1971 : 326) [examined]. O. minuta Car! Oxya minuta Carl, 1916: 472. Lectotype g, Java (MNH, Geneva), designated by Hollis (1971 : 293) [examined]. O. grandis Willemse Oxya grandis Willemse, 1925) : 36, figs 34, 35. Holotype g, INp1a (BMNH) [examined]. 222 DD» HOELIS O. glabra (Ramme) comb. n. Zulua glabra Ramme, 1929 : 327, pl. 3, figs 3, 4, fig. 44. Holotype g, SourH Arrica, (BMNH) [examined]. O. agavisa Tsai Oxya agavisa Tsai, 1931 : 437, fig. 1. Holotype 2, Cu1na (MNHU, Berlin) [examined]. O. ningpoensis Chang Oxya ningpoensis Chang, 1934 : 190, figs I-3. Syntypes ¢ and 9, CuiNna (partly in BMNH) [partly examined]. O. stresemanni Ramme Oxya stresemanni Ramme, 1941 : 213. Holotype g, SuLAwEs1 (MNHU, Berlin) [examined]. O. oxyura Uvarov comb. rev. Oxya oxyura Uvarov, 1953 : 68, fig. 71. Holotype J, ANGoLa, (BMNH) [examined]. O. bolaangensis Hollis Oxya bolaangensis Hollis, 1971 : 312, figs 25, 138, 150-159. Holotype g, SULAWEs! (coll. Willemse) [examined]. Discussion. When Oxya was revised by the:author (Hollis, 1971) it was realised that some of the species then included in the genus would have to be removed to other genera and this is done here with the removal of O. diminuta and O. paravicina to Caryanda (p. 217), and O. bidentata, sinobidentata and javana to the new genus Oxyina (p.228). It was also noted that the position of Zulua, a purely African genus, as a discrete taxon, was not satisfactory. Zulua was differentiated by Dirsh (1965 : 235) from Oxya by its more acute head profile, shorter antennae, the presence of a weak median longitudinal carinula on the fastigium, the subcylindrical pronotum and a tendency to shortened elytra. These features, although serving to separate Zulua species from Oxya hyla hyla, break down when Zulua is compared with the Oriental Oxya species and the two genera are synonymised. Bumacris Willemse, 1931 (Text-fig. 57) Bumacris Willemse, 1931b : 348. Type-species: Bumacris flavomaculata Willemse, by monotypy. Diacnosts. Head conical; fastigium of vertex, from above, pentagonal, wider than long, without median longitudinal carinula; frontal ridge sulcate, complete almost to clypeus; lateral facial keels complete. Eyes bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process conical with subacute apex. Dorsum of pronotum rounded or weakly flattened, shallowly or more deeply crossed by three transverse sulci, median carina weak, lateral carinae absent; mesosternal interspace as long as or slightly longer than wide. Tegmina and hind wings fully developed or slightly reduced and only extending two-thirds the length of abdomen; no trace of transverse parallel stridulatory veinlets. Lower genicular lobe of hind femur spined; hind tibia expanded in apical half, and with acute upper margins; external apical spine of hind tibia present. In male 1oth abdominal tergite without furcula; epiphallus with broad, divided bridge, without ancorae and with only the outer pair of lophi. Female ovipositor valves very long, slender, evenly toothed. DISTRIBUTION. Solomon Is. REVIEW OF THE OXYINZ 223 SPECIES INCLUDED Bumacris flavomaculata Willemse Bumacris flavomaculata Willemse, 1931b : 350, fig. 4. Holotype g, SoLtomon Is. (Malaita) (NM, Basle). B. monotona Willemse Bumacris monotona Willemse, 1935) : 166. Holotype 9, Sotomon Is. (Guadalcanal) (BMNH) [examined]. B. pagdeni Willemse Bumacris pagdeni Willemse, 1935b : 167. Holotype g, Soromon Is. (Vella Iavella) (BMNH) [examined]. B. leveri Uvarov Bumacris leveri Uvarov, 1937 :17. Holotype 9, Sotomon Is. (Isabel) (BMNH) [examined]. B. bougainvillea Ramme Bumacris bougainvillea Ramme, 1941 :118. Holotype g, Soromon Is. (Bougainville) (MNHU, Berlin). B. georgica Willemse Bumacris georgica Willemse, 1962 : 51, fig. 4. Holotype g, Sotomon Is. (St. George) (MNHN, Paris). Discussion. Dr Fer Willemse will shortly be publishing a thorough revision of this genus. Digentia Stal, 1878 (Text-figs 61, 62) Digentia Stal, 1878 : 47. Type-species: Oxya punctatissima Stal, by monotypy. Genditia I. Bolivar, 1911 : 301. Type-species: Genditia rufogeniculata I. Bolivar, by monotypy. Syn. n. Diacnosis. Head conical; fastigium, from above, rounded pentagonal, wider than long, with or without median longitudinal carinula; frontal ridge sulcate, complete to clypeus; lateral facial keels complete. Eyes moderately bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process simple, conical with subacute apex. Dorsum of pronotum rugulose and coarsely pitted, rounded or weakly flattened, moderately deeply crossed by three transverse sulci; median carina weak, lateral carinae absent; mesosternal interspace about as wide as long. Tegmina and hind wings very reduced, former minute and not extending beyond mesonotum or scale-like and extending to 1st abdominal tergite. Lower genicular lobe of hind femur spined; hind tibia weakly or moderately expanded in apical part and with acute upper margins apically; external apical spine of hind tibia present. In male posterodorsal margin of 1oth abdominal tergite with a weak furcula or if without then it is expanded dorso-laterally; epiphallus with incompletely divided bridge and without ancorae. Female ovipositor valves long, slender, smooth or evenly and weakly serrate in apical half. DISTRIBUTION. W. and C. Africa. SPECIES INCLUDED Digentia punctatissima (Stal) Oxya punctatissima Stal, 1875 : 33. Holotype ¢, W. Arrica (NR, Stockholm) [examined]. 224 D. HOLLIS D. rufogeniculata (1. Bolivar) comb. n. Genditia rufogeniculata I. Bolivar, 1911 : 301. Holotype 9, ZArRE (Kasai) (MRAC, Tervuren). D. fasciata Ramme Digentia fasciata Ramme, 1929 : 323, figs 40, 41b. Holotype g, CAmMERouN (MNHU, Berlin) [examined]. D. karnyi Ramme sp. rev. Digentia karnyi Ramme, 1929 : 331, pl. 7, fig. 9. Holotype 9, ZarrE (MRAC, Tervuren) [1 2 paratype examined]. Discussion. Prior to this review Digentia included two. recognised species, the type-species punctatissima and fasciata. Genditia was erected for rufogeniculata and Dirsh (1952) added versicolor (Ramme). Considering the. two type-species there seem to be no differences upon which to base a generic separation. D. punctatissima has more reduced organs of flight, less expanded hind tibiae and smooth ovipositor valves, features which can be clearly associated with a more or less complete adaptation to a forest habitat. Furthermore, as both punctatissima and rufogeniculata have the male epiphallus with an incompletely divided bridge, the two genera are synonymised. D. versicolor is significantly different from the two former species as the male epiphallus has a completely divided bridge, the integument is much less coarsely pitted, the antennae are very short and the overall appearance is very different to the other Digentia species. It is therefore removed from Digentia and replaced in its original combination as Cylindrotiltus versicolor Ramme, 1929. In the female paratype of Digentia karnyi Ramme the tegmina appear narrower than in rufogeniculata, also the posterior margin of the subgenital plate is sharply triangular and the ventral concavity more rounded. For these reasons Dirsh’s synonymy (1952 : 268) is not accepted. Cercina Stal, 1878 (Text-figs I, 56) Cercina Stal, 1878 : 49, 97. Type-species: Cercina obtusa Stal, by monotypy. Cledva I. Bolivar, 1918 : 25. Type-species: Cledva simoni I. Bolivar, by monotypy. [Synony- mised by Uvarov, 1927 : I05.] DiaGnosis. Head conical; fastigium, from above, short triangular or pentagonal, wider than long, median longitudinal carinula absent; frontal ridge weakly sulcate and not quite extending to clypeus; lateral facial keels complete. Eyes normal. Antenna shorter than combined lengths of head and pronotum. Prosternal process subconical, antero-posteriorly flattened with subacute apex. Dorsum of pronotum flattened, shallowly crossed by two or three transverse sulci, median carina very weak, lateral carinae absent; mesosternal interspace longer than wide. Tegmina and hind wings reduced, scale-like, former not extending beyond 3rd abdominal tergite. Lower genicular lobe of hind femur pointed or spined; hind tibia moderately expanded in apical half; with acute upper margins;-external apical spine of hind tibia present. In male roth abdominal tergite without furcula; epiphallus with narrow divided bridge, without ancorae and with only the outer pair of lophi. Female ovipositor valves long, slender, evenly toothed. DISTRIBUTION. Ceylon. REVIEW OF THE OXYINZ 225 SPECIES INCLUDED Cercina obtusa Stal Cercina obtusa Stal, 1878 :97. Holotype 9, CEYLon (NM, Vienna). Cledva simoni I. Bolivar, 1918 :26. Holotype g, CrEyLon (IEE, Madrid) [examined}. [Synonymised by Uvarov, 1927 : 105]. Caryanda platycera Willemse, 1924:157, fig. 1. Holotype g, Cryton (NM, Vienna) [examined]. Syn. n. C. phillipsi Henry Cercina phillipsi Henry, 1933 : 184, pl. 33, figs 1,2. Holotype g, CEyLon, (BMNH) [examined]. Discussion. An examination of the type of Caryanda platycera has shown that this species should be transferred to Cercina and that it is synonymous with C. obtusa Cylindrotiltus Ramme, 1929 gen. rev. (Text-figs 2, 60) Cylindrotiltus Ramme, 1929: 319. Type-species: Cylindrotiltus versicolor Ramme, by monotypy. Dracnosis. Head conical ; fastigium, from above, rounded pentagonal, wider than long, without median longitudinal carinula; frontal ridge sulcate, evanescing just above clypeus; lateral facial keels complete. Eyes moderately bulbous. Antenna not longer than combined lengths of head and pronotum. Prosternal process conical, posteriorly flattened, with subacute apex. Dorsum of pronotum with smooth shiny integument except in metazona where it is more coarsely pitted, rounded, deeply crossed by three transverse sulci; median carina almost completely absent, lateral carinae absent; mesosternal interspace as wide as or longer than wide. Tegmina and hind wing reduced, scale-like, former only just extending beyond hind margin of 1st abdominal tergite. Lower genicular lobe of hind femur spined; hind tibia slightly expanded in apical half and with acute upper margins; external apical spine of hind tibia present. In male 1oth abdominal tergite without furcula; epiphallus with divided bridge, without ancorae. Female ovipositor valves long, slender, evenly and finely serrate. DISTRIBUTION W. Africa. SPECIES INCLUDED Cylindrotiltus versicolor Ramme comb. rev. Cylindrotiltus versicoloy Ramme, 1929 : 320, figs 37, 38, pl. 6, figs 24, 25. Holotype gj, CAMEROUN (MNHU, Berlin) [examined}. Discussion. The reasons for resurrecting this monobasic genus from synonymy with Genditia are givenon p. 224. Very little is known of the biology of C. versicolor but it would appear to be a forest-living species. On the evidence from external morphology and the male phallic complex it seems to be most closely related to Pterotiltus, being a little less specialised to forest life than the latter. 226 D: HOLLIS Pterotiltus Karsch, 1893 (Text-figs 63-69) Pygostolus Karsch, 1891 : 192. Type-species: Pygostolus impennis Karsch, by monotypy. [Homonym of Pygostolus Haliday, 1833 : 263]. Pterotiltus Karsch, 1893 : 108. [Replacement name for Pygostolus Karsch.] Diacnosis. Head conical; fastigium, from above, pentagonal, about as wide as long, median longitudinal carinula weakly present or absent; frontal ridge sulcate, evanescing below median ocellus; lateral facial keels complete. Eyes bulbous. Antenna longer than combined lengths of head and pronotum. Prosternal process simple, conical with acute apex. Dorsum of pronotum cylindrical, deeply crossed by three transverse sulci, median carina very weak, lateral carinae absent; mesosternal interspace as wide as or longer than wide. Tegmina and hind wings extremely reduced, represented by small folds of integument on meso- and metathorax. Lower genicular lobe of hind femur spined; hind tibia not expanded apically and with rounded margins; external apical spine of hind tibia present. In male postero- dorsal margin of 1oth abdominal tergite with a furcula or an unpaired structure resulting from fusion of the two lobes of furcula; epiphallus with divided bridge, without ancorae and with one or two pairs of lophi. Female ovipositor valves long, slender, smooth. DISTRIBUTION. W. and C. Africa. SPECIES INCLUDED Pterotiltus impennis (Karsch) Pygostolus impennis Karsch, 1891 : 193, figs. Syntypes J and 9, Cameroun (MNHU, Berlin) [examined]. P. inuncatus (Karsch) Pygostolus inuncatus Karsch, 1892:72. Syntypes g and 9, Cameroun (MNHU, Berlin) [examined]. P. miniatulus Karsch Pterotiltus miniatulus Karsch, 1893: 108. Syntypes g¢ and 9, Toco (MNHU, Berlin) [examined]. P. finoti Dominique Pterotiltus finoti Dominique, 1900 : 207, pl. 3, figs 11-13. Holotype J, Conco (BRAZZAVILLE) (? Nantes Museum). P. apicalis 1. Bolivar Pterotiltus apicalis I. Bolivar, 1905 : 226. Holotype Q, Frernanpo Po (IEE, Madrid) [examined]. P. coeruleocephalus I. Bolivar Pterotiltus coeruleocephalus 1. Bolivar, 1905 : 227. Holotype g, CAMEROUN (IEE, Madrid) [examined]. P. femoratus Ramme Pterotiltus femovatus Ramme, 1929 : 314, figs 34d, 35c. Holotype g, CAMERouN (BMNH) [examined]. P. berlandti Ramme Pterotiltus berlandti Ramme, 1929 : 316. Holotype ¢, ZatRE (? MNHN, Paris or MRAC, Tervuren). P. giorgii Ramme Pterotiltus giorgii Ramme, 1929 : 317. Holotype 9, ZarrE (MRAC, Tervuren). REVIEW OF THE OXYINZ Fics 63-69. Pterotiltus species, male epiphalli. 63, P. apicalis I. Bolivar; 64, P. impennis (Karsch); 65, P. miniatulus Karsch; 66 P. occipitalis Ramme; 67, P. inuncatus (Karsch) ; 68, P. minimus Ramme; 60, Pterotiltus sp., Uganda. 228 DP. HOLDS P. occipitalis Ramme Pterotiltus occipitalis Ramme, 1929 : 317. Holotype 2, CAMEROUN (MNHU, Berlin) [examined]. P. minimus Ramme Pterotiltus minimus Ramme, 1929 : 318, fig. 36. Holotype g, CAMERouN (MNHU, Berlin) [examined]. Discussion. Species of this genus are forest-living and seem to represent the culmination of adaptation to forest life in the Oxyinae. The wings are completely reduced, many of the species have adopted aposematic coloration, the hind tibiae are cylindrical and have lost all trace of flattening or expansion, and the ovipositor valves have become smooth and obviously adapted to specialised oviposition sites. Because of this high degree of specialisation the affinities of Pterotiltus are obscure, but the epiphallus resembles that of Cylindrotiltus and Cercina most closely. Oxyina gen. n. (Text-figs 45-47) Type-species: Oxya bidentata Willemse, 1925. Diacnosis. In external appearance very similar to Oxya. Differs in that male epiphallus is broad, plate-like, bears strongly developed ancorae and the ectophallic membrane is thickened anteriorly between the halves of the bridge. DisTRIBUTION. Afghanistan, Iran, Pakistan, E. China, Java. SPECIES INCLUDED Oxyina bidentata (Willemse) comb. n. Oxya bidentata Willemse, 19250 : 24, figs 16, 17. Holotype g, Pakistan (BMNH) [examined]. O. javana (Willemse) comb. n. Oxya javana Willemse, 1956 : 148, fig. 87, pl. 1. Holotype 9, Java (RNH, Leiden) [examined]. O. sinobidentata (Hollis) comb. n. Oxya sinobidentata Hollis, 1971 : 330, figs 228-233, 245. Holotype g, CHina (BMNH) [examined]. Discussion. Superficially this genus closely resembles Oxya but the male epiphallus is so distinct it is doubtful that the two genera have a common origin and their similarity is probably a result of convergence. In fact the epiphallus in Oxyina is distinct from all other Oxyinae and is similar to that of Hieroglyphodes (see Mason, 1973), a genus currently placed in the Hemiacridinae. However, Oxyina has more features common to the Oxyinae than to the Hemiacridinae and is placed in the former subfamily. REVIEW OF THE OXYINE 229 REFERENCES AKBAR, S. S. & Batockx, H. A. 1970. A new species of Gesonula Uvarov, 1940 (Orthoptera: Acridoidea: Acrididae) from West Pakistan. Pakistan J. Zool. 2 : 11-15, 2 pls. AUDINET-SERVILLE, J. G. 1831. Revue méthodique des insectes de l’ordre des Orthoptéres. Annls Sci. nat. 22 : 262-292. 1839. Histotve naturelle des insectes. Orvthoptéres. 776 pp., 14 pls. Paris. Borivar, I. 1898. Contributions a l'étude des Acridiens. Espéces de la faune Indo et Austro-Malaisienne du Museo Civico di Storia Naturale di Genova. Annali Mus. civ. Stor. nat. Giacomo Doria 19(39) : 1-36. 1905. Ortépteros Acridioideos de la Guinea Espafiola. Mems R. Soc. esp. Hist. nat.. 1 : 209-240. 1908. Acridiens d’Afrique du Musée royal d’Histoire naturelle de Belgique. Mém. Soc. ent. Belg. 16 : 83-126. 1911. Orthoptéres nouveaux du Congo Belge des collections du Musée de Tervuren. Annls Soc. ent. Belg. 55 : 298-306. 1918. Estudios entomoldégicos. Tercera parte. Seccién Oxyae. (Orth. Acrididae o Locustidae). Tvab. Mus. nac. Cienc. nat., Madr. Zool. 34 : 1-43. BRANCSIK, C. 1897. Series Orthopterorum novorum. Jh. naturw. Ver. Trencsiner Kom. 19-20 : 52-85. BRUNNER VON WATTENWYL, G. 1893. Reévision du Systéme des Orthoptéres et description des espéces rapportées par M. Leonardo Fea de Birmanie. Annali Mus. civ. Stor. nat. Giacomo Doria (2) 13 : 1-230, 6 pls. 1898. Orthopteren des Malayischen Archipels, gesammelt von Prof. Dr. W. Kiikenthal in dem Jahren 1893 und 1894. ‘ , ; < : A . : 237 INTRODUCTION ; : ‘ : ‘ : ‘ s ; 238 ABBREVIATIONS OF MUSEUMS. : ; : ; ‘ : ; 238 MEASUREMENTS AND INDICES. , : , : é ; ; 239 DEFINITION OF THE GENUS ; : : : : : . ‘ 239 NOTES ON THE GENUS ‘ : ‘ : : ; ‘ : ‘ 240 The synonymy of Leptogenys ; ‘ ; ‘ ‘ F : 240 Females of the species . : : ; é P ‘ 3 : 242 Mandibular structures . : : ; : i ; ; 242 THE SPECIES OF THE ETHIOPIAN REGION : 5 F ‘ ; ° 242 LisT OF SPECIES . : ; ‘ : F ; i s ; 242 KEy TO SPECIES (Workers) . F ; , : , ; ‘ 244 The conradti-group. ; : : , F ‘ : , 250 The mavzillosa-group - : : : : : : ‘ 252 The stuhlmanni-group . . : ; ; : , ? 256 The sulcinoda-group ok : ‘ : ; : 7 : 264 The attenuata-group . : : , : ‘ : : : 274 The guineensis-group : : : : : : : ‘ 277. The havilandi-group ; : ‘ : : : ‘ : 280 The nitida-group : 4 : . : 283 REVIEW OF THE SPECIES OF THE ‘Maracasy Rucion , , 2 i 293 LIST OF SPECIES . é P : ; : ‘ : 293 PROVISIONAL KEY TO SPECIES (Workers) . : , 5 ‘ : 293 The truncatirostris- -group ‘é . ‘ a 7 s A 295 The attenuata-group . : 2 é Z A : 2 P 296 The saussuret-group 5 A . A : ; : : 296 The incisa-group ; . . . ° : : 297 A SPECIES EXCLUDED FROM Leptogenys , ; : : ‘ : 298 ACKNOWLEDGEMENTS : A 3 ‘ = : : : 3 298 REFERENCES . : , ; : : fs é . P ; 299 INDEX - : : : 5 ‘ : : - P . : 304 SYNOPSIS The ant genus Leptogenys is revised for the Ethiopian Region and the species of the Malagasy Region are reviewed. Separate species-lists and keys for the two regions are given. Seventy species (56 Ethiopian, 14 Malagasy) are recognized, of which 26 are new. Twenty-five new 238 B. BOLTON synonyms, mostly infraspecific, are established. No subgenera are recognized. The Mada- gascan species jonesti (Forel), known only from the male, is transferred to the genus Pachycondyla F. Smith. INTRODUCTION THE genus Leptogenys comprises a reasonably compact and large genus within the tribe Ponerini. It was originally placed in a tribe of its own, along with Prionogenys Emery but as Brown (1963) pointed out, the traditional characters for separating a tribe Leptogenyini as independent are insufficient. At present there are some 280 named forms within the genus (including synonyms, etc.), which may represent about 175 valid species. Of these the Ethiopian and Malagasy regions have 56 and 14 species respectively, a total of 70 species or about 40 per cent of the world total. The genus is distributed throughout the world’s tropics and a few species occur in the subtropical zones. Wheeler (1922) gave a map of the distribution of the genus which has not radically altered since that time. Previous revisionary studies of the genus do not exist for the species of the Ethiopian and Malagasy regions although Arnold (1915) presented a key to the South African species and later gave additional notes and comments on a number of forms (Arnold, 1926). Very little is known of the biology of the species. Nest sites are usually in the ground, in rotton wood or in compressed leaf litter (Bolton, 1973) although a few subarboreal species are now known which nest in dead wood at some distance above the ground and forage upon the tree-trunks. In other zoogeographical regions some species are semi-nomadic, without a fixed nest-site, but it is not known if this applies to any species from the regions under consideration at present. Food records are few, the species for which prey has been recorded show that the main diet is either of termites or terrestrial Isopoda. Many species are nocturnal and are only rarely to be found during daylight by normal collecting methods. In the vast majority of species only the worker caste has been described. Females (queens) of only a few species are known and the males are virtually all unknown as collections of this caste in association with workers are extremely rare. The known males are all normal, winged forms but the females are apterous and in the main very ergatoid. Most species are more or less restricted in distribution but a number of them, particularly some members of the maxillosa-group, are well-known tramp species, having been spread throughout the tropics by human commerce (Wilson & Taylor, 1967). ABBREVIATIONS OF MUSEUMS AMNH, New York The American Museum of Natural History, New York, U.S.A. BMNH British Museum (Natural History), London, U.K. IE, Naples ~ Istituto di Entomologia, Naples, Italy. MCSN, Genoa Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy. MCZ, Cambridge Museum of Comparative Zoology, Cambridge, Mass., U.S.A. MHN, Geneva Muséum d’Historie Naturelle, Geneva, Switzerland. MNHN, Paris Muséum National d’Histoire Naturelle, Paris, France. REVISION OF GENUS LEPTOGENYS 239 MNHU, Berlin Museum fiir Naturkunde der Humboldt-Universitat, Berlin, Germany (D.D.R.). MRAC, Tervuren Musée Royal de 1’Afrique Centrale, Tervuren, Belgium. NM, Basle Naturhistorisches Museum, Basle, Switzerland. NM, Bulawayo National Museum, Bulawayo, Rhodesia. NM, Vienna Naturhistorisches Museum, Vienna, Austria. SAM, Cape Town South African Museum, Cape Town, South Africa. UFC, Yaounde Université Fédéral du Cameroun, Yaounde, Cameroun. UM, Oxford University Museum, Oxford, U.K. MEASUREMENTS AND INDICES Total Length (TL). The total outstretched length of the individual, from the mandibular apex to the gastral apex. Head Length (HL). The straight-line length of the head in perfect full-face view, measured from the mid-point of the anterior clypeal margin to the mid-point of the occipital margin. Head Width (HW). The maximum width of the head measured behind the eyes in full-face view. HW x 100 HL Scape Length (SL). The straight-line length of the antennal scape, excluding the basal constriction or neck. SL x 100 HW Pronotal Width (PW). The maximum width of the pronotum measured in dorsal view. Petiole Height (PH). The height of the petiole measured in profile from the apex of the ventral (subpetiolar) process vertically to a line intersecting the dorsalmost point of the node. Petiole Length (PL). The length of the petiole from the anterior process to the posteriormost point of the tergite, where it surrounds the gastral articulation. Cephalic Index (CI). Scape Index (SI). Lateral Petiole Index (LPI). sees Dorsal Petiole Width (DPW). The maximum width of the petiole in dorsal view. Dorsal Petiole Index (DPI), =~ *°° All measurements are expressed in millimetres. DEFINITION OF THE GENUS LEPTOGENYS Roger Leptogenys Roger, 1861 : 41. Type-species: Leptogenys falcigera Roger, 1861 : 42; by subse- quent designation of Bingham, 1903 : 52. 240 B. BOLTON Lobopelta Mayr, 1862 : 733. Type-species: Ponera diminuta F. Smith, 1857 : 69; by oe designation of Bingham, 1903 : 54. Syn.n. Lobopelta Mayr [as subgenus of Leptogenys]; Emery, 1911 : 101, et auctt. Machaerogenys Emery, 1911 : 100 [subgenus of Lepiogenys]. Type-species: Leptogenys trunca- tivostris Forel, 1897 : 195; by original designation. Syn. n. Dorylozelus Forel, 1915b : 24. Type-species: Dorylozelus mjobergi Forel, 1915b : 25; by monotypy (=Leptogenys tricosa Taylor, 1969 : 132, substitute name for D. mjobergi due to secondary homonymy in Leptogenys). [Synonymy by Taylor, 1969 : 132.] Microbolbos Donisthorpe, 1948 : 170. Type-species: Microbolbos testaceus Donisthorpe, 1948 : 170; by original designation. [Synonymy by Wilson, 1955: 136.] The synonymy of Lobopelta and Machaerogenys has been derived independently by Prof. W. L. Brown and will be recorded by him in his forthcoming revision of genera of tribe Ponerini. Worker. Minute to large monomorphic ants belonging to the tribe Ponerini. Mandibles variously shaped, may be subtriangular, short linear, or strongly elongate so that they are incapable of closing against the clypeus, but in all cases they are edentate or nearly so. Maximum number of teeth in species of Ethiopian and Malagasy Regions is three, consisting of apical plus two preapical; more usually only with one preapical or entirely without preapical teeth. Median portion of clypeus usually projecting as a distinct lobe, with a longitudinal median carina (absent only in truncatirostris). Palp formula 4,4. Lobes of frontal carinae small, only partially covering the antennal insertions, the major portion of which is visible in dorsal view. Antennae 12-segmented, the antennal scapes usually long, SI > 100, very rarely shorter. Eyes present in all known species, varying from very large to minute. Dorsal alitrunk with promesonotal suture well developed; metanotal groove present, usually distinct, reduced or indistinct only in some minute species. Middle and hind tibiae each with a large pectinate spur and a smaller simple spur. Pretarsal claws distinctly pectinate in most species, the pectination reduced to two, three or four teeth in some, very rarely absent in some minute species. Petiole a node of varying configuration, very rarely a thick, tapering scale (spandax). Gaster usually constricted between first and second segments, the constriction very faint or absent in some cases (stuhlmanni-group, guineensis-group). Sting long and stout. Female. Usually ergatoid and with the same characters as the workers, but with the gaster larger. In one species (ergatogyna) the female is apterous but not ergatoid, with a complete set of thoracic sclerites and a full complement of sutures. Ocelli are present in this species but not in the other known females. Male. Known only for a few species. Mandibles reduced, partially or wholly concealed by the broad clypeus when closed. Palp formula 6,4 (apparently 5,4 in some). Antennae 13-segmented, the scape short, at most about as long as the second funicular segment, often shorter. Eyes large, ocelli well developed. Lobes of frontal carinae widely separated, small, not concealing the antennal insertions. Mesoscutum with notauli and parapsidal furrows present; in profile this sclerite usually not swollen, not overhanging the pronotum in front. Hind wings without an anal lobe. Middle and hind tibiae with one large pectinate and one small simple spur. Pretarsal claws pectinate in all known species. Pygidium not projecting into a curved spine posteriorly. NOTES ON THE GENUS THE SYNONYMY OF Leptogenys. Of the four names listed above as synonyms of Leptogenys two were previously recorded. The first of these, Dorylozelus, need not concern us here as it applies to an Australian species, and is fully documented by Taylor (1969). Muicrobolbos was a name erected by Donisthorpe (1948) to cover a single West African species REVISION OF GENUS LEPTOGENYS 241 now known to be closely related to guineensis. Leaning heavily on the traditional characters used to define Leptogenys, Donisthorpe noted that in his species the claws were not pectinate and the mandible had three teeth (apical plus two preapical). Other small species are now known in which pectination is lost and a more or less complete series from fully to not pectinate can be constructed. Within the guineensis-group itself both microps and testacea lack pectination whilst guineensis has usually two minute teeth at least on the hind claws, and spandax has complete pectination. A similar phenomenon may be observed among the smaller species of the nitida-group. As pointed out by Wilson (1955), who synonymized Microbolbos to Leptogenys, the presence of an ‘extra’ preapical tooth is scarcely a useful character as in a number of species of the Oriental and Indo-Australian regions numerous preapical teeth are developed (processionalis-group). The two names newly synonymized above were originally separated from Leptogenys on mandibular structure. Lobopelta was erected by Mayr (1862) at a time when only very few species were known of Leptogenys, and all placed therein had elongate, linear mandibles. Recognizing that the two were closely related, Mayr defined Lobopelta as the short mandibulate forms, Leptogenys as the long mandi- bulate species. The two were retained as separate genera by a number of authors (e.g. Bingham, 1903) but as more species were described it became apparent that the distinction was by no means as sharp as was at first thought, and eventually Emery (Ig11) reduced Lobopelta to subgenus of Leptogenys. Since that time other species have been described which cannot easily be placed in either subgenus as their mandibles are intermediate in form, and in the present paper the thirteen species of the sulcinoda-group show a series in which the mandibles become more elongate and the median lobe of the clypeus becomes shorter and narrower. The result is that at one end of the group are species in which the mandibles close tightly against the clypeus, and at the other end are forms which have a large gap between clypeus and mandibles when the latter are closed. Thus one has the strange situation of having both subgenera within the same species-group. That the process of mandibular variation is not necessarily unidirectional is possibly illustrated by zapyxis of the stuhlmanni-group. In all other species of this group the mandibles are long and quite narrow. In zapyxis, however, a convex lamelliform extension has developed which runs most of the length of the inner mandibular margin and is, in effect, on its way back to a secondary short mandi- bulate condition as, if this lamella becomes much larger, then the mandibles will meet along their length when closed and will close against the clypeus although still projecting far in front of it. In view of these facts it was decided that Lobopelta was not viable even as a subgenus and consequently it has been placed in the synonymy. In the case of the final name, subgenus Machaerogenys, I cannot comprehend why a separate subgenus was ever thought necessary for truncatirostris and its allies. Emery’s (1911) original description of the subgenus was based only on the mandibular structure of truncatirostris itself, which is admittedly very derivative. However, this diagnosis does not apply to the other two species usually placed here, arcirostris and ridens, which have more normal mandibles and hence must be removed from 242 B. BOLTON Machaerogenys although sharing many other characters with truncatirostris (and showing a number of resemblances to the maxillosa-group). It is my opinion that a separate subgenus is unnecessary to contain a single species merely because its mandibles are more specialized than those of its closest relatives, and Machaerogenys has been abandoned. FEMALES OF THE SPECIES. Of the 56 species presently known from the Ethiopian Region, females are known for only six or seven, and all these except one are extremely ergatoid with the alitrunk and head similar to that of the worker. Ocelli and the thoracic sutures usually associated with this caste are absent. However, the female may be determined as the gaster is considerably more bulky than in the worker and the node of the petiole is usually antero-posteriorly compressed and considerably broader in dorsal view than in workers of the same species. The queen of evgatogyna is the only female yet known from the genus in which flight sclerites and ocelli are distinct and well developed, although the queen of this species appears never to develop wings, and again bears a remarkable similarity to the worker caste. MANDIBULAR STRUCTURES. Regardless of the shape or length of mandible in individuals, many species of the Ethiopian Region have a mandibular groove which has its origin dorsolaterally at the extreme base of the blade. It runs diagonally to the outer margin of the mandible and continues down the ventral surface. In some species this groove takes the form of a broad channel (attenuata) whilst in others it is quite small (furtiva). In other species it may be reduced, vestigial or absent (mactans, guineensis-group), but it is particularly marked in members of the stuhlmanni-group. In species in which this groove is present there is an apparently associated thin- spot or membranous area on the dorsal surface of the mandible, close to the base of the groove and on its inner side (towards the inner margin of the mandible). In the long mandibulate species the thin-spot is usually visible even when the mandibles are closed, but in others the spot is usually concealed by the leading edge of the clypeus unless the mandibles are open. The function of these structures is not known, nor can any reason be suggested for their absence in some species. SPECIES OF THE ETHIOPIAN REGION LIST OF SPECIES conradti-group conradti Forel crustosa Santschi conradti var. rufipes Santschi syn. n. africanus Weber syn. n. REVISION OF GENUS LEPTOGENYS 243 maxillosa-group faicigera Roger insularis F, Smith jeanneli Santschi maxillosa (F. Smith) falcata Roger vinsonnella (Dufour) syn. n. cribvata Emery (provisional synonym) pavesii Emery maxtllosa subsp. sericea Weber syn. n. stuhlmanni-group camerunensis Stitz stuhlmannii race camerunensis var. angusticeps Forel [unavailable] stuhlmanni subsp. camerunensis var. opalescens Wheeler [unavailable] erythraea Emery stat. n. leiothorax Prins. stat. n. nebra_ sp.n. regis sp.n. schwabi Forel sterops sp.n. stuhlmanni Mayr comorensis Forel syn. n. titan sp.n. vindicis sp. n. zapyxis sp.n. sulcinoda-group bellii Emery elegans sp. n. excellens sp.n. Serrarii Forel ferrarii st. dentatula Santschi syn. n. longiceps Santschi mastax sp.n. nuserra sp. n. occidentalis Bernard stat. n. ferrarit subsp. sulcinodis Bernard syn. n. ferrarii subsp. bernardi Baroni Urbani syn. n. princeps sp.n. ravida sp.n. sulcinoda (André) terroni sp.n. trilobata Santschi attenuata-group attenuata (F. Smith) jaegerskjoeldi Santschi bubastis sp.n. crassinoda Arnold stat. n. ergatogyna Wheeler cursor Arnold syn. n. guineensis-group guineensis Santschi microps sp. n. spandax sp.n. testacea (Donisthorpe) 244 B. BOLTON havilandi-group arnoldi Forel furtiva Arnold stat. n. havilandi Forel peringueyi Forel nitida-group amon sp. n. ankhesa_ sp. n. buyssoni Forel castanea (Mayr) parva Forel syn. n. parva var. bellua Forel syn. n. parva var. dispar Santschi syn. n. hewitti Santschi syn. n. cryptica sp.n. diatra sp.n. honoria sp.n. khaura_ sp.n. mactans sp.n. nitida (F. Smith) intermedia Emery syn. n. tenuis Stitz nitida var. adpressa Forel syn. n. nitida var. aena Forel syn. n. nitida var. gracilis Santschi syn. n. nitida st. insinuata Santschi syn. n. nitida var. gyandiory Forel syn. n. nitida race brevinodis Forel syn. n. nitida st. speculans Santschi syn. n. brevinodis var. deflocata Santschi syn. n. nitida var. capensis Baroni Urbani syn. n. piroskae Forel strator sp.n. Striatidens sp.n. stygia sp.n. Note. A number of the synonyms given above in nitida and maxillosa-groups have also been determined independently by Prof. W. L. Brown and will be recorded by him in his forthcoming revision of the genera of tribe Ponerini. KEY TO SPECIES (Workers) 1 With petiole in eas the posterodorsal angle projecting as a spine or tooth oe 24) — With petiole in ‘profile the ‘posterodorsal angle not projecting as a spine or tooth 2 Rugosity of head and alitrunk overlaid by a dense puncturation. Scapes, legs, and at least the first gastral tergite reticulate and dull. (Guinea, Gabon, Cameroun, Zaire, Sudan . : crustosa (p. 251) — Rugosity of head and alitrank not ‘overlaid by puncturation. Scapes and legs usually shiny, not reticulate; first aap a ssid satel (Ivory Coast, Ghana, Cameroun) : : . conradti (p. 2 12 13 REVISION OF GENUS LEPTOGENYS Entirety of head, alitrunk and gaster densely shagreened and opaque, with scattered larger punctures and short, dense, greyish, pruinose pubescence : Not entirely shagreened, usually not opaque, at least the gaster usually shiniai. If finely and densely reticulate-punctate everywhere then scattered larger punctures absent and dense pruinose pubescence lacking ° Dorsum of head, alitrunk and first two gastral tergites with numerous standing hairs. Larger species, HL > 1°75. (Tanzania) : d jeanneli (p. Dorsum of head, alitrunk and first two gastral tergites without standing hairs. Smaller species, HL < 1-75 Fringing lamella of clypeus broad and broadly rounded. Outer margin of mandible straight or concave in its distal half (Text-fig. 1). Sides of head not strongly divergent anteriorly in full-face view. (Somalia, Sudan, Cameroun, Kenya, Tanzania; tramp species in Old World tropics) F . pavesii (p. Fringing lamella of clypeus narrow, either narrowly rounded or obtusely angulate medially. Outer margin of mandible convex throughout its length aan fig. 2). Sides of head divergent anteriorly i in full-face view . ‘ ‘ Node of petiole in dorsal view distinctly broader than long, DPI > 105. (Zaire, Rhodesia, S.W. Africa; pantropical tramp species) . : maxillosa (p. Node of petiole in dorsal view at least as long as broad, usually distinctly longer than broad, DPI 100 or less (pantropical tramp species) . ; falcigera (p. First and second gastral tergites without hairs, with sparse, strongly adpressed pubescence. Eyes minute, their maximum diameter less than the maximum width of the scape. Gaster very weakly or not constricted between the first and second segments : F : First and second gastral tergites with hairs, often also “with pubescence. If gastral hairs sparse then either eyes larger, their maximum diameter greater than the maximum width of the scape, or gaster strongly constricted between first and second segments, or both Pronotal dorsum with a pair of long, erect hairs anteriorly, much longer than the remaining pilosity. Minute species, HL < 0-70, PW < 0-42, with minute eyes, diameter < 0:05. (Ivory Coast) : : : microps (p. Pronotal dorsum without a pair of long, erect hairs anteriorly, pilosity absent or hairs of approximately equal length. Larger dean HE > 0-75; PW > 0-45, with larger eyes, diameter > 0:07 . : : ; ° ; : Pronotal dorsum without short, standing hairs. Petiole in profile a thick scale, tapering dorsally (Text-fig. 28). Larger species with relatively longer antennal scapes, HL > 1:25, SI > 140. (Ghana) - : spandax (p. Pronotal dorsum with numerous short, standing hairs. " Petiole in profile not a thick scale but a high, narrow node. Smaller species with relatively shorter scapes, HL < b10; Si -< 130 . : Apex of mandible with three teeth; apical plus ine preapical (Text- fig. 2 3). (Ghana) testacea (p. Apex of mandible with two teeth, apical plus one preapical. (Guinea, Ghana) guineensis (p. Petiole in profile or in dorsal view with the upper halves of the sides longitudinally rugose or sulcate . Petiole in profile or in dorsal view ; with ‘the upper halves of the sides not longitudinally rugose or sulcate Pronotal dorsum unsculptured apart from pits from which hairs arise Pronotal dorsum sculptured, at least anteriorly In full-face view the occipital margin of the head feebliy convex (Text- fig. 4), Petiole node in profile longer, LPI < 140. (Gabon). ; : sulcinoda (p. In full-face view the occipital margin of the head broadly concave (Text-fig. 13). Petiole node in profile shorter, LPI > 145. (Guinea) ' . occidentalis (p. 245 253) It 278) 272) 270) 246 B. BOLTON 14 Larger species, HL > 1-00, PW > 0:58, with relatively longer head and scapes, CI < 65, SI > 140. (Ivory Coast, Ghana) : mastax (p. 268) — Smaller species, HL < 1-00, PW < 0°58, with relatively shorter. head and scapes, Cl > 65, SI <135. (Ivory Coast, Ghana) : nuserra (p. 269) 15 Mandibles extended into elongate, narrow, linear or - sublinear blades, the inner margins of which are often concave. Mandibles not capable of closing tightly against the median lobe of the clypeus, which is often reduced, enclosing a ae (sometimes large) between themselves and the clypeus (Text-figs 7-11) ; 16 — Mandibles short and usually quite stout, their inner margins usually distinctly convex. Mandibles capable of closing tightly against the median lobe of the clypeus, never enclosing a space between themselves and the clypeus (Text-figs 13-18) : > 35 16 Antennal scapes with SL >1.90. Gaster very weakly or not constricted between the first and second segments. Metanotal groove not cross-ribbed or sculptured en 4 Antennal scapes with SL < 1-70. Gaster distinctly constricted between first and second segments. Metanotal groove distinctly cross-ribbed or otherwise sculptured 27 17 With head in full-face view eyes not breaking the outline of the sides of the head (Text-fig. 8). Antennal scapes with dense pubescence but without erect hairs which are as long as the maximum width of the scape. Large species with relatively short scapes, HL > 2-10, SI < 170. A 18 — With head in full-face view eyes breaking the outline of the sides of the ‘head (Text- figs 4, 7). Antennal scapes with numerous erect hairs which are as long as or longer than the maximum width of the scape; pubescence on scapes usually sparse. Smaller species with relatively long scapes, HL < 2:10, SI > 180 : A 19 18 Dorsum of pronotum and propodeum unsculptured apart from pits from which hairs arise. Maximum diameter of eye about 0.66. (Cameroun) ; . titan (p. 263) — Dorsum of pronotum and propodeum, especially the latter, sculptured. Maximum diameter of eye about 0:50. (Kenya, Tanzania) ; . regis (p. 259) 19 Entire dorsum of alitrunk completely unsculptured. Sides of alitrunk unsculptured apart from cross-ribbing at meso-metapleural suture and a few rugae around the bulla of the metapleural gland. Full adult colour brown or deep red-brown, with bluish or violet opalescence, which may be patchy . P : ioe") — Propodeal dorsum at least weakly, usually strongly sculptured ; ronnie sometimes sculptured, often not. Sides of alitrunk with plurae and propodeum striate or rugose. Full adult colour black or brown but without opalescence . Zt 20 Larger species with antennal scapes both absolutely and relatively longer, SL > 2: os 5, SI > 230. (Cameroun, Zaire) camerunensis (p. 256) — Smaller species with antennal scapes both absolutely and relatively shorter, SL < 2-60, SI < 220. (Mozambique) . , ‘ . leiothorax (p. 258) 21 Inner margin of mandible with a convex, semi- scansiieent lamella (Text-fig. 21). (Ghana) : ; zapyxis (p. 264) — Inner margin of maudible sithiont a convex, semi- Panslibent lamella (Text-figs 6,20) 22 22 Antennal scapes relatively shorter, SI < 200 : ; : : : ; ees — Antennal scapes relatively longer, SI > 200 ‘ : : et 23 Larger species, HL > 1-80, with maximum diameter of eye 0:50 0r more. Pronotal dorsum weakly sculptured. With mandibular apices overlapping the basal portion of each blade passing through a curve before its junction with the clypeus. (Sudan, Kenya, Mozambique, Comoro Islands) . ; : . stuhlmanni (p. 261) — Smaller species, HL < 1-60, with maximum diameter of eye < 0°45 (ca 0-40). Pronotal dorsum unsculptured apart from minute hair-pits. With mandibular apices overlapping the basal portion of each blade not passing through a curve before its junction with the clypeus. (Ivory Coast) . ; , sterops (p. 261) 24 Dorsum of head unsculptured apart from minute hair-pits: pronotal dorsum usually the same but rarely with some faint sculpturation. . : 4 : : ~ 25 25 26 27 28 29 30 31 32 33 34 REVISION OF GENUS LEPTOGENYS Dorsum of head and pronotum distinctly sculptured everywhere. (Cameroun) vindicis (p. Larger species with very large eyes, HL > 1-90, maximum diameter of eye 0°54 or more. Full adult colour brown. (Ethiopia) . ; , ; erythraea (p. Smaller species with smaller eyes, HL < 1 ae maximum diameter of eye 0:48 or less. Full adult colour black Eyes larger, maximum diameter in the range 0- 44-0: 48 (in HW range of I- car oar Width of head across eyes the same as or only slightly less than the width across the clypeus from corner to corner. (Rhodesia, South Africa) . . . schwabi (p. Eyes smaller, maximum diameter about 0-36 (with HW ca 1-22). Width of head across eyes much less than width across the clypeus from corner to corner. (Cameroun) . are nebra (p. Dorsum of head finely rugose or peiculstestd soot: dine ieemmipal spaces with numerous fine punctures ; Dorsum of head not rugose nor reticulate-rugose sther punctate renpuinte: eomacbate or smooth and shining Pronotum with a fine and very Wate reticulation! whieh covers the eatre dstedin also with scattered hair-pits. aes cae BL > 5730, SL > 1-40, Sl > 125: (Cameroun) ; : . terroni (p. Pronotum without a fine reticulation, at eee with a bey Bea rounded rugulae. Smaller species, HL < 1-20, SL < 1-20, SI < 120. (Cameroun, Zaire) trilobata (p. Sides of head below and behind eyes and lateral portions of ventral surface of head strongly sculptured. (Zaire) ‘ : excellens (p. Sides of head below and behind eyes and ister Portions of Senta surface of head unsculptured except for hair-pits and very rarely some weak, superficial reticulation . : Dorsum of head coarsely aa closely reticulate: spanoete the pafictires tot eee by smooth areas. Larger species, HL >1-:20, PW >0:°-75. Node of petiole massive, PL > 0:60, PH > 0°75. . - ‘ princeps (p. Dorsum of head with scattered punctures which are reeparated by smooth areas. Smaller species, HL < 1:15, PW < 0-70. Node of Beta smaller, PL < 0°55, FH <.6-70 ; Mesopleuron completely reticulate. spalictate, with numerous ieangvene eupaiaes Petiole dorsum and upper halves of sides reticulate-punctate superimposed upon a coarser rugulation. (Senegal, Ghana) . ; ; ; longiceps (p. Mesopleuron not reticulate-punctate; a few puncnires and transverse rugulae may be present but there are extensive unsculptured areas. Petiole dorsum and sides not reticulate-punctate Middle of posterodorsal margin of peabis slightly projecting as an p ebtane tabercis, (Ethiopia) . : Z . bellii (p. Middle of posterodorsal tare of pstinte HE projecting asa saber : Eyes smaller, their maximum diameter approximately equal to the length of tte third funicular segment (Text-fig. 10). Head black, very polished, with only a few scattered, minute punctures. (Zaire) : : . ravida (p. Eyes larger, their maximum diameter markedly sieator thank the length of the third funicular segment (Text-fig. 9). Head brown or black, if black then with numerous quite distinct punctures . : Larger species, HL > 0:95, SL > 0-85, with Neaber pecs LPI < ee Full 258) 273) 273) 266) 30 271) 31 268) 32 265) 33 272) 34 adult colour black. (Ivory Coast, Ghana, Nigeria, Cameroun) . elegans (p. 265) Smaller species, HL < 0-90, SL < 0-85, with shorter aac LPI > 140. Fulladult colour brown. (Zaire, Rhodesia) . : : ; ferrarii (p . 267) 248 36 37. 43 44 45 46 B. BOLTON Propodeal dorsum with a longitudinal groove medially, and anteriorly with a broad, transverse impression posterior to the metanotal groove. Mandibles very broad, massively developed (Text-fig. 22). Antennal scapes relatively very long, SI > 220. (Ghana, Uganda, Cameroun, Zaire) : : . ergatogyna (p. Propodeal dorsum without a median groove and transverse a ae Sete: Mandibles not as above. Antennal scapes shorter, SI < 200 Dorsum of head coarsely ai aa eae or very densely punctulate c over most or all of its surface . E : Dorsum of head either aioe and inte with aaly deatiacid punetnies from which hairs arise, or rarely superficially finely reticulate ° ‘ Minute species, HL < 0-85, PW < 0.50. Eyes small or minute, wee maximum diameter much less than the maximum width of the scape. Second funicular segment at most as long as third, usually much shorter than third Larger species, HL > 1-10, PW >0:-65. Eyes large, their maximum gianna greater than the maximum width of the scape. Second funicular segment distinctly longer than the third Pronotal dorsum unsculptured, smooth and shining (Nigeria) . khaura (p. Pronotal dorsum strongly sculptured . : P , : : Antennal scapes longer, SI 119 or more. Eyes eee with more than to facets. Full adult colour black. (Nigeria, Cameroun) : : . Sstygia (p Antennal scapes shorter, SI 115 or less. Eyes smaller, its less than to facets. Full adult colour yellow-brown. (Ghana) ‘ cryptica (p. First and second gastral tergites covered with a fine, aan. Pelcuinte. pun tan tion or granulation, without smooth, shining areas. (Rhodesia) ; . furtiva (p. First and second gastral tergites a oa smooth and go apart from pits from which hairs arise Pronotal dorsum with scattered small cnits scpaeiicd by ator areas, eathout a ans blanketing sculpture on any part of the sclerite. (South Africa) peringueyi (p. Pronotal dorsum with a fine, dense sculpturation, blanketing the sclerite at least in part. : : . s : c ° : : : < : Fringing lamella of clypeus narrow and indistinct. Antennal scapes relatively short, SI < 140. Eyes large, maximum diameter 0-34 or more. (Rhodesia) arnoldi (p. Fringing lamella of clypeus broad and distinct, usually opaque. Antennal scapes relatively long, SI > 140. Eyes smaller, maximum diameter 0-30 or less. (South Africa) . ; ; : havilandi (p. Ventral process of petal comnles. agabie cote ste of a large lobe or tooth antero- ventrally and a smaller lobe or tooth situated more pore the two separated by a distinct space or indentation (Text-fig. 25) Ventral process of petiole simple, single, consisting only of a ie or Footh antero: ventrally, of varying configuration Sides and ventral surface of head sculptured. Node of pete in ‘profile decteusing slightly in thickness from base to apex. (Zaire) ; : . Strator (p. Sides and ventral surface of head not sculptured. Node of petiole’ in profile increasing slightly in thickness from base to apex Pronotal dorsum with numerous shallow, foveolate impressions saparaeed by smooth areas. Smaller species, SL < 1-10, with relatively shorter antennal scapes, SI < 135. (Cameroun) : - : , honoria (p. Pronotal dorsum smooth, iiihout Eepraclats impressions. Larger species, SL > 1°10, with relatively longer antennal scapes, SI > 135. (Zaire) Antennal scapes with SL > 1:40 Antennal scapes with SL < 1-30 ankhesa (p. 276) 36 37 43 38 40 288) 39 Pp. 292) 286) 281) 41 282) 42 281) 282) 44 46 291) 45 287) 284) 47 50 47 48 49 REVISION OF GENUS LEPTOGENYS Maximum diameter of eye in full-face view less than maximum width of scape. Ventral process of petiole a broad lobe with a posteroventral tooth. Full adult colour red-brown. (Ethiopia) . ; ‘ buyssoni (p Maximum diameter of eye in full-face view "greater than maximum width of scape. Ventral process of petiole a simple or acute lobe. Full adult colour black . Larger species with relatively longer head and antennal scapes and larger eyes; HL 1:50 or more, CI < 65, SI > 170, ocular diameter 0:36 or more. Node of petiole in dorsal view very strongly narrowed anteriorly, much broader behind than in front, the sides feebly concave (Text-fig. : Head and body usually with bluish opalescence. (South Africa) : . attenuata (p Smaller species with relatively shorter head and antennal scapes and smaller eyes; HL < 1-45, CI > 65, SI < 165, ocular diameter 0-32 or less. Node of petiole in dorsal view only slightly narrower in front than behind pins oo): Head and body shiny black, without bluish opalescence : é ‘ Mesonotum and propodeum without scattered large pits or " punctures, Propodeal dorsum shorter, about 1-10 in profile from metanotal groove to apex of prominence on declivity in workers with HW range 0-96—1-o00. (South Africa) crassinoda (p. Mesonotum and propodeum with numerous scattered large pits or punctures. Propodeal dorsum longer, about 1-30 in profile from metanotal groove to apex of prominence on declivity in workers with HW range 0:96-1:00. (Cameroun) bubastis (p. Antennal scapes relatively very short, SI < 95. (South ee ‘ mactans (p Antennal scapes relatively long, SI 100 or more ‘ : ; é : Mandibles covered with dense, fine, longitudinal striation. Dorsum of head and pronotum with dense but superficial reticulation. (Cameroun) ._ striatidens (p Mandibles smooth and shining, not striate. Dorsum of head and pronotum smooth, without superficial reticulation Circumocular impression very broad and deep in front of, above and ‘behind eye, so that the eye appears to be sunk in a concavity in the side of the head (Text-fig. 18). Node of petiole in dorsal view much ee than broad, DPI < 70; head long and narrow, CI < 65. (Cameroun) : . amon (p Circumocular impression very weak or absent, the eye not sunk into a concavity in the side of the head. Node of petiole much broader, DPI 80 or more; head shorter and broader. CI 69 or more Antennal scapes both relatively and absolutely shorter, SL 0°75 (range 0:80-1:08), SI 120 or more (range 120-130) : : : : ; Smaller species, HL ca 0:86, HW ca 0:62, SL ca 0-80. "Ventral process of petiole an elongate, recurved tooth. (Cameroun) . . diatra (p Largerspecies, HL > 0:90, HW > 0:65, SL>0°8 5. Ventral process of petiole a lobe, the posteroventral angle of which may be dentiform, but not as above Eyes larger, ocular diameter greater than 0-11, the eyes set well forwards on the Sides of the head. The maximum diameter of the eye equal to or greater than the maximum width of the scape in full-face view. (Eastern and southern Africa) nitida (p. Eyes small, ocular diameter ca 0-09, the eyes set well back on the sides of the head (Text-fig. 15). The maximum diameter of the eye much less than the maximum width of the scape in full-face view. (Ethiopia) ‘ . ° piroskae (p 249 . 285) 48 - 274) 49 276) 275) . 288) 5ST . 292) 52 . 283) 53 285) 54 . 287) 55 289) - 290) 250 B. BOLTON THE CONRA DTI-Grovup Mandibles elongate and linear, curved, not closing tightly against the clypeus. Eyes large, their maximum diameter in full-face view greater than the maximum width of the scape. Second funicular segment elongate, distinctly longer than the third. Head strongly broadened anteriorly, much broader in front than behind. Posterodorsal angle of petiole in profile drawn out into a tooth or spine (Text-fig. 24). Sculpture coarse, rugose upon the head and alitrunk. Gaster constricted between the first and second segments. Clypeus with a prominent median lobe, without a fringing lamella. Standing hairs present on all dorsal surfaces of head and body. The two species constituting this small group are easily distinguished from their congeners by the characters listed above, the fifth of which is diagnostic. Related species with this character are also known from the Neotropics (unistimulosa Roger and its allies). The two species of the Ethiopian Region seem to be confined to forested zones but are widely distributed. Leptogenys conradti Forel Leptogenys convadti Forel, 1913b : 666. Syntype workers, CAMEROUN: (Conradt) (MHN, Geneva) [examined]. DIAGNOSIS OF WORKER. Posterodorsal angle of petiole in profile drawn out into a spineor tooth. Head and alitrunk rugose, not overlaid by punctures. First gastral tergite smooth and shining. FURTHER DESCRIPTION. Worker. TL 9:4~-10-4, HL 1:66-1:76, HW 1-40-1-48, CI 81-85, SL 1-92-2:20, SI 145-155, PW 1°18—1-24, PH 1-10—-1-18, PL 0-g0o—-0-96, LPI 117-125, DPW 0°76-0:84, DPI 83-87 (12 measured). Head considerably broader in front than behind, the occipital border acutely margined and broadly but shallowly concave. Eyes large, maximum diameter ca 0-40-0°44, their greatest diameter in full-face view greater than the maximum width of the scape. Circumocular impression present on inner and posterior margins of eyes. Clypeus with a rounded prominent median lobe. Mandibles elongate, becoming slightly broader from base to apex, most strongly curved near the base and enclosing a large space between them and the clypeus when their apices are overlapping. Second funicular segment long, the respective lengths of segments I-3 Ca 0*26-0°30, 0-44—0'50, 0°36-0°40. Metanotal groove impressed and cross-ribbed. Petiole in profile massive, the posterodorsal angle produced into a tooth which is variously developed amongst individuals but is usually distinctly dentiform. Head, alitrunk and petiole coarsely rugose, the rugae on the head predominantly longitudinal, more sharply developed in some individuals than in others. Gaster unsculptured, smooth and highly polished. Legs and antennae usually shining, but rarely with some fine punctures. Standing hairs abundant upon all surfaces. This species and crustosa are characterized by the projection of the posterior portion of the petiole node into a spine or tooth. They are separable by the charac- ters given in the key and the diagnoses above and also by the fact that crustosa averages smaller than contradti although this may be found to be incorrect as the species become better known. A single specimen in the MCZ, Cambridge collection may represent a third species in this group. It resembles conradiz closely but is smaller, HL 1-38, HW 1-14, (C183), SL 1-38, (SI 121), PW 0-96. Note also that the scapes are relatively much shorter. The posterodorsal prominence of the petiole is short and blunt and the clypeus is less acutely carinate than in conradtt. The sculpture of the cephalic dorsum in REVISION OF GENUS LEPTOGENYS 251 this specimen consists of numerous fine rugulae, not arranged into any pattern or direction. This specimen may represent a distinct species or may be an extremely aberrant or first-brood specimen of conradti, but as only a single worker is available it is not possible at present to decide upon its status. Accordingly I have labelled it ‘con- vadti?’ in the MCZ, Cambridge collection. MATERIAL EXAMINED. Ivory Coast: Lamto, Toumodi(J. Lévieux) ;O.R.S.T.O.M. Exp. Sta., Abidjan (W.L. Brown). GHANA: Tafo (B. Bolton); Bosuso (D. Leston); Mt Atewa (D. Leston); Legon (D. Leston); Sajimasi (D. Leston); Aburi (D. Leston). CAMEROUN: Ototomo (G. Terron); Mbale Mejo to Ekingli (G. Schwab). Leptogenys crustosa Santschi (Text-fig. 24) Leptogenys crustosa Santschi, 1914a : 326, fig. 12. Syntype workers, GUINEA: Conakry, 8.viii.1913 (F. Silvestvi) (IE, Naples). Leptogenys (Leptogenys) conradti var. rufipes Santschi, 1937 : 75. Holotype worker [not female], Gason: Samkita (F. Faure) (NM, Basle) [examined]. Syn. n. Leptogenys (Leptogenys) africanus Weber, 1942 : 47, fig. 8. Holotype and paratype workers, SupDAN: Lotti Forest, W. slopes of Imatong Mountains, 5.viii.1939 (N. A. Weber) (MCZ, Cam- bridge) [examined]. Syn. n. DIAGNOSIS OF WORKER. Posterodorsal angle of petiole node in profile drawn out into a blunt spine. Sculpturation of head and alitrunk rugose, overlaid by a fine, dense puncturation. First gastral tergite coarsely reticulate. FURTHER DESCRIPTION. Worker. TL 7-3-9:1, HL 1-34-1-64, HW 1-08—-1-38, CI 79-84, SL 1°48-1-90, SI 136-137, PW 0-92-1-20, PH 0:86-1-12, PL 0°74-1-00, LPI 112-116, DPW 0:64-0:86, DPI 86-87 (4 measured). Head markedly broader in front than behind, in the largest specimen examined width of head in front of eyes at maximum was I-60, and width across occipital margin 0-96. Eyes large, maximum diameter 0-33—0-46, surrounded on their internal margins by a distinct circumocular impression. Maximum diameter of eye much greater than maximum width of scape. Clypeus with a rounded median lobe. Mandibles elongate and narrow, not closing tightly against the clypeus, enclosing a large gap when their apices are in contact. Mandibular blades most strongly curved basally. Second funicular segment elongate, the respective lengths of segments I-3 Ca 0*30—0°40, 0°34—0'44, 0:28-0°30. Petiole in profile with the posterodorsal angle produced into a short spine. Basic sculpturation of head, alitrunk and petiole a coarse rugosity, pre- dominantly longitudinal upon the head, but everywhere the rugosity is overlaid by a dense puncturation, which is more strongly developed in large than in small individuals. First and second gastral tergites, or at least their basal halves, with a relatively coarse superficial reticu- lation. This is also present upon the legs and scapes, which are consequently dull. Standing hairs numerous upon all surfaces. Colour black, with the mandibles, antennae and legs red- brown or orange-brown. MATERIAL EXAMINED. CAMEROUN: no loc. (G. Terron). ZAIRE: Elisabetha (Tinant): Ituri, Okondo (A. Collart). 252 B. BOLTON THE MAXILLOSA-GrovupP (Text-figs I, 2, 3, 31) Mandibles elongate and linear, curved, not closing tightly against the clypeus. Eyes large, their maximum diameter in full-face view greater than the maximum width of the scape. Second funicular segment not, or only very slightly longer than the third. Head slightly or strongly broadened anteriorly, varying among species. Sculpture of a dense shagreening everywhere with scattered larger punctures. Entire body with dense, greyish pruinose pube- scence. Gaster constricted between first and second segments. Clypeus without a prominent median lobe; with a fringing lamella. A further character of value in separating three of the four species of this group (pavesit, maxillosa and falcigera) from all others of the Ethiopian Region except for some members of the guineensis-group is that they lack standing hairs on the dorsum of the head, alitrunk, petiole and first two or three gastral tergites. This character also applies to pruinosa Forel, an extralimital species of the maxillosa-group described from Sri Lanka. Forel (1g01b : 81) stated that pruinosa was found on wood imported into Hamburg from East Africa and, as I am of the opinion that the maxillosa-group is of African origin, I would not be unduly surprised if pruinosa were discovered at some time in the future in Kenya or Tanzania. All species of this group in the Ethiopian region with the exception of the large geannelt are tramp species in the tropical zones of the world and probably rely on human commerce for transportation. To judge from the number of collections examined, falcigera has been able to establish itself in some strength in the Hawaian Islands, whilst maxillosa has been recorded from both Cuba and Brazil. The maxillosa-group appears to be related to truncatirostris and its allies of the Malagasy Region, but in these species standing hairs are present on the head and body and the mandibles tend to be shorter and broader though linear or sublinear in form. Leptogenys falcigera Roger (Text-fig. 2) Leptogenys falcigera Roger, 1861 : 42. Holotype worker, Sr1 LANKA (H. Nietner) (location of type not known). Leptogenys insularis F. Smith, 1879 : 675. Syntype workers, Hawalr: Oahu Is. (T. Blackburn) (BMNH) [examined]. [Synonymy by Wilson & Taylor, 1967 : 30.] DIAGNOSIS OF WORKER. Densely shagreened with scattered larger punctures. Without standing hairs. Petiole in dorsal view as long as or longer than broad. Lamelliform margin of clypeus narrow and translucent. Outer margin of mandible convex throughout its length, FURTHER DESCRIPTION. Worker. TL 7:0-7:8, HL 1:40-1°52, HW 1°32-1°46, CI 93-06, SL 1-32-1-48, SI 98-104, PW 0:92-1:02, PH 0-84-0-90, PL 0:66-0:78, LPI 115-127, DPW 0:68—0-70, DPI 90-100 (8 measured). Head broadened in front of the eyes, the sides weakly convex but diverging anteriorly throughout their length. Eyes large, their maximum diameter ca 0:32-0:38. In full-face view their maximum diameter much greater than the maximum width of the scape. Anterior border of clypeus with a narrow, fringing lamella which is translucent and is narrowly rounded or forms an obtuse angle anteromedially. Mandibles long and linear, their outer margins convex throughout the length of the blade, most strongly convex near the base. Metanotal REVISION OF GENUS LEPTOGENYS 253 groove weak, virtually obliterated. Node of petiole in dorsal view usually longer than broad, more rarely about as long as broad. Entire body densely shagreened and with numerous scattered larger punctures. Pruinose pubescence present everywhere but standing hairs absent except on the clypeus and gastral apex. Within this species-group jeannelt is easily separated by its possession of standing hairs upon the dorsal surfaces of the head and body, but the other three members form a complex of closely related species which in all likelihood arose in the Ethiopian Region although all three are now widely distributed tramp species; and there is at least one other relative (prwinosa Forel) at present unknown in the Ethiopian Region. Both falcigera and maxillosa have a relatively narrow clypeal lamella which is translucent and is narrowly rounded or forms an obtuse, rounded angle medially. Also in both these species the outer margin of the mandible is convex throughout its length and the mandibles are long and narrow. The head is strongly broadened anteriorly and the sides are not as convex as in pavesii. In pavesii the clypeal lamella is broad and broadly and evenly rounded, not forming an angle medially, and is usually opaque. The outer margin of the mandible is convex basally but distal to the convexity is usually flat or concave for part of its length. The mandi- bular blades themselves are shorter and stouter than in maxillosa or falcigera. The head is not strongly broadened anteriorly and the sides are more convex in pavesii. The separation of falcigera from maxillosa rests most convincingly upon the shape of the petiole node in dorsal view. In maxillosa the node is always distinctly broader than long, with DPI > 105, whilst in falcigera the node is at most as broad as long, usually distinctly longer than broad, and with a maximum DPI of roo. In the present study /falcigera has not been recorded from the mainland of the Ethiopian Region but its range is expected to extend at least into the eastern parts of the continent. It has been previously recorded from Madagascar, Aldabra, Sri Lanka and the Hawaian Islands. MATERIAL EXAMINED. ALDABRA Is.: South Is. (J. G. Frazier). Sri LANKA: Peraden (C. T. Bingham). Hawa: Wailuku, Maui (R. C. L. Perkins); Hilo (G. Poinar); Honolulu (K. Oshiro). Leptogenys jeanneli Santschi (Text-figs 3, 31) Leptogenys jeanneli Santschi, 1914c : 55, fig. 3. Holotype worker, TANZANIA: Grotte B. du Kulumuzi, a Tanga (st. no. 74), iv. 1912 (Alluaud & Jeannel) (NM, Basle) [examined]. DIAGNOSIS OF WORKER. Entirety of head and body densely shagreened and opaque, with scattered larger punctures. Numerous standing hairs present on the dorsal surfaces of the head and body. FURTHER DESCRIPTION. Worker. TL 9:6, HL 1:86, HW 1°72, CI 92, SL 1-94, SI 113, PW 1:08, PH 0-98, PL 0-94, LPI 104, DPW 0-72, DPI 76. Head broader in front than behind, maximum width in front of eyes > 1-80, width across occipital margin <1-40. Eyes large, ocular diameter at maximum 0-42, the maximum diameter much greater than the maximum width of the scape. Eyes with a pronounced 254 B. BOLTON circumocular impression on their inner margins. Clypeus medially with a narrow, translucent fringing lamella. Mandibles long, narrow and curved, leaving a large gap between them at full closure. Outer margins of mandibular blades convex throughout their length, most strongly curved basally; tapering from base to apex and terminating in an acute apical tooth. Preapical teeth absent. Respective lengths of funicular segments I-3 ca 0:28, 0-40, 0:34. Metanotal groove poorly developed, only weakly impressed and not cross-ribbed. Petiole in profile nodi- form. Gaster strongly impressed between first and second segments. All surfaces of head and body are densely shagreened, with scattered larger punctures. Short, standing hairs numerous upon all dorsal surfaces. Short, greyish, dense pruinose pubescence abundant everywhere. Black, dull; the antennae, legs and mandibles brown. This quite distinctive species is apparently known only from the holotype. It is easily separated from the other members of the group by its possession of standing hairs on the dorsal head and body, and by its large size. Leptogenys maxillosa (F. Smith) Ponera maxillosa F. Smith, 1858 : 93. Syntype workers, males, Mauritius: (Dr Beke) (BMNH) [examined]. Leptogenys maxillosa (F. Smith); Roger, 1861 : 43. Leptogenys falcata Roger, 1861 : 42. Syntype workers, CuBA & Brazi (location of types not known). [Synonymy by Forel, 1901b : 46.] Formica vinsonnella Dufour, 1864 : 210. Holotype ? worker, REUNION (location of type not known). Syn. n. Leptogenys maxillosa var. vinsonella (Dufour); Emery, 1894 : 68; Forel, 1912b : 159 (Seychelles): Leptogenys cribvata Emery, 1895a : 20. Syntype workers, SoutH AFRica: Vrijburg (E. Simon) (MCSN, Genoa). Provisional synonym. DIAGNOSIS OF WORKER. Densely shagreened with scattered larger punctures. Without standing hairs. Petiole in dorsal view broader than long. Outer margin of mandible convex throughout its length. FURTHER DESCRIPTION. Worker. TL 6-8-7:5, HL 1:38-1:48, HW 1:28-1:46, CI 91-08, SL 1+30-1:60, SI 101-110, PW 0:96-1:04, PH 0:84-0:90, PL 0:64-0:68, LPI 126-140, DPW 0:68-0:76, DPI 106-113 (8 measured). Head broadened in front, the sides feebly convex and diverging anteriorly throughout their length. Eyes large, maximum diameter ca 0-32-0-36, considerably greater than the maximum width of the scape. Anterior clypeal margin with a narrow, translucent fringing lamella. Mandibles long and slender, the outer margins of the blades convex throughout their length. Node of petiole in dorsal view distinctly broader than long. Body and head everywhere densely shagreened with scattered larger punctures. Dense, pruinose pubescence present everywhere but standing hairs restricted to the clypeus and gastral apex. L. maxillosa is a reasonably common tramp species throughout both the Old and New World Tropics, but appears to originate in the central and southern portions of the Ethiopian Region where it is a widely distributed but relatively uncommon species. Its closest relative within the group is falcigera, but in this species the petiole node is at least as long as broad, usually distinctly longer than broad, whilst in maxillosa the node is always considerably broader than long. In maxillosa the scapes are usually somewhat longer, SI 101-110, as opposed to SI g8—104 in falcigera and the LPI overlaps slightly, falcigera LPI 115-127 (mean 119), maxillosa LPI 126-140 (mean 133). I have not been able to see the types of vinsonnella or cribrata but in the case of REVISION OF GENUS LEPTOGENYS 255 the former I have examined series from numerous islands and systems near Réunion and all the specimens have been maxillosa. Similarly, maxillosa is the only species which fits the original description of cribrata and to the best of my knowledge is the only species of this group occurring in southern Africa. Consequently I have placed these two names in the synonymy of maxillosa, but I place cribrata there only provisionally as a future examination of the types may prove the present contention to be incorrect. MATERIAL EXAMINED. ZAIRE: 13 mi. N. of Kienge (EF. S. Ross & R. E. Leech). RHODESIA: Bulawayo (G. Arnold); Honde Valley, Mtarazi R. (W. L. Brown). SouTH WEsT AFRICA: Lontpansberg (Vernay-Lang). SoutH Arrica: Natal, Dukuduku (Grobler). SEy- CHELLES: Bird Is. (V. Fitzgerald); Dennis Is., Amirante Is., Coetivy Is. (P. Sladen expedition). Mauritius: (J. E. M. Brown). CHaGos ARCHIPELAGO (M. J. D. Hirons). CusBa: Soledad, Cienfuegos (N. A. Weber); no loc. (coll. F. Smith). Leptogenys pavesii Emery (Text-fig. 1) Leptogenys pavesii Emery, 1892 : 111. Syntype workers, SomaLia (Robecchi) (MCSN, Genoa). Leptogenys (Leptogenys) maxillosa subsp. sericea Weber, 1942 : 46. Syntype workers, SUDAN: Imatong Mts, E. slope ca 4100 ft, 31.vii.1939 (N. A. Weber), (MCZ Cambridge) [examined]. Syn. n. DIAGNOSIS OF WORKER. Densely shagreened with scattered larger punctures. Without standing hairs. Petiole in dorsal view longer than broad. Lamelliform margin of clypeus broad and often opaque. Outer margin of mandible straight or concave in part of its length. FURTHER DESCRIPTION. Worker. TL 6:8-7:6, HL 1-40-1-62, HW 1:24-1°44, CI 88-90, SL 1-36-1.64, SI 107-114, PW 092-108, PH 0:80-0.92, PL 0-66-0-72, LPI 121-127, DPW 0:60-0:72, DPI 90-97 (6 measured). Sides of head convex, the head not strongly broadened in front of the eyes, the sides more nearly parallel than in other members of the group. Eyes large, maximum diameter ca 0-32— 0-36, greater than the maximum width of the scape. Fringing lamella of anterior clypeal margin broad and broadly and evenly rounded, usually opaque or semi-opaque. Mandibles curved basally, the outer margins convex for some distance beyond the basal curve, but begin- ning at about the midlength the outer margin becoming flat or more usually concave. Mandibles shorter and stouter than in other members of the group. Node of petiole in dorsal view longer than broad. Body densely shagreened as in the other species but the larger punctures are more sparse and usually less distinct than in mavillosa or falcigera. Standing hairs present only on the clypeus and gastral apex. Notes on the separation of this species from its closest relatives are given under falcigera. The range of pavesii does not appear to overlap that of maxillosa in the Ethiopian Region. All specimens of the former examined have come from the countries of east and north-east Africa whilst those of the latter originated in the central and southern parts of the continent. Like other members of this group pavesiz is a tramp species and in this study I have examined specimens from Saipan, though whether this is Saipan in the Marianas or Saipan, Palau Is. is not clear. 256 B. BOLTON MATERIAL EXAMINED. SoMALIA: Dolo (Vatova). KENyA: Miongave, nr Mombasa (L. F. Brown); Ngong Forest Reserve (G. Arnold?). TANZANIA: Morogoro (A. Loveridge). CAm- EROUN: no loc. (G. Terron). Guam: Ladrone Is. (?). MARIANA or PALAU Is.?: Saipan (R. K. Enders). THE STUHLMANNI-Group (Text-figs 4-8, 19-21, 32) Mandibles elongate, linear, usually increasing in width from base to apex, not closing against theclypeus. Eyes large or very large, their maximum diameter much greater than the maximum width of the scape. Second funicular segment considerably longer than the third, sometimes twice as long. Head broader in front than behind. Clypeus with a prominent median lobe. General shape of petiole as shown in Text-figs 19, 32. Metanotal groove impressed but without cross-ribs or sculpture. Gaster not or only very weakly constricted between the first and second segments. Of the eleven species of this group seven are large or very large. Of these, five are distributed throughout eastern and southern Africa, with one also occurring on the Comoro Islands, and two are of central African origin. The four smaller species are known from West and Central Africa, with two in each area. The group falls into two complexes of species, depending on the shape of the basal portion of the mandible. In camerunensis and its allies (nebra, sterops, vindicis and zapyxis) the mandible does not pass through a curve basally before its articulation (Text-fig. 5), and the blades of the long mandibles are usually approximately straight, without feebly convex outer margins. These five species are all of West or Central African origin. The remaining species allied to stuhlmanni (erythraea, letothorax, schwabi, regis and titan) are all from East or South Africa except the last, which comes from Cameroun. They have the base of the mandible curved before its articulation (Text-fig. 6), although this is indistinct in some populations of schwabi. The outer margin of the blade is usually slightly convex throughout its length in full-face view. Leptogenys camerunensis Stitz (Text-figs 5, 7, 32) Leptogenys camerunenis Stitz, 1910: 130. Syntype workers, CAMEROUN: Barombistation (Preuss) (NM, Basle) [examined]. Leptogenys stuhlmanni race camerunensis var. angusticeps Forel, 1916 : 399. Holotype worker, ZAIRE, St Gabriel (Kohl) (MHN, Geneva) [examined]. [Name not available.] Leptogenys stuhlmannai subsp. camerunensis var. opalescens Wheeler, 1922 : 94. Syntype workers. ZAIRE: Akenge (Lang & Chapin) (MCZ, Cambridge) [examined]. [Name not available. ] DIAGNOSIS OF WORKER. Alitrunk without sculpture dorsally and only with cross-ribbing at meso-metapleural suture laterally. With blue or violet opalescence. Large species with very long antennal scapes, HL > 1°80, SI > 230. FURTHER DESCRIPTION. Worker. TL 1t-o—-12-0, HL 1-g0-2:02, HW 1:24—-1°32, CI 65-66, REVISION OF GENUS LEPTOGENYS 257 SL 2-92-3°38, SI 235-256, PW 1:24-1-30, PH 1:°12—1-20, PL 1:06—1-12, LPI 105-107, DPW 0*72-0'74, DPI 64-70 (3 measured). Head markedly broader in front than behind, the maximum width anteriorly in syntypes > 1-60, width across occipital ridge < 0-90. Eyes very large, maximum diameter ca 0-60, much greater than the maximum width of the scape. Inner margins of eyes with a circumocular impression. Mandibles long, more or less straight, gradually increasing in width apically and with a single preapical tooth. Mandibular blades not closing tightly against the clypeus, not curved basally before their articulation. Second funicular segment very long, respective lengths of segments I-3 Ca 0°50, I‘I0, 0:56. Metanotal groove weakly impressed, not cross-ribbed. Gaster not constricted between first and second segments. Entire body smooth and shining, unsculptured apart from small pits from which hairs arise and some cross-ribs at the meso- metapleural suture. Erect hairs present on all dorsal surfaces. Colour red-brown or brown, the appendages lighter, and the head and alitrunk with weak blue or violet opalescence or reflections, which may be patchy. One of the larger species of the group, camerunensis is distinguished primarily by its size and lack of sculpturation. The mandibles have a virtually straight outer margin and the blades are not curved outwards along their length. In this respect camerunensis shows affinities with zapyxis and sterops. MATERIAL EXAMINED. ZAIRE: Wombali (P. Vanderyst). CAMEROUN: no loc. (G. Terron). Leptogenys erythraea Emery stat. n. Leptogenys stuhlmanni subsp. erylhraea [sic] Emery, 1902 : 33. Syntype workers, ETHIOPIA: Eritrea (Belli) MCSN, Genoa)., Leptogenys stuhlmanni subsp. erythraea Emery; Forel, 1907a:131 et auctt. [Justified emendation. ] DIAGNOSIS OF WORKER. Metapleuron sculptured. Eyes large, maximum diameter 0-54 or more. Antennal scapes long, SI > 200. Full adult colour brown. Large species, HL > 1-80. FURTHER DESCRIPTION. Worker. TL 9:7-10°8, HL 1:94-2:00, HW 1:28-1-34, CI 65-67, SL 2:72-2:80, SI 209-214, PW 1-20-1:24, PH 1:14, PL 0-94, LPI 121, DPW 0-68-0-70, DPI 73- 74 (3 measured). Head broader in front than behind, broadest across the clypeus. Eyes large, maximum diameter 0-54—0'56, much greater than the maximum width of the scape; the eyes breaking the outline of the sides of the head in full-face view. Clypeus with a prominent median lobe. Mandibles elongate and narrow, slightly curved along their length and passing through a more marked curve basally before their articulation. Funicular segments 1-3 with respective lengths Ca 0°42—0°44, 0°84-0°86, 0°52-0'54. Gaster not constricted between first and second segments. Elongate standing hairs present upon all surfaces and on appendages. Head, pronotum and mesonotum unsculptured apart from minute pits from which hairs arise. Propodeal dorsum with feeble, nearly effaced sculpturation, with smooth areas. Meso- and metapleurae striate, strongest on metapleuron and extending onto the lower portions of the sides of the propodeum. Gaster unsculptured, smooth and shining. Full adult colour brown. I have not seen the types of this species but have examined a series compared with type by Prof. W. L. Brown. These indicate that erythraea is a distinct species and not a subspecies of stuhlmanni. The male has been described by Forel (19072). MATERIAL EXAMINED. EtuiopiA: Eritrea, Ghinda (Staudinger). 258 B. BOLTON Leptogenys leiothorax Prins stat. n. Leptogenys stuhlmanni subsp. leiothorax Prins, 1965 : 154, fig. 1. Holotype and paratype workers, MOZAMBIQUE: Dondo 1I0.xii.1960 (G. Arnold) (SAM, Cape Town) [examined]. DIAGNOSIS OF WORKER. Dorsal alitrunk completely unsculptured. With blue or violet opalescence. Smaller species with shorter antennal scapes, HL < 1°75, SI < 220. FURTHER DESCRIPTION. Worker. TL 8-3-9:1, HL 1-62-1-66, HW I-10—1°14, CI 67-60, SL 2-34-2°42, SI 212-214, PW 1:04-1:08, PH 0:96-0:98, PL 0-88-0-90, LPI 109-111, DPW 0°54—0'58, DPI 61-64 (3 measured). Head broader in front than behind, broader across the clypeus from corner to corner than across the eyes; sides of head convex. Eyes large, maximum diameter ca 0-44—0°50, breaking the outline of the sides of the head in full-face view. Clypeus with a prominent median lobe. Mandibles elongate, in full-face view weakly curved along their length, slightly more strongly curved basally before their articulations. Respective lengths of funicular segments 1-3 ca 0:32~ 0*40, 0*74-0°78, 0°42-0'46. Metanotal groove impressed but not cross-ribbed; gaster only extremely weakly constricted between first and second segments. All surfaces of body and appendages with numerous standing hairs. Dorsal surfaces of head and body completely unsculptured, smooth and shining except for small pits from which hairs arise. Propodeal declivity with a few transverse rugae. Sides of alitrunk unsculptured apart from cross-ribbing at the meso-metapleural suture and some rugosity around the area of the metapleural glands. Full adult colour deep red-brown with patchy bluish or violet opalescence. At first sight this species is reminiscent of a small version of camerunensis as it is similarly sculptured and coloured, but its true affinities lie with schwabi which, apart from its lack of sculpturation, letothorax closely resembles. In general shape and build and also in dimensions leiothorax is very close to schwabi but the lack of sculpture on the propodeum in the former will separate them. Also, in leiothorax the width of the head across the clypeus is greater than the width across the eyes, whereas in schwabi the two widths are approximately equal. Leptogenys nebra sp. n. DIAGNOSIS OF WORKER. Propodeum sculptured. Antennal scapes relatively long, SI > 200; eyes small, maximum diameter ca 0:36. FURTHER DESCRIPTION. Holotype worker. TL 9:1, HL 1°84, HW 1:20, CI 65, SL 2-60, SI 217, PW 1-06, PH 0-98, PL 0-86, LPI 114, DPW 0:62, DPI 72. Sides of head convex, broadest in front, distinctly broader across the clypeus from corner to corner than across the eyes. Eyes quite small for a member of this group, maximum diameter ca 0°36, but still distinctly greater than the maximum width of the scape. Clypeus with a prominent median lobe. Mandibles elongate, more or less straight in full-face view, increasing in width from base to apex and not passing through a curve basally before their articulations. Respective lengths of funicular segments 1-3 ca 0°32, 0:70, 0°48. Metanotal groove impressed but without cross-ribs. Gaster not constricted between first and second segments. All surfaces of head, body and appendages with numerous long, standing hairs, the longest of which on the scape are longer than the maximum scapal width. Dorsum of head, pronotum and gaster smooth and shining, unsculptured apart from hair-pits. Mesonotum and propodeal dorsum tranversely rugose, the latter more coarsely than the former. Mesopleuron with some rugae above but the median portion tending to be smooth. Metapleuron and sides of propodeum rugose. Colour black, the appendages brown. Holotype worker, CAMEROUN: Ototomo, 17.vi.1969, JLA (G. Terron) (BMHN). This relatively small-eyed species is related to sterops and zapyxis but the combi- REVISION OF GENUS LEPTOGENYS 259 nation of characters noted above, plus the lack of a convex lamella on the inner margin of the mandible will separate nebra from all other species of the group. Leptogenys regis sp. n. (Text-fig. 8) DIAGNOSIS OF WORKER. Propodeum and pronotum sculptured. Eyes not breaking outline of sides of head in full-face view. Large species with relatively short antennal scapes, HL > 2-00, st < 170. FURTHER DESCRIPTION. Holotype worker. TL 11-2, HL 2:24, HW 1:66, CI 74, SL 2°54, SI 153, PW 1-28, PH 1-14, PL 1-02, LPI 112, DPW 0-66, DPI 64. Head much broader in front than behind, the width of the head across the eyes noticeably less than the width across the clypeus from corner to corner. Eyes large, maximum diameter 0-50 but not so strongly convex or prominent as in other members of the group so that the eyes in full-face view fail to break the outline of the sides of the head. Clypeus with a prominent median lobe. Mandibles elongate, feebly curved along their length and passing through a curve basally before their articulations. Lengths of funicular segments 1-3 ca 0-40, 0°76, 0°46. Dorsal surfaces of body with numerous short, standing hairs; hairs on the antennal scapes very short, scarcely longer than the underlying pubescence, without long hairs such as are found in other species of the group which are almost or quite as long as the scape is broad. Pits from which hairs arise conspicuous upon the cephalic dorsum. Pronotum, mesonotum and propodeum sculptured, the last much more strongly than the rest. Sides of alitrunk sculptured, with rugae on the pleurae. Gaster smooth and shining except for pits from which hairs arise. Paratype worker. As holotype but slightly larger, TL 11-4, HL 2:24, HW 1-64, CI 73, SL 2-62, SI 159, PW 1-28, PH 1:24, PL 1-12, LPI 110, DPW 0-68, DPI 61. Eyes ca 0-50 at maximum; funicular segments I—3 ca 0-42, 0-80, 0°50 respectively. Holotype worker, Kenya: Shimba Forest Reserve, vi.1957 (N. L. H. Krauss) (BMNH). Paratype worker, TANZANIA: Dar-es-Salaam, College Campus, 1969 (Ilse Walker) (MCZ, Cambridge). This large species and the related titan are easily differentiated from other members of the group by the failure of the eyes to break the outline of the sides of the head in full-face view and by the lack of long, standing hairs on the antennal scapes. In all other large species of the group the scape is equipped with numerous long hairs, the longest of which are at least equal to or more usually greater than the maximum width of the scape, whereas in regis and titan the hairs which are present are short, often not much longer than the scapal pubescence. The two species, regis and titan, are separated by characters of size and differences of sculpturation, as the latter is much larger than the former and has the pronotum unsculptured. Leptogenys schwabi Forel Leptogenys schwabi Forel, 1913¢ : 208. Holotype worker, RHODESIA: Bulawayo (G. Arnold) (MHN, Geneva) [examined]. 260 B. BOLTON DIAGNOSIS OF WORKER. Propodeum sculptured. Maximum diameter of eye < 0-50, the eyes breaking the outline of the sides of the head in full-face view. Antennal scapes long, SI > 200. Width of head across eyes the same as or only slightly less than the width across the clypeus from corner to corner. FURTHER DESCRIPTION. Worker. TL 8:+4-9°8, HL 1:60-1:82, HW 1-08-1-22, CI 64-68, SL 2:28-2:54, SI 203-211, PW 1:04-1-14, PH 0-92-1-10, PL 0-88—1-00, LPI 104-113, DPW 0:54-0'64, DPI 62-64 (12 measured). Head broader in front than behind. In front of the eyes the sides of the head slightly convex so that the width of the head across the clypeus from corner to corner anteriorly is the same as or only slightly more than the width across the centre of the eyes. Eyes large and convex but not very large as in some members of the group, maximum diameter ca 0-44-0'48. Clypeus with a prominent median lobe. Mandibles elongate, in most specimens feebly curved along their length and passing basally through a curve before their articulations. Funicular segments I-3 Ca 0:34-0:40, 0°64—-76, 0:42-0:46 respectively. All surfaces of body and appendages with numerous standing hairs. Dorsum of head smooth and shining, unsculptured apart from small pits from which hairs arise. Pronotal dorsum similar but very rarely with faint traces of sculpturation on the extreme posterior portion of the sclerite. Propodeal dorsum with transverse rugae. Pleurae and sides of propodeum sculptured. Gaster unsculptured apart from hair-pits. In the past this species has been confused with stuhlmanni. The misidentification of specimens of schwabi as stuhlmanni and the consequent misapplication of the latter name is common in collections. In the BMNH collection, for example, series which were all schwabi had been labelled as such and also as stuhlmanni. Prins (1965) reduced schwabi to a variety of stuhlmanni, basing his argument on specimens collected by Arnold and Grobler in Rhodesia and South Africa respectively. All these specimens were determined by Arnold and were in fact all schwabi (stuhlmannt remains unknown in Rhodesia and South Africa). Prins’s argument that he could find very little difference between the two ‘species’ in these series was therefore correct as all were schwabi but some of them had been misidentified as stwhlmannt. The cause of this confusion appears to stem from Arnold’s (1915) record of stwhl- mannt from Durban. I have examined this series and they are in fact schwabi, though slightly larger than is usual in this species. The two are separable on a number of characters, many of which were in fact noted in the original descriptions of Mayr and Forel but subsequently not used. L. stuhlmanni is a larger, more stockily built species than schwabi, with relatively shorter antennal scapes (compare measurements). The head is broader in front in stuhlmanni and the eyes are less prominent, so that the width of the head across the eyes is noticeably less than the width across the clypeus from corner to corner. The pronotal dorsum is completely sculptured in stuhlmanni whereas in schwabi it is usually smooth, although in some specimens faint traces of sculpture are present posteromedially. Pits from which hairs arise on the cephalic dorsum are small and inconspicuous in schwabt but larger and more distinct in stuhlmannt. MATERIAL EXAMINED. RuopEsIA: Bulawayo (G. Arnold); Chirinda Forest (G. Arnold); Umtali (G. Arnold). SoutH Arrica: Natal, Umhlanga (G. Arnold?); Natal, Durban (G. Arnold); Durban (C. B. Cooper); Zululand, Eshowe (R. E. Turner). REVISION OF GENUS LEPTOGENYS 261 Leptogenys sterops sp. n. (Text-fig. 20) DIAGNOSIS OF WORKER. Smaller species, HL < 1:60. Propodeum and metapleurae sculptured. Mandibles without a lamelliform extension of the inner margin. FURTHER DESCRIPTION. Holotype worker. TL 8-2, HL 1-56, HW 1:06, CI 68, SL 1-98 SI 183, PW 0-96, PH 0-86, PL 0-72, LPI 119, DPW 0:52, DPI 72. Head broader in front than behind but not strongly broadened in front of the eyes; the sides feebly convex, the occipital border marginate. Ocular diameter 0-40 at maximum, greater than the maximum width of the scape. Clypeus with a prominent median lobe. Mandibles elongate but not linear, increasing in width from base to the level of the single preapical tooth then narrowing again to the apex; the inner margin feebly convex basad of the preapical tooth. Mandibular blades not passing through a curve basally before their articulations. Second funicular segment very long, the respective lengths of segments 1-3 ca 0-30, 0°52, 0-36. Alitrunk elongate; metanotal groove not cross-ribbed. Node of petiole in profile high and narrow, the anterior and posterior faces both feebly convex and converging dorsally, the anterior and dorsal faces forming a single convex curve. Gaster very weakly constricted between the first and second segments. Standing hairs numerous on all surfaces. Head, pronotum, petiole and gaster unsculptured apart from minute pits from which hairs arise. Propodeal dorsum rugose, predominantly transversely so. Metapleuron and sides of propodeum regularly rugose. Mesopleuron with rugae developed only in the upper and extreme lower portions, the greater part of the sclerite between these areas with the sculpture wholly or partially effaced and with smooth, shining areas. Colour principally dark brown, the appendages lighter, the head and propodeum darker, black in places. Paratype workers. TL 8-0-8-2, HL 1°54-1-56, HW 1:02-1:06, CI 66-69, SL 1:98-2:00, SI 183-194, PW 0-94-0'98, PH ca 0-86 PL 0:68-0:72, LPI 119-126, DPW 0:52-0:58, DPI 72-85 (5 measured). Ocular diameter ca 0-40; length of funicular segments 1-3 ca 0-28—0-30, 0°48— 0°52, 0°34-0°36. As holotype but in one specimen with the mesopleuron almost covered with very fine striation. Holotype worker, Ivory Coast: Banco Forest, nr Abidjan, 13. i. 1963, no. A72 (W. L. Brown) (MCZ, Cambridge). Paratype workers. Ivory Coast: 5 with same data as holotype; 1 worker, O.R.S.T.O.M. Res. Sta., 17km W. of Abidjan, 7.i.1963, no. Art (W. L. Brown) (BMNH; MCZ, Cambridge). This species, the smallest yet discovered in the stuhlmanni-group, is most closely related to zapyxis, also from West Africa. The two are easily distinguished as in sterops the mandible increases in width from the base towards the apex and is widest at the preapical tooth. Also, the inner margin of the blade lacks a lamelliform extension and is only feebly convex at most. In zapyxis on the other hand the inner margin of each mandibular blade is equipped wth a semi-translucent, lamella- like extension which is convex in outline, and the mandible usually attains its maximum width at a point proximal to the preapical tooth. Leptogenys stuhlmanni Mayr (Text-fig. 6) Leptogenys (Lobopelta) stuhlmanni Mayr, 1893 : 198. Syntype workers, MozAMBIQUE: Quilimane (Stuhlmann) (location of types not known). 262 B. BOLTON Leptogenys comorensis Forel, 1907): 76. Syntype workers, Comoro IsLANDs: Mohali (Voelizkow) (NM, Basle; MCZ, Cambridge) [examined]. Syn. n. DIAGNOSIS OF WORKER. Propodeum sculptured. Large species with relatively short antennal scapes, HL > 1-75, SI < 200. Pronotal dorsum feebly sculptured. Eyes breaking outline of sides of head in full-face view. FURTHER DESCRIPTION. Worker. TL 9:6-10-4, HL 1°82-2:00, HW 13°2-14°6, CI 68-74, SL 2-+32-2:70, SI 180-194, PW 1-16-1:26, PH 1-10-1-18, PL 1-02-1:06, LPI 104-115, DPW 0:62-0:70, DPI 61-68 (6 measured). Head broader in front than behind, distinctly broadened in front of the eyes. Eyes large. their maximum diameter 0-50—0-54, much greater than the maximum width of the scape. In full-face view the eyes breaking the outline of the sides of the head. Clypeus with a prominent median lobe. Mandibles long, usually very slightly curved along their length, characteristically passing through a curve basally before their articulations. Respective lengths of funicular segments I-3 ca 0:40-0:42, 0:74-0'76, 0:46-0'48. Metanotal groove not cross-ribbed. Gaster not constricted between first and second segments. All surfaces of body and appendages with numerous standing hairs. Pits on dorsum of head from which hairs arise usually large and conspicuous. Dorsum of pronotum and sometimes also the sides weakly sculptured. Dorsum and sides of propodeum and the pleurae rugose. Dorsum of petiole usually feebly sculptured. Gaster smooth and shining, unsculptured apart from hair- pits. One of the three large, black species of the group which inhabit eastern and southern Africa and its offshore islands, stuhlmanni is distinguished from regis by size, the convexity of the eyes, which break the outline of the sides in stuhlmanni but not in regis, and by the presence of numerous long hairs upon the antennal scapes which are absent in vegis. The other closely related species, schwabi, tends to be smaller and less stockily built than stuhlmanni, with relatively longer antennal scapes and finer sculpturation. On the dorsum of the head the pits from which hairs arise are distinct and quite large in stwhlmanni and the pronotal dorsum is sculptured, whereas in schwabi the hair-pits are small and inconspicuous and the pronotum is usually unsculptured, although some individuals may have slight sculpturation. Finally, in schwabi the width of the head across the eyes is the same as or only slightly less than the width across the clypeus from corner to corner, whereas in stuhlmanni the corner to corner clypeal width is distinctly greater than the width across the eyes as the head is more strongly broadened in front in this species. There seems to be little doubt that stuhlmanni and schwabi were confused by earlier authors (see under schwabi) and records of distribution of the species published to the present should be used with some caution. I have not been able to find the types of stuhlmanni but Dr M. Fischer (NM, Vienna) has lent me two workers collected by Stuhlmann in Mozambique and named by Mayr himself, which are taken as the basis for my interpretation of this species. MATERIAL EXAMINED. SUDAN: Imatong Mts (N. A. Weber). KENya: Nakuru (E. S. Ross & R. E. Leech); Mombasa (N. A. Weber). MOZAMBIQUE: (Stuhlmann, ex coll. G. Mayr). REVISION OF GENUS LEPTOGENYS 263 Leptogenys titan sp. n. DIAGNOSIS OF WORKER. Propodeum unsculptured. Eyes not breaking outline of sides of head in full-face view. Very large species with relatively short antennal scapes, HL > 2:10, SI < 170. FURTHER DESCRIPTION. Holotype worker. TL 13:6, HL 2:30, HW 2:02, CI 88, SL 3:24, SI 160, PW 1-62, PH 1°36, PL 1°18, LPI 115, DPW 0:76, DPI 64. Sides of head strongly divergent anteriorly, the head much broader across the clypeus than across the eyes. Eyes large, maximum diameter ca 0-66, distinctly failing to break the outline of the sides of the head in full-face view. Maximum diameter of the eyes considerably greater than the maximum width of the scapes. Clypeus with a prominent median lobe. Mandibles very long, only slightly broadened apically, the blades passing though a distinct curve basally before their articulations. Respective lengths of funicular segments I-3 ca 0:50, I-02, 0:66. Metanotal groove without cross-ribs. Gaster weakly impressed between first and second segments. Head, body and appendages with numerous, relatively short, standing hairs. On the antennal scapes in dorsal view no hairs are present which exceed the maximum width of the scape. Dorsum of head, pronotum, propodeum and gaster unsculptured apart from hair-pits. Mesonotum with a number of very faint transverse striae, particularly upon the median portion of the sclerite. Sides of alitrunk unsculptured except for cross-ribbing at the meso-metapleural suture and a few rugae on the bulla of the metapleural glands. Colour black, appendages dark brown. Holotype worker, CAMEROUN: Ngoa-Ekelé 4.i.1969 (G. Terron) (BMNH). L. titan is most closely related to regis of east Africa but is larger than that species and lacks propodeal sculpturation. These two species are the only members of the group in which the eyes fail to break the outline of the sides of the head in full-face view. Leptogenys vindicis sp. n. DIAGNOSIS OF WORKER. Head and dorsal alitrunk sculptured. Antennal scapes relatively long, SI > 200; smaller species, HL < 1°75. FURTHER DESCRIPTION. Holotype worker. TL 9:0, HL 1-64, HW 1-14, CI 60, SL 2-38, SI 209, PW 1:12, PH 0-96, PL 0-80, LPI 120, DPW 0:68, DPI 85. Head broader in front than behind but the width of the head across the clypeus only very slightly more (ca 0-02) than the width across the eyes in full-face view. Eyes large, maximum diameter ca 0-50, much greater than the maximum width of the scape. Clypeus with a prominent median lobe. Mandibles long, slightly broader apically than basally, passing through a very feeble curve near the base. Funicular segments I-3 ca 0:34, 0°68, 0:38 respectively. Gaster feebly constricted between first and second segments. All surfaces of head, body and appendages with numerous standings hairs. All dorsal surfaces of head and body finely sculptured except for the gaster, metanotal groove and area immediately behind the prome- sonotal suture, which are smooth and shining. Sculpture takes the form of fine, disorganised rugulation with a few larger rugae, so that the surfaces appear matt or only dully shining. Sides of alitrunk with more distinctly separated rugae and with some shining areas on the pronotum and pleurae. Colour a very deep brown, almost black, the gaster lighter brown, the appendages mid-brown and shining. Holotype worker, CAMEROUN: Mt Nkolodon, JLA, 19.v.1969, 900-1000 m (G. Terron) (BMNH). 264 B. BOLTON Leptogenys zapyxis sp. n. (Text-figs 4, 19, 21) DIAGNOSIS OF WORKER. Propodeum and metapleuron sculptured. Inner margin of mandible extended by a convex, semi-translucent lamella. FURTHER DESCRIPTION. Holotype worker. TL 9:0, HL 1-78, HW 1:16, CI 65, SL 2-40, SI 207, PW 1:08, PH 0-92, PL 0:82, LPI 112, DPW 0:62, DPI 76. Head broader in front than behind, the sides weakly convex and diverging anteriorly. Occipital border marginate. Eyes with maximum diameter ca 0-42, much greater than the maximum width of the scape. Clypeus with a prominent, truncated median lobe. Mandibles elongate, their outer margins straight to feebly concave, the inner margins convex, with a semi-translucent, lamellate extension forming the true inner margin and imparting the convexity to the border. Second funicular segment elongate, respective lengths of segments 1-3 ca 0°36, 0:64, 0°42. Metanotal groove not cross-ribbed. Petiole in profile with the anterior and posterior faces convex and sloping towards one another dorsally, the anterior and dorsal faces united ina continuous curve. Gaster very feebly constricted between first and second segments, All surfaces with numerous standing hairs. Head, pronotum, petiole and gaster unsculptured apart from hair-pits. Dorsum and sides of propodeum and metapleuron rugose. Mesopleuron with some effaced sculpturation but with extensive smooth areas. Black, the appendages lighter. Paratype workers. TL 8-1-9:0, HL 1:62-1:80, HW 1:04-1:18, CI 64-66, SL 2:18-2:44, SI 200-209, PW 0-96—-1:08, PH 0-88—-0-92, PL 0-76—0-84, LPI 109-118, DPW 0:58-0:64, DPI 73-77 (7 measured). Ocular diameter at maximum ca 0-38—0-44; respective lengths of funicular segments I-3 Ca 0°30—0°36, 0:60-0:64, 0:38-0-42. As holotype but colour varying from dark brown to black. Holotype worker, GHANA: Mt Atewa, 22.x.1968 (C. A. Collingwood) (BMNH). Paratypes. GHANA: I worker, same data as holotype; 6 workers, Tafo, 26.vi. 1970, in wet-rotten log (B. Bolton); 3 workers, Tafo 3.viii.1966, ant ecology sample 150C (D. Leston); 2 workers, Tafo 2.viii.1966, ant ecology sample 144A (D. Leston) (BMNH; MCZ, Cambridge; NM, Basle; MNHN, Paris). THE SULCINODA-GRovuP (Text-figs 9-14, 26, 27) Mandibles variously developed, either short and closing tightly against the clypeus or linear and curved, enclosing a space between themselves and the clypeus when closed. Eyes medium or large, but always with their greatest diameter in full-face view greater than the maximum width of the scape. Second funicular segment equal to or slightly longer than the third. Head usually broader in front than behind; more markedly so in some species than in others. Sculpture variable but never with the entire dorsal head and alitrunk rugose, nor shagreened. Gaster constricted between first and second segments. Metanotal groove cross-ribbed or otherwise sculptured. The sulcinoda-group includes 13 species which fall roughly into two complexes depending upon whether the mandibles are capable of closing tightly against the clypeus or not. In general there are two criteria which determine whether full closure is possible, the first being the length of the mandible itself and the second the size of the median lobe of the clypeus. These seem interdependent and modifi- cations occur simultaneously, for as the mandible lengthens and narrows the clypeal lobe becomes shorter and narrower. REVISION OF GENUS LEPTOGENYS 265 Species with relatively shorter, broader mandibles and longer, broader clypeal lobes are connected to those with longer, narrower mandibles and shorter, narrower clypeal lobes by intermediates such as princeps, and illustrate and support the con- tention that long-mandibulate forms have arisen from short-mandibulate stock several times within the genus. All members of the group are small to medium-sized species and the majority appear to be confined to rather densely forested areas, especially in West and Central Africa. Leptogenys bellii Emery Leptogenys belliti Emery, 1901 : 59. Syntype workers, Etniopia: Eritrea, Ghinda (Belli) (MCSN, Genoa). I have not been able to examine the types of this species nor have I seen any material which matches the original description, but the description is good enough to recognise bellii as a member of the sulcinoda-group. The following description is modified from the original. Worker. T6-0. Head longer than broad, narrower behind than in front, with the occipital margin straight or feebly concave. Eyes large. Median lobe of clypeus short, blunt (trun- cated?) apically. Mandibles weakly curved, narrow, slightly broadening apically where they terminate in a point. Scapes surpassing occipital margin by less than twice their diameter. Node of petiole a little longer than broad, almost as high as long; in profile the anterior and posterior faces vertical. “Anterodorsal angle rounded, posterodorsal angle acute. In the middle of the dorsum is a narrow, shining zone, a little elevated but not cariniform. Posterior face of petiole bordered by a sharp corner, a little projecting in the middle of the superior margin where it forms an obtuse tubercle. Vertex smooth, with small piligerous punctures which become larger and more numerous towards the sides and may even be confluent behind the eyes. On pro- and mesonotum the punctures are separate on the dorsum, confluent on the sides and larger than those of the head; larger and confluent on sides of propodeum. Meso- and metapleurae partly sculptured. Sides of petiole with confluent punctures. Gaster shining, with minute piligerous punctures. Standing hairs present on dorsal surfaces. Black, shining, the appendages reddish. The description gives the impression of an average-sized member of the sulcinoda- group most probably related to elegans and ferrarit but separated from them by details of petiolar structure and sculpturation. Leptogenys elegans sp. n. (Text-fig. 9) DIAGNOSIS OF WORKER. Upper halves of sides of petiole not sulcate. Dorsum of head, alitrunk and first gastral tergite unsculptured apart from large hair-pits. FURTHER DESCRIPTION. Holotype worker. TL 4:8, HL 1-00, HW 0-72, CI 72, SL o-go, SI 125, PW 0°62, PH 0:62, PL 0:46, LPI 135, DPW 0°48, DPI 104. Head slightly wider in front of the eyes than behind, the sides feebly convex. Eyes with maximum diameter ca 0-24, greater than the maximum width of the scape in full-face view. Median lobe of clypeus prominent, broadest basally, rounded anteriorly and with convex sides. Mandibles elongate and narrow, not closing tightly against the clypeus, enclosing a large space when their apices are overlapping. Lengths of funicular segments I—3 respectively 266 B. BOLTON ca 0'16, 0:16, 0-14. Gaster strongly constricted between first and second segments. Dorsum of head with numerous shallowly incised punctures separated by smooth, shining interspaces. Sides and ventral surface of head unsculptured apart from scattered hair-pits. Dorsal alitrunk unsculptured apart from crossribbing in the metanotal groove and scattered, large, pit-like punctures from which hairs arise. Sides of alitrunk also with pit-like punctures and the metapleuron and extreme lower region of the mesopleuron with longitudinal rugulae. Upper halves of sides of petiole with some rugulation. First and second gastral tergites with scattered but strongly marked large hair-pits, otherwise unsculptured. All surfaces of body and append- ages with standing hairs. Colour black, shining; the legs, antennae and mandibles brown. Paratype workers. As holotype but with size range as follows: TL 4+5-4:9, HL 0:96-1:02, HW 0:68-0:72, CI 70-72, SL 0:86-0:94, SI 125-138, PW 0-58—0-62, PH 0:56-0:62, PL 0:44—0-48, LPI 127-135, DPW 0-42-0°48, DPI 95-105 (8 measured). Ocular diameter 0-20—0-24, and size range of funicular segments 1-3 ca 0-16—0-18, 0-14-0°16, 0*12—0'14 respectively. Sculpturation of metapleuron and sides of propodeum variable. In most specimens it is as in the holotype but in some the rugulae are reduced or absent in places whilst in others the rugulae are more extensive and also occur on the propodeal sides. Holotype worker, NIGERIA: Gambari, 10.ix.1969, in rotten stump (B. Bolton) (BMNH). Paratypes. 2 workers with same data as holotype. Ivory Coast: 5 workers, 40 km W. of Abidjan, 14.1.1963, Ag7, Plantation Niecky (W. L. Brown). GHANA: 1 worker, Pankese Cocoa Station, 10.ix.1969 (C. A. Collingwood); 4 workers, Bunso, 7.vii.1969, pyrethrum knockdown B2/2 (D. Leston); 2 workers, Bunso, 30.vii. 1969, pyrethrum knockdown B8/1 (D. Leston); 1 worker, Tafo, 8.x.1966, ant ecology sample 318F (D. Leston); 1 worker, Asikaisu, 9.ix.1969, ant ecology sample AM7o (D. Leston) (BMHN; MCZ, Cambridge; NM, Basle). The nest from which the holotype was taken was situated in the stump of a rotten tree branch about 2 feet above ground level. This species appears to be subarboreal as a number of specimens were collected by Leston as they were running on tree trunks, and some were acquired by pyrethrum knockdown from cocoa trees. This species appears to be closely related to ferrariz, and notes on their separation are given under the latter. Dr G. Terron (UFC, Yaounde) has sent me specimens of this species from Byime- Assi, in Cameroun. Leptogenys excellens sp. n. DIAGNOSIS OF WORKERS. Sides of head below and behind eyes and lateral portions of ventral surface of head sculptured. Upper halves of sides of petiole not sulcate. FURTHER DESCRIPTION. Holotype worker. TL 5-0, HL 1-00, HW 0:72, CI 72, SL 0-92, SI 128, PW 0-66, PH 0:66, PL 0-44, LPI 150, DPW 0-46, DPI 104. Head broader in front than behind, the sides feebly convex, the occipital margin in full-face view extremely shallowly concave, almost straight. Eyes large, maximum diameter 0-24, their diameter greater than the maximum width of the scape. Inner and posterior margins of eye with a weak circumocular impression. Median lobe of clypeus short, triangular, narrowly rounded apically. Mandibles long and curved, not closing against the clypeus. Relative lengths of funicular segments 1-3 ca 0:16, 0°18, 0-16. Petiole in profile with the anterior face vertical, the posterior face feebly sinuate and higher than the anterior, so that the dorsum slopes upwards posteriorly. Anterodorsal angle broadly rounded, posterodorsal angle also rounded but noticeably more acute. Petiole in dorsal view slightly broader than long, scarcely narrowing anteriorly. Gaster constricted between first and second segment. All surfaces REVISION OF GENUS LEPTOGENYS 267 of body and appendages with numerous standing hairs. Dorsum of head with numerous coarse punctures, the spaces between them feebly reticulate. Sides of head behind and below the eyes with weak rugosity, the ventral surfaces of the head complexely sculptured on their lateral portions by a number of short, shallow indentations and numerous fine punctures. Dorsum of alitrunk with scattered large punctures, the posterior margins of many of which are indistinct on the pronotum. Meso- and metapleurae reticulate-punctate but each with a shining median area, more extensive on the former. Upper halves of sides of petiole in profile reticulate-rugose with many small punctures. Gaster unsculptured apart from _hair-pits. Black, shining, the gaster dark brown and polished, the appendages lighter brown; the antennal funiculae lighter in colour than the scapes. Paratype worker. As holotype, with dimensions as follows; TL 5-0, HL 0-98, HW 0-72, CI 73, SL 0-92, SI 128, PW 0-62, PH 0-64, PL 0-44, LPI 145, DPW 0-44, DPI 100. Diameter of eye 0-24, lengths of funicular segments 1-3 ca 0°16, 0-16, 0°14. Holotype worker, ZAIRE: Yangambi Reserve, 0949 (A. Raignier & J. van Boven) (MCZ, Cambridge). Paratype. I worker, same data as holotype (BMNH). Leptogenys ferrarii Forel Leptogenys ferrarit Forel, 1913¢ : 209. Syntype workers, RHODESIA: Bulawayo, 23.ii.1913 (G. Arnold) (MHN, Geneva) [examined]. Leptogenys fervarii st. dentatula Santschi, 1915 : 246. Holotype worker, ZAIRE: Katanga, Kataki (Gérard) (NM, Basle )[examined]. Syn. n. Leptogenys ferrarit race dentulata Santschi; Bernard, 1952 : 210. [Lapsus.] DIAGNOSIS OF WORKER. Upper halves of sides of petiole not longitudinally sulcate. Hairs on first gastral tergite not arising from large pits. FURTHER DESCRIPTION. Worker. TL 4:-1-4:3, HL 0-84-0-86, HW 0:64-0:66, CI 74-78, SL 0:80-0:82, SI 124-125, PW 0:54-0-60, PH 0-50-0-58, PL 0:36-0:40, LPI 142-147, DPW 0:40—0°46, DPI 105-115 (5 measured). Head only slightly broader in front of eyes than behind, the sides convex; the occipital margin in full-face view weakly concave. Eyes large, maximum diameter 0°18—-o'20, with a weak circumocular impression on the internal and posterior margins. Diameter of eye much greater than maximum width of scape. Median lobe of clypeus triangular, narrowed anteriorly, almost pointed. Mandibles elongate, not closing against the clypeus. Respective lengths of funicular segments I—3 Ca 0°16, 0-14, 0°12. Petiole in dorsal view broader than long, broadest posteriorly. Node narrow and high in profile, the dorsal surface sloping upwards posteriorly but only slightly so. Gaster impressed between first and second segments. Dorsum of head only with small pits from which hairs arise. Dorsal alitrunk with larger shallow punctures which are best defined on the pronotum. In some specimens they are absent from the propodeal dorsum and may be effaced on the pronotum so that the dorsal alitrunk is largely smooth and shining. Meso- and metapleurae with weak rugulation, sometimes also finely punctate, but always with extensive unsculptured areas. Gaster unsculptured, smooth and shining apart from minute hair-pits. Full adult colour dark brown, the appendages lighter. This species seems most closely related to elegans but the two are separated by the construction of the petiole, as reflected by their indices, and also by their sculptura- tion and colour, as noted in the key. Also, the sides of the head are less convex in elegans and the head broadens more markedly anteriorly. MATERIAL EXAMINED. RuOopEs!IA: Lonely Mine (H. Swale); Cecil Kop nr Umtali (W. L. Brown). 268 B. BOLTON Leptogenys longiceps Santschi Leptogenys longiceps Santschi, 1914a : 328, fig. 13. Holotype worker, SENEGAL: Dakar (F. Silvestri) (IE, Naples). DIAGNOSIS OF WORKER. Mesopleuron completely sculptured. Median clypeal lobe very short and bluntly rounded. FURTHER DESCRIPTION. Worker. TL 4:2-4:5, HL 0:88-0:96, HW 0:70-0:72, CI 74-80, SL 0-76-0°86, SI 108-120, PW 0-58—0-62, PH 0-54-0-60, PL 0-42-0:46, LPI 128-136, DPW 0*40-0°46, DPI 95-100 (7 measured). Head slightly broadened in front of eyes. Maximum diameter of eyes 0-18-0-22. Median lobe of clypeus developed as in trilobata (Text-fig. 11) but much shorter and narrower. Mandibles elongate, not closing against the clypeus. Length of funicular segments 1-3 ca 0-16, 0°16, 0-14 respectively. Head everywhere with scattered fine punctures dorsally, anteriorly also with a very fine superficial reticulation which extends from the posterior clypeal margin almost to the level of the posterior margin of the eye. Laterally and ventrally the head unsculptured apart from hair-pits. Dorsal alitrunk with numerous low, rounded rugulae overlaid by a dense reticulate-puncturation. Posterior portion of pronotal dorsum unsculptured, smooth and shining. Sides of pronotum very finely rugulose, mesopleuron, metapleuron and sides of propodeum more coarsely rugulose and also covered with a dense reticulate-puncturation. Upper halves of sides of petiole reticulate-punctate, this sculpture superimposed upon a rather coarser rugulation. Gaster smooth and shining apart from hair-pits. The shape of the median lobe of the clypeus in this species resembles that found in trilobata but in a much reduced condition. However, the marked differences in sculpture between the two easily serve to separate them. In terms of sculpturation the species most closely resembling Jongiceps is excellens, but in this latter species the sculpture of the cephalic dorsum is continued on the sides and ventre. MATERIAL EXAMINED. GHANA: Legon (D. Leston); Legon (Mkhize). Leptogenys mastax sp. n. (Text-figs 12, 27) DIAGNOSIS OF WORKER. Petiole node with upper halves of sides longitudinally sulcate. Dorsum of head reticulate-punctate. Pronotum strongly sculptured. HL > 1-00, SI > 145. FURTHER DESCRIPTION. Holotype worker. TL 4:9, HL 1:06, HW 0-66, CI 62, SL 1:02, SI 154, PW 0-62, PH 0:66, PL 0-46, LPI 143, DPW 0-44, DPI 95. Head elongate, narrow, slightly broadened in front, the sides more or less straight and the occipital margin in full-face view weakly concave. Maximum diameter of eyes 0-24, greater than the maximum width of the scape. Inner and posterior borders of eyes with a circumocular impression. Median clypeal lobe long, triangular and quite broad. Mandibles relatively short, capable of closing tightly against the clypeus. Antennal scapes relatively long (SI >140); lengths of funicular segments 1-3 respectively ca 0-18, 0-20, 0-16. Node of petiole in dorsal view broadest posteriorly, the upper halves of the sides converging strongly dorsally so that the actual dorsal surface of the node is a narrow strip. In profile the posterior face of the node feebly concave, the anterior feebly convex; the posterior face much higher than the anterior so that the dorsal surface slopes upwards posteriorly and is slightly convex. Gaster constricted between first and second segments. All surfaces of body and appendages with standing hairs. Dorsum of head reticulate-punctate, the sides behind the eyes with a fine, superficial reticulation, the ventral surface unsculptured. Pronotal dorsum rough, with dense punctulation. The REVISION OF GENUS LEPTOGENYS 269 extreme posterior portion of the pronotum smooth and shining and with finger-like projections leading anteriorly into the sculptured area. Remainder of dorsum densely punctulate and with a few fine rugulae. Sides of pronotum finely reticulate, the meso- and metapleurae with some fine rugulae and a few punctures but with extensive smooth areas, especially upon the former. Upper halves of sides of petiole strongly longitudinally sulcate, the lower halves shiny and unsculptured. Gaster with only minute hair-pits. Black, the appendages brown. Paratype workers. TUL 4:7-5:0, HL 1:04-1:10, HW 0-66-0-70, CI 62-64, SL 0:98-1:08, SI 148-156, PW 0-62-0-64, PH 0:62-0:66, PL 0-42-0-46, LPI 139-147, DPW 0-42-0-44, DPI 92-100 (7 measured), Ocular diameter 0-22—0-26; lengths of funicular segments 1-3 ca 0-16—0°18, 0'20—0°22, 0*16—0-:20.' Otherwise as holotype. Paratype females. Ergatoid; fitting description of worker but the petiole is modified and the gaster swollen. In the females examined the second gastral segment measured 0-92—0-96 high in profile and 0-86—0-92 wide in dorsal view. This is compared to measurements of 0-76-0-78 and 0-72-0-76 respectively in workers of the same series. Dimensions of the females are: HL 1:06-1:08, HW 0-68-0-70, CI 63-65, SL 0:96-1:02, SI 141-147, PW 0:62-0:64, PH 0:68-0:70, PL 0:40-0:42, LPI 162-171, DPW 0:48-0:50, DPI 119-125, ocular diameter 0:24-0:26 (4 measured). These compare well with the worker except that the scapes are relatively slightly shorter and the petiole is noticeably narrower in profile and broader in dorsal view. Ocelli are absent and the alitrunk is similar to that of the worker as flight sclerites are not developed. Holotype worker, GHANA: Mt Atewa, 12.vi. 1970, under rotten twig on path (B. Bolton) (BMNH). Paratypes. GHANA: 5 workers, Mt Atewa, ant ecology sample 138E, 29.vii.1966 (2 workers); ant ecology sample 139C, 29.vii.1966 (I worker); ant ecology sample I61E, 5.vill.1966 (I worker); ant ecology sample 164C, 5.viii.1966 (x worker) (D. Leston). Ivory Coast: 6 workers, 4 females, Banco Forest, Abidjan, 8.1.1963, no. A6 (W. L. Brown) (BMNH; MCZ, Cambridge; NM, Basle). It is interesting to note the number of females present in the paratype series from Ivory Coast. As this caste is very poorly known in the African species of Lepiogenys it is impossible to say whether this multigynous condition holds in any other species or is peculiar to members of the sul/cinoda-group or merely to this one species. L. mastax is a member of a small complex of species within the group centring on sulcinoda and including also nuserra and occidentalis. All share the characters of relatively short mandibles which close tightly against the clypeus and a strong sulcate sculpturation on the upper halves of the sides of the petiole. They are separated by the characters given in the key and their respective diagnoses. Leptogenys nuserra sp. n. DIAGNOSIS OF WORKER. Upper halves of sides of petiole sulcate. Dorsum of head densely punctate. Pronotum sculptured. Small species with relatively short scapes, HL < 1-00, SI < 140. FURTHER DESCRIPTION. Holotype worker. TL 4:0, HL 0-90, HW 0-60, CI 67, SL 0-78, SI 130, PW 0:52, PH 0°52, PL 0-34, LPI 150, DPW 0-38, DPI 111. Head not broadened in front of eyes, the sides feebly convex throughout their length. Maximum diameter of eye 0-20, in full-face view as long as the maximum width of the scape. Median lobe of clypeus triangular, broad and quite long. Mandibles relatively short, capable of closing tightly against the clypeus. Funicular segments 1-3 respectively 0-12, 0-14, 0-12. Petiole in profile high and narrow, the anterior and posterior faces virtually parallel but the 270 B. BOLTON latter longer than the former so that the dorsal surface slopes upwards posteriorly. In dorsal view the node broadest posteriorly, narrowing in front, and the upper halves of the sides strongly convergent dorsally so that the dorsal surface proper is very narrow. Gaster constricted between first and second segments. All surfaces and appendages with numerous standing hairs. Dorsum of head densely but shallowly punctate, the sides and ventral surfaces unsculptured. Pronotal dorsum densely but finely reticulate or shagreened, the extreme posterior portion unsculptured and with finger-like projections radiating forwards. Remainder of dorsal alitrunk with scattered punctures and some very fine shagreening. Metapleuron with some fine striation and punctures, the former extending slightly onto the mesopleuron; otherwise sides of alitrunk unsculptured. Upper halves of sides of petiole longitudinally sulcate. Gaster smooth apart from minute hair-pits. Black, the legs, mandibles and scapes brown, the funiculi yellow-brown. Paratypes workers. TL 3*5-4:1, HL 0:80-0:90, HW 0:56-0:62, CI 66-72, SL 0:66-0:78, SI 118-130, PW 0-46-0°54, PH 0-46-0°54, PL 0:30-0:36, LPI 150-159, DPW 0:32-0:38, DPI 100-112 (6 measured). Maximum diameter of eye 0:16-0:20; respective lengths of funicular segments I-3 Ca 0-I2—0°14, 0*14—0°16, O-I12—0'14. Paratype females. TL 3-6—-3-7, HL 0:86-0:88, HW 0:58-0:60, CI 66-70, SL 0-72, SI 120-124, PW 0:51-0:52, PH 0°52, PL 0-26, LPI 200, DPW 0-42, DPI 161 (2 measured). Ocular diameter 0:20. Ergatoid, answering the description of the worker but with the petiole differently constructed as can be seen in the measurements given. The node in profile is strongly compressed antero-posteriorly and in dorsal view is relatively very broad. Gaster swollen; in the two females studied the height of the second segment in profile was 0-72—-0-76 whilst its width in dorsal view was 0:74-0:76. In workers the height of this segment is 0:54—0-60 and the width 0°52—0°56. Holotype worker, GHANA: Mt Atewa, 29.vii.1966, ant ecology sample 141E (D. Leston) (BMHN). Paratypes. GHANA: 2 workers with same data as holotype; 1 female, Tafo, 22.vi. 1966, ant ecology sample 45 (D. Leston); 1 worker, Tafo, 18.x.1966, ant ecology sample 335F (D. Leston); 1 worker, Mt Atewa, 8.viii.1969, pyrethrum knockdown sample A13/6 (D. Leston); 1 worker, Sajimasi, 25.vii.1g69, on trunk (D. Leston). Ivory Coast: I worker, 1 female, Banco Forest, nr Abidjan, 8.1.1963, sample Arr (W. L. Brown); 5 workers, Banco Forest, nr Abidjan, i.1963, sample A (W. L. Brown) (BMNH; MCZ, Cambridge; NM, Basle.) This species is most closely related to mastax but is smaller and the head is less strongly sculptured. It is the smallest of the complex of species immediately related to sulcinoda in which the upper halves of the sides of the petiole are longitu- dinally sulcate. Leptogenys occidentalis Bernard stat. n. (Text-fig. 13) Leptogenys ferrarit subsp. sulcinodis Bernard, 1952 : 210, fig. 5D. Holotype worker, GUINEA: Ravin 1 du Mont To (st.B1 28) (Lamotte) (MNHN, Paris) [examined]. [Junior homonym of sulcinoda André, 1892 : 48.] Leptogenys (Lobopelta) nitida subsp. occidentalis Bernard, 1952 : 211, fig.5 F. Syntype workers, GuINEA: To Foret 21-2. Bz 41 ravin 1 (Lamotte) (MNHN, Paris) [examined]. [Synonym of the above and hence valid name.] Letogenys ferrarit subsp. bernardi Baroni Urbani, 1971 : 360. [Replacement name for sulcinodis Bernard.] Syn. n. REVISION OF GENUS LEPTOGENYS 271 DIAGNOSIS OF WORKER. Upper halves of sides of petiole longitudinally sulcate. Pronotum unsculptured apart from hair-pits. Occipital margin in full-face view concave. FURTHER DESCRIPTION. Worker. TL 4:5-4°6, HL 1:04-1:06, HW 0:72-0:76, CI 69-72, SL 0:94-0:98, SI 126-130, PW 0:62-0:66, PH 0:60-0:64, PL 0:40-0:42, LPI 150-155, DPW 0*42-0°44, DPI 100-105 (4 measured). Head slightly broader in front than behind, the occipital margin broadly concave in full-face view. Maximum diameter of eye ca 0:20-0:24. Median lobe of clypeus broad, narrowly rounded anteriorly. Mandibles short, capable of closing against the clypeus. Length of funicular segments 1-3 respectively ca 0-17, 0°18, 0:16. Node of petiole in dorsal view as broad as or slightly broader than long, much broader behind than in front. Sides of petiole converging dorsally so that the dorsal surface is quite narrow. Petiole high and narrow in profile, the anterior and posterior faces vertical and more or less parallel but the former shorter than the latter. Gaster constricted between first and second segments. Dorsum of head with numerous small, shallow punctures, the spaces between them smooth and shining. Sides and ventral surface of head and sides and dorsum of alitrunk unsculptured apart from hair-pits; propodeal declivity with transverse rugae. Upper halves of sides of petiole strongly longi- tudinally sulcate. Gaster unsculptured apart from minute hair-pits. Black, with appendages brown. The two Bernard names given above are straight synonyms despite the fact that he related one to ferrarit, an elongate-mandible form, and the other to mitida, a short-mandible form. The only reason which I can see for this procedure is that in Bernard’s sulcinodis the mandibles are open, whilst in his occidentalis they are closed. Measurement of the mandibles shows that they are of equal dimensions in both but the fact that they are open in one and closed in the other gives the illusion that those of the former are actually longer than those of the latter. This species is of course related to ferrarii and its allies and not to nitida. Its closest relative within the group appears to be sulcinoda but the two are separable by the shape of the occipital margin and the dimensions of the petiole, as noted in the key. Leptogenys princeps sp. n. (Text-fig. 26) DIAGNOSIS OF WORKER. Head reticulate-punctate. Dorsal alitrunk completely sculptured. Node of petiole massive. Large species, HL > 1:20, PW > 0°75. FURTHER DESCRIPTION. Holotype worker. TL 5:9, HL 1:26, HW o-go, CI 71, SL 1-30, SI 144, PW 0-82, PH 0-86, PL 0-62, LPI 138, DPW 0-68, DPI tog. Head slightly broadened in front of the eyes, the sides of the head feebly convex, the occipital margin in full-face view feebly concave. Maximum diameter of eye 0-32, greater than the maximum width of the scape. Eyes with a weak circumocular impression on their inner and posterior margins. Respective lengths of funicular segments 1-3 ca 0-18, 0:26, 0-24. Node of petiole massively constructed, in profile nearly as large as the first gastral segment. Posterior face of node approximately vertical, meeting the dorsal surface in an acute angle which slightly overhangs the posterior face. In dorsal view the nodes as long as the first gastral tergite, broader behind than in front. Hairs present on all surfaces of head, body and appendages. Dorsum of head coarsely reticulate-punctate, the sides and ventral surface unsculptured apart from a number of hair-pits. Alitrunk everywhere finely rugulose, the dorsal alitrunk and sides of the pronotum also with scattered large punctures. Propodeal declivity only with transverse rugulae. Upper halves of sides of petiole finely punctate with some larger punctures, the lower halves smooth and shiny. Posterior one-quarter of the sides 272 B. BOLTON of the node with a vertical series of short, coarse, longitudinal sulci. Gaster unsculptured apart from small hair-pits, smooth and shining. Colour dark brown, the mandibles, antennae and legs lighter, the node black. Holotype worker, GHANA: Mt Atewa, 4.vili.1966 (D. Leston) (BMNH). This is one of the larger species of the group and its size and massive petiole node make it easy to identify. Leptogenys ravida sp. n. (Text-fig. 10) DIAGNOSIS OF WORKER. Eyes relatively small, their maximum diameter about equal to the length of the third funicular segment. Median clypeal lobe narrow, short and acute. FURTHER DESCRIPTION. Holotype worker. TL 5-1, HL 1-08, HW 0-78, CI 72, SL 1:02, SI 130, PW 0-66, PH 0-64, PL 0-48, LPI 133, DPW 0-46, DPI 96. Head broader in front than behind, the sides feebly convex and diverging anteriorly through- out their length. Occipital margin in full-face view broadly and very shallowly concave. Eyes relatively small for a member of this group, maximum diameter ca 0-18, about equal in length to the third funicular segment and only very slightly larger than the maximum width of the scape (ca 0:14). Median lobe of clypeus short, narrow and acute. Mandibles elongate and narrow, not closing tightly against the clypeus. Node of petiole with the anterior and posterior faces approximately vertical in profile, but the former shorter than the latter so that the dorsal surface slopes upwards posteriorly. Dorsum of head smooth and shining, unsculp- tured apart from a number of minute, widely scattered hair-pits. Dorsal alitrunk similar except that the hair-pits are much larger and the extreme posterior portion of the propodeum has a few faint, transverse striae. Propodeal declivity rugose. Sides of pronotum unsculptured apart from scattered pits. Mesopleuron with a few faint rugulae and extensive smooth, shining areas; metapleuron and sides of propodeum minutely rugulose. Node of petiole with a few faint striae on the upper halves of the sides. Gaster smooth and shiny apart from minute hair-pits. Black, shining, the appendages brown. Holotype worker, ZAIRE: (‘Belg. Congo’ on data label) Mulungu Res. Sta., 1650 m, 28.iv.1948 (A. Emerson), ‘Ants from mound of Odontotermes patruus (Sjostedt)’ (MCZ, Cambridge). Leptogenys sulcinoda (André) (Text-fig. 14) Lobopelta sulcinoda André, 1892 : 48. Holotype worker, GABON (Mocquerys) (MNHN, Paris) [examined]. Leptogenys (Lobopelta) sulcinoda (André); Emery ,1911 : 102. DIAGNOSIS OF WORKER. Upper halves of sides of petiole sulcate. Pronotum unsculptured apart from hair-pits. Occipital margin in full-face view feebly convex. FURTHER DESCRIPTION. Worker. TL 4:6, HL 1:02, HW 0-72, CI 70, SL o-go, SI 125, PW 0:58, PH 0:64, PL 0-48, LPI 133, DPW 0-46, DPI 96. Head only slightly broadened anteriorly, the occipital margin feebly convex. Eyes large, maximum diameter 0-24. Median lobe of clypeus broad and broadly rounded anteriorly. Mandibles short, capable of closing tightly against the clypeus. Petiole in profile with the posterior face very weakly sinuate, much longer than the anterior face so that the dorsal surface slopes upwards posteriorly. Anterior face of petiole grades into dorsum through a broad convexity; the angle separating dorsum and posterior face more acute though still REVISION OF GENUS LEPTOGENYS 273 rounded. In dorsal view the node slightly longer than broad, broadest posteriorly. Upper halves of sides of petiole node converge dorsally so that the dorsal surface is very narrow. Gaster strongly constricted between first and second segment. Head with numerous fine, shallow punctures, separated by shining interspaces; sides and ventral surface unsculptured. Alitrunk mostly unsculptured apart from hair-pits but the declivity of the propodeum has a few weak transverse rugae, and a few weak rugae or vestiges of such are present upon the propodeum and metapleuron. Upper halves of sides of petiole strongly longitudinally sulcate. Gaster unsculptured apart from minute hair-pits. Colour black, the appendages brown. This species is apparently only known from the original collection of a single specimen. It is closest related to occidentalis but separated from it by the possession of a feebly convex occipital margin, and by the other characters noted above and in the key. Two other species are known in which the upper halves of the sides of the petiole are longitudinally sulcate, mastax and nuserra, but in both of these species the pronotal dorsum is strongly sculptured. Leptogenys terroni sp. n. DIAGNOSIS OF WORKER. Dorsum of head and alitrunk covered with dense sculpturation. Large species, HL ca 1-48, SL ca 1-62. FURTHER DESCRIPTION. Holotype worker. TL 7:8, HL 1-48, HW 1:18, CI 80, SL 1-62, SI 137, PW 0-98, PH 0-90 PL 0°74, LPI 121, DPW 0:66, DPI go. Head much broader in front than behind, the width across the clypeus considerably greater than across the eyes. Sides of head more or less straight, diverging strongly from back to front. Eyes large, maximum diameter 0-42, much greater than the maximum width of the scape and with a distinct circumocular impression above, in front of and behind the eye, but not below. Clypeus with a prominent, truncated median lobe. Mandibular blades very long, only feebly curved along their length, increasing slightly in width from base to apex. Funicular segments I—3 Ca 0°30, 0:32, 0-18 respectively. Metanotal groove impressed and crossribbed. Petiole with the upper halves of the sides converging dorsally so that the true dorsal surface is a narrow, longitudinal strip. Gaster strongly impressed between first and second segments. All surfaces of head, body and appendages with numerous standing hairs. Dorsum of head, especially between the eyes, finely and closely reticulate-rugulose, the spaces between the rugulae with numerous small punctures. Pronotum and mesonotum covered with a dense, fine reticu- lation and with scattered hair-pits. Propodeum and upper halves of sides of petiole similarly sculptured but also with larger rugae. Sides of pronotum as dorsum, the pleurae rugose. Legs, gaster and lower halves of sides of petiole smooth and shining, with hair-pits but otherwise unsculptured. Colour black, the gaster and appendages brown, the former darker in colour than the latter. Holotype worker, CAMEROUN: no. 1519 (no further data) (G. Terron) (BMNH). This is the largest and one of the most distinctive species yet known in the sulcinoda-group. Its large size and characteristic sculpturation render it easily distinguishable from other members of the group. Leptogenys trilobata Santschi (Text-fig. 11) Leptogenys trilobatus Santschi, 1924 : 196, fig. 1. Holotype worker, ZAIRE: Tschela, 2.xi.1920 (H. Schouteden) (MRAC, Tervuren) [examined]. 274 B. BOLTON DIAGNOSIS OF WORKER. Dorsum of head rugose, dorsum of alitrunk not densely sculptured, mostly shining. Median clypeal lobe truncated apically. FURTHER DESCRIPTION. Worker. TL 5:4-5:6, HL 1:02-1:10, HW 0-76-0-84, CI 75-76, SL 0:86-0:94, SI 112-113, PW 0:°62-0-68, PH 0:54-0:60, PL 0°56—0-62, LPI 96-98, DPW 0-50— 0:52, DPI 84-89 (2 measured). Head broader in front than behind. Maximum diameter of eyes 0:22-0:24, greater than the maximum width of the scape. A weak circumocular impression present on inner and posterior margins of eyes. Median lobe of clypeus strongly prominent, with almost parallel sides and a broadly arcuate, truncated apex. Mandibles long and narrow, not closing against the clypeus. Respective lengths of funicular segments 2—3 ca 0:16-0:18, 0-14-0'16. Metanotal groove weakly cross-ribbed. Node of petiole massive; in dorsal view longer than wide, widest posteriorly; in profile very nearly as long as high. Gaster constricted between first and second segments. Dorsum of head rugose, with numerous minute punctures; the median strip of the ventral surface unsculptured. Dorsal alitrunk with scattered foveolate punctures, the pronotum with a few weak, rounded rugae, the propodeum also with a few small punctures. Meso- and metapleurae striate, with punctures between the striae. Gaster smooth and shining. Standing hairs numerous on all surfaces. Black with the appendages light brown. MATERIAL EXAMINED. CAMEROUN: Yolé Massif, Kala (G. Terron) THE ATTENUATA-Grovup (Text-figs 22, 29, 30) Mandibles short, closing tightly against the clypeus. Eyes medium to large, their maximum diameter greater than the maximum width of the antennal scape. Second funicular segment distinctly longer than the third. Head slightly or not broadened anteriorly. Node of petiole in dorsal view much longer than broad. Sculpture only of fine punctures dorsally from which hairs arise, with large shining spaces between the punctures. Gaster constricted between first and second segments. Clypeus without a prominent median lobe, without a fringing lamella. Of the four species placed in this small group bubastis and crassinoda are certainly closely related, but the remaining species, attenuata and ergatogyna, are very different. However, in view of the characters above it is probably best to leave them in one group until the genus as a whole is better understood. In the four species, crassinoda and attenuata are known only from South Africa whilst bubastis is known only from Cameroun. The last species, ergatogyna, is more widely distributed, having been recorded from Ghana, Uganda and Zaire, and appears to be strictly a forest species. Three other species referable to this group and related either to attenuata or crassinoda are known from the Malagasy Region. Leptogenys attenuata (F. Smith) (Text-fig. 29) Poneva attenuata F. Smith, 1858 : 91. LECTOTYPE and paralectotype workers, by present designation, SourH Arrica: Cape of Good Hope, no. 1490 (Drege) (UM, Oxford; BMNH) {examined}. REVISION OF GENUS LEPTOGENYS 275 Lobopelta attenuata (F. Smith); Roger, 1863 : 19. Leptogenys (Lobopelta) jaegerskjoeldi Santschi, 1914b:9. Syntype workers, SOUTH AFRICA: Natal, Richmond, 24.iii.1905 (I. Trdgdévdh) NM, Basle) [examined]. [Synonymy by Arnold, 1926 : 210.] Leptogenys jagerskiéldi Santschi; Santschi, 1917 : 297. [Misspelling.] DIAGNOSIS OF WORKER. Petiole in dorsal view elongate and strongly compressed from side to side, much narrower anteriorly than posteriorly. Body often with bluish or violet reflections. Antennal scapes moderately long, SI > 175. FURTHER DESCRIPTION. Worker. TL 7:5—7:8, HL 1:50-1:62, HW 0-94-1-02, CI 59-63, SL 1-72-1:98, SI 183-194, PW 0:84-0:94, PH 0:66-0:80, PL 0-76-0-88, LPI 86-91, DPW 0:38-0:46, DPI 50—53 (8 measured). Head elongate, quite narrow, broadest across the eyes which are situated approximately at the midlength of the sides of the head. Occipital margin concave in full-face view. Eyes large, maximum diameter ca 0-36—-0'42, greater than the maximum width of the scape. Clypeus triangular, the mandibles closing tightly against it. Respective lengths of funicular segments I-3 ca 0:26-0:28, 0:46—0°48, 0:36—-0'40. Metanotal groove impressed but not cross- ribbed. Node of petiole in dorsal view elongate, strongly narrowed anteriorly, much broader behind than in front. In profile the anterior and dorsal faces confluent through a broad curve. Gastral constriction between first and second segments distinct. Sculpture lacking except on the mandibles where a few very fine striae are usually present. All surfaces of body with standing hairs. Colour black or blackish brown, usually with bluish or violet reflections. Legs, mandibles and funiculi lighter. This quite distinctive species, the male of which was described by Arnold (1915), appears to be restricted to South Africa. Some apparently closely related forms are known from the Oriental and Indo-Australian Regions, such as peuquett André and assamensis Forel, and these species resemble attenuata more closely than any species present in the Ethiopian Region. MATERIAL EXAMINED. Soutu Arrica; E. Cape Prov., Katberg (R. E. Turner); Grahamstown (W. L. Brown); E. Cape Prov., Alexandria (W. L. Brown); Natal, Durban (G. Arnold) ; Durban (H. B. Marley); Durban (C. B. Cooper); W. Grahamstown (fF. Jacot- Gutllarmod); Cape Prov., Knysna (B. Malkin). Leptogenys bubastis sp. n. DIAGNOSIS OF WORKER. Propodeal dorsum without a transverse impression anteriorly but with scattered large pits. Petiole node in dorsal view not strongly compressed from side to side. Scapes relatively short, SI < 165. FURTHER DESCRIPTION. Holotype worker. TL 7-0, HL 1-42, HW 0-98, CI 69, SL 1°50, SI 153, PW 0-86, PH 0:72, PL 0-70, LPI 103, DPW 0-46, DPI 66. Head broadest across the eyes, not much broader in front than behind, the sides feebly convex. Maximum diameter of eye ca 0-30, greater than the maximum width of the scape. Clypeus broadly triangular, the mandibles closing tightly against it. Funicular segments I-3 Ca 0:26, 0°42, 0-30 respectively. Metanotal groove a narrow, incised line, not cross-ribbed. Propodeal dorsum relatively much longer than in the related crassinoda (see key, couplet 49). Node of petiole in dorsal view longer than broad, narrowing anteriorly but the sides not strongly compressed. Gaster constricted strongly between first and second segments. Dorsal surfaces of head and body with short, standing hairs. Head with numerous minute punctures from which hairs arise, the pronotum with very few such pits, but the mesonotum and 276 B. BOLTON propodeum with a number of large pits. First and second gastral tergites abundantly supplied with distinct hair-pits. Propodeal declivity not transversely rugose. Sides of alitrunk with some cross-ribbing or rugosity at the suture lines and on the bulla of the metapleural glands; otherwise the body smooth. Colour black, the appendages brown. Holotype worker, CAMEROUN: series MZ (G. Terron) (BMNH). Most closely related to crassinoda; the differences between the two are given under that species and in the key. Leptogenys crassinoda Arnold stat. n. (Text-fig. 30) Leptogenys attenuata var. crassinoda Arnold, 1926: 210. Syntype workers, SOUTH AFRICA: East London (Rattray) (BMNH) [examined]. DIAGNOSIS OF WORKER. Propodeal dorsum without a transverse impression anteriorly, with scattered large pits. Petiole node in dorsal view not strongly compressed from side to side. Antennal scapes relatively short, SI < 165. FURTHER DESCRIPTION. Worker. TL 6-2-7:2, HL 1:38-1:42, HW 0:94-1:00, CI 68-70, SL 1-48-1:52, SI 152-157, PW 0:82-0:86, PH 0:70-0:72, PL 0-64-0:66, LPI 109-110, DPW 0:40-0:44, DPI 60-68 (2 measured). Head broadest across the eyes, not markedly broader in front than behind. Occipital margin broadly but shallowly concave in full-face view. Maximum diameter of eye ca 0-28— 0:30, greater than the maximum width of the antennal scape. Clypeus triangular, the mandibles closing tightly against it. Metanotal groove impressed but not cross-ribbed. Node of petiole in dorsal view elongate, broadest behind and narrowing slightly anteriorly. Constriction between first and second gastral segments distinct. Body unsculptured. Standing hairs present upon all surfaces. Colour black, the appendages brown or yellow-brown. Arnold originally described this species as a variety of attenuata but it is smaller, with a broader head, smaller eyes, shorter scapes and a shorter, thicker petiole. As far as can be ascertained crassinoda is still only known from the type-collection. The closest related species to crassinoda is bubastis, which it closely resembles. However, in bubastis the mesonotum and propodeum have numerous scattered, large punctures or pits which are absent from crassinoda, and the propodeal dorsum is relatively much longer in bubastis, as is noted in the key. Leptogenys ergatogyna Wheeler (Text-fig. 22) Leptogenys (Lobopelta) ergatogyna Wheeler, 1922 : 95, fig. 20. Holotype female, ZAIRE: Medje (Lang & Chapin) (AMNH, New York) [examined]. Leptogenys (Lobopelta) cursor Arnold, 1954 : 293, fig. 4. Holotype worker, UGANDA: Zika Forest, near Entebbe, 18.v.1952 (G. Arnold) (NM, Bulawayo) [examined]. Syn. n. DIAGNOSIS OF WORKER. Mandibles massively developed, very broad. Propodeal dorsum with a broad, transverse impression anteriorly and a longitudinal groove which is most distinct posteriorly. Antennal scapes very long, SI > 200. - FURTHER DESCRIPTION. Worker. TL 8:0-8-4, HL 1-60—1:62, HW o0-98-1-02, CI 61-62, SL 2-34-2°44, SI 238-244, PW 0:96-1:02, PH 0-76-0-80, PL 0-71-0-72, LPI 106-111, DPW 0-48-0-50, DPI 66—69 (4 measured). REVISION OF GENUS LEPTOGENYS 277 Head elongate and narrow, somewhat broader in front than behind; occipital margin short and only extremely weakly concave. Eyes large, maximum diameter ca 0-40-0-42, the eyes situated just in front of the midlength of the sides of the head. Clypeus triangular. Mandibles massive, broad and heavy, their basal borders closing tightly against the clypeus. Apical (masticatory) margin very long, meeting the basal margin through a broad curve. A single preapical tooth present, close to the apical. Antennal scapes very long; lengths of funicular segments 1-3 respectively ca 0+30-0:34, 0:60-0:64, 0:46-0°50. Metanotal groove present but not cross-ribbed. Propodeal dorsum with a longitudinal groove, most distinct posteriorly, and also with a transverse impression anteriorly, just posterior to the metanotal groove. Node of petiole in dorsal view elongate, broadest behind and narrowing slightly anteriorly. Constriction between first and second gastral segments distinct. Meso-metapleural suture cross-ribbed, otherwise body unsculptured apart from hair-pits, which are abundant on the head. Standing hairs present on all dorsal surfaces; pubescence quite dense on dorsum of head and on legs. Colour black, the appendages brown. Female. TUL 8-2, HL 1-62, HW 1:00, CI 62, SL 2:26, SI 226, PW 1-02, PH 0-84, PL 0-72, LPI 116, DPW 0-52, DPI 72. Head similar to that of worker but with three ocelli, of which the median (anterior) is the best developed. Scapes relatively slightly shorter than in worker but eyes about the same size, maximum diameter ca 0-42. Alitrunk with developed flight sclerites but wings apparently never developed. Parapsidal furrows present; posterior arm of notauli present, the anterior arms absent. Node of petiole slightly higher and broader than in worker, the gaster swollen beyond the size of the worker. In the female the height of the second segment in profile is 1-20 whilst in the worker it is 1-04, and the dorsal width of the second segment is 1-10 in the female, 0-95 in the worker. The female has some fine, transverse striation across the anterior portion or covering all of the propodeal dorsum which is not present in the worker, and lacks the transverse propodeal impression present in the worker, but otherwise the two are similar. The association of the worker and female castes of this species described respectively as cursor and ergatogyna, was permitted by a series collected at Tafo, Ghana by D. Leston, which contained a number of workers and a single female. The holotype of ergatogyna and the female in the Ghana series are similar in most respects but the former is more strongly sculptured. The fine punctures of the head are rather more dense and are also abundant on the pronotum, which in the case of the Ghana female is mostly smooth and shining. Transverse striae are restricted to the anterior half of the propodeal dorsum in the Ghana specimen whilst in the holotype of ergatogyna they occur on the entirety of the segment. MATERIAL EXAMINED. GHANA: Tafo (D. Leston); Mt Atewa (D. Leston); Nkwantanan (D. Leston). CAMEROUN: Ototomo (G. Terron). THE GUINEENSIS-Grovup (Text-figs 16, 23, 28) Mandibles short, closing tightly against the clypeus. Eyes small or minute, their maximum diameter always less than the maximum width of the antennal scape. Second funicular segment as long as or shorter than the third. Head only slightly or not broadened anteriorly. Sculpture usually only of fine punctures dorsally. Gaster not or only very feebly constricted between the first and second segments. Standing hairs absent from dorsum of propodeum, petiole and first and second gastral tergites. 278 B. BOLTON The four related species forming this group are presently known only from West Africa; three of the species are forest inhabitants whilst the fourth (microps) is a savannah species. Leptogenys guineensis Santschi (Text-fig. 16) Leptogenys (Lobopelta) guineensis Santschi, 1914a : 329, fig. 14. Syntype workers, GUINEA: Mamou (F. Silvestri) (NM, Basle) [examined]. DIAGNOSIS OF WORKER. Apex of mandible with two teeth. Larger species, HL > 0°88. Petiole in profile not a dorsally tapered, thick scale. FURTHER DESCRIPTION. Worker. TL 4:3-4:5, HL 0:92-0:96, HW 0-68-0-70, CI 73-74, SL 0:80-0:82, SI 117-118, PW 0:54-0:56, PH 0:52, PL 0:32-0:36, LPI 145-160, DPW 0:36-0:38, DPI 105-112 (6 measured). Head not broadened anteriorly, broadest just behind the eyes, the sides shallowly convex. Eyes small, maximum diameter ca 0:08; in full-face view their maximum diameter less than the greatest width of the scape. Second funicular segment usually slightly shorter than the third, sometimes the same length; the relative lengths of segments 1-3 ca 0:12—0'14, 010-012, 0:12. Mandibles short, with a single preapical tooth and closing tightly against the clypeus. Metanotal groove impressed and with a very fine cross-ribbing so that the base of the groove appears to be traversed by a row of punctures. Node of petiole high and narrow in profile, with a slightly convex anterior face and a virtually straight, sloping posterior face. Gaster only very weakly impressed between first and second segments. Body unsculptured apart from transverse rugae on propodeal declivity and cross-ribbing on the meso-metapleural suture. Short, standing hairs absent from propodeum and first two gastral tergites. Colour brown,.glossy, the appendages lighter. This species is very closely related to testacea, and further collections may prove them to be inseparable. However, in the material examined during the course of this study specimens of guineensis have consistently been larger and have lacked the third (middle) tooth on the mandibular apex which is characteristic of testacea. Another feature which appears to be typical of guineensis is the presence on the anterior portion of the median clypeal lobe of a yellow-coloured patch which is usually about the same colour as the mandibles. Two long setae with distinct black pits project from the upper portion of this area. In ¢estacea this area is slightly lighter in colour but does not form a strong contrast with its surroundings as it does in guineensis. MATERIAL EXAMINED. GHANA: Mt Atewa (C. A. Collingwood); Mt Atewa (D. Leston) ; Tafo (D. Leston). Leptogenys microps sp. n. DIAGNOSIS OF WORKER. Apex of mandible with two teeth. Minute species, HL < 0°70; eyes minute, maximum diameter < 0:05. FURTHER DESCRIPTION. Holotype worker. TL 2:7, HL 0:66, HW 0:48, CI 73, SL 0-50, SI 104, PW 0:38, PH 0-34, PL 0-28, LPI 121, DPW 0-22, DPI 78. Head not broader in front than behind, broadest at about its midlength, the sides feebly convex. Occipital margin very weakly concave medially in full-face view. Clypeus with a REVISION OF GENUS LEPTOGENYS 279 relatively short, bluntly triangular median lobe, the apex of which is rounded. Mandibles short, closing tightly against the clypeus, the inner margins of the blades convex, the outer slightly concave in full-face view. Eyes minute, maximum diameter 0-04, much less than the maximum width of the scape, with very few facets. Funiculus with segments 1-3 ca o-1Io, 0:06, 0:07 respectively, the second funicular segment very nearly as broad as long. Petiole in profile with the anterior face slightly convex, rounding into the dorsal surface; the latter meeting the almost straight posterior face in a more acute angle. Body of node inclined forward so that the anterior face is much shorter than the posterior and so that when the ventral surface is horizontal the anterior and posterior faces slope anteriorly. Gaster weakly constricted between first and second segments. Standing hairs absent from dorsum of propodeum, petiole and first two gastral tergites, which do, however, have a strongly adpressed pubescence. Pronotum with a pair of long, erect hairs anteriorly and with shorter standing hairs and pubescence. Dorsum of head with minute, faint superficial punctures. Dorsal alitrunk with scattered punctures, larger than those on the head and more distinct, most obvious on the propodeum. Propodeal declivity with a few fine, transverse striae. Gastral tergites with scattered minute punctures. Colour uniform dull yellow. Paratype workers. TL 2-7-2:9, HL 0-64-0:66, HW 0:48-0:50, CI 73-78, SL 0:50-0:52, SI 104-108, PW 0:38, PH 0:34-0:36, PL 0-28, LPI 121-128, DPW 0:22-0:24, DPI 78-86 (2 measured). As holotype, with eyes the same size, but in one the occipital margin in full-face view is nearly straight and the punctures of the propodeal dorsum are shallower and less well defined. Holotype worker, Ivory Coast: Lamto (Toumodi), 15.ix.1965, AA129 (probably collected by J. Lévieux) (MCZ, Cambridge). Paratypes. I worker with same data as holotype, and 1 worker from the same locality but dated 10.ix.1965, AA 124 (BMNH; MCZ, Cambridge). This minute species is the smallest of the guineensis-group and one of the smallest species known of the genus. In some ways it is convergent with the very small species of the nitida-group (khaura, cryptica, stygia) but these forms have numerous standing hairs on the propodeum, petiole and gaster, absent from microps. Leptogenys spandax sp. n. (Text-fig. 28) DIAGNOSIS OF WORKER. Apex of mandible with two teeth. Larger species, HL > 1:00. Petiole in profile a thick scale, tapering dorsally. FURTHER DESCRIPTION. Holotype. TL 6-8, HL 1-38, HW 0-98, CI 71, SL 1°46, SI 149, PW 0-82, PH 0:82, PL 0-60, LPI 137, DPW 0-54, DPI 90. Occipital margin feebly convex, sides of head very weakly convex and slightly diverging anteriorly so that the front of the head is somewhat wider than the back. Eyes small, maximum diameter ca 0-14, failing to break the outline of the sides of the head in full-face view. Maximum diameter of eye distinctly less than maximum width of scape. Clypeus with a broadly triangular median lobe. Mandibles short, closing tightly against the clypeus, their inner margins convex, outer margins slightly concave. Second and third funicular segments of equal length, ca 0:24. Metanotal groove impressed and weakly cross-ribbed. In profile the propodeal declivity with a short tooth on each side, projecting just above the bulla of the metapleural glands. Node of petiole in profile a thick scale, tapering strongly in its upper half. Anterior face almost vertical, posterior face sloping strongly anteriad, the dorsum very short. In dorsal view the node short and broad, with a narrow, wide dorsal surface, a slightly convex anterior and slightly concave posterior face. Gaster not constricted between first and second segments. Standing hairs absent from dorsal surfaces of head, alitrunk, gaster and appendages, present only around 280 B. BOLTON the mouth and gastral apex. Dorsum of head finely punctate. Pronotum similarly sculptured but the punctures larger and more dispersed, with reticulate sculpture between them. Propodeal dorsum finely rugulose, the declivity very coarsely transversely rugose. Sides of propodeum and metapleuron rugose, mesopleuron with patches of rather effaced sculpture, mostly smooth and shining. First gastral tergite with numerous minute punctures. Brown, the legs, antennae and mandibles yellow-brown. Paratype worker. As holotype but with HL 1-36, HW 0:98, CI 72, SL 1-44, SI 147, PW 0-78, PH 0:78, PL 0°58, LPI 134, DPW 0°54, DPI 93. Holotype worker, GHANA: Tafo, 24.ix.1966, ant ecology sample 282E (D. Leston) (BMNH). Paratype. I worker with same data as holotype (MCZ, Cambridge). Leptogenys testacea (Donisthorpe) (Text-fig. 23) Microbolbos testaceus Donisthorpe, 1948 :170. Holotype and paratype workers, GHANA: Eastern Region, New Tafo, vii. 1945, no. F463 (Strickland) (BMNH) [examined]. Leptogenys testacea (Donisthorpe) ; Wilson, 1955 : 136. DIAGNOSIS OF WORKER. Apex of mandible with three teeth. Smaller species, HL 140. Fringing lamella of clypeus distinct, broad and often opaque. FURTHER DESCRIPTION. Worker. TL 6:5-6:°8, HL 1:30-1:40, HW 0:92-1:00, CI 70-72, SL 1°38-1-48, SI 147-150, PW 0-80-0:86, PH 0-74-0°80, PL 0:56-0:60, LPI 130-133, DPW 0°52-0°58, DPI 92-07 (6 measured). Head only very slightly broader in front than behind, the sides very feebly convex, almost 282 B. BOLTON straight. Frontal groove short, not extending to level of posterior margins of eyes. Eyes medium, maximum diameter ca 0:28-0:32. Clypeus broad, with a distinct and usually opaque, broad fringing lamella on the anterior margin. Mandibles short, closing tightly against the clypeus. Respective lengths of funicular segments 1-3 ca 0:22-0:24, 0°32-0°36, 0:24-0:26. Metanotal groove deeply impressed; gaster very strongly constricted between first and second segments. All surfaces of body with numerous standing hairs. Dorsum of head, alitrunk and first two gastral tergites very finely and very densely reticulate-punctate or punctulate, the individual punctures so small and closely packed that the surfaces appear coarsely granular. Propodeal dorsum also with a few weak rugulae. Pleurae and sides of propodeum more coarsely sculptured, with distinct rugae. This species is separated from others of the group by its strongly sculptured gaster, but in all other respects it is very close to havilandi. In the original descrip- tion Arnold (1926) noted that the workers were carrying termites in their jaws. Leptogenys havilandi Forel Leptogenys (Lobopelta) havilandi Forel, 1901a : 332: Holotype worker, SoutH AFRIca: Natal (Haviland) (MHN, Geneva) [examined]. DIAGNOSIS OF WORKER. First and second gastral tergites unsculptured. Antennal scapes longer, SI > 140. Fringing lamella of clypeus distinct, broad. Maximum diameter of eye < 0°34. FURTHER DESCRIPTION. Worker. TL 6:7—-7:2, HL 1:32-1:42, HW 0-96-1-00, CI 70-73, SL 1:40-1:60, SI 146-160, PW 0:80-0:88, PH 0-76-0:80, PL 0:60-0:62, LPI 126-130, DPW 0:50-0:56, DPI 83-91 (3 measured). Head only very slightly broader in front than behind, the sides feebly convex, almost straight. Frontal groove distinct, extending to the level of the posterior margin of the eyes or further. Maximum diameter of eyes ca 0-26—-0-30. Clypeus broadly triangular and rounded, with a distinct fringing lamella on the anterior margin. Mandibles closing tightly against clypeus. Respective lengths of funicular segments I-3 ca 0:22-0:24, 0-34—0:38, 0:26-0:28. Metanotal groove deeply impressed; gaster strongly constricted between the first and second segments. All surfaces of body with numerous standing hairs. Dorsum of head reticulate-punctate. Pronotum with small, crowded rugulae and some puncturation, similarly the mesonotum and propodeal dorsum but on the latter the rugulae tend to be coarser. Sides of propodeum and pleurae coarsely rugose, the pronotum finely sculptured. First and second gastral tergites unsculptured apart from hair-pits. Most closely related to furtiva and peringueyt, from which it is separated by details of sculpturation; havilandi may be found to be inseparable from them when more collections have been made. At present havilandi represents a mid-point between the very heavily sculptured furtiva and the lightly sculptured peringueyi, but these three forms may well be different expressions of a single variable species. Leptogenys peringueyi Forel Leptogenys (Lobopelta) havilandi st. peringueyt Forel, 1913c¢ : 210. Holotype worker, SouTH AFrRIca: Cape Prov., Table Mountain (L. Peringuey) (MHN, Geneva) [examined]. Leptogenys (Lobopelta) peringueyi Forel; Arnold, 1915 : 96. [Raised to species.] DIAGNOSIS OF WORKER. First and second gastral tergites unsculptured. Pronotal dorsum sculptured with punctures separated by smooth, shining spaces. REVISION OF GENUS LEPTOGENYS 283 FURTHER DESCRIPTION. Worker. TL 6:7, HL 1-30, HW o-go, CI 70, SL 1-22, SI 135, PW 0:74, PH, PL — not measurable, DPW 0:50. Head not broadened in front of eyes, the sides approximately straight. Maximum diameter of eyes ca 0-26, greater than the maximum width of the scape. Frontal groove extending to level of posterior margins of eyes. Clypeus broadly triangular, with a narrow, fringing lamella. Mandibles short and stout, closing tightly against the clypeus. Metanotal groove deeply impressed; gaster strongly constricted between first and second segments. All surfaces of body with numerous standing hairs. Dorsum of head densely punctate, the pronotal dorsum more sparsely so, with distinct smooth, shining spaces between the individual punctures, broader than those upon the head. Mesonotum similarly sculptured but propodeal dorsum also retaining traces of rugose sculpturation. Sides of pronotum very lightly sculptured; pleurae and sides of propodeum more strongly so, with distinctrugosity. Gaster unsculptured apart from hair-pits. This species is closely related to havilandt, the major differences separating the two belonging to the sculptural patterns of the head and alitrunk. When they are better known peringueyi and havilandi may be found to be conspecific. THE NITI DA-GrRoup (Text-figs 15, 17, 18, 25) Mandibles short, closing tightly against the clypeus. Eyes medium to minute, their maximum diameter in full-face view ranging from just greater than the maximum width of the scape to considerably less than the scape width. Second funicular segment at most about as long as the third, usually shorter than the third. Head not or only slightly broadened anteriorly. Sculpture usually only of scattered fine punctures on the dorsal surfaces; reticulate-punctate in some species, rarely otherwise. Gaster constricted between first and segments segments. This group of fourteen species divides itself into a number of complexes of more or less closely related forms. The first, including cryptica, khaura and stygia, is West African in distribution and is characterised by strong reticulate-punctate sculpture on the cephalic dorsum and by its small size. The second, including buyssoni, ankhesa, honoria and strator are known from Zaire, Cameroun and Ethiopia. They are larger species with relatively blocky petiole nodes and strongly developed subpetiolar processes. They appear to feed on termites. The remaining species are found in central, eastern and southern Africa, are medium-sized, lightly sculptured and with usually a simple lobiform subpetiolar process. One of them (mactans) appears to have convergently acquired a number of characters possessed by the processtonalis-group of the Oriental and Indo-Australian Regions. Leptogenys amon sp. n. (Text-fig. 18) DIAGNOSIS OF WORKER. Head relatively long and narrow, CI ca 62. Circumocular groove very strongly developed. Petiole long and narrow, DPI < 70. FURTHER DESCRIPTION. Holotype worker. TL 5:2, HL 1-12, HW 0-70, CI 62, SL 0°84, SI 120, PW 0°58, PH 0-52, PL 0:54, LPI 97, DPW 0-34, DPI 66. Head elongate and narrow, increasing slightly in width anteriorly; occipital margin very shallowly concave, the sides of the head more or less straight. Maximum diameter of eyes 284 B. BOLTON ca 0-18, about equal to the maximum width of the strongly dorsoventrally flattened scapes. Circumocular groove strongly developed in front of, above and behind the eye so that the eye appears to be sunk in an impression in the side of the head. Median portion of clypeus a broad, rounded lobe. Mandibles short and stout, closing tightly against the clypeus. Funicular segments I—3 respectively ca 0:16, 0°18, 0:16. Metanotal groove an incised line; cross-ribbing is present in the groove but the individual ribs are so small that the spaces between them appear as a line of small punctures across the dorsum. Node of petiole in dorsal view longer than broad, broadest behind and narrowing anteriorly. Gaster strongly constricted between first and second segments. Short, standing hairs present on all surfaces. Completely devoid of sculpture except for hair-pits and cross-ribbing on the meso-metapleural suture. A few weak rugulae are present on the bulla of the metapleural glands and the propodeal declivity is transversely rugose, though only 2-3 complete rugae are present. Colour medium brown, the gaster lighter, the appendages a dark yellow-brown. Holotype worker, CAMEROUN: Ototomo, 20.v.1968, JLA (G. Terron) (BMNH). This species is easily distinguished from the other members of the nitida-group by its possession of marked circumocular grooves, elongate head and long, narrow petiole. Leptogenys ankhesa sp. n. (Text-fig. 25) DIAGNOSIS OF WORKER. Subpetiolar process double, with separate anterior and posterior portions. Propodeal dorsum unsculptured. Scapes relatively long, SI > 135. FURTHER DESCRIPTION. Holotype worker. TL 6:1, HL 1-22, HW 0-84, CI 69, SL 1:20, SI 143, PW 0-76, PH 0-78, PL 0°58, LPI 134, DPW 0-62, DPI 107. Head only very slightly broader in front than behind, the occipital margin straight to feebly concave, the sides more or less straight. Maximum diameter of eye ca 0-20, slightly greater than the maximum width of the scape in full-face view (ca 0-12-0-14). Clypeus broadly triangular, blunt apically. Mandibles short, closing tightly against the clypeus. Respective lengths of funicular segments I-3 ca 0:20, 0:18, 0:20. Mesonotum distinctly broader than long. Metanotal groove impressed and cross-ribbed. Node of petiole in dorsal view broader than long, the anterior and posterior faces virtually parallel, the sides diverging posteriorly. In profile the node of the petiole broadening slightly from base to apex. Subpetiolar process double, complex, consisting of a lobe situated anteriorly, the posteroventral corner of which projects as a short tooth, and a more posteriorly situated short, blunt tooth, separated from the anterior lobe by a marked gap. Gaster strongly constricted between first and second segment. All dorsal surfaces of head and body with standing hairs, though these are sparse upon the first gastral tergite and propodeal. Unsculptured, smooth and shining except for small hair-pits and transverse rugulation on the propodeal declivity. Some faint rugae also present on the extreme lower portion of the metapleuron. Colour uniform red-brown. Paratype worker. TL 5:9, HL 1-20, HW 0-84, CI 70, SL 1-20, SI 143, PW 0-78, PH 0-80, PL 0-56, LPI 142, DPW 0:62, DPI 111. Maximum diameter of eye 0-18; funicular segments I-3 Ca 0°18, 0:20, 0:20. Otherwise as holotype. Holotype worker, ZAIRE (‘Belg. Congo’ on data label), Sona Mpungu, Forest, 21km N. Lufu, 10.iv. 1948, ‘ants collecting termites’ (A, Emerson) (MCZ, Cam- bridge). Paratype. I worker, same data as holotype (BMNH). Related to buyssoni and strator, it is separated from the former by size and the more complex development of the subpetiolar process (the second tooth is absent REVISION OF GENUS LEPTOGENYS 285 in buyssoni), and from the latter by details of sculpturation and shape of petiole node as given in the key and descriptions. Leptogenys buyssoni Forel Leptogenys (Lobopelta) buyssont Forel, 1907a : 131. Holotype worker, Etuiopia: Hieka Bourka, 1905 (Rothschild) (MHN, Geneva) [examined]. DIAGNOSIS OF WORKER. Larger species, HL > 1-45; scapes relatively long, SI > 140. Eyes small, maximum diameter ca 0:16. Subpetiolar process with a posteroventral tooth. FURTHER DESCRIPTION. Worker. TL 7-4, HL 1°52, HW 1-06, CI 70, SL 1:66, SI 156, PW 0-90, PH 0°84, PL 0-70, LPI 120, DPW 0-60, DPI 87. Head not broadened anteriorly, broadest at the level of the eyes. Occipital margin straight to feebly convex, the sides of the head straight. Eyes small, only very weakly convex, maximum diameter ca 0-16, less than the maximum width of the scape. Clypeus broadly triangular. Mandibles short, closing tightly against the clypeus. Second funicular segment slightly longer than the third, respective lengths of segments I-3 ca 0°24, 0°34, 0:30. Metanotal groove impressed and cross-ribbed, the propodeal dorsum with a slight longitudinal impression medially, most distinct at the midlength of the sclerite. Subpetiolar process a broad lobe, developed into a short tooth posteroventrally which projects from the body of the lobe. Gaster strongly constricted between first and second segments. All surfaces of body with short, standing hairs. Dorsal surfaces of head, alitrunk and gaster unsculptured apart from minute hair-pits. Sides of alitrunk with cross-ribbing at meso-metapleural suture and some short rugae near the metapleural gland, but otherwise as dorsum. Full adult colour red-brown. This species is closest related to stvator and ankhesa, but although the subpetiolar process is strongly developed in bwyssoni it is not followed by a second lobe or tooth posteriorly as is the case in the other two species. Also bwyssonz is distinctly larger than either strator or ankhesa. Leptogenys castanea (Mayr) Lobopelta castanea Mayr, 1862 : 734. Holotype worker, SoutH Arrica: Cape of Good Hope (BMNH) [examined]. Leptogenys (Lobopelta) parva Forel, 1901a : 330. Syntype workers, male, SouTH AFRICA: Natal (Haviland) (MHN, Geneva) [examined]. Syn. n. Leptogenys (Lobopelta) parva var. bellua Forel, 1914 : 214. Syntype workers, SouTH AFRICA: Natal, Durban (G. Arnold) (MHN, Geneva) [examined]. Syn. n. Leptogenys (Lobopelta) parva var. dispar Santschi, 1914b:10. Syntype workers, SOUTH Arrica: Natal, Zululand (I. Tragdrdh) (NM, Basle; MRAC, Tervuren) [examined]. Syn. n. Leptogenys hewitti Santschi, 1923 : 381, fig. 1. Syntype workers, SoutH AFRICA: C.P., Grahamstown, 1920 (J. Hewitt) (BMNH) [examined]. Syn. n. DIAGNOSIS OF WORKER. Small species with relatively short antennal scapes, HL < 0-90, SL < 0-75, SI < 120. Dorsal surfaces of body unsculptured. FURTHER DESCRIPTION. Worker. TL 3:5-4:1, HL 0-74-0-88, HW 0:54-0:68, CI 70-79, SL 0°62-0-70, SI 100-117, PW 0-44-0°52, PH 0:40-0:44, PL 0-28-0-32, LPI 131-143, DPW 0:28—0-34, DPI 93-104 (12 measured). Head not broader in front than behind, broadest behind the eyes. Occipital margin straight to feebly concave, the sides of the head shallowly convex. Eyes small, maximum diameter ca 0:08-0:12, equal to or slightly less than the maximum width of the scape in full-face view. Clypeus narrowly triangular medially. Mandibles short, closing tightly against the clypeus. Lengths of funicular segments 1—3 respectively ca 0:10-0:12, 0-08—0-10, 0-08-o-10. Subpetiolar 286 B. BOLTON process a broad and strongly developed simple lobe. Gaster strongly constricted between first and second segments. All surfaces of body with numerous short, standing hairs. Unsculptured apart from transverse rugae on propodeal dorsum. Colour light yellow-brown to light brown. This small species may be confused with the smaller, light-coloured workers of some populations of nitida but castanea may be distinguished by its scapes, which are consistently shorter, both relatively and absolutely, than those of nitida. Also, in the smallest niteda workers the eyes are about 0-12 in maximum diameter, whereas the eyes of only the largest examples of castanea approach this size. Finally, the HL of the largest castanea worker examined in this survey was 0-88, whereas that of the smallest nitida worker which could be found was 0-92. MATERIAL EXAMINED. RHODESIA: Bulawayo (G. Avnold). SouTH AFRICA: Natal, Durban (C. B. Cooper); Durban (I. Bevis); Krantz Kloof (H. B. Marley); Zululand, Mfongosi (G. Arnold), E. Cape Prov., Beggar’s Bush nr Grahamstown (W. L. Brown); Grahamstown, Fern Kloof (W. L. Brown); Alexandria Forest Reserve (W. L. Brown); Grahamstown (F. Jacot-Guillarmod). Leptogenys cryptica sp. n. DIAGNOSIS OF WORKER. Small species, HL < 0:75. Head densely reticulate-punctate; pronotum sculptured. Eyes very small, < 0-07; SI in the range 113-115. FURTHER DESCRIPTION. Holotype worker. TL 2:8, HL 0:66, HW 0-46, CI 70, SL 0:52, SI 113, PW 0-38, PH 0:36, PL 0-24, LPI 150, DPW 0-24, DPI Ioo. Head not broader in front than behind, broadest across the eyes. Sides of head shallowly convex, occipital margin straight to feebly concave medially. Eyes very small, maximum diameter ca 0:05, less than the maximum width of the scape in full-face view and with less than 10 facets. Clypeus narrowly triangular medially. Mandibles relatively short, closing tightly against the clypeus. Funicular segments 1-3 respectively ca 0-10, 0-08, 0-08. Metanotal groove reduced to a weak transverse impression, in profile scarcely more marked than the promesonotal suture; metanotal groove not cross-ribbed, in dorsal view less distinct than the promesonotal suture. Subpetiolar process a large subrectangular lobe without prominences or teeth. Gaster strongly constricted between first and second segments. All surfaces of body with numerous standing hairs. Dorsum of head coarsely and closely reticulate-punctate, the ventral surface with a number of foveolate punctures which are separated by smooth inter- spaces. Dorsal alitrunk strongly sculptured with a mixture of punctures and scattered rugulae, the latter most distinct upon the pronotum but not forming a pattern. Propodeal declivity transversely rugose. Sides of alitrunk as dorsum but more weakly so. Gaster smooth and shining, unsculptured. Colour uniform yellow-brown to mid-brown. Paratype workers. TL 2-6-3-1, HL 0-64-0:68, HW 0-44-0-46, CI 67-70, SL 0-50-0:52, SI 113-115, PW 0:36-0:38, PH ca 0:36, PL 0-22-0-24, LPI 150-163, DPW 0:20-0:24, DPI 100-109 (4 measured). Diameter of eye 0:04-0:06; respective lengths of funicular segments I-3 Ca O*10—0'12, 0:06-0:08, 0-:06—0-08. Otherwise as holotype. Holotype worker, GHANA: Tafo, I.x.1970, ‘rotten log’ (B. Bolton) (BMNH). Paratypes. 4 workers, same data as holotype (BMNH; MCZ, Cambridge; NM, Basle). The combination of light colour, small size, strong sculpture and very small eyes should render this species easily recognizable. It is most closely related to REVISION OF GENUS LEPTOGENYS 287 stygia which it resembles in many ways, but this latter species is larger, black, with larger eyes and with longer antennal scapes. Leptogenys diatra sp. n. DIAGNOSIS OF WORKER. Subpetiolar process a recurved tooth. Sides of head in front of eyes feebly concave. Small species, HL < 1-00. FURTHER DESCRIPTION. Holotype worker. TL 3:9, HL 0-86, HW 0-62, CI 72, SL 0-80, SI 129, PW 0:48, PH 0-44, PL 0-34, LPI 129, DPW 0-30, DPI 88. Head not broadened in front, broadest just behind the eyes. Sides of head behind eyes weakly convex, in front of eyes feebly concave. Occipital margin transverse, almost straight, with an extremely feeble convexity. Eyes with maximum diameter ca 0-10, equal to or very slightly less than the maximum width of the scape. Clypeus with an apically truncated, narrow, median triangular lobe, the anterior and lateral portions of which are translucent and lamellate. Mandibles short, closing tightly against the clypeus. Funicular segments 1-3 ca O*I4, 0°12, 0-12 respectively. Metanotal groove impressed and cross-ribbed. Subpetiolar process a broad-based, back-curved tooth or short spine. Gaster strongly constricted between the first and second segments. Standing hairs present upon all dorsal surfaces and on appen- dages. Unsculptured except for some fine punctures on the head anteriorly, between eye and frontal carinae, cross-ribbing in metanotal groove and transverse rugae on the propodeal declivity. Colour a uniform deep brown, shining. Holotype worker, CAMEROUN: no. 1726 (no further data) (G. Terron) (BMNH). This small species is closest related to stvator and its allies but is separated from them by its lack of a second subpetiolar process. Because of this diatra runs out in the key with nztida and its immediate relatives but the characters given there will serve to separate the species. Leptogenys honoria sp. n. DIAGNOSIS OF WORKER. Subpetiolar process double. Ventral surface of head not sculptured. Pronotal dorsum with shallow foveolate impressions. FURTHER DESCRIPTION. Holotype worker. TL 5:4, HL 1-08, HW 0-76, CI 70, SL 0-96, SI 126, PW 0-66, PH 0:64, PL 0-50, LPI 128, DPW 0:56, DPI 112. Head not broader in front than behind, the sides more or less straight in full-face view, the occipital margin very feebly concave, almost straight. Maximum diameter of eye ca 0-16, equal to or slightly greater than the maximum width of the scape. Median lobe of clypeus triangular and prominent. Mandibles short, closing tightly against the clypeus. Funicular segments I—3 Ca 0-16, 0-14, 0-12 respectively. Metanotal groove impressed and cross-ribbed. Subpetiolar process double, consisting of an anterior lobe, the ventral angle of which is acute, and a posterior tooth, the two separated by a distinct semicircular notch or impression. Node of petiole in profile increasing slightly in thickness from base to apex, noticeably higher than long; in dorsal view the node is much broader than long. Gaster strongly constricted between first and second segments. Standing hairs present on all dorsal surfaces but very short and sparse on the first and second gastral tergites, reduced to a row across the posterior margin of each segment. Mostly smooth and unsculptured but with the following: pronotal dorsum with a number of broad, shallow impressions; meso-metapleural suture cross-ribbed; lower portion of metapleuron and bulla of metapleural glands finely rugose; propodeal declivity transversely rugose. Colour uniform red-brown, the appendages lighter. Holotype worker, CAMEROUN: no. 1222 (no further data) (G. Terron) (BMNH). The form of the petiole and particularly its ventral process allies this species 288 B. BOLTON closely to strator and ankhesa, more distantly to diatra and buyssont. The characters given in the key and diagnoses will separate these related species. Leptogenys khaura sp. n. DIAGNOSIS OF WORKER. Dorsum of head finely and densely punctulate, pronotal dorsum unsculptured. Eyes minute, diameter < 0-04. Minute species, HL < 0-60, PW < 0°35. FURTHER DESCRIPTION. Holotype worker. TL 2:1, HL 0-52, HW 0:38, CI 73, SL o-42, SI 110, PW 0:30, PH 0:28, PL 0-16, LPI 175, DPW 0-22, DPK\137. Head not broadened in front, broadest behind eyes. Occipital margin feebly concave, sides of head shallowly convex. Eyes minute, with less than ro facets, their maximum diameter 0-04, considerably less than the maximum width of the scape. Clypeus narrowly triangular medially. Mandibles short, closing tightly against the clypeus. Funicular segments I-3 ca 0:08, 0:04, 0:05 respectively; the third, fourth and fifth segments as broad as or broader than long. Metanotal groove impressed and cross-ribbed but the ribs so short that the groove appears to have a series of punctures traversing the alitrunk. Node of petiole in dorsal view much broader than long, anteroposteriorly compressed, the anterior face slightly convex, the posterior slightly concave. Subpetiolar process broad and subrectangular in shape, without projections or teeth. Node in profile somewhat inclined forwards. Gaster strongly impressed between first and second segments. All surfaces of body with numerous standing hairs. Dorsum of head shallowly, finely and densely punctulate, the ventral surface unsculptured. Pronotal dorsum unsculptured, smooth and shining, as are the sides of the pronotum, mesono- tum, mesopleuron and gaster. Propodeal dorsum and sides and metapleuron mostly unsculp- tured but with some very fine, superficial regulation and a few punctures; propodeal declivity transversely rugose. Colour yellow-brown. Paratype worker. TL 2-3, HL 0:54, HW o-40, CI 74, SL 0-42, SI 105, PW 0:30, PH 0:30, PL 0-18, LPI 166, DPW 0:22, DPI 123. As holotype but propodeal sculpture much more reduced, the surface only superficially marked and with extensive smooth areas. Holotype worker, N1GERIA: Ile-Ife, 5.vili.t971 (J. T. Medler) (BMNH). Paratype. 1 worker as holotype but dated 21.vii.1g71 (MCZ, Cambridge). This species is the smallest member of the genus yet known from the Ethiopian Region. The combination of characters given in the diagnosis above will distin- guish it from all other small or minute species of the region. In size it is approached only by microps of the guineensis-group but the differences between their respective groups separate them. Other small species such as cryPtica and stygia are strongly sculptured and easily separable. Leptogenys mactans sp. n. (Text-fig. 17) DIAGNOSIS OF WORKER. Antennal scapes relatively very short, SI < 95. External margins of mandibles convex. Subpetiolar process with a posteroventral blunt tooth. FURTHER DESCRIPTION. Holotype worker. TL 4:5, HL 0-94, HW 0-76, CI 81, SL 0-68, SI 90, PW 0:58, PH 0°48, PL 0°42, LPI 114, DPW 0°34, DPI 81. Head not broader in front than behind, broadest just behind the eyes. Occipital margin shallowly concave, sides of head feebly convex. Eyes moderate, maximum diameter 0-16, slightly greater than the maximum width of the scape in full-face view. Eyes just failing to break the outline of the sides of the head in full-face view. Median lobe of clypeus short and broad, broadly truncated apically. Mandibles short and stout, their external margins REVISION OF GENUS LEPTOGENYS 289 markedly convex, closing tightly against the clypeus. Antennal scapes very short, scarcely surpassing the midpoint of the occipital margin. Funicular segments I-3 ca 0-12, 0-10, 0-IO respectively. Metanotal groove weakly impressed. Subpetiolar process a broad lobe with a short, rounded, posteroventral tooth. Gaster strongly constricted between first and second segments. All surfaces of head and body with numerous standing hairs. Unsculptured, completely smooth and shining apart from small hair-pits and a few faint transverse rugulae on the propodeal declivity. Colour red-brown. Paratype worker. TL 4:8, HL 0-98, HW 0-78, CI 80, SL 0-72, SI 92, PW 0:60, PH 0:46, PL o-40, LPI 115, DPW 0:34, DPI 85. Maximum diameter of eye 0:16. Otherwise as holotype. Holotype worker, SoutH AFRIcA: Cape Province, Somerset East, x.1930 (R. E. Turner) (BMNH). Paratype. I worker with same data as holotype (MCZ, Cambridge). The short scapes, broad head, strongly curved short mandibles and the shape of the subpetiolar process serve to separate this species from its relatives. In some respects mactans appears to be convergent upon the species of the processionalis- group of the Oriental and Indo-Australian regions but in these species the mandibles have numerous teeth and the petiole tends to be anteroposteriorly compressed to form a thick scale. In mactans the mandibles have only two teeth (apical and preapical) and the petiole and its ventral process suggest that the species is related to buyssont and its allies ankhesa and strator. Leptogenys nitida (F. Smith) Ponera nitida F. Smith, 1858 : 92. Syntype workers, Sourn AFrica: ‘Port Natal’ (= Durban) (BMNH) [examined]. Lobopelta nitida (F. Smith); Mayr, 1866 : 358. Leptogenys intermedia Emery, 1902 : 32. Syntype workers, SouTH AFRICA: Cape, Willowmore (Brauns) (MCSN, Genoa). Syn. n. Leptogenys tenuis Stitz, 1911 : 376, fig. 1. Syntype workers, ZAIRE: Lake Kivu (Mecklenburg) (probably in MNHU, Berlin). [Synonymy by Forel, 19124 : 52.] Leptogenys (Lobopelta) nitida var. adpressa Forel, 1914 : 214. Syntype workers, SouTH AFRICA: Grahamstown (Hewitt) (MHN, Geneva) [examined]. Syn. n. Leptogenys (Lobopelta) nitida var. aena Forel, 1914 : 215. Syntype workers, SouTH AFRICA: Willowmore (Brauns) (MHN, Geneva) [examined]. Syn. n. Leptogenys (Lobopelta) nitida var. gracilis Santschi, 1914b:11. Holotype worker, ZAIRE: Lake Kivu (Mecklenburg) [attributed to Stitz] (NM, Basle) [examined]. [Junior homonym of gracilis Emery, 1899 :271.] Syn. n. Leptogenys (Lobopelta) nitida st. insinuata Santschi, 1914b:11. Holotype worker, SouTH Arrica: Natal, Richmond, 24.iii.1905 (I. Trdgavdh) (location of types not known). Syn. n. Leptogenys (Lobopelta) nitida var. grandior Forel, 1915a : 335. Syntype workers, SouTH Arrica: Natal, Krantz Kloof (H. B. Marley) (MHN, Geneva) [examined]. Syn. n. Leptogenys (Lobopelta) nitida race brevinodis Forel, 1915@ : 335. Syntype workers, SouTH AFRICA: Cape Province (Brauns) (MHN, Geneva; MRAC, Tervuren) [examined]. [Junior homonym of brevinoda Emery, 1914.] Syn. n. Leptogenys (Lobopelta) nitida st. speculans Santschi, 1926a : 209. Syntype workers, ZAIRE: Lugumbe (Gérard) (NM, Basle) [examined]. Syn. n. Leptogenys (Lobopelta) brevinodis var. deflocata Santschi, 1926a:209. Syntype workers, MozaMBIQUE: Forét d’Amatonga, ii. 1917 (G. Arnold) (NM, Basle) [examined]. Syn. n. Leptogenys nitida subsp. capensis Baroni Urbani, 1971 : 360. [Replacement name for brevinodis Forel, 1915.] Syn. n. 290 B: BOLTON Note. Many of the above synonyms have been determined independently by Professor W. L. Brown and will be recorded by him in his forthcoming revision of the genera of tribe Ponerini. DIAGNOSIS OF WORKER. Large species, HL > 0-90, with SI > 120. Eyes > 0-11 maximum diameter. Dorsal surfaces unsculptured except for hair-pits. FURTHER DESCRIPTION. Worker. TL 4:6—5+7, HL 0:92-1:20, HW 0:66-0-88, CI 69-75, SL 0:86-1:08, SI 120-130, PW 0:52-0:70, PH 0:44-0:60, PL 0:40-0:48, LPI 110-125, DPW 0+32—0:40, DPI 80—95 (12 measured). Head not broader in front than behind, broadest at about the level of the eyes. Occipital margin straight to feebly convex, sides of head shallowly convex. Maximum diameter of eyes ca 0:12-0:22, usually equal to or slightly greater than the maximum width of the scape in full-face view, rarely otherwise. Median lobe of clypeus triangular, blunt apically. Mandibles closing tightly against the clypeus, their outer margins somewhat concave. Funicular segments I—3 Ca 0*12—0'20, 0-10—-0'18, 0-10—0-16 respectively. Subpetiolar process a simple lobe or with a small posteroventral tooth, very variable in size. Gaster strongly constricted between first and second segments. Short, standing hairs numerous upon all dorsal surfaces of head and body. Unsculptured, smooth and shining except for minute hair-pits and usually with a few tranverse rugae on the propodeal declivity. Colour very variable, ranging from jet-black to light brown with all intermediate shades. In terms of colour and size nitida is probably the most variable species of the genus in the Ethiopian Region; no two series examined in the course of this survey were exactly alike in all respects. The larger or more darkly coloured individuals of nitida are unlikely to be con- fused with any other species but the smaller, more lightly coloured specimens may be mistaken for castanea. The characters given in the key and the notes under castanea will serve to distinguish the two species. MATERIAL EXAMINED. KENYA: (no data). ZAMBIA: Kipushi (H. S. Evans). Botswana: Malepi Stream (G. Arnold); Mamathes (C. Jacot-Guillarmod). RHODESIA: Chirinda Forest (G. Arnold); Salisbury (G. H. Bunzli); Vumba Mts (G. Arnold); Umtali (G. Arnold). SouTH AFRICA: Orange Free State (Wroughton); Transvaal, Piet Retief (H. Brauns); Ladismith (H. Brauns); Cape. Prov., Oudtshoorn (B. Malkin); George (H. Brauns); Cape Prov., Montagu Pass (A. Mackie); Katberg (R. E. Turner); Grahamstown (J. Hewitt); Mossel Bay (R. E. Turner); Natal (G. B. King); Durban (G. Arnold), (C. B. Cooper), (H. B. Marley); Krantz Kloof (H. B. Marley); Natal, Slievyre (Haviland); Natal, Estcourt (G. Arnold); Natal, Eshowe (H. B. Marley). Leptogenys piroskae Forel (Text-fig. 15) Leptogenys (Lobopelta) piroskae Forel, 1910a: 247. Syntype workers, Erniopia: Eritrea, Ghinda (Escherich) (NM, Basle) [examined]. DIAGNOSIS OF WORKER. Larger species, HL > 0:90, with SI > 120. Maximum diameter of eye 100), the anterior and posterior faces almost parallel, the dorsum weakly convex. In dorsal view the node slightly broader behind than in front, with a convex anterior face and almost straight posterior face. Gaster strongly constricted between first and second segments. Head and body everywhere smooth and shining, unsculptured apart from hair-pits and except for the propodeal declivity which is transversely rugose and the metanotal groove and meso-metanotal suture, which are cross-ribbed. Standing hairs present on all dorsal surfaces. Colour uniform orange-brown or light brown. Leptogenys strator sp. n. DIAGNOSIS OF WORKER. Subpetiolar process double, with separate anterior and posterior portions. Propodeal dorsum rugose. FURTHER DESCRIPTION. Holotype worker. TL 5:8, HL 1:22, HW 0-86, CI 70, SL not measurable, PW 0:72, PH 0:77, PL 0°54, LPI 133, DPW 0°52, DPI 96. Head not broader in front than behind, broadest behind the eyes. Occipital margin shallowly concave, sides of head feebly convex. Eyes with maximum diameter ca 0-16 (antennae missing from all members of type-series). Median lobe of clypeus bluntly triangular. Mandibles short, closing tightly against the clypeus. Mesonotum broader than long; metanotal groove deeply impressed and cross-ribbed. Node of petiole in dorsal view about as broad as long, with the anterior and posterior faces almost parallel and the sides diverging posteriorly. In profile the node is broadest basally and tapers slightly above. Subpetiolar process double, complex, consisting of a broad lobe anteriorly, the posteroventral corner of which is acute, and a secondary blunt, short tooth, situated behind the lobe and separated from it by a distinct notch. Gaster strongly constricted between first and second segments. Standing hairs present on all dorsal surfaces, sparse (some may have been lost by abrasion). Dorsum of head finely and quite densely punctured in the space between the eye and frontal carinae, the remainder unsculptured apart from hair-pits. Ventral surface of head with large, shallow, poorly defined punctures, giving the surface an undulate appearance. Pronotal and propodeal dorsa rugose, the rugae broad, shallow and rounded, the propodeum more strongly sculptured than the pronotum. Meso-metapleural suture strongly cross-ribbed, the metapleuron and sides of the propodeum with some rugulation. Gaster smooth. Colour red-brown. Paratype workers. TL 5:8-5-9, HL 1:24, HW 0°88, CI 71, (antennae missing), PW 0-70-0°74, PH 0:70-0:72, PL 0:54, LPI 129-130, DPW 0:54, DPI 100 (2 measured). Diameter of eye ca O-14—0'10. Holotype worker, ZAIRE: (‘Belg. Congo’ on specimen) Mikeno, 24.vii.1938 (C. P. Haskins) (MCZ, Cambridge). Paratypes. 2 workers, same data as holotype (BMNH; MCZ, Cambridge). Closest related to ankhesa, strator is distinguished by its sculpturation, which is absent from the former, and by the shape of the node which, in profile, is broader basally whereas in ankhesa the node is slightly narrower basally than it is above. 292 B. BOLTON Leptogenys striatidens sp. n. DIAGNOSIS OF WORKER. Mandibles densely striate. Head and pronotum covered with dense superficial reticulation. FURTHER DESCRIPTION. Holotype worker. TL 5:6, HL 1°36, HW 0-92, CI 68, SL 1°14, SI 124, PW 0:76, PH 0-66, PL 0:54, LPI 122, DPW 0-52, DPI 96. Head broadest in front, increasing slightly in width from back to front, the sides very weakly convex, the occipital margin feebly concave. Eyes small, their maximum diameter ca 0-14, less than the maximum width of the antennal scape. Median lobe of clypeus long, blunt apically and with weakly convex sides. Mandibles closing tightly against the clypeus, the angle at the junction of the inner and apical margins concealed by the lobe of the clypeus. Funicular segments I-3 ca 0-16, 0°18, 018 respectively. Metanotal groove broad and shallow, very finely cross-ribbed. Gaster strongly constricted between first and second segments. Dorsal surfaces of head, body and appendages with numerous standing hairs. Mandibles finely and densely but very distinctly longitudinally striate. Dorsal surfaces of head and alitrunk covered by a fine, superficial reticulation. Sides of head below level of eyes similarly sculptured but also with elongate, shallow impressions. Pronotum with the reticulation petering out on the sides so that the lower halves are smooth. Lower halves of meso- and metapleurae rugose; propodeal declivity transversely rugose. Sides of petiole superficially reticulate but the posterior half of the dorsum smooth. Gaster smooth and shining. Colour black with faint traces of bluish opalescence; the appendages brown. Holotype worker, CAMEROUN: Mt Kala, 820 m, 22.11.1973 (G. Terron) (BMNH). This species is unique amongst the members of the nitzda-group known at present as it is the only one with strong sculpturation over the entire head and alitrunk, and possesses distinctly striate mandibles. Leptogenys stygia sp. n. DIAGNOSIS OF WORKER. Smaller species, HL < 0-80. Head densely reticulate-punctate. Pronotum sculptured. Diameter of eye 0-08—0-09; SI in the range 119-122. FURTHER DESCRIPTION. Holotype worker. TL 3:2, HL 0-70, HW 0-48, CI 68, SL 0°58, SI 121, PW 0-38, PH 0-38, PL 0-26, LPI 146, DPW, 0-22, DPI 85. Head not broader in front than behind, broadest behind the eyes. In full-face view the occipital margin straight to feebly concave; sides of head weakly convex. Maximum diameter of eye ca 0:08, in full-face view less than the maximum width of the scape. Median lobe of clypeus narrowly triangular. Mandibles short, closing tightly against the clypeus. Funicular segments I-3 Ca 0:10, 0:08, 0-08 respectively. Metanotal groove shallow, only weakly impressed, less distinct than the promesonotal suture. Subpetiolar process a broad, rounded lobe, without projections or teeth. Gaster strongly constricted between first and second segments. Stand- ing hairs numerous upon all surfaces of head and body. Dorsum of head densely and quite coarsely reticulate-punctate, the ventral surface with scattered foveolate punctures separated by smooth, shining interspaces. Dorsal alitrunk with scattered punctures and fine rugulae, densest on the propodeum. Propodeal declivity transversely rugose. Sides of alitrunk sculptured as dorsum. Gaster shining, unsculptured apart from hair-pits. Black, the legs and scapes brown, the funicular segments yellow-brown. Paratype workers. TL 3:0-3:2, HL 0-68-0-74, HW 0-48-0-52, CI 68-71, SI 119-122, PW 0°38—0'42, PH 0:36-0'40, PL 0:24-0:28, LPI 143-153, DPW 0:20-0:24, DPI 83-92 (7 measured). Maximum diameter of eye ca 0-08-o0-09. Otherwise as holotype. Holotype worker, NIGERIA: Gambari, 3.xi.1969, ‘under fallen banana stem’ (B. Bolton) (BMNA). REVISION OF GENUS LEPTOGENYS 293 Paratypes. 7 workers, with same data as holotype (BMNH; MCZ, Cambridge; NM, Basle). Most closely related to cryptica, from which it differs in size and colour, cryptica being smaller and yellow-brown with relatively shorter antennal scapes and smaller eyes. The specimens of stygia described above were discovered in a small piece of wet-rotten wood under a fallen and rotting bananastem. The species is also present in Cameroun as G. Terron (UFC, Yaounde) has sent me a specimen recently. REVIEW OF SPECIES OF MALAGASY REGION As the Malagasy fauna is so poorly known and collections of the genus from this region are virtually non-existent, a formal revision of the species has not been under- taken. Instead a list and provisional key are presented together with notes on the groups and types of species which I have examined, in the hope that the review will be of some benefit to further studies of the regional fauna. LIST OF SPECIES Species shared with the Ethiopian Region (falcigera, maxillosa, stuhlmanni) are not included. truncatirostris-group arcirostris Santschi ridens Forel truncatirostris Forel attenuata-group angusta (Forel) coerulescens Emery grandidieri Forel saussurei-group acutirostris Santschi stat. n. oswaldi Forel saussurei (Forel) incisa-group alluaudi Emery antogilensis Emery gracilis Emery incisa Forel incisa var. imerinensis Forel syn. n. incisa subsp. suavensis Emery syn. n. voeltzkowi Forel PROVISIONAL KEY TO SPECIES (Workers) The key is based for the greater part on type-specimens and original descriptions only and consequently should be used with some care as no indication of degree of 294 B. BOLTON variability of the species is known. All species are from Madagascar except where otherwise stated. 1 Standing hairs absent from dorsum of head, alitrunk and first two gastral tergites. Cuticle densely shagreened, with dense, pruinose pubescence. ; ; ; 2 — Standing hairs present on some or all dorsal surfaces. Cuticle not densely shagreened, without pruinose pubescence : 3 2 Node of petiole in dorsal view yekeread broader than long, DPW > 105 (pantropical tramp species) . ; maxillosa — Node of petiole in dorsal view ‘at least as ‘long : as broad, usually distinctly longer than broad, DPW too or less (pantropical tramp species) . : : . falcigera 3 Anterior clypeal margin broadly rounded or transverse, not projecting into a triangular lobe medially. Mandibles elongate, narrow, but capable of closing against the clypeus. Head strongly broadened in front, the Lg not breaking the outline of the sides of the head in full-face view ‘ 4 — Anterior clypeal margin projecting into a triangular lobe medially. Mandibles short or elongate, but if elongate then not capable of closing against the clypeus. Head usually not broadened anteriorly, the eyes usually breaking the outline of the sides of the head in full-face view . : : : : : ; : ‘ 6 4 Clypeus without a median, longitudinal carina : : j . truncatirostris — Clypeus with a median, longitudinal carina : j 5 5 Antennal scapes both relatively and absolutely shorter, SL < I: 75, SI < 12 5 arcirostris — Antennal scapes both relatively and absolutely longer, SL > 1°75, SI > 135 ridens 6 Mandibles short, not linear, capable of closing tightly against the clypeus; without a gap between themselves and the clypeus when closed . 7 — Mandibles elongate and linear, not capable of closing against the clypeus: with a large gap between themselves and the clypeus when closed . : 12 7 Dorsum of head and alitrunk unsculptured apart from minute pits from which hairs arise . : : , : : 8 — Dorsum of head, alitrunk, or both, distinctly sculptured : , : ; : Io 8 Propodeal declivity smooth and shining, not sleet rugose . . d angusta — Propodeal declivity transversely rugose . : 9 9 Smaller species, HL ca 1-40; black, without ‘bluish reflections or opalescence. Median lobe of clypeus not distinctly longitudinally striate : ; grandidieri — Larger species, HL > 1-50; black with bluish reflections or opalescence. Median lobe of clypeus longitudinally striate. : ‘ coerulescens 1o Dorsum of head very densely and very finely reticulate-rugose, the interspaces minutely punctulate, without smooth shining areas. Small species, HL < 2-20, diameter of eye ca 0°44 ; oswaldi — Dorsum of head with scattered large punctures or foveolae, the spaces between which may have some faint striae but are mostly smooth and sSpretnsk ance species, HL > 2-50, diameter of eye > 0-50 . : It Ir Posterior border of petiole strongly emarginate in dorsal view. Larger species, HL > 2°85, with larger eyes, maximum diameter > 0-70. . : . saussurei — Posterior border of petiole not emarginate in dorsal view. Smaller species, HL < 2:80, with smaller eyes, maximum diameter < 0-65 ; acutirostris 12 Dorsum of head and first gastral tergite unsculptured apart from pits from which hairs arise. Gaster not constricted between first and second segments. (East Africa, Comoro Islands) ‘ ; stuhlmanni — Dorsum of head sculptured, first gastral tergite usually with at least a fine, superficial reticulation. Gaster constricted between first and second segments incisa-group [including incisa, alluaudi, voeltzkowi, gracilis, antongilensis} REVISION OF GENUS LEPTOGENYS 295 THE TRUNCATIROST RIS-Grovup Mandibles elongate but closing tightly against the clypeus, with considerable overlap. Clypeus with a fringing lamella on the anterior margin, not projecting into a triangular lobe medially, either broadly rounded or transverse. Eyes large, their maximum diameter greater than the maximum width of the scape. Second funicular segment slightly longer than the third. Head broadened in front; the eyes not breaking the outline of the sides of the head in full-face view. Entirety of head and body sculptured, either reticulate-punctate, densely rugulose or densely but irregularly punctulate. Standing hairs present on all dorsal surfaces. Gaster strongly constricted between first and second segments. The three species of this small group formerly constitued the subgenus Machaero- genys, now placed in the synonymy of Leptogenys (see p. 240). Species included Leptogenys arcirostris Santschi Leptogenys (Machaerogenys) arcirostris Santschi, 1926b : 25. Holotype worker, MADAGASCAR; Moramanga (Descarpentries) (NM, Basle) [examined]. Leptogenys ridens Forel Leptogenys ridens Forel, 19106 : 16. Holotype worker, Mapacascar: Fort Dauphin (Sikora) (MHN, Geneva) [examined]. Leptogenys truncatirostris Forel Leptogenys truncatirostris Forel, 1897 : 195, fig. 2. Syntype workers, MapaGascar: Nossi Bé (Voeltzkow) (MHN, Geneva; NM, Basle) [examined]. Of the three species included in the group, trwncatirostris is easily distinguished as it is larger (HL 1-88, HW 1-66, CI 88, SL 1-86, SI 112, PW 1-10) and lacks a median, longitudinal clypeal carina, which is present in all other species of both the Malagasy and Ethiopian regions. Also in truncatirostris the anterior margin of the clypeus is transverse, not convexly arched, and the mandibles are elongate and strongly curved basally. Each blade is broadened in the distal half of its length and then tapers to a long acute apex. A single, strongly developed preapical tooth is present which is curved in the direction of the mandibular apex. The two remaining species, arcirostris and ridens, are smaller (ridens: HL 1-70, HW 1:36, CI 80, SL 1-98, SI 145, PW 1:08; arcirostris: HL 1-48, HW 1-28, CI 86, SL 1-50, SI 117, PW 0-96) with a median clypeal carina and a convex anterior clypeal margin. The mandibles in both species are elongate, curved slightly along their length, increasing slightly in thickness from base to apex and terminating in an abrupt apical surface. Preapical teeth are not present, the apical surface joins the inner margin through a curve. The two are easily separated as videns is a larger species with relatively much longer scapes, as indicated in the measurements above. 296 B. BOLTON THE ATTENUATA-GRovuP Characteristics of the group given on p. 274. Species included Leptogenys angusta (Forel) Lobopelta angusta Forel, 1892: 519. Syntype workers, MapAGascaR: Imerina, Andrangoloaka Forest (Sikora) (MHN, Geneva) [examined]. Leptogenys coerulescens Emery Leptogenys coerulescens Emery, 18956 : 339. Syntype workers, MapaGascar: Diego Suarez, 1893 (Ch. Alluaud) (probably in MCSN, Genoa). Leptogenys grandidieri Forel Leptogenys (Lobopelta) grandidieri Forel, 1910b:17. Holotype worker, MapaGascar: Fort Dauphin (Sikora) (MHN, Geneva) [examined]. L. angusta is a large species, with CI and SI within the range of attenuata, which it resembles (angusta: HL 2:24, HW 1-34, CI 60, SL 2:56, SI 191). As in attenuata the petiole in dorsal view is much longer than broad, strongly narrowed in its anterior half. The smooth integument has a scattered, bluish opalescence and the propodeal declivity is smooth, without transverse rugae. The median lobe of the clypeus is not longitudinally striate. L. coerulescens appears to be related to angusta and also has bluish opalescence, but in this species the propodeal declivity is transversely rugose and the median clypeal lobe is longitudinally striate. The last known species of the group, grandidieri, is smaller (HL 1-40, HW 0:92, CI 66, SL 1-30, SI 141) and on the whole appears to be more closely related to crassinoda than to any of the above. The petiole in dorsal view is longer than broad, but is only slightly broader behind than in front. The propodeal declivity is transversely rugose but the median lobe of the clypeus lacks striae, apart from a few on the extreme lateral portions. THE SAUSSU REI-Group Mandibles short, closing tightly against the clypeus. Eyes large, their maximum diameter greater than the maximum scape width. Second funicular segment only slightly longer than the third. Head not or only slightly broadened anteriorly. Dorsum of head and alitrunk sculptured. Of the three species in the group two are closely related whilst the third, oswaldi, may not actually belong to the group as the gaster is constricted between the first and second segments, a feature much reduced in saussurei and acutirostris. All three species are large. REVISION OF GENUS LEPTOGENYS 297 Species included Leptogenys acutirostris Santschi stat. n. Leptogenys saussurei st. acutirostris Santschi, 1912 : 150. Holotype worker, MADAGASCAR (Le Moult) (NM, Basle) [examined]. Leptogenys oswaldi Forel Leptogenys o’swaldi Forel, 1891a : 119, pl. 4, fig. 2. Holotype worker, MADAGASCAR: 30 milles au Nord-Ouest de Tamatave (O’Swald) (MHN, Geneva) [examined]. Leptogenys saussurei (Forel) Lobopelta saussurei Forel, 1891a : 121. Holotype worker, Mapacascar: Mahanoro, céte Est de Madagascar (Saussure) (MHN, Geneva) [examined]. L. oswaldi is the smallest species of this group (HL 2-08, HW 1:68, CI 80, SL 2-06, SI 124, PW 1-32) and the head is relatively broad. The gaster is strongly con- stricted between the first and second segments and the eyes have a maximum dia- meter ca 0-44. Dorsum of head densely sculptured with a fine rugoreticulum, the interspaces of which are minutely punctulate. The remaining two species are larger and less strongly sculptured, with narrower heads and the gastral constriction very weak indeed. The larger of the two, saussurei (HL 2-94, HW 1-98, CI 67, PW 1°84; (antennae missing from holotype) has an ocular diameter of 0-80 at maximum. The clypeus and gena to the level of the eye is longitudinally rugose but the dorsum of the head is coarsely punctured, the punctures becoming larger, shallower and more widely spaced posteriorly, separated by smooth interspaces. Petiole in dorsal view has the posterior margin strongly concave, the posterolateral angles prominent. L. acutirostris is similarly sculptured but is smaller (HL 2-76, HW 1-84, CI 63, SL 2-34, SI 127, PW 1-60) with smaller eyes, maximum diameter ca 0-58. The posterior margin of the petiole in dorsal view is not excised as in saussuret. Tue TINCISA-Grovup Mandibles elongate, curved, not capable of closing against the clypeus. Eyes medium to large; maximum diameter greater than maximum width of scape, if only slightly so. Second funicular segment longer than third. Head usually broadest at about the level of the eyes, the sides convex. Gaster constricted between first and second segments. Dorsum of head densely sculptured, either reticulate-punctate or with a fine, dense rugoreticulum with punctulate interspaces. First gastral tergite usually sculptured, at least with a superficial reticulation. Dorsal alitrunk sculptured. Species included Leptogenys alluaudi Emery Leptogenys alluaudi Emery, 18956 : 338. Holotype worker, MapaGascar: Diego Suarez, iv-vili. 1893 (Alluaud) (probably in MCSN, Genoa). 298 B. BOLTON Leptogenys antongilensis Emery Leptogenys incisa var. antongilensis Emery, 1899 : 272, fig. Holotype worker, MADAGASCAR: Baia di Antongil, 1897-98 (A. Mocquerys) (probably in MCSN, Genoa). Leptogenys antongilensis Emery; Emery, 1911 : 100. [Raised to species. ] Leptogenys gracilis Emery Leptogenys gracilis Emery, 1899: 271. Syntype workers, MADAGASCAR: Bia di Antongil, 1897-98 (A. Mocquerys) (probably in MCSN, Genoa). Leptogenys incisa Forel Leptogenys incisa Forel, 1891a : 113, pl. 4, figs 1, 1a. Syntype workers, MADAGASCAR: Montagne de Lokobé, a Nosibé (O’Swald) (location of types not known). Leptogenys incisa var. imerinensis Forel, 1891b : 242. Syntype workers male, MADAGASCAR: Forét d’Andrangoloaka (Sikora) (MHN, Geneva) [examined]. Syn. n. Leptogenys incisa subsp. suarensis Emery, 1895) : 338. Syntype workers, MADAGASCAR: Diege Suarez, 1893 (Alluaud) (probably in MCSN, Genoa). Syn. n. Leptogenys voeltzkowi Forel Leptogenys voeltzkowi Forel, 1897 : 194. Syntype workers, MapaAcascar: Nossi Bé (Voeltzkow) (MHN, Geneva; MCZ, Cambridge) [examined]. The few specimens and types of this group which I have been able to examine show that the species are closely related. It will not be possible to sort the species accurately until all the types can be examined and compared, and until some other collections have been made. A SPECIES EXCLUDED FROM LEPTOGENYS Pachycondyla jonesii (Forel) comb. n. Lobopelta jonesii Forel, 1891a : 219. Syntype males, MapaGascar: Forét d’Andrangoloaka (Sikora) (MHN, Geneva) [examined]. This species, known from two males, was originally placed by Forel in Lobopelta even though the pretarsal claws were not pectinate, as he pointed out in the original description. In fact, each claw has a single small tooth at about the midlength. Other definitive characters besides this, leading to the exclusion of jonesit from Leptogenys and its placement in Pachycondyla, include the curved, spiniform exten- sion of the pygidium (absent from Leptogenys), the absence of notauli (usually present in Leptogenys), and the presence of an anal lobe on the hind wing (absent from Leptogenys). ACKNOWLEDGEMENTS I would like to express my thanks and gratitude to the following, who provided REVISION OF GENUS LEPTOGENYS 299 types or other material during the course of this study, and which greatly facilitated its completion. Dr Cl. Besuchet (MHN, Geneva); Prof. W. L. Brown (Cornell University, Ithaca) ; Dr J. Decelle (MRAC, Tervuren); Mrs M. Favreau (AMNH, New York); Dr M. Fischer (NM, Vienna); Dr E. K6nigsmann (MNHU, Berlin); Mr F. C. de Moor (NM, Bulawayo) ; Dr A. J. Prins (SAM, Cape Town); Mr E. Taylor (UM, Oxford); Dr G. Terron (UFC, Yaounde); Dr C. Baroni Urbani (NM, Basle); Mm J. C. Weulersse( MNHN, Paris); Ms J. C. White (MCZ, Cambridge). REFERENCES ANDRE, E. 1892. Matériaux myrmécologiques. Revue Ent. 11 : 45-56. ARNOLD, G. 1915. A monograph of the Formicidae of South Africa, part 1. Ann. S. Afr. Mus. 14 : 1-159, 1 pl. 1926. Ibidem: appendix. Ann. S. Afr. Mus. 23 : 191-295, 23 figs. 1954. New Formicidae from Kenya and Uganda. Annis Mus. r. Congo Belg. (N.S. in 4°) Zool. 1 : 291-295, 6 figs. BERNARD, F. 1952. La réserve naturelle intégrale du Mt. Nimba, part 11. Hyménopteéres, Formicidae. Mém. Inst. fr. Afr. noive 19 : 165-270, 3 pls, 15 figs. Bineuam, C. T. 1903. Fauna of British India, including Ceylon and Burma. Hymenoptera 2, Ants and Cuckoo Wasps. London. 506 pp., 161 figs, 1 pl. Botton, B. 1973. The ant genera of West Africa: a synonymic synopsis with keys. Bull. Br. Mus. nat. Hist. (Ent.) 27 : 317-368, 1 fig. Brown, W.L. 1963. Characters and synonymies among the genera of ants, part 3. Some members of the tribe Ponerini. Breviova no. 190 : 1-10. DONISTHORPE, H. St. J. K. 1948. Microbolbos testaceus, a new genus and species of ponerine ant. Entomologist 81 : 170-171. Durour, L. 1864. Note sur une nouvelle espéce de fourmi (Formica vinsonnella). Annls Soc. ent. Fr. (4) 4: 210. Emery, C. 1892. Sopra alcune formiche raccolte dall’Ingegnere L. Bricchetti Robecchi nel paese dei Somalia. Annali Mus. civ. Stor. nat. Giacomo Doria (2) 12 [32] : 110-122, 3 figs. 1894. Mission scientifique de M Ch. Alluaud aux iles Séchelles (mars, avril, mai 1892). Annls Soc. ent. Fr. 63 : 67-72, figs. 1895a. Voyage de M E. Simon dans |’ Afrique australe (janvier—avril 1893). Formicides. Annls Soc, ent. Fr. 64 : 15-56, pl. 2. 18950. Mission scientifique de M Ch. Alluaud dans le territoire de Diego-Suarez (Madaga- scar Nord) (avril—aofit 1893). Annis Soc. ent. Belg. 39 : 336-345, 2 figs. 1899. Formiche di Madagascar raccolte dal Sig. A. Mocquerys nei pressi della Baia di Antongil (1897-98). Boll. Soc. ent. ital. 31 : 263-290, figs. 1901. Spicilegio mirmecologico. Boll. Soc. ent. ital. 33 : 57-63. — 1902. Note mirmecologiche. Rc. Sess. Accad. Sci. Ist. Bologna (N.S.) 6 : 22-34. 1911. In Wytsman, Genera Insectorum. Hymenoptera, Fam. Formicidae, subfam. Ponerinae. fasc. 118 : 125 pp., 3 pls. Foret, A. 1891a. In Grandidier, Histoire physique, naturelle et politique de Madagascar 20. Histoire naturelle des Hyménoptéres, part 2, Les Formicides: 237 pp., 7 pls. Paris. 1891b. Ibidem. Supplement au 28° fascicule: 229—-280. 1892. Nouvelles espéces de formicides de Madagascar (recoltées par M. Sikora) Annis Soc. ent. Belg. 36 : 516-535. 1897. Ameisen aus Nossi-Be, Majunga, Juan de Nova (Madagaskar), den Aldabra- Inseln und Sansibar. Gesammelt von Herrn Dr. A. Voeltzkow aus Berlin. Mit einem anhang iiber die von Herrn Privatdocenten Dr A. Brauer in Marburg auf den Seychellen und von Herrn Perrot auf Ste. Marie (Madagaskar) gesammelten Ameisen. Abh. Senckenb. naturforsch. Ges. 21 : 185-208, 3 figs. 300 B. BOLTON 1g01a. Nouvelles espéces de Ponerinae. Revue suisse Zool. 9 : 325-353. 1901b. Formiciden des Naturhistorischen Museums zu Hamburg. Neue Calyptomyrmex, Dacryon, Podomyrma und Echinopla-Arten. Mitt. Naturh. Mus. Hamburg 18: 43-82. 1907a. Fourmis d’Ethiopie; recoltées par M le Baron Maurice de Rothschild en 1905. Revue Ent. 26 : 129-144. o 1907b. In Voeltzkow, Reise in Ostafrika in den Jahren 1903-1905. Wissenschaftliche Ergebnisse Bd. 2. Systematische Arbeiten 2 : 39-178. 1910a@. Ameisen aus der kolonie Erythraa. Gesammelt von Prof. Dr K. Escherich. Zool. Jb. Syst. 29 : 243-274. 1910b. Glanures myrmécologiques. Annls Soc. ent. Belg. 54 : 6-32. 1912a. E/inige neue und interessante Ameisenformen aus Sumatra etc. Zool. Jb. Syst. 15 suppl.: 51-78. 1912b. The Percy Sladen Trust expedition to the Indian Ocean in 1905. 4 no. ITI. Fourmis des Seychelles et des Aldabra, regues de M. Hugh Scott. Tvans. Linn. Soc. Lond. 15 : 159-167. 1913a. Fourmis de Rhodesia etc., recoltées par M. G. Arnold, le Dr H. Brauns et K. Fikendey. Avnnls Soc. ent. Belg. 57 : 108-147. 1913b. Quelques fourmis des Indes, du Japon et d’Afrique. Revue suisse Zool. 21: 659-673, I fig. 1913c. Ameisen aus Rhodesia, Kapland usw. Gesammelt von Herren G. Arnold, Dr H. Brauns und anderen. Dt. ent. Z. 1913 (Beiheft) : 203-225. 1914. Formicides d’Afrique et d’Amerique nouveaux ou peu connus. Bull. Soc. vaud. Sci. nat. 50 : 211-288. 1915a. Formicides d’Afrique et d’Amerique nouveaux ou peu connus, part 2, Bull. Soc. vaud. Sci. nat. 50 : 335-364. 19156. Results of Dr E. Mjoberg’s Swedish scientific expeditions to Australia 1910-1913. 2 Ameisen. Ark. Zool. 9 (16) : 1-119, 6 figs. 1916. Fourmis du Congo et d’autres provenances recoltées par MM Hermann Kohl, Luja, Mayné etc. Revue suisse Zool. 24 : 397-460, 7 figs. Mayr, G. 1862. Myrmecologische studien. Verh. zool.-bot. Ges. Wien 12 : 649-776, I pl. 1886. Notizen iiber die Formiciden-"Sammlung des British Museum in London. Verh. z0ol.-bot. Ges. Wien 36 : 353-368. 1893. Formiciden von Herrn Dr F. Stuhlmann in Ost-Afrika gesammelt. Jb. hamb. wiss. Anst. 10 : 193-201. . Prins, A. J. 1965. Notes on African Formicidae 1. J. ent. Soc. sth. Afr. 27 : 153-159, 10 figs. RocGErR, J. 1861. Die Ponera-artigen Ameisen. Berl. ent. Z. 5 : 1-54 1863. Verzeichniss der Formiciden-Gattungen und Arten. Berl. ent. Z. 7 suppl. : 1-65. SANTSCHI, F. 1912. Fourmis d’Afrique et de Madagascar. Avnnls Soc. ent. Belg. 56 : 150-167, figs A—D. 1914a. Formicides de |’Afrique occidentale et australe du voyage de M le Professeur F. Silvestri. Boll. Lab. Zool. gen. agr. Portici 8 : 307-385, 34 figs. 1914b. Meddelanden fran Géteborgs Musei Zoologiska Afdeling no. 3. Fourmis du Natal et du Zululand recoltées par le Dr I. Tragardh; avec un appendice: Notes biologiques par I. Tragardh. Gdéteborgs K. Vetensk.-o. VitterhSamh. Handl. 15 : 1-47, 10 figs. 1g14c. In Voyage de Ch. Alluaud et R. Jeannel en Afrique orientale (1911-1912). Insectes, Hyménoptéres 2 Formicidae: 43-148, 2 pls, 30 figs. Paris. 1915. Nouvelles fourmis d’Afrique. Annls Soc. ent. Fr. 84 : 244-282, 15 figs. 1917. Fourmis nouvelles de la Colonie du Cap, du Natal et de Rhodesia. Amnnls Soc. ent. Fr. 85 (1916) : 279-296. 1924. Descriptions de nouveaux formicides africains et notes diverses 2. Revue Zool. Bot. afr. 12 : 195-224, 10 figs. 1926a. Descriptions de nouveaux formicides Ethiopiens, part 3. Revue Zool. Bot. afr. 13 : 207-267, 6 figs. 1926b. Trois notes myrmécologiques. Annls Soc. ent. Fr. 95 : 13-28. REVISION OF GENUS LEPTOGENYS 301 1932. Formicides sud-africains. Soc. ent. Fr. livre du centenaire, 1932 : 381-392, 12 figs. 1937. Fourmis du Congo et régions limitrophes. Revue Zool. Bot. afr. 30 : 71-85, 6 figs. SMITH, F. 1857. Catalogue of the Hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and at Singapore by A. R. Wallace. J. Linn. Soc. (Zool.) 2 : 42-130, 2 pis. 1858. Catalogue of Hymenopterous Insects in the collection of the British Museum, part 6 Formicidae: 216 pp., 14 pls. 1879. Descriptions of new species of Aculeate Hymenoptera collected by the Rev. Thos. Blackburn in the Sandwich Islands. J. Linn. Soc. (Zool). 14 : 674-685. Stitz, H. 1910. Westafrikanische Ameisen 1. Mitt. Zool. Mus. Berl. 5 : 125-151, 11 figs. 1911. Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika Expedition, 1907-1908. Band 3, Zoology 1, Lieferung 9 Formicidae: 375-392, 8 figs. Leipzig. Taytor, R. W. 1969. The identity of Dorylozelus mjobergi Forel. J. Aust. ent. Soc. 8 : 131- 133, 2 figs. URBANI, C. BARONI. 1971. Einige homonymien in der familie Formicidae. Mitt. schweiz. ent. Ges. 44 : 360-362. WEBER, N. A. 1942. New doryline, cerapachyine and ponerine ants from the Imatong Mts., Anglo-Egyptian Sudan. Proc. ent. Soc. Wash. 44 : 40-49, pl. 5. WHEELER, W.M. 1922. Ants of the Belgian Congo, part 2. Ants collected by the American Museum Congo Expedition. Bull. Am. Mus. nat. Hist. 45 : 39-269, 23 pls., 76 figs. Witson, E. O. 1955. The status of the ant genus Microbolbos Donisthorpe. Psyche, Camb. 62 : 136. Witson, E. O. & TayLor, R. W. 1967. The ants of Polynesia. Pacif. Insects Monogr. 14 : 1-109, 84 figs. 302 B. BOLTON ove & Fics 1-15. 1. Anterior portion of head and left mandible of Leptogenys pavesit. 2. Anterior portion of head and left mandible of L. falcigera. 3. Outline of head of L. jeanneli, pilosity and antennae omitted. 4. Outline of head of L. zapyvxis, pilosity and antennae omitted. 5. Base of left mandible in L. camerunensis and allies. 6. Base of left mandible in L. stuhimanni and allies. 7-15. Outlines of heads of (7) L. camerunensis ; (8) L. vegis; (9) L. elegans (offset shows size of 3rd funicular segment); (10) L. ravida (offset shows size of 3rd funicular segment); (11) L. tribolata; (12) L. mastax; (13) L. occidentalis ; (14) L. sulcinoda; (15) L. piroskae; pilosity and antennae omitted, mandibles of 12-15 slightly opened. REVISION OF GENUS LEPTOGENYS 303 Fics 16-32. 16-18. Outlines of heads of (16) Leptogenys guineensis; (17) L. mactans; (18) L. amon; pilosity and antennae omitted. 19. Lateral petiole node shape of L. zapyxis, anterior face to right. 20-22. Mandibular shape in (20) L. stevops; (21) L. zapyxis; (22) L. ergatogyna. 23. Apex of mandible of L. testacea. 24-32. Lateral petiole node shape of (24) L. crustosa; (25) L. ankhesa, dorsal view above; (26) L. princeps; (27) L. mastax ; (28) L. spandax; (29) L. attenuata, dorsal view above; (30) L. crassinoda, dorsal view above; (31) L. jeanneli ; (32) L. camerunensis; anterior face of petiole to the right in all cases. 304 B. BOLTON INDEX Synonyms are in itfalics. acutirostris, 297 adpressa, 289 aena, 289 africanus, 251 alluaudi, 297 amon, 283 angusta, 296 angusticeps, 256 ankhesa, 284 antongilensis, 298 arcirostris, 295 arnoldi, 281 attenuata, 274 bellii, 265 bellua, 285 bernardi, 270 brevinodis, 289 bubastis, 275 buyssoni, 285 camerunensis, 256 capensis, 289 castanea, 285. coerulescens, 296 comorensis, 262 conradti, 250 crassinoda, 276 cribrata, 254 crustosa, 251 cryptica, 286 cursor, 276 deflocata, 289 dentatula, 267 dentulata, 267 diatra, 287 dispar, 285 Dorylozelus, 240 elegans, 265 ergatogyna, 276 erylhraea, 257 erythraea, 257 excellens, 266 falcata, 254 falcigera, 252 ferrarii, 267 furtiva, 281 gracilis, 298 gracilis, 289 gradidieri, 296 grandior, 289 guineensis, 278 havilandi, 282 hewitti, 285 honoria, 287 wmerinensis, 298 incisa, 298 insinuata, 289 insularis, 252 intermedia, 289 jaegerskjoeldi, 275 jagerskidldi, 275 jeanneli, 253 jonesii, 298 khaura, 288 leiothorax, 258 Leptogenys, 239 Lobopelta, 240 longiceps, 268 Machaerogenys, 240 mactans, 288 mastax, 268 maxillosa, 254 Microbolbos, 240 microps, 278 nebra, 258 nitida, 289 nuserra, 269 occidentalis, 270 opalescens, 256 oswaldi, 297 parva, 285 pavesii, 255 peringueyi, 282 piroskae, 290 princeps, 271 ravida, 272 regis, 259 ridens, 295 rufipes, 251 saussurei, 297 schwabi, 259 sericea, 255 spandax, 279 speculans, 289 sterops, 261 strator, 291 striatidens, 292 stuhlmanni, 261 REVISION OF GENUS LEPTOGENYS stygia, 292 suarensis, 298 sulcinoda, 272 sulcinodis, 270 tenuis, 289 terroni, 273 testacea, 280 titan, 263 trilobata, 273 truncatirostris, 295 vindicis, 263 vinsonnella, 254 voeltzkowi, 298 zapyxis, 264 395 B. Botton, B.Sc., A.R.C.S. Department of Entomology BriTIsH MusEuM (NATURAL History) CROMWELL RoapD Lonpon SW7 5BD eS NER , AA “ ’ Besa Wii suas eQyey oye . 4 Belo ¥ al es be ye mult se : ( ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August 1965. £4.20. . SanDs, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. OxapDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50. . HemminG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8. . Mounp, L. A. A review of R. S. Bagnal)'s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. {5. . AriFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. £5. a . CrosskEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. WHALLEY, P. E. S. The Thyrididae of Africa and its Tslarids,. Pp. 198: 68 plates, 15 text-figures. October, 1971. {12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CrosskEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . voN Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CrosskEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND Ts | eet ae A CATALOGUE OF THE 9 PRIMARY TYPES OF MANTODEA (DICTYOPTERA) IN THE BRITISH MUSEUM (NATURAL HISTORY) J. A. MARSHALL BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 8 LONDON : 1975 A CATALOGUE COFCTHE PRIMARY EYPES::O8 MANTODEA (DICTYOPTERA) IN THE BRITISH MUSEUM (NATURAL HISTORY) BY JUDITH ANNE MARSHALL IS Pp. 307-329 BULEELIN~OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 8 LONDON: % 1075 an of Ue Tae Pe THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Depariment. _ This paper is Vol. 31 No. 8 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) ISSN 0007-1501 © Trustees of the British Museum (Natural History), 1975 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 12 February, 1975 Price £1.35 & Cn TALOGUE OF THE PRIMARY. [TYPES OF MANTODEA (DICTYOPTERA) IN THE BRITISH MUSEUM (NATURAL HISTORY) By J. A. MARSHALL CONTENTS Page SYNOPSIS , ‘ ‘ ‘ : : ‘ ; : ‘ : 309 INTRODUCTION ; : : : : , : : : : 309 CATALOGUE . : . ; é , : ‘ : ‘ F 311 REFERENCES . . : ; : 2 : , ‘ : : 327 SYNOPSIS An alphabetical catalogue is given of the holotypes, lectotypes (and associated paralectotypes) and syntypes of Mantodea in the collection of the British Museum (Natural History); 278 nominal species are represented by such type-material. Two lectotypes are newly designated. INTRODUCTION KIRBY (1904)) indicated in his Synonymic Catalogue that there were at that time 78 nominal species of Mantodea represented by holotypes or syntypes in the British Museum (Natural History). In the course of re-curating the Mantodea collection I have found that the types of a further 200 nominal species have since been added, and this would seem an appropriate time to publish a new list. The primary types, comprising holotypes, lectotypes (and their associated paralectotypes) and syntypes, are listed alphabetically by their original specific, subspecific or varietal name. The information for each type is given, when available, in the following standard sequence. Specific name, original genus in parentheses, author, date and page reference of original publication. : Status, sex, locality, any bionomic information, date of collection and collector of type(s); reference to lectotype designation if applicable; mention of any significant damage. Any further annotations, in a separate paragraph. In listing the data for each specimen the name of the country, in its correct form, is always given first, in capitals; the remaining items of locality data are then listed in descending order of magnitude, irrespective of the order in which they may appear on the label(s). When the country name given on the label differs from the one in current use, the older name given on the label is added in inverted commas. When the country name has been omitted from the label but is known, it is enclosed in 310 J. A. MARSHALL square brackets. If a place name given on the label differs in any way from the orthography in current English use, the name is quoted in inverted commas; if such a name is a misspelling or a name no longer in use, the correct spelling or currently used name is added in square brackets. The date of collection is always given in standard form, with Arabic numerals for the day and year, and Roman numerals for the month. The collector’s name is given in parentheses and italicized. Where a specimen is from a particular person’s collection but may not have been collected by that person the name of the person, followed by ‘Coll.’ is given, in parentheses but not italicized The data listed for each specimen is usually that taken from the label(s) pinned under the specimen. However, all the specimens have been checked against the original descriptions and in a few cases the published information augments the data on the labels; such information has been added to the data listed and marked as ‘[published information]’. Discrepancies between the published information and the label data have also been noted, and so too has a mistaken statement of the sex of a specimen; however, no attempt has been made to detect inaccuracies in the morphological descriptions of the specimens. Where a label on a specimen indicates that it has been the subject of a published illustration, this information is included with the data. All the specimens bear a British Museum (Natural History) registration number, which includes the year in which the specimen was received by the Museum. As this number is usually shared with many other specimens, often belonging to quite unrelated species, it is not quoted unless there is a special reason for doing so. Several older specimens bear a registration number but no date of collection, and some authors, notably Beier (1930a; 1930); 1932; 19374; 1937), consistently quoted the year of registration as the year of collection. The status of each type-specimen has been carefully checked against the original description, and all the types bear the circular colour-coded status labels used in the British Museum (Natural History): red for a holotype, purple for a lectotype, and blue for a paralectotype or syntype. Lectotypes have only been newly desig- nated in the two cases where it was thought particularly desirable to do so (Mantis lucubrans Burchell and Theopompa westwoodi Kirby). When only part of a syntypic series is present in the British Museum (Natural History) collection, the published number, sex and depository of the other syntypes are noted, to the extent that such information is available. The depositories are abbreviated as follows. BMNH British Museum (Natural History). FRI Forest Research Institute, Dehra-Dun. ISSS Istituto Superiore di Storia della Scienza, Florence. IZPAN Instytut Zoologiczny, Polska Akademia Nauk, Warsaw. MCSN Museo Civico di Storia Naturale, Genoa. MHN Muséum d’Histoire Naturelle, Geneva. MIZSU_ Museo ed Istituto di Zoologia Sistematica dell’Universita, Turin. MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin. MRAC Musée Royal de 1’Afrique Centrale, Tervuren. NR Naturhistoriska Riksmuseum, Stockholm. UM University Museum, Oxford. PRIMARY TYPES OF MANTODEA 311 Many type-specimens bear identification labels by the original author. In those cases where there is no such label but type status has been established, an identification label bearing the original name, status and sex, with ‘det. J. A. Marshall, 1974.’, has been added. In addition to the 78 nominal species correctly listed as types by Kirby (1904)), were two which he erroneously considered to be represented by type-material. The single (J) specimen of Phyllocrania westwoodi Wood-Mason, 1876, and the two (I gf, I 9) specimens of Melliera apotogamia Saussure, 1892, in the British Museum (Natural History) collection, are in both cases specimens seen by the authors and recognized as conspecific with the type-material, but not part of the type-species. Of the Fabricius type-material, maintained in the Banks Collection at the British Museum (Natural History), the types of two species, Mantis cancellata Fabricius, 1775, and Mantis urbana Fabricius, 1775, cannot be traced, and were not indicated as present by Kirby (19046). The types of three other species cannot be traced in the British Museum (Natural History) collection, the holotype (9) of Metviomantis pilosella Giglio-Tos, 1915, a syntype (g) of Leptomantis indica Giglio-Tos, 1915, and the two (I J, I 9) syntypes of Evemiaphila cairina Giglio-Tos, 1917. My thanks are due to Dr D. R. Ragge for his advice and assistance during the preparation of this work. CATALOGUE abyssinicus (Pseudoharpax) Beier, 1930): 457. Holotype g, Etuiopia: ‘Abyssinia’, plains N. of Lake Zwai, 5500-6000 ft, 3—4.xi.1926 (Dr H. Scott). acuminatus (Catasigerpes) Kevan, 1954: 467. Holotype 3, Kenya: N.F.D., El Wak, desert grass and thorn-bush, 5.xii.1944 (D. K. Kevan). affinis (Coptopteryx) Giglio-Tos, 1915b : 65. 1d syntype?, ARGENTINA (O. W. Thomas). Although this specimen was labelled by the author as ‘cotyp.’, the published data included only specimens from Brazil in the MNHU, Berlin and from Uruguay in the ISS, Florence. aliena (Amantis) Beier, 1930) : 439. Holotype 2, [BuRMA:] Tenasserim. ammonita (Eremiaphila) Uvarov, 1933a@ : 664. Holotype 9, JoRDAN: ‘Palestine’, Amman, E. of Jordan, 25.viii.1921 (P. A. Buxton). androgyna (Stagmomantis) Saussure & Zehntner, 1894 :147. Holotype 3, [BELIzE:] Belize (Blancaneaux). apotogamia (Melliera) Saussure, 1892 : 123. See page 311. arabica (Ameles) Uvarov, 19394:551. Holotype ¢, Saupr ArRasiA: ‘Arabia’, Ahlal [published in error as Abhah] 15.vi.1931 (H. St J. B. Philby). arabica (Nilomantis) Beier, 1930b : 438. Holotype g, OMAN: ‘Muscat’, Pass of Khubain, 1926 (J. Fernandez). arabicus (Tarachodes) Uvarov, 1922 : 352. Holotype g, SAuDI ARABIA: ‘S. W. Arabia’, El Kubar (G. W. Bury). Uvarov mistakenly quotes the data of the holotype as identical with that of the paratypes, i.e. SAupDI ARABIA: ‘Arabia’, Ktubu (G. W. Bury). 312 J. A. MARSHALL The head and pronotum on the ¢ paratype is that of the 2 paratype (head ‘mutilated’ as stated by Uvarov), and probably the head and pronotum on the 9 paratype is that of the ¢ paratype. argentina (Leptomiopteryx) Beier, 1930b:444. Holotype g, ARGENTINA: ‘Argentine Republic’, Villa Ana, F.C.S.Fe., ii.1926 (K. J. Hayward). aruana (Hierodula) Westwood, 1889 : 35. Holotype 9, [INDoNnEsIA:] Aru Island, ‘Westw. Mantd. pl. 4, f. 4.’ (Wallace). assamensis (Indothespis) Werner, 1935: 497. Holotype g, [Inp1A:] Assam, Goalpara [published in error as ‘Goalpore’], Kochugaon, v. 1906 [tip of abdomen missing]. atra (Gonypetella) Giglio-Tos, 1915¢ :143. Holotype g, UGanpa: ‘Uganda Prot.’, plains N.E. of Lake Edward, 3200 ft, 15—16.x.1911 (S. A. Neave). atrocephala (Gonypetella) Beier, 1930b : 442. Holotype 3, [SourH AFrRica:] Johannesburg, Bloksberg [error for Boksburg] (C. H. Pead). aurea (Sphodromantis) Giglio-Tos, 1917 : 63. 1 ¢ syntype, [GHANA:] ‘W. Africa’, Fantee country [published in error as ‘Fantec’], Bosso (Dr S. H. Jones); 1 § syntype, LIBERIA: ‘Liberian Coast’, Nanna Cru (W. P. Lowe); 1 2 syntype, GHANA: ‘Gold Coast’, Bibianaha, ix-x1i.1909 (Dr Spurrell). australiana (Orthodera) Giglio-Tos, 1917: 107. 1 g¢ syntype, AusTRALIA: ‘N. Australia’, Alexandria, i.1906 (W. Stalker); 1 9 syntype, AUSTRALIA: ‘Cent. Australia’, Hermannsburg (H. J. Hillier). australis (Miomantis) Beier, 1930b : 451. Holotype g, SourH West Arrica: Okahandja, 19-29.i11.1928 (R. E. Turner). : baetica (Mantis) Rambur, [1838] : 19. Lectotype 9, [Spain]: Malaga [published information] ; designated by Uvarov (1948 : 371). Paralectotypes. 1 $, [SPAIN:] Malaga [published information]; 1 9, [Sparn:] ‘Grenade’. Rambur said he found the species near Malaga, and Uvarov included the 9 from Granada as a paralectotype in the belief that it was labelled as being from Malaga. bandanus (Tarachodes) Giglio-Tos, 1917: 98. Holotype 3, GHANA: ‘Gold Coast’, N. Territories, Banda, 24.v.1913 (Jas. J. Simpson). bidens (Spanionyx) Saussure & Zehntner, 1894 :177. Holotype 9, [HonpuRAs:] ‘Ruatan’ Island, ‘figuré’ (Gaumer). bispinosa (Diacanthomantis) Giglio-Tos, 1915): 43. Holotype 9 nymph, Sri LANKA: ‘Ceylon’, near Kandy (A. Willey). boliviana (Musonia) Beier, 1930) : 443. Holotype g, Bortvia (J. Steinbach). borneana (Paratoxodera cornicollis) Beier, 1932 : 154. Holotype g, BRUNEI: I.vi.192I (Pres. by Mrs L. A. Allen). brevicollis (Oxyothespis) Beier, 1930b : 448. Holotype 3, Eturiopia: ‘Abyssinia’, S.E. of Lake Zwai, circa 5500 ft, Io-13.x1.1926 (J. Omer-Cooper). brevipennis (Oxyothespis) Beier, 19300: 448. Holotype g, GHANA: ‘Gold Coast’, Makango, 18.vi.1928 (A. W. J. Pomeroy). brevis (Mantis) Rambur, [1838] :21. Lectotype g, [Spatn:] ‘Grenade’; designated by Uvarov (1948 : 371) [abdomen detached and stuck on card]. Paralectotypes. 2 9, [SpAIN:] ‘Grenade’ [published information]. PRIMARY TYPES OF MANTODEA 313 brunneri (Humbertiella (?)) Kirby, 1904a: 83. 2 g¢ syntypes, [INDoNEsIA:] Lombok, Sapit 2000 ft, v-vi. 1896 (Heyene Coll.); 1 9 syntype [no locality label] (Heyene Coll.). burri (Oxypilus (Setoxypilus)) Roy, 1969: 1119. Holotype g, ANGora: Villa Luso, 3.x1.1927 (M. Burr). cairina (Eremiaphila) Giglio-Tos, 1917 : 81. See page 311. cancellata (Mantis) Fabricius, 1775 : 276. See page 311. capensis (Galepsus) Beier, 1930) : 436. Holotype g, SourH Arrica: Cape Province, Ceres, ii. 1925 (R. E. Turner). cattulus (Creobroter) Westwood, 1889: 42. Holotype g, [INDoNEsIA:] Java, ‘Westw. Mon. Mant. pl. 9, f. 10’ (E. I. C.). centralis (Phthersigena) Giglio-Tos, 1915) : 32. Holotype 2, AusTRALIA: ‘Central Australia’, Hermannsburg (H. J. Hillier). ceylonica (Compsomantis) Uvarov, 1927a : 88. Holotype 9, Sri LanKa: ‘Ceylon’, from the old collection of Colombo Museum, no exact data [published information]. championi (Acontista) Kirby, 1904@ : 233. See quadrimaculata (Acontista mexicana vat.) Saussure & Zehntner. championi (Angela) Saussure & Zehntner, 1894 :158. Holotype 9, [GUATEMALA:] Vera Paz, Cubilguitz (Champion) [abdomen missing]. championi (Phasmomantis) Saussure & Zehntner, 1894 :149. Holotype 9, [PANAMA:] V. de Chiriqui, 25—4000 ft (Champion). coeca (Iris oratoria) Uvarov, 19314 : 235. Holotype g, SupAN: Khor Arbaat Delta, iv-v.1926 (H. B. Johnston). collenettei (Eremiaphila) Beier, 1930a:193. Holotype g, SoMALI ReEPpuBLic: ‘British Somaliland’, Buran, 3000 ft, 10°13’ N, 48°47’ E, 19.xii.1929 (C. L. Collenette). congica (Hoplocorypha) Giglio-Tos, 1916: 18. 1 syntype, [ZAIRE:] S.E. Katanga, 4000 ft, 8.xii.1907 [head and all legs missing]. 2g and 1 9 syntypes in the MRAC, Tervuren. congica (Metentella) Giglio-Tos, 1915c : 178. 1 syntype, [ZAIRE:] S.E. Katanga, 4000 ft, 10-I1.xii.1907 (Neave Coll.); 2 2 syntypes, [ZAIRE:] Fwambo. cornuta (Theoclytes chlorophaea var.) Saussure & Zehntner, 1894: I9I. I Q syntype, [GUATEMALA:] Capetillio (G. C. Champion). Additional syntype(s) in the MHN, Geneva. coronatus (Galepsus) La Greca, 1952 :12. Holotype 9, Eruiop1a: Didessa River, 4000 ft, v.1946 (K. Guichard). costalis (Carvilia) Kirby, 1904a : 85. Holotype 9, Erniopia: ‘Abyss.’. crassa (Popa) Giglio-Tos, 1917 : 73. Holotype ?, Matawt: ‘Nyassaland’ (Dr J. E. S, Old). culicivora (Ligariona) Giglio-Tos, 1915c : 175. Holotype g, ANGoLa: W. Africa, ‘“‘Stated by natives to prey on mosquitoes.’’ (Note by donor.)’, 1905 (Dr F. C. Wellman). cycloptera (Eremiaphila) Uvarov, 19394 :550. Holotype 9, SaupI ARABIA: ‘Arabia’, S. Hedjaz, Daka, 17.x.1931 (H. St J. B. Philby). cypria (Ameles) Uvarov, 1936): 507. Holotype g, Cyprus: Limassol, 3.v.1923 (G. A. Mavromoustakis). 314 J. A. MARSHALL deserti (Iris) Uvarov, 1923: 60. Holotype 9, ALGERIA: ‘Alger. C. Sahara’, S. Oued Mya, 1912 (Hartert & Hilg.). Uvarov mistakenly quotes the data of the holotype as identical with that of the 2 ¢ paratypes, i.e. [ALGERIA:] North of In Salah, Ain Guettara, 12-14.iv.1912 (Hartert & Hilg.). He also states that he did not see any specimens from S. Oued Mya, although he knew of their existence; this must be a mistake in the case of the single female, which is labelled, described and illustrated as the holotype. dhufarica (Sphrodromantis) Uvarov, 1933): 261. Holotype 9, Oman: ‘S.E. Arabia’, Qara Mts, Fuzul, 1350 ft, 18.xi.1930 (B. S. Thomas). dioscoris (Teddia) Burr, 1899 : 43. Holotype g, [SouUTHERN YEMEN:] Sokotra, Dahamish, 21.xii.1898 (Dr H. O. Forbes & W. R. Ogilvie-Grant). dispar (Hierodula) Kirby, 1900:146. I g syntype, CuRistmMAs Is.: West Coast, x.1897 (C. W. Andrews); 1 2 syntype, CuRisTMAs Is.: neighbourhood of Flying Fish Cove, 20.ix- zo.x.— (C. W. Andrews); 3 2 & 1 nymphal 2 syntypes, Curistmas Is.: (C. W. Andrews); 1 9 ‘pupa-skin’ syntype, Curistmas Is.: Flying Fish Cove (C. W. Andrews). 35, 2 2 syntypes are missing. distanti (Solygia (?)) Kirby, 1899: 350. 1¢syntype, [SouTH AFRIcaA]: Barberton (P. Rendall) ; Ig, 2 29 syntypes [SouTH AFRIcA:] Zululand (Rev. W. H. Heale). distinctus (Oxypilus) Beier, 1930b : 447. Holotype 3, SIERRA LEONE: Moyamba, 29.i.1925 (E. Hargreaves). distinguenda (Hoplocorypha) Beier, 1930b: 444. Holotype g, Nicreria: ‘N. Nigeria’, Azare, vi.1924 (Dr Ll. Lloyd). . doriana (Rhombodera) Laidlaw, 1931 : 88. Holotype g, [Inp1A:] Behar, Ranchi, iii.1928 (W. B. R. Laidlaw). dyaka (Hierodula) Westwood, 1889 : 34. Holotype g, [?INDONEsIA:] ‘Borneo’, ‘Westw. Mantd. pl. 6, f. 4’. enei (Oxypilus (Oxypilus)) Roy, 1966:1440. Holotype g, NIGERIA: Western Prov., 34 mls N. of Oyo, near Idode, 16.xii.1960. evansi (Microthespis) Uvarov, 19314: 418. Holotype g, [PAKISTAN:] Baluchistan, Quetta, 27.vii-8.viii.1929 (J. W. Evans). exilis (Micrentella) Giglio-Tos, 1915c : 180. Holotype g, [?SoutH AFrRica:] ‘S’tn.’ [Simons- town?], ii.1892. eximia (Callimantis) Pascoe, 1882 : 423. 1 9 syntype, [BRAzIL:] Para, near Nazaré (Ff. P. Pascoe); I 2 syntype, VENEZUELA. Pascoe describes the single specimen which he collected in Brazil, and adds that in the British Museum the species bears the manuscript name ‘Callimantis venezuelae Bates’; the single 2 from Venezuela bears this label. fallax (Calidomantis) Giglio-Tos, 1917: 49. 1 ¢ syntype, Kenya: ‘B.E.A.’, Nairobi, 20.ili.1906 (W. L. Sclater) [abdomen missing]; 1 9 syntype, [KENyA:] ‘Brit. E. Africa’, Narossura R. (W. P. Lowe). fasciata (Pyrgomantis) Giglio-Tos, 1917: 106. 2 ¢ syntypes, [ZAIRE:] S.E. Katanga, 29.xi.1907 (Neave Coll.) [abdomen missing on one specimen, head missing on the other]. feminina (Miomantis) Beier, 1930b : 451. Holotype 9, Eruiop1a: ‘Abyssinia’, Abbai Affat, v-vi.1902 (Degen) [right elytron missing]. PRIMARY TYPES OF MANTODEA 315 Semorata (Musonia) Saussure & Zehntner, 1894 : 166. Holotype 2, NicaraGcua: Chontales (Janson) [tip of abdomen missing]. Jenestrata (Mantis) Fabricius, 1781 : 349. Holotype g, Soutn Arrica: ‘S. Africa’, ‘Africa aequinoctiali’ [published information]. In the Banks Collection, BMNH. ferus (Mionyx) Saussure & Zehntner, 1894 : 167. Holotype g, [PANAMA:] V. de Chiriqui, 25-4000 ft, ‘figuré’ (Champion) [tip of abdomen missing}. festae (Stauromantis) Giglio-Tos, 1917 : 55. 1d syntype, Ecuapor: Guayaquil (Rosenberg). I 2 syntype in the MIZSU, Turin. Ffraseri (Eremiaphila) Uvarov, 1921 :175. Holotype 9, [IRAg:] Mesopotamia, R. Euphrates, Anah, ‘‘‘Lived ro days in captivity, with nothing to eat or drink. A few seen in desert; run very fast.’’ (Note by collector)’, 15.vii.1920 (A. D. Fraser). fraseri (Pararivetina) Beier, 1931 : 361. Holotype g, Inp1A: Nilgiris [published as ‘Nilgiri Hills’), 1922 (Lt. Col. F. C. Fraser). fraterna (Acontista) Saussure & Zehntner, 1894 : 136. Holotype 2, Costa Rica: Cache (H. Rogers). fraterna (Stagmomantis) Saussure & Zehntner, 1894:144. I ¢ syntype, [MExico:] Tabasco, Teapa, iv.i— (H. H. Smith); 1 Q syntype, BrELizE: ‘B. Honduras’, R. Sarstoon, ‘figuré’ (Blancaneaux); 1 2 syntype, BELIZE: ‘British Honduras’ (Blancaneaux). Additional syntype(s) in the MHN, Geneva and the IZPAN, Warsaw. fratricida (Hierodula (Rhombodera)) Wood-Mason, 1878 : 581. Holotype g, [no data] [Inp1a:] Malabar [published information]. fuliginosa (Cimantis) Werner, 1931 : 1330. Holotype g, [Inp1A:] Madras, Anamalai Hills, 2400 ft, (46), 5.v.1930 [published in error as ‘6.v.1930’] (J. C. M. Gardner). fumosa (Cimantis) Giglio-Tos, 1915¢ : 155. Holotype g, [Inp1A:] Darjeeling. fumosa (Hoplocorypha) Giglio-Tos, 1916:11. 1 9 syntype, ZamBi1A: ‘N.E. Rhodesia’, Mid-Luangwa Vy., 3-1800 ft, 23-31.viii.1g1o (S. A. Neave) [abdomen missing]. The 3 syntype is missing. fumosus (Thrinaconyx) Saussure & Zehntner, 1894 :179. I ¢ syntype, [PANAMA:] V. de Chiriqui, 2—3000 ft, ‘figuré’ (Champion); 3 ¢ syntypes, [PANAMA:] V. de Chiriqui, 2—3000 ft (Champion). fusca (Bantiella) Giglio-Tos, 1915c :187. 1 ¢ syntype, Trin1IDAD: Port of Spain, 30.i.1912 (G. A. K. Marshall); 1 2 syntype, TrrnrpaD: Port of Spain (H. Caracciola). fusca (Gonypetella) Giglio-Tos, 1915c : 144. I ¢ syntype, UGANDA: Entebbe, 20.vii.1gI1I (C. C. Gowdey). I g syntype in the MRAC, Tervuren. gardneri (Deiphobella) Werner, 1935 : 496. Holotype 3, [INp1A:] U.P., Chakata Range, Haldwani, (95), 1.v.1930 (S. NN. Chatterjee). gardneri (Memantis) Werner, 1931 : 1331. 1 ¢ syntype, [INDIA:] Tarakhat, (26), v.1919 (Student’s Coll.). I g syntype in the FRI, Dehra-Dun. garuana (Hoplocorypha) Giglio-Tos, 1916 : 15. 1 syntype, NicERIA: ‘N. Nigeria’, Sokoto Dist., xii.tg10 (C. E. S. Watson); 1 3 syntype, Nicrria: ‘N. Nigeria’, Baro, 30.x.19Ito 316 J. A. MARSHALL (Dr W. Morrison) [head, pronotum and forelegs missing]; 1 nymphal 2 syntype, LipeRia: (W. P. Lowe) [abdomen missing]. I nymphal 2 syntype missing; additional syntypes in the MNHU, Berlin and the MCSN, Genoa. gigas (Acithespis) Giglio-Tos, 1916: 43. Holotype g, Niceria: ‘N. Nigeria’, Zungeru, x-xi.1910 (J. W. Scott-Macfie). gigas (Eremiaphila) Beier, 1930b : 432. Holotype 9, SupDAN: ‘Sudan Govt.’, Nikeila, 1.xii.1927 (Mr Shaw). godmani (Acanthops) Saussure & Zehntner, 1894 :181. Holotype 3, [BELIzE:] Belize, ‘figuré’ (Blancaneaux). gracilicollis (Parasphendale) Beier, 1930b : 449. Holotype 3, Kenya: ‘Brit. E. Africa’, E. Lake Baringo (R. Ford). gracilicollis (Sphodromantis) Beier, 1930b : 452. 46, 1 2 syntypes, GHANA: ‘Gold Coast’, Bibianaha, ix-xii.1909 (Dr Spurrell). The 2 and 1 3 were listed and labelled by Beier as ‘types’, but the description was based on these two specimens and 4 g ‘paratypes’, of which one is missing. gracilis (Brunneria) Giglio-Tos, 1915b:69. 1 ¢ syntype, Brazit: Parana (E. Dukinfield Jones). Additional syntypes in the MNHU, Berlin and MCSN, Genoa. gracilis (Neocilnia) Beier, 1930b: 450. Holotype 9, [SourH AFRIcA:] Cape Province, Ceres, iii.1925 (R. E. Turner). . gracilis (Orthodera) Giglio-Tos, 1917: 107. 1 g, 2 2 syntypes, AusTRALIA: Killalpanima, too miles E. of L. Eyre (H. J. Hillier). grandis (Acromantis) Beier, 1930b : 454. Holotype 9, [VierNnam (NortH):] Tonkin, Than- Moi, vi-vii.— (H. Fruhstorfer). gryps (Harpagonyx) Saussure & Zehntner, 1894 :176. I g syntype, [GUATEMALA:] Vera Paz, Pancina [published in error as ‘Panima’], ‘figuré’ (Champion). Additional syntype(s), depository not named. guichardi (Hypsicorypha) La Greca, 1952 :52. Holotype ¢, Somatt ReEpustic: ‘Brit. Somaliland’, Biyo Dader, 15.iii.1949 (K. M. Guichard). harpyia (Hierodula) Westwood, 1889 : 35. Holotype 9, Srr LANKA: ‘Ceylon’, ‘Westw. Mon. Mantd. pl. 5, f. 3’. henryi (Pezomantis) Uvarov, 1927a: 87. Holotype 2, Sri LanKa: ‘Ceylon’, Hakgalla, 4.iv.1924 (G. M. Henry). heterogamia (Stagmomantis) Saussure & Zehntner, 1894: 142. 14, 1Qsyntypes, [PANAMA:] Bugaba, 800-1500 ft, ‘figuré’ (Champion); 1 g.syntype, [PANAMA:] Bugaba, 800-1500 ft (Champion). heteromorpha (Oxyelaea) Beier, 1930b : 433. Holotype g, Kenya: ‘B. E. Africa’, Maziwa Mitatu & Maungu, 14.iii & 4.iv.1897 (C. S. Betton). incisa (Deiphobe) Werner, 1933 : 900. 14 syntype, [INDIA:] U.P., Chakata Range, Haldwani, at light, (57), 20.vi.1930 (S. N. Chatterjee) [published in error as ‘(47)’ and ‘N. C. Chatterjee’}. Ig and 1 2 syntypes in the FRI, Dehra-Dun. indica (Amorphoscelis) Giglio-Tos, 1915 : 33. Holotype 3, Inp1a: Ratnagiri (Dr Jayakar). indica (Deiphobe) Giglio-Tos, 1916 : 25. Holotype 9, Inp1a: Mhow. PRIMARY TYPES OF MANTODEA 317 indica (Leptomantis) Giglio-Tos, 1915) : 89. See page 311. inermis (Fischeria) Uvarov, 1922 : 353. Holotype g, SaAupi ARABIA: ‘S. W. Arabia’, El Kubar (G. W. Bury). infuscata (Pseudomiopteryx) Saussure & Zehntner, 1894 : 163. 1 ¢ syntype, [PANAMA:] V. de Chiriqui, 2—3000 ft, ‘figuré’ (Champion); 1 3 syntype, Mexico: Cordova (Hoege) ; I g syntype, [(GUATEMALA:] V. de Atitlan, 2500-3500 ft (Champion); 1 g syntype, [PANAMA:] Bugaba, 800-1500 ft (Champion). Additional syntypes, depository not named. inquinata (Acontista mexicana var.) Saussure & Zehntner, 1894 :136. 1 g syntype, Mexico: Cordova, ‘figuré’ (Hoege); 1 g syntype, [MExico:] Guerrero, Acaguizotla, 3500 ft, x.— (H. H. Smith). Additional syntype(s), depository not named. insignis (Phyllocrania) Westwood, 1843 :51. 1 2 syntype, SIERRA LEONE: ‘Westw. Arc. Ent. t. 62, f. 1’ (Revd. D. F. Morgan). I 2 syntype missing, I 9 syntype in the UM, Oxford. iridipennis (Miopteryx) Westwood, 1889 : 38. 1 Q syntype, Inp1a: ‘Ind. ?’ [abdomen missing]; I 2 syntype ?, Inp1a: ‘S. Ind.’. Additional syntype(s) in the UM, Oxford. jonesi (Pyrgomantis) Kirby, 1904a : 83. Lectotype g, NicErta: ‘S. Nigeria’ (Dr S. A. Jones); designated by Roy (1967 : 1542). Paralectotypes. 3, data as lectotype. judithae (Congoharpax) Roy, 1972b : 863. Holotype 3, Nicerta: ‘N. Nigeria’, Zaria, Samaru, m.v. trap, 9.v.1966 (J. C. Deeming). kilimandjarica (Miomantis) Sjéstedt, 1909 : 64. 1 3 syntype, [TANZANIA:] Kilimandjaro, ix.1905-6 (Sjéstedt); 1 g syntype, [TANZANIA:] Kilimandjaro, Kibonoto, 1300-1900 m, 24.1V.1905—6 (Sjdstedt); 1 2 syntype, [TANZANIA:] Kilimandjaro, 12.ix.1905—6 (Sjéstedt). Additional syntypes in the NR, Stockholm. kimberleyensis (Truxomantis) Sjéstedt, 1918 : 33. 1 g syntype, AusTrRaLta: ‘N, V. Austr.’, Kimberley District, ii.tg1o-1913 (Mjéberg) [tip of abdomen missing]. 93, I 2 syntypes in the NR, Stockholm. laevifrons (Eremiaphila) Uvarov, 1922 : 351. Holotype 9, OMAN: ‘Arabia’, Muscat (A. S. G. Jayakar). laosana (Humbertiella) Beier, 1930b : 434. Holotype 9, Laos: ‘French Laos’, Bandat (J. F. Godfrey). Beier gives the date as ‘1920-24’, misinterpreting the registration number of ‘1920-244’. lateralis (Cliomantis) Hinton, 1939 : 284. Holotypeg, [AusTRALIA:] N. Queensland, Flinders R., Glendower Stn, 1700 ft, 12.xi.1923 (G. H. Wilkins) [head missing]. laticollis (Pseudoharpax) Beier, 1930) : 458. Holotype 2, [RHoprEsIA:] Mashonaland, Salisbury, xi.1899 (G. A. K. Marshail). laurifolia (Phyllomantis) Saussure, 1892:124. Holotype 9, PANAMA: Bugaba, ‘figuré’ (Champion). liliputana (Haldwania) Beier, 1930b : 441. Holotype g, Inp1a: Haldwani Dist. [published in error as ‘Haldwania Dist.’], Kumaon, 1923 (H. G. Champion). limbata (Sibylla) Giglio-Tos, 1915b :92. 1 g¢ syntype, GHANA: ‘Gold Coast’, Bibianaha, 700 ft, 1912 (H. G. F. Spurreil). I 2 syntype in the MNHU, Berlin. 318 J. A. MARSHALL lobata (Mantis) Fabricius, 1781 : 350. Holotype 9, [mo data] [SourH Arrica:] ‘ad Cap. bon. sp.’ [published information]. In the Banks Collection, BMNH. lobipes (Parastagmatoptera) WRedtenbacher, 1892: 206. Holotype g, West INDIEs: ‘W.I.’, St. Vincent, S. end of island, open swampy land near sea, beaten from bushes, 27.ix.— (H. H. Smith). longipennis (Amantis) Beier, 1930) : 439. Holotype g, [VIETNAM (NortH):] Tonkin, Bao Ha, x-xii.1923 (H. Stevens). : lowi (Bolbe) La Greca, 1969 : 636. Holotype g, AusTRALIA: ‘N. Australia’, Queensland, nr Cairns, Yorkeys Knob, 4.ix.1966 (A. Walford-Huggins). lucubrans (Mantis) Burchell, 1822 : 418, 465. LECTOTYPE 4d, [no data} SoutH AFRICA: near Klaarwater (W. J. Burchell) [published information], here designated. Burchell gives the species description on p. 465, but has previously named the species on p. 418, where he also states that he ‘never took more than five’. Although the single specimen has been regarded as a holotype, it is now designated as the lectotype as other specimens may still be in existence. luteola (Mantoida) Westwood, 1889: 28. 1 ¢ ? syntype, [BRAzIL:] Santarem [abdomen missing]. Additional syntype(s) in the UM, Oxford. mabirica (Plistospilota) Giglio-Tos, 1917 :61. Holotype g, UGANDA: ‘Uganda Prot.’, Mabira Forest, Chagwe, 3500~—3800 ft, 16—25.vi.1g911I (S. A. Neave). maculata (Hapalopeza) Kirby, 1904a : 84. 1 g syntype, Sri LANKA: ‘Ceylon’, Kaltura, ii.1895 (£. E. Green) [no mid or hind legs]; 1 9 syntype, Sr1 LANKA: ‘Ceylon’, Dickoya, 1.xl1.1881. Further syntypes may exist, as Kirby does not state the number of specimens seen but mentions ‘some’ specimens. maculata (Junodia) Roy, 1972a: 566. Holotype g, ZAamsBia: ‘N. Rhodesia’, N’Changa (C. T. Macnamara). madagascariensis (Idolomorpha) Westwood, 1889: 46. I Q syntype, MADAGASCAR: “Westw. Mantd. pl. 6, f. 7.’ [abdomen missing]. Additional syntypes(s) in the UM, Oxford. malayana (Xanthomantis) Beier, 1932 :150. Holotype g, Mataysia: ‘Malay Penin.’, Selangor, Bukit Kutu, 3500 ft, at light, 13.iv.1926 (H. M. Pendlebury). malayensis (Psychomantis) Beier, 1932 : 152. Holotype 3, Maraysia: ‘Malay Penin.’, Kedah Peak, 3300 ft, [published as ‘at light’] 24.iii.1928 (H. M. Pendlebury). maroccana (Ameles) Uvarov, 1930b : 210. Holotype ?, Morocco: Ifrane, viii. 1930 (Miss A. Mackie). maya (Harpagonyx) Saussure & Zehntner, 1894 : 176. Holotype g ?, [MExico:] N. Yucatan, Temax (Gaumer) [abdomen missing]. maya (Liturgousa cayennensis var.) Saussure & Zehntner, 1894: 160. 2 3, I 9 syntypes, [Mexico:] N. Yucatan, Temax (Gaumer). maya (Mantoida) Saussure & Zehntner, 1894 : 125. 2 2 syntypes, [MExico:] N. Yucatan, Temax (Gaumer). maya (Stagmomantis) Saussure & Zehntner, 1894 :145. I Q syntype, Brxize: ‘British Honduras’ (Blancaneaux); 1 9 syntype, [Mexico:] N. Yucatan, Temax (Gaumer). I g§ syntype in the IZPAN, Warsaw. PRIMARY TYPES OF MANTODEA 319 maya (Theoclytes) Saussure & Zehntner, 1894: 192. I g¢ syntype, [MrExico:] Tabasco, Teapa, iv.— (H. H. S.); 1 2 syntype, [Mexico:} N. Yucatan, Temax (Gaumer). media (Diamusonia) Giglio-Tos, 1916: 7. 1g syntype, TRINIDAD (P. Rendall). 2 2 syntypes in the MNHU, Berlin. media (Photina) Giglio-Tos, 1915) : 73. 1 Q syntype, BRaziL: Parana (E. Dukinfield Jones). The ¢ syntype from Paraguay is missing. meggitti (Paratoxodera) Uvarov, 1927b : 658. Holotype 3, Burma: (F. J. Meggitt). mexicana (Acontista) Saussure & Zehntner, 1894 : 135. 1 2 syntype, [PANAMA:] Bugaba, 800-1500 ft (Champion) ; 1 9 syntype, [MExico:] Vera Cruz, Atoyac (Schumann); 1 2 syntype, [GUATEMALA:] V. de Chiriqui, 2—-3000 ft (Champion). There are also 3 2 with identification labels by Saussure and similar data to the above 2 from Guatemala, except for a different collection height of ‘below 4000 ft.’ Only ‘2—3000 ft’ is mentioned in the description and these specimens have therefore not been regarded as syntypes. Additional 2 syntypes, depository not named. mexicanus (Oligonyx) Saussure & Zehntner, 1894 :172. I ¢ syntype, Mexico: Presidio, ‘figuré’ (Forrer); 1 3 syntype, [GUATEMALA:] Vera Paz, Cubulco (Champion). micans (Amorphoscelis) Beier, 1937) : 177. Holotype 3, [MaLaysia:] Sarawak, Mt Dulit, 4000 ft, moss forest, house clearing, sheet (light), 8 p.m.-1o p.m., 29.x.1932 (Oxford Univ. Exp. B. M. Hobby & A. W. Moore). microptera (Eremoplana) Westwood, 1889 : 32. 1 2 syntype, [INpIA:] Burdwan, ‘Westw. Mantd. pl. 3, f. 5.’ (Hardwicke Bequest) ; 1 9 syntype, [INp1A:] Burdwan (Hardwicke Bequest) ; I 9 syntype, NEPAL: ‘India’ (Hardwicke Bequest). ministralis (Mantis) Fabricius, 1775 :277. 1 ¢ syntype, AUSTRALIA; 2 2 syntypes, [no data] [AUSTRALIA:] ‘nova Hollandia’ [published information] [one specimen with abdomen pinned separately and elytra missing, the other with abdomen missing, elytra and wings broken short]. In the Banks Collection, BMNH. minor (Archimantis) Giglio-Tos, 1917 : 43. 1 Q syntype?, [AUSTRALIA:] King George's Sound [published as ‘Kings River’]; 1 9 syntype?, [AusTRALIA:] Swan River [published as ‘Kings River’). Neither of the above specimens bears the usual hand-written identification label by Giglio-Tos, but it is possible that one or both of the specimens is syntypic. I g syntype in the MNHU, Berlin. minor (Blepharodes cornuta) Kevan, 1954: 471. Holotype 2, Kenya: N.P., Mt Kulal, 5500-6000 ft, 02°35’ N, 36°55’ E, x.1947 (J. G. Williams). minor (Memantis) Werner, 1931 : 1330. Holotype g, [INp1A:] Madras, Vizagapatam, (28), xii. 1928 (EZ. A. Glennie). minuta (Coptopteryx) Giglio-Tos, 1915b : 69. Holotype 9, CHILE: ‘Chili’, (R. M. Middleton). monstrosa (Archimantis) Wood-Mason, 1878 : 583. Holotype 9, AusTRALta: ‘North Australia’, Victoria River, 10.iii.1856 (I. R. Elsey). monstrosus (Tarachodes) Giglio-Tos, 1917:97. 1 ¢ syntype, UGaNpDa: Palebek, 24.11.1913 (W. P. Lowe); 1 2 syntype, Somatrt Repustic: ‘Brit. Somaliland’ (R. E. Drake-Brockman) {abdomen missing]. montana (Polyspilota) Beier, 1931 : 360. Holotype 9, Eruiopia: ‘Abyssinia’, Djem-Djem, circa 8000 ft, 6.x.1926 (Dr H. Scott). 320 J. A. MARSHALL montana (Stagmomantis) Saussure & Zehntner, 1894 :146. I ¢ syntype, [GUATEMALA:] V. de Atitlan, 2500-3000 ft, ‘figuré’ (Champion) [abdomen missing]; 1 ¢ syntype, [GUATE- MALA:] V. de Atitlan, 2500-3000 ft (Champion); 1 3 syntype, [MExico:] Guerrero, Chilpan- cingo, 4000 ft, vi— (H. H. Smith); 1 2 syntype, [MExico:] Guerrero, Acapulco, ix.— (H. H. Smith); 1 9 syntype, [Mexico:] Guerrero, Tepetlapa, 3000 ft, x.— (H. H. Smith). montanus (Muscimantis) Henry, 1931 :124. Holotype g, Sri LanKa: ‘Ceylon’, Ohiya, 6800 ft, 20.iv.1928 (G. M. Henry). nana (Iris) Uvarov, 1930a : 632. Holotype g, Iraq: (Baghdad ?) (O. Theodor) [abdomen missing]. natalica (Miomantis) Beier, 1930b : 450. Holotype g, [SourH Arrica:] Natal, Durban, 5.1v.1894 (H. W. Whyte). nebulosa (Sinaiella) Uvarov, 1924 :3. Holotype g, [IsraEL]: N. Sinai, Mohammedia, 23.x.1916 (A. W. Boyd). nicaraguae (Macromantis ovalifolia var.) Saussure & Zehntner, 1894:150. Holotype 6, NICARAGUA: Chontales (Janson). nigerica (Hoplocorypha) Beier, 1930b : 446. Holotype $, NicEr1a: ‘N. Nigeria’, Azare, vi.1924 (Dr Ll. Lloyd). nigerica (Plistospilota) Giglio-Tos, 1917 : 61. Holotype 2, NicErt1a: ‘N. Nigeria’, Antipa, 6.i.1911 (J. J. Simpson). nigerica (Pyrogomantis) Giglio-Tos, 1917 : 105. Holotypeg, NicERtA: ‘N. Nigeria’, Zungeru, X-x1.1910 (J. W. Scott-Macfie). nigericus (Oxypilus) Beier, 1930b : 447. Holotype 3, NicERtIA: ‘N. Nigeria’, Azare, 1925 (Dr LI. Lloyd). nigericus (Paragalepsus) Beier, 1930b : 435. Holotype 3, NicErRiA: ‘N. P. Nigeria’, Akwanga Div., Mama Dist., ii. 1928 (Dr Ll. Lloyd). nigericus (Paroxyophthalmus) Giglio-Tos, 1917 : 104. 2 syntypes, NicERIA: ‘N. Nigeria’, Zungeru, x-xi.1g10 (J. W. Scott-Macfie). nigericus (Pseudoharpax) Giglio-Tos, 1915b:105. 1 ¢ syntype, NicERIA: ‘N. Nigeria’, Baro, 30.x.1910 (Dr W. Morrison); 1g syntype, SIERRA LEONE: ‘Protectorate’, 1911 (Dr H.E. Arbuckle) ; 1 9 syntype, NicER1A: ‘N. Nigeria’, Kogin Sirikin, Pawa, 8.xii.1g10 (J. J. Simpson). nigericus (Sigerpes) Giglio-Tos, 19154 :13. 1 syntype, NicErRta: ‘S. Nigeria’, Cross River, Afikpo, 7.vi.1910 (J. J. Simpson). I g syntype in the MNHU, Berlin. nigriventer (Amorphoscelis) Beier, 1930b : 432. Holotype g, GHANA: ‘Gold Coast’, Bibianaha, ix-xii.1909 (Dr. Spurreill). nigropicta (Theopompa) Westwood, 1889 : 29. 14 syntype, [no data] AUSTRALIA [published information], ‘Westw. Mantd. pl. 2, f. 6’. Additional syntype(s) in the UM, Oxford. nilotica (Oxyothespis) Giglio-Tos, 1916 : 42. Holotype g, [SUDAN or Ernror1a:] Blue Nile (E. S. Crespin) [abdomen missing]. niloticus (Sigerpes) Giglio-Tos, 1915a:12. Holotype g, [SupaAN:] ‘Egypt’, White Nile (L. Loat). nimulensis (Galepsus) Giglio-Tos, 1917: 101. 1 Q syntype, NIGERIA: ‘N. Nigeria’, Baro, 14—-16.x.1910 (J. J. Simpson). . 2 3 syntypes in the MCSN, Genoa and 1 ¢ syntype in the MNHU, Berlin. PRIMARY TYPES OF MANTODEA 321 nordica (Stagmomantis) Giglio-Tos, 1917 : 53. I g, 1 2 syntypes, [U.S.A.:] ‘Am Bor’, ‘Virga.’ (Fry Coll.). nova (Plistospilota) Beier, 1930b : 452. Holotype 9, [INp1A:] Cachar (W. Grant). Beier quoted ‘18.x.—’ as the date of collection, misinterpreting the handwritten label ‘78-70’, which is the registration number 1878-70. obesa (Trachymantis) Giglio-Tos, 1917 : 46. Holotype 9, AustTraLia: Central Australia, Hermannsburg (H. J. Hillier). obscura (Cliomantis) Hinton, 1939 : 286. Holotype 2, AusTRALIA: ‘W. Australia’, Merredin [published in error as ‘Merreden’], 13.xii.1935 (R. E. Turner). obscura (Mantis) Fabricius, 1781 : 349. Holotype g, ‘ArrRica’: ‘Africa aequinoctiali’ [published information]. In the Banks Collection, BMNH. occidentalis (Galinthias) Beier, 1930b : 456. Holotype 9, SIERRA LEONE: Nijala [published in error as ‘Njaba’], xii.1926 (E. Hargreaves). occidentalis (Pyrgomantis) Giglio-Tos, 1917: 105. 1 g¢ syntype, NIGERIA: ‘N. Nigeria’, Minna, i.1910 (Dr. J. W. S. Macfie). I g syntype in the MNHU, Berlin. occidentalis (Sigerpes) Wood-Mason, 1879: 259. Holotype 3g, [no data] SIERRA LEONE [published information]. occipitalis (Hapalopeza) Westwood, 1889 : 36. 1d, 1 2 syntype, Hone Kona; 1 g syntype, HoncG Kona [only abdomen and metathorax with hind wings and left hind leg}. Additional syntype(s) in the UM, Oxford. ocellata (Statilia) Uvarov, 1922 : 353. Holotype 2, [PAKISTAN:] Quetta. ocularis (Litaneutria) Saussure, 1892 : 124. Holotype 3, Mexico: N. Sonora (Morrison) [hind legs missing]. oculata (Mantis) Fabricius, 1781 : 348. Holotype ¢, ‘Arrica’: ‘Africa aequinoctiali’ [published information] [abdomen missing]. In the Banks Collection, BMNH. orientalis (Bolbena) Beier, 1930a : 196. Holotype, Somat Repustic: ‘British Somaliland’, Buran, 3000 ft, 10°13’ N, 48°47’ E, 19.xii.1929 (C. L. Collenette). orientalis (Heterochaeta) Kirby, 1904a: 87. 1 9 syntype, ‘E. AFRICA’; I 3g ? syntype, ‘AFR. OR.’: Luitpoldkette (Janson) [abdomen missing]; 1 nymphal syntype, ‘E. AFR.’ [abdomen missing]. The @ syntype bears an incorrect registration number, 1892-22 (referring to ‘3 galls on oak, S. Spain’); the next entry in the Register, 1892-23 includes ‘1 Mantis’ from ‘E. Africa (German)’, and is probably the correct registration number for the specimen. orientalis (Metentella) Giglio-Tos, 1915¢:178. 3 3 syntypes, MozAMBIQUE: ‘Port. E. Africa’, Valley of Kola R., nr E. Mt Chiperone, 1500-2000 ft, 6.iv.1913 (S. A. Neave); 13 syntype, Marawi: ‘Nyasaland’, S.E. shore L. Nyasa, btwn Ft Maquire & Fort Johnston, 6—-17.iii.1910 (S. A. Neave) [head, pronotum and forelegs missing]. Additional syntypes in the MNHU, Berlin. orientalis (Oligomantis) Giglio-Tos, 1915a: 4. 1 syntype, StncaporE (H. N. Ridley). I 2 syntype in the MNHU, Berlin and 1 2 syntype in the MCSN, Genoa. oxynotus (Tarachodes) La Greca, 1952: 6. Holotype 9, Somat1 Repustic: ‘Somaliland’. Hargeisa, 09°33’ N, 44°03’ E, i.1947 (W. H. Wood). 322 J. A. MARSHALL oxyops (Metacromantis) Beier, 1930b : 455. Holotype 2, Sri Lanxa: ‘Ceylon’, Haragama, 18.vii.1910 (E. E. Green). oxyops (Paragalepsus) Beier, 1930) : 436. Holotype g, UGanpa: Kamule, 20.xii.1921 (H. Hargreaves). pallida (Beesoniella) Werner, 1935 : 498. Holotype g, [INnpIA:] Madras, Amarampalam R. Nilambur, (94), 12.iv.1933 (C. F. C. Beeson). pallida (Pyrgomantis) Giglio-Tos, 1917: 106. 1 ¢ syntype, GHANA: ‘Gold Coast’, N. Territories, 6.v.1913 (Jas. J. Simpson). palmonii (Xenomantis) Uvarov, 1939b : 219. Holotype ?, JorDAN: ‘Palestine’, E. bank of R. Jordan, Lake Genazareth, 620 ft below sea, 1938 (J. Palmoni). parumspinosus (Blepharodes) Beier, 1930b : 460. 1 g syntype, NiGERIA: ‘N. Nigeria’, Azare, 1925 (Dr Li. Lloyd); 1 9 syntype, NIGERIA: ‘N. Nigeria’, Azare, 1926 (Dr LI. Lloyd). The above syntypes were listed and labelled by Beier as ‘types’, but the description of the species was based on these specimens plus 1 ¢ ‘paratype’ with the same data and 1 9 ‘paratype’ from the Sudan, both of which are missing. Two further specimens with the same locality and collector as the ‘types’ but different collection dates are present, but were not included in the description. parva (Camelomantis) Beier, 1932 : 151. Holotype g, Mataysia: ‘Malay Penin.’, Kedah Peak, 3300 ft, 27.iii.1928 (H. M. Pendlebury). parva (Leptomantis) Werner, 1933 : 899. I ¢ syntype, [INpDIA:] Dehra Dun, Student’s Coll., (68), 15.ix.1925. Ig, I 2 syntype in the FRI, Dehra-Dun. parva (Oxyothespis) Beier, 1937a:378. Holotype 3, UGanpa: Turkana Prov., v.1934 (D. R. Buxton) [abdomen missing]. pauperata (Mantis) Fabricius, 1781 : 346. Holotype 92, [no data] [INp1a:] Coromandel {published information] [head and abdomen missing]. In the Banks Collection, BMNH. pectoralis (Rhombodera) Westwood, 1889 : 33. Holotype 9, [no data] PapuA NEw GUINEA: ‘Papua’ [published information]. penangica (Camelomantis) Giglio-Tos, 1917 : 67. 2 syntypes, [MaLAysIA:] Penang Hills, 1800-2500 ft, iii.1898 (S. S. Flower); 1 g syntype, [MataysiA:] Penang Hills, 2000- 2500 ft, iii.1898 (S. S. Flower); 1 g syntype, [MaLaysiA:] Penang, 2260 ft, 25.xi.1896 (S. Flower); 4 3 syntypes, [MALaAysIA:] Penang. I 2 syntype, depository not named. persa (Iris) Uvarov, 1922 : 355. 1 syntype, [IRAN:] Bushire, viii.1890 (W. D. Cumming) ; I g syntype, [IRAN:] Bushire, 4.ix.1890 (W. D. Cumming); 1 § syntype, [IRAN:] Bushire, ix.1890 (W. D. Cumming); 1 § syntype, [IRAN:] Bushire (W. D. Cumming). Uvarov incorrectly states that all the specimens were collected in September, 1890. peruviana (Liturgusa) Giglio-Tos, 1915b : 77. Holotype 2, PERu: Paleazu. philbyi (Attalia) Uvarov, 1936a : 533. Holotype g, Saup1 ARaABta: ‘Arabia’, S. Hedjaz, Ashaira, 12.viii.1931 (H. St J. B. Philby). philippina (Amantis) Giglio-Tos, 1915c : 152. Holotype g, PHILIppINEs (C. S. Banks). philippina (Haania) Giglio-Tos, 1915b : 200. Holotype g, PHILippINes (C. S. Banks). picea (Hoplocorypha) Giglio-Tos, 1916: 10. 1, I 2 syntypes, [KENyA:] ‘Brit. E. Africa’, Narossura R. (W. P. Lowe). PRIMARY TYPES OF MANTODEA 323 pilosella (Metriomantis) Giglio-Tos, 1915b: 70. See page 311. planivertex (Miomantis) Sjéstedt, 1909 :64. I 3 syntype, [TANzANIA:] Meru, ‘Nieder’, Ngare na nyuki, i (Sjéstedt) [abdomen and hind legs missing]. 24, 2 2 syntypes in the NR, Stockholm. pluvisilvae (Aetaella) Henry, 1931 : 126. Holotype g, Sri LAnKa: ‘Ceylon’, Ratnapura’ 16.ix.1929, ‘Reared to maturity in Colombo, 4.ii.1930’ (G. M. Henry). punctata (Gonypetella) Giglio-Tos, 1915¢:144. 1 g syntype, UGANDA: ‘Uganda Prot.’, Mabira Forest, Chagwe, 3500-3800 ft. 2 $6 syntypes in the MCSN, Genoa. punctillata (Oxymantis) Werner, 1931 : 1333. Holotype g, [INp1A:] Madras, Palghat, Sappal, (43), 1700 ft, 15.v.1930 (J. C. M. Gardner). punctulatus (Oligonyx) Saussure & Zehntner, 1894 :174. Holotype 3g, [MExico:] Guerrero, Dos Arroyos, 1000 ft, ‘figuré’, ix.— (H. H. Smith). pustulifera (Hierodula (Rhombodera)) Wood-Mason, 1878 : 583. 1 9 syntype, [no data] [AUSTRALIA:] ‘one of the islands in the Torres Straits’ (Rev. McFarlane) [published informa- tion]. pygmaea (Calidomantis) Giglio-Tos, 1917 : 52. 14 syntype, UGANDA: Entebbe, I-4.v1.1912 | (C. C. Gowdey). I g syntype in the MNHU, Berlin. quadrimaculata (Acontista mexicana var.) Saussure & Zehntner, 1894 : 136. Holotype 6, [GUATEMALA:] Vera Paz, Lanquin, ‘figuré’ (Champion). See championi (Acontista) Kirby, replacement name for the above, which is a junior secondary homonym of Acontista quadrimaculata (Serville, [1838)). queenslandica (Truxomantis) Sjéstedt, 1918 : 31. 1 g syntype, [AUSTRALIA:] ‘Queensl.’, Alice River, ix.— [1910-1913] (Mjéberg). 46, 2 2 syntypes in the NR, Stockholm. radians (Iris) Uvarov, 1931a : 234. Holotype g, [PakisTANn:] Lyallpur, light collection, 1.ix.1929 [abdomen missing]. raggei (Oxypilus (Oxypilus)) Roy, 1969: 1115. Holotype 3, SomMaLtt REpuBLic: ‘Brit. Somaliland’, Hargeisa Locust Camp, 14.xi.1949 (K. M. Guichard). rapax (Mionyx) Saussure & Zehntner, 1894:167. Holotype 92, Costa Rica: Cache (H. Rogers). rehni (Oxyophthalmellus) La Greca, 1952 :15. Holotype g, Somat! ReEpusBLic: ‘Brit. Somaliland’, Haud, 8°15’ N, 46°15’ E, 2100 ft, night, after rainy season, 3.iv.1932 (R. H. R. Taylor Coll.). rendalli (Acanthomantis) Kirby, 1899 : 352. Holotype ¢ [not 9, as stated by Kirby], Matawt: ‘Nyassaland’, Fort Johnston (P. Rendall). rhodesiaca (Bolbella) Beier, 1930b : 438. Holotypeg, Ruopesia: ‘S. Rhodesia’, Mt Chirinda, x-xl.1g11 (C. F. M. Swynnerion). rhodesica (Pyrgomantis) Giglio-Tos, 1917: 106. Holotype g, ZampBia: ‘N. Rhodesia’, Mkushi, iii.1910 (Silverlock Coll.) [abdomen missing]. robusta (Sphendale) Kirby, 1904a : 86. Holotype 9, NEpaL (Hardwicke Bequest). robusta (Tarachodes) Beier, 1930b : 434. Holotype g, Nicer: ‘N. P. Nigeria’, Matyoro Lakes, 25 miles N. of Gombe, iii.1929 (Dr Li. Lloyd). 324 J. A. MARSHALL rotundipennis (Eremiaphila) Kirby, 1904b : 210. 1 2 syntype, (no data] Ecypt [published information]; 1 9 syntype, [no data] ‘Westw. Mant. pl. 1, f. 11’; 1 nymphal syntype, Ecypr. Additional syntype(s) in the MHN, Geneva. rubrobrunneus (Theopropus elegans) Beier, 1932 : 153. Holotype g, MAraysia: ‘Malay. Penin.’, F. M. S., Pahang, Cameron Highlands, Tanah Rata, 4800 ft, 25.xi.1924 (I. Kedit). rufipennis (Eremiaphila) Uvarov, 1929: 92. Holotype 9, [ISRAEL:] Sinai, Wadi Tarfa (Bodenheimer & Theodor). rungsi (Tenodera) Uvarov, 19354: 41. Holotype 2, Morocco: ‘Maroc’, Rabat, 15.xi.1933 (Rungs Coll.). rustica (Mantis) Fabricius, 1781 : 350. 1 g syntype, [ARGENTINA:] Patagonia [elytra and wings very damaged]; 1 g syntype, [no data] [ARGENTINA:] ‘ad littora Patagonia’ [published information] [head, pronotum and forelegs missing]. In the Banks Collection, BMNH. sabulosa (Sinaiella) Uvarov, 1939a : 552. Holotype g, SAupi AraBiA: ‘Arabia’, Shudhaib, viii.1936 (H. St J. B. Philby). saevus (Mionyx) Saussure & Zehntner, 1894: 167. I g syntype, [PANAMA:] V. de Chiriqui, 2500-4000 ft (Champion) ; 1 § syntype, [PANAMA:] Bugaba, 800-1500 ft (Champion) [abdomen missing]. saussurei (Alphamantis) Giglio-Tos, 1915c:171. I g syntype, LzsotHo: ‘Basutoland’, Maseru (L. Wroughton). I ¢ syntype in the MHN, Geneva. semirufa (Acontistes) Westwood, 1889 : 45. Holotype 3, [BraziL:] Ega. septentrionalis (Stagmatoptera) Saussure & Zehntner, 1894 : 186. Holotype 2, [PANAMA:] Bugaba, 800-1000 ft, ‘figuré’ (Champion). serricornis (Parastagmatoptera) Kirby, 1904b : 298. 1 3 syntype, [BRaziL:] Santarem, “Westw. Mantd. pl. 6, f. 1.’. Additional syntypes in the MHN, Geneva and possibly in other depositories. shelfordi (Deroplatys) Kirby, 1903 : 304. Holotype 9, [MArays1a:] Sarawak, vi.1896. signifer (Harpax) Walker, 1859. Holotype ?, Sr1 LANKa: ‘Ceylon’ [left elytron only present]. similis (Gonypetella) Giglio-Tos, 1915¢: 145. I g syntype, ZAmBia: ‘N. E. Rhodesia’, Serenje Dist., about 4500 ft, 14.xii.1907 (Neave Coll.); 1g syntype, ZAMBIA: ‘N. E. Rhodesia’, Serenje Dist., about 4500 ft, 15.xii.1907 (Neave Coll.) ; 1g syntype, [RHODEsIA:] Mashonaland, Salisbury, xii.1899 (G. A. K. Marshall). simplex (Gonypeta) Beier, 1930b : 440. Holotype 3, SINGAPORE (H. N. Ridley). sinensis (Antissa) Giglio-Tos, 1915b:99. 2 4 syntypes, Cu1na: Shense Prov., Taipaishan, 10.x.1905.; I g@ syntype, CHINA: Shense Prov., Taipaishan.; 1 9 syntype, CHINA: Shense Prov., Taipaishan 12.x.1905. I ¢ syntype missing. singaporana (Amorphoscelis) Giglio-Tos, 1915) : 32. Holotype g, SInGAporE: (H. N. (Ridley). siporana (Acromantis) Giglio-Tos, 1915a:6. 1 g syntype, [INDoNEsIA:] Sumatra, Nias (Fry Coll.) [abdomen missing]. I 9 syntype in the MCSN, Genoa. PRIMARY TYPES OF MANTODEA 325 somalica (Dorymantis) Beier, 1930a:197. Holotype g, SomaLt ReEpusBtic: ‘British Somaliland’, Buran, 3000 ft, 10°13’ N, 48°47’ E, 19.xii.1929 (C. L. Collenette). speciosa (Chrysomantis) Giglio-Tos, 1915b : 103. Holotype 3, GHANa: ‘Gold Coast’, Aburi, 1913 (W. H. Patterson). spinicollis (Sphaeromantis) Beier, 19304 :194. Holotype g, Somatt Repustic: ‘British Somaliland’, Buran, 3000 ft, 10°13’ N, 48°47’ E, 19.xii.1929 (C. L. Collenette). splendida (Iris) Uvarov, 1922 : 355. Holotype 9, ArGHANISTAN: ‘Afghan.’ (Dr Aitchison). steelae (Oreomantis) Uvarov, 1935b:208. 2 9, 1 nymphal syntypes, [CAMEROUN:] Mt Cameroon, 2nd plateau, 9300 ft, 11.i.1932 (M. Steele); 1 nymphal syntype, [CAMEROUN:] Mt Cameroon, Ist plateau, 10,000—13,000 ft, 12.i.1932 (M. Steele). I nymphal syntype missing. strachani (Pseudochaeta) Kirby, 1904a : 88. Holotype ¢ [not 9, as stated by Kirby], [NIGERIA:] Lagos (Dr H. Strachan). straminea (Cheddikulama) Henry, 1932 :14. Holotype 9, Srr Lanka: ‘Ceylon’, Kallar, 3 miles from Cheddikulam, ‘Reared in Colombo fron a young larva. Killed vi.1931.’, 12.vill.1930 (K. L. A. Perera). striata (Hierodula) Giglio-Tos, 1917 : 64. Holotype 9, [Maraysia:] Perak (Doherty). striata (Hoplocorypha) Beier, 1930b : 445. Holotype J, Soura West Arrica: Okahandja, 10-16.ii.1928 (R. E. Turner). strigipennis (Oxypilus) Westwood, 1889 : 44. Holotype g, [SourH Arrica:] ‘Port Natal’ [Durban], ‘Westw. pl. 9, f. 7.’. striolatus (Oligonyx) Saussure & Zehntner, 1894 :173. 1 ¢ syntype, GUATEMALA: S. Geronimo, ‘figuré’ (Champion) [abdomen missing]; 1 ¢ syntype, GUATEMALA: S. Geronimo, (Champion) ; 1 § syntype, GUATEMALA: Duenas (Champion) [head missing]. subcornuta (Calamothespis) La Greca, 1952: 45. Holotype g, SoMALI REPuBLIc: ‘Brit. Somaliland’, Haud, 8°26’ N, 45°43’ E, 2300 ft, day, end of rainy season, 17.v.1932 (R. H. R. Taylor Coll.) [head and fragments of hind legs detached and stuck on card]. sudanensis (Oxyothespis) Giglio-Tos, 1916: 42. 1 g syntype, [SUDAN:] Red Sea, Port ‘Soudan’ (Mrs N. Waterfield) [abdomen missing]. 2 nymphal 2? syntypes, one each in the MIZSU, Turin and the ISSS, Florence. superstitiosa (Mantis) Fabricius, 1781 : 348. Holotype 9, ‘Arrica’: ‘Africa aequinoctiali’ [published information] [abdomen missing]. In the Banks Collection, BMNH. tanzanicus (Oxypilus (Oxypilus)) Roy, 1969 : 1110. Holotype J, Tanzania: ‘Tanganyika’, Rukwa Rift, Kafukola, 25.xi.1954 (R. F. Chapman). taylori (Calamothespis) La Greca, 1952:40. Holotype g, Somatr Repustic: ‘Brit. Somaliland’, Haud, 4000 ft, rainy season, ix.1932 (Capt. R. H. Taylor). tessellatus (Oligonyx) Saussure & Zehntner, 1894 :174. Holotype g, [Mexico:] Guerrero, Omilteme, 8000 ft, ‘figuré’, vii— (H. H. Smith). testacea (Cimantis) Werner, 1931 : 1330. Holotype, [INp14:] Bengal, Tista Valley, Darjeel- ing, 700 ft, (27), 27.ix.1929 (J. C. M. Gardner). toltecus (Oligonyx) Saussure & Zehntner, 1894 :175. Holotype 2, [Mextico:] Guerrero, Chilpancingo, 4600 ft, viii— (H. H. Smith). 326 J. A. MARSHALL trilobus (Galepsus) Giglio-Tos, 1917: 102. 2 g syntypes, [TANZANIA:] ‘G. E. Africa’, Vy. Ruaha R., 18—21.xii.1g10 (S. A. Neave). I ¢ syntype in the MNHU, Berlin. trinitatis (Bantiella) Giglio-Tos, 1915c : 186. 1 3, 2 9 syntypes, TRINIDAD: Port of Spain (H. Caracciola). tristis (Uvaromantis) Beier, 1930) : 457. Holotype g, GHANA: ‘Gold Coast’, Makango, 18.vi.1928 (A. W. J. Pomeroy) [head missing]. turcica (Eremiaphila) Westwood, 1889: 29. 4 2 syntypes, TURKEY. turneri (Hoplocorypha) Beier, 1930b : 445. Holotype g, SourH West AFrRica: Okahandja, 10-16.ii.1928 (R. E. Turner). ugandana (Hoplocorypha) Beier, 1930) : 444. Holotype 3, UGaNnpDa: Chua, xi-xii.1925 (Dr G. D. H. Carpenter). ugandanus (Amorphoscelis) Beier, 1931 : 359. Holotype 9, UGANDA: Kampala, on Eucalyptus, 27.ix.1928 (G. L. R. Hancock) [head missing]. ugandanus (Pseudoharpax) Giglio-Tos, 1915b: 106. 1 syntype, [RHODESIA:] Mashonaland, Salisbury, xi.1899 (G. A. K. Marshall); 1 2 syntype, UGANDA: Entebbe, 28.x.1912 (C. C. Gowdey); I 9 syntype, UGANDA: Mabira Forest, 3.viii.1913 (C. C. Gowdey). The ¢ from Rhodesia is labelled by Giglio-Tos as ‘typus’, although the only locality given in the description is Uganda. Additional syntypes, g and 9, in the MCSN, Genoa. umtalica (Ligariella) Giglio-Tos, 1915c:172. Holotype g, [RuHoprsi1a:] Mashonaland, Umtali, 1900 (Marshall Coll.). urbana (Mantis) Fabricius, 1775 : 278. See page 311. uvana (Dysaules) Henry, 1932: 2. Holotype g, Sri Lanka: ‘Ceylon’, Uva, Wellawaya, ‘Reared to maturity in Colombo, 6.vili.1931’, iv.1931 (K. L. A. Perera). uvarovi (Empusa) Chopard, 1921 : 52. 1 ¢ syntype, [IRAQ:] Mesopotamia, on alhagi at margin of dried-up marsh 12 miles below Amara nr Tigris, 12.ix.1918 (W. Edgar Evans); 1 9 syntype, [IRAQ:] Mesopotamia, Amara, 10.ix.1918 (P. A. Buxton). uvarovi (Eremiaphila) Bodenheimer, 1933: 79. Holotype 9, JoRDAN: ‘Transjordan’, Maan, 1.xi.1932. uvarovi (Gonypetella) Beier, 1930b : 442. Holotype 3, [KENyA:] ‘Brit. E. Africa’, Mogorr R. Valley, vi.1913 (A. O. Luckman). vansomereni (Junodia) Roy, 1972a : 573. Holotype g, UGanpa: Bwamba (H), iii.1948 (V. G. L. van Someren). venusta (Stagmomantis) Saussure & Zehntner, 1894 :145. I g¢ syntype, [GUATEMALA:] Vera Paz, Teleman, ‘figuré’ (Champion) [tip of abdomen missing]; 1 9 syntype, [GUATEMALA:] Vera Paz, Sinanja, ‘figuré’ (Champion); 1 2 syntype, GUATEMALA: Panima (Champion) [abdomen missing]. versicolor (Hierodula) Henry, 1932: 10. Holotype 3, Srt LanKa: ‘Ceylon’, ‘Reared in Colombo ex eggs laid by a 2 captured at Sigiriya, Ceylon, 30.xi.1931’, ‘First laboratory bred generation’, ‘Killed 23.iii.1932’ (P. D. Ambrose). victorina (Pseudomantis) Westwood, 1889 : 36. Holotype 2, AustRALt1A: ‘North Austr.’, Victoria River Depot, ‘Westw. Mantd. pl. 9, f. 6.’, iii.1856. PRIMARY TYPES OF MANTODEA 327 viridis (Coptopteryx) Giglio-Tos, 1915b : 66. 1g syntype, [ARGENTINA:] Buenos Aires. Additional syntypes in the MNHU, Berlin. vitrea (Acontista) Saussure & Zehntner, 1894 : 138. 1 syntype, [PANAMA:] V. de Chiriqui, 2-3000 ft, ‘figuré’ (Champion). There are also 2 § with identification labels by Saussure and similar data to the above except for collection heights of ‘3—4000 ft’ and ‘below 4000 ft.’ Only ‘2—3000 ft’ is mentioned in the description and these specimens have therefore not been regarded as syntypes. Additional syntype(s) in the MHN, Geneva and perhaps in other depositories. westwoodi (Acontista) Saussure & Zehntner, 1894 : 134. 1 ¢ syntype, [BRaziL:] Tapajos; I 9 syntype, [BRAzIL:] Para. Additional 9 syntype(s) in the MHN, Geneva. westwoodi (Hierodula) Kirby, 1904b : 247. 1 3 syntype, NEPAL: ‘Westw. Mant. pl. 4, fig. 2.” (Hardwicke Bequest); 1 g, 2 9 syntypes, NEPAL: (Hardwicke Bequest). westwoodi (Phyllocrania) Wood-Mason, 1876 : 176. See page 311. westwoodi (Theopompa) Kirby, 1904a : 83. LECTOTYPE ¢ [not 9, as stated by Kirby], GHANA: ‘Gold Coast’, Tamsoo; here designated. Paralectotypes. 1 g [not 9, as stated by Kirby], data as lectotype [abdomen missing]; I g [not 9, as stated by Kirby], [GHANA:] Ashanti; 2 g, data as lectotype [different species, see below]. The 2 3 paralectotypes thought by Kirby to represent the ¢ of his supposed westwoodi 9, are now identified as Theopompella chopardi Roy, 1961. woodmasoni (Hierodula) Werner, 1931 : 1331. Holotype 9, [INp1A:] Madras, Nilambur, Arevacode, (42), 6.1ii.1925 (S. K. Pillar) [published as ‘Pillai’]. REFERENCES BEIER, M. 1930a. Die Mantiden der Ausbeute Collenette aus Britische-Somaliland. Zool. Anz. 91 : 193-198. 1930b. New and rare Mantodea (Orthoptera) in the British Museum. Ann. Mag. nat. Hist. (10) 6 : 432-460. 1931. Three new species of Mantidae (Orthoptera). Ann. Mag. nat. Hist. (10) 7 : 359- 361. 1932. Neue und interessante Mantiden. Bull. Raffles Mus. 6 : 149-154. 19374. Mantodea from the Lake Rudolf Rift Valley Expedition, 1934. Ann. Mag. nat. Hist. (10) 19 : 376-3709. 1937b. Results of the Oxford University Expedition to Sarawak (Borneo), 1932. Mantodea. Proc. R. ent. Soc. Lond. (B) 6 : 177-181. BODENHEIMER, F. S. 1933. Eine neue Evemiaphila-Art (Orth. Mant.). Mutt. dt. ent. Ges. 4 : 79-80. pened W.J. 1822. Travels in the interior of southern Africa. 1. viii + 582 pp. London. Burr, M. 1899. VIII. Descriptions of two new genera and six new species of Orthoptera, collected by Dr. Henry O. Forbes and Mr. W. R. Ogilvie-Grant in the islands of Abd-el- Kuri and Socotra. Bull. Lpool Mus. 2 : 42-45. CuHoparD, L. 1921. Report on the Orthoptera of Mesopotamia and Persia. J. Bombay nat. Hist. Soc. 27 : 47-59, 3 pls. Fapricius, J.C. 1775. Systema entomologiae. xxviii + 832 pp. Flensburgi et Lipsiae. —— 1781. Species insectorum. 1. vi + 552 pp. Hamburgi et Kilonii. Gietio-Tos, E. 1915a. Mantidi esotici. VIII. Acromantinae. Boll. Muset Zool. Anat. comp. R. Univ. Torino 30 : 1-16. 328 J. A. MARSHALL GicLio-Tos, E. 19156. Mantidi esotici genere e specie nuove. Boll. Soc. ent. ital. 46 : 31-108. 1915c. Mantidi esotici genere e specie nuove. Boll. Soc. ent. ital. 46 : 134-200. 1916. Mantidi esotici genere e specie nuove. Boll. Soc. ent. ital. 47 : 3-44. 1917. Mantidi esotici genere e specie nuove. Boll. Soc. ent. ital. 48 : 43-108. Henry, G. M. 1931. New Ceylonese Mantidae. Spolia zeylan. 16 : 123-128, 2 pls. 1932. Observations on some Ceylonese Mantids with descriptions of new species. Spotla zeylan. 17 : 1-19. Hinton, H. E. 1939. Notes on Australian Mantidae (Orthoptera). Ann. Mag. nat. Hist. (11) 4 : 282-289. Kevan, D. K. McE. 1954. Mantodea from northern Kenya and Jubaland. Ann. Mag. nat. Hist. (12) 7 : 455-473. Kirpy, W. F. 1899. On a collection of Mantidae from the Transvaal &c. formed by Mr. W.L. Distant. Ann. Mag. nat. Hist. (7) 4 : 344-353. 1900. A monograph of Christmas Island. xiii + 337 pp., 21 pls. London. 1903. Description of a new species of Mantidae referred to in the above paper. [Shelford, 1903]. Ann. Mag. nat. Hist. (4) 7 : 304. 1904a. IX. - Notes on Mantidae in the collection of the British Museum (Natural History), South Kensington, with descriptions of new species. Ann. Mag. nat. Hist. (7) 13 : 81-88. 19046. A synonymic catalogue of Orthoptera. 1. x + 501 pp. London. La Greca, M. 1952. Specie nuove e note critiche su Mantodei dell’Africa orientale. Annuar. Ist. Mus. Zool. Univ. Napoli 4 : 1-54. 1969. Su alcuni Mantodei dell’Australia e delle Filippine. Annali Mus. civ. Stor. nat. Giacomo Doria 77 : 633-639. LaIpLaw, W. B. R. 1931. A new mantis from Behar, India. Entomologist’s mon. Mag. 67: 88-89, I pl. Pascog, F. P. 1882. Description of a new species of Mantidae. Ann. Mag. nat. Hist. (5) 9 : 423-424. RAMBUR, M. P. 1838-1840. Faune entomologique del’ Andalousie. 2. 336pp., 16 pls. Paris. REDTENBACHER, J. 1892. On the Orthoptera of the island of St. Vincent, West Indies. 2. Familia Mantodea. Proc. zool. Soc. Lond. 1892 : 206-207. Roy, R. 1966. Revision préliminaire du genre Oxypilus Audinet-Serville [Mantodea]. Bull. Inst. fond. Afr. notre (A) 28 : 1420-1486. 1967. Récoltes du Dr. M. Gaillard 4 Kolda, Senegal (1963-1966). Orthoptéres et ordres voisins. Bull. Inst. fond. Afr. noive (A) 29 : 1538-1567. 1969. Compléments a la connaisance du genre Oxypilus. [Mantodea]. Bull. Inst. fond. Afr. notre (A) 31 : 1105-1127. 1972a. Contribution a la connaissance du genre Junodia (Mantodea Hymenopodidae). Bull. Inst. fond. Afr. notre (A) 34 : 550-589. 1972b. Contribution a la connaissance du genre Congoharpax (Mantodea Hymenopo- didae). Bull. Inst. fond. Afr. noire (A) 34 : 857-868. SAUSSURE, H. DE. 1892. Orthoptera Centrali-Americana. Societas ent. 7 : 121-124. SAUSSURE, H. DE & ZEHNTNER, L. 1893. Suborder Orthoptera genuina, fam. Mantidae. Biologia cent.-am. 1 : 123-197, pls 6-10. SHELFORD, R. 1903. Bionomical notes on some Bornean Mantidae. Ann. Mag. nat. Hist. (4) 7 : 293-304. ss SJ6sTEDT, Y. 1909. Waéissenschaftliche Ergebnisse der Schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deuisch-Ostafrikas. 1905-1906. 3(17). 7-147 pp., 3 pls. Stockholm. 1918. Results of Dr. E. Mjéberg’s Swedish scientific expeditions to Australia 1910-1913. 17. Mantidae und Phasmidae. Ark. Zool. 11 : 1-61, 7 pls. Uvarov, B. P. 1921. Evemiaphila fraseri, sp. n., a new Mantid from Mesopotamia. Entomologist’s mon. Mag. 57 : 175-176. 1922. Records and descriptions of Orthoptera from S.W. Asia. J. Bombay nat. Hist. Soc. 28 : 351-370. PRIMARY TYPES OF MANTODEA 329 Uvarov, B. P. 1923. Records and descriptions of Orthoptera from north-west Africa. Novit. zool. 30 : 59-78, 1 pl. 1924. Some new and interesting Orthoptera in the collection of the Ministry of Agriculture, Cairo. Bull. Minist. Agric. Egypt tech. scient. Serv. No. 41, iv + 41 pp., 3 pls. 19274. Some Orthoptera of the families Mantidae, Tettigoniidae and Acrididae from Ceylon. Spholia zeylan. 14 : 85-114, 1 pl. 1927b. A new Mantid from Burma. Ann. Mag. nat. Hist. (9) 19 : 658-659. 1929. IV A. Orthoptera collected in Sinai by Dr. F. S. Bodenheimer and Dr. O. Theodor. In Ergebnisse der Sinai Expedition 1927. viii + 143 pp. Leipzig. 1930a. Notes on Palaearctic Mantidae (Orthoptera). Ann. Mag. nat. Hist. (10) 5 : 631- 633, 1 pl. 19306. Orthoptera collected by Professor T. D. A. Cockerell in Morocco. Bull. Soc. Sci. nat. Maroc 10 : 210-214. 1931a. Notes on the genus Jvis Saussure (Orthoptera, Mantidae). Ann. Mag. nat. Hist. (10) 7 : 234-238, 1 pl. —— 19316. A new Mantid from Baluchistan. Ann. Mag. nat. Hist. (10) 8 : 418-419. 19334. Notes on new and little-known Orthoptera from Palestine. Ann. Mag. nat. Hist. (10) 11 : 663-672. 19336. Orthoptera collected by Mr. Bertram Thomas in Southern Arabia. Proc. zool. Soc. Lond. 1933 : 259-271. 19354. Two new Orthoptera from Morocco. Bull. Soc. Sci. nat. Maroc 15 : 41-43. 19350. Some Orthoptera from the high regions of the Mt. Cameroon, West Africa. Ann. Mag. nat. Hist. (10) 16 : 205-210. 1936a. Studies in the Arabian Orthoptera.—I. Descriptions of new genera, species and subspecies. J. Linn. Soc. Lond. (Zool.) 39 : 531-554, 2 pls. 1936b. New Orthoptera from Cyprus. Ann. Mag. nat. Hist. (10) 18 : 505-515. 1939a. Studies in the Arabian Orthoptera.—II. New and little-known Mantidae and Phasmidae. J. Linn. Soc. Lond. (Zool.) 40 : 547-559. 19396. New and less-known Palestinian Orthoptera. Ann. Mag. nat. Hist. (11) 4 : 216- 229; 7.0 1948. Andalusian Orthoptera described by Rambur. Eos 24 : 369-390. WALKER, F. 1859. Characters of some apparently undescribed Ceylon insects. Order Orthoptera. Ann. Mag. nat. Hist. (3) 4 : 220-221. WERNER, F. 1931. Further notes on Indian Mantids or praying insects. Proc. zool. Soc. Lond. 1931 : 1329-1334, 1 pl. 1933. Third contribution to the knowledge of Indian Mantids, or praying insects. Proc. zool. Soc. Lond. 1933 : 897-901. 1935. Further communication on Indian Mantids or praying insects. Proc. zool. Soc. Lond. 1935 : 495-408. WESTWOOD, J. O. 1843-45. Arcana entomologica, or illustrations of new, rave and interesting insects. 2. 192 pp., 47 pls. London. —— 1889. Revisio insectorum familiae Mantidarum, speciebus novis aut minus cognitis descriptis et delineatis, 52 + iii pp., 14 pls. London. Woop-Mason, J. 1878. On new and little-known Mantidae. Proc. zool. Soc. Lond. 1878: 580-587, 2 pls. 1879. Description of Sigerpes occidentalis, the type of a new genus of Mantodea from West Africa. Proc. Asiat. Soc. Beng. 1879 : 257-259. Mrs, J. A. MARSHALL, B.Sc. Department of Entomology BRITISH MusEuM (NATURAL History) CROMWELL Roap Lonpon SW7 5BD ENTOMOLOGY SUPPLEMENTS Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August, 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OxapDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, 9 maps. February, 1967. £3.50. . Hemminc, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8. . Mounp, L. A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . ArirI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 51 text- figures. December, 1968. £5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Extot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. £4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 108: 68 plates, 15 text-figures. October, 1971. £12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera : Termitidae). Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. . CrosskEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . von Hayex, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CrossKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND De aD Be THE PROTURA Coin wo OF THE BISMARCK ARCHIPELAGO. AND SOLOMON ISLANDS S. L. TUXEN AND G. IMADATE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 9 LONDON : 1975 THE PROTURA OF THE BISMARCK ARCHIPELAGO AND SOLOMON ISLANDS BY: SOREN LUDVIG TUXEN | Universitetets Zoologiske Museum, Copenhagen AND GENTARO IMADATE Tokyo Medical and Dental University Ph. 331-375; 76 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 31 No. 9 LONDON : 1975 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, ts issued in five series corresponding to the Departments of the Museum, and an Historical series. Paris will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary serves of longer papers was instituted, numbered serially for each Department. This paper is Vol. 31 No. 9 of the Entomological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Pertodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Ent.) ISSN 05 24-6431 © Trustees of the British Museum (Natural History), 1975 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 24 April, 1975 Price £3.00 THE PROTURA OF THE BISMARCK ARCHIPELAGO AND SOLOMON ISLANDS By S.L. TUXEN & G. IMADATE CONTENTS Page SYNOPSIS : ‘ : . 7 : : : 7 . . 333 INTRODUCTION ; ; : ‘ : : : , : : 333 LIsT OF LOCALITIES . : : : : : : : . : 334 KEY TO THE KNOWN MELANESIAN PROTURA ‘ : ; : : 335 ACERENTOMIDAE F : F . : é : : : : 330 PROTENTOMIDAE ; : : ; , : : ; a. : 346 EOSENTOMIDAE : : : j : : , : : : 350 GENERAL DISCUSSION : , ‘ ; 4 : : : : 370 Ecology : : ; ; : : : ‘ ‘ * ; 370 . Zoogeography : : : ‘ é : é F . : 371 Taxonomy . : ‘ , ‘ ‘ ; ‘ : : ‘ B73 REFERENCES . ; - : : ; , , ‘ ‘ : 373 INDEx . ‘ : ‘ : : : ‘ , ; ; : 375 SYNOPSIS An account is given, with keys, of the Protura of the Bismarck Archipelago and Solomon Islands, making use of recently acquired material in the Zoological Museum, Copenhagen, and the British Museum (Natural History). Twelve species, eleven of them new, are added to the single species previously known from this region. INTRODUCTION AT THE time of the senior author’s revision of the world species of Protura (Tuxen, 1964) only one Melanesian species was known, Silvestridia solomonis (Imadaté) from Guadalcanal. Since then, much more material has become available from the following sources. I. Noona Dan Expedition of the Zoological Museum, Copenhagen (1962), soil fauna samples from various localities in the Bismarck Archipelago and Solomon Islands (Petersen, 1966; Wolff, 1968). Specimens placed in Zoo- logical Museum, Copenhagen. 2. Royal Society Expedition to the Solomon Islands (1965), soil samples placed in British Museum (Natural History), collections made by Mr P. N. Lawrence and Dr P. J. M. and Mrs P. Greenslade. 3. Collections of soil arthropods made by Dr and Mrs Greenslade in the Solomon Islands during the years 1964-1966, specimens placed in British Museum (Natural History). 334 S.L. TUXEN & G. IMADATE We are grateful to Dr and Mrs Greenslade and to Dr Theresa Clay for making the last two groups of soil samples available to us. In total, these samples contained 450 specimens of Protura from more than 35 different localities and proved to comprise 13 species belonging to three families. It is now possible for a synoptic study to be made of the species from these islands following the lines of Tuxen (1964), where details of the genera will be found, includ- ing diagnoses and keys for their separation. All figures are drawn from the holotypes. We have followed the schemes proposed by Tuxen (1949) and Imadaté (1965 : 198), and for each species set out the chaetotaxy as follows: if all stages are known, setae are marked primary (‘prim.’), secondary (‘sec.’), tertiary (‘tert.’) and complementary (‘compl.’) for larva I, II, maturus junior and adult respectively. If larva I is not known, the chaetotaxy of the first known stage is marked ‘composition of setae’. In the first column for every stage is given the formula according to Tuxen’s system. It should be noted that what Tuxen called no. 3 in the anterior row on the abdomi- nal terga is here called ‘P 3’, namely no. 3 in the posterior row. Tuxen (1964 : 62) noted that ‘ ‘‘3”’ of the anterior row is often situated behind the other setae of this row and may in terg. VII lie so far back that it can be reckoned as belonging to the posterior row’. As shown by Imadaté (1965 : 197) from a study of postembryonal development of the chaetotaxy, actually the reverse is the case: it is P 3 from the posterior row which is placed anteriorly in some of the abdominal tergites. This should be taken into consideration when comparing our ‘formulae’ with the earlier ones by Tuxen and others. The following abbreviations are used for depositories of material cited in this paper. BMNH British Museum (Natural History), London KU Kyoto University, Kyoto, Japan. UZM Universitetets Zoologiske Museum, Copenhagen, Denmark. LIST OF LOCALITIES On the map (Text-fig. 76) the collecting sites are numbered as follows (the island in italics). BISMARCK ARCHIPELAGO (Noona Dan Expedition). 1. Boliu, Mussau, 4.vi.1962. Soil with rotten leaves. Malakata, Mussau, I1.vi.1962. Soil and half-rotten leaves, primary forest. 3. Banatam, Lavongai, 20-26.iii.19g62. Half-rotten leaves and soil. Mangrove swamp (20.iii) and primary forest (26.iii). 4-5. Sumuna and Kollepine, Dyaul, 7—-12.iii.1962. Half-rotten leaves and soil, primary forest. 6. Lemkamin, New Ireland, 23.iv.1962. Grass tufts and soil, g00 m. 7. Manuan, Duke of York, 21.vii.1962. Soil, secondary forest. 8. Valoka, New Britain, 13.vii.1962. Soil and litter, primary forest. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 335 SOLOMON IsLAnps (P. J. M. & P. Greenslade and P. N. Lawrence collections as Q. 10: 5 a 12. 13. I4. 15. 16. I6a. r7; 18. IQ. 20. ai. 22. 23. 24. 25. 26. 277. 28. 29. 30. ST; a2. 33: 33a. 34. 35. well as two sites from Noona Dan Expedition (No. 28 & 34)). Falomai village, Mono, Treasury group, 20.ix.1965. Soil. Kuturasele, Choisewl, 23.xii.1965. Soil. Wagina, Choiseul group, 25.v.1965. Gizo, Nusatupe, New Georgia group, 4.vi.1965. Litter. Kuzi, Kolombangara, New Georgia group, 6.xi.1965. Buala, Santa Isabel, 14.xii.1964. Litter. Tatamba, Santa Isabel, 27.1x.1965. Yandina, Russell, 15.vill.1966. Russell, 18.11.1965. Forest soil. Vatilau, Nggela, Florida group. 14.vi.1965. Soso, Nggela, 2.x11.1965. Forest litter. Koela, Savo, Guadalcanal group. 1.vi.1965. Litter. One Naika, Savo, I.vi.1965. Litter. Mt Austen, Guadalcanal, 330 m, 24.x.1964-10.viii.1966. Forest litter. Mt Jonapau, Guadalcanal, 1100 m, 6.v.1965. Forest litter. Mt Popamanaseu, Guadalcanal, 26.vi.1965. Forest litter. Nr Malukuna, Guadalcanal, 9.vi.1966. Betimatu, Guadalcanal, 31.xii.1964. Forest litter, bush. Berande Point, Guadalcanal, 30.iv.1965. Forest litter, sea level. Kukum, Guadalcanal, 15.xil.1964—-14.vii.1966. Honiara, Guadalcanal (Noona Dan Expedition), 4.viii.1962. Lunga, Guadalcanal, viii.1966. Dala, Malaita, 16.11.1965. Forest litter. Warahitor, San Christoval, 1.viii.1965. Bellona, Rennell group, 28.v.1965. Garden litter. Kagaba Bay, Rennell, 6.iv.1965. Litter and soil from Coconut-grove. Kagaba Bay, Rennell, 6.iv.1965. Rotten tree trunk, primary forest. Niupani, Rennell (Noona Dan Expedition), 18-19.viii.1962. Graciosa Bay, Ndeni, Santa Cruz Group, 6.iv.1965. KEY TO THE KNOWN MELANESIAN PROTURA Spiracles absent; only one or two pairs of the abdominal legs two-segmented (Acer- entomoidea) i 2 Spiracles present; all three pairs of abdominal legs ver ‘segmented (Bosentomoidea; only genus Eosentomon) ; ‘ 6 The two first pairs of abdominal legs two- -segmented é _ Condeellum crucis G 346) Only the first pair two-segmented . : . . 3 Second and third pair of abdominal legs with only one 2 seta Silvestridia solomonis (p. 336) These abdominal legs with two setae, a long subapical seta and a very short apical one . ; : ; ; , 4 Striate band of abdominal segment Vv II! well developed Gracilentulus greensladei (p. 340) Striate band reduced (Berberentulus) : ’ ° ‘ ‘ : ‘ ; 5 336 S.L. TUXEN & G. IMADATE 5 Body length ab.800y4m; sensilla e of foretarsus much shorter than c; A 1 missing in tergite V . ‘ . B. buchi (p. 340) — Body length ab. 900 pm; ‘sensilla eas 5 Jong as C; A I present i in tergite V B. rennellensis (p. 338) 6 Caput processus of female squama genitalis semicircularly rounded; P 2 on abdominal terg. II-VI displaced anteriorly. Foretarsal sensilla ¢-1 situated close to « 3, b’—1 absent, P 2 and 1” on terg. VIII not displaced anteriorly, foretarsus 63 wm E. noonadanae (p. 367) — Caput processus bent against median edge of stylus in a sharp nl P 2 on terg. II- VI situated in a row with P 1 and/or P 3 : 7 7 Foretarsal sensilla t-1 placed at about midway between x 3 ‘and a 3’, bx present; P 2 and 1” on abdominal terg. VIII not eee anteriorly, stern. VIII with o—7 setae . : : 8 — Foretarsal sensilla ix placed elase to 4 3', b’-1 absent; P 2 and 1” on terg. VII displaced anteriorly, stern. VIII with 2-7 setae : 12 8 Accessory setae P ta on abdominal terg.VI short, less than one-fifth of P I and ; situated in row with P 1 and 2. Abdominal terg.IV with A 4 and 5, terg.V-VI without A 1, 2 and 3 or A 1, 2, 3 and 4, terg. VII without A 1, 2, 3 and 4, terg. X-XI with 8 setae, foretarsus 94-104 wm. : ; E. oceaniae (p. 350) — Accessory setae P 1a on terg.VI long, more than P 1 and situated in a row with the other accessory setae . 9 9 Abdominal terg. X—XI with 8 setae, P Ia on n tere. VII placed i in a row with the other accessory setae. Terg.IV without A 3, terg.V—-VI without A 1, 2 and 3, terg. VII without A 1, 2, 3 and 4, foretarsus 97-98 wm : E. solomonense (p. 361) — Abdominal terg. X—XI with less than 8 setae, P 1a on terg. VII acai in row with Ptand 2 10 10 6Terg.X with 4 setae (I and 4). " Terg. IV with 10 anterior setae (4 £2; ss 4 and 5), terg.V—VI without A 1, 2 and 3, ele VII without A 1, 2, 3 and 4, foretarsus 85-95 um. 5 ' i : : E. sakura (p. 359) — Terg.X with only 2 setae in : : ; ; : . : 1 11 Terg.IV with to anterior setae. Terg.V—VI without A 1, 2 and 3, terg.VII without A I, 2, 3, 4 and 5; female squama re with latero-proximal sclerotization, foretarsus 70-76 wm . : . E. solare (p. 356) — Terg.IV without A 1, 2 and 3 or A 1, 2, 3 and Pm Terg. V-VI without At, 2,3:ana4 or A 1, 2 and 3, terg. VII without A 1, 2, 3 and 4; female squama genitalis without latero-proximal sclerotization, foretarsus 73-77 wm ; E. melanesiense (p. 352) 12 Empodium of hind tarsus short and less than one-fifth claw length; foretarsal sensilla a’ placed nearer to « 3’ than to « 3. Terg. II-VI without A 3, terg.VII without A 1 and 3, foretarsus 95 wm . ; ‘ . E. guadalcanalense (p. 363) — Empodium of hind tarsus long and more than one-third claw length; foretarsal sensilla a’ placed nearer to « 3 than to a 3’.. Terg. II-VI with 10 anterior setae, terg.VII without A 1 and 3, foretarsus 57-62 wm . ‘ F . E. notiale (p. 365) ACERENTOMIDAE One-hundred and twenty-three specimens from 14 localities are found in the present collection. They consist of four species, viz. Silvestridia solomonis (Imadaté), Berberentulus rennellensis sp. n., B. buchi sp. n. and Gracilentulus greensladet sp. n. Silvestridia solomonis (Imadaté) Acerentulus solomonis Imadaté, 1960 : 5-8. Holotype 9, Sotomon Is.: Guadalcanal, River Tantu, 20.ix.1958 (T. Tokioka) (lost). PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 337 Silvestvidia solomonis (Imadaté) Tuxen, 1964 : 328-329. Silvestridia solomonis (Imadaté); Tuxen & Imadaté, 1974 : 85-88 (redescription). SPECIMENS EXAMINED. SoLomon IsLanpDs: Savo, Koela, 1.vi.1965 (P. J. M. & P. Greenslade), 1 3, 4 2 (BMNH); Guadalcanal, Mt Austen, I1.ii.1965, I 2; 2I.iv.1965, I 9; 24.v.1965, 1 2 Fics 1, 2. Berberentulus rennellensis. Yoretarsus, (1) interior and (2) exterior views. 338 S. L. TUXEN & G. IMADATE I Q; 8.11.1966, 1 Q (all P. J. M. & P. Greenslade) (all BMNH); G., nr Malukuna, g.vi.1966 (P. J. M. & P. Greenslade), 1 9 (BMNH); Ndeni, Graciosa Bay, 6.iv.1965 (P. J. M. & P. Greenslade), 1 29 (BMNH). Differs from other species of the genus Silvestridia by the absence of foretarsal sensilla 0’. Berberentulus rennellensis sp. n. (Text-figs I-10) Body length 900 wm in expanded adults. Mouthparts small, sensillae on maxillary palpus slightly broadened, that on labial palpus distinctly broadened; pseudoculus almost circular, PR = 13-14; canal of the maxillary gland of the normal shape, its proximal part rather short. Foretarsus 68 um in length; claw with no inner flap; TR = 3:3; empodium short, EU = 0*12-0'14; position and shape of foretarsal sensillae similar to that of B. yodai Imadaté. Dorsal sensilla t-1 claviform, BS = 0-41; #2 thin and ¢-3 small. Exterior sensilla a relatively long, apex almost reaching base of y 3; b slightly broadened and extremely long, its apex slightly surpassing the base of y 4; c situated close to d and subequal to d in length; apices of c and d not surpassing base of « 5; e as long as c and by far surpassing base of g and ¢-3; f thin and long and close to e; g slightly surpassing the tarsus. Interior sensilla a’ broad and located in Fics 3-10. Berberentulus rennellensis. 3, pseudoculus; 4, maxillary palpus; 5, labial palpus; 6, canal of maxillary gland; 7, abdominal appendage II; 8, dorsal chaetotaxy of abd. VII-VIII; 9, comb on terg. VIII; 10, female squama genitalis. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 339 TABLE I Chaetotaxy of Berberentulus rennellensis Larva II Adult (Dorsal) Formula Comp. of setae Formula Tert. and compl. setae ans 4 r2 4 II-III 6 A 2,4, M 6 12 I 4224,3 455 Te Pita Abd.I oO 6 At;2,5 Io P 1,2,24,3,5 ‘12 Pia II-III fe) 6 A: £5255 12 P 1,2,24,3,4,5 16 P 1a,4a IV fe) 6 A 1,2,5 Ta P 1,2,24,3,4,44,5 16 Pia V fe) 6 A 1,2,5 74 P 1,2,24,3,4,44,5 16 Pia Vi oO 4 War | vi P 1,2,24,3,4,4@,5 16 Pia VII oO 2 A5 14 P 1,2,24,3,4,4@,5 16 Pia VIII 2-6 A 3, M 2,3.4 4-7 A 5, Mic 8 FP 1;2,3,4 8 IX 8 1,3,4,5 14 2,34,4a xX 12 1,2,3,34,4,5 XI 6 Telson 9 9 (Ventral) Th.I 2-2 A1,M1 4-2 A2 hap P1,2 “6. P3 II-III 5-2 A ¢,2,3, M 5(7)-2 (A 4) 2 foie 4 EZ Abd. I 3 A 6,2 3 2 yes Pi te II-III I Ac 3 A:'2 5 F6,2,;3 5 IV-VII I Ac 3 eB 6 P 1,2,3 8 Pia VIII 4 4 oO ° IX 4 4 x 4 XI 4(6) Telson 8 6 340 S. L. TUXEN & G. IMADATE row with ¢-1; b’ absent; c’ thin and a little distal to « 5, apex slightly surpassing the tarsus. Chaetotaxy (Table 1) similar to that of B. nitidus Imadaté & Yosii, but P 2 on abdominal stern. I present and A 1 on terg. VI absent, but present on terg. I-V. Abdominal appendages II-III with two setae each, the apical one less than half the subapical. The striate band on abd. VIII reduced, no distinct striae; comb on terg. VIII oblique, consisting of 12-13 minute teeth; female squama genitalis with pointed acrostylus. Larva II. ¥oretarsus 50 wm. Chaetotaxy typical for the genus. Holotype 3, SOLOMON IsLANDs: Rennell Island, Kagaba Bay, 6.iv.1965 (P. J. M. & P. Greenslade) (BMNH). Paratypes. Same data as holotype, 1 g, 1 9 (BMNH). This species closely resembles B. yodat Imadaté from southern Thailand in such characters as the position and shape of most of the foretarsal sensillae, in the absence of b’ as well as in the tergal chaetotaxy of the abdomen, but it may be distinguished by the presence of A 1 on abdominal terg. IV-V. Berberentulus buchi sp. n. Body length 770-800 wm in expanded adults. Mouthparts as in preceding species; pseudoculus almost circular, PR = 11-13; canal of maxillary gland of normal shape, its proximal part rather short. Foretarsus 51-56 um in length; claw with no inner flap, 7R = 3-1-3:2; empodium short, EU = 0:16-0:18; position and shape of foretarsal sensillae similar to that of the preceding species, except that sensilla e is short, much shorter than c and just reaching base of #3. The chaetotaxy (Table 2) similar to that of the preceding species, but A 1 is always missing on terg. V, and sometimes also on terg. ITI-IV or even II (2 f and 19 from Valoka). Abdominal appendages II-III, the striate band and comb on abdomen VIII and female squama Sous as in the preceding species. Muturus junior. Foretarsus 43-48 um. Important characters such as foretarsal sensillae, canal of maxillary gland, etc. as in adult. Larva II. Foretarsus 37-44 wm. Chaetotaxy typical. Holotype 9, BisMARCK ARCHIPELAGO: New Britain, Valoka, 13.vii.1962 (Noona Dan Expedition) (UZM). Paratypes. BISMARCK ARCHIPELAGO: Mussau, Boliu, 4.vi.1962 (Noona Dan Expedition), 2 9, 1 1II (UZM); New Britain, Valoka, 13.vii.1962 (Noona Dan Expedt- tion) 8 J, 8 9, 2 m.j., 21 II (UZM). The present species is named in honour of Mr Will. Buch who made the soil collections (as well as many others) on the Noona Dan Expedition. It is closely similar to the preceding species and B. yodai Imadaté from southern Thailand but, because of the difference in length of sensilla e, the body size and the different chaetotaxy of terg. IV—V, we prefer to regard it as a distinct species. Gracilentulus greensladei sp. n. (Text-figs 11-21) Body length 700-810 wm in expanded adults. Mouthparts of normal type, sensillae on maxillary palpus slightly broadened, that on labial palpus also broadened; pseudoculus almost circular, PR = 11-13; canal of maxillary gland of normal shape, its proximal part rather short. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 341 TABLE 2 Chaetotaxy of Berberentulus buchi Larva II Mat. jun. Adult ‘Devoall) Form. Comp. of setae Form. Tert. setae Form. Compl. setae Th.I 4 12 4 4 II-III 6 A 2,4,M 6 6 I2 P 1,2,24,3,4,5 ta: Pia 14 Abd.I fe) 6 A i235 6 10 P 1,2,24,3,5 To 12 Pia II-III fo) 6 A t:205 6 12 F-1,2,24,3)4,5 TA P 4a 16 Pia IV oO 6 At 25 6 Fa. P4,2;28,3;4,48;5 ae 16 ‘Pera V oO 4 AKZ.s 4 14 P 1,2,24,3,4,44,5 14 16 Pia Vi fo) 4 A 25 4 14 P 1,2,24,3,4,4@,5 14 16 1a VII fe) 2 As 14 P 1,2,24,3,4,44,5 14 16 Pia VIII 2-6 A 3,M 2,3,4 4-7 As5Mc 4-7 a F552.3:4 “Bo 8 IX 8 1,3,4,5 12 2,4a 14 3a xX 8 1,3,4,5 12 2,3a XI 6 Telson 9 9 9 (Ventral) Th.I 2-2 A1,Mr1 4-2 Ar? 4-2 ye P1,2 26> P3 ce. II-III 5-2 A ¢,2,3,M 5-2 5(7)-2 (A 4) hae Pi i as Fa 4 Abd. I 3 Ac,2 3 3 2 2 ae 4 E2 4 II-III I Ac 3 Aa 3 5 P c,2,3 5 5 IvV-VIlI I Ac 3 A2 3 6 F8,2 3 s. Pia 8 VIII 4 4 4 ° ry ° IX 4 4 4 x 4 4 XI 0(2) 4(6) Telson 8 6 6 342 S. L. TUXEN & G. IMADATE Foretarsus 51-56 wm in length. Among 76 adult specimens from seven different localities no local variation was found in the foretarsal length. Claw with no inner flap, TR = 2-8—3:1; empodium short, EU = o-11-0-16; dorsal sensilla #1 claviform, BS = 0-42-0-44; #2 thin and ¢-3 small; exterior sensilla a slightly broadened and long, apex almost reaching the base of « 4; 6 also a little broadened and long, its apex reaching the base of y 4; ¢ situated close to d and subequal to d in length; c and d shorter than b; e and f close together; apices of f and g slightly surpass the tarsus. Interior sensilla a’ slightly distal to ¢-1 and broadened, b’ thin and situated in a row with #2; c’ thin. Abdominal chaetotaxy (Table 3) peculiar by the reduction of tergal anterior setae: A 1 on abdominal terg. II-VI absent as in Silvestridia; on terg. VII only a single pair of anterior setae, A 5, present; terg. VIII with two pairs of anterior setae, A 3 and 5. Abdominal appen- dages II-III have two setae each, the apical one being less than half the subapical. Striate az 12 Fics 11, 12. Gvracilentulus greensladei. Foretarsus, (11) interior and (12) exterior views. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 343 13 14 19 »\ T. Fics 13-21. Gracilentulus greensladei. 13, maxillary palpus; 14, labial palpus; 15, pseudo- culus; 16, canal of maxillary gland; 17, dorsal chaetotaxy of abd. VI-VIII; 18, striae on the band of abd. VIII; 19, comb on terg. VIII; 20, abdominal appendage IT; 21, female squama genitalis. 344 S. L. TUXEN & G. IMADATE band on abdomen VIII reduced to some degree; striae distinct in the anterior part of the band, but not visible in the posterior part; comb on terg. VIII consisting of about 10 small teeth; female squama genitalis with long and pointed acrostylus. Maturus junior. Foretarsus 47-48 um. Important characters such as foretarsal sensillae, comb on abdominal terg. VIII, etc. as in adult. Larva II. Foretarsus 42-43 wm. Foretarsal sensilla b’ not found in the three specimens examined; striae in band on abdomen VIII fainter than those of the adult. Larva I. Foretarsus 37 wm. Position and shape of foretarsal sensillae, the striate band on abdomen VIII, etc. as in the larva II. Holotype g, SoLomon IsLanps: Guadalcanal, Kukum, 15.xii.1964 (P. J. M. & P. Greenslade) (BMNH). Paratypes. SOLOMON ISLANDs: Santa Isabel, Buala, 14.xii.1964 (P. J. M. & P. Greenslade), 49 (BMNH); S.I., Tatamba, 27.ix.1965 (P. N. Lawrence), 19 (BMNH); Neggela, Vatilau, 14.vi.1965, 1 9; N., Soso, 2.xi.1965, 19 (all P. J. M. & P. Greenslade) (all BMNH); Guadalcanal, Kukum, 15.xii.1964, 16 9, 2 m.j., 3 1 II; G., Kukum, vi.—vii.1966, 6 9 (all P. J. M. & P. Greenslade) (all BMNH); G., Lunga, viii.1966 (P. J. M. & P. Greenslade), 46 9, 1 m.j.,111(BMNH). BisMARCK ARCHIPELAGO: Duke of York, Manuan, 21.vii.1962 (Noona Dan Expedition) 1 9 (UZM). This new species is named in honour of Dr P. J. M. and Mrs P. Greenslade, who made such a rich proturan collection in the Solomon Islands. G. greensladet is peculiar, among all known forms of this genus, in the position of foretarsal sensillae c, d, e, and f, as well as in the size of a and b. KEY TO THE KNOWN SPECIES OF GRACILENTULUS 1 Canal of maxillary gland with long proximal part with two or three oval dilatations along the very narrow tube . , : Canal of maxillary gland with short proximal part with end dilatation : Sensilla 6 shorter than half of a : : ‘ Sensilla b longer than a % Sensilla b extremely short and thin, only reaching v2; 10 and a ina line Sensilla b almost reaching base of y 3 and surpassing t-2; b’ midway between ¢-2 and % 4; @ surpassing ¢-2; terg. I-VI with six anterior setae japonicus (Imadaté, 1961) 4 Sensilla a only just reaching c; terg. V-VI with only four anterior setae malaysiensis Imadaté, 1964 — Sensilla a reaching t-2; terg.V—VI with six anterior setae (or eight) kenyanus (Condé, 1948) 5 Sensilla b very long, almost ae gia ae y 3; smaller, foretarsus about 50 wm : : ohyamai Imadaté, 1964 — Sensilla b shorter, hardly reaching é;a reaching yar bigger, foretarsus about 75 wm sanjianus Imadaté, 1964 wits | kW An 6 Sternite VIII with two posterior setae ‘ : : ‘ é : . . 7 — Sternite VIII with only the four anterior setae . : : ; 8 7 Canal of maxillary gland long, as long as proximal branch ‘of fulcrum: b’ present; terg. VII with eight anterior setae . , : 3 tasmanicus Tuxen, 1967 - Canal of maxillary gland short, half the proximal branch of fulcrum; b’ absent; in terg.VII A 1 is missing 3 . similis Tuxen, 1967 8 Sensilla b’ missing; b as long as c and d; comb VIII with short teeth floridanus (Ewing, 1924) PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 345 TABLE 3 Chaetotaxy of Gracilentulus greensladei Larva I Larva II Mat. jun. Adult Form. Prim. setae Form. Sec. setae Form. Tert. set. Form. Compl. setae (Dorsal) pu ay 22-2 4 4 II—III 4 A2,M 6 A4 8 P1,2,3,5 (4?) 14(12) Pra, (2a),4 14 14 Abd.I o oO 6 Al t2;5 6 8 Oo Fe Fa Io Ea To 12 Pra II-III o oO 4 A 2,5 4 TO P1,2;3,4;5.- 12(14) fF 24;(4a) I4 16 Pia IV-VI o fe) 4 Aes 4 IO PE 23 as (TA(t2)) 22a; (aa) Tas 16 Pxka VII o fe) 2 A5 2 10 Pi5,25354;5 1 P 2a,4a 16 P 3a 18 P ia VIII 0-6 M 2,3,4 2-6 As 4-7 A5,Mc 4-7 6 Boge BU Ey 8 P.3 8 8 IX 8 1,3,4,5 10 2 12 3a xX 8 1,3,4,5 12(10) 2,(3a) XI 6 6 Telson 9 9 9 9 (Ventral) Sick. of a2 Avy, Mt 4-2 A2 4-2 4 Pt,2 “6 P3 x II-III ? 5-2 Ac,2,3,M 5-2 7(5)-2 (A 4) 2 ek 4 en 4 Abd.I o 3 A052 3 3 2 Py 2 2 4 y da II-III o 1(0.) Ac 3 Az 3 3 Pe,3 5(3) P2 5 5 IV-VII 1 Ac I 3 Az 3 re P23 6 Y gas 8 P ta 8 Willi 2 2 4 I 4 4 ° ° ° O Ix 4 4 4 x 4 4 XI o(2) (2) 64) 14.3 Telson 8 8 6 6 346 S. L. TUXEN & G. IMADATE — Sensilla b’ present; b shorter or longer thanc . 9 9 Sensilla b longer than c, reaching ¢-3 ; b-c—d in a line; f close to é; terg. II-VI with four anterior setae, terg.VII with only two (A 5) . . greensladei sp.n (p. 340) — Sensilla 6 shorter than c; d distal to b-c; terg. II-VI with six anterior setae . Z 10 Io Sensilla f close to e ; % ; 2 : ~ sachikoae Imadaté, 1964 — Sensilla f midway between e and Boos s : , , ‘ A II 11 Sensilla b short, just surpassing base of d . : ‘ , meridianus (Condé, 1945) — Sensilla b longer, reaching base of e . ; : : . gracilis (Berlese, 1908) PROTENTOMIDAE Twelve protentomid specimens belonging to a single species of the genus Condeellum are present. It is interesting to note that they were all collected from Mt Austen (330 m) on Guadalcanal Island. Condeellum crucis sp. n. (Text-figs 22-35) Body length 790-820 wm in well expanded adults. Mouthparts rather broad; mandible not pointed, but has a blunt end with possibly a few minute teeth; maxillary and labial palpi ornamented with a tuft of setae apically; lobus internus of maxilla rather broad and lobus externus pointed distally; labium with five setae: three relatively long and situated along the outer margin, one short seta placed near to inner margin and a little distal to the palpus, and one short seta at the tip, as in all other members of this genus. Pseudoculus oval, PR = 9-10; canal of the maxillary gland of the normal shape for the genus, proximal part sharply bent at about halfway between the heart-shaped dilatation and the end. Foretarsus 36-44 um in length, claw with an inner tooth, TR = 2+7-2-9; empodium short, EU = 0-1; position and shape of foretarsal sensillae similar to that of other members of the genus. Dorsal sensilla ¢-1 not broadened, BS = 1:0; #2 a little thinner than ¢-1 and pointed apically; #3 broad; exterior sensilla a subequal to ¢-3 in shape and length; b and f slightly broadened, but smaller than a; f placed a little ventrally and distally to y 4; c, d, eand g absent; interior sensilla a’ proximal to f-1, short and slightly broad; b’ and c’ absent. Middle and hind tarsi have long empodia and tunica-lobes; middle tarsus 18-21 wm, and its claw 10-13 pm; hind tarsus 19-22 wm and its claw 11-14 pm. Thorax I with two pairs of dorsal setae, pair 1a absent. Thorax II-III with dorsal P 5 rudimentary. Abdominal terg. I-VI with only a single pair of anterior setae, A 1, terg. VII no anterior setae. Stern. II-VII with a central seta, P c, present in the posterior row. Ab- dominal appendages I-II two-segmented with four setae; III rudimentary with two short apical and one long subapical seta. Comb on abdomen VIII consists of about to small teeth; female squama genitalis with pointed acrostylus. Chaetotaxy as in Table 4. Maturus junioy. Foretarsus 33-35 wm. Important characters such as foretarsal sensillae, canal of the maxillary gland, etc., as in the adult. In comparison with the chaetotaxy of the adult, setae 2 on abd. terg. X, setae 1 and 3 on stern. XI, are absent. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 347 24 23 Fics 22-25. Condeellum crucis. 22, pseudoculus and canal of maxillary gland; 23, hindtarsus; 24, 25, foretarsus, (24) interior and (25) exterior views. 348 S. L. TUXEN & G. IMADATE Fics 26-35. Condeellum crucis. 26, 27, chaetotaxy of abd. VI-XII, (26) dorsal and (27) ventral views; 28, comb of terg. VIII; 29, female squama genitalis; 30, abdominal appendage II; 31, abdominal appendage III; 32, mouthparts, ventral view; 33-35, labium in lateral view of (33) C. ishiianum, (34) C. matobai and (35) C. regale. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 349 Holotype 2, SoLomon IsLAnps: Guadalcanal, Mt Austen, 330m, I1.ii.1966 (BMNH). Paratypes. Same data as holotype, 6 9, I m.j.; 28.11.1966, I 9; 7—-10.vi.1966, I 9; 10.viii.1966, 2 9 2 m.j. (all P. J. M. & P. Greenslade) (all BMNH). The species name is derived from the Latin word ‘crux’ in reference to the Southern Cross, the famous constellation of the southern hemisphere. This new species is peculiar among Condeellum in the absence of anterior setae on abdominal terg. VII. All known species of this genus may be separated by the chaetotaxy of the abdominal terg. II—-VII, as given in the following key. TABLE 4 Chaetotaxy of Condeellum crucis Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) Tht 4 1,2 ‘hel 2-4 A M 1,2 I-III 6 A 2,4, M 6 P 1,2,3 12 P 1 ,14,2,28,3;4 II 4-2 A 1,2, M Abd.I 2 At 4 ae Tr P 1,14,2,24,3,4,5 III 6—2 A 1,2,3, M II-VI 2 At 4 ee 14 P 1,2,24,3,4,4@,5 Abd.I 4 A235 VU ° Z ses 18 P 1,14,2,24,3,34,4,4@,5 FL-1E 4 A 2,3 VIII 6-6 A 1,3,5, M 2,3,4 5(3) Pe, (2),3 = FP 3,2;3,4 IV-VI 4 A; 28 IX 12 157:3,30.455 7(9) P.0,%, (1a);:2;3 xX 10 1,2,3,4,5 Vit 4 A 2,3 9 P €,1,14,2,3 XI 6 VIII 6 I,1@,2 Telson 9 IX-X 4 >. Gi 6 12,3 Telson 6 KEY TO THE KNOWN SPECIES OF CONDEELLUM 1 Abdominal terg. II-VI with a single pair of anterior setae, AI . 2 — Abdominal terg. II-VI with two or three pairs of anterior setae (cluding a pleural pair, A 5) ° 2 Terg. VII with two pairs of anterior setae, A I and 53 " foretarsus 46 am (Japan) matobai Imadaté, 1973 350 S. L. TUXEN & G. IMADATE — Terg. VII without anterior setae; foretarsus 36-44 wm (Solomon Is.) crucis sp.n. (p. 346) 3. Terg. II-VI with three pairs of anterior setae, A 1, 2 and 5; terg. VII with A 4 and 5; foretarsus 42-54 um (Thailand, Borneo) 2 . ishiianum Imadaté, 1965 — Terg. II-VI with two pairs of anterior setae A I and 5; terg. VII with A 4 (as in Condé, 1958) and 5; foretarsus 50-55 wm (Réunion, Nepal and Thailand) regale (Condé, 1958) EOSENTOMIDAE There are 315 specimens from more than 35 different localities in the present collection. Six of these are indeterminable, owing to their condition; the 309 remaining specimens have been placed in eight species as follows: Eosentomon oceaniae sp. n., E. melanesiense sp. n., E. solare sp. n., E. sakura Imadaté & Yosii, E. solomonense sp. n., E. guadalcanalense sp. n.,-E. notiale sp. n., E. noonadanae sp. n. The first species, oceaniae, belongs to the ‘swani-group’ (Tuxen, 1964 : 132) which has been recorded from East Asia, Australia and South America, and the next five to the ‘kumei-group’ (loc. cit.: 139), found in both the tropical and temperate areas of East Asia (Imadaté, 1965). The seventh species, notiale, is regarded as a member of the ‘saharense-group’ on account of the long empodium of the hind leg (Tuxen 1964 : 145), although as in E. udagawai Imadaté it may be more natural to arrange it into the ‘kwmei-group’ on the structure of the female squama genitalis. The last species, noonadanae, is peculiar among all known forms of eosentomids, not only in the structure of the female squama genitalis, but also in the chaetotaxy. Eosentomon oceaniae sp. n. (Text-figs 36-40) [Eosentomon tankoktongi Imadaté sensu Tuxen, 1967: 6. Misidentification.] Body length usually 1000-1200 wm in expanded adults. Mouthparts normal; labral setae present; clypeal apodeme distinct; pseudoculus rather small, PR = 12-13. Foretarsus usually 94-104 wm in length, TR = 5:2-5-4 and EU = 1:0, dorsal sensilla ¢-1 placed at about halfway between « 3 and a@ 3’, BS = 1-2-1°3; t-2 thin; apex of ¢-3 almost reach- ing base of « 7; exterior sensilla a relatively long, apex reaching level of a 3; 6 and ¢ normal; d situated a little proximal to y; spatulate e and g present; f—1 seta-like and f-2 short; interior sensilla a’ stout; b’-1 placed about level with « 3’; b’-2 normal and c’ slightly thickened. Hind tarsal empodium short and less than one-fifth the claw length. Thoracic chaetotaxy normal. On abdominal terg. IV-VI, A 1, 2 and 3 absent; A 4 often missing on terg. VI or V and VI; on terg. VII A 1, 2, 3 and 4 lacking. Tergal accessory setae P 1a on terg. VI-VII short, not filiform, less than one-fifth of P 1 and situated in the same row as P1and2. P2and 1” on terg. VIII not displaced. Female squama genitalis resembles that of E. swani Womersley and its allies; caput processus bent against the median edge of stylus in a sharp angle, lateral sclerotization present. Maturus junior. Foretarsus 80-88 um. Important characters such as foretarsal sensillae, chaetotaxy, etc. as in the adult with the exception of the absence of setae 2 and 4 on abdominal stern. XI. Larva II. Foretarsus 68-74 wm. Clypeal apodeme indistinct, position and shape of the foretarsal sensillae are not different from those of the adult. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 351 Larva I. Foretarsus 72 wm in the specimen from Betimatu, Guadalcanal Island. Clypeal apodeme indistinct; position and shape of the foretarsal sensillae similar to those of larva II and adult. Chaetotaxy as in Table 5. 39 40 Fics 36-40. Eosentomon oceaniae. 36, female squama genitalis; 37, mouthparts, dorsal view (ls = labral setae); 38, tergal chaetotaxy of abd. III-VIII; 39, 40, foretarsus, (39) interior and (40) exterior views. 352 Ss. L. TUXEN & G. IMADATE Holotype 9, SoLomMon IsLANDs: Guadalcanal, Mt Austen, 11.ii.1966 (P. J. M. & P. Greenslade) (BMNH). Paratypes. BISMARCK ARCHIPELAGO: Mussau, Boliu, 4.vi.1962, 1 9; Lavongai, Banatam, 20.ili.1962, I m.j.; 26.11.1962, 1 g; Dyaul, Sumuna, 7.iii.1962, 1 g, 3 9; D., Kollepine, 12.ii1.1962, 1 9; New Ireland, Lemkamin, 23.iv.1962, r gf, 1 9, 31 II; Duke of York, Manuan, 21.vii.1962, 1 g, 29, 11 II; New Britain, Valoka, 13.vii.1962, Ig, 109,3 mj., 2111, 111 (all Noona Dan Expedition) (all UZM). SoLomon IsLANDs: Choiseul, Katurasele, 23.x11.1965 (P. J. M. & P. Greenslade), 1 3; Kolom- bangara, north of Kuzi, 6.ix.1965 (P. N. Lawrence), r 3; Santa Isabel, Buala, 14.xii.1964, 4 5, 1 1 II; 12.11.1965, 1 J (all P. J. M. & P. Greenslade); Russell, Yan- dina, 15.viii.1966, 1 g; R., not localized, 18.iii.1965 (P. J. M. & P. Greenslade), I m.j.; Nggela, Vatilau, 14.vi.1965 (P. J. M. & P. Greenslade), 1 3, 1 9, I mj.; Savo, One Naika, 1.vi.1965 (P. J. M. & P. Greenslade), 2 3, 49, 11 II; Guadalcanal, MtAusten; 13:x1.1964, 1°, 2-2; 24.x.1904, 1°9; 21-11.19066, 9.6,.5.9; 2:1m.J,, 2111; 10.vill. 1966, 19,5 9,2m.j. (all P. J. M. & P. Greenslade); G., Mt Popamanase, 2100 m, 26.vi.1965 (P. J. M. & P. Greenslade), 1 m.j.; G., Betimatu, 30.iv.1965 (P. J. M. & P. Greenslade), 11 1 (all BMNH); G., Honiara, 4.viii.1g62 (Noona Dan Expedition), I 2 (UZM); G., Lunga, viii. 1966 (P. J. M. & P. Greenslade), 4 3; Malaita, Dala, 16.11.1965 (P. J. M. & P. Greenslade), 1 g; Bellona, 28.v.1965 (P. J. M. & P. Greenslade), I 3, I m.j.; Rennell, Kagaba Bay, 6.iv.1965 (P. J. M. & P. Greenslade), I 2 (all BMNH); R., Niupani, 18.viii.1962, 3 9, 2 9; 19.viii.1962, 1 3 (all Noona Dan Expedition) (all UZM). FURTHER DISTRIBUTION. Australia (Tuxen, 1967, as tankoktongi Imadateé). It is likely that this form is the commonest eosentomid in Melanesia. £. oceaniae is similar to E. tankoktongt Imadaté from Borneo. The important characters such as the position and shape of the foretarsal sensillae and of the tergal accessory setae P ta on abdominal terg. VI-VII are almost the same in both, but the tergal setae A I, 2, and 3 on abdominal terg. IV are always absent in the Melanesian specimens examined as well as in the Australian specimen examined by Tuxen (1967). The above difference in chaetotaxy seems to be significant and it is reason- able to regard the Melanesian and Australian form as different from E. tankoktongt. Eosentomon melanesiense sp. n. (Text-figs 41-46) Body length is usually 700-900 wm in expanded adults. Mouthparts normal; labral setae present; clypeal apodeme distinct; pseudoculus rather small, PR = 11-13. Foretarsus 73-77 wm in length, TR = 4:7~-5-1 and EU = 1-0; position and shape of fore- tarsal sensillae similar to those of E. oceaniae. Dorsal sensilla ¢-1 situated at about halfway between « 3 and « 3’, BS = 1-2; apex of exterior sensilla a slightly surpassing level of « 3; spatulate e and g normal; interior sensilla b’-1 present; empodium of hind tarsus short and less than one-fifth claw length. Thoracic chaetotaxy normal. On abdominal terg. IV-VI A 1, 2 and 3 absent and A 4 often also missing; on terg. VII A 1, 2, 3 and 4 absent and P ra short, not filiform, less than one-fifth TABLE 5 Chaetotaxy of Eosentomon oceaniae Larva I Larva II Mat. jun. Adult Form. Prim. setae Form. Sec. setae Form. Tert. setae Form. Compl. setae (Dorsal) Tn. 2 I 4 2 4 4 i Gee A2,M 6 A4 6 6 12 P1,1a,2,3,4.5 14 P2a 16 P3a 16 III 4 A2M 6 A 4 6 6 Io P1,1a,2,3,4 12 £P2a 16 P3a,5 16 Abd.I o oO 4 At; 2 4 6? «=P 1,1a,2 8? «=P 3? To P4 To ri oO 2 As5 Io A.1:2;3-4 10 I2 et 1G, 203-4. 5 16 P 2a,4a 16 16 Ill 4 A 4,5 10 AT25 10 12 P 1,14,2,3,4,5 16 P 2a,4a 16 ‘16 sh Ee AG pls zs I2 P4,34,2,3,4,5 16 P 2a,4a 16 16 V=VE 0: 4(2) A 5, (4) 4(2) 4(2) I2 Pt516)2,;3-4:5 36 P 2a,4a 16 16 VIT--o 2 A5 2 2 12 P1,1a,2,3,4,5 16 P2a,4a 16 16 ViiTior 6 As Tas 6 6 6 7 IPIClI2 24 re) eat 9 9 IX 8 8 8 X-XI 8 8 Telson 9 9 9 9 (Ventral) Th.I-II 4-2 A1,2,M 6-2 A3 6-2 6-2 “2. P2 "Oe Piss Os “6 III 4-2 A1m,z,M1 62 A3 6-4 M2 6-4 “2. P2 “6 P13 “8 P2a ee Abd.I 4 A 1,2 4 4 4 “2 Pr 4 P2 ei 4 II-III 2 Atl 4 Az 6 A 6 a. rie 4 4 4 IV-VII 2 Al 4 A 2 6 Ans 6 6 Piss 8 P 2a 10 £28) To VIII o oO oO oO 5 Pe,1,2 7 Pia 7 oi IX 4 152 4 4 X 4 1,2 4 XI 4 1,3 8 2,4 Telson 12 12 12 12 354 S.L. TUXEN & G. IMADATE of P 1 and situated in the same row as P 1 and 2; P 2 and 1” on terg. VIII not displaced; setae I, 2 and 3 on terg. X and setae 1 and 2 on terg. XI absent. Female squama genitalis characterized by the distinctive S-like shape of the combined caput and corpus processus in the processus sternalis. Maturus junior. Foretarsus 63-72 wm. Important characters such as foretarsal sensillae, chaetotaxy, etc. not different from those of adult except for absence of setae 2 and 4 on abdomi- nal stern. XI. Larva II. Foretarsus 51-66 wm; _ position and shape of foretarsal sensillae similar to those 41 42 Fics 41, 42. Eosentomon melanesiense. Foretarsus, (41) interior and (42) exterior views. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 355 of maturus junior. Except for shape and position of P 1a on abdominal terg. VI, chaetotaxy not different from that of same stage of E. oceaniae. Larva I. Foretarsus 45-52 wm. Except for shape and position of P 1a on abdominal terg. VI, chaetotaxy as in same stage of EF. oceaniae. Chaetotaxy as in Table 6. Holotype 3, Sotomon IsL_AnpDs: Guadalcanal, Mt Austen, 11.11.1966 (P. J. M. & P. Greenslade) (BMNH). Paratypes. BISMARCK ARCHIPELAGO: Mussau, Malakata, I1I.vi.1g62 (Noona Dan Expedition) 1 9; Lavongai, Banatam, 20.ii1.1962, 3 3, 5 9, 2 m.j.; 23.11.1962, 2$19,1m.j.; 26.11.1962, 7 J, 109, 11 I (all Noona Dan Expedition); New Britain, Valoka, 13.vii.1962 (Noona Dan Expedition), 11 II (all UZM). SoLomon IsLanps: Wagina, 25.v.1966 (P. J. M. & P. Greenslade), 1 3, 1 9; Kolombangara, north of Kuzi, 6.1x.1965 (P. N. Lawrence), 1 2; Santa Isabel, Buala, 14.xii.1964 (P. J. M. & P. Greenslade), 1 3, 1 9, 11 II; Savo, Koela, 1.vi.1965 (P. J. M. & P. Greenslade), I 9; Guadalcanal, Mt Austen, 24.viii.1965 (P. N. Lawrence), I 2; 24.v.1965, I J, 39, 210.}j,5 111,1900, 13d, 18 9, 3 m.4.5 10.vill.1006,.7 6, 18 9, 21 Lal. JOM. Ss P. Greenslade); G., Mt Jonapau, 1100 m, 6.v.1965 (P. J. M. & P. Greenslade), 1 9; G., Mt Popamanaseu, 1500 m, x.1965 (P. J. M. & P. Greenslade), 1 9; G., Berande Point, 30.iv.1965 (P. J. M. & P. Greenslade), 1 3; G., Lunga, viii. 1966 (P. J. M. & P. Greenslade), 3 9, 1 m.j., 11 I (all BMNH). Fics 43-46. Eosentomon melanesiense. 43, female squama genitalis; 44, 45, chaetotaxy of abd. IX—XII, (44) dorsal and (45) ventral views; 46, tergal chaetotaxy of abd. V—-VII. 356 S. L. TUXEN & G. IMADATE TABLE 6 Chaetotaxy of Eosentomon melanesiense Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) Thek 4 T2 Th.I-Il 6-2 A D3 II-III 6 A 2,4, M 6 P 1,2,3 T6 1,14,2,2@,3,34,4,5 III 6-8 CUD Be i eae a Be Abd.I 4 Aas 8 P 1,2,2a,3 Io | See Om 0 ale Hie ar Abd.I 4 AGT? II-III Io A 1, 2,3;455 4 PAE? 16 P 1,14,2,24,3,4,44,5 II-III 6 A s32;3 IV-VI 2(4) A's, (4) 4 1 eas ipo 16 P 1,14,2,2a,3,4,4a,5 IV-VII 6 A 1,2,3 VII 2 A5 ae ead, 16 P 1,1@,2,24,3;4,;44,5 VIII fo) VIII 6 7 PP CHIn Ta? 9 ix x 4 IX 8 1,2,3,4 XI 8 x a 4 Telson 12 XI 4 3.4 Telson 9 The present new species and the following species, E. solare, are closely similar to E. dawsont Condé from New Zealand and E. wygodzinskyi Bonet from Brazil in important characters such as the peculiar pattern of the chaetotaxy on abdomen X-XI as well as the shape and position of foretarsal sensillae. E. melanesiense is distinguishable by the absence of A 1, 2 and 3 on abdominal terg. IV and by the shape of the female squama genitalis (from E. dawsoni). Significant difference of body size is also denotable between this species and E. dawsonit. Eosentomon solare sp. n. (Text-figs 47-50) Body length usually 700-900 um in expanded adults. Mouthparts normal; clypeal apodeme distinct; labral setae present; pseudoculus of medium size, PR = 10-11. The foretarsus is 70-76 wm in length, TR = 4-8-5:1 and EU = 1-0; position and shape of foretarsal sensillae similar to those in FE. melanesiense; dorsal sensilla t-1 situated about halfway between « 3 and « 3’, BS = 1-1-1-2; apex of exterior sensilla a clearly surpasses level of « 3; spatulate sensillae e and g normal; interior sensilla b’—-1 present. Empodium of hind tarsus short and less than one-fifth claw length. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 357 Thoracic chaetotaxy normal. Abdominal chaetotaxy similar to that of E. wygodzinskyt Bonet and E. dawsoni Condé. On abdominal terg. II-IV five pairs of anterior setae (A 1, 2, 3, 4 and 5) present, but A 1, 2 and 3 on terg. V-VI absent; on terg. VII A 1, 2, 3 and 4 missing and P ta short, not filiform, less than one-fifth of P 1 and situated in the same row as P 1 and 2; P 2 and 1” on terg. VIII not displaced; setae 1, 2, and 3 on terg. X and setae 1, 2 on terg. XI absent. Female squama genitalis similar to that of E. melanesiense, but with latero-proximal sclerotization. Maturus junior. Foretarsus 59-62 wm; except for absence of setae 2 and 4 on abdominal stern. XI, important features such as foretarsal sensillae, chaetotaxy, etc., do not differ from those of adult. Larva II. Foretarsus 52-53 um; position and shape of foretarsal sensillae as in maturus junior. Chaetotaxy as in Table 7. 47 48 Fics 47-50. Eosentomon solare. 47, 48, foretarsus, (47) interior and (48) exterior views; 49, female squama genitalis; 50, clypeal apodeme. 358 S. L. TUXEN & G. IMADATE Holotype 2, SoLomon IsLanps: Choiseul, Katurasele, 23.xii.1965 (P. J. M. & P. Greenslade) (BMNH). Paratypes. BIsMARCK ARCHIPELAGO: Dyaul, Sumuna, I1.iii.1962 (Noona Dan Expedition), 1 3, 1 1 II; D., Kollepine, 12.iii.1962 (Noona Dan Expedition), 2 9, 11 II; Duke of York, Manuan, 21.vii.1g62 (Noona Dan Expedition), 4 3,59, 111] (all UZM). Sotomon IsLanps: Choiseul, Katurasele, 23.xii.1965 (P. J. M. & P. Greenslade), 2 9; Russell, Yandina, 15.viii.1966 (P. J. M. & P. Greenslade), 1 9; Nggela, Soso, 2.xii.1965 (P. J. M. & P. Greenslade), 2 3, 2 9; Guadalcanal, Lunga, viii. 1966 (P. J. M. & P. Greenslade), 1 3, 5 9, 2 m.j.; Bellona, 28.V.1965 (P. J. M. & P. Greenslade), 1 9; Rennell, Kagaba Bay, 6.iv.1965 (P. J. M. & P. Greenslade), 1 2 (all BMNH). TABLE 7 Chaetotaxy of Eosentomon solare Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) Thok 4 12 Th.I-II 6-2 Ar > DE aes a 6 A 2,4,M SOF MPa a3 16 P 1, 1@,2,24,3,34,4;5 Ill 6-4 Az 2 3,201 1,2 Abd. 1 4 Ai,2 Be Rae za Io Dass ep ts ode Abd.I 4 Aye? Tet ve 46 A 1,2,3,4,5 4 Pag 16 PUIG: 2. 20;3.4.40;5 II-III 6 4) 3723 V-VI 4 A 4,5 4 Pex;2 16 P 1,14,2,24,3,4,44,5 IV-VII 6 A 5355 VII 2 A5 10 P%,2,20,20 53 16 P 1,14,2,24,3,4,4a,5 VIII oO VIII a 7 P ¢,1,1a,2 9 ix 4 1,2 IX 8 1,2,3,4 XI 8 Xx 2 4 Telson 12 XI 4 3.4 Telson As mentioned in the description of the preceding species, the present species is very similar to E. melanesiense, E. dawsoni Condé and especially E. wygodzinskyt Bonet. From E. melanesiense it is distinguished by the presence of five pairs of anterior setae on abdominal terg. IV as well as by the presence of a latero-proximal sclerotization in the female squama genitalis. The shape of the caput processus of the female squama genitalis is different from that of E. dawsoni. The difference between E. solare and E. wygodzinskyi, such as the relative length of the filum processus of the female squama genitalis and of the foretarsal sensilla c’, may be significant, although these two forms closely resemble each other. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 359 Eosentomon sakura Imadaté & Yosii (Text-figs 51-55) Eosentomon sakura Imadaté & Yosii, 1959: 7, pl. 2, figs 8-10. Holotype 9, JAPAN: Nara, Mt Yoshino, 30.vii.1954 (G. Imadaté) (KU). Eosentomon sakura Imadaté & Yosii; Imadaté, 1974 : 261-268. 55 51 52 Fics 51-55. Eosentomon sakura. 51, 52, foretarsus, (51) interior and (52) exterior views; 53, 54, chaetotaxy of abd. IX—XII, (53) dorsal and (54) ventral views; 55, female squama genitalis. 360 S. L. TUXEN & G. IMADATE SPECIMENS EXAMINED. BISMARCK ARCHIPELAGO: New Britain, Valoka, 13.vii.1962 (Noona Dan Expedi- tion), 19,1111, 111(UZM). SoLomon IsLanps: Nggela, Vatilau, 14.vi.1965 (P. J. M. & P. Greenslade), 1 3; Savo, Koela, 1.vi.1965 (P. J. M. & P. Greenslade), 1 m.j.; Guadalcanal, Mt Austen, 11.11.1966 (P. J. M. & P. Greenslade), 1 3, 1 9, I mj. (all BMNH); G., Honiara, 4.viii.1962 (Noona Dan Expedition), 1 9, I m.j., 1 1 II, 11 I (UZM); San Christoval, Warahitor, I.viii.1966 (P. N. Lawrence) (BMNH). FURTHER DISTRIBUTION. Japan and Formosa. As shown in Table 8 and Text-figs 51-55, the important specific features such as chaetotaxy, position and shape of foretarsal sensillae, structure of the female squama genitalis, etc. of the Melanesian specimens examined agree well with those of E. sakura. The measurements of the Melanesian specimens are as follows: foretarsus 83-87 pm (adult), 74-76 wm (maturus junior), 63-65 pum (larva II) and 54-57 pm (larva I), PR, TR, EU and BS in adults are 11-12, 5-0, 1-0 and I-I-I+2 respectively. E. collarum Yin from China and E. imbutum Imadaté from tropical Asia closely resemble E. sakura. Whether they are both really distinct from it will only be solved when a considerable number of specimens from different places in Asia are subjected to minute examination. TABLE 8 Chaetotaxy of Eosentomon sakura from Melanesia Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) AM ols 4 142 Th.I-II 6-2 A t.2;57 Me 11-111 6 A 2,4,M *65 P 1,2,3 16 P 1,14,2,24,3,34,4,5 III 6-4 A 1,2,3, M 1,2 Abd.I 4 A 1,2 “8 P 1,2,2a,3 ‘10 P 1,14,2,3,4 Abd.I 4 A 1,2 II-IV Io A 1,2,3,4,5 £ PST 16 P-L, 14;2;24;3;4;44;5 II-III 6 A 1,2,3 V-VI 4 A 4,5 4 Pe 16 P 1,14,2,24,3,4,44,5 IV-VII 6 A 1,2,3 VII 2 A5 To PI 48.245 16 P 1,14,2,24,3,4,4@,5 VIIl oO VIII 6 7 P ¢,1,1a,2 “9 IX-X 4 152 IX 4 1523354 XI 8 x 4 tf Telson 12 XI 4 3.4 Telson 9 PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 361 Eosentomon solomonense sp. n. (Text-figs 56-60) Body length Iooo—-1100 wm in expanded adults. Mouthparts normal; clypeal apodeme distinct; labral setae present; pseudoculus rather large, PR = 8-9. Foretarsus 97-98 wm in length, TR = 5-4-5:8 and EU = 1-0; position and shape of fore- tarsal sensillae similar to those of preceding species. Dorsal sensilla ¢-1 situated about halfway between « 3 and a 3’, BS = 1:1; apex of exterior sensilla a surpasses level of « 3; interior sensilla b’-1 present. Empodium of hind tarsus short and less than one-fifth claw length. Chaetotaxy (Table 9) similar to that of E. gimangi Imadaté from Borneo except for absence of A 3 on abdominal terg. IV; on terg. V-VI A 1, 2 and 3 absent; on terg. VII P 1a situated at posterior margin with the other accessory setae and A 1, 2, 3 and 4 absent; P 2 and 1” not displaced anteriorly on terg. VIII; on terg. X—XI four pairs of tergal setae, 1, 2, 3 and 4 present. Female squama genitalis similar to that of E. melanesiense and sakura. 56 57 Fics 56, 57. Eosentomon solomonense. Foretarsus, (56) interior and (57) exterior views. 362 S. L. TUXEN & G. IMADATE Holotype 3, SOLOMON IsLANDs: Guadalcanal, Mt Austen, 13.xi.1964 (P. J. M. & P. Greenslade) (BMNH). Paratypes. SoLomMon IsLaAnps: Mono, Falomai, 20.ix.1965 (P. J. M. & P. Greenslade), 1 3, 1 1 I; Wagina, 25.v.1966 (P. J. M. & P. Greenslade), 1 9; Nusatupe, Gizo, 4.vi.1965 (P. J. M. & P. Greenslade), 1 2; Nggela, Soso, 2.xii.1965 (P. J. M. & P. Greenslade), ? 1 1 1; Guadalcanal, Mt Austen, 13.xi.1964, 1 4, II.ii.1966, 2 9 (all P. J. M. & P. Greenslade) (all BMNH). This species is closely related to the preceding three species in many respects, e.g., the position and shape of the foretarsal sensillae, the structure of the female squama genitalis, etc., but it is different in the presence of four pairs of tergal setae on abdomen X—XI, in the absence of A 3 on terg. IV, in the position of P 1a on terg. VII and in the relative size of the pseudoculus. On the other hand, this species is also similar to E. gimangi Imadaté from Borneo, but it is distinguished A4 ae IV ag: #2 po / \ 2 et la Vi ee Ag eR, Vv re Ce ee coca Vi ve As -————__\ Vil | ' 58 60 Fics 58-60. Eosentomon solomonense. 58, 59, tergal chaetotaxy of (58) abd. IV—VII and (59) abd. IX—XII; 60, female squama genitalis. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 363 from the latter by the relative length of foretarsal sensilla a and the size of pseudo- culus as well as by the absence of A 3 on terg. IV. One female specimen from Honiara, Guadalcanal, 4.viii.1g62 (Noona Dan Expedi- tion) is apparently a related form of this species, but it is rather small in size (fore- tarsus 60 mm) and the chaetotaxy is consistent with that of E. udorni Imadaté from Thailand except for the presence of A 2 on abdominal terg. V—VI. TABLE 9 Chaetotaxy of Eosentomon solomonense Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) hk 4 ros AQoR Sh t 6-2 At, 2.3 I-III 6 A 2,4, M 76: Bij25 16 JP 1 ,30;2520;5,34,455 Ill 6—4 Avt,2,3) 0 i2 Abd.I a A 1,2 ‘Sy + GP 12205 Io 1 ,10,2;354 Abd. I 4 ‘A 1,2 TI-Itt 10 A 13,2,3:455 zl Pie 16 P 1,1@,2,24,3,4,44,5 II-III 6 AYr23 IV 8 A 1,2,4,5 4 Pie 16 P3514 ,2;24;9;4;4455 LVeVil 6 Wer 2.5 J-VI 4 A 4,5 10 Pt. 220,20 73 16 P 1,%4,2,24,3,4,4@,5 VIII fe) VII 2 A 5 7 PAG UTA. 2 16 P 1,14,2,24,3,4,4a,5 IX-X 4 1,2 VIII 6 XI 8 9 Telson 12 IX-XI 8 1,2,3,4 Telson 9 Eosentomon guadalcanalense sp. n. (Text-figs 61-64) Body length go0o—1200 wm in expanded adults. Mouthparts normal, but labral setae not visible; clypeal apodeme indistinct; pseudoculus of medium size, PR = to. Foretarsus 95 mm in length, TR = 5-9 and EU = 0°8; position and shape of foretarsal sensillae similar to those of E. imadatei Tuxen from Australia; dorsal sensilla ¢-1 placed slightly proximally to « 3’, BS = 1-4; #2 is thin; t~3 rather small; exterior sensilla a normal and apex slightly surpassing level of « 3; c a little broadened, f-2 is short and situated distally to f-1; spatulate e and g normal; interior sensilla a’ stout and placed almost level with ¢-1; b’—1 absent; c’ relatively short. Empodium of hind tarsus short and less than one-fifth claw length. Thoracic chaetotaxy normal. On abdominal terg. II-III, P 4a not visible; A 3 on terg. II-VI and A 1 and 3 on terg. VII absent; P 1a on terg. VII situated at posterior margin with 364 S. L. TUXEN & G. IMADATE 61 62 Fics 61-64. Eosentomon guadalcanalense. 61, 62, foretarsus, (61) interior and (62) exterior views; 63, female squama genitalis; 64, chaetotaxy of abdominal terg. VIII. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 365 the other accessory setae; P 2 and 1” on terg. VIII displaced anteriorly as shown in Text-fig. 64; two anterior setae present on stern. VIII. Female squama genitalis similar to that of E. solare and solomonense. Maturus junior. Foretarsus 82 wm, BS = 1:4 and TR = 5:8; the specific characters such as position and shape of foretarsal sensillae, chaetotaxy (except for stern. XI), etc., similar to those of adult. Larva I. Foretarsus 66 wm; position and shape of foretarsal sensillae similar to those of maturus junior. Chaetotaxy as in Table 1o. Holotype 9, SoLomon IsLANDs: Guadalcanal, Lunga, viii. 1966 (P. J. M. & P. Greenslade) (BMNH). Paratypes. Same data as holotype, 5 9, 1 m.j., 111. E. guadalcanalense is similar to E. imadatet Tuxen from Australia, but differs from it in the displacement of P 2 and 1” on abdominal terg. VIII, in the presence of A 1 on terg. VI and in the shape of the corpus processus of female squama geni- talis. TABLE 10 Chaetotaxy of Eosentomon guadalcanalense Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) Thot 4 1,2 Th.I-II 6-2 A 1,2,3, M II-T8T 6 A 2,4, M 6 Paes 16 P 1,14,2,24,3,34,4,5 Lt 6—4 Ad, 2,3; A552 Abd.I 4 A 1,2 8. P12,20,5 10 PP 4,14;2 354 Abd.I 4 ALY2 II-III 8 A 1,2,4,5 4 Pee 14 i 1;10;2,20,3,4,5 II-III 6 A ;2;3 IV-VI 8 A 1,2,4,5 4 Pie 16 P 1,14,2,24,3,4,44@,5 IV-VII 6 Ack.2;3 VII 6 A 2,4,5 10 P.3;2,24,28'3 16 P 1,14,2,2@,3,4,44,5 Vit 2 A VIII 6 7 PCT Ta\2 9 1b ead. 4 IX-XI 8 1.2,54,4 XI 8 Telson 9 Telson 12 Eosentomon notiale sp. n. (Text-figs 65-68) Body length 670-750 wm in expanded adults. Mouthparts normal; labral setae not visible, but clypeal apodeme distinct; pseudoculus of medium size, PR = Io. Foretarsus 57-62 ym in length, TR = 5-7-6-1 and EU = 0:8; position and shape of fore- tarsal sensillae similar to those of E. guadalcanalense ; dorsal sensilla t-1 placed slightly proximally 366 S. L. TUXEN & G. IMADATE to « 3’, BS = 1:3; interior sensilla a’ situated about halfway between « 3 and «3’; b’-1 absent. Empodium of hind tarsus distinctly elongate and more than one-third claw length. Thoracic chaetotaxy normal. Abdominal terg. II-VI with five pairs of anterior setae; A 1 and 3 missing on terg. VII and P 1a is situated at posterior margin with the other accessory setae; P 2 and 1” on terg. VIII displaced anteriorly; two anterior setae present on stern. VIII. Female squama genitalis similar to that of the previous species. Maturus junior. Foretarsus 48-50 wm. Except for absence of setae 2 and 4 on abdominal stern. XI, the specific features such as chaetotaxy, position and shape of foretarsal sensillae similar to those of adult. Chaetotaxy as in Table 11. Holotype 9, Sotomon IsLanps: Santa Isabel, Buala, 14.xii.1964 (P. J. M. & P. Greenslade) (BMNH). Paratypes. SoLomon IsLanps: Santa Isabel, Buala, 14.xii.1964 (P. J. M. & P. Greenslade), 1 9; Guadalcanal, Mt Austen, I1.ii.1966, I g, IQ, 10.viii.1966, 2 9 (all P.J.M. & P. Greenslade); G., Kukum, vi.—vii. 1966 (P. J. M. & P. Greenslade), I 2 (all BMNH). 65 68 66 Fics 65-68. Eosentomon notiale. 65, 66, foretarsus, (65) interior and (66) exterior views; 67, distal part of hindtarsus; 68, female squama genitalis. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 367 This species is similar to the preceding species, E. guadalcanalense, in some impor- tant features such as the position of the foretarsal sensilla /-1 and the displacement of P 2 and 1” on abdominal terg. VIII as well as in the presence of anterior setae on stern. VIII, but it differs in the long empodium of the hind tarsus and in the position of the foretarsal sensilla a’ as well as in body size. TABLE II Chaetotaxy of Eosentomon notiale Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) Tht 4 12) Th.I-II 6-2 AS 2,95 ME II-III 6 A 2,4,M Ginter 16 P 1,1a,2,24,3,34,4,5 III 6-4 A-1,2,3,. 14 1.2 Abd.I 4 Ai,2 1B o P2203 Io Pita234 Abd. I 4 A 1,2 II-VI 10 A 1,2,3,4,5 4 Pilg 16 P 1,14,2,24,3,4,44,5 II-III 6 A i255 VII 6 A 2,4,5 4 P 1,2 16 P 1,1@,2,24,3,4,4@,5 IV-VII 6 A 1,2,3 VIII 6 if) 2, 20,204 9 VIII 2 A FX-xXt 8 1,2,3.4 7 P ¢,1,1a,2 Telson 9 IX-X 4 1,2 XI 8 Telson 12 The combination of unusual characters such as the position of ¢-1, the displace- ment of P2 and 1” on terg. VIII and the long empodium of hind tarsus, etc., are also found in FE. saharense Condé and E. simile Condé from Africa, although both African forms have b’—1 on the foretarsus. One female specimen (foretarsus 50 um) and a larva II from New Britain, Valoka, 13.vii.1962 (Noona Dan Expedition) have similar chaetotaxy to this species, but the displacement of P 2 and 1” on terg. VIII and the long empodium of the hind tarsus are not present. Eosentomon noonadanae sp. n. (Text-figs 69-75) Body length 800 wm in the holotype. Mouthparts normal, but not very protruded in the holotype; labral setae present; clypeal apodeme distinct; pseudoculus of medium size, PR = It. Foretarsus 63 wm in length, TR = 5-7 and EU = 1-0; dorsal sensilla ¢-1 close to « 3, BS = 0-9; #2 and #3 normal; exterior sensilla a relatively short; c slightly broadened; spatulate 368 S. L. TUXEN & G. IMADATE sensillae e and g present; interior sensilla a’ stout; b’-1 absent. Hind tarsal empodium short and less than one-fifth claw length. Thoracic chaetotaxy normal. On abdominal terg. II-VI the principal setae, P 1, 3 and 4 on posterior row, placed at posterior margin in line with accessory setae P 1a and 2a, which are a little longer than the principal ones, but P 2 and 5 are situated distinctly anterior to the other posterior setae. A 2 and 3 on terg. III and A 2, 3 and 5 on terg. IV—VI lacking. On terg. VII, P 1, 2, 4 and 5 situated anterior to the other posterior setae, P 1a, 2a, 3 and 4a, and all anterior setae absent. P 2 and 1” on terg. VIII not displaced. On abdominal stern. I-VII, the pair of P 1 missing and one central seta, Pc, is present instead of P 1; on stern. IV-VII, A 3 lacking. Abdominal appendages I-III normal and provided with 5 setae each. Female squama genitalis characterized by shape of caput processus, which is semi-circularly rounded, corpus processus divided by deep incisions and coalescent with lateral sclerotizations of stylus to form a broad structure terminating in the filum processus. Chaetotaxy as in Table 12. 69 70 72 Fics 69-72. Eosentomon noonadanae. 69, 70, foretarsus, (69) interior and (70 exterior views; 71, mouthparts, ventral view; 72, female squama genitalis. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 369 Holotype ?, BisMARCK ARCHIPELAGO: New Britain, Valoka, 13.vii.1962 (Noona Dan Expedition) (UZM). This species differs strikingly from all other eosentomids in the peculiar chaetotaxy of its abdominal segments, viz. the displacement of P 2 on abdominal terg. II-VI, the absence of A 3 on stern. IV—VII and of P 1 on stern. I-VII, the presence of a central seta, P c, on stern. I-VII, etc., and in the shape of the female squama genitalis, which shows some slight similarity to that of E. yosemitense Ewing from California. It seems likely that these characters are significant enough to establish a new genus, but we prefer not to describe it as such, since only one specimen is known. Fics 73-75. Eosentomon noonadanae. 73, dorsolateral view; 74, chaetotaxy of abd, II; 75, chaetotaxy of abd. VII (t = tergum, s = sternum). 370 S. L. TUXEN & G. IMADATE TABLE 12 Chaetotaxy of Eosentomon noonadanae Formula Composition of setae Formula Composition of setae (Dorsal) (Ventral) is 4 12 Th.I-III ? II-III 6 A 2,4, M Abd. I 4 A &;2 16 P 1,14,2,2a,3,34,4,5 3 Bsa ole Abd.I 4 A 1,2 II-III 6 ANE23 To PRIA 2s S04 3 P62 II Io AT 2.2 5455 IV-VII 4 AI? 16 P-5,14,2;28,3)4544,5 9 7'0,2.208 24): 3 Ill 6 A 1,4,5 VIII fe) 16 P 1,14,2,2a,3,4,44,5 7 ‘PCM 1a;2 IV-VI 4 A 1,4 IX-X 4 12 16 P 1,1a,2,24,3,4,4a,5 XI mm Vil fe) Telson 12 16 P 1,14@,2,2@,3,4,4@,5 VIII 6 9 IX-XI 8 Telson 9 GENERAL DISCUSSION EcOLoGy. Very little can be said about the ecology, first of all because of the relatively small numbers of each species. For almost all habitats where its character is mentioned, be it for the Noona Dan or Greenslade collections, only ‘litter and soil’ is stated. According to Greenslade & Greenslade (1968) ‘litter’ refers to the upper, organic, litter and humus layers, and ‘soil’ to mineral material below. This is not very informative as to the whereabouts of the Protura, but we should think that they are found, as elsewhere, particularly in the uppermost 5 cm, the layer with a high organic content being shallow in tropical regions (Greenslade, 1969, and the authors’ personal observations). A little more may be said about the localities. Almost all the Noona Dan samples and most of the Greenslade samples are from lowland forest soil, but a few are from higher altitudes: Mt Austen on Guadalcanal (c. 300m), Lemkamin on New Ireland (g00m), Mt Jonapau (1100m) and Mt Popamanaseu on Guadalcanal (1500m and 2100m). On Mt Austen almost all the known species have been found; it is, in fact, the only locality for Condeellum crucis. But at higher altitudes only two species have been found, viz. Eosentomon oceaniae and melanesiense. These two are the commonest of all Proturan species mentioned in this paper and are found in almost all localities, highland and lowland. So it appears that a PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 371 special mountain fauna can be distinguished simply by only a few of the most eurytopic and widespread species being able to withstand the higher altitudes. It might be considered wise to compare the fauna at different times of the year, though only Eosentomon oceaniae yields enough material of all stages for a tentative evaluation. The result is, as might have been expected, that all stages seem to be found all the year round. Further ecological information must await more thorough and methodical collecting. ZOOGEOGRAPHY. The distribution of the 13 species in the Bismarck Archipelago and Solomon Islands is shown in Table 13 (p. 372). Only two of these species are known from outside the Bismarck-Solomon group: Eosentomon oceaniae, which is known from North Queensland, Australia, and Eosentomon sakura, known from Japan and Formosa and probably also China and Thailand. Both are widespread over the Bismarck-Solomon group, the first named being the commoner. In his papers (1968; 1969) P. J. M. Greenslade proposes an origin of the Solomon fauna in New Guinea, spreading west to east via the Bismarck Archipelago. Further, he suggests that colonization has been from larger to smaller islands and depends on the width of the water gaps. He states possible ways of expansion and concludes (1969: 275) for the Solomon Islands that there is ‘a high incidence of endemics on San Christoval’ and that some ‘taxa are restricted to two or more of the islands extending from Bougainville to Nggela’; also that there is a ‘fauna size’ (number of taxa) correlation with the area of the island. Unfortunately the Proturan fauna of New Guinea is not known, but the material in the present collection (124 specimens from the Bismarck Archipelago and 326 from the Solomon Islands) might be thought sufficiently large and widespread (37 localities on 21 islands) to show or at least confirm some parts of Greenslade’s propositions. This seems not to be the case. There are four endemic species among the twelve. (1) Condeellum crucis, known from several specimens from only one locality on Guadalcanal. (2) Berberentulus vennellensis, known from three specimens from Rennell. (3) Eosentomon guadal- canalense, known from several specimens from only one locality (another) on Guadalcanal. (4) Eosentomon noonadanae, known from one specimen from New Britain. On the other hand San Christoval has only one species so far, viz. Eosen- tomon sakura, widespread in the area and beyond. Almost all the species (ten of the thirteen) have been found on Guadalcanal. This leaves a special faunal element on Bougainville - Nggela out of the question, and it does not indicate a fauna size-area correlation either. Two species show, however, an interesting distribution. (a) S#lvestridia solomonis which was originally found on Guadalcanal in 1958; ten more specimens have been found during the present investigation from Guadalcanal, and an additional, single specimen on Ndeni of the Santa Cruz group to the far east of the Solomon Islands. It is furthermore the only species and specimen found on these islands. (b) Berber- 372 S. L. TUXEN & G. IMADATE TABLE I3 Distribution of the known Melanesian Protura ” 2 eg & 3 eos 5 =~ “3 2 2 3 $ N SN . > S a 3 5 S & o Island Pee eine ie semen ee eee I-2. Mussau x x x 3. Lavongai x x 4-5. Dyaul x x 6. New Ireland x Js Duke of York x x x 8. New Britain x x x x Q. Mono 10, Choiseul Xx x 11. Wagina x 12. Nusatupa 13; Kolombangara x x 14-15. Santa Isabel x x x 16. Russell x Xx 17-18. Nggela x x x x 19-20. Savo x x < x 21-29. Guadalcanal x x * x x x x 30. Malaita — Xx 31. San Christoval x ine. Bellona x 33-34. Rennell oa < x 35- Ndeni x E. solomonense E. guadalcanalense E. noonadanae E. notiale = PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 373 entulus buchi (several specimens) found on Mussau and (especially) New Britain of the Bismarck Archipelago, but not on the Solomon Islands in spite of the fairly large collection. On the other hand it is closely related to Berberentulus rennellensis from Rennell. Whether this confirms or not Greenslade’s (1968 : 191) supposition of a Louisade affinity of the Rennell fauna we are unable to say since the Proturan fauna of New Guinea, as mentioned above, and of the Louisades is quite unknown, as is that of New Hebrides and New Caledonia. Such a relationship seems not to be upheld by various authors in the volumes of The Natural History of Rennell. Concluding it may be said that there is a distinct faunal connection between the Bismarck Archipelago and the Solomon Islands, both faunas probably originating in East Asia, though links are also shown with the Australian fauna. Guadalcanal harbours the richest fauna and in addition has two endemics. It is probable that in forming these conclusions we have overstretched the amount of evidence available. TAXONOMY. Both as to number of species and number of specimens the family Eosentomidae by far outnumbers the two other families. This seems reasonable since, generally speaking, the Eosentomon species are eurytopic, whilst the other Protura are more exacting in their demands and are stenotopic (Tuxen, 1949). The differences ao | 160°E ee Fic. 76. Map showing the distribution of the collecting sites. The numbers correspond to those cited in the list of localities (p. 334). 374 S. L. TUXEN & G. IMADATE between the majority of species within this genus are also smaller, often merely a matter of chaetotaxy, than in the other families, so a species relationship is in many cases difficult to state. It is, however, interesting to note that some of the species here mentioned are fairly closely related inter se; this holds true, for example, of Eosentomon melanesiense, solare, sakura, and solomonense, forming as it seems a small group of their own. But this is not expressed in their geographical distribution where they do not always occur together nor do they always replace each other. A curious tendency may be seen in the acerentomids, namely a convergence in the evolution of the foretarsal sensillae and the chaetotaxy in the four species of the three genera Silvestridia, Berberentulus and Gracilentulus of the present fauna. The position of the foretarsal sensillae c and d as well as the size of a and b are com- mon to the four species of these three different genera and more or less different from other species of the same genera. Also a distinct reduction of anterior setae on the abdominal terga V—VIII is common to these four species. These common features are clearly due to convergence, but the reason is not obvious. It is likely that such convergence will be seen not only in morphological characters, but also in their biology. It is also significant that no two (or more) acerentomids have been found coexist- ing at the same collecting site in this collection. There seems thus to be a distinct habitat segregation in the same way as it was shown for Japanese Protura (Imadaté, 1973). REFERENCES GREENSLADE, P. J. M. 1968. The distribution of some insects of the Solomon Islands. Proc. Linn. Soc. Lond. 179 : 189-196. 1969. Land fauna. Insect distribution patterns in the Solomon Islands. Phil. Tvans. R. Soc. (B)255 : 271-284. 1969. Part II. Ecology [of the Collembola of the Solomon Islands]. Phil. Trans. R. Soc. (B)255 : 313-320. GREENSLADE, P. J. M. & GREENSLADE, P. 1968. Soil and litter fauna densities in the Solomon Islands. Pedobiologia 7 : 362-370. ImapaTE, G. 1960. A new species of Protura from the Solomon Islands: Acerentulus solomonis sp. nov. Bull. Osaka Mus. nat. Hist. 12 : 5-8. 1965. Proturans-fauna of south-east Asia. Nature Life S.E. Asia 4 : 195-302. 1973. Habitat segregation between proturan species. Z. Zool. Syst. EvolForsch. 11 : 287-303. 1974. Protura. Fauna Japonica. 351 pp. Tokyo. ImapaTf£, G. & Yost, R. 1959. A synopsis of Japanese species of Protura. Contr. biol. Lab. Kyoto Univ. 6 : 1-43. PETERSEN, B. 1966. The Noona Dan Expedition, 1961-62. Insects and other land arthro- pods. Ent. Meddr 34 : 283-304. Tuxen, S. L. 1949. Uber den Lebenszyklus und die postembryonale Entwicklung zweier danischer Proturengattungen. K. dansk. Vidensk. Selsk. Skr. 6 (3) : 1-49. 1964. The Protuva. A Revision of the species of the world. With keys for determination. 360 pp. Paris. 1967. Australian Protura, their phylogeny and zoogeography. Z. Zool. Syst. EvolForsch. 5 : I-53. PROTURA OF BISMARCK ARCHIPELAGO & SOLOMON ISLANDS 375 TuxeEN, S. L. & ImapaTE, G. 1974. The Silvestridia complex within Protura. Entomologica Scand. 5 : 81-94. Wo rr, T. 1968. The Noona Dan Expedition (Rennell Section, 1962) and the Danish Rennell Expedition, 1965. Nat. Hist. Rennell Isl., 5 : 9-37. INDEX Principal page references are in bold; pages on which there are figures are in italics. Berberentulus, 336, 337, 374 buchi, 336, 340, 373 collarum, 360 Condeellum, 335, 346, 349 crucis, 335, 346, 347-348, 350, 370, 371 dawsoni, 356, 358 _Eosentomon, 336, 350, 373 floridanus, 344 gimangi, 362 Gracilentulus, 335, 340, 344, 374 gracilis, 346 greensladei, 335, 340, 342-343, 346 guadalcanalense, 336, 357, 363, 364, 367, 371 imadatei, 363, 365 imbutum, 360 ishiianum, 348, 350 japonicus, 344 kenyanus, 344 kumei-group, 350 malaysiensis, 344 matobai, 348, 349 melanesiense, 336, 352, 354-355, 358, 379, 374 meridianus, 346 S, L. LuxEn, Dr-Phil: UNIVERSITETETS ZOOLOGISKE MUSEUM UNIVERSITETSPARKEN 15 2100 K@BENHAVN @ DENMARK Prof. G. ImapatTE, D.Sc. BIOLOGICAL LABORATORY KONODAI COLLEGE Tokyo MEDICAL AND DENTAL UNIVERSITY ICHIKAWA CHIBA, 272 JAPAN noonadanae, 336, 350, 367, 368-369, 371 notiale, 336, 350, 365, 366 oceaniae, 336, 350, 351, 370, 371 ohyamai, 344 regale, 348, 350 rennellensis, 336, 337-338, 338, 371, 373 sachikoae, 346 saharense, 367 saharense-group, 350 sakura, 336, 359, 359, 371, 374 sanjianus, 344 Silvestridia, 335, 336, 374 simile, 367 similis, 344 solare, 336, 356, 357, 374 solomonense, 336, 361, 361-362, 374 solomonis, 335, 336, 371 swani-group, 350 tankoktongi, 350, 352 tasmanicus, 344 udagawal, 350 udorni, 363 wygodzinskyi, 358 yodai, 340 yosemitense, 369 ENTOMOLOGY SUPPLEMENTS . Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 18 plates, 270 text-figures. August 1965. £4.20. . SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 1965. £3.25. . OKADA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- philidae. Pp. 129: 328 text-figures. May, 1966. £3. . GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. £3.15. . FLETCHER, D. S. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 text-figures, g maps. February, 1967. £3.50. HeEmMInG, A. F. The Generic Names of the Butterflies and their type-species (Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972. . STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- palocera). Pp. 322: 348 text-figures. August, 1967. £8. . Mounn, L.A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172: 82 text-figures. May, 1968. £4. . Watson, A. The Taxonomy of the Drepaninae represented in China, with an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. November, 1968. £5. . AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- figures. December, 1968. 5. . CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. . Exiot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). Pp. 155: 3 plates, ror text-figures. September, 1969. {4. . GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe (Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. £19. . WuattEy, P. E. S. The Thyrididae of Africa and its Islands. Pp. 108: 68 plates, 15 text-figures. October, 1971. £12. . SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). Pp. 244: g plates, 661 text-figures. July, 1972. £9.90. . CrosskEY, R. W. A Revisionary Classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. February, 1973. £6.50. . VON HayexK, C. M. F. A Reclassification of the Subfamily Agrypninae (Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. . CROSSKEY, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Pp. 221: 95 text-figures. December, 1973. £9.55. 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