BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY VOL. VII 19581959 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1959 DATES OF PUBLICATION OF THE PARTS No. i. 22 July . .... 1958 No. 2. 26 August I958 No. 3. 26 August . . . . . . I95 g No. 4. 12 September ...... 19 cg No. 5. 24 October IQ58 No. 6. 31 October No. 7. 14 October No. 8. 9 January No. 9. 30 January IQ5g PRINTED IN GREAT BRITAIN AT THE BARTHOLOMEW PRESS DORKING BY ADLARD AND SON, LTD. CONTENTS ENTOMOLOGY VOLUME VII No. i. The Mealy-Bugs (Pseudococcidae : Homoptera) described by W. J. Hall, F. Laing and A. H. Strickland from the Ethiopian Region. By D. J. WILLIAMS i No. 2. The African species of Stivalius, a genus of Siphonaptera. By F. G. A. M. SMIT 39 No. 3. The Pseudococcidae (Horn. : Coccoidea) described by C. K. Brain from South Africa. By G. DE LOTTO 77 No. 4. Revisions of Mallophaga Genera. Degeeriella from the falconiformes. By THERESA CLAY 121 No. 5. Revision du genre Exocentrus Mulsant (Col., Cerambycidae) . By S. BREUNING 209 No. 6. New and little known Emesinae (Reduviidae, Hemiptera) in the British Museum (Natural History), London. By P. WYGODZINSKY 329 No. 7. New species and subspecies of Odonata and some Trichoptera from S. Rhodesia and Portuguese East Africa. By D. E. KIMMINS 347 No. 8. New or little known Butterflies from Malaya. By Lr.-CoL. J. N. ELIOT 369 No. 9. A study of the New Zealand Chironomidae (Diptera, Nematocera). By PAUL FREEMAN 393 Index to Volume VII 439 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) DESCRIBED BY W. J. HALL, F. LAING AND A. H. STRICKLAND FROM THE ETHIOPIAN REGION D. J. WILLIAMS BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. i LONDON: 1958 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) DESCRIBED BY W. J. HALL, F. LAING AND A. H. STRICKLAND FROM THE ETHIOPIAN REGION BY D. J. WILLIAMS -" Commonwealth Institute of Entomol Pp. 1-37 ; J 5 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. i LONDON : 1958 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 7, No. i of the Entomological series. Trustees of the British Museum 1958 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued July, 1958 Price Ten Shillings THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) DESCRIBED BY W. J. HALL, F. LAING AND I A. H. STRICKLAND FROM THE ETHIOPIAN REGION By D. J. WILLIAMS Commonwealth Institute of Entomology THIS paper is one of a series to describe and illustrate, where necessary, all the mealy-bugs of the Ethiopian Region. The work is being undertaken jointly by the writer and by Mr. G. De Lotto of the Department of Agriculture, Nairobi, Kenya. One paper has already appeared by De Lotto (1957) dealing with the species described by James from East Africa and another has been completed (Williams, 1958) on the species described by Maskell, Newstead, Cockerell and Green from the Ethiopian Region. Altogether twenty-three species are herein discussed, of which eighteen are regarded as valid and of these, illustrations are given of fifteen. Excellent illustrations of the remaining three species have been given already by other authors and these will be mentioned in the appropriate places. The redescription in each case is based on the holotype and type material held in the British Museum (Natural History). Following the system in my earlier paper, no attempt will be made, in this instance, to erect new genera as the main purpose is to redescribe the species. A species will be placed in a different known genus, however, should this be thought necessary. The terms used are those in current use and are drawn mainly from Ferris (1950), Ezzatt & McConnell (1956) and Borkhsenius (1949). Although many of the species discussed are known from the original discovery only, a few have been collected in other localities in recent years. No attempt is made here to list all localities and hosts as it is hoped to incorporate this aspect in a final revision of the Pseudococcidae. I am indebted to Mr. G. De Lotto for kindly comparing material of Antonina indica panica Hall with the related species described by Brain. THE SPECIES DESCRIBED BY W. J. HALL Hall described five mealy-bugs from Southern Rhodesia and two from South Africa. One of these, Trionymus pterocauloni , described from Southern Rhodesia, is here regarded as being identical with T. sanguineus James. Hall (1937) recorded Trionymus masrensis Hall from Southern Rhodesia but this material is not the same ENTOM. 7. I. I 4 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) as the type from Egypt. An interesting species described by Hall (1941) as Molluscococcus fibrillae from Southern Rhodesia and listed as a Pseudococcine form is here considered to belong to the family Dactylopiidae, recently denned by Ferris (19550) . Two species described from Egypt, Antonina indica panica and Phenacoccus inermis are discussed because the latter is here recorded from the Republic of the Sudan and the former was recorded by Hall (1937) from Southern Rhodesia. Antonina indica panica Hall Antonina indica var. panica Hall, 1925, Bull. Minist. Agric. Egypt, 64 : 6. Antonina indica var. panica Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 125. Hall described this species from Egypt on Panicum turgidum and later recorded it from Theydon, Southern Rhodesia on the roots of Eragrostis sp. near chalcantha. It seems probable that it is the same as a species described by Brain from South Africa. Mr. De Lotto, who is reviewing the species described by Brain will deal with this question in a future paper. Mirococcus inermis (Hall) (Text-fig, i) Phenacoccus inermis Hall, 1925, Bull. Minist. Agric. Egypt, 64 : 7. Mirococcus inermis (Hall), Borkhsenius, 1947, Proc. Acad. Sci. Armen. S.S.R. 7 : 142. HABIT. Originally described by Hall from material collected near Helwan, Egypt, on the roots of Cleome arabica, Cressa cretica, Frankenia pulvemlenta and Zygophyllum simplex. It has since been recorded throughout North Africa and Borkhsenius (1949) has recorded the species from Ukraine, Armenia, Azerbaijan, Uzbekistan and Tadjikistan in the U.S.S.R. Material is at hand from the Ethiopian Region collected at Khartoum, Sudan on Portulaca sp. Hall gives the following description of the external appearance : " Naples-yellow in colour, covered some- what sparsely but uniformly with white pulverulent secretionary matter. Marginal filaments wanting. Skin delicate." RECOGNITION CHARACTERS. The shape of the adult female varies considerably according to the age of the individual. In the young adult the shape is elongate-oval but later becomes more rounded and some specimens become globose ; older speci- mens attaining a length of 3-5 mm. Posterior end of body rounded, anal lobes obsolete. Antennae short, 9-segmented, the terminal segment rounded. Legs short and slender, with a denticle on the plantar surface of the claw. Circulus rather large. Ostioles poorly developed with three or four trilocular pores and an occasional seta on each lip. Anal ring with six setae which are slightly shorter than the diameter of the ring. The outer ring is composed of small pores giving the whole ring a narrow appearance. Cerarii absent. Dorsal setae all short and slender, not numerous. Multilocular disc pores distributed over dorsum, scattered on head and thorax but they occupy transverse rows on the abdomen. Tubular ducts small, of the oral collar type, present on the abdomen only where they are sparse and are arranged more or less in transverse rows, there being scarcely more than ten on any THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) *>'" ' >*> //.'/'/ ' .' *> -'.<'*'.''"' ':'" * o FIG. i. 6 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) one segment and usually less on the posterior segments. Trilocular pores few, evenly distributed. Ventral surface with a pair of long, stout, apical setae. Ventral setae having a similar distribution to those on the dorsum but they tend to be longer, especially on the posterior segments. Multilocular disc pores scattered on the head and thorax but there is a noticeable group posterior to each spiracle. On the abdomen they lie in transverse rows and lateral groups and are more numerous than on the dorsum. Tubular ducts sparse on the thorax where they are present mainly on the margins and between the first legs. They form transverse rows on the abdominal segments and become more numerous posteriorly. Trilocular pores evenly but sparsely distributed. NOTES. This species has been made the type of the genus Mirococcus. The total absence of cerarii, the g-segmented antennae and the denticle on the claw serve to distinguish the species. Octococcus pentziae Hall (Text-fig. 2) Octococcus pentziae Hall, 1939, /. ent. Soc, S. Afr. 2 : 93. HABIT. Described from Grootfontein School of Agriculture, Middelburg, Cape, South Africa on Pentzia sp. (Compositae) . Hall gives the following description of the insect in life : " Adult female enclosed in a closely felted sac which is white or more often dirty white on account of extraneous matter which has become incor- porated. The sac is broadly ovoid and convex, almost globular, with a small orifice towards one extremity." RECOGNITION CHARACTERS. A small oval species rarely exceeding 1-5 mm. in length. Antennae g-segmented. Legs long and slender with a few translucent pores on the hind coxae. Small conical setae are situated on the coxae, trochanters and tibiae of the second and third pairs of legs. Claw with a minute denticle. Circulus absent. Ostioles poorly developed, there being a posterior pair only, each of which is in the form of a narrow slit with sclerotized lips. Anal ring with six setae which are about one and a half times as long as the diameter of the ring. The anal ring is often located at a short distance from the apex of the body and as the cisanal setae are of similar size and shape to the anal ring setae and lie very close to the posterior end of the ring, the impression is given of a ring with eight setae. Cerarii confined to the two posterior segments. Anal lobe cerarii each composed of a pair of short conical setae accompanied by about four long stout blunt setae and two or three trilocular pores surrounded by an elongate sclerotized area. Penultimate cerarii each with two conical setae and two long auxiliary setae and with one or two trilocular pores, surrounded by a small oval area of sclerotization. Dorsal setae not numerous, of various sizes but all rather stout and blunt. The longer setae tend to be more numerous on the posterior abdominal segments. Dorsal multilocular disc pores absent. Tubular ducts of two sizes, the largest with an oral rim and with the duct of a large diameter. There is also another rim encircling the middle of the tube. These ducts are not numerous and occupy THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) FIG. 2. 8 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) single transverse rows. The smaller type of duct is rather slender but has a wide flat oral rim which is not heavily sclerotized. They lie in transverse rows on the abdomen but become scattered on the thorax and head. Trilocular pores sparse. Ventral surface with a curved area of sclerotization on the anal lobes which is continuous with the dorsal sclerotization. Apical setae slightly longer than anal ring setae. Ventral setae of various sizes but more slender than those on the dorsum, not numerous. Multilocular disc pores present mainly on the abdomen where they are situated hi single transverse rows on the anterior and posterior edges of the segments. A few pores are located on the thorax. Tubular ducts similar to the small type on the dorsal surface, are present in no definite arrangement but they tend to occupy transverse rows on the abdomen. A few are present on the thorax especially around the margins. Trilocular pores not numerous. NOTES. This species was made the type of the genus Octococcus Hall on the basis of the anal ring with eight setae. An examination of a number of specimens has shown that the two posterior setae are detached from the ring and are the cisanal setae which often lie on the dorsal surface when the ring is located even a short distance from the apex of the body. Hall has stated that the claw is without a denticle but in all the specimens seen there is a small but distinct denticle at the distal end. This is quite a distinctive species and there is no doubt that the genus is valid although another species Puto africanus Brain which Hall assigned to it does not seem to be congeneric. Paracoccus proteae (Hall) (comb, nov.) (Text-fig. 3) Pseudococcus proteae Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 128. HABIT. Described originally from Inyazura, Southern Rhodesia on Protea sp. Hall gave the following account of the species in life : "A small ovate species in which the brownish colour is obscured by a coating of white pulverulent matter. Four short and stout caudal filaments ; these are about 1/3 of the length of the body of the insect. A few successively shorter marginal filaments occur on the abdominal segments, but these are poorly developed in some individuals. Ovisac of indeter- minate shape. Eggs very pale brown almost yellow." RECOGNITION CHARACTERS. Adult female ovate, a rather small species measuring approximately 2-5 mm. X 1-5 mm. Antennae 8-segmented. Legs long and slender for the size of the insect. Dorsal ostioles well developed, the lips with a few setae and trilocular pores and the inner edges moderately sclerotized. Circulus absent. Anal ring with six setae, these longer than the diameter of the ring and longer than the cisanal setae. Dorsal surface with a reduced number of cerarii there being seven to nine pairs present. Each cerarius consists of two setae surrounded by a few trilocular pores and without auxiliary setae, the cerarian setae becoming more slender anteriorly so that the anteriormost resemble the other setae on the dorsum. Dorsal setae not numerous but all short and slender. Multilocular disc pores absent. Tubular ducts present of the oral rim type only, these arranged singly near the THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) '-- co FIG. 3 io THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) margins of each segment except the last. The penultimate segment has usually a group of two or three. A single oral rim duct is usually situated in the mid-region of the fifth to eighth abdominal segments. Trilocular pores not numerous, evenly distributed. Ventral surface with a small lightly sclerotized anal bar with a short bar seta. Apical setae detached from the bar. Ventral setae not numerous, of similar shape and size to those on the dorsum but on the abdomen and between the antenna! bases there are longer setae present. Multilocular disc pores on the abdomen only, arranged in more or less single transverse rows at the posterior edges of the fourth and posterior segments. Ventral tubular ducts of two types. Some of the oral rims ducts similar to those on the dorsum are situated mainly in a submarginal zone on the thorax, there being usually a noticeable group lateral to the first spiracles. Smaller tubular ducts of the oral collar type are distributed on the prevulvar abdominal segments in transverse rows and also in marginal groups on all the abdominal segments. They are very sparse on the thorax and absent on the head. Trilocular pores not numerous, evenly distributed. NOTES. This species seems to be referable to the genus Paracoccus Ezzatt & McConnell and belongs to the group with seven definite pairs of cerarii. It comes close to P. solani Ezzatt & McConnell described from Arizona both species lacking a circulus, but differs in possessing fewer dorsal oral rim ducts on the head and thorax. Pseudococcus barleriae Hall (Text-fig. 4) Pseudococcus barleriae Hall, 1939, /. ent. Soc. S. Afr. 2 : 96. HABIT. Described from Pretoria, South Africa on Barleria macrostegia (Acan- thaceae) , in the curled leaves at the end of twigs. Hall gives the following description of the habit : " Adult female small and rarely exceeding 1-5 mm. in length, oval in shape, pale brown in colour and sparsely coated with white pulverulent matter. No marginal or caudal filaments apparent. Eggs yellow and in some individuals they were observed to emerge joined together like a string of sausages." RECOGNITION CHARACTERS. Body of mounted female oval and measuring approximately 1-5 mm. long. Antennae 7-segmented. Legs normal except the hind coxae which are noticeably large in comparison to the other coxae, the junction of the coxae to the derm being rather indistinct ; each hind coxa and tibia with a num- ber of translucent pores. Circulus absent. Ostioles represented by a poorly developed posterior pair only, with a few trilocular pores on each lip and apparently without setae. Hall has stated in his original description that the anterior pair is also present but this has not been seen in any of the specimens examined. Length of anal ring setae about one and a half times the diameter of the ring. Cerarii confined to the anal lobes although there is often a single cerarian seta on the pen- ultimate segment. Anal lobe cerarius composed of two medium sized setae and a few trilocular pores ; one or two auxiliary setae are also present. Dorsal setae not numerous, all short and slender. Multilocular disc pores arranged in single trans- THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) n o FIG. 4 ENTOM. 7. I. I 12 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) verse rows at the posterior edges of the thoracic and abdominal segments. Dorsal tubular ducts, small and confined to the margins in small groups. Trilocular pores sparse following the pattern of the dorsal setae. Ventral surface with a pair of apical setae, these nearly twice as long as the anal ring setae. Ventral setae rather sparse, short and slender but they tend to be longer than the dorsal setae. Multilocular disc pores scattered on the head and thorax where they are not numerous. On the anterior abdominal segments they are also scattered but posteriorly they lie in transverse rows on the anterior and posterior edges of the segments ; there are about twenty-five pores posterior to the vulva. Tubular ducts, similar to those on the dorsum, are present in transverse rows on the three prevulvar segments and apart from a few scattered ducts they are mainly arranged in submarginal groups on the thorax and abdomen. Trilocular pores sparse but evenly distributed. NOTES. The distinctive features of this species are the 7-segmented antennae, the reduced number of cerarii, the distribution of the multilocular disc pores on both the dorsal and ventral surfaces and the rather large hind coxae. It does not belong to the genus Pseudococcus as now defined but it is retained in this genus for the time being until further study has been made of the African species. Pseudococcus mazoeensis Hall (Text-fig. 5) Pseudococcus mazoeensis Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 127. HABIT. This species was described from Mazoe, Southern Rhodesia, on Acacia sp. (Leguminosae) and Zizyphus jujuba (Rhamnaceae). The habit is given by Hall as follows : " Adult female, globose, usually brown in colour but some indi- viduals show a tinge of pink. It has a somewhat dense covering of white pulverulent matter which in old specimens has often been worn off to some extent. Marginal filaments confined to the abdominal region ; they are short and stout increasing in size towards the caudal extremity but even the caudal pair are short. Adult female viviparous." RECOGNITION CHARACTERS. Adult female broadly oval, the older specimens attaining a length of 3 mm. Antennae 8-segmented. Legs rather short and stout with a few translucent pores on the hind coxae and tibiae. Circulus present, well developed. Ostioles large with the inner edges of the lips sclerotized and each lip with about three to six setae and a few trilocular pores. Anal ring with six setae, these about twice as long as the diameter of the ring. Cerarii confined to the last six abdominal segments, although there are sometimes seven present. Anal lobe cerarii usually composed of three conical setae surrounded by a number of trilocular pores. Penultimate cerarii each with about nine conical setae which vary in size. The cerarii of the seventh abdominal segment are similar to the penultimate but anteriorly they become smaller so that the anteriormost cerarii each have about five setae or less and a small number of trilocular pores. Dorsal surface beset with small slender setae. The only dorsal pores present are trilocular which are somewhat abundant, and also a few small circular disc pores. THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) FIG. 5 i 4 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) Ventral surface with a faintly sclerotized anal lobe bar and a slender bar seta. The apical seta is detached from the bar and is about twice the length of the anal ring setae. There is apparently a pair of cisanal setae present which are nearly as long as the anal ring setae and also a shorter pair of obanal setae. Ventral setae rather numerous, of various sizes, mainly short and slender but they are generally longer than those on the dorsum. Multttocular disc pores on all segments posterior to the circulus. On the fifth and sixth segments they form single transverse rows but on the seventh and eighth segments they lie in double transverse rows and do not extend to the margins. Ventral tubular ducts of the oral collar type situated in transverse rows and lateral groups on the three prevulvar segments. On the fifth segment they form small lateral groups only and a few are present around the anal lobes. Trilocular pores evenly distributed, not so numerous as on dorsal surface. Small circular disc pores scattered. NOTES. This species does not belong to the genus Pseudococcus as now understood and bears characters which link it with the genus Cataenococcus as recently described by Ferris (1955) and especially to C. phoradendri (Cockerell). It differs from all the known species of Cataenococcus, however, in having the anal ring located at the apex of the body instead of being set at about its own length from the apex of the body. Pseudococcus rhodesiensis Hall (Text-fig. 6) Pseudococcus rhodesiensis Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 130. HABIT. This species was described from South Marendellas, Southern Rhodesia on grass roots. Hall gave the following description of the habit : " Adult female ovate to elongate oval and pale to bright yellow in colour. The segmentation is distinct and the dermis is sparsely dusted with a little white pulverulent matter. No marginal or caudal filaments apparent. Later the female becomes enclosed in a cell of white fibres of indeterminate shape, the inside of which is comparatively smooth and matted." RECOGNITION CHARACTERS. Adult female as mounted on the slide, elongate-oval measuring approximately 3-5 mm. long, the posterior end of the body rounded. Antennae very short with either six, seven or eight segments. Legs small in com- parison to the size of the body, claws without a denticle. Circulus absent. Anterior and posterior ostioles absent. Anal ring with six setae, their lengths being nearly twice as long as the diameter of the ring. Outer ring of anal ring pores rather numerous, giving the ring a wide appearance. Spiracles with wide apodemal plates but without a crescentic band of pores on the spiracular opening. Cerarii represented by a single pair on the anal lobes each usually composed of a single short conical seta surrounded by a few long stout auxiliary setae but without trilocular pores. Dorsal setae not numerous of various lengths but all slender and not lanceolate. Apart from the group of long auxiliary setae surrounding each anal lobe cerarius there is another group on the margin of the penultimate segment. A few other long marginal setae are present on some of the other abdominal segments. THE MEALY-BUGS (PSEUDOCOCCIDAE ; HOMOPTERA) 16 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) Dorsal multilocular disc pores in groups of up to twelve ; numerous across the abdominal segments but present on the thorax mainly on the margins. Each group of pores surrounds a small slender tubular duct but there is often more than one duct present probably because the groups are merged. Single pores are often scattered between the groups. On the last three segments there is a variable number of sieve-like disc pores. These are usually larger than the multilocular disc pores and the shape varies from circular to oval. Trilocular pores about the same size as the multilocular disc pores, always round ; usually distributed along the anterior and posterior margins of the abdominal segments. They are sparser on the thorax and head. Small circular disc pores scattered. Ventral surface with a few long setae on the margins of the abdominal segments. Other setae short and slender, not numerous. Multilocular disc pores in similar groups to those on the dorsal surface ; numerous in transverse rows on the abdomen and around the margins. Sieve-like disc pores present on the three posterior segments. Trilocular pores more numerous than on the dorsum there being notice- able concentrations around the spiracular openings. Small circular disc pores present in no definite arrangement. NOTES. The groups of multilocular disc pores each surrounding a slender tubular duct would suggest a relationship with the genus Peliococcus Borkhsenius. Nevertheless this genus belongs to the Phenacoccus series with g-segmented antennae, with a denticle on the claw and with ventral quinquelocular pores. As rhodesiensis has none of these characters and as it possesses sieve-like disc pores on the abdomen and spiracles with rather wide apodemal plates it may be that it has some relationship, however remote, to the grass feeding genera centred around Antonina Signoret and Antoninoides Ferris. It is significant that Antoninoides parrotti (Cockerell) has extremely small antennae and legs similar to those of rhodesiensis. Trionymus inyazurae Hall (Text-fig. 7) Trionymus inyazurae Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 131. HABIT. Originally described from Inyazura, Southern Rhodesia on grass just underground at the base of the aerial shoots. Hall notes the external appearance as follows : " Adult female ovate, pale brown to maroon, but the colour is somewhat obscured by a very fine and uniform film of white pulverulent matter. Segmentation distinct. There are no marginal filaments, and in the absence of these the four very short caudal filaments are readily seen. Ovisac of indeterminate form but composed of fibres that are capable of being drawn out to a considerable length. Eggs pale brown. Young adult females are relatively elongate filling out and becoming more ovate later." RECOGNITION CHARACTERS. Adult female in prepared specimens, ovate and measuring approximately 2-5 mm. long. Posterior edge of body rounded. Antennae 8-segmented. Legs normal, rather slender with a few translucent pores on hind coxae. Circulus absent. Anterior and posterior ostioles moderately developed, THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) FIG. 7 i8 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) the lips with a few trilocular pores but apparently without setae. Anal ring with six setae, the lengths of which are nearly twice the diameter of the ring. Cerarii confined to the anal lobes only, each consisting of two small conical setae surrounded by a small cluster of trilocular pores and three or four short, slender, auxiliary setae surrounded by a lightly sclerotized area. Dorsal setae evenly distributed, not numerous and all short and slender. Multilocular disc pores present on the dorsum, these rather scattered on the head and thorax but on the abdomen they are distri- buted in transverse rows at the anterior and posterior edges of the segments ; they are absent on the last segment. Tubular ducts distributed over entire dorsum, of three sizes all of the oral collar type. A large type, few in number, is distributed mainly on the anterior head region and singly on the margins ; other single ducts are present on the dorsum but these are not constant in number or position. An intermediate size present over entire dorsum, rather numerous and arranged more or less in transverse rows across the segments. Small tubular ducts on the last four segments only, these in transverse rows at the posterior edges of the segments except the last where there is a small marginal group, the latter being the only ducts on the last segment. Trilocular pores not numerous but evenly distributed. Ventral surface with a pair of apical setae which are stouter and longer than the anal ring setae. Ventral setae similar to those on dorsum. Multilocular disc pores rather numerous on the abdomen at the anterior and posterior edges of the segments, becoming less numerous anteriorly on the thorax. They are sparse on the head. Tubular ducts of the same three sizes as those on the dorsum. An occasional duct of the large type is present on the margins and on the anterior head region. The intermediate size duct is the most numerous, these scattered on the thorax, but on the abdomen they lie in transverse rows and lateral groups. Numerous small tubular ducts confined to the last three segments. Trilocular pores sparse. NOTES. This species does not seem to be a typical Trionymus but it bears a close similarity to T. magnus (Cockerell & Cockerell) known only from North West Mexico and recently redescribed by Ferris (1953). The latter species has been placed in Trionymus but it differs from inyazurae mainly in having a small circulus and 7-segmented antennae instead of 8-segmented antennae. It is significant that inyazurae is a grass-feeding species, as are most species of Trionymus. Trionymus pterocauloni Hall = Trionymus sanguineus James (SYN. NOV.) Trionymus pterocauloni Hall, 1937, Trans. R. ent. Soc. Land. 86 : 133. This species was described by Hall from Salisbury, Southern Rhodesia on Pterocaulon decurrens and Trifolium sp. It is identical, however, with Trionymus sanguineus described by James (1936) to which the name Trionymus pterocauloni Hall is here sunk as a synonym. De Lotto (1957) has already redescribed sanguineus and given an illustration. THE SPECIES DESCRIBED BY F. LAING Seven species have been described from the Ethiopian Region by Laing at one time or another. Two names have been sunk as synonyms already and a further THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) 19 name is synonymized herein. Ezzatt & McConnell (1956) have redescribed and illustrated Planococcoides njalensis. In the accompanying pages, illustrations and descriptions are given of Heliococcus phaseoli, Pseudococcus hargreavesi and Pseudococcus ugandae. Heliococcus phaseoli (Laing) (Text-fig. 8) Phenacoccus phaseoli Laing, 1929, Ann. Mag. nat. Hist. (10) 4 : 475. Heliococcus phaseoli (Laing), Goux, 1934, Bull. soc. ent. Fr. 39 : 171. HABIT. This species was described from Hill Station, Sierra Leone on dwarf beans. Laing was unable to give any description of the external appearance as the specimens were preserved in alcohol. RECOGNITION CHARACTERS. Adult female ovate measuring approximately 3-5 mm. long. Anal lobes moderately sclerotized on the dorsal surface. Antennae 9-segmented. Legs stout and long, with a denticle on the claw. Circulus rather large and wide. Ostioles moderately developed, each lip with about two setae and a few trilocular pores. Anal ring with six setae, these being slightly longer than the diameter of the ring. Cerarii numbering thirteen pairs and borne at the apices of small membranous tubercles. There is a cerarius on the margin of each abdominal segment and the remaining cerarii are situated evenly on the margins of the thorax and head. Each cerarius composed of a pair of lanceolate setae surrounded by a few trilocular pores. Dorsal surface with minute lanceolate setae which are quite sparse. Dorsal multilocular disc pores numerous, in definite transverse rows on all abdominal segments except the last, on the three thoracic segments and one row on the head. Crateriform ducts present in transverse rows, these being of three sizes. There are two pairs of large crateriform ducts on the anal lobes, each duct having three or four setae around the base of the duct prominence. Intermediate sized crateriform ducts are situated mainly in groups of two or three around the margins and an occasional duct is present in the mid-region. These ducts have three, or occasionally four, setae around the base of the duct prominence. Small crateriform ducts each with a single seta at the base of the duct prominence are present in single transverse rows on the abdomen whilst on the head and thorax they form irregular rows. A few small tubular ducts of the oral collar type are present in marginal groups on the seventh and eighth segments. Trilocular pores not numerous, evenly distributed. Ventral surface with a pair of long apical setae accompanied by two shorter setae. In some specimens there is a small area of faint sclerotization near each apical seta but this is indistinct. Ventral setae of various sizes, there being some long setae especially in the mid-region interspersed with shorter setae. Minute lanceolate setae similar to those on the dorsum are located around the margins. Multilocular disc pores numerous. On the abdominal segments they occupy dense transverse rows at the anterior and posterior edges of the segments and on. the head and thorax they form irregular rows. Quinquelocular pores sparse, there being a few between the transverse rows of multilocular disc pores on the abdomen and groups between 20 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) 'fewfel^=fefc^^ r ' -- *"< *- t Mk ''\* '. / : . fV .' ; ^ x <^' * /* ' * ' ' --0-Ctf * ' ' V-- * J* .'. .//. > '." ." . -""V: '',- '". *' " o FIG. 8 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) 21 the coxae. Small crateriform ducts similar to the small type on the dorsal surface are distributed around the margins but they are not numerous. There are noticeable groups posterior to each spiracle. Small oral collar tubular ducts in transverse rows between the rows of multilocular disc pores on the abdomen. They are fairly numerous on the posterior segments but become fewer anteriorly ; a few are present between the coxae. Trilocular pores sparse. NOTES. Goux (1934) included this species in the genus Heliococcus Sulc on the basis of the crateriform pores. It seems to belong to this genus although the type species has eighteen pairs of cerarii whilst phaseoli has only thirteen pairs. Borkhsenius (1949), however, has described a number of species with two to eighteen pairs of cerarii and it seems that the number can be variable. Paraputo ritchiei Laing = Paraputo anomala (Newstead) Paraputo ritchiei Laing, 1929, Ann. Mag. nat. Hist. (10) 4 : 473. In an earlier paper of this series (Williams, 1958) it has been established that this species is identical with Paraputo anomala (Newstead) to which the name ritchiei was sunk as a synonym. Planococcoides njalensis (Laing) Pseudococcus njalensis Laing, 1929, Ann. Mag. nat. Hist. (10) 4 : 472. Pseudococcus exitiabilis Laing, 1944, Bull. ent. Res. 35 : 91. Pseudococcus njalensis Laing, Hall, 1945, Bull. ent. Res. 36 : 305. Planococcoides njalensis (Laing), Ezzatt & McConnell, 1956, Univ. Maryland Agric. Exp. Sta. Bull. A 84 : 55. Laing described njalensis in 1929 from Sierra Leone and in 1944 described exitiabilis from Gold Coast. It has been shown by Hall (1945) that the latter name is a synonym of njalensis and that the species is extremely variable. Great interest has been shown in this species in recent years as it is a vector of the " Swollen Shoot " virus disease of cacao. It is probably widespread throughout West Africa and the reader is referred to the paper by Hall who discussed its distribution and host records. Ezzatt & McConnell (1956) have recently made it the type species of the genus Planococcoides and have given an excellent illustration. Pseudococcus bukobensis Laing = Pseudococcus hargreavesi Laing An examination of type material of Pseudococcus bukobensis Laing described in 1929 has shown that it is identical with Pseudococcus hargreavesi Laing described in 1925 and the synonymy is given in the discussion of the latter species. Pseudococcus exitiabilis Laing = Pseudococcus njalensis (Laing) As previously stated this species has been shown by Hall (1945) to be the same as njalensis and it is listed here purely for reference. 22 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) Pseudococcus hargreavesi Laing (Text-fig. 9) Pseudococcus hargreavesi Laing, 1925, Bull. ent. Res. 16 : 52. Pseudococcus bukobensis Laing, 1929, Ann. Mag. nat. Hist. (10) 4 : 471 (SYN. NOV.). HABIT. Described originally from Kampala, Uganda on Bauhinia sp. and again under P. bukobensis from Bukoba, Tanganyika Territory on coffee. In neither case is there any description of the insect in life due to the specimens having been preserved in alcohol. RECOGNITION CHARACTERS. Body of adult female oval, attaining a length of 4 mm. Antennae 8-segmented. Legs long and slender without a denticle on the claw, hind coxae and tibiae with a few translucent pores. Circulus present. Ostioles moderately developed, with the inner edges of the lips slightly sclerotized and with two or three setae and a few trilocular pores on each lip. Anal ring with six setae which are nearly twice as long as the diameter of the ring. Spiracles with a rather short, broad, apodemal plate. Cerarii numbering eighteen pairs. Anal lobe cerarii each with about seven conical setae of various sizes, with a few trilocular pores and one or two auxiliary setae surrounded by a characteristic sclerotized area. Penultimate cerarii similar to anal lobe cerarii each surrounded by a smaller area of sclerotization. The anterior cerarii are each composed of a few conical setae, there being rarely less than four setae and sometimes as many as seven but their numbers vary in different specimens. Dorsal surface with minute lanceolate setae which are not numerous. Trilocular pores accompany these setae in definite areas only there being thus some areas devoid of pores and setae as illustrated. A few tubular ducts of the oral collar type are usually scattered on the thorax. Ventral surface of anal lobes each with a small sclerotized anal bar and a long slender bar seta. The apical seta is detached from the bar and is stouter and longer than the anal ring setae. Ventral setae not lanceolate, mainly long and slender but not numerous. Multilocular disc pores present on all segments posterior to the circulus, situated in the mid-region in transverse rows. On the fifth and sixth segments they form single rows at the posterior edges and on the seventh and eighth segments they form double rows. There are a few pores on the anterior edge of the seventh segment and a more or less double row on the anterior edge of the eighth segment. They are numerous between the anal lobes. Tubular ducts in transverse rows on the fourth to eighth abdominal segments and in marginal groups from the thorax to the anal lobes. Trilocular pores sparse. NOTES, an examination of type material of Pseudococcus bukobensis Laing has shown that it is the same as P. hargreavesi Laing and the former name is here sunk as a synonym. This is a distinctive species which seems to belong to the tribe Planococcini as defined by Ezzatt & McConnell (1956) . The dorsal setae are typically lanceolate resembling those of the Phenacoccus series, nevertheless the antennae are 8-segmented and there is no denticle on the claw. THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) '..' /.*< . .'; : ".*.*. vV ' :r - .' FIG. 9 24 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) Pseudococcus ugandae Laing (Text-fig. 10) Pseudococcus ugandae Laing, 1925, Bull. ent. Res. 16 : 53. HABIT. Described from Kakumiro, Uganda on Grevillea robusta. Laing has given no indication of the external covering, presumably because his type material had been preserved in alcohol. RECOGNITION CHARACTERS. Adult female broadly oval, attaining a length of 3-5 mm. Anal lobes rather small, the dorsal surface sclerotized. Antennae 9- segmented. Legs slender, claw without a denticle. Circulus moderately developed. Ostioles present, the inner edges of the lips slightly sclerotized and the lips with a few trilocular pores and rarely with setae. Anal ring with six setae which are about twice as long as the diameter of the ring. Cerarii on the five posterior segments only. Anal lobe cerarii each with a pair of conical setae surrounded by a few tri- locular pores and two auxiliary setae. Anterior cerarii similar to those on anal lobes, with a few trilocular pores, but the two anteriormost cerarii are each usually composed of one seta which is smaller than the other cerarian setae. Dorsal surface with slender setae of moderate length but not numerous. Dorsal multilocular disc pores absent. Tubular ducts of three sizes. Large tubular ducts present, of the oral rim type, distributed mainly in marginal groups of two or three or even five on the posterior segments, there being also a few on the mid-region of the thorax. These large ducts are often in pairs and are thus easily noticeable. An intermediate size tubular duct with an oral rim is distributed sparsely over the dorsum mainly in irregular transverse rows. Small tubular ducts of the oral collar type are present among the intermediate type but they are not numerous. Trilocular pores sparse, evenly distributed. Ventral surface with a pair of long stout apical setae which are longer than the anal ring setae. There is a wide sclerotized anal bar continuous with the dorsal sclerotization of each anal lobe, bearing a pair of long setae. Other ventral setae of moderate length and similar to those on the dorsum. Multilocular disc pores on all segments posterior to the circulus. On the fifth segment they occupy a double transverse row on the posterior edge. Posteriorly they are numerous in transverse rows at the anterior and posterior edges of the segments. Small tubular ducts each with an oral collar, not numerous, distributed mainly in irregular transverse rows between the multilocular disc pores on the abdomen ; anteriorly they are scattered. Trilocular pores sparse. NOTES. This species does not belong to the genus Pseudococcus as now understood. It seems to have close affinities to Phenacoccus hirsutus Green in possessing 9- segmented antennae, only five pairs of cerarii and with numerous oral rim ducts. In hirsutus there are large oral rim ducts on the ventral surface which are absent in ugandae. Laing has stated in his original description that the antennae are 8- segmented and that there is an obscure division across the eighth segment suggesting a tendency to a g-segmented form. In all the specimens seen the antennae are distinctly g-segmented. THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) FIG. 10 26 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) THE SPECIES DESCRIBED BY A. H. STRICKLAND Strickland collected some important scale insects in the Gold Coast during a period of research into the vectors of the virus causing " Swollen Shoot " disease of cacao. In two papers, Strickland (1947, 19470) described seven new mealy-bugs, mainly from cacao. These are very interesting and give some indication of what might be expected after further intensive collecting in West Africa. Two of these species have been adequately redescribed and illustrated recently by other workers. The remaining five species are redescribed in the following pages. Cataenococcus loranthi (Strickland) Farinococcus loranthi Strickland, 1947, Bull. ent. Res. 38 : 515. Catenococcus loranthi (Strickland), Balachowsky, 1954, ^ ev - Path veg. 33 : 247 (mis-spelling). Strickland described this species in the genus Farinococcus from Tafo, Gold Coast, on Loranthus bangwensis attended by an undetermined ant of the genus Cremato- gaster. Ferris (1955) erected the genus Cataenococcus with Dactylopius olivaceus Cockerell as type, mainly on the basis of the 8-segmented antennae and the anal ring with six to ten setae. Balachowsky (1954) has placed loranthi in the genus Cataenococcus and given an excellent illustration from specimens collected in French Guinea on Rhizophora racemosa. Delococcus tafoensis (Strickland) Formicococcus tafoensis Strickland, 1947, Bull. ent. Res. 38 : 513. Delococcus tafoensis (Strickland), Ferris, 1955, Microentomology, 20 : 5. Originally described from Tafo, Eastern Province, Gold Coast on Theobroma cacao. It has been made the type of the genus Delococcus by Ferris (1955) on the basis of the 6-segmented antennae and the numerous setae on the anal ring. Ferris has also illustrated this species. Planococcus celtis (Strickland) (comb, nov.) (Text-fig, u) Pseudococcus celtis Strickland, 1947, Proc. R. ent. Soc. Lond. (B) 16 : 154. HABIT. Described originally from Tafo, Eastern Province, Gold Coast, on Celtis sp. (Urticaceae). The habit has been described by Strickland as follows : " Oval to sub-circular in shape, completely covered dorsally and ventrally with white wax, thinner along the intersegmental membranes, and ventrally, especially around the beak and coxae. With seventeen or eighteen pairs of stout lateral wax filaments, the anal pair being slightly longer than the abdominal pairs, and these longer than those anteriorly placed. No dorsal median wax-free stripe. Body colour when wax removed, a dull lemon yellow." RECOGNITION CHARACTERS. A broadly oval species measuring approximately 2 mm. long on the slide. Antennae 8-segmented. Legs short and stout, the hind legs with some translucent pores on the coxa and tibia. Circulus present, normal for THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) 27 o FIG. ii 28 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) the genus. Ostioles well developed with the inner edges of the lips sclerotized and with about seven setae and a few trilocular pores on each lip. Anal ring with six setae which are only slightly longer than the diameter of the ring but they are longer than the cisanal setae. Dorsal surface of anal lobes moderately sclerotized. Cerarii numbering eighteen pairs. Each cerarius composed of a pair of stout conical setae which are pointed or flagellate distally, surrounded by a group of trilocular pores and often one or two slender auxiliary setae. The third cerarius usually has one or two extra conical setae which are smaller than the main pair. Dorsal tubular ducts and multilocular disc pores absent. Trilocular pores with an even distribution ; there are often one or two trilocular pores at the bases of some of the setae on the thorax but these are not to be confused with dorsal cerarii. Ventral surface with a pair of long apical setae which are over twice as long as the anal ring setae. Anal lobe bar reaching to the apical seta, with the bar seta as long as an anal ring seta. Ventral setae not numerous but generally longer than those on the dorsum. Multilocular disc pores confined to the abdominal segments posterior to the circulus, in single transverse rows except on the first prevulvar segment where they are in a double row. They are situated mainly in the mid-region of the segments but often reach to the margins. Approximate numbers of pores on each segment as follows : V 8, VI 22, VII 22, VIII 28, IX 20. Tubular ducts of the oral collar type mainly present in submarginal groups on the seventh and eighth segments. There is also an occasional duct in the mid-regions of these segments. Trilocular pores sparse. NOTES. This species seems to be intermediate between two species described by De Lotto from Kenya as P. rotundatus and P. subukiaensis. It is related to the former species by the cerarii possessing auxiliary setae but differs in possessing more multilocular disc pores. The arrangement of the multilocular disc pores is similar to that of subukiaensis but the latter has no auxiliary setae with the cerarii. Rhizoecus spelaea (Strickland) (comb, nov.) (Text-fig. 12) Coccidella spelaea Strickland, 1947, Bull. ent. Res. 38 : 502. HABIT. Originally described from Tafo, Gold Coast on the roots of Theobroma cacao. Strickland gives the following description of the external appearance : " Covered with a thin layer of finely particulate white wax, thinner along the inter- segmental membranes and around the beak and coxae. With one pair of short, stout, wax filaments on the last abdominal segment." RECOGNITION CHARACTERS. An oval species, widest on the thorax and with the abdomen tapering gradually. Length as mounted on the slide approximately 2 mm. Antennae 6-segmented, strongly geniculate, there being four stout, curved, blunt setae on the two apical segments. Legs with long slender claws and with the claw digitules reduced to small slender setae. Circuli three in number being rather large for the genus, each having a reticulated surface. The anterior circulus lying between the hind coxae and the other two circuli on the two posterior segments ; the middle THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) 29 O FIG. 12 30 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) circulus being the largest and the posterior circulus the smallest. Dorsal ostioles poorly developed with sclerotized lips but without setae or trilocular pores. Anal ring wide with irregular oval pores, setae about twice as long as the diameter of the ring. Eyes and cephalic plate apparently absent. Anal lobes with faint sclero- tization and each with two dorsal and one ventral setae, these quite long. Dorsal surface with few setae, all short and slender. Dorsal multilocular disc pores absent. Tubular ducts often situated on the margins of the anterior abdominal segments, these rather small and their shape being somewhat difficult to determine. A few tritubular pores are present on the head margin. Trilocular pores sparse. Ventral surface with small slender setae which are quite sparse. Multilocular disc pores absent. Tubular ducts similar to those on dorsum in transverse rows on the abdomen where they are more numerous on the anterior segments. Tritubular pores situated between the coxae, varying in number but there are usually about six pairs present. Trilocular pores sparse. NOTES. This species was originally described in the genus Coccidella Hambleton but is here placed in the genus Rhizoecus following the redefinition of this genus by Ferris (1953). The species may be readily distinguished from the others in the genus by the three prominent circuli and by the arrangement of the characteristic tubular ducts. Tylococcus boafoensis Strickland (Text-fig. 13) Tylococcus boafoensis Strickland, 1947, Proc. R. ent. Soc. Lond. (B) 16 : 151. HABIT. This species was described from Tafo, Eastern Province, Gold Coast on Musanga smithii (Moraceae). The description of the insect in life has been given as follows : " Elongate oval in shape, covered with white wax, thinner along the intersegmental membranes and around the beak and coxae, without a dorsal wax-free stripe. With eighteen pairs of lateral wax filaments, the anal pair longer than the rest. Body colour, when wax removed, light yellow." RECOGNITION CHARACTERS. Adult female oval measuring approximately 1-5 mm. long. Antennae 8-segmented. Legs long and slender without a denticle on the claw, hind coxae and tibiae with a few translucent pores. Circulus present, moderately developed. Anterior and posterior ostioles with the inner edges of the lips sclerotized and each lip with one or two setae and a few trilocular pores. Anal ring with six setae which are more than twice as long as the diameter of the ring. Cerarii numbering eighteen pairs, each cerarius situated at the apex of a small slightly sclerotized tubercle, except the anal lobe cerarii which are borne at the apices of larger tubercles representing the anal lobes. Each anal lobe cerarius with two stout, conical setae surrounded by a cluster of trilocular pores and usually with two auxiliary setae. The anterior cerarii similar to the anal lobe cerarii often with an auxiliary seta. Dorsal surface with a small number of slender setae many of which are very noticeable by having one to four trilocular pores at their bases and thus resembling dorsal cerarii. These setae are, however, much more slender than the cerarian setae. Dorsal multilocular disc pores and tubular ducts absent. Trilocular pores sparse. THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) 31 o FIG. 13 32 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) Ventral surface with a small sclerotized anal lobe bar with a bar seta shorter than the anal ring setae. The apical seta is detached from the anal lobe bar and is longer than the anal ring setae. There appears to be a pair of cisanal and obanal setae which are roughly of similar shape and size but are shorter than the anal ring setae. Other ventral setae short and slender but they tend to be longer than those on the dorsal surface. Ventral multilocular disc pores very few, confined to the last three segments in single transverse rows, there being but four to six on each of the two prevulvar segments and one to three on the last segment. Ventral tubular ducts very sparse on the seventh and eighth segments only. They are situated lateral to the multilocular disc pores and there are usually four or five on the seventh segment and three or four on the eighth segment. Trilocular pores sparse. NOTES. This species is certainly not congeneric with the type of Tylococcus. It seems to have a connection with the tribe Planococcini as defined by Ezzatt & McConnell (1956) by having eighteen pairs of cerarii and with the sclerotized anal lobe bars but differs from all species so far placed in that tribe by the cerarii being situated at the apices of small sclerotized tubercles. Apart from these characters the species is easily recognizable by the sparse microscopical characters such as setae and pores. Tylococcus malacanthae Strickland (Text-fig. 14) Tylococcus malacanthae Strickland, 1947, Proc. R. ent. Soc. Lond. (B) 16 : 149. HABIT. Described from Tafo, Eastern Province, Gold Coast on Malacantha sp. (Sapotaceae). Strickland gives the following description of the adult female in life : " Anterior segments lightly powdered with a fine, dusty, yellow wax, the posterior segments nude of wax. With eighteen pairs of groups of thin, glossy, yellow wax filaments laterally, each group consisting of two or three individual filaments issuing from a cerarius. Body colour, when wax removed, ochreous yellow." RECOGNITION CHARACTERS. A broadly oval species, the largest measuring approximately 2-5 mm. long x 1-5 mm. wide. Antennae 8-segmented. Legs short and stout. Circulus large, dumb-bell shaped. Ostioles well developed, the lips heavily sclerotized and bearing an occasional seta but apparently without trilocular pores. Anal ring set at a distance of about one and a half times its diameter from the apex of the body ; with six setae which are only slightly longer than the diameter of the ring. The ring is surrounded by a sclerotized band containing a few short setae and trilocular pores belonging to the ninth segment. Cerarii numbering eighteen pairs, each borne at the apex of a sclerotized tubercle of variable size. Anal lobe cerarius consisting of three prominent and slightly lanceolate setae at the apex of a large tubercle representing the anal lobe. The anal lobe tubercle is the largest and bears about three auxiliary setae and one or two trilocular pores. The penultimate and antepenultimate cerarii each bear four cerarian setae at the apex of a tubercle. Anteriorly the cerarii are each composed of two large setae except the ocular cerarius which usually contains but one seta. The tubercles become smaller anteriorly but the frontal cerarius is often large and lies on the ventral THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) 33 o FIG. 14 34 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) surface. Each tubercle bears from one to five extremely long setae and one or two trilocular pores ; occasionally there is also a circular disc pore either on the dorsal or ventral side. Dorsal setae not numerous, of various sizes, but all slender. Some of these setae are very long and become flagellate distally ; on the anterior part of the body they form groups as illustrated. Dorsal multilocular disc pores and tubular ducts absent. Circular disc pores, larger than the trilocular pores are distributed over the surface and they lie in definite groups on the anterior half of the body associated with the groups of setae. Posteriorly there are a few in the mid-region of each segment and a few laterally. Trilocular pores sparse, associated with the groups of setae. Ventral surface with a pair of long, apical setae. As the anal lobe tubercles are heavily sclerotized it is possible that there is an anal lobe bar which is masked ; an examination of young adult females would possibly show if this is correct. Ventral setae of various sizes but all slender, not numerous. Multilocular disc pores confined to the last three segments, there being eleven to seventeen on the seventh segment, nine to thirteen on the eighth segment and five to eight between the anal lobes. On the two prevulvar segments they occupy single transverse rows and altogether there are scarcely more than thirty-five present. Circular disc pores similar to those on the dorsum, sparsely scattered. There is usually one on the margin of each abdominal segment and others in marginal groups anteriorly and between the coxae. Ventral tubular ducts absent. Trilocular pores sparse. NOTES. Although this species has close affinities to the genus Tylococcus it is not certain whether it is congeneric. It differs from T. madagascariensis Newstead, the type of genus, in possessing numerous circular disc pores on the dorsal surface and very long setae with flagellate ends. These characters are shared with T. westwoodi Strickland, a discussion of which follows but the species differs from westwoodi in possessing eighteen pairs of cerarii instead of eleven or twelve. Tylococcus westwoodi Strickland (Text-fig. 15) Tylococcus westwoodi Strickland, 1947, Bull. ent. Res. 38 : 510. HABIT. Described from Atikpale, Eastern Province, Gold Coast on Theobroma cacao attended by an undetermined ant of the genus Crematogaster. Strickland gave the following account of the adult female : " Body colour apparently dark red, but material preserved two days in alcohol before examination, so no field description is possible. Specimens with a few strands of a dark red wax still adhering to the dorsum." RECOGNITION CHARACTERS. Adult female broadly oval measuring approximately 1-5 mm. long. Antennae 8-segmented. Legs short and stout with a few translucent pores on the hind coxa and tibia. Circulus present. Ostioles well developed, with the inner edges of the lips heavily sclerotized and each lip with two or three long setae and three or four trilocular pores. Anal ring lying about one and a half times its diameter from the apex of the abdomen, with six setae which are only a little longer than the diameter of the ring. Cerarii numbering eleven or twelve pairs. THE MEALY-BUGS ( PSEUDOCOCCID AE : HOMOPTERA) 35 ./' /..; W( : '<'',' /A (' e ' / A . /x ' /?. / ./.' V/ . //>//. /*./ v' FIG. 15 36 THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) Anal lobe cerarii each composed of two stout conical setae which are blunt at the apices, surrounded by about three trilocular pores and two long, stout, auxiliary setae. The anal lobes form prominent sclerotized tubercles. Anteriorly there is a pair of cerarii to each abdominal segment and the other cerarii are located on the thorax and head. Most of these cerarii are composed of two conical setae often of unequal size but some of the anterior cerarii may have only one seta. The posterior cerarii are borne at the apices of poorly developed tubercles and the anterior cerarii are each surrounded by a circular sclerotized area bearing an occasional trilocular pore and one or two auxiliary setae, these often extremely long. Dorsal setae of various lengths, mainly slender. Many of these setae are extremely long with the distal end flagellate, present on the margins and irregularly over the surface especially on the head and thorax. Dorsal multilocular disc pores and tubular ducts absent. Circular disc pores numerous in mid-regional groups on each segment. They become scattered laterally. Trilocular pores sparse. Ventral surface with the anal lobes sclerotized on the margins and with a small sclerotized anal lobe bar and a bar seta which is of similar length to the anal ring setae. Apical seta detached from the anal bar, nearly twice as long as anal ring setae. Ventral setae of various sizes mainly short and slender but there are a few long setae around the margins similar to those on the dorsal surface. Multilocular disc pores confined to the four posterior segments. On the sixth segment there are usually one or two, on the seventh and eighth segments there are single transverse rows of about six pores and on the last segment there are one or two pores. Altogether there are scarcely more than twenty pores present. Tubular ducts distributed mainly in marginal groups on the fourth to eighth abdominal segments but others may be present on the mid-region of the abdominal segments. Circular disc pores not so numerous as on dorsum ; they are rather scattered and do not form definite groups. Trilocular pores not numerous. NOTES. All the species seen appear to be young adults and it may be that the marginal tubercles bearing the cerarii at the apices become more developed in the older specimens. This species seems to be congeneric with T. malacanthae Strickland, already discussed, in possessing numerous dorsal circular disc pores and extremely long setae with the distal ends flagellate. REFERENCES BALACHOWSKY, A. S. 1954. Sur 1'Indigenat et le Statut de Catenococcus loranthi Strickl. (Coccoidea : Pseudococcini) en Afrique Occidentale. Rev. Path. veg. 33 : 247. BORKHSENIUS, N. S. I94Q. Fauna of the U.S.S.R. Homoptera, Coccoidea, Pseudococcidae. Biological Institute of the U.S.S.R., New Series, No. 38. DE LOTTO, G. L. 1957. The Pseudococcidae (Horn. : Coccoidea) described by H. C. James from. East Africa. Bull. Brit. Mus. (Nat. Hist.) Ent. 5 : 185-232. EZZATT, Y. M. & McCoNNELL, H. S. 1956. The Mealybug Tribe Planococcini (Pseudo- coccidae, Homoptera). Univ. Maryland Agric. Exp. Sta. Bull. A 84. FERRIS, G. F. 1950. Atlas of Scale Insects of North America, 5. Stanford University, California. 1953. Ibid. 6. Stanford University, California. 1955- On some genera of the Pseudococcidae. Microentomology , 20 : 1-6. I 955- Atlas of Scale Insects of North America, 7. Stanford University, California. THE MEALY-BUGS (PSEUDOCOCCID AE : HOMOPTERA) 37 Goux, L. 1934. Notes sur les Coccides (Hem.) de la France. (9* note). Contribution a 1 e"tude du genre Heliococcus avec description de deux especes nouvelles. Bull. soc. ent. Fr. 39 : 164-171. HALL, W. J. 1937. Observations on the Coccidae of Southern Rhodesia. Trans. R. ent. Soc. Lond. 86 : 119-134. 1941. On some new species and two new genera of Coccidae (Homoptera) from Southern Rhodesia. /. ent. Soc. S. Afr. 4 : 237. 1945. The identity of a Mealybug vector of " Swollen Shoot " virus disease of cacao in West Africa. Bull. ent. Res. 36 : 305-313. JAMES, H. C. 1936. New mealybugs from East Africa. Trans. R. ent. Soc. Lond. 85 :igj-2i6. STRICKLAND, A. H. 1947. Coccids attacking Cacao (Theobroma cacao L.), in West Africa, with descriptions of five new species. Bull. ent. Res. 38 : 497-523. 19470. Three new species of Coccoidea (Hemiptera : Homoptera) from the Gold Coast, British West Africa. Proc. R. ent. Soc. Lond. (B) 16 : 149-156. WILLIAMS, D. J. (1958). The mealy-bugs (Pseudococcidae : Homoptera) described by W. M. Maskell, T. D. A. Cockerell, R. Newstead and E. E. Green from the Ethiopian Region. Bull. Brit. Mus. (Nat. Hist.) Ent. 6 : 205-236. ENTOM. 7. I, THE AFRICAN SPECIES OF STIVALIUS, A GENUS OF SIPHONAPTERA F. G. A. M. SMIT BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. 2 LONDON: 1958 THE AFRICAN SPECIES OF STIVALIUS, A GENUS OF SIPHONAPTERA BY F. G. A. M. SMIT Pp. 39-76 ; 65 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. 2 LONDON: 1958 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 7, No. 2 of the Entomological series. Trustees of the British Museum, 1958 Issued August, 1958 Price Ten Shillings THE AFRICAN SPECIES OF STIVALIUS, A GENUS OF SIPHONAPTERA By F. G. A. M. SMIT THE seven species of the large genus Stivalius (Family Pygiopsyllidae) which were hitherto known to occur in Africa 1 are redescribed and figured in the present paper, while six new species of Stivalius from Africa are described and also the hitherto unknown male of 5. sellatus ; a key is provided for the identification of these thirteen species. The specimens examined are in the Rothschild and British Museum collection of fleas at Tring, unless stated otherwise. The thirteen African species of Stivalius belong to two groups which can be distin- guished as follows : (a) Genal margin below the eye divided into two partly overlapping small lobes (Text- fig, i) ; cJ movable process of clasper without a dense group of thin setae on inner side (Text-fig. 5) ; tendons of phallosome very short, not or hardly reaching beyond the apex of the aedeagal apodeme (Text-fig. 16) ; dorsal margin of aedeagal apodeme nearly straight, not deeply concave preapically (Text-fig. 16) ; - no paired sclerotic structure alongside the bursa copulatrix (Text-fig. 25) ; dilated part of ductus spermathecae slender, with a number of thick sclerotic internal rings, giving this part of the duct a strong resemblance to a tape-worm (Text-fig. 25)" ; bulga of spermatheca with a thin wall and internal striae (Text-fig. 25) /mMS-group, p. 42 (b) Genal margin below the eye entire (Text-fig. 2) ; <$ movable process of clasper with a dense group of straight and thin setae on inner side and bordering the ventral margin (Text-figs. 8-15) ; tendons of phallosome making at least half a convolution (Text-fig. 17) and often much more (Text -fig. 18) ; apical half of dorsal margin of aedeagal apodeme strongly concave (Text-figs. 17, 18) ; $ bursa copulatrix in most species with a dark sclerotic structure on each side (Text-figs. 26, 28-37) the dilated part of the ductus spermathecae with a large number of very thin internal rings (Text-figs. 26, 28-37) ' bulga of spermatheca with a thick wall and without internal striae (Text-figs. 26, 28-37) .... fonws-group, p. 47 1 Jordan & Rothschild (1922, Ectoparasites, 1 : 252, 254) recorded Stivalius ahalae and S. aporus from Mfongosi, Zululand ; these two rat-parasites do not belong to the African fauna, but to that of India and Burma. In all probability this record was due to some error, perhaps mislabelling. 2 The ductus spermathecae is similarly ringed in females belonging to the following groups of Stivalius : robinsoni-group (squirrel-parasites : S. robinsoni (Rothschild) (Malaya, Sumatra), S. javanus Jordan (Java), S. rhaebus Jordan (Borneo) and 5. lonchus Jordan (Borneo)) ; a/ia/ae-group (rat-parasites : S. ahalae (Rothschild) (India), S. aporus Jordan & Rothschild (India, Ceylon), S. phoberus Jordan & Rothschild (Ceylon) and S. cognatus Jordan & Rothschild (Java)) ; jacobsoni-group (rat-parasites : S. jacobsoni (Jordan & Rothschild) (Java, Sumatra) and S. klossi (Jordan & Rothschild) (Annam, Thai- land, Malaya, Sumatra, Java)) ; squirrel and Tupaia parasite S. mjobergi Jordan (Borneo). ENTOM. 7, 2 3 42 THE AFRICAN SPECIES OF STIVALIUS FERINUS-GROVP The new species described below is the only known representative of the ferinus- group in Africa. Of the other three species, belonging to this group, two occur in the Oriental Region (Ceylon, India, Malaya) and one in Japan. Stivalius alienus sp. n. (Text-figs, i, 3, 5-7, 16, 25) TYPE MATERIAL. Male holotype, female allotype and 8 male paratypes from Calonne plantation, nr. Elisabethville, Belgian Congo, from a nest (probably of a gerbil) , 9 . vi . 1953 ; i female paratype from the same locality, from Rattus (Mastomys) natalensis, vi . 1953 ; 2 female paratypes, nr. Elisabethville, from Rattus (Mastomys) natalensis, viii . 1953 ; all collected by Dr. R. Devignat, to whom one pair of paratypes has been returned ; i male paratype from the neighbourhood of Elisabethville, from Crocidura pilosa, 1957 ; i female paratype, same locality, from Mus triton, 1957 ; i female paratype, same locality, from Pelomys fallax, 1957 these three specimens were collected by P. L. Pirlot and the male and one female are in the Musee Royal du Congo Beige, Tervuren ; i male paratype from the Suji Valley, 6,000 ft., S. Pare Mts., Tanganyika, i.i957, from Arvicanthis sp., collected by J. G. Halcrow. DIAGNOSIS. A member of the/mVms-group, which includes S.ferinus (Rothschild) (a shrew-parasite from Ceylon and India), 5. insolli Traub (a bird-parasite from Malaya) and 5. aestivalis Jameson & Sakaguti (a wood-mouse (Apodemus) parasite from Japan). The new species differs from 5. ferinus by the unmodified (not sub- spiniform) setae in the submarginal frontal row, from S. insolli and S. aestivalis by the absence of a row of short setae preceding the main row of setae on the pronotum, while in 5. insolli the number of pronotal spines is about 30 as against 20 in 5. alienus. There are also differences in the genitalia between these species. DESCRIPTION. HEAD (Text-fig, i). Frontoclypeal margin smoothly rounded. Preoral tuber short. Submarginal frontal row consisting of six setae in both sexes ; between this row and the eye there are about a dozen large and fairly large setae and numerous minute setae on the preantennal region of the head, the minute ones absent on the gena. Genal margin below the eye divided into two short lobes the anterior of which partly overlaps the posterior. Frontal area of micropores relatively narrow. Eye well developed, kidney-shaped. Maxillary palps not quite reaching to the middle of the anterior margin of the fore coxa ; the first segment longer than the second, while the third segment is the shortest of the four. The laciniae are smooth basally and extremely finely serrated apically. The labial palp, reaching to about two-thirds the length of the fore coxa, consists of five segments. Scapus of antenna on the outer side of its widened portion with five to six thin setae in the male and three to four in the female ; pedicellus in both sexes with six slender setae, several of which reach to or a little beyond the first segment of the clava ; the clava consists of the usual nine segments (excluding the petiolus). Postantennal region of head with three rows of setae (the displaced seta between the first and second row in THE AFRICAN SPECIES OF STIVALIUS 43 Text-fig, i is an abnormality) and a large seta about mid- way between the lowest seta of the second and third row ; the first row consists of five setae each side in the male and six in the female, while the second row normally has six setae each side in both sexes as has also the third row. Bordering the antennal fossa posteriorly are about 11-14 small setae in both sexes. THORAX. Pronotum (Text-fig, i) narrow, with one row of six setae each side and a ctenidium of 20 slightly curved spines which are longer than the pronotum. Mesonotum with a main row of five setae each side, preceded by two rows of more numerous small and irregularly placed setae ; two fairly long pseudosetae dorsally under the collar of the mesonotum. Mesepisternum with three to four setae, of which one or two are usually small ; mesepimeron normally with six (sometimes seven) setae. Metanotum with three rows of setae ; the first row consists of two to three setae in the male, four to six in the female, the second and third rows in both sexes with seven to ten and six setae respectively (the lowest seta of the main row much smaller than the others in the row) ; in the female the first metanotal row is preceded by one or two small dorsal setae. Pleural arch well-developed. Met- episternum with one large and one or two minute setae ; metasternum dorso- posteriorly with one large seta ; metepimeron with one to three small and eight to nine large setae in the male, three to four small and nine to eleven large setae in the female. LEGS. Fore coxa with numerous setae all over the outer side ; mid coxa with setae along the lower half of the anterior margin, a patch of setae on the outer side of the ventro-anterior part and two (sometimes three) ventro-posterior setae ; the oblique suture of the outer surface of the mid coxa is uninterrupted ; chaetotaxy of hind coxa similar to that of mid coxa, but in addition there is a small group of short setae ventro-anteriorly on the inner side. Fore femur, apart from marginal setae, with 12-16 lateral setae on the outer side and only one very small seta on the basal part of the inner side. Mid and hind femora with the usual marginal and submarginal setae, but without lateral setae. All tibiae with seven notches in the posterior (dorsal) margin, the most dorsal one bearing only two smallish setae ; chaetotaxy of the hind tibia as shown in Text-fig. 3. Fifth segment of all tarsi with six pairs of lateral plantar setae, arranged as is usual in the genus, namely the first and third pairs shifted on to the plantar surface in the fore and mid tarsus, while in the hind tarsus only the third pair is shifted on to the planta ; in the male the last segment of fore and mid tarsi has four short and stout subapical plantar setae. The two preapical lateral setae are short on the fifth segment of all tarsi, reaching to about the middle of the claws. ABDOMEN. Tergum I with three distinct rows of setae and a few dorsal setae in front of the first row ; terga II-VII with two distinct rows of setae and in addition several dorsal setae in front of the first row and these may form an irregular short row. The numbers of setae in the main row on each side of terga I-VII are in the male : 4, 7, 7, 7, 7, 7, 7 respectively ; in the female : 4, 7, 8, 8, 8, 7, 4 (or 5). Terga II-V in both sexes each with one marginal spinelet on each side near the dorsum. Both sexes with two antesensilial setae, the lower of which is more than twice the length of the upper ; in the female (Text-fig. 6) the margin of tergum VII between ENTOM. 7, 2 3 THE AFRICAN SPECIES OF STIVALIUS FIGS, i, 2. Head and pronotum of: i. Stivalius alienus sp. n. (female paratype, plantation Calonne). 2. S. torvus (Rotschild) (female, Kisii, Kenya). Figs. 3, 4. Hind tibia of : 3. S. alienus sp. n. (female allotype). 4. S. torvus (Rothschild) (female, Keruguya, Kenya). THE AFRICAN SPECIES OF STIVALIUS 45 FIGS. 5-7. Stivalius alienus sp. n. 5. Clasper and sternum IX (holotype). 6. Termi- nalia (allotype). 7. Outline of sternum VII of female (a paratype, nr. Elisabeth vi lie, b paratype, plantation Calonne). 46 THE AFRICAN SPECIES OF STIVALIUS the two sets of antesensilials is produced into a short triangular lobe ; below the antesensilials the margin forms in the female an angulate lobe and below this the margin is slightly concave for a considerable distance. Basal abdominal sternum with a lateral patch of two to four setae in the male and 13-17 in the female, and with two setae each side along the ventral margin of which one is placed in front of the other. Sterna III-VII in the male normally with three setae each side in the main row, in the female the main row of sterna III-VI consists of four setae ; in both sexes these main rows are preceded by a patch of numerous smaller setae. MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 5, 16). Tergum VIII with 0-3 setae each side anterior to the vertical part of the spiracular fossa. Sternum VIII with about 30-35 setae each side. Apodeme of tergum IX narrow, ventrally not solidly fused with the dorso-caudal part of the manubrium, the latter basally very broad and tapering gradually to an upturned tip (Text-fig. 5). Fixed process of clasper with two (one short, one long) acetabular setae. Movable process (Text-fig. 5) of a shape characteristic for the majority of the representatives of the genus, with relatively few setae along the ventral (posterior) margin and a group of three large setae and one smaller one placed along this margin just before the bend. Proximal arm of sternum IX (Text-fig. 5) fairly broad ; the distal arm of this sternum narrow in its basal half whence it widens gradually, its dorso-apical portion smoothly rounded ; four or five of the setae along the apical part of the ventral margin much stouter than the other setae of the apical portion. Phallosome as in Text-fig. 16 ; note the very short tendons of the phallosome, the straight and simple inner tube, the dorso-apical aedeagal sclerite with two sharp apical projections, and the long and narrow caudally curved and membranous lobe of the ventral lateral wall. FEMALE (Text-figs. 6, 7, 25). Posterior margin of sternum VII (Text-figs. 6,7) with a double sinus, the upper bay of which is much smaller than the lower ; the main row consists of five strong setae and is divided by a gap between two dorsal setae and three ventral ones. In front of this row are numerous smaller setae. Vari- ation in the outline of the posterior margin of sternum VII as shown in Text-figs. 6, 7. Tergum VIII with three to six setae in front of the widened vertical part of the spiracular fossa ; chaetotaxy of the ventral part of tergum VIII as in Text-fig. 6. Sternum VIII apically narrow and with several minute setae at and near the apex. Anal segment as in Text-fig. 6 ; anal stylet about thrice as long as its maximum width, with one long apical seta and two minute preapical ones. Bulga of sperma- theca (Text-figs. 6, 25) longer than wide, with a dorsal hump ; the hilla protrudes deeply into the lumen of the bulga and bears apically a papilla. Ductus bursae curved, bursa copulatrix with a longish posterior internal sclerotization. The basal half of the ductus spermathecae is internally reinforced by numerous sclerotic rings (Text-fig. 25). LENGTH. <$ 2|-2| mm., $ 3-3! mm. REMARKS. Even without the host records it would have been possible to deduce Stivalius alienus to be probably a parasite of rodents. The species of the ferinus- group provide an excellent example of the modifications of the pronotal ctenidium which arise in response to the nature of host-relationships. In S. alienus (Text- fig.i) THE AFRICAN SPECIES OF STIVALIUS 47 and in 5. aestivalis (Text-fig. 63) (a flea of Apodemus) the pronotal ctenidium consists of fairly straight spines this is the usual type of ctenidium in a large number of rodent-fleas. In 5. ferinus (Text-fig. 64), a parasite of shrews, the spines of the pronotal ctenidium are blunt and distinctly curved and are longer than the pronotum this is characteristic of a number of shrew-fleas. In 5. insolli (Text-fig. 65), a bird-parasite, the number of pronotal spines has increased considerably about 30 as against 18-20 in the three mammal parasites of this group ; fleas of the super- family Ceratophylloidea which have become parasites of birds always have a larger number of pronotal spines (usually more than 24) than related forms living on mammals. TORVUS-GROUP The members of this group, which is confined to Africa, are rather uniform in the structure of the head, thorax, legs and unmodified abdominal segments ; the main differences between the species are in the genitalia. Stivalius torvus, by far the commonest and most widespread member of the group, is described in detail, and the other species are described in comparison with this species. Stivalius torvus (Rothschild), 1908 (Text-figs. 2, 4, 8, 17, 26, 28, 38, 39, 53, 54) Pygiopsylla torvus Rothschild, 1908, Ent. mon. Mag. 44 : 77. Pygiopsylla " afer " Jordan & Rothschild, 1913, Novit. zool. 20 : 537 (err. det., i $ from Kagamba, Uganda ; see Jordan & Rothschild, 1922, Ectoparasites, 1 : 252). Stivalius " afer " Symes & Hopkins, 1932, Rec. Med. Res. Lab. Nairobi (1) : 18, 19, 40, 44, 56. Stivalius torvus Jordan & Rothschild, 1922, Ectoparasites, 1 : 251, 264, fig. 241 ; Dalla Torre, 1924, Ber. naturw. med. Ver. Innsbruck, 39 : n ; Jordan, 1936, Novit. zool. 39 : 297, figs. 54-56; Jordan, 1937, Novit. zool. 40 : 290 ; Hopkins, 1947, Uganda J. 11 (Suppl.) : 155 ; Jordan, 1948, in Smart, Insects of medical importance (London) : 240 ; Hopkins, 1949, Rep. rats, fleas, plague, Uganda: 9, tables 2, 6, 10, 12, 20. MATERIAL EXAMINED. TANGANYIKA : Tengeru, Rattus (Mastomys) natalensis, 26. ii. 1952, i <$, i $. KENYA : Keruguya, Rattus (Mastomys) natalensis, Lophuromys flavopunctatus aquilus, Lemniscomys sp., Otomys sp., 1935-36, 10 ^ 8 - ; without locality and host, 1927, i $, and 1913, I ?. D \ AAV Li \ o / o / * ,? \ o r ' ? ,/ ;r FIG. 9. Pseudococcus vnallyi Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 99 spicuous with lips membranous and without trilocular glands or setae. Circulus missing. Legs all well developed with a small denticle on claw ; tarsal digitules finely pointed ; ungual ones knobbed at apex. Antennae with eight short and stout joints. Anal ring entire with six robost setae. Pseudococcus mirabilis Brain (Text-fig. 10) Pseudococcus mirabilis Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 121. Four slides of which three containing a single specimen and one with five specimens were available. Two slides were labelled : " Pseudococcus mirabilis Br.; on 1 ; Ceres, C.P.; Oct. 1898 ; B. 54 ; C.K.B." Remaining two slides marked only " Pseudococcus mirabilis ; B. 54, C.K.B." All specimens were adult females in very poor condition, partly broken and badly distorted. The ovisacs are creamy-white or slightly buff-coloured, and are aggregated on the leaf-cluster bases in conspicuous masses . . . The adult is apparently vivi- parous . . . When fully distended it is no more than 1-5 mm. long." (Brain, I.e.) . Mounted specimens elongate to very broadly elliptical with dermis membranous. Marginal cerarii present only on last three occasionally four abdominal segments. Each cerarius is formed by two stout conical spines surrounded by many tubular ducts, without any grouping of trilocular pores ; auxiliary setae absent ; area about cerarian spines not chitinized. Third cerarius anterior to anal lobe one, when present, reduced to a single spine. Ventral side of each anal cerarius with a robust apical seta about same size as those of anal ring ; chitinized bar absent. Multilocular disc pores arranged in five ventral groups. The number of pores in one specimen was as follows : (v) 17 ; (vi) 25 ; (vii) 26 ; (viii) 27 ; (ix + x) 29. On segments anterior to genital opening they are set in fairly linear transverse rows along distal margin. A few pores are scattered on dorsum of abdomen. Tubular ducts with oral collar of two different sizes. Large ones set in groups on ventral and dorsal marginal areas as far as head ; a few others arranged on dorsum along distal margin of abdomi- nal segments anterior to anal lobes. Tubular ducts of smaller size occurring in trans- verse irregular rows on either side of abdomen ; others scattered all over body. Trilocular pores very few and uniformly distributed. Circular disc pores about half the size of trilocular pores, few and scattered on dorsum and venter. Dorsal and ventral setae about same size, all rather short, stout to very stout, not numerous ; a few setae on dorsum of last abdominal segments similar hi shape and size to those of marginal cerarii. Anterior dorsal ostioles not recognizable ; posterior ones fairly prominent with lips membranous and provided with a few small setae and trilocular pores. Circulus absent. Legs well developed, with some translucent pores on hind femur and coxa ; claw without denticle ; ungual and tarsal digitules apically knobbed. Anal ring V-shaped with six setae. Antennae 7-jointed with a pseudo- articulation on apical joint. 1 The name of the host plant is omitted. According to the original description it is Borbonia cordata loo THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA O FIG. 10. Pseudococcus mirabilis Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 101 Pseudococcus muraltiae Brain (Text-fig, n) Pseudococcus muraltiae Brain, 1912, Ann. ent. Soc. Amer. 5 : 184. Two slides were seen in Pretoria, one of which containing larvae ; the second with a single specimen distorted and partly broken labelled : " Pseudococcus muraltiae Brain; on Muraltia heisteria; Newlands : 22.x. 1910; paratype ; 52, C.K.B." Another slide with two specimens bearing the same collecting data was previously examined at the British Museum (Natural History), London. " Adult female : small ; largest specimens, with completed ovisac, was 1-9 mm. long by 1-13 mm. broad, slatey-grey in colour ; waxy secretion scant but segmenta- tion conspicuous. Lateral appendages were absent but usually four caudal ones present." (Brain, I.e.}. Body of mounted specimens rather broadly elliptical, membranous. Margin of body with seventeen pairs of cerarii. Anal lobe cerarii each with two moderately robust conical spines beset by few trilocular pores and four to six small slender auxiliary setae. Area about spines not chitinized. Each of remaining cerarii with two spines which tend to be more slender anteriorly, where they attain about same size and shape as dorsal setae ; each cerarius is surrounded by a group of two to five trilocular pores without auxiliary setae ; one or two thoracic cerarii reduced to a single spine. Ventral side of each anal lobe without chitinized bar ; apical seta robust, longer than those of anal ring ; subapical one much shorter. Multilocular disc pores few, occurring on ventral side of last five abdominal segments. The number of pores in one specimen was as follows : (v) 3 ; (vi) 19 ; (vii) 17 ; (viii) 45 ; (ix + x) 16. On segments anterior to genital opening they are arranged in transverse rows along distal margin only. Ventral tubular ducts with oral collar not numerous and mostly distributed on marginal area of last five or six abdominal segments ; a few occur on median and submedian areas in association with multilocular disc pores. Dorsal tubular ducts with oral rim very few. One duct is normally associated with each frontal cerarius and one with each preanal cerarius ; two or three occasionally occur on thorax and one on median area of preanal segment. Trilocular pores not abundant but evenly distributed. Circular disc pores noticeably smaller than trilocular pores, very few and scattered. Ventral setae moderately long and slender; dorsal ones much shorter. Circulus absent. Anterior and posterior dorsal ostioles inconspicuous with lips membranous and provided with two or three small setae and a few trilocular pores. Legs well developed but rather short with a few translucent pores on hind coxa. Antennae with eight joints. The synonymy of Pseudococcus simulator James with muraltiae published in our previous paper (De Lotto, 1957) has to be rejected because in muraltiae the circulus is absent. P. simulator is instead a synonym of burnerae as pointed out in our notes on dealing with that species. Pseudococcus natalensis Brain (= Nipaecoccus graminis (Maskell)) Pseudococcus natalensis Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 100. We have been kindly informed by Dr. D. J. Williams of the Commonwealth Institute of Entomology, London, that this species is identical with Nipaecoccus 102 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA FIG. ii. Pseudococcus muraltiae Brain THE PSEUDOCOCCIDAE (ROM. COCCOIDEA) FROM SOUTH AFRICA 103 graminis (Maskell). Recently he examined type material of Maskell's species and this new synonymy is dealt with in his paper on Pseudococcidae described by Maskell, Cockerell, Newstead and Green from the Ethiopian region (1958). Pseudococcus nitidus Brain (Text-fig. 12) Pseudococcus nitidus Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 143. The material examined was represented by six slides, three of which each contained a single specimen labelled : " Pseudococcus nitidus Brain ; on Acacia caffra ; Pretoria: Nov. 1914 ; paratype ; 39, C.K.B." All specimens were in very poor condition, distorted and partly broken, having been mounted from dry material. The remaining three slides contained only larvae and males. The ovisacs . . . are closely felted, smooth, and in the majority of cases have the shape and approximate size of an adult insect, being about 3 mm. long and so smoothly felted on the exterior as to look like a piece of white kid . . . Adult female 2 '5 mm. long, translucent brown in colour ; legs and antennae of the same colour. No waxy secretion and no filaments except two extremely short caudal ones which appear as two white specks." (Brain, I.e.). Body of mounted specimens rather elongate elliptical with anal lobes strongly developed ; dermis at maturity membranous. Margin of body with only five pairs of cerarii on last abdominal segments. Anal lobe cerarii each formed with two small conical spines set apart from each other, without auxiliary setae or grouping of trilocular pores ; area about spines clear. Remaining cerarii each with two spines which tend to be smaller and more widely separate from each other anteriorly ; on fourth cerarius anterior to the anal lobe one they attain shape and size of dorsal setae. Ventral side of each anal lobe with a long and robust apical seta and a much shorter subapical one ; chitinized bar absent. Multilocular disc pores few and present only on ventral side of last three abdominal segments. The number of pores in one specimen was as follows: (vii) ii ; (viii) 24 ; (ix + x) 32. On segments (vii) and (viii) they are arranged in transverse rows along distal margin only. Tubular ducts with oral collar very few on ventral side of last abdominal segments, mostly in association with multilocular disc pores. Tubular ducts with oral rim very numerous on dorsum and extending all along marginal area of venter. Quinquelocular pores fairly abundant on median and submedian areas of venter except on segment posterior to genital opening, where they are missing. Trilocular pores not numerous and evenly distributed. Circular disc pores apparently absent. Anterior and posterior dorsal ostioles not detected. Circulus absent. Legs long and rather slender, with a small denticle on claw ; ungual digitules knobbed at apex ; tarsal ones finely pointed ; no translucent pores on hind legs. Anal ring of Pseudococcid type, with six setae. Stigmatic openings unusually large. Dorsal setae rather few, small, spiniform ; ventral ones longer and slender. Antennae formed by nine long slender joints. io 4 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA FIG. 12. Pseudococcus nitidus Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 105 Pseudococcus quaesitus Brain (Text-fig. 13) Pseudococcus quaesitus Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 123. Pseudococcus elisabethae Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 126 (SYN. NOV.). Pseudococcus trichiliae Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 131 (SYN. NOV.). The material available consisted on one slide labelled : " larvae of Pseudococcus quaesitus Brain ; Pretoria: Nov. 1914 ; larvae; 60, C.K.B."; another slide labelled: " cJ of Pseudococcus quaesitus Brain ; Pretoria ; $ paratype ; $ 60 ; Nov. 16, 1914." Six more slides were marked with the serial number " 60, C.K.B." one of which with larvae only ; the remaining five slides contained altogether eleven old adult females. Three more slides with altogether nine old adult females bore the follow- ing data : " Pseudococcus quaesitus Brain ; Acacia horrida ; Grahamstown : Jan. 1899 ; dry material ; 63 ". " The ovisacs are often collected into masses which remind one of Ps. filamentosus Ckll., but present a pinkish tinge rather than yellow or greyish. Seen singly as in cavities in tree-trunks, the ovisacs are usually more or less button-shaped, with straight vertical sides and a rounded top. The largest observed measured approxi- mately 3 mm. in diameter . . . Adult female may reach 4 mm. in length, pinkish coloured at first and later purplish, with dense, white, powdery secretion. Lateral filaments short and fragile. Caudal filaments two in number, stout, may attain one third the length of the body." (Brain, I.e.}. Body of mounted specimens elliptical. Margin of body with a series of seventeen pairs of cerarii. Anal lobe cerarii each formed by two conical spines surrounded by a small group of trilocular pores and a few slender auxiliary setae ; area about spines not chitinized. Remaining cerarii each with two spines which tend to be noticeably more slender anteriorly. Each cerarius is associated with a grouping of a few trilocular pores but no auxiliary setae. Ventral side of each anal lobe with a robust apical seta, longer than those of anal ring ; subapical seta very short ; chitinized bar absent. Multilocular disc pores rather few and arranged in five groups on ventral side of last abdominal segments. The number of pores in one specimen was as follows : (v) 24 ; (vi) 19 ; (vii) 21 ; (viii) 29 ; (ix + x) 21. On segments anterior to genital opening they are set in transverse rows along distal margin. Tubular ducts with oral collar set in five or six small groups on ventral marginal area of abdomen. Tubular ducts with oral rim fairly numerous on dorsum and a few on venter on marginal and submarginal areas of thorax. Trilocular pores moderately numerous and uniformly distributed on both sides of body. Circular disc pores apparently absent. Dorsal setae short ; ventral ones longer but slender, in either case not numerous. Anterior and posterior dorsal ostioles well developed with lips membranous having some trilocular pores and a few short setae. Circulus absent. Legs well developed, robust, with a few translucent pores on hind femur. Anal ring normal, with six setae. Antennae 8-jointed, at times with a pseudoarticu- lation on apical joint. Pseudococcus elisabethae and trichiliae described by Brain in the same paper are synonyms of quaesitus. 106 THE PSEUDOCOCCIDAE (ROM. COCCOIDEA) FROM SOUTH AFRICA FIG. 13. Pseudococcus quaesitus Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 107 Pseudococcus segnis Brain (Text-fig. 14) Pseudococcus segnis Brain, 1915, Trans, roy Soc, S. Afr. 5 : 145. One slide with a single very old adult female in fairly good condition was made available from the U.S. National collection of Coccidae, Washington, D.C. It bore the following data: "Pseudococcus segnis Brain; Stellenbosch : I7.xii.i9i4; B. 55, C.K.B." Although the collecting data is not exactly the same recorded in Brain's paper, that is I3th December, 1914, the serial number is identical and we can assume that the specimen acually belongs to the original type series. " Adult female : the four specimens range from 2-8 to 3-4 mm. in length, and are dark olivaceous-green in colour. There are no lateral or caudal filaments, and only a slight trace of white secretion, the insects appearing rather greasy or slug-like. This absence of secretion may be due to shaking in transit." (Brain, I.e.). Body elliptical with eighteen pairs of cerarii. Anal lobe cerarii each apparently formed by two spines 1 surrounded by eight to ten trilocular pores ; auxiliary setae absent ; area about spines not chitinized. Remaining cerarii each provided with two small and slender conical spines which on most anterior pairs are not differentiated from setae of dorsum. Each cerarius is beset by five to eight trilocular pores. Ventral side of each anal lobe without chitinized bar ; apical seta about as long as those of anal ring ; subapical one shorter. Multilocular disc pores, tubular ducts and circular disc pores entirely absent. Trilocular pores fairly abundant and uniformly distributed. Dorsal setae few and very small, spiniform ; ventral setae also few but much longer. Anterior and posterior dorsal ostioles well developed, lips membranous with many trilocular pores and a few minute setae. Circulus absent. Legs normal with a denticle on claw ; ungual and tarsal digitules apparently pointed ; hind legs without trans- lucent pores. Antennae g-jointed. Pseudococcus socialis Brain (Text-fig. 15) Pseudococcus socialis Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 103. Three slides, one with ova and two with altogether three specimens in fairly good condition were seen. All were labelled : " Pseudococcus solitarius sp. n.; Pretoria, Union Buildings : 20. xi. 1914; C.K.B. ; on grass ; C.K.B.; 526." Even in this instance the specific name under which the species was described does not agree with that originally assigned on the type slides. Nevertheless there were no doubts that the material belonged to socialis as all collecting data and serial number are exactly the same as this species referred by Brain in his paper. " Ovisac : an irregular mass of white cottony secretion, about 3 mm. in diameter . . . The adult female is elongate, of a purplish-brown colour, the whole body being slightly powdered with white. There were no signs of lateral or caudal filaments . . . The average size of female insects (containing ova) when mounted is 1-7 mm. 1 In the specimens examined all cerarian spines of the anal lobes were broken away. io8 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA FIG. 14. Pseudococcus segnis Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 109 ^'- D FIG. 15. Pseudococcus socialis Brain no THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA long and 0-8 mm. broad." (Brain, I.e.). Body elongate elliptical, membranous. Margi- nal cerarii recognizable only on last two abdominal segments. Anal lobe cerarii each with two rather slender conical spines beset by a few trilocular pores ; area about cerarian spines not chitinized ; auxiliary setae absent. Preanal cerarii each with two spines more slender and set somewhat apart from each other, without grouping of trilocular pores or auxiliary setae. Ventral side of each anal lobe with a robust apical seta noticeably longer than those of anal ring; subapical seta much shorter ; chitinized bar absent. Multilocular disc pores arranged in five groups on ventral side of last abdominal segments. The number of pores in one specimen was as follows : (v) 14 ; (vi) 35 ; (vii) 64 ; (viii) 75 ; (ix + x) 26. A few pores scattered on marginal and submarginal ventral areas of thorax and head. Dorsal tubular ducts with oral rim few. One occurring on median line of each abdominal segment (iv) to (viii) ; one on marginal area of each abdominal and thoracic segment ; a few more widely scattered. Tubular ducts with oral collar set in transverse irregular rows on last abdominal segments in association with multi- locular disc pores and a few widely scattered all over ventral side of body. Trilocular pores not numerous and evenly distributed all over body. Circular disc pores about same size as trilocular pores, having a granulate surface ; they are few and apparently present only on ventral side of abdomen. Dorsal and ventral setae rather few, slender. Anterior and posterior dorsal ostioles not prominent but with lips membran- ous and with a grouping of a few trilocular pores and two to four minute setae. Circulus small, rounded, with border fairly highly chitinized, set near basal margin of fifth abdominal segment. Legs all short otherwise normal, with some translucent pores. Antennae short, built up with seven joints. Pseudococcus solitarius Brain (= Nipaecoccus vastator (Maskell)) (SYN. NOV.) Pseudococcus solitarius Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 104. Three slides containing altogether six specimens, all badly distorted and broken were examined. They bore the following label : " Pseudococcus solitarius sp. n.; Transvaal, Pretoria and dist.: Sept.-Oct. 1914 ; C.K.B.; on thorn tree ; C.K.B., 65." Carefully examined this species was found identical with Nipaecoccus vastator (Maskell) with which it is synonymized. Pseudococcus stelli Brain (Text-fig. 16) Pseudococcus stelli Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 146. Of three slides seen, only one contained a single adult female in fairly good con- dition. It was labelled : " Pseudococcus stelli Brain ; on Borbonia cordata Linn.; Stellenbosch : Dec. 17, 1914 ; paratypes ; B. 56, C.K.B." The remaining two slides contained males, larvae and ova only. " Ovisac : the ovisacs are rounded masses of cottony material ... In form they appear almost spherical . . . The greatest diameter averages approximately 2-5 mm. . . . The adult female is pale canary-yellow in colour, about 2 mm. THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA ni FIG. 1 6. Pseudococcits stelli Brain ii2 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA to 2-5 mm. long. The lateral filaments are very short, but distinct and gradually increase in length posteriorly. The caudal ones, two in number, are also short, about twice as long as the next pair, stout at the base and tapering towards the tip." (Brain, I.e.). Mounted specimens elliptical, membranous. Margin of body with a complete series of eighteen pairs of cerarii. One of anal lobe cerarii was provided with two conical spines ; opposite one with three ; both surrounded by a group of a few trilocular pores ; auxiliary setae missing ; area about spines not chitinized. All remaining cerarii each formed with two spines more slender than those of anal lobe ones, beset by three to seven trilocular pores, without auxiliary setae. Ventral side of each anal lobe with a robust apical seta longer than those of anal ring ; sub- apical seta much shorter ; chitinized bar absent. Multilocular disc pores abundant on both sides of body, particularly so on dorsum ; on abdomen they tend to be crowded in transverse segmental rows along distal margin of last segments. Quinquelocular pores rather few and scattered on median and submedian ventral areas of thorax and first two abdominal segments. Trilocular pores not numerous and evenly distributed. Circular disc pores apparently absent. Tubular ducts with oral rim entirely missing. Tubular ducts with oral collar of two sizes. Small ones rather few on ventral marginal area of all abdominal segments and in associatiation with ventral abdominal multilocular disc pores. One occasionally two tubular ducts of large size occur on dorsal and ventral marginal areas near each cerarius ; others are scattered all over dorsum. Dorsal setae very short ; ventral ones much longer but slender ; in neither case abundant. Anterior and posterior dorsal ostioles not prominent, with lips membranous having a cluster of a few trilocular pores and two to four small setae. Circulus may or may not be present because the area where normally it occurs was broken away on specimen examined. Legs all well developed, without translucent pores ; claw with a small denticle ; ungual digitules short and knobbed apically ; tarsal ones finely pointed. Antennae with nine joints. Pseudococcus stelli tylococciformis Brain ( Pseudococcus stelli Brain) (SYN. NOV.) Pseudococcus stelli tylococciformis Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 149. This variety was described on a few specimens collected on the same twigs as those attacked by P. stelli with which, according to Brain, they were identical except that they were smaller and the marginal cerarii were inserted on small tubercles a character peculiar to almost all species at the beginning of the adult stage. Although no types, paratypes or else could be seen, we definitely regard this variety as based on very young adult females of stelli with which it is here synonymized. Pseudococcus transvaalensis Brain Pseudococcus transvaalensis Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 129. Trionymus sanguineus James, 1936, Trans. R. ent. Soc. Lond. 85 : 197 (SYN. NOV.). The material of this species examined was as follows : six slides one of which contained larvae and the remaining five with altogether nine specimens, all marked THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 113 with Brain's serial number " B. 46, C.K.B." Two other slides contained altogether three specimens marked with the serial number only " B. 47, C.K.B." Four more slides of which three with a single specimen and one with larvae, all labelled : " Pseudo- coccus transvaalensis Brain ; roots of cornflower ; Pretoria ; 27.xii.igi4 ; C.K.B.; B. 47a, C.K.B." In the original paper the collecting data of these specimens is reported as 28 Dec. 1914, which has to be explained as a pen slip made by Brain on labelling the slides. All above listed material was carefully compared with specimens of Trionymus sanguineus James from Kenya and found structurally identical. The James species which has been redescribed and illustrated in our previous paper (De Lotto, 1957) has to be understood as a synonym of P. transvaalensis Brain. Pseudococcus trichiliae Brain (= Pseudococcus quaesitus Brain) Pseudococcus trichiliae Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 131. Three slides containing altogether six specimens labelled : " Pseudococcus trichiliae Brain ; on Trichilia sp.; Durban, Natal ; 27.x. 1914 ; paratypes ; B. 51, C.K.B." were seen. This species is a synonym of P. quaesitus which Brain described in the same paper on an earlier page. All paratypes of trichiliae are large and very old adult females, stucturally they cannot be differentiated from those of quaesitus. Pseudococcus wachendorfiae Brain (Text-fig. 17) Pseudococcus wachendorfiae Brain, 1912, Ann. ent. Soc. Amer. 5 : 183. Two specimens were examined. One was labelled : " Pseudococcus wachendorfiae Brain; paratype ; on Wachendorfia paniculata ; Newlands Flats: 3.x. 1910; 53." The second bore the following data: "Pseudococcus wachendorfiae Brain; on Wachendorfia paniculata Lin.; Newlands Flats: 2.x. 1910; paratype; 53." Both paratypes were distorted, partly broken and with many body setae missing. " Ovisac : no definite ovisac was found, although where the adult female was situated a definite white granular patch of waxy secretion was noticed on the plant. Adult female : largest specimen found measured while alive 4-1 mm. long and 1-9 mm. broad. The body was finely covered with granular secretion, white, but segmentation was still conspicuous. Lateral appendages of wax were absent, but a short caudal tuft was generally noticeable." (Brain, I.e.). Mounted specimens elongate elliptical, membranous. Cerarii confined to a single pair on anal lobes, each built up with two strong conical spines, beset by several trilocular pores and a few robust auxiliary setae ; area about the spines not chitinized. Ventral side of each anal lobe provided with a long robust apical seta, longer than those of anal ring ; subapical one much shorter ; no chitinized bar. Multilocular disc pores in three groups on ventral side of last abdominal segments. The number of pores in one specimen was as follows : (vii) 15 ; (viii) 34 ; (ix + x) 26. On ii 4 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA O FIG. 17. Pseudococcus wachcndorfiac Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 115 segments anterior to genital opening they are arranged in transverse rows along distal margin. Trilocular pores numerous all over body. Circular disc pores smaller than trilocular pores and abundantly distributed on either side of body. Tubular ducts with oral rim very numerous on dorsum and venter, except on last abdominal segment where they are missing. Tubular ducts with oral collar few on ventral side of abdominal segments only. Dorsal and ventral setae very numerous and unusually long and robust. Anterior and posterior dorsal ostioles rather conspicuous, lips membranous having a cluster of a few trilocular pores and small setae. Circulus absent. Legs well developed ; hind tibiae with numerous small translucent pores. Antennae with eight joints. Puto (?) africanus Brain (Text-fig. 18). Puto (?) africanus Brain, 1915, Trans, roy. Soc. S. Afr. 5: 151. One slide with two adult females in poor condition being very badly distorted was made available from the U.S. National Collection of Coccidae, Washington, B.C. The slide bore the following label : " Puto africanus Brain ; on Tamarix articulata ; Cape Town : Jan. 1898 ; paratype ; B. 70, C.K.B." " Adult female enclosed in a dense felted or papery sac, which is generally white or yellowish in colour . . . The ovisacs, when not deformed by massing together, are regularly elongate oval about 2 mm. long and 1-2 mm. in diameter . . . The adult female as recovered from dry material is merely a black shrivelled mass without indication of secretionary covering of any kind, and without lateral or caudal filaments ... In mounted specimens the body averages 1-7 mm. in length and 0-9 mm. breadth." (Brain, I.e.}. Mounted females oval to broadly oval in outline, membranous. Marginal cerarii recognizable only on anal and preanal segments. Anal lobe cerarii each formed by two very robust spines, somewhat lanceolate in shape, surrounded by a loose group of a few trilocular pores and one or two fairly long stout auxiliary setae ; chitinized area large and extending to ventral side. Preanal cerarii each with two spines of same shape and size as those of anal cerarii, beset by a few trilocular pores ; auxiliary setae missing ; area about spines not chitinized. Ventral side of each anal lobe provided with an apparently robust long apical seta 1 and two to four shorter ones. Multilocular disc pores present only on median and submedian ventral areas as far as prothorax, set widely apart from one another. Tubular ducts of two types. One type being rather long and slender without usual collar but the opening being instead surrounded by a small chitinized keel. These ducts occur on both sides of body, apparently without any particular pattern. Other ducts distinctly larger with a narrow rim occurring on dorsum only. Trilocular pores few. Circular disc pores larger than trilocular pores, not numerous and distributed all over body. Dorsal setae not numerous, fairly stout ; a few on abdomen similar in shape and size to cerarian spines ; ventral ones slender. Anterior dorsal ostioles not recognizable on specimens examined ; posterior ones inconspicuous without setae or grouping of trilocular pores on their lips. 1 Both missing in the specimens examined. n6 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA FIG. 18. Puto africanus Brain THE PSEUDOCOCCIDAE (ROM. COCCOIDEA) FROM SOUTH AFRICA 117 Circulus absent. Legs normal ; claw without denticle ; hind legs without trans- lucent pores ; a few setae on median and hind legs stoutly spiniform. Anal ring of usual Pseudococcid type, opened posteriorly, with six robust setae. 1 Antennae with eight or nine joints. Rhizoecus africanus Brain (= Rhi zoecus falcifer Kuenckel) Rhizoecus africanus Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 89. Three slides containing altogether seven adult and two preadult females were seen. They were labelled : " Rhizoecus africanus Brain ; roots of plants ; Cape Town : Feb. 1906 ; paratype ; 62." Brain's paratypes were compared with the redescription and figure of R. falcifer Kuenckel recently published by Ferris (1953) and our conclusion fully agrees with Hambleton's opinion (1946) that the Brain species is a synonym of falcifer. Tylococcus chrysocomae Brain (Text-fig. 19) Tylococcus chrysocomae Brain, 1915, Trans, roy. Soc. S. Afr. 5 : 93. The material examined was represented by four slides with altogether eight specimens, all old adults partly distorted, labelled as follows : " Tylococcus chrysocomae sp. n.; Grahamstown : 4th March 1915 ; A. Kelly ; on Chrysocoma tenuifolia ; C.K.B., 61." " Ovisac : white, dense, elongate oval, may reach 5 mm. long and 2 mm. in diameter. The ovisac may be single or clustered. The adult female is found at one extremity of the ovisac and often appears as though partly enclosed owing to the median dorsal keel of white secretion . . . When cleared, stained, and mounted, the adult female is 2 mm. to 2-5 mm. long." (Brain, I.e.}. Body of mounted specimens very broadly elliptical, nearly circular ; membranous. Margin of body with cerarii reduced to sixteen or seventeen pairs owing to absence of one or two pairs on thorax. Cerarii on head and last abdominal segments tending to be inserted in a small very broadly rounded prominence. All cerarii formed by two to four robust conical spines, without any grouping of trilocular pores or auxiliary setae ; area about spines not chitinized. Ventral side of each anal lobe with a stout apical seta about same length as those of anal ring ; subapical one much shorter ; chitinized bar absent. Multilocular disc pores fairly numerous on either side of body, distributed without any regular pattern. Quinquelocular pores not abundant and scattered on dorsum and venter. Tubular ducts somewhat departing in their structure from those normally found in Pseudococcidae. They are provided neither with oral rim nor oral collar, the opening being instead surrounded by a small circular chitinized keel, similar to those seen in Puto (?) africanus. They occur abundantly on both sides of body. Trilocular pores few but evenly distributed, circular disc pores apparently absent. Ventral setae rather short and slender ; dorsal ones about same 1 The two setae posterior to the anal ring actually do not belong to it, as Brain stated. They are the cisanal setae which in this species lie unusually close to the anal ring. n8 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA rrr-*-^! \* ' * ; ' .\*.. J , * -^ 1 " o . o oz. f rr -p- x FIG. 19. Tylococcus chrysocomae Brain THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 119 length but robust ; a couple of stout spines similar to those of marginal cerarii occur on median area of thoracic and abdominal segments. Anterior dorsal ostioles not recognizable ; posterior ones very poorly marked. Circulus absent. Legs well developed, without translucent pores on hind pair ; claw with a small denticle ; ungual and tarsal digitules short and finely pointed. Antennae normally 7- jointed ; but in one specimen one antenna had seven joints, the other eight. SUMMARY The author deals with the identity of the Pseudococcidae described from South Africa by C. K. Brain. Twenty-two species are retained as valid and are redescribed or reviewed. Four species and one variety are synonymized in the course of the paper. ACKNOWLEDGMENTS We wish to convey our sincere thanks to Dr. T. J. Naude, Chief Entomologist, Department of Agriculture, Pretoria, South Africa, for his permission to carry out the examination of all Brain's typical material stored in the Division of Entomology ; to Dr. H. Morrison, Senior Entomologist, U.S. Department of Agriculture, Washing- ton, D.C., who made available slides of four species apparently missing in Pretoria. Sincere thanks are also due to Dr. W. J. Hall, Director, and to Dr. D. J. Williams, Entomologist, Commonwealth Institute of Entomology, London, for their invaluable help connected with some bibliographical references. Our sincere thanks are particularly extended to Dr. H. K. Munro, Department of Agriculture, Pretoria, for his kindness and assistance given during our visit to South Africa and in whose laboratory the main part of the present work was carried out. We are greatly indebted to Dr. R. H. Le Pelley, Senior Entomologist, Department of Agriculture, Kenya, who as usual undertook the task of going through the manu- script, suggesting valuable amendments. REFERENCES BRAIN, C. K. 1912. Contribution to the knowledge of Mealy Bugs, genus Pseudococcus, in the vicinity of Cape Town, South Africa. Ann. ent. Soc. Amer. 5 : 177-189. 1915. The Coccidae of South Africa, I. Trans, roy. Soc. S. Afr. 5 : 65-194. 1918. The Coccidae of South Africa, II. Bull. ent. Res. 9 : 107-139. DE LOTTO, G. 1957. The Pseudococcidae (Homopt. : Coccoidea) described by H. C. James from East Africa. Bull. British Mus. (Nat. Hist.} Ent. 5 : 183-232. ESSIG, E. O. 1942. College Entomology, pp. vii -f- 900, McMillan Co., New York. EZZAT, Y. M. & MCCONNELL, H. S. 1956. A classification of the Mealybug Tribe Plano- coccini (Pseudococcidae, Homoptera). Bull, agric. exp. Stn., Univ. Maryland, A 84, pp. 108. FERRIS, G. F. 1918. The California species of Mealy Bugs. Stanf. Univ. Publ. pp. 78. 1948-50. Atlas of the Scale Insects of North America, vols. v and vi, Stanford Univ. Press. GREEN, E. E. 1915. Observations on British Coccidae in 1914 with descriptions of new species. Ent. mon. Mag. 51 : 175-185. 120 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA HALL, W. J. 1937. Observations on the Coccidae of Southern Rhodesia. VIII. Trans. R. ent. Soc. Lond. 86 : 119-134. HAMBLETON, E. J. 1946. Studies of hypogeic Mealybugs. Rev. Ent. 17 : 1-77. JAMES, H. C. 1933. Taxonomic notes on the Coffee Mealybugs of Kenya Colony. Bull. ent. Res. 24 : 429-436. 1936. New Mealybugs from East Africa. Trans. R. ent. Soc. Lond. 85 : 197-216. JOUBERT, C. J. 1928. Pseudococcus gahani Green, in South Africa. Bull. ent. Res. 29 : 209. MORRISON, H. 1945. The Mealybug Genus Heterococcus and some of its relatives (Homoptera : Coccoidea). /. Wash. A cad. Sci. 35 : 38-55. WILLIAMS, D. J. 1958. The Mealybugs (Pseudococcidae : Homoptera) described by W. M. Maskell, T. D. A. Cockerell, R. Newstead and E. E. Green from the Ethiopian Region. Bull. Brit. Mus. (Nat. Hist.) Ent. 6 : 205-236. ZIMMERMAN, E. C. 1958. Insects of Hawaii, vol. 5 : Homoptera Sternorhyncha, pp. 464, Honolulu, Univ. Hawaii Press. REVISIONS OF MALLOPHAGA GENERA. DEGEERIELLA FROM THE FALCONIFORMES THERESA CLAY BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. 4 LONDON: 1958 REVISIONS OF MALLOPHAGA GENERA. DEGEERIELLA FROM THE FALCONIFORMES BY THERESA CLAY Xv British Museum (Natural History) Pp. 121-207 ; 9 Plates ; 164 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7, No. 4 LONDON: 1958 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in jive series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 7, No. 4 of the Entomological series. Trustees of the British Museum, 1958 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued September, 1958 Price Thirty Shillings REVISIONS OF MALLOPHAGA GENERA. DEGEERIELLA FROM THE FALCONIFORMES By THERESA CLAY INTRODUCTION BEFORE attempting to define the genus Degeeriella, the type species of which para- sitizes one of the Falconiformes, it is necessary to consider shortly the whole of the Degeeriella-complex. THE Degeeriella-coMPLEX. It is difficult to delimit this group exactly but the following genera should probably be included : Degeeriella Neumann ( = Kelerinirmus Eichler), Acutifrons Guimaraes, Austrophilopterus Ewing, Capraiella Conci, Cotin- gacola Carriker, Cuculicola Clay & Meinertzhagen, Lagopecus Waterston (= Colini- cola Carriker), Picicola Clay & Meinertzhagen (= Tyrannicola Carriker), Trogoni- nirmus Eichler, Upupicola Clay & Meinertzhagen, a group of undescribed species from the Bucerotidae, and an undescribed species from the Megapodidae, probably an aberrant Lagopecus. Buceronirmus Hopkins and Hopkinsiella Clay & Meinertz- hagen should also perhaps be included here. Possible derivatives from this group include Syrrhaptoecus Waterston, Tinamotaecola Carriker, some of the Ischnocera from the Bucerotidae and also possibly Penenirmus. The complex (omitting the doubtful members) can be defined as follows : Ischnocera with marginal carina of head usually complete dorsally but may be partially interrupted anteriorly, and also partially interrupted each side when a dorsal preantennal suture is present ; ventrally it may be complete or interrupted medially. Hyaline margin absent or small, never greatly enlarged and never con- tinuous with hyaline area delimiting a complete dorsal anterior plate. Ventral carina never forms a semicircular band, but is interrupted medially ; usually the two carinae pass towards the anterior margin of the head but never form well defined bands continuous with the marginal carina, and only rarely have the strongly sclerotized parallel surfaces to which are attached lobes of the pulvinus as in Bruelia (Clay, 1951) ; pulvinus usually in the form of a single sac-like structure. 1 Ocular seta (except in Trogoninirmus and Austrophilopterus) and at least two of the temporal 1 At one time it was thought possible to use the characters of the ventral carinae and pulvinus to separate the Degeeriella- and Bruelia-complex.es, however the species of Degeeriella from Chelictina and Elanoides have the characters of these structures as in some species of Bruelia (see Clay, 1958). ENTOM. 7, 4. 5 124 REVISIONS OF MALLOPHAGA GENERA setae elongated. Prothorax with one marginal seta each side (except in Lagopoecus meinertzhageni Clay) ; third episternum fully sclerotized laterally. Abdomen with postspiracular setae on segments III-VII 1 (exceptionally on IV-V only) with sensillae on III or IV-V. Tergal plates entire or separated medially ; sternal plates median, lateral or absent. In the male segments IX-X with a single fused tergal plate (entire or divided medially), separated from XI by a suture and setae ; anal and genital openings close together on dorsal surface, dorsal part of XI narrowed with the 3+3 anal setae on the dorsal surface of the abdomen (see Clay, 1953). Male genitalia usually with sclerotized penis, short curved parameres, the outer and inner edges of which are continuous with the basal apodeme, and endomeral plate of characteristic form (PI. 8, fig. 7). This basic form which is found in some species of most of the genera is also found modified to a greater or lesser extent in a few species belonging to many of the genera and in some species the basic degeerielline pattern can no longer be recognized, for instance, there may be an articulation between the parameres and the basal apodeme. The genitalia do not provide good group characters in this complex ; these structures in Acutifrons megalopterus Carriker and Degeeriella rufa (Burmeister) for instance, being more similar to each other than are those of A. megalopterus and A. caracarensis (Kellogg & Mann) ; and those of Capraiella subcuspidata (Burmeister) are nearer those of D. fulva (Giebel) than are those of D. fulva and D. mookerjeei Clay. The internal male genitalia are too various, even within one related group (see below, p. 127) to be used as diagnostic characters for the complex, but a general type similar to that of D. fulva from Buteo (Text-fig, i) with or without the lateral lobes is found in some of the species of many of the genera ; all the species examined with one exception (a Picicola from one of the Tyrannidae) have the ductus ejacu- latorius long and coiled. An examination has been made of the internal male genitalia of about 150 species belonging to 73 genera of the Ischnocera ; it was hoped that the characters of these organs might help in the generic or suprageneric classification of this superfamily. The members of the Gonides-complex (including only those found on the Galliformes and Columbidae) have the vesicular apparatus (see further below) formed of two simple lobes, not joined medially and the ductus ejaculatorius modified in some way, they differ in these characters from Austro- gonoides, Osculotes, Chelopistes and members of the Heptapsogaster-complex. The Otidoecus-complex (Otidoecus, Rhynonirmus and Cuclotogaster) have an unpaired diverticulum arising from the ductus ejaculatorius 2 . Apart from these two groups it has not been possible to find characters of generic or suprageneric importance, although they may be of specific or of species group value. Recently Blagovesht- chensky (1956) has published a most useful and extensive account with many figures of the internal genitalia of both Ischnocera and Amblycera. In the female the genital plate (when present) does not reach to the upper margin of the vulva (cf . Bruelia) ; genital region without lateral spine-like setae (cf . Rallicola) or clump of setae on tubercle-like area (cf. Bruelia). Inner genital sclerites and 1 As in previous publications roman numerals are used for the true segments, see below, p. 126. 2 The presence of this diverticulum and other characters make it certain that the " Lipeurus variabilis " in Strindberg, 1918 : 633 was in fact Cuclotogaster heterographus (Nitzsch). REVISIONS OF MALLOPHAGA GENERA 125 subvulval sclerites present (Text-figs. 96, 97). Spermatheca with sclerotized calyx and simple thin- walled sac. It has not been possible to find any characters separating the females of the Degeeriella-complex from those of the Otidoecus-complex (i.e. Cuclotogaster, Otidoecus and Rhynonirmus) except that in the latter the calyx of the spermatheca is never apparent and it has not been possible to find any sign of a spermatheca in dissected specimens (no sections examined). The males are quite distinct : in the Otidoecus- complex the genital opening is terminal or ventro-terminal, intertergital sclerites are present and the ductus ejaculatorius has an unpaired diverticulum not yet found elsewhere amongst the Ischnocera. The present distribution of the Degeeriella-complex suggests that an ancestral stock must have been present on birds at an early stage of their evolution and that the Mallophaga have diverged with their hosts. On some host groups there are more than one species group belonging to the Degeeriella-complex, these presumably having diverged from each other on the host group in question ; these species groups are either sympatric and probably restricted to different ecological niches on the host, or allopatric and restricted to different taxonomic divisions of the host group. The species belonging to one of these groups have large heads and rounded abdomens with the characters frequently found in this type : that is a dorsal preantennal suture, temporal carinae, pleural thickening less well developed, and the tergites and sternites narrowed or interrupted medially ; the other is the more elongate form as found in Degeeriella fulva (PL i, fig. i). There appears to have been a considerable amount of parallel evolution in the degeerielline stocks resulting in a superficial resemblance between the species groups on different host groups. For instance, an undescribed species of Lagopoecus from the Megapodidae, Acutifrons viemi Guimaraes from the Accipitridae and Cuculicola acutus (Rudow) from the Cuculidae all have large heads pointed anteriorly, preantennal dorsal sutures and partial or complete temporal carinae passing posteriorly from the preantennal nodus ; the genitalia are all of the typical degeerielline type or modifications of it. Again Cuculicola latirostris from Cuculus canorus resembles superficially such species of Degeeriella as D. rufa from Falco tinnunculus, while the Cuculicola species from Geococcyx resembles Acutifrons megalopterus Carriker from a hawk (Phalcoboenus) in the broad head and abdomen and the form of the preantennal suture, in both genitalia are of the degeerielline type. In all these cases the species have retained the basic form of the abdominal tergites and sometimes the sternites : the species from the Galliform.es have the divided tergal and sternal plates, those from the Falconiformes have the entire tergal and sternal plates, while those from the Cucu- lidae have the anterior tergal plates at least, divided. The stability of certain characters and the divergence of the ancestral degeerielline stock on the various host groups together with parallel evolution makes it impossible to define a subfamily for the Degeeriella-complex, and further causes great difficulty in generic separation. It is possible with further study based on more material that some of the genera now recognized will have to be re-incorporated in Degeeriella. Degeeriella as found on the Falconiformes is here defined in detail and the charac- 126 REVISIONS OF MALLOPHAGA GENERA ters found throughout the genus will not be repeated in the descriptions of the indi- vidual species which follow. DEGEERIELLA Neumann, 1906 Nirmus. Nitzsch, 1818, Germar's Mag. Ent. 3 : 291 (nee Hermann, 1804). Degeeriella. Neumann, 1906, Bull. Soc. zool. Fr. 20 : 60. Nomen novum for Nirmus Nitzsch nee Hermann. Type species by subsequent designation, Johnston & Harrison, 1911, Proc. Linn. Soc. N.S.W. 36 : 326 : " D. discocephalus N." Kelerinirmus. Eichler, 1940, Zool. Am. 130 : 101. Type species : " Nirmus fuscus Nitzsch in Denny." Ischnocera not exceeding 3 mm. in length ; usually without marked sexual dimorphism, but the females average larger. Usually well pigmented species, the colour pattern sometimes forming a taxonomic character. Shape of head various, anterior margin varies from pointed (D. meinertzhageni) , flattened (D. fulva) or rounded (D. leucopleura, D. discocephalus). Marginal carina entire dorsally ; ven- trally may be interrupted medially to a greater or lesser extent ; hyaline margin may be apparent as a narrow rim round the anterior margin of the head. Dorsal preantennal suture and a true dorsal anterior plate never present in adult ; the dorsal preantennal region may have thickened areas or surface sculpturing ; dorsal post- antennal sutures rarely present (D. punctifer). Ventral carina never forms a complete semicircular band but is interrupted medially and the two carinae pass anteriorly ; at the aijterior edge of the pulvinus they merge with the general sclerotization of the head and a ventral suture (the ventral preantennal suture) is carried forward to or near the anterior margin of the head. Pulvinus usually appears as a simple lobe, but in a few species (e.g. D. guimaraesi) each ventral carina has a sclerotized flattened part parallel to that of the other carina to which is attached a lobe of the pulvinus (see Clay, 1958). Temporal carinae absent. Mandibles similar throughout the genus ; hypopharyngeal sclerites and gular plate well developed. Male antenna usually similar to that of female, but may show marked sexual dimorphism (D. mookerjeei). Chaetotaxy of the head of the basic ischnoceran type (Clay, 1951) ; ocular seta and at least two of the marginal temporal setae each side elongated. Prothorax similar throughout the group with rounded or parallel lateral margins and straight posterior margin ; one posterolateral or posterior elongated seta each side. Pterothorax may or may not show lateral indication of meso-metathoracic junction ; third episternum fully sclerotized laterally. Sternal plate narrowed anteriorly, normally with three setae each side. Dorsal pterothoracic setae usually comprise two lateral setae, one elongated and one spine-like, and four elongated setae each side of the posterior margin arranged in two pairs ; some species (D. discocephalus) may have a greater and more irregular number. Abdomen with nine apparent segments in the male and eight in the female ; these are interpreted as follows : the first apparent segment, probably I and II fused, is referred to as II, the second to the seventh (the spiracle bearing segments) as III-VIII. In the male the eighth segment represents IX-X fused, the ninth is XI ; in the female the last apparent segment is IX-XI fused. Segment II is always shorter than III. The tergal plates of II-X in the male and II-XI in the female are in the form of single plates across each segment ; tergites II-III may show partial REVISIONS OF MALLOPHAGA GENERA 127 division into two sclerites. The tergite of fused IX-X in the male is usually arched and narrowed medially to a greater or lesser extent and tergal plate XI when present is a single or double sclerite immediately anterior to the anal and genital openings. Keler (1939) has been followed in considering the dorsal plates as representing the fused tergal and pleural plates. At the lateral edge of these plates of some or all of segments II-VIII there is, in most species of Degeeriella, a characteristic internal thickening. This thickening, here called the pleural thickening, usually consists of an internal sclerotized buttress along the edge of each segment which is continued inwards a short way along the inner anterior margin of the dorsal plate ; there is usually a characteristic anterior part passing into the segment above, known as the re-entrant head (Waterston, 1928). Sternal plates II-VI in the form of median sclerites in both sexes ; in the male the terminal sternites form a single fused genital plate. Anal and genital openings of male on dorsal surface of the abdomen with the 3 + 3 anal setae as described above under the definition of the Degeeriella- complex. The genital region of the female comprises the genital plate (i.e. sternal plate VII) usually not differing greatly from the anterior plates, but sometimes (D. rufa) with a median posterior prolongation. It is not possible to be certain to which segments the remaining sclerites of the genital region belong. Below the genital plate is an uncoloured area of the integument with a sclerite each side, perhaps those of VIII. The integument passes to the vulva and turns in to form the ventral wall of the genital chamber. On this wall are two sclerites, sometimes fused to a greater or lesser extent in the mid-line ; these are perhaps the median sclerites of VIII and are here called the inner genital sclerites (Text-fig. 96, ig.). On the dorsal wall of the genital chamber there is a sclerite each side which projects beyond the vulva ; this is perhaps the sclerite of X or IX and X fused and is here called the subvulval sclerite (Text-fig. 97, sv). The opening of the spermathecal tube (os.) lies between the subvulval sclerites in the dorsal wall of the genital chamber. The spermatheca is a simple thin-walled sac and the calyx is lightly sclerotized. The external male genitalia (see Clay, 1956) comprise a flattened basal apodeme ; short curved parameres, the outer and inner edges of which are continuous with the basal apodeme without a point of articulation ; an endomeral plate, rather thick dorsoventrally, with diverging dorsal arms (Text-fig. 59, da.) which may or may not join the basal apodeme each side and two ventral arms (Text-fig. 52, va.) with setae. Centrally there is a sclerotized tube-like penis which usually has at its base an irregular area of sclerotization (shown in the figures by an interrupted line) joined to the basal apodeme by a narrow sclerite (the penial sclerite, PS.) ; at this junction there is usually a curved arm each side (the penial arm, pa.) bearing a seta (the penial seta, pst.). The dorsal and ventral endomeral arms are joined by an area passing ventro-dorsally (and not always visible) to a line of thickening each side of the ventral surface of the plate (Text-fig. 50, a.). Internal male genitalia have been examined from 40 specimens of Degeeriella from only 19 species of hosts belonging to the Falconiformes, but even these show considerable variation. In D. fulva from Buteo vulpinus and Buteo buteo these structures conform in general characters to those of Columbicola columbae (Linn.) as described by Schmutz (1955). The vesicular apparatus comprises four separate 128 REVISIONS OF MALLOPHAGA GENERA lobes united into a single organ ; as in all members of the Degeeriella-complex examined the two lateral lobes (Text-fig, i, la.) are shorter than the median lobes (me) ; in some species of Syrrhaptoecus however, the lateral lobes are considerably longer than the median ones. Within each median lobe two chambers can be dis- tinguished, the upper containing spermatozoa and the lower what is presumed to be a secretion. Each vas deferens (vd.) enters separately into each of the median lobes. The vesicular apparatus is continued into the ductus ejaculatorius (de.) which is strongly muscular near its base. The testes and vasa deferentia are similar throughout the complex, and do not differ significantly within any of the Philop- teridae examined. Variation of the vesicular apparatus and the ductus ejaculatorius and the point of entry of the vasa deferentia, within the species of Degeeriella examined are shown in Text-figs. 1-8. \ FIGS. 1-8. Internal male genitalia of Degeeriella. i. D. fulva from Buteo vulpinus. 2. D. beaufacies. 3. D. fusca from Circus aeruginosus. 4. D. elani from Elanus caeruleus vociferus. 5. D. rufa from Falco t. rupicolus. 6. D. r. regalis from Milvus migrans. 7. D. mookerjeei from Pernis ptilorhyncus gurneyi. 8. D. punctifer. me. median lobe of vesicular apparatus ; la. lateral lobe of vesicular apparatus ; vd. vas deferens; de. ductus ejaculatorius. Line ==0-5 mm. REVISIONS OF MALLOPHAGA GENERA 129 The length of the median lobes and the relative size of the lateral lobes may vary considerably in different species ; no lateral lobes could be seen in D. mookerjeei. In some cases the material was not in sufficiently good condition to distinguish the internal chambers of the median lobes, but there seems no doubt that in both D. regalis and D. mookerjeei there is only a single chamber, as spermatozoa could be seen filling the whole of the median lobes as figured for " Lipeurus variabilis " (? = Cuclotogaster heterographus, see footnote on p. 124) by Schmutz (1955 : 303). However, material suitable for sectioning is required before an accurate account can be given of the internal chambers of the median lobes. The ductus ejaculatorius is long and coiled in all species, and in all except D. rufa only a short basal portion is strongly muscular, in this latter species this muscular part is carried nearly to the end. The chaetotaxy of the abdomen has the following features common to all species ; the anterior tergal setae of segment II (probably those of the suppressed segment I) are two in number and elongated ; postspiracular setae are present on segments III-VII with sensillae on III-V ; VIII has the usual lateral seta in a sunken alveolus. All the above setae are omitted from the specific descriptions. Terga II-VIII and sterna II-VI each with a single line of setae ; pleural setae present on some or all of segments III-X. Vulva with some spine-like setae and with a varying number of sensilli ; posterior to the vulva there are, apart from the three anal setae, a single spine-like seta and one to three elongated setae (the pleural setae of X) each side. Nymphs. The three nymphal instars are easily separable by the chaetotaxy of the posterodorsal margin of the pterothorax (see Clay, 1955), as shown in Text-figs. 11-14 '> the third instar has the setae arranged as in the adult but at least two of the setae are thinner. The head does not take on the full adult characters until after the final moult. Boetticher & Eichler (1954) have shown the differences between the shape of the preantennal region of the head in nymphs and adults in Degeeriella and based some of their phylogenetic deductions on these findings. The present study of Degeeriella nymphs from 24 species of hawks shows that the curvature of the anterior margin of the head may be approximately the same in nymph and adult as in D. punctifer and D. discocephalus ; may be more rounded in the nymph as in D. fulva (Text-figs. 15-17) or more pointed as in D. rufa (Text-figs. 18-20). In D. rufa the anterior margin of the first instar (Text-fig. 18) resembles that of D. fulva to a greater extent than do those of the second or third instars (Text-figs. 19-20). The ventral carinae are sometimes better defined in the nymph than in the adult and in those of D. rufa (Text-fig. 10) there is a definite inner projection to which is attached a lobe of the pulvinus as in the nymphs and adults of D. guimardesi ; thus D. rufa resembles this latter species to a greater extent in the nymph than in the adult (see Clay, 1958). The second and third instars of D. rufa have a semicircular anterior dorsal thickening and a preantennal dorsal suture with a partial lateral break in the marginal carina each side (Text-fig. 10) ; these characters are not visible in the adults. In some species e.g. D. nisus frater both nymphs and adults have a similar dorsal anterior thickening. There may be considerable differences between the nymphs of two species : thus, although D. rufa and D. fulva are super- 1 3 o REVISIONS OF MALLOPHAGA GENERA ficially rather similar the nymphs of each are markedly different (Text-figs. 9, 10). These differences are also reflected in the adults in the characters of the male genitalia, female genital region and chaetotaxy of the abdomen. D. discocephalus and D. fulva superficially distinct have rather similar nymphs. The greater similarity of the head of rufa in the nymphs than in the adult to that of the adults and nymphs of 11 FIGS. 9-14. 9-10. Heads of third instar nymphs. 9. D. fulva from Buteo jamaicensis. 10. D. rufa from Falco tinnunculus. 11-14. Posterior margin of pterothorax of D. fulva from Buteo buteo. n. First instar. 12. Second instar. 13. Third instar. 14. Adult male. D. guimaraesi is also reflected in the adults of rufa which have other characters in common with guimaraesi not found elsewhere amongst the Degeeriella. Populations within a species may also differ from each other to a greater extent in the nymphal than in the adult stage : the third instars of D. rufa, for instance, from Falco rusticolus candicans and F. r. islandus (Text-figs. 21, 22) are more different than are the adults, which in some specimens are hardly separable (figs. 143, 147). This suggests that the superficial similarity of the majority of the species of Degeeriella on the Falconi- formes is a secondary adaptation to the environment found on this group of birds REVISIONS OF MALLOPHAGA GENERA 131 and that the characters of the nymphs may be useful in the elucidation of relation- ships within the genus. Some of the difficulties of understanding these relationships are mentioned below under Host Relationships. Apart from the species of the Degeeriella-complex found on the Falconiformes there are two other groups which have been given generic status, but fall within the definition of Degeeriella as given above, these are Capraiella and part of Picicola. Capraiella Conci, 1941. This genus was erected for Nirmus subcuspidatus Bur- meister from Coracias garrulus mainly on the character of the pointed head. As will be seen below some Degeeriella from the hawks also have heads pointed anteriorly. It has not been possible to find any characters on which subcuspidata can be separated from Degeeriella, in fact the male genitalia considered alone would place this species near D. fulva. It is doubtful, therefore, whether Capraiella can be kept as a separate genus but further species may be found on other members of Coraciidae which may throw more light on the relationships of this group. Picicola Clay & Meinertzhagen, 1938, and the subgenus Tyrannicola Carriker, 1956. This genus contains species found on the Pici and the Passeriformes, some of which can be included in the definitions of Degeeriella as given below. The species may lack the preantennal suture and have the tergites entire as in Degeeriella sens. sir., or may have a preantennal suture and divided tergal plates as in Cuculicola. The genitalia may be of the type found in D. fulva or a modification of this. These differences cut across the host divisions ; for instance, the species from Geocolaptes and Thripias belonging to the Pici and those from Colonia and Sayornis belonging to the Passeriformes have the tergites entire ; in the two former species the male genitalia are of the D. fulva type. The species from Dendrocopus (Pici) and Pitta (Passeriformes) have the tergites divided ; the latter species has the genitalia of the D. fulva type, the former the modified form. The species found on the Pici are in general less heavily sclerotized than those found on the Passeriformes and Falconi- formes. It is doubtful whether the erection of numerous subgenera is the best solution of this problem. Within the species of the Degeeriella-complex found on the Falconiformes three genera have been erected : Degeeriella Neumann, 1906 type species Nirmus disco- cephalus Burmeister ; Kelerinirmus Eichler, 1940, type species Nirmus fuscus Denny , and Acutifrons Guimaraes, 1942, type species A. vierai Guimaraes. Keleri- nirmus was described to include the species with elongate heads and abdomen and to separate them from the species with round heads and abdomens represented by D. discocephalus, the type species of Degeeriella. This division, however, appears to be a purely artificial one and places together D. discocephalus and punctifer purely on shape of head and abdomen together with certain characters directly correlated with this shape and of little phylogenetic importance (Clay, 1951). In fact, the characters of the carinae and sutures of the head, the male genitalia and female genital region show that these two species are not closely related. It appears that the discocephalus group and fulva group of species, both found on the same host groups, are nearly related to each other and perhaps derived from a common ancestor on these host groups. These two species groups are for instance, more closely related to each other than either is to rufa in spite of the superficial 132 REVISIONS OF MALLOPHAGA GENERA similarity of rufa and fulva. Thus, it is not possible to recognize Kelerinirmus as a generic division of Degeeriella. Carriker (1956 : iT-4) 1 suggests the possibility that Acutifrons should be included in the discocephalus group of Degeeriella, but here again the similarity is purely superficial and it is unlikely that the Acutifrons group of species are particularly nearly related to discocephalus. It is possible that Acuti- frons is not a monophyletic group, the characters distinguishing the species ; the anterior dorsal suture, the temporal carinae and the enlarged head and abdomen having been developed more than once in different but related stocks. Until more is known about the distribution of these species Acutifrons should probably be maintained as a distinct genus. Nirmus splendidus Kellogg, 1899. While agreeing with Carriker (1956 : 126) that the identity of this species must await the examination of the types, the des- cription and figure are those of a Cuculicola not Acutifrons. No known species belonging to the Degeeriella-complex from hawks have the abdominal tergites divided medially, a character which is found throughout Cuculicola, further, the figure, except for the lateral margins of the temples represents the species found on Geococcyx calif orniensis. Since writing this, Carriker (1957) has been seen in which a figure is given of the male genitalia of a paratype of Nirmus splendidus, this seems to represent those of the species from Geococcyx californiensis in a somewhat com- pressed condition, as usually seen within the specimen. There seems little doubt that this is the true host of Nirmus splendidus Kellogg as figured originally by Kellogg (1899) and recently by Carriker (1957), and that the species should be included in Cuculicola. THE SPECIES, SUBSPECIES AND LOCAL POPULATION. In Degeeriella there is the difficulty, as always in the case of a widely-distributed homogenous group, of deciding whether any given form should be considered as a species or subspecies or whether some merit taxonomic rank at all. As Mayr (1951 : 93) has said, the subspecies is primarily a taxonomic concept which cannot be delimited from the local population on one hand and the species on the other. In the Mallophaga the application of the subspecific concept has been most haphazard and practically no attention has been paid to the amount of variability within populations from the same host form, and it seems that the time has come to consider this problem as a whole and to try to get some conformity within the suborder. In the distribution of the Mallophaga it is usual for an order or suborder of birds to be parasitized throughout by the same genus (or genera) of Mallophaga. The populations 2 of this genus on the different species of birds may be apparently indis- tinguishable, only statistically distinguishable, or may comprise individuals which are slightly but constantly different, or which are markedly different. The present 1 I should like to draw attention to a misquotation in this paper ; on p. 114 it is stated that I use the shape of the abdomen as the principle generic character of Oxylipeurus ; I have never considered shape of either abdomen or head as of any phylogenetic importance and as the whole of the passage to which Mr. Carriker refers was an attempt to demonstrate the dangers of using shape as a generic character in the Mallophaga it is apparent that Mr. Carriker has misunderstood what I was attempting to say, as is also shown by his remarks on p. 115, paragraph three. 2 The word population is here used for all the individuals of a taxa of Mallophaga found on one host form which can potentially interbreed because their hosts are potentially capable of interbreeding. Thus, all the Degeeriella from Buteo b. buteo throughout its range would be considered as comprising one population. REVISIONS OF MALLOPHAGA GENERA 133 distribution and relationships of the mallophagan genera suggest that these allopatric populations have, in general, been separated from each other by the splitting and species formation of the host stock and are thus analogous to populations of free- living animals on a group of continental islands which have been isolated by the disappearance of land connections (Clay, 1949). As in the case of such populations of free-living animals each of the mallophagan populations is an isolated unit without zones of contact with any other populations. Thus, as with all isolated allopatric FIGS. 15-25. Heads of nymphs. 15-17. D. fulva from Buteo buteo. 15. First instar. 1 6. Second instar. 17. Third instar. 18-20. D. rufa from Falco tinnunculus. 18. First instar. 19. Second instar. 20. Third instar. 21. Third instar D. rufa from Falco rusticolus islandus. 22. Third instar D. rufa from Falco rusticolus candicans. 23. Third instar D. beaufacies. 24. Third instar D. n. nisus frim Accipiter nisus. 25. Third instar D. n. f rater from Accipiter badius, Thailand. populations where there is no evidence available on the degree of reproductive isolation, there are no criteria for separating the polytypic species from the super- species except morphological ones. As these populations are obviously allopatric replacements of each other on the different host group it might be possible in many cases to consider them as belonging to one polytypic species. But this is to ignore the morphological evidence and obscures the fact that while some show marked morphological differences others are hardly separable taxonomically. Further, some distinct populations may each have a number of related morphologically similar 134 REVISIONS OF MALLOPHAGA GENERA populations, making it more convenient to call each of these population groups, a species divided into a number of subspecies. It seems probable that there have been two tendencies in the evolution of the Mallophaga. One was to speciate rather rapidly, perhaps due to the original occu- pation of an empty ecological niche provided by the feathered bodies of birds, a changing environment due to the evolution of the birds themselves, together with the increasing isolation of the populations of the Mallophaga. The second tendency was a conservatism due to the later similarity of the environments afforded by birds belonging to one group, and to the close adaptation to a host which results from the parasitic habit and perhaps limits subsequent morphological change. The first tendency was probably responsible for the formation, in the Ischnocera, of the large number of genera and distinctive species groups, while the second has led to the similarity of the forms comprising these groups. The uniform environment and the necessity of being able to move easily through the feathers has probably been responsible for the relatively smooth uniform surface of the Mallophaga without the development of feathered setae, scales and other modifications of the exoskeleton which provide useful taxonomic characters in many groups of insects. It is rather frequent in the genera of the Ischnocera to find a series of populations superficially similar and differing mainly in the characters of the male genitalia, the uniformity of the environment having led to superficial similarity and the isolation of the popu- lations being shown in differences in such non-adaptive characters as the male genitalia. It must be expected in the Mallophaga that the character differences between related groups will be small, and these must of course be judged for each group of related species ; similar character differences cannot of course be used in separating analogous taxa in the Ischnocera and Amblycera, for instance. The degree and time of isolation cannot be used to determine the specific or subspecific status of a population : the populations of Degeeriella (D. regalis regalis) on Milvus and on Buteo galapagoensis are separated by host and geographical distri- bution, and although there cannot have been any gene flow between these populations over great periods of time, and although the gene pools must now be distinct, there is no clear cut morphological difference between these populations. If distribution is taken into account there is a further difficulty that the exact relationship between the hosts is not always known, so that on the analogy of the free living allopatric populations the exact position of the locality of any one population in relation to another is uncertain and deductions of which are the most nearly related populations cannot always be made. Thus, although some groups may show gradients in such characters as the size and shape of head and number of abdominal setae these cannot be equated with clinal variation in free-living populations, as the populations are isolated and they have a host (not geographical) distribution, the most similar forms not necessarily being most nearly related. For instance, in Degeeriella there are examples of Harrison's law that in related populations those parasitizing the larger hosts tend to comprise larger individuals ; correlated with this increase in size there is a tendency towards larger heads, broader anteriorly, and sometimes to a greater number of abdominal setae. This tendency is seen in some of the popu- lations of D. rufa on Falco, D. nisus on Accipiter and D. fulva on Buteo. Thus, REVISIONS OF MALLOPHAGA GENERA 135 sometimes the similarity of characters is partly due to ecological factors and not relationship (although in some cases of course the former may be dependent on the latter). Some of the subspecies in the Mallophaga differ from each other in only one character or in two or three correlated characters perhaps associated with size differences which are themselves dependent on host size. Thus, there may be populations, not very closely related, which are indistinguishable from each other and must be included in the same subspecies (Mayr, Linsley, Usinger, 1953 : 32) ; these are analogous to the polytopic subspecies of the free-living animal. It has been suggested (Mayr et al., 1953 : 104) that the morphological differences between sympatric species of the same genus might give an indication of the correct status of isolated populations, but Brown & Wilson (1956 : 49) have shown that when two species of animals overlap geographically the difference between them is accent- uated in the zone of sympatry and weakened or lost entirely in parts of their range outside this zone. This might explain the differences between the species of Degeeriella (a genus in which sympatry is rare) on Pernis, which are so much greater than is usual between species found on hawks belonging to the same genus. The two species, Pernis apivorus and P. ptilorhynchus might originally have had the same species of Degeeriella, the populations of which split into two and diverged sufficiently to remain distinct when they later became sympatric (see Clay, 1949) ; if the fact that that they had become sympatric caused them to diverge to a greater extent and if D. phlyctopygus became extinct on Pernis ptilorhynchus and D. mookerjeei on P. apivorus, the differences between these two species of Degeeriella would be more marked than if they had not formerly been sympatric. This explanation is partly supported by the fact that these two species are separated by the characters of the male antennae, a common difference between sympatric species of the same genus (Clay, 1949). It is perhaps for this reason that differences between allopatric species are sometimes much smaller than those distinguishing sympatric species. If we accept the definition of subspecies as populations which would interbreed under natural conditions if they occurred sympatrically, then any morphological differences which might prevent interbreeding should be considered as specific characters. It seems reasonable to suppose that at least some of the character differences between closely related sympatric species are those which prevent or discourage cross-breeding. In the Ischnocera closely related sympatric species may be distinguished by the male genitalia, male antennae and in one genus (Osculotes) the legs of the male, and in size and proportions of the head and abdomen. The former characters would probably all prevent or discourage cross-breeding, while the last two characters might mean that the populations were partly restricted to different ecological niches on the body of the bird resulting in partial isolation from each other. An example of this in the Anoplura is provided by Pediculus humanus humanus and P. h. capitis, whose occupation of different ecological niches on man has resulted in impaired fertility when they are crossed (Hopkins, 1949 : 419). Even gross differences in the form of the male genitalia in insects may not form a mechanical bar to successful copulation (Dobzhansky, 1955 : 189). Jordan (1896) in his analysis of the genitalia of Papilio showed that in general each of the species was distinguishable by the form of the male genitalia ; he also showed that there was geographical variation 136 REVISIONS OF MALLOPHAGA GENERA FIGS. 26-34. Endomeral plate, dorsal view. 26. D. fulva from Aquila chrysaetos. 27. D. carrikeri. 28. D. emersoni. 29. D. n. nisus from Accipiter n. nisus. 30. D.fusca from Circus aeruginosus. 31-32. D. n. frater from Accipiter badius, Thailand ; 2 specimens from the same host individual to show variation. 33. D. hopkinsi. 34. D. leucopleura. REVISIONS OF MALLOPHAGA GENERA 137 in the structure of the male genitalia and concluded that it was not possible to draw a distinction between specific and subspecific characters and that a peculiarity of a structure might be an individual aberration, a subspecific or a specific character. Jordan did, however, believe that divergence in the organs of copulation was a means of preventing intercrossing. Within the Mallophaga the genitalia may be uniform, with only minor or no apparent differences throughout genera, or large species-groups, examples of this are found mainly in the Amblycera (e.g. Colpo- cephalum and Actornithophilus) , and in some Ischnocera (e.g. Anaticola} ; in others there may be relatively small but constant differences in the population from nearly every host species, as in some groups of Quadraceps, and in other genera the differences may be so great that it is difficult to homologize the sclerites forming the genitalia of the different species. It must, therefore be presumed that the differentiation of the genitalia has taken place at different rates in different groups and that simi- larity of genitalia cannot always be used as a criterion of conspecificity, this is especially so in some genera of Amblycera. In Anaticola again, where the genitalia are similar throughout the genus it would seem to be necessary to use the characters of the preantennal region of the head for specific divisions. It is clear therefore, that the delimination of the specific and subspecific categories in the Mallophaga must be based on criteria which differ in each genus and that a study of the whole genus is necessary before a decision on these categories is made. Moreover, it is necessary to choose completely arbitrary criteria for the decision as to whether a population should have specific or subspecific rank, and this, in spite of some of the criticisms mentioned above, must be based on the characters of the genus as a whole, the number of character differences present, the characters separating sympatric species of the same genus and to a lesser degree host distribution. It is apparent that the male genitalia of the Degeeriella-complex (see above p. 124) are rather constant in character, those of Capraiella, for instance, being quite near ihefulva group of species, and that small differences in these structures may therefore be of significance in distinguishing species. A population has been considered as a full species if the individuals comprising it show one of the following qualifications : 1. Male genitalia quite distinct. 2. Male genitalia differ to a lesser extent, sometimes to a rather minor degree, but there are also a number of other character differences, such as the presence or absence of the pleural setae on certain of the abdominal segments, 4 or more setae on each of the sterna II I- VI, together with other morphological characters such as the form of the pleural thickening, marginal carina and ventral suture of the head, and marked differences in the shape of the nymphal heads. 3. Male genitalia apparently indistinguishable but the individuals differ in many of the other characters listed above. Populations are treated as subspecies when the male genitalia are apparently identical or only differ to a minor degree (e.g. number of setae as in D. rufa caruthi), which differ in the shape and proportions of the head and in a minor way in the breadth of the temporal marginal carinae and pleural thickenings or the shape of various sclerites. The second, and more controversial problem is that of deciding whether certain ENTOM. 7, 4. 6 138 REVISIONS OF MALLOPHAGA GENERA populations should be recognized taxonomically at all. Various procedures have been adopted within the classification of the Mallophaga, one is to describe as new every population occurring on a different host species, in the hope, it is presumed, that 50% or more may prove to be valid and leaving some other worker to find out. Another method is to take single specimens and to describe them as new species or subspecies on some minor character which is likely to be variable within the popu- lation or even an artefact due to method of preservation and of no taxonomic signi- ficance. These two examples of bad taxonomic procedure, unfortunately still rather frequent in the systematics of the Mallophaga, need not be further considered. The difficult cases are those where there are differences and where the populations must be genetically quite distinct, but it is considered unsatisfactory to recognize them taxonomically. In Degeeriella it is possible to distinguish three categories, apart from those where insufficient material is available, the taxonomic recognition of which it is considered would not assist in the classification of the group : 1. Certain populations are only separable from each other statistically ; here no useful purpose would be served in naming these microsubspecies, where many individuals would not be identifiable. 2. Certain populations may appear to differ, but when large series are examined too much variation is found to support the divisions, an example of this are the popu- lations from Aquila and the various species of Buteo (see p. 146). 3. The populations from two hosts may be distinctly separable on such characters as size and shape of head, but if between these two there is a series of populations from related hosts showing a character gradient in these characters it is not considered that any of the populations should be recognized. The populations of D. rufa from Falco provide a particularly difficult problem (see further p. 183) as they differ from each other in the outline of the anterior margin of the head, showing all degrees from marked differences to very slight ones, the latter being complicated by individual variation. If any subspecies are recognized then the classification of the populations showing minor differences becomes almost impossible ; this may be a case where subspecies should not be recognized. This policy of not overwhelming the classification by naming every statistically or barely separable population is not to discourage the study of populations, their variation and host distribution ; there is much interesting information on these subjects to be gathered from detailed statistical analysis, all that is here recommended is that these populations should not be given names. In this present paper an attempt has been made to sort out the populations deserving taxonomic rank and not to study detailed variation. VARIATION AND ARTEFACTS. The populations of Degeeriella from the various hawks are frequently very similar to each other and forms are separated on rather slight character differences, for this reason it is necessary to consider the amount of variation within populations from one host form. Further, it is necessary to work with specimens which have been treated in various ways so that they can be examined under high-power microscopes and this may cause various artefacts dependent on the methods used. Individuals in populations of Mallophaga, unlike some other groups of insects, tend to show little variation in size and external characters, due no doubt REVISIONS OF MALLOPHAGA GENERA 139 to the similarity of the environment in all stages of development and to the unlimited and easily accessible food supply. The reliability of the morphological characters which have been used in the taxonomy of the group are discussed below. i. Size. As already mentioned populations of Mallophaga tend to be rather constant in size. A number of experiments were carried out to see which was the most reliable measurement in Degeeriella. A male and female of each of D. fuva, D. r. regalis and D. rufa were measured at each stage of the following treatments : 40 FIGS. 35-41. Endomeral plate, dorsal view. 35. D. d. discocephalus. 36. D. elani. 37. D. tendeiroi. 38. D. rufa from Falco tinnunculus. 39. D. r. regalis from Milvus milvus. 40. D. r. deignani. 41. D. punctifer. (a) In 80% alcohol after two to three years storage ; (b) in a saturated solution of phenol in 70% alcohol, warmed to clear ; (c) after 22 hours in cold 10% caustic potash, body contents removed, cleared in clove oil and mounted in Canada balsam ; (d) after immersion for 15 minutes in 10% caustic potash in a boiling water bath, mounted in canada balsam and the cover glass pressed well down. It was found the 140 breadth of the temples remained either constant under the different treatments or changed no more than 0^004 mm., while other measurements especially total length, breadth of pterothorax and length and breadth of abdomen were rather variable due to contraction or expansion between the different regions of the body or changes in shape due to pressure by the cover slip. This means that in Degeeriella the breadth of the head is a measurement that can be taken quickly and accurately in any media, does not alter with the age of the adult, the abdomen for example in teneral females is usually smaller (Clay, 1956) and it is possible to compare the measurements of other workers as there is no ambiguity about the exact position of where the measurement is taken. The whole of the collected populations from one bird can be measured in phenol solution without the labour of mounting them in Canada balsam and ensures that not only the larger specimens, which consciously or unconsciously are likely to be picked out for permanent mounts, are measured. This measurement, therefore, is useful in comparing populations from various hosts which differ only in absolute size ; the size of the temple breadth being roughly proportional to the total size. It cannot of course be used in comparisons of populations which differ in the shape of the postantennal region of the head. It should be noted that this is a different problem from the consideration of which measurements show the least variation in a population from one host where all individuals have been treated in the same way. Tj0nneland (1955) compiled the variation coefficients for various measurements of 40 males and 40 females of Degeeriella d. aquilarum Eichler taken from the same host individual and subjected to the same treatment ; it was found that the measurement of the head showed the least variation within the populations. It has been found in numbers of specimens belonging to one species that those from the different hosts may differ in average measurements of head breadth, but that the ranges overlap ; it is important therefore to give the range and number of specimens measured. For reasons discussed elsewhere (p. 138) time has not been spent on statistical analysis of the measurements of the different populations of Degeeriella mentioned in this paper. It is doubtful whether subspecies should be recognized on size differences alone and certainly not when only two or three specimens are available. 2. Pigment and sclerolization. As it is frequently necessary to work with material which has been kept long in alcohol or over treated with caustic potash it is difficult to use the characters in the comparison of all species. However, the amount and arrangements of pigment may be a taxonomic character (see D.fusca). The scleroti- zed plates may vary in outline as some of these, especially the thoracic sternal plates and the male and female genital plates, may have part of the plate more lightly sclerotized and in some specimens, either naturally or due to treatment, the lighter part is not apparent, thus altering the outline. 3. Head. The shape and size of the head is a useful character and as shown above the breadth is not affected by the various methods of preparation. However, in some populations there appears to be a certain amount of variation in the curvature of the anterior margin (see under D. fulva). The thickness of the marginal carina and the presence or absence of an inner median indentation are usually reliable characters, but there may be intrapopulation variation in these characters, sometimes due to methods of preparation ; teneral specimens especially may be misleading in REVISIONS OF MALLOPHAGA GENERA 141 these characters (see below under D. pseudophaea] . Thus, specimens of the fulva- group from the same host form may have the typical flattened anterior margin, with broad marginal carina and well marked inner marginal indentation, whereas other specimens may have rather rounded anterior margins, rather narrow marginal carina and the inner marginal indentation not apparent. Specimens which have been left too long in caustic potash may have the marginal carina appearing narrower, the head often more rounded anteriorly and the anterior end of the ventral suture not distinguishable ; the extent of the hyaline margin may also vary in mounted specimens according to the position and pressure exerted. The proportions of the head, that is ratio of length of preantennal region to postantennal region and ratio of breadth to length of these regions may be misleading as these show variation within popu- lations. Reduction of these proportions to mathematical terms is unsatisfactory owing to the difficulty of finding exact points of measurement and a more accurate method is that described in Clay & Hopkins (1954 : 230) in which an outline of a head is drawn with a camera lucida and comparisons of other heads made by projecting them on to this outline ; by adjusting the magnification it is possible to get a fixed measurement such as the breadth of the temples and thus compare the proportions of the head (see also p. 184). If this is done with a large number of specimens from one host it will be seen that proportions are often variable and cannot be used for subspecific divisions. Both the marginal and temporal marginal carinae may have indentations, the number, shape and position of which show some individual varia- tion. However, the thickness and outline of these carinae, with the reservations discussed above, may be of taxonomic importance. The conus tends to be variable in shape and length mainly due to the position in mounting and except where the differences are strongly marked (e.g. D. punctifer], this structure has not been used as a taxonomic character. The position of the second ventral submarginal seta (Clay, 1951) shows individual variation being found either above or below the level of the inner margin of the marginal carina. 4. Thorax. The number of sternal setae and the shape of the sternal plate show individual variation (Text-figs. 112-118). and is of no taxonomic value amongst nearly related species. There are usually eight elongated posterodorsal setae on the pterothorax arranged in two groups of two each side, but there may be individual variation in the number and position. 5. Abdomen. In general the presence or absence of a partial division of tergal plate II-III is constant for a given taxa, but there are cases where this character especially in II shows individual variation. The width and the dorsal and ventral outline of the pleural thickening is often a useful character, but in mounted specimens is liable to distortion ; this distortion is particularly marked in the shape and details of the re-entrant head ; teneral specimens do not always show the normal characters of the adult pleural thickening. In the female the subvulval sclerites may show slight individual variation. 6. Male genitalia. The basal apodeme may show slight variation in outline either due to individual variation or to distortion in preparation. The shape of the tips of the parameres must again be used with caution as the appearance of these is dependent on position of mounting. The sclerotization round the penial sclerite and penial 142 REVISIONS OF MALLOPHAGA GENERA arms is irregular and rather variable in outline. The fusion or not of the dorsal endomeral arms with the basal apodeme may be a specific character or it may show individual variation and differ on the two sides of the same individual. There is also frequently considerable variation in the shape and length of the distal ends of 45 FIGS. 42-48. Male genitalia, ventral view of distal area. 42. D. fulva from Aquila chrysaetos. 43. D. beaufacies. 44. D. carrikeri. 45. D. emersoni. 46. D. n. nisus from Accipiter nisus. 47. D. n. frater from Accipiter badius, Kenya. 48. D. fusca from Circus aeruginosus. the ventral endomeral arms and the position of the setae, the two sides in one indi- vidual often being asymmetrical in these characters. It is not always possible to see the relationship of the dorsal and ventral parts of the mesosome to each other unless the genitalia are dissected and mounted separately on the slide. 7. Abdominal chaetotaxy. This frequently forms a useful taxonomic character, but it is important to consider the amount of individual variation. The presence or absence of pleural setae on some of the anterior segments and on X in the male can be used to separate species or species groups, but occasionally an individual will REVISIONS OF MALLOPHAGA GENERA 143 be found with one seta present on one side of a segment when its absence is characteri- stic of the species. Another character which can be used to separate species groups is whether the sterna of III-VI normally have 4 or more setae ; however, the species characterized by the presence of 4 setae may have the occasional specimen with one or two segments with 3 or 5 setae. The number of tergal setae may also be taxonomically important but here again there is individual variation and a more useful character is the range in the total number of setae found on segments III-VI I. The dorsal setae on segment X of the male may vary from 1-3 each side, but in some populations one each side seems to be the rule with occasional exceptions. Characters of taxonomic importance. The following characters have been found to be of taxonomic importance in Degeeriella and should be given together with their variation in all descriptions of new taxa : Shape of head : form of marginal and ventral carinae, and anterior extension of ventral suture ; thickness of temporal marginal carinae ; number of elongated marginal temporal setae ; presence or absence of postantennal sutures. Form of tergal plates of segments II-III, and XI in male and IX-XI in female ; width, and dorsal and ventral outline, and develop- ment of re-entrant head of pleural thickening. Outline of female genital plate, inner genital sclerites and subvulval sclerites. Presence and absence of pleural setae on segments II-VI and of X in male and numbers of sternal setae. The male genitalia should be figured to show the length of the penis and the form of the penial arms, and a dorsal and ventral view of the endomeral plate. SYSTEMATIC SURVEY OF THE SPECIES OF DEGEERIELLA PARASITIC ON THE FALCONIFORMES For convenience in classification and to avoid frequent repetition in descriptions, the species are divided into a number of species groups. There is naturally not always a clear cut distinction between the species groups and these may not always form natural phylogenetic assemblages. The groups are based mainly on the characters of the head, abdominal chaetotaxy and the male genitalia. Characters given under the definition of the genus (p. 126) and for the species groups are not usually repeated again in the descriptions of the species. The fulva Species Group 1. Head index less than 0-90. 2. Dorsal head sutures not apparent. 3. Two of the marginal temporal setae each side elongated. 4. Thoracic sternal plate and chaetotaxy as in Text-figs. 112-118 ; this plate shows individual variation in shape and in the number of associated setae. 5. Thorax and abdomen with general shape as in PI. i, fig. i. 6. Tergal plates of segment XI not apparent in male. 7. Terga of segments IX-XI in female as in Text-fig. 105. 8. Pleural thickening of segments III-VI usually with well developed re-entrant heads. 144 REVISIONS OF MALLOPHAGA GENERA 9. Sternites of II-VI in the form of quadrilateral median plates. 10. Male genital plate of irregular and variable outline. 11. Female genital plate without median posterior prolongation. 12. Female inner genital sclerites never fused in mid-line. 13. Male genitalia of type shown in PI. 8, fig. 3 ; penial sclerite present. 14. Setae each side of posterodorsal margin of pterothorax : i (lateral) spine- like seta, i elongated seta, 2 pairs of elongated setae (as Text-fig. 14). 15. Pleural setae absent on segments II-III and usually IV, and on segment X in male. 16. Sternocentral setae of segments III-IV normally 4. 17. Ventral chaetotaxy of male segments VII-XI as in Text-fig. 102 ; in some specimens one or both of the outer setae on segment VII may be absent. Degeeriellafulva (Giebel), 1874 Type host : Aquila chrysaetos (Linn.) (PI. i, figs. 1-7 ; PI. 8, fig. 3 ; Text-figs, i, 9, 11-17, 26, 42, 70, 84, 102, 105, 109, 112-118) Nirmus fuscus Nitzsch, 1861, nee Nirmus fuscus Denny, 1842. In Giebel, Z. ges. Nat Wiss. 17 : 525. Host : Buteo vulgaris = Buteo b. buteo (Linn.). Nirmus fulvus Giebel, 1874. Insecta epizoa : 124. Host : Aquila fulva = A. chrysaetos (Linn.). Nirmus angustus Giebel, 1874. Insecta epizoa : 126. Host : Buteo lagopus (Pontoppidan). Nirmus flavidus Giebel, 1874. Insecta epizoa : 301, Host : Buteo jaktal Buteo r. rufofuscus (J. R. Forster). Degeeriella giebeli Hopkins, 1947. Entomologist, 80 : 77. Host : Buteo b. buteo (Linn.). Degeeriella borealis Carriker, 1956. Florida Ent. 39 : 41, figs. Host : Buteo jamaicensis borealis (Gmelin). Degeeriella genitalis Carriker, 1956. Florida Ent. 39 : 43, figs. Host : Buteo regalis (G. R. Gray). The description, figure and host of D. fusca (Nitzsch) make it certain that this name, already preoccupied, is a synonym of D. fulva (see also Hopkins, 1947 : 76). The original description of D. fulva together with the fact that it was placed between fuscus and rufus, both figured, show that Giebel's original specimen must have been the elongated type (PL i, fig. i) of Degeeriella found on Aquila not the round-bodied type (PI. 9, fig. 2). As this species appears to be indistinguishable from that on Buteo there seemed a possibility that the known specimens might have been strag- glers from this latter genus. However, an examination of all the available material from Aquila, that is 18 <^, 36 $ from nine individuals of seven species of Aquila, shows there is no doubt that Aquila was the true host of at least three of these records; the hosts of the remainder cannot now be confirmed. It can be assumed, therefore, that the species described below is a natural parasite of Aquila. Degeeriella fulva is distinguishable from other species in the species group by a combination of the characters of the marginal carina, ventral suture, tergites II-III, pleural thickening, number of pleural setae and the details of the male genitalia. REVISIONS OF MALLOPHAGA GENERA 145 MALE. Inner dorsal margin of marginal carina indented medially ; ventral suture passes to anterior margin of head (Text-fig. 109, v.). Tergite II only with definite median unsclerotized indentation. Pleural thickening narrow with inner edges comparatively straight. Genital plate as in Text-fig. 102. Genitalia as in PI. 8, fig. 3 and Text-figs. 26, 42 ; there is some variation in the shape of the basal apodeme pst- FIGS. 49-53- Male genitalia, ventral view of distal area. 49. D. hopkinsi. 50. D. leuco- pleura. 51. D. d. discocephalus . 52. D. elbeli. 53. D. tendeiroi. ps. penial sclerite ; pa. penial arm ; pst. penial setae ; va. ventral endomeral arm. and of the base of the endomeral plate which does not always show an inner inden- tation. Internal genitalia as in Text-fig, i. FEMALE. Terga of segments IX-XI as in Text-fig. 105 and genital region as in Text-figs. 70, 84. CHAETOTAXY OF ABDOMEN. Tergocentral setae : II normally 6 1 range 4-7 ; III-V normally 8, range 6-8 ; VI-VIII normally 6, range 4-8. X in the male has 1 In this and all subsequent descriptions the two anterior setae always found on II are omitted. I 4 6 REVISIONS OF MALLOPHAGA GENERA from 1-3 setae each side ; of 16 males from Aquila n had one (i + i) each side, 3 had 1 + 2 and two had 2 + 2 ; in 80 males taken at random from various species of Buteo, 8 had i + i, 19 had i + 2, 44 had 2 + 2, 7 had 2 + 3 and 2 had 3 + 3. Tergal setae of segments X-XI of female as in Text-fig. 105. Pleural setae : II-IV o ; V, i on each side ; VI-VII, 2 ; VIII, 3. In the male IX has 2 each side and X, o. In the female IX and X each have 1-3 each side. Sternocentral setae of II-VI normally 4 with the occasional segment of the occasional specimen with 3 or 5. In the male total number of marginal setae of last segment dorsal and ventral, varies from 9-14. NYMPHS. No nymphs have been seen from any species of Aquila ; pterothoracic setae and heads of the three instars of specimens taken from Buteo are shown in Text-figs. 9, 11-13, i5-!7- VARIATION AND HOST DISTRIBUTION. The detailed comparison made by Tendeiro (1955 : 590) between specimens from Buteo buteo and Aquila chrysaetos has been studied closely, but the conclusions reached are different ; this is probably due to the availability of specimens from a greater number of species of Aquila and Buteo. Through the kindness of Dr Tendeiro it has been possible to examine three males and six females from Aquila chrysaetos ; these have been compared with 15 males and 30 females from six other species of Aquila and about 350 males and 400 females from 17 forms of Buteo. Certain characters were found to be too variable within the Head Prothorax Pterothorax Abdomen Total Genitalia* Head index Head Prothorax Pterothorax Abdomen Total Head index Length Breadth o-53 0-43 0-30 0-47 1-18 0-58 2 -06 0-81 0-60 0-47 0-30 0-50 0-78 Measurements in mm. Male B Length Breadth X >. f " \ t \ Range Mean Range Mean 0-50-0-58 (17) o*55 0-38-0-45 (17) '43 0-25-0-30 (12) 0-27 0-42-0-47 (12) '45 I -O2-I -25 (lo) 1-14 0-50-0-67 (10) 0-58 I-83-2-20 (ll) 2 -O2 0-34-0-38 ( 4 ) o-37 0-75-0-79 (17) 0-77 Female 0-58-0-62 (12) o-59 0-45-0-49 (12) 0-47 0-28-0-33 (10) 0-29 o-47- '53 (10) 0-50 1-13-1-43 (9) 1-17 0-60-0-70 (10) 0-65 2-03-2-35 (9) 2-23 0-77-0-81 (ll) 0-79 A. Single specimen from A. chrysaetos. B. Specimens from Buteo lagopus. * Length of genitalia of male taken from anterior margin of basal apodeme to posterior margin of endomeral plate. Number of specimens measured given in brackets. Head index breadth : length. REVISIONS OF MALLOPHAGA GENERA C Male Length Head (10) . Prothorax (7) Pterothorax (7) Abdomen (7) Total (7) . Genitalia (4) C.I. (10) . Head (9) . Prothorax (9) Pterothorax (8) Abdomen (7) Total (7) . C.I. (9) . Range Mean 0-50-0-55 o-53 Breadth A*. I -03-1 -22 i -80-2-10 ' 77"- ' 1-13 1-96 0-79 Female 8 0-57 Range 0-40-0-43 (12) 0-25-0 -28 0-40-0-47 0-50-0-60 1-27-1-38 2-15-2-30 0-76-0-80 1-32 2-22 0-78 o 44-0 46 O-27-O-28 o 460 49 0-58-0-62 C. Specimens from Aquila clangula and A. wahlbergi. D Male Mean 0-42 0-26 o-43 o-54 o-45 0-28 o-47 0-60 Head Prothorax Pterothorax Abdomen Total C.I. Length 0-50-0-58 i -oo-i -32 1-75-2-23 0-75-0-81 Breadth 0-38-0-47 0-23-0-30 0-38-0-48 0-48-0-67 Female Breadth 0-40-0-50 D. Maximum and minimum measurements of specimens from Buteo species. populations from one host species to be used for taxonomic divisions. These are : exact curvature of the anterior margin of the head and thickness of the marginal carinae, both these characters also seem to be affected by the method of treatment (see p. 140) ; outline of gular plate and thoracic sternal plates and the number of associated setae (Text-figs. 112-118) ; shape and extent of unsclerotized area of tergite II ; central narrowing of fused tergite IX-X in male ; outline of male genital plate ; exact outline of basal apodeme, differences in its total length and ratio of its length to that of the mesosome ; shape of penial arms ; ratio of height to breadth of female genital plate and exact outline of subvulval sclerites. PI. i, figs. 1-7, show the variation in the shape of the head of specimens from various hosts ; figs. 3-4 are specimens from the same host individual and mounted on the same slide. As already discussed above (p. 134) there is a tendency for the populations on larger i 4 8 REVISIONS OF MALLOPHAGA GENERA host species to have a greater number of larger individuals. This is true of the populations from the species of Buteo : measurements of head breadth of 321 males from 17 forms of Buteo and 396 females from 16 forms show a difference in the average breadth of the head between some of these populations. Thus, in males (53 speci- mens) from Buteo buteo (the smaller bird) the average is 0-41 mm., while in those FIGS. 54-57. Male genitalia, ventral view of distal area. 54. D. elani. 55. D. rufa from F. tinnunculus. 56. D. r. regalis from Milvus milvus. 57. D. punctifer. (41 specimens) from Buteo jamaicensis borealis (the larger bird) the average is 0*44 mm. Populations from other species of Buteo have intermediate averages and there is overlap in measurements of individuals of all populations. In addition to size some of the populations are composed of individuals in which the head tends to be more rounded anteriorly, such as that from Buteo jamaicensis (PL I, fig. 7), although even in this case there are individuals indistinguishable from those from other REVISIONS OF MALLOPHAGA GENERA 149 hosts. For these reasons it has not been found possible to recognize taxonomically the populations from the different species of Buteo and to separate these from the populations from Aquila ; this is also true of the population parasitic on Geranoaetus. There are considerable difficulties in placing the populations from the following hosts : Ichthyophaga, Polemaetus, Lophaetus, Hieraaetus, and Spilornis. Specimens from Ichthyophaga and Polemaetus can probably be included with fulva, but the avail- able material is not in sufficiently good condition for exact comparison. Those from Lophaetus and Hieraaetus are rather similar and have the anterior inner margin of the marginal carina sloping posterolaterally instead of being nearly parallel with the anterior margin as in typical fulva ; however in some specimens the difference is less marked and a similar condition is found in some specimens from Aquila wahlbergi. In the specimens from Hieraaetus the shape of the penial arms differ slightly from those of typical fulva, but in Lophaetus both types occur. The population from Melierax musicus poliopterus resembles that from Lophaetus in the characters of the margin carina, but specimens from some subspecies of Melierax metabates are intermediate between the latter and fulva. Specimens from Spilornis resemble the Lophaetus population but have a broader head anteriorly and may differ in colour pattern but the material is not in sufficiently good condition for identification. Nymphs are available from the Melierax metabates population only ; these resemble those from Buteo. Taking all these facts into consideration it does not seem that at the present time the classification will be simplified by giving subspecific names to all these poorly separable, perhaps inseparable, populations (see above p. 138) and these are, therefore, here kept for the present under the name fulva. The material available from Melierax is confusing : as shown above that from M. musicus poliopterus (5 $ from 3 individuals from Kenya) and that from some forms of M. metabates ((n <$ from Portugese E. Africa, Aden (1,000 <$ and 2,068 $ in spirit), Morocco and SW. Africa)) are near fulva while 9 <$ from M. gabar and 7 <$ from M . metabates from two individuals from Uganda differ constantly in the characters of the male genitalia and cannot be included in fulva. Specimens from Hypomorphus urubitinga belong to the fulva species group but seem to differ in the details of the male genitalia but the available material is not in sufficiently good condition for a decision on this. MATERIAL EXAMINED. Three <$, 6 $ from Aquila chrysaetos (Linn.), Portugal; 3 $ from Aquila heliaca Savigny, Kurdestan ; 2 $ from Aquila rapax (Temminck), Rajputana and Kenya ; 3 $, 5 $ from Aquila clanga Pallas, Czechoslovakia and Germany ; 4 <, 5 $ from Aquila verreauxii Lesson, Rondebosch, South Africa ; 7 (, 7 $ from Aquila wahlbergi, Sundevall, Uganda ; i <$, 8 $ from Aquila pomarina Brehm, no data. Many males and females from the following forms of Buteo : B. rufinus rufmus (Cretzschmar), B. rufinus cirtensis (Levaillant), B. rufofuscus (R. J. Forster), B. r. augur Riippell, B. hemilasius Temminck & Schlegel, B. regalis (G. R. Gray), B. jamaicensis alascensis Grinnell, B. j. borealis (Gmelin), B. j. kriderii Hoopes, B. j. costaricensis Ridgway, B. harlani (Audubon), B. 1. lineatus (Gmelin), B. b. buteo (Linn.), (including holotype, allotype and paratypes of D. giebeli Hopkins), B. v. vulpinus (Gloger), B. b. burmanicus Hume, B. 1. lagopus (Pontoppidan) , B. I. s.- johannis (Gmelin). Four <, 25 $ from Geranoaetus melanoleucus australis Swan, i 5 o REVISIONS OF MALLOPHAGA GENERA Chile. Eighteen $, 25 $ from Icthyophaga ichthyaetus ichthyaetus (Horsfield), Deccan, India. Eight $, 7 $ from Lophaetus occipitalis (Daudin), Sudan, Uganda, Kenya. Seven $> 7 ? from Hieraaetus ayresii (Gurney), Uganda ; i ^, i $ from H. pennatus (Gmelin), Palestine. Sixteen <, 38 $ from Spilornis cheela albidus (Temminck), Raj- putana ; 2 $, i $ from Spilornis c. cheela (Latham), Nepal, 2 $, i $ from Spilornis c. burmanicus Swan, Thailand. Nine <, 7 $ from Polemaetus bellicosus (Daudin), Natal and Zoo. Five <, 16 $ from Melierax musicus poliopterus Cabanis, Kenya ; 11 c 5 ? from Melierax metabates subspp. from Aden, Morocco, SW. Africa, Portugese E. Africa. Degeeriella rima sp. n. Type host : Kaupifalco monogrammicus (Temminck) (Text-figs. 101, 123) This subspecies is distinguished from fulva by the head being narrower and more rounded anteriorly (Text-fig. 123), by the pleural thickening of at least some of the segments having the ventral outline rounded and in the male by having a definite lateral slit each side of the basal apodeme (Text-fig. 101). This last character should not be confused with a displacement of the lateral thickening of the basal apodeme at the usual slight interruption of this thickening, which may be found in any of the species. MATERIAL EXAMINED. Eleven <$, 8 $ from the type host from Uganda and N. Rhodesia. Holotype male and allotype female, slide No. 629 in the British Museum from Kaupifalco monogrammicus (Temminck) from Bunyoro, Uganda collected by W. J. Eggeling, 4.^.1940 and presented by G. H. E. Hopkins. Paratypes : 10 <, 7 $ from the same host species with data as given above. Measurements in mm. Male Length Breadth Range Mean Range Mean Head (10) . . . 0-49-0-53 0-52 . 0-37-0-40 0-38 Prothorax (2) . 0-23-0-25 Pterothorax (2) . 0-38-0-40 Abdomen (2) . . 0-97-1-02 . 0-51-0-52 Total (2) . . . I-73-I-83 Genitalia (i) . .0-34 C.I. (10) . . . 0-72-0-77 0-74 Female Head (8) ... 0-53-0-59 0-57 . 0-40-0-45 0-42 C.I. (8) ... 0-735-0-775 0-76 . REVISIONS OF MALLOPHAGA GENERA 151 Degeeriella africana sp. n. Type host : Stephanoaetus coronatus (Linn.) (PL 2, fig. i ; text-fig. 85) This form is distinguished from fulva by the shape of the head and marginal carina and the absence of a pleural seta on segment V. MALE. Head with inner dorsal edge of marginal carina indented medially, ventral suture reaches to anterior margin of head. Tergum II with median indentation, FIGS. 58-61. Male genitalia. 58-59. D. guimardesi. 58. Ventral. 59. Dorsal. 60-61. D. meinertzhageni. 60. Ventral. 61. Dorsal, da. dorsal endomeral arm. Ill with small median concavity of varying depth. Width of pleural thickening as vn. fulva, but that of segment VII has a smaller re-entrant head. Genitalia as in fulva, except that on the available material the sides of the basal apodeme appear to be straighter. FEMALE. Terga of IX-XI as in fulva. Genital plate relatively broader in the anteroposterior line and subvulval sclerites shorter and blunter (Text-fig. 85) . CHAETOTAXY OF ABDOMEN. As in fulva except that tergocentral setae of III-V are normally 6, range 5-8, and there is no pleural seta each side of V. MATERIAL EXAMINED. Six <, 8 $ from Stephanoaetus coronatus (Linn.), Nairobi, Kenya, 5.^.1917 (skin in Nairobi Museum) collected by G. H. E. Hopkins. 152 REVISIONS OF MALLOPHAGA GENERA Holotype male and allotype female, slide no 624 in the British Museum (Natural History) from Stephanoaetus coronatus with data as given above, presented by Mr. G. H. E. Hopkins. Paratypes : 5 <$, 7 $ from the same host individual. Measurements in mm. Male (5) Length Breadth Range Mean Range Mean Head .... 0-58-0-60 0-58 . 0-47-0-48 0-47 Prothorax ... . 0-28-0-33 '3 2 Pterothorax ... . 0-470-50 0-49 Abdomen . . . 1-20-1-33 I-2 5 0-60-0-67 ' 02 Total .... 2-12-2-31 2 -20 Genitalia (i) . .0-43 C.I. .... 0-79-0-82 0-80 Female Head (7) ... 0-60-0-63 0-62 . 0-48-0-52 0-50 Prothorax (8) . 0-31-0-35 0-33 Pterothorax (8) . . . 0-49-0-55 0-52 Abdomen (6) . . 1-22-1-50 i'37 0-63-0-70 0-67 Total (6) . . . 2-17-2-53 2-33 C.I. (7) ... 0-79-0-83 0-81 Degeeriella beaufacies Ansari, 1955 Type Host : Butastur teesa (Franklin) (PL 8, fig. 4 ; Text-figs. 2, 23, 43, 77) Degeeriella beaufacies Ansari, 1955. Proc. Vllth Pakistan Sci. Conf., Biol. : 43. Host : Butastur teesa. Degeeriella beaufacies Ansari, 1956. Indian Journ. Entom. 17 : 395 (1955). Host : Butastur teesa. It is being assumed that the specimens available from Butastur teesa are this species, although in the first reference the few words of description do not distinguish the species from any other Degeeriella, and the second reference, in which the species is also referred to as new, is even less informative. This species is distinguished from fulva by the form of the ventral suture, internal and external male genitalia and the nymphs. MALE. Shape of head similar to that of fulva, anterior margin varies from flattened to somewhat rounded ; ventral suture does not reach anterior margin of head ; marginal carinae of temples as in fulva. Tergites and pleurites as in fulva. Geni- talia differ from those of fulva in details of the mesosome (PI. 8, fig. 4, Text-fig. 43). Internal genitalia as in Text-fig. 2. REVISIONS OF MALLOPHAGA GENERA 153 FEMALE. Terga of segments IX-XI and genital region as in fulva, but inner genital sclerites somewhat narrower (Text-fig. 77). MEASUREMENTS. These fall within the range for specimens of D. fulva from Buteo species. The measurements given by Ansari for the types of beaufacies are markedly smaller. CHAETOTAXY OF ABDOMEN. As in fulva but the total number of marginal setae on the last segment of the male varies from 11-17. O ne female has 2 tergocentral setae on the anterior margin of IX as in discocephalus. FIGS. 62-63. Male genitalia, ventral view of distal area. 62. D. phlyctopygus. 63. D. mookerjeei. NYMPHS. Third instar nymphs differ from those of fulva in having the preantennal region narrowed to a greater extent anteriorly and somewhat pointed (Text-fig. 23) . MATERIAL EXAMINED. Eighty-one <$, 65 $ from Butastur teesa from various locali- ties in India ; i <$ from Butastur liventer (Temminck) from Burma. Degeeriella carrikeri sp. n. Type host : Leucopternis polionota Kaup. (PI. 2, fig. 2, Text-figs. 27, 44, 86) This species is distinguished from fulva by the sculpturing of the dorsal surface of the head, pleural thickening and details of the male genitalia. MALE. Head similar to that of fulva, but flattened anteriorly with slight median concavity ; inner margin of marginal carina with median indentation ; dorsal ENTOM. 7, 4. 7 154 REVISIONS OF MALLOPHAGA GENERA sculpturing more marked and forming semicircular patch near anterior margin of head ; ventral suture passes nearly to anterior margin and is broad anteriorly ; marginal carinae of temples as in fulva. Tergites as in fulva. Pleural thickening broader than in fulva, with ventral outline of segments III-VII and dorsal outline of segments V-VII convex. Genitalia differ from those of fulva in the shape of the basal apodeme and details of mesosome. FEMALE. Terga of IX-XI and genital region as in fulva except for the shape of the subvulval sclerites (Text-fig. 86). CHAETOTAXY OF ABDOMEN. As in fulva except for the smaller number of tergo- central setae on segments II-V : II normally 4, range 3-5, III-V normally 6, range 4-7. In the male total number of marginal setae on last segment varies from 13-18. MATERIAL EXAMINED. Fifteen ^, n ? from Leucopternis polionota Kaup from S. Paulo, Brazil collected by S. Lima, November, 1949. Holotype male and allotype female in the collection of Dr. L. R. Guimaraes from Leucopternis polionota with the above data. Pamtypes : 14 <, 10 $ from the same host individual. Named in honour of Mr. M. A. Carriker. Measurements in mm. Male Length Breadth Range Mean Range Mean Head (15) . . . 0-60-0 -62 0-61 . 0-46-0-49 0-47 Prothorax (10) . . . 0-31-0-35 0-33 Pterothorax (10) . . . 0-45-0-53 0-48 Abdomen ( I o) . . 1-13-1-27 1-19 . 0-58-0-65 0-60 Total ( i o) . . . 2-15-2-30 2-19 Genitalia (3) . . . 0-408-0-412 C.I. (15) . . . 0-770-0-795 0-786 Female Head (10) . . . 0-62-0-67 0-65 . 0-48-0-52 0-^51 C.I. (10) . . . 0-790-0-815 0-796 Degeeriella emersoni sp. n. Type host : Buteogallus gundlachii (Cabanis) (Text-figs. 28, 45, 78, 87) This species is distinguished from fulva by the form of the marginal carina, pleural thickening and male genitalia. MALE. Head with general outline as in fulva, but anterior margin of marginal carina flattened and slightly concave medially ; inner margin of marginal carina indented medially ; ventral suture reaches to or nearly to anterior margin of head ; marginal carinae of temples as in fulva. Abdominal tergites as in fulva. Pleural REVISIONS OF MALLOPHAGA GENERA 155 thickening ventrally as in fulva, that is narrow with straight margin ; dorsal outline broader and curved. Genitalia similar to those of fulva, but differ in details of penial arms and endomeral plate. FEMALE. Terga of IX-XI and genital region as in fulva ; genital plate and sub- vulval sclerites as in Text-figs. 78, 87. CHAETOTAXY OF ABDOMEN. Tergocentral setae with range as in fulva, but segments II-V normally have 6-7 rarely 8. Pleural and sternal setae as in fulva. In the male the total number of marginal setae of the last segment varies from 11-15. Measurements fall within the range as given for specimens from Buteo lagopus (see table). MATERIAL EXAMINED. Twenty-three <, 34 $ from Buteogallus gundlachii Cabanis from Doce Legues, Cuba (collected by H. S. Peters). 4 <, 3 ? in rather poor condition from Parabuleo unicinctus (Temminck) seem to belong to this species. Holotype <$ and allotype $ in U.S. Bureau of Entomology, Washington from Buteo- gallus gundlachii with data as above. Paratypes : 22 <$, 33 $ from the same host individual. This species is named in honour of Dr. K. C. Emerson. Degeeriella nisus (Giebel) Specimens of Degeeriella have been seen from only nine species of Accipiter out of the 44 listed by Peters (1931), but even this small number shows more diversity in the populations from the different host species than in the case of the populations from Buteo. Four forms are here recognized and placed as subspecies of nisus, although when a greater amount of material is available from Accipiter it may be necessary to recognize some of the populations as species. For instance, haydocki and j 'rater are rather different from nisus and vagans and could perhaps be considered as specifically distinct. It should be noted that there tends to be some variation in the outline of the endomeral plate. Degeeriella nisus nisus (Giebel), 1866 Type host : Accipiter n. nisus (Linn.) (PL 3, fig. i ; PI. 8, fig. 5 ; Text-figs. 24, 29, 46, 88, no) Nirmus nisus Giebel, 1866. Z. ges. NatWiss. 28 : 364. Host : Astur nisus = Accipiter n. nisus (Linn.). This species is distinguished from fulva by the shape of the head, the form of the marginal carina, the pleural thickening and the details of the male genitalia and from fusca as given under that species. MALE. Inner edge of marginal carina straight or with slight median indentation ; small area of dorsal thickening immediately below marginal carina ; ventral suture does not reach to anterior margin of head (Text-fig, no). Marginal temporal carinae broad with many indentations. Terga II-III indented medially. Pleural 156 REVISIONS OF MALLOPHAGA GENERA thickening broad with ventral outline convex. Genitalia similar to those of fulva but differ in detail (PL 8, fig. 5 ; Text-figs. 29, 46) ; there is some variation in the shape of the dorsal endomeral and penial arms. Internal genitalia, represented by one example in rather poor condition, appear to be the same as those of D. fulva from Buteo buteo. FEMALE. Abdominal terga of IX-X as in fulva. Genital region similar to that of fulva ; subvulval sclerites as in Text-fig. 88. 68^- 69 FIGS. 64-69. 64-67. D. phlyctopygus. 64. Male genitalia. 65. Pleural thickening of segment IV. 66. Male thoracic sternal plate. 67. Dorsal arms of endomeral plate. 68-69. D. mookerjeei. 68. Pleural thickening of segment IV. 69. Dorsal arms of endomeral plate. CHAETOTAXY OF ABDOMEN. Tergocentral setae : II normally 4, range 3-5 ; III-VI normally 6, range 4-7, VIII range 4-6 ; X in the male with i seta each side (58 specimens examined), in the female 2 each side ; in the male total number of marginal setae, dorsal and ventral on last segment varies from 5-12. Pleural and sternal setae as in fulva. NYMPHS. Anterior margin of head of third instar rather less flattened than in adult (Text-fig. 24). MATERIAL EXAMINED. Sixty-six <, 113 $ from various subspecies of Accipiter nisus (Linn.) from the British Isles, Hungary, Cyprus, Saudi Arabia, Afghanistan and REVISIONS OF MALLOPHAGA GENERA 157 Pakistan. Two $, 4 $ from Accipiter striatus velox (Wilson) from British Columbia and U.S.A. are included under nisus nisus, although in the small number of specimens available the marginal temporal carinae are somewhat narrower. Neotype of Nirmus nisus Giebel : Male, slide no. 627, in the British Museum (Natural History) from Accipiter n. nisus (Linn.) from Kildare, Ireland, presented by Mr. G. H. E. Hopkins. Degeeriella nisus vagans (Giebel), 1874 Type host : Accipiter gentilis (Linn.) (PI. 3, ng. 2) Nirmus vagans Giebel, 1874. Insecta epizoa : 126. Host : Astur palumbarius = Accipiter gentilis (Linn.). This differs from the nominate form in the larger average size of both sexes, the shape of the head, the inner edge of the marginal carina, which is usually rather more indented medially, the narrower and less indented marginal carinae of the temples and the number of tergocentral setae. Tergocentral setae : II normally 6 (rarely 5 or 8, often 7) ; III-V normally 8 ; VI-VIII normally 6 ; thus in nisus the total number of tergocentral setae on segments III-V is 15-20, normally 18 and in vagans 22-26, normally 24. NYMPHS. Third instar nymphs with head similar to those of nisus nisus, but differ slightly reflecting the differences in the adult heads. MATERIAL EXAMINED. Forty-one <$, 44 $ from Accipiter gentilis (Linn.) from Germany, Switzerland, Czechoslovakia, Canada and Alaska. Fourteen <$, 48 $ from Accipiter cooperii (Bonaparte) from United States of America and British Columbia are not separable from vagans. Neotype of Nirmus vagans (Giebel), 1874 : Male, slide no. 628 in the British Museum (Natural History) from Accipiter gentilis from Rheinfelden, Switzerland, 15. ii. 1943 presented by Mr. G. H. E. Hopkins. Measurements in mm. Male D. n. nisus Length Breadth Range Mean Range Mean Head (50) . . . 0-45-0-52 0-49 . 0-33-0-39 0-36 Prothorax (10) . . . 0-22-0-25 0-23 Pterothorax (10) , . 0-33-0-38 0-36 Abdomen ( i o) . . 0-90-1-10 0-99 . 0-42-0-53 0-48 Total ( i o) . . . i -56-1 -87 1-72 Genitalia (2) . . 0-29-0-31 C.I. (50) . . . 0-73-0-79 0-76 . 158 REVISIONS OF MALLOPHAGA GENERA Head (12) . Pro thorax (10) Pterothorax (n] Abdomen (10) Total (10) . Genitalia (i) C.I. (12) . Head (10) . Pro thorax (10) Pterothorax (10) Abdomen (10) Total (10) . C.I. (10) . Head (12) . Prothorax (10) . Pterothorax (10) Abdomen (10) Total (10) . C.I. (12) . , Male D. n. vagans 0-52-0-56 0-54 1-05-1-17 i 83-2 oo 0-32 0-77-0-80 i -ii 1-94 0-79 Female D. n. nisus 0-52-0-57 0-54 1-13-1-36 1-91-2-23 0-73-0-78 1-27 2 -09 1-21-1-35 2-07-2-25 0-74-0-81 Female D. n. vagans 8 0-57 1-30 2-19 0-78 0-39-0-43 (30) 0-42 0-27-0-29 0-28 0-420-47 '44 o-55-o-59 0-57 0-37-0-42 (30) 0-40 0-22-0-27 0-26 o-37-o-43 0-41 0-45-0-58 0-54 0-42-0-46 (30) 0-45 0-28-0-32 0-30 0-47-0-49 0*48 0-57-0-63 0-62 Degeeriella nisus f rater (Piaget), 1880 Type host (emended) : Accipiter badius (Gmelin) (PI. 3, fig. 3 : Text-figs. 25, 31-32, 47) Nirmus frater Piaget, 1880. Pediculines : 145, pi. 12, fig. 2. Host : Lamprotornis amethystina. Error. Nirmus frater is represented in the Piaget collection by a single male labelled as from the original host with " Habesh " in brackets, perhaps Habesh in N. Syria. It has not been possible to separate specimens from Accipiter badius (African and Syrian birds, see below) from the type of frater (although exact comparison of the male genitalia is not possible) and it is presumed that this bird was the original host. This subspecies is distinguished from the nominate form by the shape of the head, by the slight concavity of the central part of the outer edge of the marginal carina, by the rather larger dorsal central thickening below the marginal carina, by the narrower and more curved penial arms and the shape of the endomeral plate. There is, however, some variation in this last character (Text-figs. 31-32). Tergum II does not have a narrow median indentation as is usual in nisus, but a shallow con- REVISIONS OF MALLOPHAGA GENERA 159 cavity which is sometimes hardly visible ; tergum III and chaetotaxy of the abdomen as in nisus. NYMPHS. Second and third stage nymphs have been seen from A badius poliopsis and third stage from A. virgatus affinis, these resemble each other and differ from 76 FIGS. 70-79. 70-76. Female genital regions. 70. D. fulva from Aquila chyrsaetos. 71. D. hopkinsi, 72. D. d. discocephalus. 73. D. meinertzhageni. 74. D. guimardesi. 75. D. rufa. 76. D. elbeli. 77. Inner genital sclerite of D. beaufacies. 78-79. Female genital plates. 78. D. emersoni. 79. D. leucopleura. those of Degeeriella n. nisus and D. n. vagans in having the anterior margin of the head more pointed and a larger thickened area anteriorly. Specimens have been seen from subspecies of Accipiter badius from Syria, Somali- land, Uganda, Kenya, Nepal and Thailand. Those from the Thailand birds (^4. 160 REVISIONS OF MALLOPHAGA GENERA badius poliopsis] tend to differ from those from the Syrian and African birds in having the outer edge of the marginal carina somewhat more concave medially and in having none or very few indentations in the inner edge of the marginal carina laterally. The specimens from Nepal resemble those from Thailand in the form of the anterior margin of the head and the African specimens in the lateral indentations of the, marginal carina. However, there are individuals from all these localities which are indistinguishable from each other ; it does not seem reasonable, therefore, to dis- guish taxonomically the populations from these different subspecies of Accipiter badius. There is also some variation in the shape of the dorsal endomeral arms. MATERIAL EXAMINED. One $ type of Nirmus frater ; males and females from Accipiter badius (Gmelin) from Syria (i <$, 3 $), from Africa (Uganda, Kenya, Somaliland, 14 <, 26 $), from Nepal (23 J, 27 $), from Burma (7 , 10 ?) and Thailand (9 <$, 14 $). Three J, 8 $ from Accipiter tachiro (Daudin) from Uganda and S. Africa. One <$, 4 ? from Accipiter virgatus affinis Hodgson from Thailand and 2 ^, 3 $ from A. virgatus gularis (Temminck & Schlegel) from Thailand. Ledotype of Nirmus frater Piaget : J (slide no. 1270) in the Piaget collection, British Museum (Natural History). Measurements in mm. Lectotype (g) of frater Length Breadth Head . . . 0-57 . 0-43 Prothorax . . . 0-29 Pterothorax . . . 0-47 Abdomen . . 1-18 . 0-58 Total . . . 2-03 Genitalia . . 0-33 C.I. ... Breadth of Head of Male Specimens from Accipiter badius East Africa (n) Nepal (22) Thailand (10) Range Mean Range Mean Range Mean 0-38-0-42 0-40 . 0-38-0-43 0-42 . 0-38-0-40 0-39 Degeeriella nisus haydocki subsp. n. Type host : Accipiter minullus (Daudin) (PI- 2, fig. 3) This form is separated from the other known subspecies of nisus with the exception of epustulata by having only four tergocentral setae on segments II-VIII. It is separated from this latter species by the size and shape of the head. It resembles frater in the characters of the anterior margin of the marginal carina ; in having a dorsal triangular-shaped thickening below the marginal carina, which is rather larger ; in the form of tergum II and in the shape of the penial arms. REVISIONS OF MALLOPHAGA GENERA 161 MATERIAL EXAMINED. Nine <$, u $ from Accipiter minullus (Daudin) from Gulu, Uganda and N. Rhodesia. Holotype male and allotype female, slide no 625 in the British Museum (Natural History), from Accipiter minullus, Mulashi, N. Rhodesia, 27. vi. 1955 collected by Major E. L. Haydock. Paratypes : 8 <$, 10 $ from the same host species with data as given above. Measurements in mm. Male Length Head (9) Pro thorax (7) Pterothorax (7) Abdomen (7) Total (7) Genitalia (2) . C.I. (9) Head (10) Prothorax (8) Pterothorax (8) Abdomen (8) Total (8) C.I. (10) Range Mean 0-49-0-53 0-50 0-95-1-05 0-97 1-71-1-83 1-75 0-325-0-330 0-69-0-73 0-71 Female o-53-o-57 0-55 1-17-1-27 2-OO-2 -IO o 68-0 74 I-2I 2-03 0-71 Breadth Range o-35-o-37 0-24-0-26 o-37-o-39 0-46-0-50 0-37-0-42 0-26-0-28 0-40-0-43 0-52-0-57 Mean 0-360 0-250 0-375 o-475 0-39 0-27 0-42 o-53 Degeeriellia nisus epustulata (Carriker), 1903 Type host : Accipiter bicolor (Vieillot) (Text-fig. 124) Nirmus fuscus epustulatus Carriker, 1903. Univ. Nebr. Stud. 3 : 133. Host : Accipiter bicolor. Through the kindness of Mr. Carriker it has been possible to examine a single female paratype of this form. It resembles haydocki in having only four tergal setae on each of segments III-VIII, but differs from this form in the shape of the head and the larger size. Measurements in mm. Female Head . Prothorax Pterothorax Abdomen Total Length 0-60 1-30 2-12 Breadth o-47 0-30 o-47 o-57 162 REVISIONS OF MALLOPHAGA GENERA Degeeriellafusca (Denny), 1842 Type host : Circus ae. aeruginosus (Linn.) (PI. 4, fig. 3 ; PI. 8, fig. 6 ; Text-figs. 3, 30, 48) Nirmus fuscus Denny, 1842. Mon. Anopl. Brit. : 49, 118. Host : Circus rufus = Circus ae. aeruginosus (Linn.). Nirmus socialis Giebel, 1874. Insecta epizoa : 127. Hosts : Circus cineraceus = C. pyargus (Linn.), and C. aeruginosus (Linn. Nirmus aeruginosi Denny, 1852. List Brit. Animals in Brit. Mus., pt. n, Anoplura : 16. Nomen novum for Nirmus fuscus Denny. Kelerinirmus circi Boetticher & Eichler, 1954. Biol. Zbl. 73 : 215. Host : Circus aeruginosus (Linn.). Hopkins (1947 : 76) has discussed the confusion which has arisen over the author of this name and the type host and shown that Denny must be considered as the sole author with Circus ae. aeruginosus as the type host. This species resembles most nearly D. n. nisus from which it is distinguished by the colour pattern and details of the male genitalia. MALE. Dorsal surface of head with an area of lighter sclerotization between the anterior dorsal setae. Inner dorsal margin of marginal carina indented medially ; ventral suture as in nisus. Tergites II-III with median indentation ; central area of tergite II more strongly pigmented than lateral areas. Pleural thickening broad and strongly pigmented with dark inner line, contrasting with the rather lightly sclerotized terga ; this character is not so marked in specimens from Circus cyaneus. Genitalia similar to those offulva but differ in detail. Internal genitalia as shown in Text-fig. 3. FEMALE. Terga of IX-XI and genital region as in fulva. CHAETOTAXY OF ABDOMEN. Tergocentral setae : II normally 6, range 5-7 ; III-IV normally 8, range 6-9 ; V normally 7-8, range 5-8 ; VI-VII normally 6, range 5-8 ; VIII normally 6, range 4-6 ; X in the male has i seta each side (58 examined), in the female 2 each side ; total number of marginal setae on last segment varies from 6-12. Pleural and sternal setae as in fulva. NYMPHS. Third instar nymphs have been seen from two host species, Circus cyaneus and C. melanoleucus ; these have the anterior margin of the head more pointed than in the adult. HOST DISTRIBUTION. There appear to be no taxonomically recognizable differ- ences between the population from the five species of Circus listed below. Eleven specimens from one host individual of Circus melanoleucus average somewhat smaller (breadth of head : 0-40 mm.). Specimens from Circus cyaneus do not seem to have the colour pattern quite typical offusca, except for the darker central area of tergum II ; it is possible that these may prove to be a distinct subspecies, but fresh material from all hosts is needed. MATERIAL EXAMINED. Fifty-five <$, 81 from Circus aeruginosus (Linn.) from Czechoslovakia, Malta, Cyprus, Saudi Arabia, India, Ceylon, Cape Colony ; 13 $, 37 $ from Circus c. cyaneus (Linn.) from Orkneys, Hungary and Czechoslovakia ; REVISIONS OF MALLOPHAGA GENERA 163 10 <$, 12 $ from Circus cyaneus hudsonius (Linn.) from British Columbia and various localities in the United States of America ; 55 $, 74 $ from Circus Pygargus (Linn.) from Cyprus and Kenya ; 16 $, 24 $ from Circus macrourus (S. G. Gmelin) from N. Africa, Saudi Arabia and Aden ; 10 $, 9 $ from Circus melanoleucus (Pennant) ^rfty FIGS. 80-95. 80-83. Female genital regions. 80-81. D. r. regalis, to show variation ; 80 from Milvus m. milvus and 81 from Milvus migrans aegyptius. 82. D. r. castanea. 83. D. punctifer. 84-95. Subvulval sclerites. 84. D. fulva from Aquila chrysaetos. 85. D. africana. 86. D. carrikeri. 87. D. emersoni. 88. D. n. nisus. 89. D. leucopleura. 90. D. elbeli. 91. D. elani. 92. D. tendeiroi. 93. D. meinertzhageni. 94. D. guima- rdesi. 95. D. rufa from Falco tinnunculus. from Assam and Thailand. In the Denny collection there are 7 $ labelled Nirmus fuscus by the person responsible for mounting this collection and who rarely kept Denny's original labels. These specimens have no host label, but three of the slides have a small circular (probably original) label with what appears to be " aeruginos ". This must refer either to aeruginosi, the new name given to fuscus by Denny in 1852 or to the name of the host, C. aeruginosus. The three females so labelled together 164 REVISIONS OF MALLOPHAGA GENERA with one other are the species usually found on C. aeruginosus : two other females are D. r. regalis and presumably came from Milvus ictinus referred to by Denny (1842 : 119) and one other female is D. fulva and presumably came from Buteo lagopus also referred to by Denny. One of the females labelled " aeruginos " will be selected as lectotype of fusca. Lectotype : $, slide no. 350, in the Denny collection, British Museum (Natural History) ; paratypes, 3 $ in the same collection. Measurements in mm. Male Length Head (50) . Pro thorax (10) Pterothorax (10) Abdomen (10) Total (10) Genitalia (2) . C.I. (50) Head . Prothorax Pterothorax Abdomen Total . C.I. Range Mean 0-51-0-57 o-54 i -08-1 -23 I -88-2 -12 1-98 0-32-0-35 0-75-0-81 0-78 Female (10) 0-57-0-60 0-58 1-31-1-40 2-22-2-42 0-75-0-80 1-36 2-28 0-78 Breadth Range 0-39-0-45 O' 25-0 -28 0-42 0-45 o 52-0 60 0-42-0-47 0-28-0-31 0-47-0-50 o 62-0 68 Mean 0-42 0-27 o-43 0-56 o-45 0-29 0-48 0-64 Degeeriella hopkinsi sp. n. Type host : Terathopius ecaudatus (Daudin) (PI. 4, fig. 2 ; Text-figs. 33, 49, 71) This species is distinguished from the rest of the fulva species group by the presence of a pleural seta on segment IV. MALE. Head broad and rounded anteriorly ; inner edge of marginal carina indented dorsally in mid-line ; ventral suture extends to or nearly to the anterior margin. Terga II-III indented ; pleural thickening narrow with re-entrant heads normal only on segment III, gradually becoming more and more reduced on the following segments. Genitalia similar to those of fulva but differ in detail (Text- n s - 33> 49)- There is some variation in the number and position of the setae associated with the ventral arms of the endomeral plate in the five males examined : two specimens had an extra seta each side anterior to the end of the arms, one had three setae on one arm and on the other one anterior to the arm and one on the arm, one specimen was normal and in one the setae could not be seen. REVISIONS OF MALLOPHAGA GENERA 165 FEMALE. Terga of IX-XI as in fulva. Genital region similar to fulva but genital plate narrower from side to side and the subvulval sclerites shorter with blunter ends (Text-fig. 71) ; there are fewer sensillae anterior to the vulval margin. CHAETOTAXY OF ABDOMEN. Tergocentral setae : II, 6 ; III-V, 6-8 ; VI- VIII, 6 ; X in the male has 1-2 each side and in the female 2. Pleural and sternal setae as in fulva except that there is a pleural seta each side of IV. MATERIAL EXAMINED. Five <, n $ from Terathopius ecaudatus (Daudin) from Lodwar, Kenya, 7.^.1934 (skin in Nairobi Museum) collected by G. H. E. Hopkins. Holotype male and allotype female slide no. 623 in the British Museum (Natural History) from Terathopius ecaudatus with data as given above, presented by Mr. Hopkins. Paratypes : 4 <$, 10 $ from the same individual. Measurements in mm. Male Length Head (5) Pro thorax (5) Pterothorax (5) Abdomen (4) Total (4) Genitalia (i) . C.I. (5) Head (n) . Prothorax (7) Pterothorax (7) Abdomen (7) Total (7) C.I. (ii) Range I-I2-I-22 I-92-2-07 0-38 0-8I-0-85 Mean o-55 1-18 2-OO 0-82 Female o-53-o-59 0-57 1-18-1-28 1-25 2-07-2-17 2-13 0-82-0-85 0-83 Breadth Range 0-43-0-48 0-28-0-30 0-43-0-49 0-58-0-65 0-45-0-50 0-28-0-30 0-47-0-50 o 60-0 63 Mean o-45 0-28 0-47 0-62 0-47 0-28 0-48 0-62 Degeeriella leucopleura (Nitzsch), 1874 Type host : Circaetus cinerascens J. W. Muller (PI. 4, fig. i ; Text-figs. 34, 50, 79, 89) Nirmus leucopleurus Nitzsch, 1874. In Giebel, Insecta epizoa : 129. Host : Falco brachy- dactylus = Circaetus cinerascens J. W. Muller. Nirmus temporalis Piaget, 1890. Tijdschr. Ent. 33 : 228. PL 8, fig. 6. Host : Buceros manil- lensis. Error. This is a distinctive species separated by the shape of the head, form of the pleural thickening and absence of pleural seta on V. MALE. Head broad with preantennal region rounded ; inner dorsal edge of marginal carina with slight median indentation ; ventral suture does not reach anterior margin. Abdomen elongated and with neither terga II nor III with a 1 66 REVISIONS OF MALLOPHAGA GENERA definite median slit-like indentation although II may show a slight concavity. In spite of the elongated abdomen the pleural thickening does not have the strongly sclerotized re-entrant heads characteristic of this species group. Genitalia as shown in Text-figs. 34, 50. FEMALE. Terga of IX-XI as in fulva. Posterior margin of genital plate deeply emarginate and subvulval sclerites stouter than in fulva (Text-figs. 79, 89). CHAETOTAXY OF ABDOMEN. Tergocentral setae : II normally 6, range 4-6 ; III-V normally 8, range 6-8 ; VI-VII normally 6, range 5-7 ; VII normally 4, range 3-4 ; X in the male with i each side, in female 2 each side. One female has 4 tergocentral setae on II and 6 on III-VII. Pleural setae : II-V, o ; VI-VII, FIGS. 96-98. Degeeriella phlyctopygus, female. 96. Genital region. 97. Subvulval sclerites. 98. Dorsal view of segments IX-XI. ig. inner genital sclerites ; os. opening of spermathecal tube ; sv. subvulval sclerite. 2 each side ; VIII, 3 ; in the male IX has 3 and X, o. In the female IX has 3 and X 2 each side. Sternocentral setae : II-VI normally 4, one male has 3 on VI and one female has 6 on V ; in the male the last segment does not have the usual spine- like seta each side and the second seta therefore, if present, is not distinguishable from the marginal setae which total from 10-15. NYMPHS. Second and third instars have the curvature of the anterior margin of the head similar to that of the adult ; the preantennal region is shorter and the sides less straight, as is usual in nymphs. Nirmus temporalis Piaget said to have come from Buceros manillensis is represented in the Piaget collection by a single male which appears to be the same as D. leucopleura and is presumably a straggler from Circaetus. MATERIAL EXAMINED. Four <$, 2 $ from Circaetus cinerascens J. W. Muller from Kapenguria, Kenya and I <$ from the same host species from E. Africa (skin) ; 7 <$, ii $ from Circaetus gallicus (Gmelin) from France, Czechoslovakia, Egypt (skin) and Cameroons (skin) ; i <, i ? from Circaetus cinereus Vieillot from Portuguese Guinea. REVISIONS OF MALLOPHAGA GENERA 167 Lectotype of Nirmus temporalis Piaget : <$ in the British Museum (Natural History), slide no. 1421. Neotype of Degeeriella leucopleura (Nitzsch) : <$ in the Meinertzhagen collection, British Museum (Natural History), slide no. 20568 from Circaetus cinerascens from Kapenguria, Kenya, March 1956. Measurements in mm. Male Length Breadth Head (5) Prothorax (5) Pterothorax (5) Abdomen (3) Total (3) Genitalia (i) . C.I. (5) Head (2) Prothorax (2) Pterothorax (2) Abdomen (i) Total (i) C.I. (2) Range Mean o 65-0 -67 o 66 Range 1-32-1-43 2-33-2-45 o-43 0-79-0-81 1-38 2-39 0-80 Female o 68-0 69 0-32-0-33 o-50-o-55 o 65-0 68 Mean 0-52 o-33 0-52 0-67 (5) i -40 2-43 o 77-0 78 o-35 o-53-o-55 0-67 The discocephalus Species Group i. Head index greater than 0-94. 2-3. As in fulva group. 4. Thoracic sternal plate as in Text-fig. 119. 5. Thorax and abdomen with shape as in PI. 9, fig. i. 6-7. As in fulva group. 8. Pleural thickening without well sclerotized re-entrant heads. 9. Sternite II in form of central triangular plate, III-VI narrow central strips of sclerotization. 10. Male genital plate small and irregular and less indented laterally than in fulva group. 11-13. As in fulva group. 14. Setae each side of posterodorsal margin of pterothorax variable in number and position. 15. Pleural setae absent on segments II-III. 16. Sternocentral setae of segments III-VI variable. 17. As in fulva group. This species group is distinguished from the fulva group by the shape of head and abdomen ; from the punctifer group by the absence of head sutures and chaetotaxy of the temples, and in the females by the dorsal chaetotaxy of tergum IX and in the male by the characters of the male genitalia. i68 REVISIONS OF MALLOPHAGA GENERA Degeeriella discocephalus discocephalus (Burmeister), 1838 Type host : Haliaeetus albicilla (Linn.) (PI. 9, fig. i ; Text figs. 35, 51, 72, 106, 119) Nirmus discocephalus Burmeister, 1838. Handb. Ent. 1 : 430. Host : Aquil. albicilla = Haliaeetus albicilla (Linn.). Nirmus discocephalus var. amblys Kellogg, 1896. Proc. Calif. Acad. Sci. (2), 6 : 499, pi. 67, fig. 6. Host : Haliaeeetus leucocephalus (Linn.). The specimens used by Burmeister for his description of this species were figured by Nitzsch in Giebel, 1874 (pi. 7, fig. 10) and represent the species described below. MALE. Marginal carina thick and entire ; ventral suture reaches to or nearly to inner margin of marginal carina. Thoracic sternal plate as in Text-fig. 119, but shows some variation in outline ; posterodorsal marginal setae of pterothorax variable in number and position, 4-6 each side (omitting the lateral spine-like seta and the seta with sunken alveolus). Tergum II with median unsclerotized area, tergum III somewhat narrowed medially. Genitalia of fulva type ; dorsal endomeral arms may or may not join basal apodeme. FEMALE. This species differs from all other known Degeeriella in having two (occasionally one) setae in the middle of the anterior region of tergite IX ; these setae are occasionally found as an abnormality in species of the fulva group. The subvulval sclerites are small and indistinct and almost covered by the vulva (Text- fig. 72). CHAETOTAXY OF ABDOMEN. Tergocentral setae : II range 6-8 ; III-V, normally 8, range 6-9 ; range 6-8 ; VII- VIII normally 6, range 6-7 ; X in the male normally 2 each side, range 2-4 ; in the female IX has 2 (rarely i) anterior setae ; X has 2 each side. Pleural setae ; II-III, o ; IV-V, i each side ; VI-VII, 2 (occasionally i or 3 on one side) ; VIII, 3 ; IX, 2 ; X in the male has o and in the female 2. Sternocentral setae irregular in number : II, 4-7 ; III-IV, 5-8 ; V, 5-7 ; VI, 4-7 ; total number of setae of segments II-VI of specimens counted varied from 24-34 ; in the male the last pair of sternal setae are both elongated. Total number of marginal setae of last segment in male varies from 12-16. NYMPH. One third instar from Haliaeetus albicilla has been seen, this resembles the adult in the shape of the head. MATERIAL EXAMINED. Fourteen <$, 12 $ from Haliaeetus albicilla (Linn.) from Germany, Czechoslovakia, Finland and Russia ; 2 J, 5 $ from Haliaeetus pelagicus (Pallas) from Siberia ; 7 <$, 3 $ from Haliaeetus I. leucocephalus (Linn.) from various localities in N. America. There appear to be no constant characters on which the population (i.e. D. amblys (Kellogg)) found on this latter host can be separated from discocephalus. Neotype of Nirmus discocephalus Burmeister : <$ in the British Museum (Natural History), slide no. 617 from Haliaeetus albicilla from Samorin, Czechoslovakia, 9.xii.i952. REVISIONS OF MALLOPHAGA GENERA 169 xi 100 103 104 106 107 108 FIGS. 99-108. 99. Male antenna, D. mookerjeei. 100. D. guimardesi, terminal segments of male abdomen. 101. D. rima, basal apodeme (lateral slits somewhat exaggerated). 102-103. Male genital plates. 102. D.fulva iromAquila chrysaetos. 103. D. punctifer. 104. D. rufa from Falco tinnunculus, pleural thickening of segment IV, ventral. 105-108. Segments IX-XI of female abdomens, dorsal. 105. D. fulva from Aquila chrysaetos. 106. D. d. discocephalus. 107. D. rufa from F. tinnunculus. 1 08. D. punctifer. ENTOM. 7, 4. i 7 o REVISIONS OF MALLOPHAGA GENERA Measurements in mm. Male Length Range Mean Range Head (n) . . . 0-44-0-47 0-46 . 0-42-0-47 0-45 Prothorax (6) . 0-25-0-28 0-27 Pterothorax (8) . '35-'43 0-40 Abdomen (7) . . 0-70-0-87 0-77 . 0-52-0-68 (8) 0-61 Total (7) . . . i -37-1 -53 1-45 Genitalia (3) . . . 0-42-0-44 C.I. (u) . . . 0-96-1-00 0-98 Female Head (7) . . . 0-50-0-53 0-51 . 0-50-0-53 0-52 Prothorax (6) . 0-28-0-33 0-31 Pterothorax (6) . 0-45-0-49 0-46 Abdomen (6) . . 0-97-1-13 1-05 . 0-68-0-77 0-73 Total (6) . . . 1-17-1-95 1-83 C.I. (7) . . . 1-00-1-03 I>01 Degeeriella discocephalus aquilarum Eichler, 1943 Type host : Aquila n. nipalensis (Hodgson) (PI. 9, %. 2) Degeeriella aquilarum Eichler, 1943. Zool. Anz. 142 : 92, fig. i. Host : Aquila n. nipalensis (Hodgson) . This subspecies is close to the nominate form, from which it can be separated by the shape of the anterior margin of the head, the slightly better developed pleural thickening, especially on segment III and the fewer number of tergo- and sternocentral setae as follows : Tergocentral II-VIII Sternocentral II-VI ^ . A Maximum Minimum Maximum Minimum D. d. discocephalus . 52 48 '. 34 24 D. d aquilarum 42 38 . 22 18 (i ? with 31) SPECIMENS EXAMINED. Nine ^, 8 $ from Aquila n. nipalensis Hodgson from Somaliland ; i <$, i $ from Aquila nipalensis orientalis Cabanis, no data ; 4 <$, 4 $ from Aquila chrysaetos (Linn.) from Norway and Serbia ; i <$, 2 $ from Aquila pomarina hastata (Lesson) from Rajputana, India and Manipur ; 8 <, 4 $ from Aquila rapax raptor A. E. Brehm from Somaliland ; 4 J, 3 $ from Aquila h. heliaca Savigny from Czechoslovakia, Egypt and Rajputana, India ; i $ from Aquila clanga Pallas from Russia. REVISIONS OF MALLOPHAGA GENERA 171 There appear to be no constant differences between the populations from these species of Aquila, although no doubt there will be found some differences in average sizes when larger numbers are available. Measurements in mm. Specimens from Aquila n. nipalensis Male (9) Length Breadth Range Mean Range Mean Head .... 0-44-0-47 0-45 . 0-43-0-47 0-45 Prothorax ... . 0-25-0-27 0-26 Pterothorax ... . 0-38-0-42 0-40 Abdomen . . . 0-81-0-93 0-89 . 0-55-0-67 0-62 Total .... 1-45-1-63 1-56 C.I. .... 0-96-1-02 0-99 The elani Species Group 1-5. As in fulva group. 6. As in fulva group : elbeli and tendeiroi. Tergal plates of segment XI apparent in male : elani, meinertzhageni, guimaraesi. 7. As in fulva group : elbeli, tendeiroi and guimaraesi. As in rufa group : elani and meinertzhageni. 8-9. As in fulva group. 10. As in fulva group : elbeli. Male genital plate laterally indented to a greater extent : elani, tendeiroi, meinertzhageni and guimaraesi. n. As in fulva group : elbeli, elani, tendeiroi, meinertzhageni. Female genital plate with median prolongation : guimaraesi. 12. As in fulva group : elbeli, elani, tendeiroi. Inner genital sclerites fused : meinertzhageni and guimaraesi. 13. As in fulva group : elbeli, elani, tendeiroi. Distinctive types : meinertzhageni and guimaraesi. 14. As in fulva group. 15. Pleural setae absent on segments II-IV : elbeli. Pleural setae absent on segments II-V : elani, tendeiroi, meinertzhageni, guimaraesi. 16-17. As i n fulva group. It is apparent that these five species do not form a very homogeneous group and are here placed together mainly on the form of the ventral carinae which show a greater development anteriorly than those of any other species groups ; this character is more marked in meinertzhageni and guimaraesi than in the others. Apart from this character elbeli and perhaps tendeiroi could be included in the fulva group ; the rest of the species share some rather distinctive characters ; guimaraesi has certain characters found elsewhere only in rufa. It is possible that these five species do not, in fact, form a related group. 172 REVISIONS OF MALLOPHAGA GENERA Degeeriella elbeli Clay, 1958 Type Host : Aviceda leuphotes burmana (W. L. Sclater) (PL 7, figs. 3, 6 ; Text-figs. 52, 76, 90) Degeeriella elbeli Clay, 1958. Proc. R. ent. Soc. Land. (B) 27 : 6, PI. i, figs. 3, 6 ; Text-figs. 3, 9, 14, 17. Host : Aviceda leuphates burmana (W. L. Sclater). This species is distinguished from the rest of the species group by the presence of pleural setae on segment V, by the shape of the head and the male genitalia. It is separated from members of ihefulva group, which it resembles in many characters, by the greater development of the ventral carinae anteriorly, and by a combination 119 120 122 FIGS. 109-122. 109-111. Central anterior margin of head. 109. D. fulva from Buteo lagopus. no. D. n. nisus. in. D. rufa. 112-122. Thoracic sternal plates. 112- 118. D. fulva (cJ from Haliaeetus leucoryphus (Pallas) from various localities in India. Neotype of Nirmus regalis Giebel : from the type host from Piche, Gabu and Mansoa in Portugese Guinea. Measurements in mm. D. r. regalis from Milvus milvus. Male Length Breadth Head (10) . Prothorax (10) Pterothorax (10) Abdomen (10) Total (10) . Genitalia (i) C.I. (10) . Head (10) . Prothorax (10) . Pterothorax (10) Abdomen (10) Total (10) . C.I. (10) . Range 0-52-0-58 1-05-1-23 1-87-2-17 0-40 o 74-0 78 Mean o-55 2-02 0-76 Range Mean 0-38-0-43 (18) 0-26-0-28 0-40 0-28 0-43-0-49 0-52-0-63 0-46 o-57 Female 0-57-0-60 0-58 1-37-1-48 2-25-2-43 o 74-0 80 1-41 2-33 0-77 0-41-0-46 (l2) 0-44 0-28-0-30 0-29 o 490 - 5 3 0-51 o 60-0 -70 0-65 Degeeriella r. deignani Male head Thailand (5) Ceylon (5) Length Breadth Range o 57-0 60 0-42-0-45 Mean 0-58 o-43 Range 0-57-0-58 0-41-0-43 Mean o-57 0-42 REVISIONS OF MALLOPHAGA GENERA Head (15) . Prothorax (10) . Pterothorax (10) Abdomen (10) Total (10) . Genitalia (3) C.I. (15) - Degeeriella v. castanea Male Length Range o 55-0 60 1-17-1-32 2-05-2-25 0-41-0-42 o 74-0 76 Mean o-57 1-24 2-14 o-75 Breadth Range 0-41-0-45 0-27-0-32 o 42-0 50 o 52-0 67 Mean o-43 0-29 0-47 o-59 Head (27) . Prothorax (10) Pterothorax (10) Abdomen (10) Total (10) . C.I. (27) . Female 0-57-0-65 0-61 i -50-1 -60 2-47-2-57 o 74-0 79 1-52 2-49 0-76 0-43-0-49 0-29-0-33 0-51-0-54 o 62-0 69 0-46 0-32 0-52 0-67 Degeeriella r. subsp. (from Tendeiro, in press) Male (2) Length Breadth Head 0-57-0-58 0-460-47 Prothorax o-34 Pterothorax 0-48-0-49 Abdomen I -2O-I -23 o-55 Total 2-I6-2-I8 C.I. 0-81 . Female (5) Length 0-61 0-80-0-82 Breadth 0-49-0-50 The phlyctopygus Species Group 1-12. As in fulva group. 13. Male genitalia of distinctive type ; penial sclerite present. 14. As in fulva group. 15. As in regalis group. 16. Sternocentral setae of segments III-VI normally more than 4. 17. As in fulva group. This species group, which has only two known species, is distinguished from all other groups by the form of the male genitalia ; in the characters of the chaetotaxy of the abdomen it resembles the regalis group. REVISIONS OF MALLOPHAGA GENERA 193 Degeeriella phlyctopygus (Nitzsch), 1861 Type Host : Pernis a. apivorus (Linn.) (PI. 8, fig. i ; Text-figs. 62, 64-67, 96-98) Nirmus phlyctopygus Nitzsch, 1861. In Giebel, Z. ges. NatWiss. 17 : 526. Host : Pernis apivorus. This species (redescribed by Clay, 19570) is distinguished from D. mookerjeei the only other species in the species group, by the antennae and the genitalia in the male and by the shape of the head in the female. MALE. Anterior margin of head slightly concave, inner margin of dorsal marginal carina slightly indented medially ; ventral suture does not reach anterior margin of head. Tergite II only with definite median indentation. Pleural thickening of segments III-VI with well-developed re-entrant heads (Text-fig. 65) . The genitalia (Text-fig. 62) are unlike any other known species of Degeeriella except those of D. mookerjeei. FEMALE. Terga of IX-XI and genital region as in Text-figs. 96-98. CHAETOTAXY OF ABDOMEN. Male. Tergocentral setae : II, 5-6 ; III-V, 8 : VI, 7 ; VII, 6 ; VIII, 6-7 ; X, 1-2 each side. Pleural setae : II-III, o ; IV-V, i each side ; VI-VII, 2 ; VIII, 3 ; IX, 3-5 ; X-XI, o. Sternocentral : II, 5-6 ; III, 6-7 ; IV-V, 6 ; VI, 5-6. Total number of marginal setae, dorsal and ventral, of last segment (i specimen) : 13. In the female tergocentral setae : II normally 6, range 5-7 ; III-V normally 8, range 7-8 ; VI range 6-8 ; VII-VIII normally 6, range 5-6 ; X, 2 each side. Pleural setae : II-VIII as in male ; IX normally 3 each side, range 3-5 ; X normally 3, range 2-3. Sternocentral setae : II normally 6, range 4-6 ; III-IV normally 6, range 6-7 ; V normally 6, range 5-6 ; VI range 4-6 ; VII-XI as in Text-fig. 96. MATERIAL EXAMINED. < neotype and 2 Milvus lineatus J X Haliastur Indus . Subfamily ACCIPITRINAE X A ccipiter nisus "\ Accipiter striatusf X A ccipiter gentilis\ A ccipiter cooperii J A ccipiter badius ~\ A ccipiter tachiro > Accipiter virgatus J X A ccipiter minullus X Accipiter bicolor . Melierax musicus Melierax metabates Melierax gabar . Geranoaetus melanoleucus' Buteo rufinus Buteo rufofuscus Buteo hemilasius Buteo regalis Buteo jamaicensis HOST-LIST FALCONIFORMES . Degeeriella species No Degeeriella. No Degeeriella. Species group Page No. { elani meinertzhageni guimardesi elbeli phlyctopygus mookerjeei r. regalis xr. deignani msus msus nisus vagans nisus frater nisus haydocki nisus epustulata fulva + M + fulva r. regalis elani phlyctopygus regalis fulva Buteo harlani Buteo lineatus Buteo buteo Buteo vulpinus Buteo burmanicus Buteo lagopus Buteo swainsoni . Buteo galapagoensis ..,,,,. * Genera and species from which no Degeeriella have been seen are not X Type host, + or subspecies. ENTOM. 7, 4. fulva regalis regalis regalis fulva regalis included. 174 176 177 172 193 194 186 190 155 157 158 1 60 161 149 149 149 144 186 144 186 REVISIONS OF MALLOPHAGA GENERA Host* Parabuteo unicinctus X Leucopternis polionota X Kaupifalco monogrammicus X Butastur teesa Hypomorphnus urubitinga X Biiteogallus gundlachii Lophaetus occipitalis X Stephanoaetus coronatus Polemaetus bellicosus Hieraaetus ayresii xAquila chrysaetos Aquila heliaca Aquila rapax X Aquila nipalensis Aquila clanga Aquila pomarina Aquila verreauxii Aquila wahlbergi xHaliaeetus vocifer Haliaeetus pelagicus Haliaeetus leucoryphus Haliaeetus leucocephalus X Haliaeetus albicilla Icthyophaga ichthyaetus Subfamily AEGYPIINAE X Gyps himalayensis X Gypohierax angolensis . X Gypaetus barbatus Subfamily CIRCINAE Circus cyaneus . Circus macrourus Circus pygargus . Circus melanoleucus X Circus aeruginosus Subfamily CIRCAETINAE X Terathopius ecaudatus . Circaetus gallicus Circaetus gallicus Circaetus cinereus X Circaetus cinerascens . Spilornis cheela . Family FALCONIDAE Subfamily POLIHIERACINAE Neohierax insignis X Gampsonyx swainsonii Degeeriella species Species group Page No. emersoni fulva 154 carrikeri fulva 153 nicus . rima ,, 150 beaufacies ,, 152 nga . ? 149 emersoni ,, 154 . . fulva-\- ,, 149 '.s . africana I5 1 fulva+ - 149 . . fulva + . 149 C fulva ,, 144 . < discocephalus discocephalus 170 [_ aquilarum f fulva fulva 144 ' "\ d. aquilarum discocephalus . J fulva . ' "Y d. aquilarum discocephalus f fulva fulva ' \ d. aquilarum discocephalus j fulva fulva . ' \ d. aquilarum discocephalus fulva fulva r. castanea regalis 190 d. discocephalus discocephalus 168 t . regalis + regalis MS . d. discocephalus discocephalus 168 fulva+ ? gypsivorum r. subsp. punctifer fusca hopkinsi leucopleura leucopleura fulva punctifer regalis punctifer fulva (Host record needs confirmation) tenderoi rufa elani 149 197 191 196 162 164 165 149 77 REVISIONS OF MALLOPHAGO GENERA Host* Subfamily FALCONINAE Falco biarmicus . . Falco cherrug . . Falco mexicanus Falco jugger . , Falco rusticolus . Falco peregrinus Falco subbuteo . . Falco cuvierii . . Falco eleonorae . Falco concolor Falco hypoleucos Falco fuscocaerulescens Falco columbarius Falco ardosiaceus . Falco vespertinus . Falco amurensis . Falco naumanni X Falco tinnunculus , Falco alopex . . X Falco sparverius . leracidea orientalis Degeeriella species Species group rufa . rufa 203 Page No. 180 rufa rufa rufa r. carruthi rufa 185 HOST RELATIONSHIPS It seems that the relationships of many of the Falconiformes are still the subject of considerable differences of opinion amongst ornithologists (Clay, 1957 : 146), and for this reason any evidence from the distribution of the parasites should be considered. However, as far as Degeeriella is concerned much of the evidence is difficult to interpret and only tentative suggestions of the relationship of the hosts can be made. Among the reasons for this are the following : (i) The rather close simi- larity of most of the species of Degeeriella, and the difficulty of judging the significance of the small character differences in relation to the time of separation of the popu- lations and thus of their hosts. (2) The difficulty of knowing which of the species groups are the most primitive. It is not yet possible to describe with any certainty the form of the primitive mallophagen head. The different genera of Ischnocera usually resemble each other in the characters of the anterior margin of the head to a greater extent in the nymph than in the adult and as both nymph and adult live in the same environment, it can be presumed that the head with the complete anterior margin (as found in the majority of nymphs) is the more primitive. It is more difficult to decide whether the primitive Ischnocera had the complete semi- circular central carina or the interrupted carina, with the two carinae passing to the anterior margin of the head. In Degeeriella the ventral carina is interrupted, but the two carinae are poorly developed anteriorly, except in the elani group and the nymphs of D. rufa. The nymph of rufa also has a dorsal preantennal suture delineating a semicircular dorsal anterior plate. It is not impossible, 204 REVISIONS OF MALLOPHAGO GENERA therefore, that the characters of the head of the fulva and discocephalus groups are secondary, even secondarily approaching the primitive condition, if the primitive Ischnoceran Mallophaga is presumed to have this type of head. Or alternatively, the species of the elani group are derived from a discocephalus type, but then it must be postulated that rufa shows a more primitive con- dition of the head in the adult than in the nymph. (3) The difficulty of dis- tinguishing between primary and secondary absences of the species of Degeeriella. Was the discocephalus type evolved on a common ancestor of Aquila and Haliaeetus after this became separated from other hawks? This would suggest a close relationship between these genera. Or did it once have a widespread distribution later becoming extinct except on these hawks? From the resemblance, perhaps of no significance, between the heads of the nymphs of fulva and the adults of discocephalus it is possible that these species were derived relatively recently from a common ancestor, suggesting an originally wider distribution for disco- cephalus. It is possible that some of the differences between the mallophagan faunas such as those of Buteo galapagoensis and B. swainsoni and the rest of Buteo is due to the extinction of a different member of an original sympatric pair (Clay, 1949 : 296). The fulva, regalis, and discocephalus types may all have been found on the ancestral Accipitridae and since become extinct on some or other of the present members of the family. However, even this may indicate relationships ; for instance the fact thai fulva is not found on any of the genera included in the Milvinae suggests that it had already become extinct (if ever present) on an ancestral stock which gave rise to these genera and thus confirms their relationships. As an indication of the relationships of the Falconiformes as accepted by at least some ornithologists the arrangement in Peters (1931) has been followed. On pp. 202-3 above is a list of hosts (arranged according to Peters) and their known species of Degeeriella together with the species groups to which these belong. It can be seen from this list that in general the distribution follows that of the arrangement of their hosts, but with some notable exceptions. The relationships between species of Degeeriella which seem to throw some light on the relationships of their hosts are discussed in the following paragraphs. 1. The Degeeriella from Elanus, Chelictinia, Elanoides and possibly Aviceda form a related group, with those from Chelictinia and Elanoides probably being the most nearly related. The species from Gamsonyx, and also possibly Falco and leracidea, all belonging to the family Falconidae, should perhaps be included in this group (Clay, 1958 : 2). The Degeeriella species from Pernis (a genus placed in the Per- ninae with Elanoides and Aviceda) are quite distinct and perhaps show affinities with those from the Milvinae. 2. The Milvinae are parasitized by a distinctive species of Degeeriella ; the suppo- sition that this may be a relic of a sympatric pair has already been mentioned, and it is therefore possible that the Milvinae are in fact rather more nearly related to the Accipitrinae than their Degeeriella suggest. 3. The Degeeriella of the Accipitrinae suggest a fairly close relationship between the members of this subfamily, especially between Aquila and Buteo ; further that there is little difference between this subfamily and the Circinae, and that Terathopius REVISIONS OF MALLOPHAGO GENERA 205 and Circaetus (but not Spilornis) of the subfamily Circaetinae are similar but rather more distinct. Buteo galapagoensis , B. swainsoni, Haliaeetus vocifer and H. leucoryphus have the same species as found on Milvus (see above). Aquila and Haliaeetus have a second species discocephalus, the resemblance of the nymphs of fulva to the adults of this species has already been mentioned and perhaps confirms the close relationship of Buteo and Aquila which is suggested by both being parasitized by fulva. 4. Few Degeeriella species are known from the Aegypiinae : Gypaetus has a distinctive species (punctifer) not closely related to any other except gypsivorum from Gyps himalayensis. This would suggest that Gypaetus and Gyps are wrongly placed between the Accipitrinae and the Circinae. Boetticher & Eichler (1954) considered that the Degeeriella species found on Aquila and Gypaetus showed a relationship between these hosts, but this was based on the erroneous assumption that discocephalus and punctifer were closely related, but the two species resemble each other only in shape. The Degeeriella of Gypohierax is a subspecies of regalis, rather near that of Haliaeetus vocifer, this suggests if no secondary infestation has taken place, that Gypohierax is wrongly placed in the Aegypiinae. 5. It seems doubtful whether the genera included in the Falconidae do in fact, form a related group. As already shown the Degeeriella from Gampsonyx and possibly also Neohierax, Falco and leracidea show a relationship to those on some of the genera included in the Elaninae and Perninae. The subfamily Polyborinae do not have any species of Degeeriella sens. sir. but are parasitized by a species of the closely related genus Acutifrons. The parasites of Microhierax and of Polihierax, belonging to the Polihieracinae, do not belong to Degeeriella and have been dealt with elsewhere (Clay, 1955). A detailed study of the other genera of Mallophaga living on the Falconiformes may give some further indications of the relationships of their hosts. ACKNOWLEDGMENTS I am indebted to a large number of people for the presentation and loan of material ; these include F. Balat, M. A. Carriker, R. Elbel, K. C. Emerson, L. Guimaraes, G. H. E. Hopkins, S. von Keler, R. Meinertzhagen, J. Tendeiro and the U.S. National Museum. I am also much indebted to Mr. G. H. E. Hopkins for advice and criti- cisms on various points and to Dr. Joao Tendeiro for much helpful co-operation. I am also grateful to the Zoological Society, Calcutta, and the Royal Entomological Society for permission to publish certain text-figures and plates which have appeared in their journals. SUMMARY The characters and distribution of the Degeeriella-complex are discussed. Degeeriella sens, str., as found on the Falconiformes, is described. Variations and artefacts, characters of taxonomic importance and the concept of the subspecies in this group are considered. A systematic survey of all known species is given followed by a key and notes on names of which the correct interpretation is doubtful. 206 REVISIONS OF MALLOPHAGO GENERA Finally some suggestions are made on possible relationships within the Falconiformes based on the distribution of the Mallophaga. REFERENCES BLAGOVESHTCHENSKY, D. I. 1956. [The structure and systematic significance of the sexual systems of the Mallophaga.] Parasitol. Shorn, zool. Inst. Akad. Nauk SSSR, Leningrad 16 : 5-88. BOETTICHER, H. VON & EiCHLER, W. IQ54- Parasitophyletische Studien zur Ornitho- systematik. II. Die Verteilung der Degeeriellidae und Falcolipeuridae bei den Accipitres. Biol. Zbl. 73 : 212-221. BROWN, W. L. & WILSON, E. O. 1956. Character displacement. Syst. Zool. 5 : 49-64. CARRIKER, M. A. 1956. Neotropical Mallophaga Miscellany, No. 9. Rev. Brasil. Ent. 5 : 111-146. 1957- Notes on some of the Kellogg types of Mallophaga. Microentomology, 22 : 95-110. CLAY, T. 1949. Some problems in the evolution of a group of ectoparasites. Evolution, 3 : 279-299. 1951. An introduction to a classification of the Avian Ischnocera (Mallophaga). Trans. R. ent. Soc. Land. 102 : 171-194. 1953. Revisions of the Genera of Mallophaga. I. The .ffa/fo'eo/a-complex. Proc. zool. Soc. Lond. 123 : 563-587. 1955- Revisions of the genera of Mallophaga. Colilipeurus and a new genus. Trans. R. ent. Soc. Lond. 107 : 169-186. 1956. Phthiraptera. In Tuxen, Taxonomist's Glossary of Genitalia in Insects : 145-148. Copenhagen. 19570. Degeeriella parasitic on Pernis. Proc. zool. Soc. Calcutta, Mookerjee Memor. vol. : 339-347- 19576. The Mallophaga of Birds. In First Symposium on Host Specificity among Para- sites of Vertebrates, Neuchatel : 120-154. 1958. Three new species of Degeeriella from the Falconiformes. Proc. R. ent. Soc. Lond. (B) 27 : 1-7. CLAY, T. & HOPKINS, G. H. E. 1954. Tne early literature on Mallophaga. Bull. Brit. Mus. (Nat. Hist.} Entom. 3 : 223-266. 1955- Notes on the Rudow Collection of Mallophaga at Hamburg. Mitt. Hamburg Zool. Mus. Inst. 53 : 49-73. DOBZHANSKY, TH. 1951. Genetics and the origin of species, 3rd ed. Columbia University Press, New York. EICHLER, W. 19410. Uber die Mallophagen von Lammergeier und Himalayageier. Schweiz. Arch. Tierheilk. 83 : 178-181. 19416. Zur Klassifikation der Lauskerfe. Arch. Naturgesch., Leipzig (N.F.) B. 10 : 345-398. HOPKINS, G. H. E. 1947. Notes on Mallophagen Nomenclature. II. Entomologist, 80 : 73-79- 1949. Host-associations of the lice of mammals. Proc. zool. Soc. Lond. 119 : 387-604. JORDAN, K. 1896. On mechanical selection and other problems. Novit. zool. 3 : 426-525. KELER, VON S. 1939. Baustoffe zu einer Monographic der Mallophagen. II. Uberfamilie Nirmoidea. Nova Acta Leop. -Carol. (N.F.), 8 : 1-254. MAYR, E. 1951. Speciation in birds. Proc. Xth Intern. Ornith. Congress, 1950 : 91-131. MAYR, E., LINSLEY, E. G. & USINGER, R. L. 1953. Methods and Principles of Systematic Zoology. New York. NEUMANN, L. G. 1922. Mallophages. In Voyage de M. le Baron de Rothschild en Ethiopie et en Afrique Orientale Anglaise, 1904-05. Resultats scientifiques, Animaux articules, 1 : 22-240. REVISIONS OF MALLOPHAGO GENERA 207 PETERS, J. L. 1931. Check-list of Birds of World. Cambridge, U.S.A. SCHMUTZ, W. 1955. Zur Konstruktionsmorphologie des mannlichen Geschlechtsapparates der Mallophagan. Zoll. Jb. (Anatomie), 74 : 189-338. STRINDBERG, H. 1918. Typstudien uber die Geschlectsorgane einiger Mallophagengattungen. Z. wissZool. 117 : 591-653. TENDEIRO, J. 1955. Estudos sobre uma colec9io de Malofagos de Aves. Bol. cult. Guine Portuguesa (1954). 9 (35) : 497~ 62 5- TJONNELAND, A. 1955- A comparison of the variation coefficients of some measurements of Degeeriella aquilarum Eichler (Mallophaga) . Naturvitenskapelig rekke, 10 Univ. Bergen : 4-9. WATERSTON, J. 1928. Mallophaga of the Sand-grouse. Proc. zool. Soc. Land. : 333-356. PLATE i FIGS. 1-7. Degeeriella fulva (<$($} from various hosts : FIG. i. Aquila chrysaetos. FIG. 2. Buteo lagopus. FIGS. 3-4. Aquila wahlbergi. FIG. 5. Buteo buteo. FIG. 6. Buteo harlani. FIG. 7. Buteo jamaicensis. PLATE 2 FIG. i. Degeeriella africana. FIG. 2. Degeeriella carrikeri. FIG. 3. Degeeriella n. haydocki. PLATE 3 FIG. i. Degeeriella n. nisus. FIG. 2. Degeeriella n. vagans. FIG. 3. Degeeriella n. f rater. Lectotype. PLATE 4 FIG. i. Degeeriella leucopleura. FIG. 2. Degeeriella hopkinsi. FIG. 3. Degeeriella fusca. PLATE 5 FIG. i. Degeeriella r. regalis from Milvus milvus. FIG. 2. Degeeriella r. deignani. FIG. 3. Degeeriella r. castanea. PLATE 6 FIG. i. Degeeriella elani. FIG. 2. Degeeriella tendeiroi. FIG. 3. Degeeriella rufa from Falco tinnunculus. PLATE 7 FIG. i. Degeeriella guimardesi. FIG. 2. Degeeriella meinertzhageni. FIG. 3. Degeeriella elbeli. FIG. 4. Male genitalia of fig. i . FIG. 5. Male genitalia of fig. 2. FIG. 6. Male genitalia of fig. 3. PLATE 8 FIG. i. Degeeriella phlyctopygus. FIG. 2. Degeeriella mookerjeei. FIG. 3. Degeeriella fulva from Aquila chrysaetos, male genitalia. FIG. 4. Degeeriella beaufacies, male genitalia. FIG. 5. Degeeriella n. nisus, male genitalia. FIG. 6. Degeeriella fusca, male genitalia. FIG. 7. Degeeriella rufa from Falco tinnunculus, male genitalia. PLATE 9 FIG. i. Degeeriella d. discocephalus from Haliaeetus albicilla. FIG. 2. Degeeriella d. aquilarum from Aquila nipalensis. FIG. 3. Degeeriella punctifer. Photographs by J. V. Brown British Museum (Natural History) Bull. B.M. (N.H.) Entom. 7, 4 PLATE i Bull. B. M.(N.H.} Entom. 7, 4 PLATE 2 i Ifc Bull. B.M. (N.H.) Entom. 7, 4 PLATE 3 Bull. B.M. (N.H.) Entom. 7, 4 PLATE 4 Bull. B.M. (N.H.) En torn. 7, 4 PLATE 5 Bull. B.M. (N.H.) Entom. 7, 4 PLATE 6 Butt. B.M. (N.H.) Entom. 7, 4 PLATE 7 Bull. B.M. (N.H.), Entom. 7, 4. PLATE 8. I M - 6 Bull. B.M. (N.H.) Entom. 7, 4. PLATE g. REVISION DU GENRE EXOCENTRUS MULSANT (COL., CERAMBYCIDAE) S. BREUNING BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol. 7 No. 5 LONDON: 1958 ... REVISION DU GENRE EXOCENTRUS MULSANT (COL., CERAMBYCIDAE) PAR S. BREUNING Pp. 209-328; 5 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Vol 7 No. 5 LONDON: 1958 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in Jive series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 7, No. 5 of the Entomological series. Trustees of the British Museum, 1958 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued October, 1958 Price Two Pounds REVISION DU GENRE EXOCENTRUS MULSANT p: (COL., CERAMBYCIDAE) Par S. BREUNING Genus EXOCENTRUS Mulsant Exocentrus Mulsant, 1839, Col. Fr., Long. : 152. Exocentrus + Camptomyme Pascoe, 1864, Trans, ent. Soc. Land. (3) 131 : 27, 43. Oligopsis Thomson, 1864, Syst. Cer. : in. Exocentrus Mulsant, Bates, 1866, Ann. Mag. nat. Hist. (3) 17 : 191. Exocentrus + Oligopsis + Camptomyme Lacordaire, 1872, Gen. col. 9 : 800, 801, 805, 806, 815. Exocentrus Mulsant, Ganglbauer, 1884, Best. Tab. 7 : 692. Ispaterus Fairmaire, 1892, Rev. d'Ent. 11 : 122. Pseudocentms Fairmaire, 1901, Rev. d'Ent. 20 : 230. Exocentrus Mulsant, Matsushita, 1933, Journ. Fac. Agr. Hokk. 34 : 389. Exocentrus Mulsant, Gressitt, 1939, Lingn. Sc. Journ. 18 : 82 ; 1940, Philipp. Journ. Sc. 72 : 182, J 83 ; 1951, Longicornia, 2 : 518. Exocentrus Mulsant, Lepesme & Breuning, 1955, Bull. Soc. ent. Fr. 60 : 127. EN ovale allonge ou allonge. Antennes normalement peu fines, un peu moins longues que le corps a moderement plus longues, frangees en dessous de poils longs ; le scape long et mince, le troisieme article plus ou moins long que le quatrieme ou que le scape, les articles a partir du cinquieme diminuant progressivement en longeur. Tubercules antenniferes non ou peu saillants. Yeux assez grossierement facettes et fortement echancres, les lobes inferieurs grands. Pronotum transverse, pourvu d'une epine laterale recourbee. Elytres plus larges que le pronotum, moderement convexes, normalement arrondis a 1'apex, pourvus souvent de points serialement disposes. Tte retractile. Saillie prosternale etroite, moins haute que les bandies, arquee. Saillie mesosternale peu large, inclinee vers 1'avant. Metasternum de longueur normale. Cavites coxales intermediaries fermees. Pattes de longueur moyenne ; femurs claviformes ; tibias intermediates pourvus d'un leger sillon dorsal ; crochets divariques. Tout les corps, les pattes et les antennes herisses de poils dresses longs. Type : Exocentrus lusitanus L. 259 especes, repandues par 1'Afrique, 1'Europe, 1'Asie et les lies orientales vers Test jusqu'aux lies Carolines, la Nouvelle Guinee et le Queensland. Mulsant, lors de sa description du genre Exocentrus, y avait place deux especes, cinereus Muls. et lusitanus L. Dans sa deuxieme edition de ses Coleopteres de France parue en 1863 il avait place lui meme le cinereus dans son genre Oplosia. Ainsi lusitanus L. devient automatiquement par elimination le type du genre Exocentrus. ENTOM. 7. 5. II 212 REVISION DU GENRE EXOCENTRUS MULSANT Les especes suivantes decrites comme appartenant au genre Exocentrus Muls. sont a exclure du genre : Uoculatus Matsumura & Matsushita, binhanus Pic, fujiyamai Matsumura & Matsushita, inhirsutus Pic, leiopodinus Marsushita, meridianus Ohbayashi, saipanensis Ohbayashi et tonsus Bates, ces huit especes se pla9ant dans le genre Aegocidnus Pascoe. antennalis Jordan et polymitus Jordan, ces deux especes se plagant dans le genre Jordanoleiopus Breuning. elegans Fairmaire appartenant au genre Parhoplomelas Breuning. inermis Gahan appartenant au genre Acartus Fahraeus comme synonyme de hirtus Fahraeus. aurovilliusi Fisher appartenant au genre Paroligopsis Breuning. hirsutulus Faldermann, espece douteuse. 1 setosus Serville appartenant au genre Cosmoiomoides Melzer. nitidulus Bates, pusillus Blanchard et setosus Serville appartenant a des genres americains. Les genres Camptomyme Pascoe, Oligopsis Thomson et Pseudocentrus Fairmaire ne forment pas des genres a part mais peuvent e'tre conserves comme noms de sous- genres. Je partage le genre Exocentrus en dix sousgenres me basant surtout aussi sur la longueur relative des premiers articles antennaires. Ces sousgenres ne forment souvent pas des groupes phylogeniquement homogenes mais facilitent le partage des tres nombreuses especes du genre qui ne sont generalement pas separes par des carteres structurels importants. Ispaterus Fairmaire est un synonyme d'Oligopsis Thomson. TABLEAU DE DETERMINATION DES SOUSGENRES 1 . Troisieme article des antennes moins long que le quatrieme .... 2 - Troisieme article des antennes tout au moins aussi long que le quatrieme . . 5 2. Chaque elytre etire en une epine apicale Sg. 10, Dentexocentrus Breuning (une espece, p. 226) - Elytres arrondis a 1'apex .......... 3 3. Troisieme article des antennes moins long que le scape Sg. 3, Centenexocentrus Breuning (Tableau, p. 221) - Troisieme article des antennes plus long que le scape ..... 4 4. Antennes fines ; les points des elytres disposes sans ordre Sg. 5, Woodlarkexocentrus Breuning (une espece, p. 221) - Antennes peu fines ; les points des elytres serialement disposes Sg. 4, Barbierexocentrus Breuning (une espece, p. 221) 5. Troisieme article des antennes aussi long que le quatrieme ..... 6 - Troisieme article des antennes plus long que le quatrieme . . . . .7 1 Astyonomus hirsutulus Faldermann, 1837, Fna. Transcauc. 2 : 272, pi. 8, fig. 5. Astyonomus hirsutulus Ganglbauer, 1884, Best. Tab. 8 : 535. Exocentrus hirsutulus Plaviltschikov, 1926, Encycl. Ent. 1, Col. 2 : 60. Plaviltschikov dit au sujet de cette espece (I.e.) : " Exocentrus hirsutulus Faldermann est sans doute un Exocentrus mais d'apres la description et la figure donnees par Faldermann il est impossible de decider a quelle espece de ce genre on doit attribuer cette description (probablement a \' Exocentrus adspersus Mulsant). Je propose de ne pas citer cette espece dans la liste des especes decrites et de la prendre comme un nomen nudum (a cause de sa description insuffisante)." REVISION DU GENRE EXOCENTRUS MULSANT 213 6. Les points des 61ytres sont plus ou moins aligned, sauf parfois dans le quart sutural Sg. 2, Camptomyme Pascoe (Tableau, p. 216) - Les points des elytres sont disposes sans ordre Sg. i, Exocentrus Mulsant s. str. (Tableau, p. 213) 7. Troisieme article des antennes un peu plus long que le quatrieme Sg. 6, Pseudocentrus Fairmaire (Tableau, p. 221) - Troisieme article des antennes beaucoup plus long que le quatrieme ... 8 8. Chaque 61ytre pourvu d'une petite bosse discale postbasilaire Sg. 9, Tuberculexocentrus Breuning (une 6spece, p. 323) - Elytres sans trace d'une bosse discale ........ 9 9. Les points des elytres sont srialement disposes sauf parfois sur le tiers sutural Sg. 7 Oligopsis Thomson (Tableau, p. 225) - Les points des elytres sont disposes sans ordre Sg. 8, Formosexocentrus Breuning (une 6spece, p. 322) TABLEAU DE DETERMINATION DES ESPECES i. Sousgenre EXOCENTRUS Mulsant s.s. 1. Elytres parsemds de tres petites taches blanches, rangers longitudinalement. . 2 - Elytres sans taches semblables ......... 7 2. Ces taches ne sont pas rangers en series longitudinales r6gulieres albomaculatus Pic (p. 234) - Ces taches sont rangers en series longitudinales r^gulieres ..... 3 3. Ces taches sont disposers en outre sous forme de deux bandes transversales, une pr6m6diane et une m6diane ..... enganensis Breuning (p. 233) - Ces taches ne sont pos disposers en forme de bandes transversales ... 4 4. Ces taches sont dispose 1 es en cinq series longitudinales sur chaque elytre lachrytnosus Pascoe (p. 232) - Ces taches sont disposers en sept series longitudinales sur chaque elytre . . 5 5. L'6pine Iat6rale du pronotum faiblement recourbe'e . . lacteolus Gahan (p. 238) - L'6pine lat^rale du pronotum fortement recourb^e ...... 6 6. Cette epine est pre"cale du pronotum pre'ce' de"e d'une bosse lat^rale obtuse rufus Breuning (p. 257) - L'e'pine late"rale du pronotum non pr6ce"d6e d'une bosse . subrufus Breuning (p. 258) 7. Pronotum rouge unicolore ; Elytres sans dessins ...... 8 Pronotum d'une autre couleur ......... 12 3. Tefp brun fonce" ....... binaluensis Breuning (p. 280) 9 philippinensis unicolor Fisher (p. 281) 10 10. Elytres violets metalliques ...... callioides Pascoe (p. 282) Elytres non metalliques . . . . . . . . . . .11 11. Elytres noirs. ......... bicolor Pascoe (p. 282) Elytres brun fonc6 avec deux bandes transversales peu apparentes explanatidens Pic (p. 271) 12. Presque unicolore noir ........... 13 Jamais presque unicolore .......... 19 13. Lobes infe'rieurs des yeux aussi longs que les joues . . . tristis Pascoe (p. 282) Lobes infe'rieurs des yeux plus longs que les joues . . . . . . 14 14. Les points des eUytres ne sont pas aligned sur le quart sutural . . . . 15 Les points des e'lytres aligne"es aussi sur le quart sutural . . . . . 16 15. Partie basilaire des femurs rouge ..... femoralis Hintz (p. 250) Partie basilaire des femurs noire ..... niger Breuning (p. 257) 1 6. Lobes infe'rieurs des yeux beaucoup moins de deux fois plus longs que les joues . 17 Lobes inf6rieurs des yeux deux fois plus longs que les joues . . . . 18 17. L'6pine lat^rale du pronotum pre^de'e d'une bosse Iat6rale obtuse armatus Hintz (p. 249) Lupine late>ale du pronotum non pr6ce'de d'une bosse . subnitens Breuning (p. 250) 1 8. L'e'pine Iat6rale du pronotum fortement recourbe'e . subarmatus Breuning (p. 249) L'e'pine lat^rale du pronotum faiblement recourbe'e maiae Lepesme & Breuning (p. 247) 19. Tete et pronotum noirs, elytres brun jaunatres . . subglaber Fisher (p. 279) Corps autrement colori6 . . ........20 20. Elytres brun fonc6, la region hume'rale et la region apicale rouges humeralis Aurivillius (p. 279) Autrement colorte . .,,,,.. 21 REVISION DU GENRE EXOCENTRUS MULSANT 217 21. Elytres cou verts de pubescence gris blanchatre exception faite de deux bandes trans versales glabres (brun fonce), une premediane et une postmediane subbidentatus Gressitt (p. 273) Elytres autrement pubescents . . . . . . . . .22 22. Rouge. Elytres ornes de taches brun fonce vagues . vagemaculatus Breuning (p. 257) Autrement colorie ............ 23 23. Elytres couverts de pubescence gris blanchatre et ornes de taches brunes floues ciliatissimus Gressitt (p. 283) Autrement colorie ............ 24 24. Elytres ornes de taches quadrangulaires jaune pale, rangees sous forme de deux tres larges bandes transversales, une premediane et une postmediane ... 25 Elytres autrements pubescents ......... 27 25. Les points des elytres sont assez irregulierement disposes sur le tiers sutural submisellus Breuning (p. 276) Les points des elytres sont serialement disposes aussi sur le tiers sutural . . 26 26. Les taches de la deuxieme bande restent distantes de 1'apex misellus Lameere (p. 275) Les taches de la deuxieme bande s'etendent jusqu'a 1'apex de 1'elytre miselloides Breuning (p. 282) 27. Chaque elytre avec trois bandes transversales jaunes ...... 28 Elytres sans ces bandes ........... 29 28. Sur chaque elytre une bande postbasilaire, une postmediane et une preapicale hupehensis Gressitt (p. 273) Sur chaque elytre une bande postbasilaire, une premediane et une postmediane trifasciellus Gressitt (p. 272) 29. Sur chaque elytre deux bandes transversales ondulees jaunes .... 30 Elytres sans ces bandes ........... 31 30. Sur chaque elytre une bande premediane et une preapicale gedeensis Breuning (p. 278) Sur chaque elytre une bande postbasilaire et une mediane javaensis Breuning (p. 278) 3 1 . Elytres ornes de bandes transversales blanchatres ou gris clair ou du moins de petites taches blanchatres ou gris clair disposees sous forme de bandes transversales . 32 Elytres sans bandes et sans taches semblables ....... 65 32. Elytres rouges, revetus de pubescence jaune paille et orn6s d'une seule bande trans- versale ondulee blanche postmediane . . . univittatus Breuning (p. 247) Elytres autrement dessin6s .......... 33 33. Elytres ornes de nombreuses petites taches ovalaires blanches ne formant pas de bandes nettes ............ 34 Elytres autrement dessines .......... 35 34. Lobes inferieurs des yeux de moitie plus longs que les joues moerens Pascoe (p. 275) Lobes inferieurs des yeux quatre fois plus longs que les joues submoerens Breuning (p. 274) 35. Elytres sans taches ou bandes blanches dans la moitie apicale .... 36 Elytres avec des taches ou des bandes blanches dans la moiti6 apicale . . 43 36. Antennes rouges ............ 37 Antennes brun fonc6 sauf parfois les premiers articles . . . . . 38 37. Les deux bandes transversales blanches de 1'elytre, la postbasilaire et la mediane, reliees pres de la suture fernandopoanus Baguena & Breuning . (p. 248) Ces deux bandes non reliees pres de la suture . . . ruftcornis Hintz (p. 252) 38. Les trois premiers articles antennaires rouge clair . . . nitens Jordan (p. 250) Les trois premiers articles antennaires brun fonce ...... 39 39. Partie basilaire des articles antennaires a partir du troisieme a pubescence blanchatre pseudonitens Breuning (p. 266) Partie basilaire des ces articles sans pubescence blanchatre .... 40 40. Bord lateral du pronotum largement aplati et reborde . explanatidens Pic (p. 271) Bord lateral du pronotum non largement aplati et reborde . . . . 41 2i8 REVISION DU GENRE EXOCENTRUS MULSANT 41. Ranches et partie basilaire des femur, noirs chevangeoni Lepesme & Breuning (p. 247) Hanches et partie basilaire des fe"murs, rouges ....... 42 42. L'6pine Iat6rale du pronotum pr6c6d6e d'une bosse Iat6rale obtuse nigrescens Breuning (p. 258) Lupine late>ale du pronotum non pr6c6d6e d'une bosse schoutedeni Breuning (p. 259) 43. Les taches blanches forment dans la moiti6 post^rieure de 1'elytre un grand demi- cercle dont le point le plus convexe touche la suture multiguttulatus Pic (p. 275) Elytres autrement dessin6s .......... 44 44. Une s6rie de taches nettes jaune blanchatre longe la suture dans le tiers apical de l'61ytre ........ sumatranus Breuning (p. 276) Elytres sans ces taches . . . . . . . . . . .45 45. Les taches e'lytrales forment une seule bande transversale ondutee m6diane . . 46 Les taches 61ytrales forment plusieures bandes transversales .... 47 46. Les poils dresses des Elytres sont noirs .... nonymoides Jordan (p. 254) Les poils dresse's des 61ytres sont dor6s . . . aureopilosus Breuning (p. 267) 47. Les taches 61ytrales forment deux ou trois bandes transversales . . . . 48 Les taches 61ytrales forment tout au moins quatre bandes transversales, dont deux sont situ^es dans la moitie" apicale . . . . . . .64 48. Couleur fondamentale de l'61ytre brun rougeatre dans la moitie" basilaire et noire dans la moiti6 apicale ...... basirufus Gressitt (p. 272) Elytres autrement colori^s .......... 49 49. Sur chaque elytre une bande postbasilaire, une m6diane et une pr^apicale . . 50 Sur chaque 61ytre une bande basilaire ou postbasilaire, une pr6m6diane et une postmdiane ............ 54 50. Elytres tres grossierement ponctue's . . grossepunctatus Breuning (p. 261) Elytres assez finement ponctu6s . . . . . . . . .51 51. Sur chaque 61ytre apres la bande pr6apicale une tache apicale jaune multivittatus Breuning (p. 256) Elytres sans tache apicale jaune ......... 52 52. Chaque 61ytre avec une troite bande suturale blanche sur le quart basilaire birmanus Breuning (p. 271) Elytres sans une bande pareille ......... 53 53. Ecusson couvert de pubescence blanche . . . alboscutellaris Breuning (p. 271) Ecusson sans pubescence blanche ..... carissae Fisher (p. 270) 54. La bande pre'm^diane remonte obliquement en direction de l'6paule 55 La bande pr^me'diane ne remonte pas obliquement en direction de l'6paule . . 56 55. La deuxie mebande de 1'elytre est une bande postm^diane rhodesianus Breuning (p. 262) La deuxieme bande de 1'elytre est une bande pr6apicale . conradti Breuning (p. 251) 56. Pronotum rouge ; Elytres brun fonc6, les bandes peu apparentes . . . 57 Autrement colorie' ............ 59 57. Tete rouge ......... explanatidens Pic (p. 271) Tete noire ............. 58 58. L'6pine lat^rale du pronotum fortement recourb6e . . vaneyeni Breuning (p. 259) L'dpine Iat6rale du pronotum faiblement recourbe'e . vagesticticus Breuning (p. 259) 59. L'e"pine lat^rale du pronotum faiblement recourb^e . . . . . 60 L'6pine lat^rale du pronotum fortement recourb6e . . . . . . 61 60. La bande postme'diane de 1'elytre est fortement ondu!6e subinclusus Hunt & Breuning (p. 264) La bande postm6diane de l'61ytre est droite subinclusus latefasciatus Hunt & Breuning (p. 265) 61. La bande postm^diane de l'61ytre est peu large et ondu!6e ..... 62 La bande postm6diane de l'61ytres est large et droite ... . . . 63 62. Corps en majeure partie brun fonc6 . . substrigosus Hunt & Breuning (p. 263) Corps entierement rouge clair ...... minimus Breuning (p. 248) REVISION DUGENRE EXOCENTRUS MULSANT 219 63. Chaque elytre avec trois bandes transversales . . . strigosus Jordan (p. 251) Chaque elytre avec seulement deux bandes transversales latefasciatipennis Breuning (p. 248) 64. L'epine laterale du pronotum fortement recourbee et precedee d'une bosse laterale obtuse ......... sjostedti Breuning (p. 265) L'epine laterale du pronotum faiblement recourbee et non precedee d'une bosse laterale ......... decorsei Breuning (p. 249) 65. Elytres ornes de bandes longitudinales grises interrompues par places insularis Fisher (p. 281) Elytres autrement dessines .......... 66 66. Elytres ornes de nombreuses etroites bandes longitudinales blanches on jaune pale 67 Elytres sans bandes pareilles .......... 73 67. Les bandes blanches confluent plus ou moins lateralement sur la moiti6 laterale de 1'elytre ......... kalshoveni Fisher (p. 277) Les bandes non confluent pas .......... 68 68. Sur chaque elytre neuf ou dix bandes longitudinales ...... 69 Sur chaque elytre sept bandes longitudinales ....... 70 69. Pronotum rouge ....... binaluensis Breuning (p. 280) Pronotum non rouge albostriatus Hintz (p. 255) 70. L'epine Iat6rale du pronotum precedee d'une bosse Iat6rale obtuse leucostriatus Breuning (p. 255) L'epine laterale du pronotum non precedee d'une bosse laterale .... 71 71. Troisieme article des antennes sensiblement moins long que le scape vittatus Fisher (p. 269) Le troisieme article des antennes tout au plus un peu moins long que le scape . 72 72. Tiers apical des tibias brun fonce ..... lineolatus Muller (p. 269) Tiers apical des tibias non brun fonce . . . leucolineatus Breuning (p. 256) 73. Tier sutural de 1'elytre couvert d'une pubescence d'un brun assez fonce : les deux tiers lateraux revetus de pubescence soyeuse brun clair ..... 74 Elytres autrement pubescents ......... 75 74. L'epine laterale du pronotum precedee de deux petites preeminences anguleuses laterales ......... costatus Breuning (p. 277) L'epine laterale du pronotum non precedee de preeminences fuscovittatus Breuning (p. 267) 75. Sur chaque elytre une bande transversale premediane jaunatre qui remonte obliquement en direction de 1'epaule . . . flavofasciatus Breuning (p. 260) Elytres sans une bande semblable ......... 76 76. Sur chaque elytre une bande transversale preapicale ochracee .... 77 Elytres sans cette bande . . . . . . . . . .78 77. Pronotum entierement couvert de pubescence rousseatre . basilanus Breuning (p. 280) Pronotum sans une pubescence pareille . . . acutispina Fairmaire (p. 283) 78. Elytres ornes de taches gris clair ou blanches, disposees en series longitudinales . 79 Elytres sans taches semblables . . . . . . . . .81 79. Elytres ornes en plus de quelques taches ochracees vives aureomaculatus Aurivillius (p. 262) Elytres sans taches ochracees .......... 80 80. Les taches elytrales blanches sont etroites . . . drescheri Fisher (p. 278) Les taches elytrales blanches sont oblongues . . . angusticollis Fisher (p. 280) 81. Sur chaque elytre sept series longitudinales de petites taches quadrangulaires blanchatres faisant defaut sur l'emplacement d'une large bande' postmediane ondu!6e transversale brun rougeatre .... sumatrensis Fisher (p. 276) Elytres autrement dessines .......... 82 82. Elytres ornes de petites taches jaune pale, rangees en s6ries longitudinales . . 83 Elytres sans taches pareilles . . . . . . . . . &7 220 REVISION DU GENRE EXOCENTRUS MULSANT 83. Sur chaque 61ytre, en plus une bande transversale postmMiane ondutee brun fonc6 et une tache brun fonc6 a cot6 de l'6cusson fuscosignatipennis Hunt & Breuning (p. 263) Elytres sans ces dessins ........... 84 84. Scape rouge clair ............ 85 Scape rouge fonc6 ou brun noir ..... chatter jeei Fisher (p. 270) 85. Articles antennaires 7 a ii rouges ..... seriatus Jordan (p. 252) Articles antennaires 7 a n brun fence" ........ 86 86. Pronotum deux fois plus large que long ..... Mrtus Fisher (p. 279) Pronotum moins large ...... subseriatus Breuning (p. 263) 87. Elytres orn6s de tres nombreuses taches quadrangulaires blanches qui couvrent pratiquement toute leur surface exception faite de taches minimes d6nude"es variegatus Duvivier (p. 260) Elytres autrement dessine's .......... 88 88. Elytres erne's de nombreuses petites taches blanches ...... 89 Elytres sans ces taches ........... 91 89. Des taches pareilles se trouvent aussi sur le tiers apical albosignatus Lepesme & Breuning (p. 253) Pas de taches blanches sur le tiers apical de 1'elytre ...... 90 90. Les deux premiers articles antennaires sont rouge clair . alluiiudi Breuning (p. 267) Les deux premiers articles antennaires sont brun fonc6 albostictipennis Breuning (p. 260) 91. Elytres revetus de pubescence jaune blanchatre et gris blanchatre et orn6s d'une bande transversale postm6diane brune ........ 92 Elytres sans une bande semblable . . . . . . . . .103 92. Elytres grossierement ponctu6s ......... 93 Elytres assez finement ponctu6s ......... 95 93. Tibias unicolores rouges ..... rufotibialis Breuning (p. 246) Tibias brun fonc6 dans la moitie" apicale et rouge clair dans la moiti6 basilaire . 94 94. Epaules rouge clair ....... echinulus Gahan (p. 261) Epaules non rouges ....... exiguus Gahan (p. 262) 95. Sur chaque lytre une large bande longitudinale lat^rale brun fonc6 parcourant les deux tiers ant^rieurs ........... 96 Elytres sans cette bande .......... 97 96. Moiti6 basilaire des articles antennaires a partir du troisieme a pubescence blanche orientalis Breuning (p. 265) Seule 1' extreme base des articles antennaires a partir du troisieme a pubescence blanche ........ guineensis Breuning (p. 246) 97. Premiers articles antennaires rouge clair . . . rufobasicornis Breuning (p. 258) Premiers articles antennaires non rouge clair ....... 98 98. La bande postm^diane brune de I'dlytre n'est pas ondulde ..... 99 La bande postme'diane brune de 1'eTytre est ondule . . . . . .100 99. Pronotum avec deux taches dories .... latevittipennis Breuning (p. 256) Pronotum sans taches dories ...... jeanneli Breuning (p. 266) 100. Bord lateral de I'e'lytre garni de poils dresse's pales ...... 101 Bord lateral de I'e'lytre garni de poils dresse's noirs . . . . . .102 1 01. Lobes infe'rieurs des yeux trois fois plus longs que les joues raffrayi Breuning (p. 268) Lobes infe'rieurs des yeux a peine deux fois plus longs que les joues gardnerianus Breuning (p. 266) 102. La bande postm6diane fonc6e de I'e'lytre interrompue par de petites taches circu- laires jaunatres fasciolatus Bates (p. 274) La bande postm6diane fonce'e de I'e'lytre non interrompue par des taches subfasciatus Jordan (p. 253) 103. Elytres converts d'une pubescence jaune pale ou brun jaunatre sauf une tres grande tache lat^rale m^diane ou postm6diane brune . . . . . .104 REVISION DU GENRE EXOCENTRUS MULSANT 221 Elytres revetus de pubescence brun clair et orn6s de taches plus fonc^es sudanicus Aurivillius (p. 268) 104. Lupine Iat6rale du pronotum faiblement recourb^e . pseudomurinus Breuning (p. 269) L'6pine lat^rale du pronotum fortement recourb^e . . . . . .105 105. Tete noire ............. 106 Tete rouge ............. 107 106. Pronotum noir ........ nigricollis Hintz (p. 269) Pronotum brun rouge ...... constricticollis Gressitt (p. 273) 107. Les points des 61ytres r6gulierement aligned dans le quart sutural ficicola Fisher (p. 270) Les points des Elytres non aligned dans le quart sutural . murinus Breuning (p. 264) 3. Sousgenre CENTENEXOCENTRUS nov. 1. Sur chaque 61ytre une grande tache postme"diane d6nud6e ..... 2 Elytres sans cette tache .......... 3 2. Tiers apical des tibias a pubescence brun fence" . . centenes Pascoe (p. 286) - Tiers apical des tibias sans cette pubescence . . centenoides Breuning (p. 286) 3. Elytres erne's de dessins blanchatres sauf sur I'emplacement d'une tache ou bande transversale postme'diane brune ......... 4 - Elytres autrement dessin^s .......... 9 4. Pronotum avec deux grandes taches discales jaunes . hispidulus Pascoe (p. 284) - Pronotum sans ces taches .......... 5 5. Lupine lat^rale du pronotum fortement recourbe ...... 6 - L'e'pine late"rale du pronotum seulement assez faiblement recourb6e ... 8 6. L'6pine late'rale du pronotum dirig^e directement vers 1'arriere tnindanaoensis Fisher (p. 286) - L'e'pine Iat6rale du pronotum dirig^e obliquement vers I'arriere .... 7 7. Pronotum deux fois plus large que long . . seriatopunctatus Aurivillius (p. 285) - Pronotum faiblement transverse ..... artocarpi Fisher (p. 286) 8. Troisieme article des antennes aussi long que le scape, hispiduloides Breuning (p. 285) - Troisieme article des antennes sensiblement moins long que le scape neopomerianus Breuning (p. 285) 9. Les deux premiers articles antennaires brun noir . bicoloripennis Breuning (p. 287) - Les deux premiers articles antennaires rouge clair rufobasiantennalis Breuning (p. 287) 4. Sousgenre BARBIEREXOCENTRUS nov. Une seule espece ........... barbieri Pic (p. 288) 5. Sousgenre WOODLARKEXOCENTRUS nov. Une seule espece ........ woodlarkianus Breuning (p. 288) 6. Sousgenre PSEUDOCENTRUS Fairmaire 1. Elytres d'un bleu m^tallique, rouge clair sur une tache hum6rale et sur une tache apicale ........ rufohumeralis Breuning (p. 290) - Elytres autrement colori^s .......... 2 2. Elytres unicolores ou tout au moins sans dessins ...... 3 - Elytres presentant tou jours des dessins . . . . . . 22 3. Elytres rouges, mais a peu pres la moitie' suturale brun fonc6 ou noire. . . 4 - Elytres autrement colori6s .......... 5 4. Toute la moiti6 suturale de 1'eTytre noire . . parterufipennis Breuning (p. 291) - Seuls les deux tiers ant&rieurs de la moiti6 suturale de 1'elytre brun fonc6 flsheri Gressitt (p. 294) 5. Elytres noirs ou brun fonc6, tout au moins en majeure partie .... 6 222 REVISION DU GENRE EXOCENTRUS MULSANT - Elytres rouges ou tout au moins plus clairs . . . . . . . 13 6. Partie basilaire de 1'elytre rouge ......... 7 - Partie basilaire de 1'elytre egalement brun fonce ou noir ..... 8 7. Seul le huitieme basilaire de 1'elytre rouge testaceus rufobasipennis Breuning (p. 300) - Tout le cinquieme basilaire de 1'elytre rouge testaceus rufoampliatus Breuning (p. 300) 8. D'un brun rougeatre fonc6 unicolore .... blbtei Breuning (p. 290) - D'une autre couleur ............ 9 9. Tete noire .............. 10 - Tete tout au moins partiellement rouge . . . . . . . . n 10. Pronotum rouge a fine pubescence dor6e .... indicola Fisher (p. 306) Pronotum rouge fonc6 sans pubescence doree . . . rufithorax Gressitt (p. 294) 11. Antennes unicolores brun noir .... tnindoroensis Breuning (p. 291) Articles antennaires anneles sur la partie basilaire de pubescence gris blanchatre . 12 12. Front et joues d'un brun noir .... testaceus diver siceps Pic (p. 300) Front et joues rouges ..... testaceus subbicolor Breuning (p. 300) 13. Pronotum brun fonc6 ou brun noir ......... 14 Pronotum rouge ............ 19 14. Elytres rouges, brun fonc6 pres de la suture sauf en arriere testaceus lateraloides Breuning (p. 300) Elytres unicolores ou presque .......... 15 15. Elytres d'un brun assez fonce, vaguement marbres de blanc crassepunctus Lepesme & Breuning (p. 313) Elytres brun rougeatre clair ou bruns ou brun jaunatre . . . . . 16 1 6. Elytres grossierement ponctues . . . mirei Lepesme & Breuning (p. 312) Elytres fmement ponctues .......... 17 17. Elytres bruns ........ saitoi Matsushita (p. 294) Elytres jaunes ............ 18 1 8. Troisieme article des antennes un peu plus long que le scape zikaweiensis Savio (p. 295) Troisieme article des antennes moins long que le scape itnmaculatus Gressitt (p. 293) 19. Pattes et antennes rouges .......... 20 Pattes et antennes noires . . . . . . . . . .21 20. Lobes inferieurs des yeux fois plus longs que les joues . tertninaliae Fisher (p. 304) Lobes inferieurs des yeux un peu plus longs que les joues . . ruflcolor Pic (p. 295) 21. Les points des elytres sont disposes sans ordre . . . granulicollis Fisher (p. 301) Les points des elytres sont serialement disposes sauf sur le quart sutural testaceus Fisher (p. 299) 22. Elytres noirs avec des dessins blancs nets . . . . . . . . 23 Elytres autrement dessines .......... 24 23. Femurs jaune rougeatre ...... flemingiae Fisher (p. 299) Femurs brun noir . . . . . . . . fouqueti Pic (p. 296) 24. Elytres marbr6s de jaune blanchatre exception faite de quelques assez grandes taches postmedianes brun rougeatre .... pilosicornis Fisher (p. 305) Elytres autrement dessins .......... 25 25. Elytres rouge fonc6 avec des taches blanchatres floues sur le disque de la moitie anterieure et du tiers apical ......... 26 Elytres autrement dessinis . .. . . . ... . . 32 26. Tibias unicolores . . . . ... . . . . . 27 Moiti6 basilaire des tibias d'un rouge clair . ... . . . . 31 27. Pronotum couvert de pubescence grise . . . . .28 Pronotum sans pubescence grise .... actinophorae Fisher (p. 290) 28. Les taches elytrales sont assez grandes et disposees en series longitudinales . . 29 Les taches elytrales sont petites ou non dispos6es en series longitudinales . . 30 29. L'epine laterale du pronotum pr6cedee d'une bosse latrale obtuse celebicus Breuning (p. 292) REVISION DU GENRE EXOCENTRUS MULSANT 223 L'epine laterale du pronotum non pr6cede d'une bosse laterale mindoroanus Breuning (p. 291) 30. Les taches elytrales agglomerees pour former une bande transversale mediane tectonae Fisher (p. 289) Les taches elytrales non agglom6r6es pour former une bande transversale subreticulatus Breuning (p. 307) 31. Troisieme article des antennes beaucoup moins long que le scape pubescens Fisher (p. 304) Troisieme article des antennes a peine moins long que le scape gardneri Fisher (p. 304) 32. Chaque elytre avec une grande tache postmediane brune ou brun fonc6 . . 33 Elytres sans cette tache . . . . . . . . . -37 33. Sur chaque Elytre aussi des taches brun fonc6 disposers en forme de bande trans- versale assez large ....... nigronotatus Pic (p. 296) Elytres sans ces taches ........... 34 34. Sur chaque elytre en plus deux taches ou bandes brun rouge ou noir, une basilaire et une apicale ............ 35 Elytres sans ces taches ou ces bandes . . . . . . . .36 35. Ces taches ou bandes brun rougeatre .... collarti Breuning (p. 301) Ces taches ou bandes noires ..... dalbergianus Gressitt (p. 294) 36. L'epine laterale du pronotum dirigee nettement vers 1'arriere reticulatus Fairmaire (p. 306) L'epine laterale du pronotum dirigee obliquement vers 1'arriere sumbawanus Breuning (p. 289) 37. Chaque elytre vaguement marbr de brun et de gris clair, avec une grande tache postmediane suturale brune. .... subgrisescens Breuning (p. 313) Elytres autrement dessines .......... 38 38. Elytres nettement marbres d'ochrac6 sur fond d'un brun assez fonc6 a/nt'Fisher (p. 301) Elytres autrement dessin6s .......... 39 39. Elytres revetus de pubescence jaune grisatre et ornes de taches longitudinales et transversales brunes ........ tonkineus Pic (p. 296) Elytres autrement dessin6s .......... 40 40. Sur chaque Elytre une 6troite bande longitudinale brun fonce au bord lateral de la moitie ant^rieure . . . . . . . . . . .41 Elytres sans une bande semblable ......... 42 41. Cette bande rejoint en arriere une etroite bande transversale postmediane ondutee brun fonc6 sublateralis Breuning (p. 311) Elytres sans bande postmediane transversale brun fonc6 . . lateralis Gahan (p. 308) 42. Elytres parsemes dans la moitie suturale des deux tiers ant6rieurs de taches blanch- atres et sur le restant de leur surface de taches jaune ocre ghesquierei Breuning (p. 313) Elytres autrement dessines .......... 43 43. Sur chaque elytre cinq ou six series longitudinales de petites taches blanchatres . 44 Elytres sans taches semblables ......... 48 44. Ces taches font d6faut sur remplacement d'une bande transversale postmediane. 45 Ces taches ne font pas defaut sur une bande semblable ..... 46 45. Antennes rouges, unicolores ...... championi Fisher (p. 302) L'extreme base des articles antennaires a partir du quatrieme a pubescence blanchatre greviae Fisher (p. 303) 46. Lobes inferieurs des yeux de moitie plus longs que les joues alboseriatus Gahan (p. 298) Lobes inf6rieurs des yeux quatre fois plus longs que les joues .... 47 47. Rebord lateral du pronotum avant 1'epine laterale largement ourle tnarginicoltis Fisher (p. 297) Le rebord lateral du pronotum non largement our!6 andamanensis Breuning (p. 298) 224 REVISION DU GENRE EXOCENTRUS MULSANT 48. Chaque elytre orn6 de petites taches ochrac6es dispos^es en forme de deux bandes transversales, une postbasilaire et une postmediane . tippmanni Breuning (p. 312) Elytres sans taches pareilles .......... 49 49. Sur chaque elytre des bandes transversales blanchatres souvent floues . . 50 Elytres sans ces bandes . . . . . . . . . .58 50. Sur chaque elytre quatre bandes transversales, une basilaire, une pr6mediane, une postmediane et une apicale . . . . . tnonticola Fisher (p. 303) Sur chaque elytre trois bandes transversales, une postbasilaire ou prem6diane, une mediane ou postmediane et une preapicale . . . . . . .51 51. Elytres assez grossierement ponctues .... dalbergiae Fisher (p. 302) Elytres finement ponctues . . . . . . . . . .52 52. Partie basilaire des articles antennaires a partir du quatrieme a pubescence blanche 53 Partie basilaire de ces articles sans pubescence blanche ..... 54 53. Ecusson a pubescence blanchatre ..... trifasciatus Fisher (p. 302) Ecusson a pubescence brune ....... theresae Pic (p. 295) 54. La bande transversale postbasilaire et la bande pr6m6diane sont largement reunies binhensis Breuning (p. 297) Ces bandes non r6unies ........... 55 55. Les points sur les elytres s6rialement disposes sauf sur le quart sutural cudraniae Fisher (p. 302) Les points sur les elytres non serialement disposes ...... 56 56. Les bandes eiytrales sont assez nettes ..... santali Fisher (p. 305) Les bandes eiytrales sont floues . . . . . . . . -57 57. L'epine laterale du pronotum mince ; troisieme article des antennes un peu plus long que le scape ........ malloti Fisher (p. 304) L'epine laterale du pronotum tres large ; troisieme article des antennes aussi long que le scape ........ kuluensis Breuning (p. 306) 58. Elytres brun fonc6 chacun avec une bande transversale postmediane ondu!6e blanchatre seticollis Fisher (p. 303) Elytres autrement dessines .......... 59 59. Elytres avec de nombreuses petites taches jaunes ou gris jaunatre disposees en series longitudinales et souvent agglomerees en une etroite bande transversale ondulee mediane ....... guttulatus Bates (p. 292) Elytres autrement dessines .......... 60 60. Sur chaque elytre une large bande transversale ochracee au tiers apical . . 61 Elytres sans une bande pareille ......... 62 61. Troisieme article des antennes un peu plus long que le scape futnosus Gahan (p. 298) Troisieme article des antennes un peu moins long que le scape saleyerianus Breuning (p. 298) 62. Sur chaque elytre une assez large bande transversale brune ou brun fonc6, situ^e juste apres le milieu ........... 63 Elytres sane une bande pareille, ornes d'etroites bandes longitudinales blanches . 70 63. Bord lateral de 1'elytre garni de poils dresses blancs ...... 64 Bord lateral de 1'eiytre garni de poils dresses noirs ...... 66 64. Elytres peu finement ponctues .... subexiguus Breuning (p. 307) Elytres tres finement ponctues . . . . . . . . . 65 65. Les points des elytres sont subalignes sur la moitie laterale ivorensis Breuning (p. 315) Les points des elytres non subalignes sur la moitie laterale subfasciatipennis Breuning (p. 305) 66. L'6pine laterale du pronotum faiblement recourbee . pseudexiguus Breuning (p. 307) L'epine laterale du pronotum fortement recourbee ...... 67 67. Les points des elytres sont disposes sans ordre . senegalensis Breuning (p. 315) Les points des elytres sont subalignes ........ 68 68. La bande transversale brune est large et transverse . . inclusus Pascoe (p. 308) REVISION DU GENRE EXOCENTRUS MULSANT 225 La bande transversale brune est etroite et descend dans la moitie suturale oblique- ment en direction de la suture ......... 69 69. Pronotum parseme de taches minimes denudees densefuscosticticus Breuning (p. 312) Pronotum sans taches semblables . . . obliquevittatus Breuning (p. 314.) 70. Pronotum orne de deux larges bandes longitudinales discales ochracees ochreovitticollis Breuning (p. 314) Pronotum sans ces bandes . . . . . . . . . .71 71. L'epine laterale du pronotum precedee d'une bosse laterale obtuse ; sur chaque elytre douze bandes lineiformes blanches ....... 72 L'epine laterale non precedee d'une bosse laterale, les elytres ornes de bandes blanches moins nombreuses. ......... 73 72. Elytres finement ponctu6s ..... multilineatus Breuning (p. 311) Elytres grossierement ponctues ..... alternans Breuning (p. 308) 73. Chaque elytre avec dix tres etroites bandes longitudinales blanches albolineatus Breuning (p. 310) Elytres avec moins de bandes blanches . . . . . . . 74 74. Chaque elytre avec six bandes longitudinales blanches vittulatus Aurivillius (p. 311) Chaque elytre avec huit ou neuf bandes blanches . . . . . . 75 75. L'epine laterale du pronotum dirigee droit vers 1'arriere albovittipennis Breuning (p. 309) L'epine laterale du pronotum dirigee obliquement vers 1'arriere albovittatus Breuning (p. 309) 7. Sousgenre OLIGOPSIS Thomson 1. Elytres d'un vert metallique unicolore . . . viridipennis Breuning (p. 317) Elytres jamais vert metallique ......... 2 2. Chaque elytre avec huit bandes longitudinales blanches sur la moitie anterieure sexseriatus Aurivilius (p. 316) Elytres sans ces bandes ........... 3 3. Sur chaque elytre, dans la moitie anterieure, deux series longitudinales de tres petites taches jaune blanchatre .... timorensis Breuning (p. 321) - Elytres autrement dessines .......... 4 4. Sur chaque elytre de petites taches blanches, disposees en sept series longitu- dinales ....... seriatotnaculatus Schwarzer (p. 322) - Elytres autrement dessines .......... 5 5. Elytres ornes de nombreuses taches jaune pale disposers serialement et agglomerees en une bande transversale postmediane ondulee ...... 6 - Elytres autrement dessines .......... 8 6. Pubescence fonciere des elytres brun fonce . . . roonwali Breuning (p. 321) - Pubescence fonciere des elytres brun rougeatre ...... 7 7. Lobes inferieurs des yeux deux fois plus longs que les joues exocentroides Thomson (p. 320) Lobes inferieurs des yeux un peu plus longs que les joues albizziae Fisher (p. 320) 8. Sur chaque elytre une bande transversale noire situee un peu apres le milieu . 9 - Elytres sans une bande pareille . . . . . . . . . u 9. En plus sur chaque elytre une grande tache basilaire discale brun fonce tneridionalis Breuning (p. 316) - Elytres parsemes de tres petites taches brun fonce, mais sans cette tache basilaire brun fonce. ............ 10 10. L'epine laterale du pronotum pointue .... longipilis Fairmaire (p. 319) L'epine laterale du pronotum trounquee a 1'apex . . major Breuning (p. 317) 11. Sur chaque elytres quatre grandes taches circulaires noiratres nigroplagiatus Breuning (p. 318) Elytres sans ces taches ........... 12 ENTOM. 7. 5. 12 22 6 REVISION DU GENRE EXOCENTRUS MULSANT 12. Sur chaque elytre une assez grande tache postmediane laterale brune bctschuanus Breuning (p. 316) Elytres sans cette tache . .. . . . . . . . 13 13. Elytres grossierement ponctu6s ..... patrizii Breuning (p. 318) Elytres finement ponctues . . . " .. . . . . 14 14. Elytres cou verts de pubescence unicolore jaune grisatre unicoloripennis Breuning (p. 318) Elytres couverts de pubescence brun fonc6 et ornes, chacun, de deux bandes trans- versales floues gris blanchatre, une postbasilaire et une mediane annamensis Breuning (p. 321) 8. Sousgenre FORMOS EXOCENTRUS Breuning Une seule espece ........ variepennis Schwarzer (p. 322) 9. Sousgenre TUBERCULEXOCENTRUS Breuning Une seule espece basituberculatus Pic (p. 323) 10. Sousgenre DENTEXOCENTRUS Breuning Une seule espece dentipes Breuning (p. 323) 1. EXOCENTRUS MULSANT SOUSGENRE EXOCENTRUS SENSU STRICTO Exocentrus Mulsant, 1839, Col. Fr., Long. : 152. Exocentrus Sousgenre s. str., Lepesme & Breuning, 1955, Bull. Soc. ent. Fr. 60 : 127. Troisieme article des antennes aussi long que le quatrieme. Elytres arrondis a 1'apex, ponctues sans ordre. Type : lusitanus Linne. i. Exocentrus lusitanus Linne Cerambyx lusitanus Linn6, 1767, Syst. Nat., ed. 12 : 1767. Cerambyx subpilosus Filler & Mitterpiller, 1783, Iter : 67, pi. 8, fig. 5. Cerambyx lusitanicus Olivier, 1790, Encycl. method., Ent. 5 : 269 ; 1795, Ent. 4, No. 67 : 120, No. 70, pi. 5, fig. 54. Cerambyx pubicornis Schrank, 1790, Naturf. 24 : 76. Cerambyx quercus Rossi, 1790, Fna. Etrusca, 1 : 143. Cerambyx crinitus Panzer, 1795, Ent. Germ. : 249. Lamia balteata Gyllenhal, 1817, Schonherr, Syn. Ins., Append. 2 : 163. Exocentrus balteus Schiodte, 1864, Naturl. Tidskr. (2) 3 : 561. Exocentrus lusitanus Linne^ Ganglbauer, 1883, Wien. ent. Ztg. 2 : 299, pi. 4, fig. 2 ; 1884, Best. Tab. 8 : 530. Antennes un peu plus longues que le corps, le troisieme article a peine aussi long que le scape. Lobes inferieurs des yeux sensiblement plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une mince epine laterale pointue fortement recourbee. Elytres tres densement et tres finement ponctues sur les trois quarts anterieurs. Rouge, couvert d'une tres fine pubescence jaunatre. Sur chaque elytre une large bande transversale postmediane brune qui n'atteint pas tout a fait la suture. Long. : 4-5 mm. ; Larg. : i mm. 1/2-2 mm. REVISION DU GENRE EXOCENTRUS MULSANT 227 Decrit sur des individus de Lusitanie. Repandu par 1'Europe sauf 1'Angleterre. E. subpilosus Pill. & Mitterp., lusitanicus Ol., pubicornis Schr., quercus Rossi, crinifer Panz, balteata Gyll. et balteus Schiodte sont des synonymes. 2. Exocentrus adspersus Mulsant Exocentrus adspersus Mulsant, 1846, Col. Fr., Long., Suppl. : (9). Exocentrus adspersus Mulsant, Fairmaire, 1864, Gen. Col. d'Eur. 4, pi. 47, fig. 217. Exocentrus adspersus Mulsant, Ganglbauer, 1884, Best. Tab. 8 : 529. Antennes un peu plus longues que le corps, le troisieme article a peine aussi long que le scape. Lobes inferieurs des yeux sensiblement plus longs que les joues. Pro- notum deux fois plus large que long, pourvu d'une petite epine laterale pointue faiblement recourbee. Elytres tres densement et tres finement ponctues sur les trois quarts anterieurs. Rouge, couvert d'une tres fine pubescence jaunatre. Elytres parsemes de petites taches blanchatres, disposees en series longitudinales, plus nombreuses surtout au quart apical ; ces taches font defaut sur remplacement d'une assez large bande transversale postmediane ondulee mais sont condensees sou vent juste avant cette bande en forme d'une etroite bande transversale ondulee. Dessous du corps plutot brun a fine pubescence blanchatre. Partie basilaire des articles antennaires 3 et 4 a pubescence blanchatre. Long : 6-8 mm. ; Larg. : 2-2 mm. 1/2 Decrit sur des individus des environs de Lyon. Europe centrale et meridionale ; Caucase, Transcaucasie (Plaviltschikov) . var. clarae Mulsant Exocentrus clarae Mulsant, 1861, Ann. Soc. Linn. Lyon (2), 8 : 206. Exocentrus adspersus var. clarae Pic, 1891, Long. 1 : 46 ; 1915, Long. 9 : 22. Comme la forme typique, mais les petites taches blanchatres des elytres inter- rompent en partie aussi la bande transversale postmediane brune. var. revelieri Mulsant & Rey Exocentrus revelieri Mulsant & Ray, 1875, Ann. Soc. Linn. Lyon (2) 21 : 413. Exocentrus adspersus var. revelieri, Pic, 1891, Long. 1 : 47. Comme la forme typique, mais avec la coloration fondamentale des elytres d'un brun fonce. 3. Exocentrus stierlini Ganglbauer Exocentrus stierlini Ganglbauer, 1883, Wien. ent. Ztg. 2 : 298, pi. 4, fig. 3 : 1884, Best. Tab. 8 : 530. Antennes un peu plus longues que le corps, le troisieme article a peine aussi long que le scape. Lobes inferieurs des yeux de moitie plus longs que les joues. Pronotum deux fois plus large que long, densement et extre"mement finement obliquement strie, pourvu d'une epine laterale pointue fortement recourbee. Elytres densement et tres finement ponctues sur les trois quarts anterieurs. 228 REVISION DU GENRE EXOCENTRUS MULSANT Rouge, couvert (Tune tres fine pubescence jaunatre. Sur chaque elytre une large bande transversale postmediane brune largement arrondie du cote de la suture qu'elle n'atteint pas tout a fait, et une bande transversale preapicale brune tres peu apparente ; chacun des points situees entre ces deux bandes entoure d'un tres petit cercle brun. Long. : 4-6 mm. ; Larg. ; i mm. 1/2-2 mm. Decrit sur des individus d'Europe meridionale. Allemagne, Autriche (Ganglbauer) ; Hongrie (Aurivillius) ; Siberie (Plaviltschikov) . 4. Exocentrus punctipennis Mulsant & Guillebeau Exocentrus punctipennis Mulsant & Guillebeau, 1856, Ann. Soc. Linn. Lyon (2) 3 : 103. Exocentrus punctipennis Mulsant & Guillebeau, Ganglbauer, 1884, Best. Tab. 8 : 530. Antennes un peu plus longues que le corps, le troisieme article a peine aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une large epine laterale pointue fortement recourbee. Elytres tres densement et tres finement ponctues sur les trois quarts anterieurs. Rouge, couvert d'une fine pubescence jaune paille. Disque du pronotum, rem- bruni sauf au bord anterieur et au bord posterieur. Elytres parsemes de tres petites taches brunes rangees en series longitudinales. Sur chaque elytre une large bande transversale ondulee brune qui n'atteint pas tout a fait la suture. Long. : 4-6 mm. ; Larg. ; i mm. 1/2-2 mm. 1/4. Decrit sur des individus de Lyon. Repandu par 1'Europe centrale et meridionale. var. signatus Mulsant & Rey Exocentrus signatus Mulsant & Ray, 1863, Ann. Soc. Linn. Lyon (2) 10 : 163. Exocentrus punctipennis var. signatus Ganglbauer, 1884, Best. Tab. 8 : 530. Exocentrus graecus Pic, 1901, Echange, 17 : 52. Comme la forme typique, mais la bande transversale brune de 1'elytre moins large et plus foncee. Grece et Turquie. 5. Exocentrus galloisi Matsushita Exocentrus galloisi Matsushita, 1933, Journ. Fac. Agric. Hokk. 34 : 396, 397, pi. 4, fig. 7. Proche de punctipennis Mulsant & Guillebeau, mais 1 'epine laterale du pronotum un peu moins fortement recourbee et les elytres sans petites taches brunes disposees en series longitudinales. Decrit sur un individu du Japon : Chiuzenji au Musee de Hokkaido. He Hondo (coll. Frey). 6. Exocentrus savioi Pic Exocentrus curtipennis Pic var. Savioi Pic, 1925, Bull. Soc. ent. Fr. : 138. Exocentrus curtipennis Pic var. Savioi Pic, Savio, 1929, Not. d'Ent. chin. : I. Exocentrus curtipennis Pic var. Savioi Pic, Gressitt, 1951, Longicornia, 2 : 527. Proche de punctipennis Mulsant & Guillebeau, mais 1'epine laterale du pronotum REVISION DU GENRE EXOCENTRUS MULSANT 229 plus mince et plus longue et la bande transversale postmediane brune de 1'elytre plus fortement ondulee et devenant beaucoup moins large dans le tiers sutural. Decrit sur un individu de Chine : Zikawei dans la coll. Pic. Provinces de Kiangsi et Kiang-su (Gressitt) ; Chekiang (coll. Frey) ; He de Formose (Gressitt). 7. Exocentrus lineatus Bates Exocentrus lineatus Bates, 1873, Ann. Mag. nat. Hist. (4) 12 : 384. Exocentrus lineatus Bates, Matsushita, 1933, Journ. Fac. Agric. Hokk. 34 : 396. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une petite epine laterale mince et pointue, dirigee obliquement vers 1'arriere. Elytres tres densement et finement ponctues sur les quatre cinquiemes anterieurs. D'un rouge assez fonce, couvert d'une fine pubescence brun grise. Elytres rev^tus de pubescence brun rougeatre et ornes, chacun, dans la moitie anterieure de cinq troites bandes longitudinales jaunes ou jaune grisatre ainsi que de trois courtes bandes pareilles dans le quart apical. Moitie apicale des tibias et les tarses brun fonce. Antennes a pubescence brun rougeatre fonce. Long. : 5-6 mm. ; Larg. : 2-2 mm. 1/4. Decrit sur des individus du Japon au Museum de Paris. lies Hondo, Shikoku et Kiushiu (Matsushita). 8. Exocentrus testudineus Matsushita Exocentrus testudineus Matsushita, 1931, Trans. Sapp. Nat. Hist. Soc. 12 : 47 ; 1933, Journ. Fac. Agric. Hokk. 34 : 396. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum faible- ment transverse, pourvu d'une assez longue epine laterale pointue a peine recourbee. Elytres tres densement et finement ponctues dans les trois quarts anterieurs, eparse"- ment et tres finement dans le quart apical. Brun fonce couvert d'une tres fine pubescence grise. Le bord anterieur et le bord posterieur du pronotum, les elytres et les antennes rouges et revetus de pubescence brun rougeatre. Ecusson a pubescence blanchatre. Sur chaque elyte une grande tache circulaire preapicale suturale blanchatre vague et deux bandes transversales de meme couleur, une postbasilaire qui remonte obliquement en direction de 1'epaule et une ondulee situee juste apres le milieu, ces deux bandes reliees ensemble par deux ou trois etroites bandes longitudinales discales de me'me couleur. Long. : 6-8 mm. ; Larg. : 2 mm. 1/4-3 mm. Decrit sur des individus du Japon : He Hokkaido, Mts. Daisetsu dans la coll. Matsushita. He Hondo (Matsushita). 9. Exocentrus brevisetosus Gressitt Exocentrus testudineus Matsushita subspecies brevisetosus Gressitt, 1938, Philipp. Journ. Sc. 64 : 167, pi. i, fig. 4. Exocentrus brevisetosus Gressitt, 1951, Longicornia, 2 : 526. 230 REVISION DU GENRE EXOCENTRUS MULSANT Proche de testudineus Matsushita, mais Tepine laterale du pronotum plus forte- ment ' recourbee, les elytres plus grossierement ponctues et garnis de polls dresses plus courts. Long. : 4 mm. ; Larg. ; i mm. 1/2. Decrit sur un individu de 1'Ile de Formose : Hori, a 1' Academic de Calif ornie. Je ne connais cette espece que d'apres la description. 10. Exocentrus pici, nom. nov. Exocentrus signatus Pic, 1933, Mater. Longic. 11:6. Antennes un peu plus longues que le corps, le troisieme article un peu plus long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum transverse, pouvu d'une longue epine laterale mince et pointue, dirigee obliquement vers 1'arriere. Elytres tres densement et finement ponctues. Brun fonce, couvert de pubescence brun rougeatre fonce. Sur chaque elytre une tres etroite bande suturale blanchatre, une bande premediane blanche peu large qui remonte obliquement en direction de 1'epaule et qui est interrompue dans sa moitie laterale par de petites taches denudees, une assez etroite bande transversale post- mediane fortement zigzaguee et une etroite tache discale preapicale blanche. La partie de 1'elytre entres les bandes premediane et postmediane rouge a pubescence brun rougeatre clair. Tibias rouge clair. Long. : 6 mm. ; Larg. ; 2 mm. 1/2. Decrit sur un individu de Chine : Province de Szetschouan, Tatsienlu, Dzaschi dans la coll. Frey, sous le nom signatus. Ce nom etant preoccupe je le change en pici. 11. Exocentrus badius Fisher Exocentrus badius Fisher, 1925, Philipp. Journ. Sc. 28 : 242. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Pronotum fortement transverse, pourvu d'une assez longue et mince epine laterale pointue, dirigee obliquement vers 1'arriere. Elytres densement et peu finement ponctues sur les trois quarts anterieurs. Brun rougeatre couvert d'une fine pubescence grise. Elytres revetus de pubescence brun rougeatre et parsemes de petites taches circulaires grises, ces taches faisant defaut sur 1'emplacement d'une grande tache transversale postmediane laterale. Long. : 5 mm. ; Larg. : 2 mm. Decrit sur un individu de 1'Ile Mindanao : Davao, au Musee de Washington. Je ne connais cette espece que d'apres la description. 12. Exocentrus albovarius Fisher Exocentrus albovarius Fisher, 1925, Philipp. Journ. Sc. 28 : 243. Proche de badius Fisher, mais le pronotum deux fois plus large que long, son epine laterale plus rapprochee de la base et dirigee vers 1'arriere, le corps d'un brun REVISION DU GENRE EXOCENTRUS MULSANT 231 plus fonce, la tache postmediane de Felytre depourvue de taches grise est de forme plus irreguliere. Long. : 3 mm. 1/2-5 mm - Larg. : i mm. 1/2-2 mm. Decrit sur des individus de 1'Ile Mindanao : Davao, au Musee de Washington. Je ne connais cette espece que d'apres la description. 13. Exocentrus philippinus Fisher Exocentrus philippinus Fisher, 1925, Philipp. Journ. Sc. 28 : 246. Antennes un peu plus longues que le corps, le troisieme article sensiblement moins long que le scape. Pronotum de moitie plus large que long, pourvu d'une longue epine laterale pointue, dirigee obliquement vers l'arriere. Elytres densement et peu finement ponctues sur les trois quarts anterieurs. Brun rougeatre, couvert d'une fine pubescence blanchatre. Tete a pubescence grise. Elytres revetus de pubescence brune et marbres de jaune pale, ces marbrures condensees en une grande tache humerale, une bande transversale au tiers apical et en quelques petites taches au quart apical rangees en series longitudinales. Dessous noiratre. Antennes a pubescence grise. Long. : 4 mm. 1/2-7 mm - > Larg. : i mm. 3/4-2 mm. 1/2. Decrit sur des individus de 1'Ile Mindanao : Davao et Zamboanga, au Musee de Washington. Je ne connais cette espece que d'apres la description. 14. Exocentrus echimys Pascoe Exocentrus echimys Pascoe, 1864, Trans, ent. Soc. Lond. (3) 3 : 30. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois et demie plus longs que les joues. Pro- notum deux fois plus large que long, eparsement et finement ponctue, pourvu d'une epine laterale pointue dirigee tres obliquement vers rarriere. Elytres densement et tres finement ponctues. Rouge, couvert d'une fine pubescence jaune pale. Elytres avec une large bande transversale postmediane rouge fonce floue. Tiers apical des tibias bran fonce. Long. : 5 mm. ; Larg. : i mm. 2/3. Decrit sur un individu de 1'Ile Morty (Morotai), au British Museum. 15. Exocentrus erinaceus Pascoe Exocentrus erinaceus Pascoe, 1863, Trans, ent. Soc. Lond. (3) 1 : 529. Antennes un peu plus longues que le corps, le troisieme article sensiblement moins long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum transverse, pourvu d'une epine laterale pointue tres fortement recourbee. Elytres densement et assez grossierement ponctues sur les trois quarts anterieurs. Brun fonce. Elytres revetus de pubescence bran rougeatre fonce et marbres un 232 REVISION DU GENRE EXOCENTRUS MULSANT peu avant le milieu et sur le tiers apical de jaune paille. L'extreme base des articles antennaires a partir du quatrieme a pubescence blanchatre. Long. : 4 mm. 1/2 ; Larg. : I mm. 1/3. Decrit sur un individu de Queensland : Port Denison, au British Museum. 16. Exocentrus lachrymosus Pascoe Exocentrus lachrymosus Pascoe, 1864, Trans, ent. Soc. Lond. (3) 3 : 29. Antennes un peu plus longues que le corps, le troisieme article sensiblement moins long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pro- notum plus de deux fois plus large que long, pourvu d'une epine laterale pointue fortement recourbee et precedee d'une bosse laterale obtuse. Elytres densement et tres finement ponctues. Brun rougeatre. Sur chaque elytre cinq rangees longitudinales de petites taches jaune blanchatre. Long. : 5 mm. 1/2 ; Larg. : 2 mm. Decrit sur un individu de Borneo : Sarawak, au British Museum. 17. Exocentrus insulicola nom. nov. Exocentrus insularis Fisher, 1934, Stylops, 3 : 36, 42. Antennes aussi longues que le corps, le troisieme article un peu moins long que le scape. Pronotum transverse, pourvu d'une petite epine laterale pointue, dirigee obliquement vers I'arriere. Elytres assez densement et peu finement ponctues sur les trois quarts anterieurs. Brun rougeatre, couvert de pubescence jaunatre. Pronotum avec deux taches obliques discales blanchatres floues. Elytres revetus de pubescence brune et ornes d'assez nombreuses taches oblongues blanches, condensees en une sorte de bande transversale mediane ondulee et faisaint defaut sur 1'emplacement d'une large bande transversale postmediane. Long. : 4 mm. 1/2 ; Larg. : i mm. 1/2. Decrit sur un individu de 1'lle Banka : Mt. Mangkol, au Musee de Washington. Je ne connais cette espece que d'apres la description. 18. Exocentrus bauhiniae Fisher Exocentrus bauhiniae Fisher, 1934, Stylops, 3 : 36, 37. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une petite epine laterale conique et pointue, dirigee obliquement vers Tamere. Elytres densement et peu finement ponctues sur les trois quarts anterieurs. Brun, couvert de pubescence jaune blanchatre. Elytres revetus de pubescence bran rougeatre et ornes, chacun, d'une etroite bande longitudinale suturale et de cinq bandes longitudinales discale blanchatres largement interrompues apres le REVISION DU GENRE EXOCENTRUS MULSANT 233 milieu. Antennes a pubescence brun jaunatre, a pubescence un peu plus claire sur la partie basilaire des articles a partir du troisieme. Long. : 4-5 mm. 1/2 ; Larg. : i mm. 1/2-2 mm. 1/4. Decrit sur un individu de Java : Kadiri, au Musee de Washington. Samarang, Buitenzoorg, etc. (Fisher). 19. Exocentrus hageni sp. n. Antennes aussi longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum plus de deux fois plus large que long, pourvu d'une epine laterale conique et pointue, dirigee vers Tamere et precedee d'une bosse laterale obtuse. Elytres tres densement et assez finement ponctues sur les quatre cinquiemes anterieurs. Rouge, couvert d'une tres eparse pubescence jaunatre. Elytres parsemes de nombreuses petites taches jaune pale, rangees, sur chacun, en forme de sept series longitudinales. Long. : 5 mm. 1/2 ; Larg. : i mm. 3/4. Type de Sumatra : Serdang, Tandjong-Morawa, leg. D. B. Hagen, au Museum de Leiden. 20. Exocentrus enganensis Breuning Exocentrus (s. s.) enganensis Breuning, 1957, Bull. Inst. voy. Sc. nat. Belg. 33, No. 8 : 8. Antennes a peine aussi longues que le corps, le troisieme article un peii moins long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une assez longue epine laterale pointue, dirigee obli- quement vers 1'arriere. Elytres tres densement et finement ponctues sur toute leur etendue. Rouge, couvert d'une fine pubescence soyeuse grise. Ecusson borde en arriere de pubescence blanche. Elytres reve'tus de pubescence brun rougeatre et ornes de tres petites taches blanches, rangees, sur chacun, en forme de sept series longitudinales regulieres ; ces taches plus nombreuses pour former aussi deux etroites bandes transversales, une premediane qui remonte obliquement en direction de la marge externe et une mediane qui descend un peu en direction de la marge externe. Partie basilaire des articles antennaires 4 et 5 a pubescence blanch atre. Long. : 5 mm. ; Larg. : 2 mm. Decrit sur un individu de 1'Ile Engano, au British Museum. 21. Exocentrus sublineatus Breuning Exocentrus (s. s.) sublineatus Breuning, 1957, Bull. Inst. roy. Sc. nat. Belg. 33, No. 8 : 7. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une petite epine laterale pointue fortement recourbee. Elytres densement et finement ponctues sur les trois quarts anterieurs. Rouge, couvert de pubescence brun rougeatre. Sur chaque elytre une etroite bande suturale, six etroites bandes longitudinales discales atteignant presque le milieu, 234 REVISION DU GENRE EXOCENTRUS MULSANT trois bandes semblables parcourant le tiers apical et quelques tres petites laches entre ces deux series de bandes, toutes de couleur jaune pale. Long. : 4-5 mm. ; Larg. : i mm. 3/4-2 mm. Decrit sur un individu de 1'Indochine : Saigon a 1'Institut Royal des Sciences Naturelles de Belgique. Tonkin : Hoa-Binh (coll. Frey). Cette espece se rapproche beaucoup de lineatus Bat., mais s'en distingue par Fepine laterale du pronotum moins troite et plus fortement recourbee, les bandes claires de la moitie anterieure de 1'elytre plus courtes, etc. 22. Exocentrus suturalis Pic Exocentrus suturalis Pic, 1926, Mel. exot. ent. 45 : 28. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum forte- ment transverse, pourvu d'une longue epine laterale pointue, dirigee directement vers Farriere et precedee d'une bosse laterale obtuse. Elytres densement et finement ponctues. Rouge fonce, couvert de pubescence grisatre. Elytres parsemes de nombreuses petites taches jaune pale, rangees, sur chacun, en cinq series longitudinales, et avec une etroite bande suturale jaune pale. Long. : 5 mm. ; Larg. ; 2 mm. 1/4. Decrit sur un individu du Tonkin a la coll. Pic. 23. Exocentrus albomaculatus Pic Exocentrus albomaculatus Pic, 1928, Mel. exot. ent. 51 : 29. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux un peu plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une petite epine laterale pointue fortement re- courbee. Elytres densement et finement ponctues sur les trois quarts anterieurs et garnis de tres longs poils dresses. Brun fonce, couvert de pubescence brune. Elytres rouges et parsemes de taches circulaires blanches. Antennes bran fonce. Long. : 3 mm. ; Larg. : 3/4 mm. Decrit sur un individu du Tonkin a la coll. Pic. 24. Exocentrus pseudandamanensis sp. n. Exocentrus pseudandamanensis Breuning, 1957, Bull. Inst. roy. Sc. nat. Belg. 33, No. 8 : 8. Exocentrus andamanensis Fisher part., 1932, Stylops, 1 : 231. Antennes un peu plus longues que le corps, le troisieme article un peu plus long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une epine laterale pointue fortement recourbee. Elytres assez densement et finement ponctues sur les trois quarts anterieurs. Rouge, couvert d'une fine pubescence jaune grise. Elytres revetus de pubescence REVISION DU GENRE EXOCENTRUS MULSANT 235 brun rougeatre et parsemes de nombreuses petites taches jaune blanche, rangees, sur chacun, en forme de sept series longitudinales. Antennes a pubescence brun rougeatre. Long : 5 mm. ; Larg. : 2 mm. Type de Birmanie : Katha, Mohngin Res., au British Museum. Fisher identifie cet exemplaire avec andamanensis Fisher tout en parlant de differences. 25. Exocentrus downingi Fisher Exocentrus downingi Fisher, 1932, Stylops, 1 : 226. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Prono- tum fortement transverse, pourvu d'une courte epine laterale peu pointue fortement recourbee. Elytres densement et extremement finement ponctues dans les deux tiers anterieurs. Brun fonce, couvert de pubescence brune. Les bords des yeux et la base du pro- notum a pubescence jaune pale. Elytres marbres de blanchatre dans le quart basilaire et dans le tiers apical et avec quelques tres petites taches blanchatres entre ces deux zones. Partie basilaire des articles antennaires 4 a 9 a pubescence blanchatre. Long. : 7 mm. ; Larg. : 2 mm. 1/2. Decrit sur un individu de Tlnde : Nilghiri Hills, au British Museum. 26. Exocentrus beesoni Fisher Exocentrus beesoni Fisher, 1933, Ind. For. Rec. (4) 18 : 3. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux aussi longs que les joues. Pronotum deux fois plus large que long, pourvu d'une longue epine laterale conique et pointue forte- ment recourbee. Elytres densement et finement ponctues dans les trois quarts anterieurs. Brun, couvert d'une fine pubescence gris clair. Ecusson a pubescence gris blan- chatre. Elytres et antennes revetus de pubescence brun rougeatre, les elytres par- semes de taches gris blanchatre floues couvrant la majeure partie de la moitie anterieure et formant une bande transversale preapicale ondulee. Partie basilaire des articles antennaires 2 a 10 a pubescence blanchatre. Long. : 5-5 mm. 1/2 ; Larg. : 2-2 mm. 1/4. Decrit sur des individus de 1'Inde : Madras, N. Salem, Ayur, au British Museum et au Musee de Washington. 27. Exocentrus transversifrons Fisher Exocentrus transversifrons Fisher, 1940, Ind. For. Rec. (2) 6 : 209. Antennes de moitie plus longues que le corps, le troisieme article un peu plus long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pro- 236 REVISION DU GENRE EXOCENTRUS MULSANT notum transverse, pourvu d'une longue epine laterale mince et pointue faiblement recourbee. Elytres tres densement et finement ponctues. Brun couvert de pubescence jaune paille. Elytres densement marbres de brun. Long. : 6-7 mm. ; Larg. : 2-3 mm. Decrit sur des individus de 1'Inde : U.P. Chakrata Range, au British Museum et au Musee de Dehra Dun. 28. Exocentrus tesselatus Perroud Exocentrus tesselatus Perroud, 1855, Ann. Soc. Linn. Lyon (2) 2 : 397 ; Mel. ex. 3 : 77. Antennes d'un quart plus longues que le corps, le troisieme article un peu plus long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une epine laterale pointue forte- ment recourbee. Elytres assez densement et extrmement finement ponctues dans les deux tiers anterieurs. Rouge, couvert de pubescence jaune paille. Tte, pronotum, ecusson et scape a pubescence rousseatre. Elytres marbres, sauf dans la region apicale, de brun. Articles antennaires a partir du troisieme rembrunis a 1'apex. Long. : 5 mm. ; Larg. : i mm. 2/3. Decrit sur un individu de 1'Inde : Pondichery, dans la coll. Pic. 29 Exocentrus parrotiae Fisher Exocentrus parrotiae Fisher, 1932, Ind. For. Rec. 16 : 296, 304. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux aussi longs que les joues. Pronotum deux fois plus large que long, tres eparsement et tres finement granule sur le disque, pourvu d'une epine laterale pointue fortement recourbee. Elytres tres densement et finement ponctues. Rouge, couvert d'une fine pubescence gris blanchatre. Sur chaque elytre une tache discale postmediane oblique brun rougeatre. Articles antennaires a partir du troisieme rev6tus de pubescence brun rougeatre, mais de pubescence blanchatre sur la partie basilaire. Les poils dresses du dessus et des antennes sont noirs, ceux du bord lateral des elytres, du dessous et des pattes blancs. Long. : 7-8 mm. 1/4 ; Larg. : 2 mm. 3/4-3 mm. 1/2. Decrit sur des individus du Kashmir : Upper Munda, au British Museum et au Musee de Dehra Dun. 30. Exocentrus klapperichi Breuning Exocentrus (s. s.) klapperichi Breuning, 1957, Ent. NachrBl. osterr. u. schweiz. Ent. 8, No. 3 : 12. Antennes d'un quart plus longues que le corps, le troisieme article un peu plus long que le scape. Lobes inferieurs des yeux un peu plus longs que les joues. Pro- notum deux fois plus large que long, pourvu d'une petite epine laterale pointue forte- REVISION DU GENRE EXOCENTRUS MULSANT 237 ment recourbee. Elytres densement et tres finement ponctues sur les trois quarts anterieurs. Brun fonce, couvert de pubescence gris jaune. Pronotum rouge au bord anterieur et au bord posterieur, revetu de pubescence brune. Sur chaque elytre deux bandes transversales zigzaguees brunes, une mediane qui s'elargit au bord lateral de facon a y occuper presque tous les deux tiers anterieurs, et une postmediane qui n'atteint pas la suture, ainsi qu'une petite tache preapicale subsuturale brune. Moitie apicale des tibias et les antennes a pubescence brune. Long. : 5-7 mm. ; Larg. : i mm. 3/4-2 mm. 1/4. Decrit sur un individu d'Afghanistan : Nuristan, Kutiau, dans la coll. Tippmann. Hindukusch : Chitral, Tal Bikir (Musee de Trieste). 31. Exocentrus kashmirensis Breuning Exocentrus kashmirensis Breuning, 1957, Ent. Arb. Mus. Frey, 8 : 277. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux sensiblement plus longs que les joues. Pro- notum deux fois plus large que long, pourvu d'une assez longue epine laterale mince et pointue, dirigee obliquement vers Farriere. Elytres tres densement et finement ponctues. Rouge, couvert d'une pubescence jaunatre. Chaque elytre avec sept series longi- tudinales de petites taches jaune pale qui s'arr6tent a peu pres au milieu tout en y formant une bande transversale fortement ondulee, et, dans le tiers apical avec trois courtes bandes longitudinales discales, jaune pale. Antennes a pubescence brun rougeatre clair, la partie basilaire des articles 3 a 8 a pubescence jaunatre. Long. : 5 mm. ; Larg. : I mm. 3/4. Type de Kaskmir : Kulu, 2300 m. alt., leg. C. Rost, dans la coll. Frey. Un Paratype (idem). 32. Exocentrus madecassus Fairmaire Exocentrus madecassus Fairmaire, 1880, Ann. Soc. ent. Fr. (5) 10 : 338. Antennes un peu moins longues que le corps, le troisieme article a peine aussi long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Tete et pronotum a ponctuation tres fine et extre'mement dense. Pronotum deux fois plus large que long, pourvu d'une petite epine laterale mince et pointue, faible- ment recourbee. Elytres densement et peu finement ponctues sur les deux tiers anterieurs. Rouge, le disque du pronotum sauf an bord anterieur et au bord posterieur, rembruni. Les cotes du pronotum et 1'ecusson converts d'une fine pubescence gris clair. Elytres revetus de pubescence brun rougeatre clair et parsemes de petites taches gris blanchatres serrees vagues qui font defaut sur 1 'emplacement d'une assez grande tache postmediane suturale vague. L'extr&ne base des articles antennaires 3 a ii a pubescence blanchatre. Long. : 5-6 mm. ; Larg. : 2 mm. 1/4-2 mm. 1/2. Decrit sur des individus de Madagascar, au Museum de Paris. Suberbieville (Museum de Paris). 238 REVISION DU GENRE EXOCENTRUS MULSANT 33. Exocentrus interruptefasciatus Hunt & Breuning Exocentrus interruptefasciatus Hunt & Breuning, 1957, Durban Mus. Novit. 5 : 64. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux de moitie plus longs que les joues. Pronotum transverse, pourvu d'une epine laterale pointue tres fortement recourbee. Elytres densement et tres finement ponctues sur toute leur etendue. Rouge fonce, couvert de pubescence brun rougeatre fonce. Ecusson et moitie anterieure des elytres revetus presque entierement de pubescence jaune blanchatre. Moitie posterieure des elytres parsemee de petites taches jaune blanchatre rangees en series longitudinales et agglomerees le long de la suture. Moitie basilaire des tibias et les tarses, rouges. Le deuxieme article des antennes et la partie basilaire des articles 3 a n rouge et converts de fine pubescence jaune. Long : 4-4 mm. 1/2 ; Larg. : i mm. 1/2-1 mm. 2/3. Decrit sur des individus de Zululand: Ubombo, Eteza (coll. Hunt). 34. Exocentrus lacteolus Distant Exocentrus lacteolus Distant, 1906, Ins. Transvaal : 166, fig. 26. Antennes aussi longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum transverse, pourvu d'une petite epine laterale pointue, peu rapprochee de la base et assez faiblement recourbee. Elytres finement et extre'mement densement ponctues. Rouge fonce. Elytres parsemes de tres petites taches blanches disposees, sur chacun, en a peu pres sept rangees longitudinales. Pattes rouge fonce. L'extreme base des articles antennaires 3 a 9 a pubescence blanche. Long. : 4 mm. 1/2 ; Large. : 2 mm. 1/4. Decrit sur des individus de Natal ; Durban au Musee de Pretoria. Rhodesia : Salisbury (British Museum). 35. Exocentrus ruficollis Lameere Exocentrus ruficollis Lameere, 1892, Ann. Soc. ent. belg. 37 : 508. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum plus de deux fois plus large que long, pourvu d'une epine laterale pointue peu forte- ment recourbee. Elytres finement et extre'mement densement ponctues. Noir, couvert de pubescence brun fonce. Pronotum, ecusson et epipleures rouges et converts de pubescence rousseatre. Tout le quart apical le 1'elytre couvert d'une fine et eparse pubescence gris jaunatre. Sur chaque elytre une etroite bande trans- versale tres ondulee gris clair situee un peu avant le milieu. L'extreme base des articles antennaires a partir du quatrieme a pubescence blanchatre. Long. : 5-7 mm. ; Larg. : i mm. 3/4-2 mm. 1/4 Decrit sur un individu du Congo a 1'Institut Royal des Sciences Naturelles de Belgique. Zambezi : Noca Choupanga pres Chemba (Museum de Paris) ; Afrique orientale anglaise : Xinavane (British Museum). REVISION DU GENRE EXOCENTRUS MULSANT 239 36. Exocentrus subruficollis Breuning Exocentrus (s. s.) subruficollis Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 129. Proche de ruficollis Lameere, mais 1'epine laterale du pronotum sensiblement plus fortement recourbee et precedee d'une bosse laterale obtuse peu accusee et les elytres revetus d'une pubescence unicolore brun fonce. Decrit sur des individus du Congo beige : Mayidi, au Musee de Tervueren. Repandu du Togo (Museum de Paris), au Victoria Nyanza (Musee de Tervueren) et au Gabon (Museum de Paris). 37. Exocentrus pseudoruficollis Breuning Exocentrus (s. s.) pseudoruficollis Breuning, 1957, Bull. Inst. roy. Sci. nat. Belg. 33, No. 8 : 8. Proche de ruficollis Lameere, mais plus petit, 1'epine laterale du pronotum plus courte et tres fortement recourbee, precedee d'une bosse laterale obtuse peu accusee, les elytres tres finement ponctues et entierement d'un brun rouge clair, de meme que le dessous du corps, les pattes et les antennes. Long. : 4-6 mm. ; Larg. : i mm. 1/2-2 mm. Decrit sur des individus du Cap, au British Museum. 38. Exocentrus trinigrovittatus Breuning Exocentrus (s. s.) trinigrovittatus Breuning, 1957, Bull. Inst. roy. Sc. nat. Belg. 33, No. 8 : 7. Antennes d'un quart plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une assez courte epine laterale pointue fortement recourbee. Elytres tres densement et finement ponctues. Noir. Tete et pronotum marbres de blanchatre et de brun noir. Ecusson et elytres revetus de pubescence blanchatre. Elytres parsemes de taches denudees minimes et ornes, chacun, de trois bandes transversales noires, une etroite basilaire s'elargissant en direction de la suture, une assez large postmediane faiblement ondulee et une apicale peu large. Tarses et antennes a pubescence noire, le deuxieme article des antennes et la partie basilaire des articles 3 a 10 a pubescence blanche. Long. : 5 mm. ; Larg. : 2 mm. 1/4. Decrit sur un individu du Tanganyika : Shinyanga, au British Museum. 39. Exocentrus asmarensis sp. n. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum forte- ment transverse, pourvu d'une assez longue epine laterale pointue fortement re- courbee. Elytres densement et tres finement ponctues. Rouge, couvert de pubescence jaune paille. Elytres ornes d'une bande transversale postmediane brun rougeatre formant sur les deux elytres un grand M. Moitie apicale des tibias et les derniers articles antennaires d'un brun assez fonce. Long. : 5 mm. 1/2 ; Larg. : 2 mm. Type d'Erithree : Asmara, dans la coll. Frey. 240 REVISION DU GENRE EXOCENTRUS MULSANT 40. Exocentrus ugandicola Breuning Exocentrus (s. s.) ugandicola Breuning, 1958, Bull. Inst. roy. Sc. nat. Belg., 34, No. 22. Antennes un peu plus longues que le corps, le troisieme article sensiblement moins long que le scape. Lobes inferieurs des yeux de moitie plus longs que les joues. Pro- notum deux f ois plus large que long, pourvu d'une assez longue epine laterale pointue et recourbee. Elytres densement et tres finement ponctues sur les trois quarts anterieurs. Brun noir, convert de pubescence jaune pale. Vertex et disque du pronotum, sauf une assez large bande longitudinale prothoracique denudee, revetus de pubescence jaune. Elytres parsemes de tres petites taches circulaires brun noir et ornes, chacun, d'une large bande transversale postmediane brun fonce s'amincissant beaucoup dans le tiers sutural, et de deux taches brun fonce : une discale proche de la suture et une laterale, preapicales. Moitie apicale des tibias, les tarses et les antennes a partir du troisieme article a pubescence brun noir, le quart basilaire des articles anten- naires 3 a n a pubescence blanchatre. Long. : 5-6 mm. ; Larg. : 2 mm. 1/4-2 mm. 1/2. Decrit sur des individus de 1'Uganda : Mpanga, au British Museum. 41. Exocentrus ochreopunctatus Breuning Exocentrus (s. s.) ochreopunctatus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 129. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une petite epine laterale pointue, dirigee oblique- ment vers Farriere. Elytres tres densement et tres finement ponctues. Noir, couvert de pubescence noire ; le bord anterieur et le bord posterieur du pronotum rougeatres. Elytres vaguement marbres de gris blanchatre et ornes de tres petites taches circulaires nettes ochracees, rangees en forme de deux etroites bandes transversales ondulees, une postbasilaire et une postmediane. Partie basilaire des articles antennaires 3 a n a pubescence blanchatre. Long. : 7 mm. 1/2 ; Larg. : 2 mm. 1/2. Decrit sur des individus de Ruanda : Gabiro, au Musee de Tervueren. 42. Exocentrus josephi Duvivier Exocentrus josephi Duvivier, 1890, Ann. Soc. ent. belg. 34 : 38 ; Bull. Soc. ent. Belg. 1890 : 197. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux de moitie plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une epine laterale mince et assez longue, dirigee vers Tamere et precedee d'une bosse laterale obtuse. Elytres densement et tres finement ponctues. Rouge, couvert de pubescence gris tirant sur le rougeatre. Sur chaque elytre une etroite bande transversale postmediane blanche faiblement ondulee. Partie basilaire des articles antennaires a partir du troisieme a pubescence blanchatre. Long. : 4 mm. 2/3 ; Larg. : 2 mm. REVISION DU GENRE EXOCENTRUS MULSANT 241 Decrit sur un individu du Congo beige : Leopoldville, a 1'Institut Royal des Sciences Naturelles de Belgique. 43. Exocentrus grisescens Jordan Exocentrus grisescens Jordan, 1894, Novit. Zool. 1 : 246. Exocentrus ortmansi Gahan, 1917, Bull. Ent. Research, 8 : 117. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une epine laterale pointue fortement recourbee. Elytres tres densement et tres finement ponctues. Brun fonce, couvert de pubescence bran fonce vaguement entremelee de gris clair sur le disque des elytres. Elytres parsemes de tres petites taches denudees, disposees en series longitudinales. Articles antennaires a partir du troisieme reve'tus de pubescence gris blanchatre. Long. : 6 mm. 1/2-7 mm. ; Larg. : 2 mm. 1/3-2 mm. 2/3. Decrit sur un individu de Gabon : Kuilu, au Museum de Paris. Mayumbe (Gahan). Exocentrus ortmansi Gah. est un synonyme. 44. Exocentrus decellei Breuning Exocentrus (s. s.) decellei Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 127. Antennes d'un quart plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois plus longs que les joues. Pronotum transverse, pourvu d'une petite epine laterale pointue, fortement recourbee. Elytres densement et tres finement ponctues. Brun fonce, couvert de pubescence soyeuse jaunatre. Elytres reve'tus de pubescence brun fonce et parsemes de petites taches de pubescence soyeuse jaunatre, ces taches plus nombreuses le long de la suture et en forme de trois larges bandes trans- versales ondulees, une postbasilaire, une mediane et une postmediane. Moitie apicale des tibias, les tarses et les antennes revetus de pubescence brun fonce, les deux premiers articles des antennes et la base des articles 3 et 4 a pubescence soyeuse jaunatre. Long. : 4 mm. 1/2-5 mm - I / 2 Larg. : 2-2 mm. 1/2. Decrit sur des individus du Congo beige : Yangambi, au Musee de Tervueren. 45. Exocentrus massarti Breuning Exocentrus (s. s.) massarti Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 127. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux deux fois et demie plus longs que les joues. Prono- tum fortement transverse, pourvu d'une assez longue epine laterale pointue, forte- ment recourbee. Elytres tres densement de tres finement ponctues. Brun fonce, couvert de pubescence brun fonce. Ecusson revetu de pubescence jaunatre. Sur chaque elytre trois larges bandes transversales ondulees, une post- ENTOM. 7. 5. 13 242 REVISION DU GENRE EXOCENTRUS MULSANT basilaire, une mediane et une postmediane, et une tache apico-suturale formees par 1'agglomeration de nombreuses petit es taches jaunatres. Long. : 4 mm. 1/2 ; Larg. : 2 mm. Decrit sur des individus du Congo beige : Lualaba, Kaniama, au Musee de Tervueren. 46. Exocentrus fuscosignatus Breuning Exocentrus (s. s.} fuscosignatus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 128. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux sensiblement plus longs que les joues. Pronotum fortement transverse, pourvu d'une epine laterale conique et pointue, dirigee oblique- ment vers 1'arriere et precedee d'une bosse laterale proeminente, mais obtuse. Elytres densement et tres finement ponctues. Brun fonce, couvert de pubescence brune. Elytres revetus de pubescence jaune pale a 1'exception d'une large bande transversale postmediane bran fonce qui n'atteint pas tout a fait la suture et de deux assez grandes taches bran fonce, une basilaire situee a cote de 1'ecusson et une latero-premediane. Tiers basilaire des articles antennaires 3 a 6 couvert de pubescence gris blanchatre. Long. : 4 mm. 1/2 ; Larg. : 2 mm. Decrit sur des individus du Congo beige : Mayidi, au Musee de Tervueren. 47. Exocentrus plagiatus Hintz Exocentrus plagiatus Hintz, 1919, Ergebn. d. II. Centr. Afr. Exped. 1 : 630. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux plus de deux fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une epine laterale pointue fortement recourbee. Elytres densement et tres finement ponctues sur les deux tiers anterieurs. Rouge, couvert de pubescence jaune paille. Elytres revetus de pubescence brun rougeatre, ornes de nombreuses taches jaune blanchatre disposees en series longi- tudinales et couvrant presque entierement la moitie laterale, parsemes de taches minimes denudees disposees egalement en series longitudinales et pourvus, chacun, d'une assez grande tache discale postmediane bran rougeatre fonce en ovale allonge. Articles antennaires a partir du quatrieme revetus de pubescence brun rougeatre, leur partie basilaire a pubescence jaune paille. Long.: 6-8 mm. ; Larg. : 2 mm. 1/4-3 mm. Decrit sur un individu du Congo beige : Sankuru, a ITnstitut Royal des Sciences Naturelles de Belgique. Mayidi (Musee de Tervueren) ; Lac Upemba region (Pares Nationaux) . 48. Exocentrus subplagiatus Breuning Exocentrus (s. s.) subplagiatus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 128. Proche de plagiatus Hintz, mais plus petit, 1'epine laterale du pronotum dirigee plus fortement vers 1'arriere, la tache postmediane foncee de 1'elytre precedee de REVISION DU GENRE EXOCENTRUS MULSANT 243 pubescence blanchatre et les articles antennaires 4 a n sans pubescence jaune paille sur la partie basilaire. Long. : 4 mm. ; Larg. : i mm. 1/2. Decrit sur un individu du Congo beige : Moto, au Musee de Tervueren. 49. Exocentrus bialbomarmoratus Breuning Exocentrus (s. s.) bialbomarmoratus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 129. Antennes aussi longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux sensiblement plus longs que les joues. Pronotum transverse, pourvu d'une petite et mince epine laterale pointue, dirigee obliquement vers 1'arriere, precedee d'une bosse laterale obtuse. Elytres densement et tres fine- ment ponctues sur les deux tiers anterieurs. Noir, couvert de pubescence noire. Ecusson a pubescence blanche. Les elytres presentent des marbrures blanchatres dans la moitie anterieure et sur une bande transversale preapicale. Long. : 3 mm. 1/2 ; Larg. : i mm. 1/4. Decrit sur un individu du Congo beige : Tshela, au Musee de Tervueren. 50. Exocentrus albosticticus Breuning Exocentrus (s. s.) albosticticus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 128. Antennes un peu plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum transverse, pourvu d'une assez longue et mince epine laterale faiblement recourbee. Elytres tres densement et assez finement ponctues. Brun noir, couvert de pubescence brun fonce. Pronotum rouge. Elytres ornes de tres petites taches blanchatres, disposees de fagon a former deux bandes transversales ondulees, un premediane et une postmediane. Partie basilaire des articles anten- naires 3 a ii a pubescence blanche. Long. : 5 mm. ; Larg. : 2 mm. Decrit sur un individu du Congo beige : Elisabeth ville, au Musee de Tervueren. 51. Exocentrus hallei Lepesme & Breuning Exocentrus (s. s.) hallei Lepesme & Breuning, 1955, Bull. Soc. ent. Fr. 60 : 128. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum fortement transverse, pourvu d'une epine laterale pointue, dirigee fortement vers I'arriere. Elytres densement et tres finement ponctues. Brun fonce, couvert de pubescence brun grisatre. Sur chaque elytre sept series longitudinales de petites taches blanchatres. Elytres ornes en plus d'une etroite bande transversale mediane brune en forme de M. Antennes a pubescence brune, le deuxieme article et la partie basilaire des articles 3 a 11 a pubescence blanchatre. Long. : 5 mm. 1/2 ; Larg. : 2 mm. Decrit sur un individu de Cote d'lvoire : Adiopodoume, dans la coll. Lepesme. 244 REVISION DU GENRE EXOCENTRUS MULSANT 52. Exocentrus m-fuscus Breuning Exocentrus (s. s.) m-fuscus Breuning, 1956, Bull. Ann. Soc. Roy. ent. Belg. 92 : 126. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux trois fois plus longs que les joues. Pronotum forte- ment transverse, pourvu d'une epine laterale pointue, dirigee vers rarriere. Elytres densement et tres finement ponctues. Bran fonce, couvert de pubescence jaune paille. Sur chaque elytre trois assez petites taches bran foncee : une latero-posthumerale, une latero-preapicale et une disco-preapicale ainsi qu'une etroite bande transversale mediane bran fonc6 formant avec celle de 1'elytre oppose un grand M ; cette bande, ainsi que la tache disco- preapicale sont bordees en avant de pubescence blanche. Tiers apical des tibias et les tarses bran fonce. Les trois quarts apicaux des articles antennaires 3 a n a pubescence bran fonce. Long. : 8 mm. 1/2 ; Larg. : 3 mm. 1/2. Decrit sur un individu du Congo beige : Tohouapa, Boende, au Musee de Tervueren. 53. Exocentrus ziczac Breuning Exocentrus (s. s.) ziczac Breuning, 1957, Bull. Inst. roy. Sc. nat. Belg. 33, No. 8 : 6, fig. 5. Antennes un peu plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux de moitie plus longs que les joues. Pronotum deux fois plus large que long, pourvu d'une epine laterale pointue fortement recourbee. Elytres densement et tres finement ponctues sur les deux tiers anterieurs. Rouge fonce, couvert de pubescence brane entremelee de jaunatre. Sur chaque elytre trois bandes transversales zigzaguees blanches, une postbasilaire tres peu apparente, une postmediane assez large descendant en oblique en direction de la suture et une preapicale. Quart basilaire des articles antennaires a partir du quat- rieme revfitu de pubescence blanche. Long. : 6 mm. 1/2 ; Larg. : 3 mm. Decrit sur un individu du Cameroun : Ja River, a 1'Institut Royal des Sciences Naturelles de Belgique. 54. Exocentrus freyi Breuning Exocentrus (s. s.) freyi Breuning, 1955, Ent. Arb. Mus. Frey, 1 : 665. Antennes d'un tiers plus longues que le corps, le troisieme article un peu moins long que le scape. Lobes inferieurs des yeux plus de deux fois plus longs que les joues. Pronotum transverse, tres eparsement et tres finement ponctue, pourvu d'une tres petite epine laterale pointue fortement recourbee. Elytres, sauf dans la partie apicale, densement et tres finement ponctues. Bran fonce couvert de pubescence jaune paille. Sur chaque elytre une bande transversale zigzaguee blanche, peu large situee juste apres le milieu. Le tiers basil- REVISION DU GENRE EXOCENTRUS MULSANT 245 aire des articles antennaires a partir du troisieme a pubescence blanche, les deux tiers apicaux de ces articles a pubescence brun fonce. Long. : 8-10 mm. ; Larg. : 3-3 mm. 3/4. Decrit sur un individu de Guinee francaise : N'Zerekore, dans la coll. Frey. Dahomey (coll. Lepesme). FIG. i. Exocentrus occidentalis Breuning. 55. Exocentrus occidentalis Breuning Exocentrus (s. s.) occidentalis Breuning, 1957, Bull. Inst. roy. Sc. nat. Belg. 33, No. 8 : 6. Antennes d'un quart plus longues que le corps, le troisieme article aussi long que le scape. Lobes inferieurs des yeux quatre fois plus longs que les joues. Pronotum fortement transverse, retreci vers la base et pourvu d'une epine laterale pointue fortement recourbee. Elytres tres densement et finement ponctues. Rouge, convert d'une fine pubescence jaune paille. Elytres parsemes de tres petites taches circulaires brun fonce et ornes, chacun, d'une grande tache