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BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY VOU. Vu

1055-1959

PRINTED BY ORDER OF THE TRUSTEES OF

THE BRITISH MUSEUM (NATURAL HISTORY)

LONDON: 1959

DATES OF PUBLICATION OF THE PARTS

No. No. No.

CS PIAWEY Ys

22 July 26 August 26 August 12 September 24 October 31 October 14 October

9 January 30 January

PRINTED IN GREAT BRITAIN AT THE BARTHOLOMEW PRESS DORKING BY ADLARD AND SON, LTD.

1958 1958 1958 1958 1958 1958 1958 1959 1959

CONTENTS ENTOMOLOGY VOLUME VII

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 The African species of Stivalius, a genus of Siphonaptera. By

P.G. A. M. Saur 39 The Pseudococcidae (Hom. : Coccoidea) described by C. K. Brain

from South Africa. By G. DE Lotto 27 Revisions of Mallophaga Genera. Degeeriella from the falconiformes.

By THERESA CLAY I2I Révision du genre Exocentrus Mulsant (Col., Cerambycidae). By S.

BREUNING 209

New and little known Emesinae (Reduviidae, Hemiptera) in the British Museum (Natural History), London. By P. WycopzInsky 329 New species and subspecies of Odonata and some Trichoptera from S. Rhodesia and Portuguese East Africa. By D. E. KImMMINS

347 New or little known Butterflies from Malaya. By Lt.-Cor. J. N. ELIOT 369 A study of the New Zealand Chironomidae (Diptera, Nematocera). By PauL FREEMAN 393

Index to Volume VII 439

rs

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. 1 LONDON: 1958

THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) Pes ripe BY W. |. HALL, F. LAING AND A. H. STRICKLAND FROM THE ETHIOPIAN REGION

BY

D. J. WILLIAMS

Commonwealth Institute of Entomology

Pp. 1-37; 15 Text-figures

BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 7 No.1 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.

Paris appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.

This paper is Vol. 7, No. 1 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 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 pterocaulont, 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. 1§

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 defined by Ferris (19554). Two species described from Egypt, Antonina indica panica and Phenacoccus imermis 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. Anionina 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. 1)

Phenacoccus inermis Hall, 1925, Bull. Minist. Agric. Egypt, 64: 7. Mirococcus inermis (Hall), Borkhsenius, 1947, Proc. Acad. Sci. Armen. S.S.R. 7: 142.

Hasit. Originally described by Hall from material collected near Helwan, Egypt, on the roots of Cleome arabica, Cressa cretica, Frankenia pulverulenta 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 ting 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) 5

Fic. I.

6 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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 Mzrococcus. The total absence of cerarii, the 9-segmented antennae and the denticle on the claw serve to distinguish the species.

Octococcus pentziae Hall (Text-fig. 2) Octococcus pentziae Hall, 1939, J. ent. Soc. S. Afr. 2: 93.

Hasit. 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 I-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) 7

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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 in 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.

Hasit. 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 15mm. 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) 9

FiG. 3

10 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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 antennal 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. solant 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, J. ent. Soc. S. Afr. 2: 96.

Hasit. 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-

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12 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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.

Hapsit. 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) 13

Fie. 5

14 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. Multilocular 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. phoradendnri (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.

Hasit. 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)

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16 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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 9-segmented antennae, with a denticle on the claw and with ventral quinquelocular pores. As rhodestensis 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 inyazuvae Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 131.

Hasit. 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) 17

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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 imyazurae 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. Lond. 86 : 133.

This species was described by Hall from Salisbury, Southern Rhodesia on Pterocaulon decurrens and Trifolium sp. It is identical, however, with Tvionymus sanguineus described by James (1936) to which the name Trionymus pterocaulon Hall is here sunk asasynonym. De Lotto (1957) has already si peat te 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 (PSEUDOCOCCIDAE : 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 AHeliococcus phaseolt, Pseudococcus hargreavest and Pseudococcus ugandae.

Heliococcus phaseoli (Laing) (Text-fig. 8)

Phenacoccus phaseoli Laing, 1929, Ann. Mag. nat. Hist. (10) 4: 475. Heliococcus phaseolt (Laing), Goux, 1934, Bull. soc. ent. Fr. 39: 171.

Hasit. 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 g-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. Maultilocular 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

HOMOPTERA)

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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 phaseolt 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 ritchiet 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 mjalensis 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.).

Hasit. 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 asasynonym. 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

Fic. 9

HOMOPTERA)

23

24 THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA)

Pseudococcus ugandae Laing (Text-fig. 10) Pseudococcus ugandae Laing, 1925, Bull. ent. Res. 16: 53.

Hapsit. 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 9-segmented.

THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) 25

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26 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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, 1947a) 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, Rev. Path vég. 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 lorantdi 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. 11) Pseudococcus celtis Strickland, 1947, Proc. R. ent. Soc. Lond. (B) 16: 154.

Hapsit. 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 (PSEUDOCOCCIDAE : HOMOPTERA) 27

Fic. Ir

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 ina doublerow. 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 swbukiaensis 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.

HasitT. 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

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30 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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.

Hapit. This species was described from Tafo, Eastern Province, Gold Coast on Musanga smithit (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 (PSEUDOCOCCIDAE : HOMOPTERA) 31

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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.

Hasit. 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

Fic. 14

34 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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.

Hasit. 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 I'5mm.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 (PSEUDOCOCCIDAE : HOMOPTERA) 35

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36 THE MEALY-BUGS (PSEUDOCOCCIDAE : 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 TJ. malacanthae Strickland, already discussed, in possessing numerous dorsal circular disc pores and extremely long setae with the distal ends flagellate.

REFERENCES

Bavacuowsky, A. S. 1954. Sur l’Indigénat et le Statut de Catenococcus lovanthi Strickl. (Coccoidea : Pseudococcini) en Afrique Occidentale. Rev. Path. vég. 33: 247.

BorKHSENIUS, N. S. 1949. 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 (Hom. : Coccoidea) described by H. C. James from East Africa. Bull. Brit. Mus. (Nat. Hist.) Ent. 5 : 185-232.

Ezzatt, Y. M. & McConne.i, 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. Jbid.6. Stanford University, California. 1955. On some genera of the Pseudococcidae. Microentomology, 20: 1-6. —— 1955a. Atlas of Scale Insects of North America, 7. Stanford University, California.

THE MEALY-BUGS (PSEUDOCOCCIDAE : HOMOPTERA) 37

Goux, L. 1934. Notes sur les Coccides (Hem.) de la France. (9% note). Contribution 4 1 étude du genre Heliococcus avec description de deux espéces nouvelles. Bull. soc. ent. Fr. 39 : 164-171.

Hatt, W. J. 1937. Observations on the Coccidae of Southern Rhodesia. Tvans. R. ent. Soc. Lond. 86 : 119-134.

— 1941. On some new species and two new genera of Coccidae (Homoptera) from Southern Rhodesia, J. 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. Tvrans. R. ent. Soc. Lond. 85 :197-216.

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.

1947a. Three new species of Coccoidea (Hemiptera : Homoptera) from the Gold Coast,

British West Africa. Proc. R. ent. Soc. Lond. (B) 16: 149-156.

Wituiams, 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.

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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

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THE AFRICAN SPECIES OF STIVALIUS, A GENUS OF SIPHONAPTERA

BY

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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), «stituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical Series.

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This paper is Vol. 7, No. 2 of the Entomological serves.

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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! 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 S. 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. 1); g—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 preapicaily (Text-fig. 16); 9Q—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)

ferinus-group, p. 42

(6) Genal margin below the eye entire (Text-fig. 2) ; g—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); 9Q—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) . . : : torvus-group, Pp. 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.

? 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. yhaebus Jordan (Borneo) and S. lonchus Jordan (Borneo)) ; ahalae-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. mjébergi Jordan (Borneo).

ENTOM. 7, 2 3

42 THE AFRICAN SPECIES OF STIVALIUS

FERINUS-GROUP

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. 1, 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 ; 1 female paratype from the same locality, from Rattus (Mastomys) natalensis, vi.1953; 2 female paratypes, nr. Elisabethville, from Rattus (Mastomys) natalensis, vill. 1953 ; all collected by Dr. R. Devignat, to whom one pair of paratypes has been returned; 1 male paratype from the neighbourhood of Elisabethville, from Crocidura pilosa, 1957; I female paratype, same locality, from Mus triton, 1957; 1 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 Musée Royal du Congo Belge, Tervuren; I male paratype from the Suji Valley, 6,000 ft., S. Pare Mts., Tanganyika, i.1957, from Arvicanthis sp., collected by J. G. Halcrow.

Dracnosis. A member of the ferinus-group, which includes S. ferinus (Rothschild) (a shrew-parasite from Ceylon and India), S. insolli Traub (a bird-parasite from Malaya) and S. aestivalis Jameson & Sakaguti (a wood-mouse (Apodemus) parasite from Japan). The new species differs from S. ferinus by the unmodified (not sub- spiniform) setae in the submarginal frontal row, from S. imsolli and S. aestivalis by the absence of a row of short setae preceding the main row of setae on the pronotum, while in S. insollt the number of pronotal spines is about 30 as against 20 in S. alienus. There are also differences in the genitalia between these species.

DESCRIPTION. HEAD (Text-fig. 1). 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. 1) 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.

Lecs. 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 38

44

THE AFRICAN SPECIES OF STIVALIUS

Fics. 1, 2. Head and pronotum of: 1. 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). 4.5. ¢ovvus (Rothschild) (female, Keruguya, Kenya).

THE AFRICAN SPECIES OF STIVALIUS

6 7

Fics. 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. Elisabethville, b—paratype, plantation Calonne).

45

40 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. <¢ 23-2? mm., 2 3-34 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.1)

THE AFRICAN SPECIES OF STIVALIUS 47

and in S. 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 S. 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 S. imsolli (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)

Pygiopsylia torvus Rothschild, 1908, Ent. mon. Mag. 44: 77. Pygiopsylla “ afer ’’ Jordan & Rothschild, 1913, Novit. zool. 20 : 537 (err. det., 1 9 from Kagamba,

Uganda; see Jordan & Rothschild, 1922, Ectoparasites, 1 : 252).

Stivalius “‘ afer’’ Symes & Hopkins, 1932, Rec. Med. Res. Lab. Nairobt (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, 39:11; 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.

vats, fleas, plague, Uganda: 9, tables 2, 6, 10, 12, 20.

MATERIAL EXAMINED. TANGANYIKA: Tengeru, Rattus (Mastomys) natalensis, 26.11.1952, 1 dg, 19. Kenya: Keruguya, Rattus (Mastomys) natalensis, Lobhuromys flavopunctatus aquilus, Lemniscomys sp., Otomys sp., 1935-36, 10 g 8 9; without locality and host, 1927, 1 2, and 1913, I g, 3 2; Machakos, house-rat, 23.v.1927, 2 2 (Jordan, 1936: 297); Kisii, house-rat, 23.v.1927, 1 9. UGANDA: Mubuku valley, E. side of Ruwenzori, 6,000 ft. Grammomys dryas, 1.iii.1906, the male holotype (Rothschild, 1908:77; Jordan & Rothschild, 1922: 252); Kanungu, Kigezi, Lophuromys flavopunctatus aquilus, 2.x.1940,1 9; same locality, Arvicanthis abyssinicus, x.1940, I 2; Kagamba, Kazara county, Ankole, Dasymys incomtus medius, 14.vii.IgiI, i 9 (Jordan & Rothschild, 1913: 537, as “ afer’’; 1922: 252) ; Damba Island, Lake Victoria, young rodent in nest, xi. Ig1I, I 9. BELGIAN CONGO (specimens in the Musée Royal du Congo Belge, Tervuren, are marked with (MCB)) : Irumu district, Ituri, Avvicanthis abyssinicus, 15.vili.1946, 3 9 (MCB); Kunga, Mt. Wago, Ituri, nest of Oenomys hypoxanthus, 23.vii.1946, 1 2 (MCB); Pikanza, Ituri, Rattus (Mastomys) natalensis ugandae, 14.v.1946, 1 g§ (MCB); Blukwa, Ituri, Rattus (Mastomys) natalensis ugandae, 13.xi1.1945, I 2 (MCB); Blukwa,

48

THE AFRICAN SPECIES OF STIVALIUS

Figs. 8,9. Clasper and sternum IX of: 8. Stivalius torvus (Rothschild) (Keruguya, Kenya). 9. S. parilis sp. n. (holotype).

THE AFRICAN SPECIES OF STIVALIUS 49

rats, 1931, 1 2 (MCB); Blukwa, 1940, 1 9 (MCB); Blukwa, Rattus (Mastomys) natalensis ugandae, ii. 1935, 1 3; Lombu nr. Blukwa, Rattus (Mastomys) natalensis ugandae, 19.xii.1945, I 2 (MCB) ; Rakwa nr. Blukwa, Rattus (Mastomys) natalensis ugandae, 9.X.1945, I g¢ (MCB) ; Drodro nr. Blukwa, Arvicanthis abyssinicus 2.ix.1945,1 6 (MCB); Lotomukubwanr. Blukwa, Arvicanthis abyssinicus, 15.X.1945, 1 ¢ (MCB); Costermansville, Kivu district, rats, 1940, 1 g, 1 9; Lubero, Kivu, field rats, xii.1940, 3 d, 3 2; Lubero, 1950, 2 g, 1 9; Boyoro region, Geti-Boga, Rattus (Mastomys) natalensis ugandae, iv.1929, I g, 1 9 (andi g, 1 9 MCB) ; Dilolo, Katanga, Tatera valida, Rattus (Mastomys) natalensis, 1934, 2 g, 2 9 (and I g, MCB) (Jordan, 1936: 297); Lukolela, Tshuapa district, Crocidura occidentalis, 1 ¢ (Jordan, 1937: 290); Elisabethville, Katanga, 1935, 1 2; the following are all from the neighbourhood of Elisabethville: Crocidura pilosa, 1 2; Steatomys pratensis, 1 3; Saccostomus campestris, 5 3, 2Q; Rattus (Praomys) morio jacksont, 24,49; Tatera valida,1 9; Aethomys chrysophilus,1 9; Mus triton, 19; Pelomys fallax, 22; Rattus (Mastomys) natalensis ugandae, 12 3, 169; Lophuromys flavo- punctatus major, 1 3, 2 2 (all these specimens were collected by P. L. Pirlot in 1957); nest of Steatomys pratensis, x.1956, 20 3, 32 2; Saccostomus campestris, 12.iv.1956, I g; Rattus (Mastomys) natalensis, iii.1956, I g, iv.1956, I ¢ (these. specimens were collected by M. Lips; part of the Pirlot and Lips material has been generously presented by the Tervuren Museum to the British Museum collection of fleas at Tring).

DESCRIPTION. HEAD (Text-fig. 2). Frontoclypeal margin smoothly rounded. Preoral tuber short, though a little longer than in S. alienus (Text-fig. 1). Submarginal frontal row consisting of six setae in both sexes ; between this row and the eye there are 12-16 large and fairly large setae, while there are many minute setae situated between the frontal row and the next irregular row and also along the anterior margin of the antennal fossa. Genal margin below the eye entire, not divided into two small lobes. Eye well-developed, kidney-shaped. Maxillary palp reaching to about the middle of the anterior margin of the fore coxa; the first two segments of this palp of subequal length, the third segment the shortest. The laciniae are smooth basally and finely serrated apically. The labial palp, reaching to about three-fourths the length of the fore coxa, consists of five segments. Scapus of antenna with six to nine setae on the outer surface of its widened portion in the male and three to four in the female ; pedicellus in both sexes with six to nine slender setae, several of which reach to or a little beyond the first segment of the clava. Postantennal region of head with three rows of setae and a large seta about mid-way between the lowest seta of the second row (the latter seta is moved considerably forward) and that of the third row ; in both sexes the first occipital row consists of six setae, the second and third rows of six or seven. Bordering the antennal fossa posteriorly are about 16-20 small setae in the male, 20-25 in the female.

TuHorax. Pronotum (Text-fig. 2) dorsally about as long as the dorsal spines of the ctenidium, with two rows of setae, the anterior of which does not extend down- wards on to the ventral half of the pronotum and consists of five to seven short setae each side, while the main row consists of six setae each side in both sexes. Pronotal ctenidium with 22-24 spines (occasionally 20, 21 or 25). Mesonotum with

50

THE AFRICAN SPECIES OF STIVALIUS

Figs. 10, 11. Clasper and sternum IX of: 10. Stivalius timanus Jordan (holotype). 11. S. fallaciosus sp. n. (holotype).

THE AFRICAN SPECIES OF STIVALIUS 51

a main row of five setae each side, preceded by three rows of smaller setae ; two fairly long pseudosetae present under the dorsal portion of the mesonotal collar. Mesepisternum with three to five small and one large seta, the latter is often placed opposite the internal rod or even beyond this and is then actually on the mesepimeron ; mesepimeron with six or seven setae. Metanotum with four rows of setae, the first row being short and irregular; the main row has six setae each side, the lowest of which is the shortest. Pleural arch well developed. Metepisternum with one large and one or two small setae ; metasternum dorso-posteriorly with one large and one or two small setae ; metepimeron with about five small and nine to ten large setae in the male, five or six small and 11-13 large setae in the female.

Lecs. Chaetotaxy and structure of coxae and femora more or less as in S. alienus. Fore tibia with seven groups of setae in notches of the posterior (dorsal) margin, the groups consisting from base to apex of 3, 3, I, 3, 3, 3 and 3 setae respectively ; mid tibia with eight groups: 2, 2, 2, 2, 3, I (usually not in a notch), 3 and 3 setae ; hind tibia (Text-fig. 4) with eight groups: 2, 2, I (or 2), 3, 3, I, 3 and 3 setae respec- tively. Fifth segment of all tarsi with the chaetotaxy usual for the genus.

ABDOMEN. Tergum I with four rows of setae; terga II-VII with three rows each, but the first row often very irregular. The numbers of setae in the main row on each side of terga I-VII are in the male 5, 7, 7, 7, 7, 7 and 7 respectively, in the female 5, 7, 8, 8, 8, 7 and 4 (or 5). Terga II-V in both sexes with one subdorsal marginal spinelet each side. Both sexes with two antesensilial setae, the lower of which is about twice the length of the upper ; in the female the margin of tergum VII between the two groups of antesensilials forms a triangular lobe; below the antesensilials the margin is angulate. Basal abdominal sternum with a lateral patch of one to five small setae in the male and 12-25 in the female, and along the ventral margin four setae each side forming a horizontal row. Sterna IJI-VII in the male normally with three large setae on each side in the rather irregular main row; in the female the main row of sterna III-VI consists of three or four setae; in both sexes these main rows are preceded by a patch or numerous smaller setae.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 8, 17). Tergum VIII with four to six setae each side anterior to the vertical part of the spiracular fossa. Sternum VIII with about 30-35 setae each side and a group of three stout and two small closely set setae apically at the ventral margin. Apodeme of tergum IX narrow in its upper part, its lower portion not solidly fused with the corpus of the clasper (Text-fig. 8). Manubrium broad, tapering only weakly, with an upturned apex. Fixed process of clasper with one long and one short acetabular seta. A dark T-shaped sclerite present between the bases of the two manubria. Movable process of the shape which is usual in the genus, long and narrow and with its apical portion turned downwards ; with an area of densely set thin and moderately long setae on the inner side bordering the ventral margin, also along this margin, but beyond the bend, are four or five (sometimes only three) long setae. The ventral margin of the movable process gently convex in the setose portion (Text-fig. 8) or practically straight. Distal arm of sternum IX (Text-fig. 8) apically widened, with numerous short setae, many of which are thickened, along the apical margin and unmodified ones on the adjoining lateral portion ; dorsally the apical part of the arm bears

ENTOM. 7, 2 38§

52

Fics. 12, 13. Clasper and sternum IX of: 12. Stivalius cacuminis sp. n. (holotype). 13. S. vichardi Jordan (holotype).

THE AFRICAN SPECIES OF STIVALIUS 53

a patch of very small setae on the inner side. Phallosome as in Text-fig. 17; the tendons of the phallosome make about half a convolution, the inner tube is short and thick-walled and the dorso-apical aedeagal sclerite is of a characteristic shape and structure ; the aedeagal apodeme is very broad but preapically the dorsal margin is strongly concave.

FEMALE (Text-figs. 26, 28, 38, 39, 53, 54). Posterior margin of sternum VII with a large sinus, which divides the main row of four or five setae into two upper and two or three lower; in front of this main row are numerous smaller setae (Text-figs. 38, 39). Variation in the outline of the posterior margin of sternum VII is as shown in Text-figs. 38, 39. Tergum VIII with about 8-12 setae in front of the vertical part of the spiracular fossa ; chaetotaxy of the ventral part of this tergum as in Text-fig. 53. The posterior margin of tergum VIII has a marked noselike projection, the variation of which is shown in Text-fig. 54. Sternum VIII as in Text-fig. 28. Anal stylet about four to five times as long as its maximum width, with one long apical and two minute preapical setae. Bulga of spermatheca (Text-figs. 26, 28) roughly rectangular, dorsally with a smoothly rounded but rather variable hump ; the hilla, bearing a papilla, is much narrower and also shorter than the bulga ; both the bulga and the the basal half of the hilla have thick or very thick walls. On each side of the bursa copulatrix is a dark and usually triangular (not longitudinal) sclerite (Text-figs. 26, 28). The relatively short dilated part of the ductus spermathecae has a thick wall and is provided internally with numerous membranous rings (Text-fig. 26).

LENGTH. ¢ 24-34 mm., 2 3-4 mm.

Stivalius richardi Jordan, 1936 (Text-figs. 13, 21, 30, 41, 61) Stivalius vichardi Jordan, 1936, Novit. zool. 39 : 298, figs. 57, 58.

MATERIAL EXAMINED. Male holotype from Dilolo, Katanga, Belgian Congo, from Malacomys longipes, 1934; female allotype, same locality and date, from Steatomys pratensis. The allotype is in the Musée Royal du Congo Belge at Tervuren.

DESCRIPTION. HEAD. Approximately as in S. torvus, but the frons is a little more strongly rounded, and there are fewer small setae bordering the antennal fossa dorsally (14-15 in the male, 17-20 in the female).

THORAX. Pronotal ctenidium in both sexes consisting of 24 spines which are somewhat shorter than the pronotum ; in other respects the thorax resembles that of S. torvus.

Lecs. Fore tibia with seven groups of setae in notches of the posterior (dorsal) margin, the groups consisting from base to apex of 2, 2, I, 3, 3, 2 and 3 setae respec- tively ; mid tibia with eight groups: 3, 2, I, 3, 3, I, 3 and 3 setae; hind tibia with eight groups: 3, 2, 2, 3, 3, I, 3 and 4 setae.

ABDOMEN. Segments I-VII as in S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 13, 21). Tergum VIII with five to eight setae each side in front of the vertical part of the spiracular fossa. Sternum VIII with about 40 setae each side and the usual ventro-

54

THE AFRICAN SPECIES OF STIVALIUS

Fics. 14, 15.

Clasper and sternum IX of: 14. Stivalius sellatus Jordan & spl unin (Torit, Sudan). 15. S. pirloti sp. n. (holotype).

THE AFRICAN SPECIES OF STIVALIUS 55

marginal group of several short and stout setae. Apodeme of tergum IX as in S. torvus; manubrium broad, tapering more distinctly than in S. torvus; fixed process of clasper with one short and one long acetabular seta; movable process (Text-fig. 13) almost like that of S. ¢torvus but longer and slenderer and the dense group of setae not extending nearly so far away from the margin in S. richardt. Apex of distal arm of sternum IX (Text-fig. 13) produced dorsally into a long and sharply pointed spike ; chaetotaxy of the distal arm as shown in Text-fig. 13. Aedeagus as in Text-fig. 21; the tendons of the phallosome make about half a convolution, the short inner tube has a very thick dorsal wall and the dorso-apical aedeagal sclerite is very large.

FEMALE (Text-figs. 30, 41, 61). Sternum VII as in Text-fig. 41. The ventral and posterior margins of tergum VIII meet at about a right angle without forming a projection ; noselike projection of the posterior margin of tergum VIII rounded-off (Text-fig. 61) ; chaetotaxy of this tergum approximately as in S. torvus. Sternum VIII rather narrow apically (Text-fig. 30). Anal stylet only three and a half times as long as its maximum width. Bulga of spermatheca (Text-fig. 30) thick-walled, almost without a dorsal hump ; hilla with a large papilla. The sclerite on each side of the bursa copulatrix is exceptionally large and dark and is shaped like the sole of a shoe, in the only known specimen, covering the short dilated part of the ductus spermathecae almost completely (Text-fig. 30).

LENGTH. ¢ 34mm., 294mm.

Stivalius curtiductus sp. n. (Text-figs. 29, 40, 62)

TYPE MATERIAL. Female holotype from Mt. Tonkoni nr. Man, Ivory Coast, g00-1,200 m., 20-30.ix.1946, A. Villiers.

DraGnosis. The female of this new species (the male is unknown) is easily distin- guishable from other species of the torvus-group by the pronotal spines being much shorter than the pronotum, the far downwards extension of the first row of setae on the pronotum, and the sharp ventro-apical angle of tergum VIII.

DeEscrIPTION. HEAD. Eye kidney-shaped, but a little more rounded than in S. torvus. The laciniae are somewhat more coarsely serrated than in the latter species, and the serration extends from the tip of the laciniae to about the middle. In other respects the head resembles that of S. torvus.

THORAX. Pronotum dorsally considerably longer than the spines of the pronotal ctenidium, with two rows of setae, the first row of eight or nine setae each side extends downwards well below the middle of the pronotum ; the second row consists of six setae each side; pronotal ctenidium with 23 spines. Mesothorax and meta- thorax essentially as in S. torvus.

Lecs. The fourth group of setae (counted from the base) of the mid tibia bears three setae (two in S. torvus) ; otherwise the legs are like those of S. torvus.

ABDOMEN. Dorsally in front of the three rows of tergal setae are several irregularly

56 THE AFRICAN SPECIES OF STIVALIUS

Fics. 16-18. Phallosome of: 16. Stivalius alienus sp. n. (holotype). 17. S. torvus (Rothschild) (Keruguya, Kenya). 18. S. timanus Jordan (holotype).

THE AFRICAN SPECIES OF STIVALIUS 57

placed setae, while the first two rows are also rather irregular. Tergum VI with eight setae in the main row each side (seven in S. torvus). The upper of the two antesensilial setae is about half the length of the lower on one side of the specimen and slightly more than half on the other. Basal abdominal sternum with a lateral patch of about 25 setae each side; the main row on sterna III-VI consists of four or five setae. In other respects the unmodified abdominal segments are like those of S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. FEMALE (Text-figs. 29, 40, 62). Posterior margin of sternum VII (Text-fig. 40) with a rather deep sinus, the lobe above it is obtuse. Tergum VIII with 12 or 13 setae in front of the spiracular fossa. Noselike projection of the posterior margin of tergum VIII weakly developed ; the ventro-apical angle of this tergum very sharp (Text-fig. 62). Sternum VIII as in Text-fig. 29. Anal stylet nearly five times as long as its maximum width, with two minute preapical setae and a fairly long apical seta. Bulga of spermatheca with a fairly thick wall, and a small dorsal hump; the sclerite on each side of the very small bursa copulatrix is almost undeveloped (Text-fig. 29). The dilated basal portion of the ductus spermathecae is broad and short and resembles that of S. torvus, but it has a thin wall; the remainder of this duct is also very short (Text-fig. 29).

LENGTH. 2° 4mm.

Stivalius vancanneyti Berteaux, 1947 (Text-figs. 31, 42, 55)

Stivalius vancanneyti Berteaux, 1947, Rev. Zool. Bot. Afr. 40: 105, fig. 9.

MATERIAL EXAMINED. Female holotype, Djugu, Ituri, Belgian Congo, from Heltosciurus ruwenzori, collected by R. van Canneyt. The holotype is in the Musée Royal du Congo Belge, Tervuren.

DESCRIPTION. HEAD. Frons rather strongly rounded ; chaetotaxy of head similar to that of S. torvus. The five-segmented labial palp reaches to the apex of the fore coxa.

THORAX. Pronotum dorsally one and a third longer than the dorsal spines of the pronotal ctenidium ; with two rows of setae, the main row consisting of five setae each side. Pronotal ctenidium consisting of 22 spines. In other respects the thorax is similar to that of S. torvus.

Lecs. Chaetotaxy approximately as in S. torvus; in the fourth notch from the base of the hind tibia the anterior of the three setae is displaced downward on both sides of the specimen, but this is possibly abnormal.

ABDOMEN. Unmodified segments as in S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. FEMALE (Text-figs. 31, 42, 55). Sternum VII as in Text-fig. 42. A patch of ten setae each side anterior to the spiracular fossa of tergum VIII; ventro-posterior angle of tergum VIII (Text-fig.

58 THE AFRICAN SPECIES OF STIVALIUS

Figs. 19-24. Aedeagus of: 19. Stivalius parilis sp. n. (holotype). 20. S. sellatus Jordan & Rothschild (Torit, Sudan). 21. S. richardi Jordan (holotype). 22. S. falla- ciosus sp. n. (holotype). 23. S. pirloti sp. n. (holotype). 24. S. cacuminis sp. n. (holotype).

THE AFRICAN SPECIES OF STIVALIUS 59

55) rounded off, not forming a projection ; posterior margin of tergum VIII with a well-developed noselike projection. Sternum VIII as in Text-fig. 31. Anal stylet about four times as long as basally broad. Genitalia as in Text-fig. 31 ; the dilated portion of the ductus spermathecae as wide as in S. torvus, richardi and curtiductus but more than twice as long as in any of these three species. Spermatheca so unfa- vourably placed that its characteristics cannot be made out satisfactorily.

LENGTH. 9 44 mm.

Stivalius afer (Rothschild), 1908 (Text-figs. 32, 43, 56)

Pygiopsylla afer Rothschild, 1908, Proc. zool. Soc. Lond. (1908) : 618, Pl. 29, figs. 7, 8.

Stivalius afer Jordan & Rothschild, 1922, Ectoparasites, 1 : 250, 264, fig. 240 (partim ; the descrip- tion and figure of the male pertain to the next new species) ; Dalla Torre, 1924, Ber. naturw. med. Ver. Innsbruck, 39: 11 ; Bedford, 1932, Rep. Vet. Res. S. Afr. 18: 449 (partim) ; Jordan, 1936, Novit. zool. 40:93 (partim); Jordan, 1948, in Smart, Insects of medical importance (London) : 240.

Stivalius afer afer Jordan, 1938, Novit. zool. 41: 115, 116.

MATERIAL EXAMINED. Female holotype from Benguela Province, Angola, 200 miles from the coast, 4,780 ft., host not known, F. C. Wellman.

DESCRIPTION. HEAD. As in S. torvus, but in the only known specimen the eye is oval, not kidney-shaped.

THorAx. As in S. torvus; pronotal ctenidium consisting of 24 spines; mete- pimeron on one side with 11 small and 11 larger setae, on the other with nine small and 12 larger setae.

Lecs. Essentially as in S. torvus, but on the dorsal margin of the mid tibia the fourth notch from the base has three setae instead of two and the single seta of the sixth group is marginal.

ABDOMEN. Terga IJ-VII with four rows of setae, but the first row is represented by only a few dorsal setae ; in other respects the unmodified abdominal segments are like those of S. torvus.

MoDIFIED ABDOMINAL SEGMENTS AND GENITALIA. FEMALE (Text-figs. 32, 43, 56). Sternum VII as in Text-fig. 43. The noselike angle of the posterior margin of tergum VIII (Text-fig. 56) is less marked than in S. torvus. Apex of sternum VIII with several fairly long setae (Text-fig. 32). Dorsal margin of the bulga of the sperma- theca without a definite hump but with a concavity defining an ill-marked postero- dorsal rounded bulge ; the base of the hilla protruding slightly into the lumen of the bulga (Text-fig. 32). The sclerite on each side of the bursa copulatrix is long and nar- row, much longer than the bursa. The dilated portion of the ductus spermathecae long and relatively narrow, with a thin wall (Text-fig. 32).

LENGTH. 9 3? mm.

60

THE AFRICAN SPECIES OF STIVALIUS

FIGs. 25, 26.

Ni]

MU Winn Ij

MN ayiy Mn MiMi Niitiaee U]) siti) Mi

Vitis ¢

Female genitalia of: 25. Stivalius alienus sp. n. (paratype, nr. Elisabethville) 26. S. torvus (Rothschild) (Keruguya, Kenya).

THE AFRICAN SPECIES OF STIVALIUS 61

Stivalius parilis sp. n.

(Text-figs. 9, 19, 33, 44, 57)

Stivalius “‘ afer’’ Jordan & Rothschild, 1922, Ectoparasites, 1: 250, 264, fig. 240 (partim ; specimens from N’Dala Tando, Angola); Bedford, 1932, Rep. Vet. Res. S. Afr. 18: 449 (partim) ; Jordan, 1936, Novit. zool. 40 : 93 (partim).

TYPE MATERIAL. Male holotype, female allotype, one male paratype and one female paratype from N’Dala Tando (also spelt Dalla Tando or Dala Tando), Loanda, Angola, from Arvicanthis niloticus rufinus, 18.xii.1908, W. J. Ansorge.

Diacnosis. Nearest related to Stivalius afer and differing from it in the female sex (the male of S. afer is unknown) especially by the fact that the bursa copulatrix is as large as the sclerites associated with it. Some other apparent differences are mentioned in the description.

DEscriPTION. Head, thorax, legs and unmodified abdominal segments as in S. afer, except that the eye is kidney-shaped instead of oval, the single seta of the sixth group of the mid tibia is not marginal and the dorso-marginal spinelets of tergum V are small or absent. The pronotal ctenidium consists of 24 spines in both sexes, as in the only known specimen of S. afer.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 9, 19). Tergum VIII with four or five setae each side anterior to the vertical part of the spiracular fossa. Sternum VIII with about 31-36 setae each side and a group of three stout and several smaller ventro-apical setae. Apodeme of tergum IX very narrow in its dorsal portion, its lower portion not solidly fused with the rest of the clasper (Text-fig. 9). Manubrium fairly broad, feebly tapering, with an upturned apex. Fixed process of clasper with one long and one short acetabular seta. The long straight part of the movable process is slightly convex dorsally and ventrally, with a dense patch of thin setae on the inner surface along the posterior part of the straight ventral margin; the ventral margin of the turned-down tip bears three or four long setae, the most distant of which is not far from the apex. Apex of distal arm of sternum IX only slightly widened, with a bluntly triangular dorso-apical angle and a number of lateral and marginal setae (Text-fig. 9). Aedeagus as in Text-fig. 19; dorsal wall of inner tube extremely thick ; ventro-lateral projection of the outer wall of the aedeagus with an acute ventro-posterior angle ; aedeagal apodeme broad but the dorsal margin strongly concave pre-apically, the apex pro- duced into a tendril-like structure; the tendons of the phallosome form several convolutions.

FEMALE (Text-figs. 33, 44, 57). Posterior margin of sternum VII (Text-fig. 44) with a large sinus, which divides the main row of six setae into two groups, each of three setae, in front of this main row are numerous smaller setae. Tergum VIII with about six to eight setae in front of the vertical part of the spiracular fossa ; chaeto- taxy of the lower part of tergum VIII similar to that of related species. Noselike angle of the posterior margin of tergum VIII rounded off (Text-fig. 57). Sternum VIII as in Text-fig. 33. Anal stylet with a long apical seta and only one minute preapical seta at the dorsal margin. Bulga of spermatheca (Text-fig. 33) rather long, with a relatively thin wall and without a marked dorsal hump ; hilla with a papilla.

62

THE AFRICAN SPECIES OF STIVALIUS

FE

died Pt ae

3|

Fic. 27. Stivalius pirloti sp. n. (allotype), terga II-IV. Figs. 28-31. Sternum VIII and IX and genitalia of: 28. S. torvus (Rothschild) (Keruguya, Kenya). 29. S. curti- ductus sp. n. (holotype). 30. S. richardi Jordan (allotype). 31. S. vancanneyti

Rartasauw fhala+éerna\

THE AFRICAN SPECIES OF STIVALIUS 63

The sclerite on each side of the bursa copulatrix is roughly ellipsoid and about as large as the bursa ; the dilated part of the ductus spermathecae is long and narrow and has fairly thick walls.

LENGTH. ¢ 34 mm., 2 34-4 mm.

Stivalius timanus Jordan, 1938 (Text-figs. 10, 18, 34, 45, 46, 58)

Stivalius afer timanus Jordan, 1938, Novit. zool. 41: 115, figs. 60, 61.

MATERIAL EXAMINED. From Georgewater Spur!, Freetown, Sierra Leone: male holotype from Rattus (Praomys) morio tullbergi1, 27.11.1937, female allotype from the same host species, I.iii.1937, one female paratype from Lophuromys sikapusi', 25.11.1937; all collected by D. H. S. Davis.

DESCRIPTION. HEAD. Frons more strongly rounded than in related species. In other respects the head resembles that of S. torvus.

THORAX. Pronotal ctenidium with 23 spines in the male holotype, 24 in the two available females ; the dorsal spines are a little shorter than the dorsal length of the pronotum. The lower of the two pseudosetae each side under the collar of the mesonotum is separated by a clear gap from the upper.

LEGS AND UNMODIFIED ABDOMINAL SEGMENTS. As in S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 10, 18). Tergum VIII of holotype with three setae on one side, four on the other ; sternum VIII with 31 setae on one side, 36 on the other, and the usual group of four to seven short setae apically along the ventral margin. Apodeme of tergum IX as in related species. Manubrium broad, with an upturned tip. Three acetabular setae on one side, two on the other (the latter is doubtless the normal number). Movable process (Text-fig. 10) long and slender, with a straight dorsal margin and a smoothly undula- ting ventral margin; with a dense patch of longish thin setae on the inner side adjacent to the convex part of the ventral margin ; the ventral margin of the turned- down apical portion with three long setae, the most apical of which is well away from the apex. Distal arm of sternum IX (Text-fig. 10) ventro-apically strongly widened without a dorso-apical projection, with a number of short setae several of which are spiniform. Phallosome as in Text-fig. 18 ; the tendons of the phallosome make two convolutions ; the aedeagal inner tube has a relatively thin dorsal wall ; the dorso-apical sclerite is rather small; the aedeagal apodeme is broad, the apical half of its dorsal margin is strongly concave, and its tip is drawn out into a long tendril-like structure.

FEMALE (Text-figs. 34, 45, 46, 58). Sternum VII with the sinus almost as in related species, but the lobe above the sinus relatively long ; variation in the outline of the posterior margin of sternum VII as in Text-figs. 45, 46. Tergum VIII with five to seven setae in front of the vertical portion of the spiracular fossa. Noselike angle of the posterior margin of tergum VIII distinct (Text-fig. 58), but less so than in S. torvus. Sternum VIII rather broad, narrowing abruptly a little before the apex

1 The names of the exact locality and of the hosts, as well as the dates of capture, were not available for the original description, but recently Dr. D. H. S. Davis has kindly supplied these additional data.

64

THE AFRICAN SPECIES OF STIVALIUS

o

Fics. 32-37. Sternum VIII and IX and genitalia of: 32, Stivalius afer (Rothschild) (holotype). 33. S. parvilis sp. n. (allotype). 34. S. timanus Jordan (allotype). 35. S. nigeriensis Jordan (holotype). 36. S. sellatus Jordan & Rothschild (holotype). 37. S. pirloti sp. n. (allotype).

THE AFRICAN SPECIES OF STIVALIUS 65

(Text-fig. 34). Anal stylet four or five times as long as wide, with the usual long apical seta but only one preapical minute seta. Bulga of spermatheca (Text-fig. 34) short, very thick-walled, posteriorly with a very prominent hump. The sclerite on each side of the bursa copulatrix is small and somewhat constricted in its middle ; it is as long as the small bursa copulatrix (Text-fig. 34). The ductus spermathecae very long, but the dilated portion of the duct (Text-fig. 34) of moderate length, not unlike that of S. vancanneyti but relatively much narrower. LENGTH. ¢ 34 mm., 2 4-4} mm.

Stivalius nigeriensis Jordan, 1938

(Text-figs. 35, 47, 48, 59)

Stivalius afer nigeriensis Jordan, 1938, Novit. zool. 41 : 116, fig. 62.

MATERIAL EXAMINED. Female holotype from Lagos, Nigeria, from a bush rat, S. L. M. Counal; one female from Adiopodoumé, Ivory Coast, from Crocidura sp., 17.iv.1953, V. Aellen.

DESCRIPTION. HEAD. Essentially as in S. torvus, but there are only about 15 small setae bordering the antennal fossa posteriorly.

THORAX. Pronotum narrow, dorsally much shorter than the length of the dorsal spines of the pronotal ctenidium, the latter consisting of 20 slightly curved spines. Though narrow, the pronotum still bears two rows of setae (contrast pronotum of S. alienus, Text-fig. 1). In other respects the thorax resembles that of S. torvus.

Lecs. The tibiae differ from those of all other species of the torvus-group by having seven groups of setae in notches of the posterior margin on all the legs, whereas in the other species the fore tibia has seven but the mid and hind tibiae have eight. The numbers of setae in these notches are: mid tibia 2, 2, 2, 2, 3, 2 and 3, hind tibia 2, 2, I, 2, 2 (or 3), 3, and 3 (the hind tibia closely resembles that of S. alienus (Text-fig. 3)).

ABDOMEN. Unmodified abdominal segments as in S. torvus, but the upper ante- sensilial seta is a little less than half the length of the lower.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. FEMALE (Text-figs. 35, 47, 48, 59). Sternum VII (Text-figs. 47, 48) of the type usual in the ¢orvus-group, the lobe above the sinus only weakly developed. Variation in outline of the posterior margin of sternum VII as shown in Text-figs. 47, 48. Tergum VIII with four to seven small setae in front of the vertical part of the spiracular fossa ; noselike projection of the posterior margin of tergum VIII nearly absent, indicated by a weak convexity of the margin (Text-fig. 59). Sternum VIII as in Text-fig. 35. Anal stylet about four times as long as its maximum width, with one or two minute preapical setae and a long apical seta. Bulga of spermatheca (Text-fig. 35) with a convex dorsal margin but without a hump. The sclerite on each side of the bursa copulatrix is small and elongate, kidney-shaped, and as long as the bursa ; dilated part of ductus spermathecae rather long and straight (Text-fig. 35).

LENGTH. @ 23-3 mm.

66 THE AFRICAN SPECIES OF STIVALIUS

Fics. 38, 39. Stivalius torvus (Rothschild). 38. Sternum VII of female (Tengeru, Tanganyika). 39. Outlines of sternum VII of females (a—Kenya, b—Keruguya, Kenya, c—Kenya, d—Lubero, Kivu district, Belgian Congo, e—Kanungu, Kigezi, Uganda, f—Machakos, Kenya, g—Damba Island, Lake Victoria). Figs. 40, 41. Sternum VII of left hand side and outline (stippled) of right hand side of same sternum. 40. S. curtiductus sp. n. (holotype). 41. S. vichardi Jordan (allotype).

mails geen

eS ee

THE AFRICAN SPECIES OF STIVALIUS 67

REMARKS. Jordan described S. nigeriensis as a subspecies of S. afer but remarked that it may represent a separate species since it agrees neither with torvus nor with afer ; he placed it provisionally as a subspecies of afer because the sclerite on each side of the bursa copulatrix is more like that of afer than that of torvus. But the considerable differences in the pronotum, pronotal ctenidium and tibiae between S. afer and S. nigertensis show that the latter is better placed as a separate species.

Stivalius fallaciosus sp. n. (Text-figs. I1, 22)

“ Pygiopsylla torvus’’ Jordan & Rothschild, rg11, Novit. zool. 18:71; Jordan & Rothschild, 1922, Ectoparasites, 1: 252.

TYPE MATERIAL. Male holotype from Bamenda, Cameroons, S.E. Nigeria, vi. 1909, leg. Adametz, without host-record. The holotype, formerly in the collection of the Zoologisches Museum der Humboldt-Universitét in Berlin, has been generously presented by Prof. Dr. F. Peus to the British Museum collection of fleas at Tring.

Dracnosis. The male of this new species can be distinguished from that of related species by the fact that the dense group of thin setae along the ventral margin of the movable process of the clasper extends along the rounded angle of the margin, by the shape of the expanded apical part of the distal arm of sternum IX and by the structure of the aedeagus. Female unknown.

DESCRIPTION. HEAD. Submarginal frontal row consisting of seven setae each side ; in other respects the head resembles that of S. torvus.

THORAX. Pronotal ctenidium consisting of 23 spines. The two dorsal pseudosetae on each side under the collar of the mesonotum are clearly separated by a gap (as in S. timanus). :

Lecs. Mid tibia with seven notches in the posterior margin, bearing 2, 2, I, 2, 3, 2 (or 3) and 3 setae respectively. Otherwise the thorax and legs are essentially as in S. torvus.

ABDOMEN. Basal abdominal sternum with one or two small setae anterodorsally ; sterna IJI-VII with a main row of four setae each side. In other respects the unmodi- fied abdominal segments agree with those of S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 11, 22). Tergum VIII with eight setae each side in front of the vertical portion of the spiracular fossa. Sternum VIII with about 40 setae each side and the usual apical ventro-marginal group of several short setae. Apodeme of tergum IX as in S. forvus ; manubrium broad ; fixed process of clasper with one long and one short acetabular seta ; movable process short for this group, the four or five long seta at the ventral margin of the bent-down apical portion are placed rather close together (Text-fig. 11). The apex of the distal arm of sternum IX with a characteristic widening, the dorsal projection rather long and its apex splayed out; the ventral spiniform setae form a dense group (Text-fig. 11). Aedeagus as in Text-fig. 22; the tendons of the phallosome make about two convolutions.

LENGTH. ¢ 34 mm.

68

THE AFRICAN SPECIES OF STIVALIUS

Fics. 42-46. Sternum VII of left hand side and outline (stippled) of right hand side of same sternum. 42. Stivalius vancanneyti Berteaux (holotype). 43. S. afer (Rothschild) (holotype). 44. S. parilis sp. n. (paratype). 45, 46. S. timanus Jordan

(45—allotype, 46—paratype).

eT se ee

THE AFRICAN SPECIES OF STIVALIUS 69

Stivalius sellatus Jordan & Rothschild, 1923 (Text-figs. 14, 20, 36, 49, 60)

Stivalius sellatus Jordan & Rothschild, 1923, Ectoparasites, 1: 304, fig. 301; Hopkins, 1947,

Uganda J. (Suppl.) 11: 155.

MATERIAL EXAMINED. Female holotype from Bubungi, North Bugishu, Eastern Uganda, 28.i1.1922, froma rat, W. N. van Someren ; one male from Torit, Equatoria, Sudan, 3,800 ft., from a mouse, 7.iv.1950, H. Hoogstraal, presented by Lt.-Col. R. Traub.

DeEscripTIon. Heap. Essentially as that of S. torvus, but the frons is somewhat more strongly rounded, and the small setae bordering the antennal fossa are less numerous.

THoRAX. Like that of S. torvus ; the pronotal ctenidium with 24 spines in both the available specimens.

LeGs. The outer side of the fore tibia with only three to five lateral setae in the male, six or seven in the female. In both the specimens examined one hind tibia has the usual eight groups of setae in notches along the posterior (dorsal) margin, but on the other side:there are nine, an extra single stout seta being present between the second and third notches (counting from the base).

ABDOMEN. Marginal spinelets of terga II-IV developed into moderately large spines which form dorsal ctenidia consisting of six spines each side on all three terga in the male, while in the female there are six and seven on the two sides of terga II and III and six on each side of tergum IV; no spines or spinelets on other terga. Dorsally the ctenidia of each side do not quite meet, but are separated by a short dorsal interspace. Basal abdominal sternum with only one lateral seta in the only male examined, but about 25 in the female holotype. In other respects the chaetotaxy of the unmodified abdominal segments resembles that of S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 14, 20). Tergum VIII anterior to the vertical part of the spiracular fossa with three short setae on one side and four on the other. Sternum VIII with about 25 setae each side and a ventro-marginal group of two stout and three small setae. Apodeme of tergum IX (Text-fig. 14) narrow and of the same construction as in other species of the torvus- group. Manubrium much narrower than the width of the basal half of the aedeagal apodeme, rather short, its ventral margin wholly convex. Fixed process of clasper

_ with one very long and slender and one very short acetabular seta. The down-bent

apical portion of the movable process bears ventrally two or three long setae and forms about a right angle with the long and straight part of the process, the usual dense patch of thin setae along the ventral margin before the bend is very narrow (Text-fig. 14). Dorsal margin of the widened apical part of the distal arm of sternum IX (Text-fig. 14) smoothly convex, without irregularities in its outline; a group of short setae, several of which are stout but none definitely spiniform, is present at the ventro-apical angle of the apex ; the dorso-apical angle is sharp. Aedeagus as in Text-fig. 20; the tendons of the phallosome form about two convolutions ; aedeagal apodeme with the deep concavity in the dorso-apical half which is charac- teristic of the torvus-group.

70 THE AFRICAN SPECIES OF STIVALIUS

eR athe tg — a b eee $ 50 5| 52

Fics. 47-49. Sternum VII of left hand side and outline (stippled) of right hand side of same sternum. 47, 48. Stivalius nigeriensis Jordan (47—holotype, 48—Adio- podoumé, Ivory Coast). 49. S. sellatus Jordan & Rothschild (holotype). Figs. 50-52. S. pirloti sp.n. 50. Sternum VII of female (allotype). 51. Outlines of sternum VII of two paratypes (nr. Elisabethville) 52. Outline of tergum VIII of female (allotype).

THE AFRICAN SPECIES OF STIVALIUS 71

FEMALE (Text-figs. 36, 49, 60). Sternum VII (Text-fig. 49) with a deep sinus of the usual shape dividing the main row of four setae into two groups of two ; chaeto- taxy of this sternum as shown in the figure. Tergum VIII with eight or nine setae in front of the vertical part of the spiracular fossa ; noselike projection of the posterior margin of this tergum rather weakly developed (Text-fig. 60). Sternum VIII as in Text-fig. 36. Anal stylet just over thrice as long as its maximum width, with a long apical seta and two minute preapical setae. Bulga of spermatheca (Text-fig. 36) with a fairly thick wall and a small dorsal hump. The sclerite each side of the extremely small bursa copulatrix is poorly sclerotised and small. The dilated part of the ductus spermathecae is not very long, in contrast with the remainder of this duct (Text-fig. 36).

LENGTH. ¢ 3 mm., 9 3 mm.

Stivalius pirloti sp. n.

(Text-figs. 15, 23, 27, 37, 50-52)

TYPE MATERIAL. From the neighbourhood of Elisabethville, Katanga, Belgian Congo, collected by Dr. P. L. Pirlot: male holotype, nine male and three female paratypes from Rattus (Praomys) morio jacksoni, 1957 ; female allotype from Kikusue near Elisabethville, from Rattus (Praomys) morio jacksoni, vi.1956, collected by M. Lips; one female paratype from Kanienge near Elisabethville, from Rattus (Praomys) morio jackson, vi.1956, M. Lips. Holotype, allotype and several para- types in the Musée Royal du Congo Belge, Tervuren.

D1aGnosis. Separable from all other African members of the genus, except S. sellatus, by the possession of dorsal ctenidia on some of the abdominal terga; from S. sellatus in the male by the possession of (a) a large area of thin setae on the inner side of the movable process, (b) only one acetabular seta (there are two such setae in all other African species), (c) a number of shortish spiniform setae on the ventro-apical margin of sternum IX, (d) sickle-shaped dorso-apical aedeagal sclerite; in the female by the presence of large, curved and rather narrow sclerites associated with the bursa copulatrix.

DESCRIPTION. HEAD. As in S. torvus.

THORAX. Pronotal ctenidium with 24-27 (normally 24) slender spines in the male, 25-26 in the female. Metanotum with a main row of seven setae on each side. In other respects the thorax, as well as the legs, resemble those of S. torvus.

ABDOMEN. All the marginal spinelets developed into spines, which form conspicu- ous dorsal ctenidia on terga II-IV (Text-fig. 27) and sometimes also on V; on tergum I there are usually no spines (occasionally one, or one on each side), on tergum II four or five on each side, on tergum III four (sometimes five), on tergum IV three or four in the male and two to four (generally three) in the female, and on tergum V one (sometimes two) in the male and one or none in the female. Basal abdominal sternum with a lateral patch of one to eight (average four) small setae in the male and 12-20 (average 17) in the female. Otherwise the unmodified abdominal segments are like those in S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 15, 23). Tergum VIII with three to six smallish setae each side anterior to the vertical portion

72

THE

AFRICAN SPECIES OF STIVALIUS

THE AFRICAN SPECIES OF STIVALIUS 73

of the spiracular fossa. Sternum VIII with 25-30 setae each side, the three most apical of the ventral setae are set closely together. Apodeme of tergum IX and manubrium (Text-fig. 15) more or less as in S. torvus. Fixed process of clasper with one long acetabular seta only. Movable process of clasper like that of S. torvus, but relatively longer, the down-bent apical portion forming an angle far greater than a right angle with the basal portion (contrast S. sellatus, Text-fig. 14). Sternum IX (Text-fig. 15) not unlike that of S. torvus, but the apical part of the distal arm is little widened, proportionally longer and the spiniform setae along the ventro- apical margin are much blunter and more widely spaced. Aedeagus as in Text-fig. 23 ; dorso-apical aedeagal sclerite sickle-shaped ; tendons of phallosome making about two convolutions ; dorsal margin of aedeagal apodeme strongly concave preapically.

FEMALE (Text-figs. 37, 50-52). Sternum VII (Text-figs. 50, 51) of the shape usual for the forvus-group and with a chaetotaxy resembling that of the other species in this group. Tergum VIII with a row of five setae in front of the vertical portion of the spiracular fossa ; chaetotaxy of the ventral part of this tergum as in other species of the group; posterior margin of tergum VIII as in Text-fig. 52. Anal segment more or less as in S. torvus. Bulga of spermatheca thick-walled, with only a small dorsal hump, hilla with a papilla (Text-fig. 37). The pair of dark sclerites associated with the bursa copulatrix long and slender and markedly curved; the dilated part of the ductus spermathecae is long, not very wide, and internally provided with the usual membranous rings (Text-fig. 37).

LENGTH. ¢ 3-34 mm., 9 34-44 mm.

REMARKS. I have pleasure in naming this new flea, a parasite of the arboreal Rattus (Praomys) mono, after Dr. P. L. Pirlot, who made extensive collections of fleas while studying and collecting mammals in the Belgian Congo.

Stivalius cacuminis sp. n. (Text-figs. 12, 24)

TYPE MATERIAL. Male holotype collected on the top of Mt. Soque (about 40-50 miles east of Lobito), Angola, from a mouse,! on 27.viii.1954, by G. Heinrich. The holotype is in the Chicago Natural History Museum.

1 Dr. R. L. Wenzel, of the Chicago Natural History Museum, kindly informed me that the following rodents were collected by Mr. Heinrich on Mt. Soque on August 27, 1954: Graphiurus sp., Otomys ivvoratus, Dendromus sp., Lophuromys sikapust, Oenomys sp., Pelomys fallax and Rattus (Praomys) namaquensis. Since Stivalius cacuminis is closely related to S. pirloti (a parasite of Rattus (Praomys) morio) it is possible that Rattus (Praomys) namaquensis is the true host of S. cacuminis.

Fics. 53, 54. Stivalius torvus (Rothschild). 53. Tergum VIII of female (Keruguya, Kenya). 54. Outlines of tergum VIII of females (a—-c: Keruguya, Kenya, d-e: Machakos, Kenya). Figs. 55-62. Outlines of tergum VIII of females of : 55. S. vancan- neytt Berteaux (holotype). 56. S. afer (Rothschild) (holotype). 57. S. parilis sp. n. (a—paratype, b—allotype). 58. S. timanus Jordan (a—allotype, b—paratype). 59. S. nigeriensis Jordan (a—holotype, b—Adiopodoumé, Ivory Coast). 60. S. sellatus Jordan & Rothschild (holotype). 61. S. vichardi Jordan (allotype). 62. S. curtiductus sp. n. (holotype). Figs. 63-65. Pronotum of females of: 63. S. aestivalis Jameson & Sakaguti (Hanase, Kyoto Prefecture, Honshu, Japan). 64. S. ferinus (Rothschild) (Namunukula, Ceylon). 65. S. imsolli Traub (Brinchong Hill, Cameron Highlands, Malaya).

74 THE AFRICAN SPECIES OF STIVALIUS

D1aGnosis. On the structure of the aedeagus this new species is nearest related to S. pirloti, from which it can be distinguished at once by the absence of ctenidia on the anterior abdominal terga.

DESCRIPTION. Head, thorax, legs and unmodified abdominal segments as in S. torvus.

MODIFIED ABDOMINAL SEGMENTS AND GENITALIA. MALE (Text-figs. 12, 24). Segment VIII as in S. torvus, except for the apical subventral group of setae on sternum VIII which consists of only three setae with four very small setae opposite these, on the inner side of the sternum. Apodeme of tergum IX, manubrium and fixed process of clasper (Text-fig. 12) similar to those in S. torvus ; movable process much longer than that of S. ¢orvus, with a relatively narrow patch of thin setae subventrally on the inner side of the long straight portion; the ventral margin of the apical bent-down portion with four long setae (Text-fig. 12). Sternum IX (Text-fig. 12) resembling that of S. pirloti, differing in some small details. Aedeagus as in Text-fig. 24; the tendons of the phallosome make nearly two convolutions ; dorso- apical aedeagal sclerite sickle-shaped ; the dorsal margin of the aedeagal apodeme is strongly concave before the tip.

LENGTH. ¢ 3} mm.

KEY TO THE AFRICAN SPECIES OF Stivalius

1. Genal margin below eye divided into two partly overlapping lobes (Text-fig. 1) ; g—tendons of phallosome short, hardly reaching beyond apex of aedeagal apodeme (Text-fig. 16) ; dorsal margin of aedeagal apodeme almost straight (Text- fig. 16) ; 9—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 (Text-fig. 25) (ferinus-group) : alienus, p. 42

— Genal margin below eye entire (Text-fig. 2) ; g—tendons of phallosome making at least half a convolution (Text-figs. 17, 18); dorsal margin of aedeagal apodeme strongly concave preapically (Text-figs. 17, 18); 9Q—bursa copulatrix in most species with a dark sclerotic structure on each side (Text-fig. 26, 28-37) ; dilated part of ductus spermathecae with a large number of membranous internal divisional rings (Text-figs. 26, 28-37) (torvus-group) ‘ ; : : : ‘ 2

2. Terga II-IV with developed dorsal ctenidia (Text-fig. 27)

— These terga with only one subdorsal marginal spinelet

3. d—Apical portion of movable process bent downwards at about a right angle ; along the ventral margin before the bend a relatively sparse group of thin setae ; fixed process of clasper with two acetabular setae (Text-fig. 14); 9—Bursa copulatrix very small, the sclerites not much darkened (Text-fig. 36)

> W

sellatus, p. 69 — g—Down-bent apical portion of movable process forming an angle considerably greater than a right angle with the basal portion, the latter part bearing a large area of thin setae on inner side; fixed process with only one acetabular seta (Text-fig. 15); 9Q—Bursa copulatrix itis = the curved sclerites very long

and narrow (Text-fig. 37) ; y pirloti, p. 71 4. Dorsal (posterior) margin of hind tibia with. seven | groupe of setae i in notches; pro- notum dorsally much shorter than dorsal pronotal spines; pronotal ctenidium

-

Io.

THE AFRICAN SPECIES OF STIVALIUS

with 20 spines (male unknown) . nigeriensis, Pp.

Dorsal margin of hind tibia with eight groups of setae : “normally at least 22 pronotal spines which are usually about as long as (or a little shorter than) pronotum

Males (this sex not known of afer, curtiductus and vancanneytt)

Females (this sex not known of fallaciosus and cacuminis) : . . ‘

Tendons of phallosome making at most half a convolution; apex of aedeagal apodeme without a tendril (Text-fig. 17) é : ‘ : , ‘

These tendons making at least a whole convolution; apex of aedeagal apodeme

drawn out into a fairly long and not strongly sclerotized tendril (Text-fig. 18)

Dorso-apical part of distal arm of sternum IX drawn out into a long and sharply

pointed projection (Text-fig. 1 or ; dorso-apical aedeagal sclerite very large (Text-

fig. 21) ; ‘ vichardi, p.

Dorso-apical part of distal arm of sternum IX not ‘forming a Jong and pointed projection (Text-fig. 8) ; dorso-apical aedeagal sclerite much smaller (Text-fig. 17)

tovuus, Pp.

Upper extension of dorso-apical aedeagal sclerite sickle-shaped (Text-fig. 24) ; ventral margin of basal part of movable process with a narrow group of thin setae

(Text-fig. 12) : . . cacuminis, p.

Upper extension of dorso- apical aedeagal Sclexite not sickle- -shaped ; area of thin setae along ventral margin of movable process much wider :

Ventro-marginal group of setae on movable process extending along curvature of margin to the first large ventral seta of down-bent portion ; apex of widened part

of distal arm of sternum IX splayed out (Text-fig. 11) . ; fallaciosus, p.

The group of thin setae on movable process not extending along curvature of margin ; apex of widened part of distal arm of sternum IX not splayed out

Proximal arm of sternum. IX subequal in length to distal arm; dorso- sical angle of club-shaped expansion of latter arm broadly rounded (Text-fig. 10) ; ventro- apical angle of lower lateral aedeagal lobe of about 90°; dorsal wall of aedeagal

inner tube relatively thin (Text-fig. 18) . , . : timanus, Pp.

Proximal arm of sternum IX distinctly longer than distal arm ; apex of latter arm not much widened (Text-fig. 9) ; apical and ventral margins of lower lateral aedeagal lobe forming a very acute angle ; dorsal wall of inner tube very thick (Text-fig. 19)

parilis, p.

Dilated part of ductus spermathecae short and thick, not or hardly longer than spermatheca (Text-fig. 26, 28-30) . ‘

Dilated part of ductus spermathecae relatively fong ana narrow ‘(Text- figs. 31-37)

Only a vestige left of the sclerites associated with the bursa copulatrix ; undilated

part of ductus spermathecae very short (Text-fig. 29) ‘ ‘ curtiductus, p.

Sclerites associated with bursa copulatrix well developed ; undilated part of ductus spermathecae much longer (Text-figs. 26, 28, 30) . : Sclerites of bursa copulatrix fairly small and roughly teenie nian (Text- figs, 26, 28)

torvus, p.

These sclerites very large and shaped like the sole of a shoe (Text-fig. 30)

vichardt, p.

Sclerites about as long as bursa copulatrix (Text-figs. 33, 34) Sclerites about twice as long as bursa copulatrix (Text-figs. 31, 32) Sclerites and bursa copulatrix relatively large, nearly as long as bulga of sperma-

theca (Text-fig. 33) ‘ : parilis, p.

Sclerite and bursa copulatrix much smaller in relation to ‘length of bulga (Text-

fig. 34) ‘ P Z timanus, p. . Dilated part of ductus spermathecae about seven . times as long as wide (Text-fig. 31) vancanneyltt, p.

Dilated part of ductus spermathecae about twelve times as long as wide (Text-

fig. 32) F : ‘ . ; ‘ 7 : : : : . afer, p.

76 THE AFRICAN SPECIES OF STIVALIUS

ACKNOWLEDGMENTS

I am deeply indebted to the following colleagues for the loan or gift of specimens of African species of Stivalius : Dr. V. Aellen (donation of a female of S. nigeriensis), Dr. P. L. G. Benoit (loan and donation of much material from the Musée Royal du Congo Belge), Dr. R. Devignat (donation of specimens of S. alienus), Prof. Dr. F. Peus (donation of holotype of S. fallactosus) and Lt.Col. R. Traub (donation of a male S. sellatus and permission to describe this sex, and for inviting me to describe a male from Angola (S. cacwmimis) which he had found among material sent to him by the Chicago Natural History Museum).

—— Ta

;

THE PSEUDOCOCCIDAE (HOM. : COCCOIDEA) DESCRIBED BY C. K. BRAIN FROM SOUTH AFRICA

G. DE LOTTO

BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 7 No. 3 LONDON: 1958

\ \ 7 py,

THE PSEUDOCOCCIDAE (HOM. : COCCOIDEA) DESCRIBED BY C. K. BRAIN FROM SOUTH AFRICA

BY

G, DE. LOTTO fh “Ky

LK

Department of Agriculture, Kenya }

Pp. 77-120 ; 19 Text-figures

BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Vol. 7 No. 3 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 Senes. ,

Parts appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.

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THE PSEUDOCOCCIDAE (HOM.:COCCOIDEA) DESCRIBED BY C.K. BRAIN FROM SOUTH AFRICA

By G. DE LOTTO Department of Agriculture, Kenya

Our knowledge in the Pseudococcid fauna of South Africa is almost entirely based on the works published by C. K. Brain, who treated altogether forty-two species and varieties, thirty-one of which were described by him as new. Omitting any criticism of the methods he used in all his descriptions, which are more or less of the standard then attained by the majority of entomologists, it is evident that none of his species can be recognized without the examination of the types. Subsequent workers dealt with a few of his species, but those remaining were still completely unrecognizable.

In the present paper all Brain’s species are reviewed or redescribed from the types, paratypes or specimens which, although marked by the serial number only, undoub- tedly belonged to the original material studied by him. The main work was carried out at the Division of Entomology of the Department of Agriculture in Pretoria, where the bulk of Brain’s collection is deposited. Material of four species apparently missing there, was made available from the U.S. National Collection of Coccidae in Washington, D.C.

With a few exceptions the original material examined was in poor and at times in very poor condition. Specimens were often broken and badly distorted and not cleaned or not properly cleaned of the body content. Furthermore some specimens were evidently mounted from dry material or overboiled when treated with KOH, so that when remounting we found very serious difficulties on making really adequate microscope preparations. The utmost care was made in studying every structure of taxonomic value in order to present adequate diagnoses and figures so as to enable the recognition of the various species and serve for their generic allocation in the final revision.

Out of twenty-nine species and two varieties described as new by Brain, three species and one variety have been already synonymized by various authors. Four more species and one variety are synonymized in the course of the following pages. The complete list of them is as follows:

Antonina transvaalensis = natalensis Brain. Pseudococcus capensis = maritimus (Ehrhorn).

ENTOM, 7, 3. 4

80 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Pseudococcus citri phenacocciformis = citri (Risso). Pseudococcus elisabethae = quaesitus Brain. Pseudococcus natalensis = graminis (Maskell). Pseudococcus segnis tylococciformis = segnis Brain. Pseudococcus solitarius = vastator (Maskell). Pseudococcus trichtliae = guaesitus Brain. Rhizoecus africanus = falcifer Kuenckel.

With the publication of this paper nearly two thirds of the described mealy-bugs of the African continent south of Sahara have been dealt with.

Antonina natalensis Brain (Text-fig. 1)

Antonina natalensis Brain, 1915, Tvans. voy Soc. S. Africa. 5: 86.

Antonina transvaalensis Brain, 1915, Trans. voy. Soc. S. Afr. 5: 87 (SYN. NOV.). Antonina indica panica Hall, 1925, Bull. Minist. Agric. Egypt. 64:6 (SYN. NOV.). Antonina indica panica Hall, 1937, Trans. R. ent. Soc. Lond. 86 : 125.

One slide with a single very adult female was available from the U.S. National Collection of Coccidae, Washington, D.C. It was labelled: ‘‘ Antonina natalensis Br.; grass; Pietermaritzburg, Nov. 15, 1914; Chas. K. Brain; 33, C.K.B.” The specimen was in fair condition, although the body skin was slightly chitinized and there were difficulties in examining some glandular structures.

“Adult female enclosed in a dense, though, felted sac at the base of grasses. Ovisac: at first sight the ovisacs appear spherical, but on closer inspection they are found to be somewhat produced at the anterior end, so as to be really pear-shaped, 4'5 mm. long and 3°8 to 4 mm. broad. In colour the sac is yellowish-white to buff. Adult female : purplish-black in colour, about 4 mm. long, distinctly semipyriform, being flattened ventrally and broadly rounded behind while the anterior end is narrowed. There is not secretionary covering, but the walls of the sac are closely adherent to the body of insect.” (Brain, /.c.). Body of mounted specimen broadly oval with posterior end chitinized. Anal and marginal cerarii entirely absent. Anal ring with six robust setae, set at inner extremity of a deep invagination of dermis. Multilocular disc pores numerous on median and submedian areas of venter and a few crowded near each stigmatic opening. Tubular ducts with inner end dome- shaped ; they are fairly abundant and uniformly distributed on either side of body. Sieve-like pores caudad of posterior stigmatic opening lacking, apparently being replaced by flat circular glands with a granulated surface, variable in size and always smaller than multilocular disc pores. Trilocular pores evenly scattered on dorsum and venter. Very small invaginations of ventral dermis mark the position where the legs would be situated. Body setae very short, except a few on posterior end of body. Atrium of each stigmatic opening with a band of trilocular pores. Antennae formed by two or three small joints, with a tuft of short setae on apex.

81

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THE PSEUDOCOCCIDAE (HOM. COCCOIDEA)

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Antonina natalensis Brain

Fic. I.

82 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Antonina transvaalensis Brain (= Antonina natalensis Brain) Antonina transvaalensis Brain, 1915, Trans. voy Soc. S. Afr. 5: 87.

One slide on loan from the U.S. National Collection of Coccidae, Washington, D.C., was seen. It contained three adult females, the last abdominal segments of another adult female and one larva. It bore the following label: ‘‘ Antonina trans- vaalensis Brain ; on roots of grass; Daspoort, Pretoria ; C. K. Brain coll., Oct. 11, 1939; Brain II 70; 70, C.K.B.”” The mistake in the year of collection is evidently due to a pen slip. Very likely this slide, as many others in Brain’s collection in Pretoria, was originally marked with the serial number only, and the label was written much later.

Antonina transvaalensis is a synonym of natalensis as no structural differences were found between the two species.

Grewiacoccus gregalis Brain (Text-fig. 2) Grewiacoccus gregalis Brain, 1918, Bull. ent. Res. 9: 108.

The material examined was represented by five slides each containing a single adult female in fairly good condition, except that the prosoma was partly distorted. All slides were labelled: ‘‘ Grewiacoccus gregalis; Transvaal, Pienaar’s River, 23 January 1917; C.K.B. and C. Fuller; on leaves of kruisbesje!; C.K.B. 305.”

“ Adult female causing a gall on the leaf of the host-plant, in which one, two or three insects may be present. The galls may be solitary, or, as is often the case, five to twelve may be found clustered around the edge of a single leaf. The mature gall is about 6 mm. in length, broadly inflated, with the sides tapering slightly to the neck, of a scrotal shape or one very much likea pitcher. The galls stand nearly erect from the distorted edge, and are of the same green colour and texture as the leaf. There is no hole at the distal end, but below, on the under-side of the leaf, is a circular opening through which project two dense white filaments accompanied by a small amount of fine white cottony matter. When obtained in January numerous young were found clustered around the necks of some of the galls. All the available space in the gall is occupied by the female—or females if these are two or three—and on the inner walls there is a slight layer of white wax, which is more plentiful near the base, or where the posterior part of the insect is. ... Adult female when alive, yellowish or slight pink, about 3:2 mm. long.” (Brain, /.c.). Mounted specimens broadly oboval, tapering posteriorly with dorsal dermis of postsoma very slightly chitinized at matur- ity. Anal and marginal cerarii absent. Anal lobes each provided on ventral side with a short stout seta. Multilocular disc pores absent. Tubular ducts very short with chitinized walls ; inner end membranous, dome-shaped. They occur on both sides of body and are more numerous on last abdominal segments. Trilocular pores distributed on either side of body; a few crowded near each stigmatic opening. Circular disc pores lacking. Posterior dorsal ostioles rather prominent, with lips

1 Afrikaans name of Grewta occidentalis L.

—

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 83

Fic. 2. Gvrewiacoccus gregalis Brain

ENTOM. 7, 3.

84 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

membranous ; anterior ones not recognizable. Circulus absent. Legs small; hind coxae broadly enlarged forming an irregularly shaped plate marked by numerous small and slightly chitinized areas. Numerous short stoutly lanceolate setae arranged dorsally all around anal ring ; on preceding segments they tend to be fewer and smaller ; on prosoma they are still more widely scattered. Ventral setae small and slender, not numerous. Antennae with six joints. Anal ring entire, with six small setae.

Natalensis fulleri Brain (Text-fig. 3) Natalensis fulleri Brain, 1915, Trans. voy. Soc. S. Afr. 5: 91.

Four slides were available, two with larvae and two containing a single specimen in very poor condition ; both being very old adults, partly broken, distorted and attacked by fungi. All slides were marked: “ 43, C.K.B.” which is the serial number of Brain’s collection. The material was originally “ collected on the roots of grass in the nests of a small red ant in the Botanic Gardens, Pietermaritzburg, Natal by Mr. Claude Fuller, 1910.” (Brain, /.c.).

‘“‘ Adult female entirely enclosed in a double-walled cyst. Cyst: outer cyst may attain 9g mm. long by 5 mm. broad, dark-coloured owing to adherent soil ; brittle when dry ... Inner cyst pale buff-coloured. This has the appearance of being more loosely constructed. The outer shape and size are those of the interior of the outer cyst, from which it separates readily ... Adult female (from dry material), after boiling in KOH dark brown in colour, broadly elliptical, 4 mm. long and 4-5 mm. broad, convex above and flat beneath.” (Brain, /.c.). Body of mounted specimens very broadly elliptical with dermis slightly chitinized at full maturity. Cerarii recognizable only on anal lobes, each being built up by two short and stout setae surrounded by a few trilocular pores. Area about setae not chitinized. Ventral side of each anal lobe without apical seta. Multilocular disc pores, tubular ducts, tritubular ducts and circular disc pores absent. Trilocular pores few and widely scattered on both side of body. Anterior and posterior dorsal ostioles, circulus and ventral cephalic plate absent. Anal ring entire, with six setae. Antennae with six joints, devoid of any falcate sensory seta. Legs very small with tarsal digitules finely pointed ; ungual ones slightly knobbed at apex. Body setae few, slender and widely distributed on either side.

Pseudococcus bantu Brain (Text-fig. 4) Pseudococcus bantu Brain, 1915, Trans. voy. Soc. S. Afr. 5: 137.

Four slides each with a single adult female in fairly good condition were seen. They bore the following label: “‘ Phenacoccus bantu Brain; grass; P.m. burg, N.?: Nov. 1914; paratype; Char. K. Brain; 34, C.K.B.”’ Another slide contained ova and larvae only.

1 Abbreviation for Pietermaritzburg, Natal.

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 85

Fic. 3. Natalensis fulleri Brain

86 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Pseudococcus bantu Brain

Fic. 4.

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 87

“ Ovisac : large clusters of irregular white ovisacs were found at the bases of the leaves and on the crowns of grass ... Adult female: small, orange in colour, with a very slight covering of powdery secretion, but without lateral or caudal filaments ... Mounted specimens average 2:3 mm. long by 1-8 mm. broad.” (Brain, /.c.).

The following redescription is made from specimens collected by E. C. G. Bedford on roots of grass at Klaserie, Transvaal, South Africa, on 20.iv.1955, which were compared with Brain’s paratypes.

Body elliptical, membranous. Margin of body with a complete series of eighteen pairs of cerarii each with two small conical spines, except one or two cerarii of thorax which at times are reduced to one spine only and the interantennal pair which is often provided with three spines. Each cerarius anterior to anal lobe ones is beset by a group of three to seven trilocular pores; auxiliary setae missing. Anal lobe cerarii each with a loose group of trilocular pores and one to three small auxiliary setae ; area about spines not chitinized. Ventral side of each anal lobe with a robust apical seta, distinctly longer than those of anal ring ; subapical seta about half the length of apical one ; chitinized bar absent. Multilocular disc pores of usual type rather abundant on ventral side of last five abdominal segments, mostly arranged in transverse rows along distal margin. Other multilocular disc pores slightly larger than normal ones, set in groups of two or three—seldom four. Each cluster of pores has associated one to four tubular ducts with oral collar, one which is always smaller and set on middle. On dorsum these clusters of pores are numerous and on thorax and abdomen they are arranged in transverse segmental rows; on ventral side they mostly occur on thorax and first two abdominal segments and on marginal area of remaining abdominal segments. Quinquelocular pores present only on median and submedian areas of venter, not abundant. Tubular ducts all with one rim, and except for those associated with clusters of large multilocular disc pores, crowded along ventral marginal area of last four abdominal segments anterior to anal lobes and intermingled with normal multilocular disc pores. Trilocular pores not very numerous andevenly distributed. Circular disc pores somewhat smaller than trilocular pores and scattered on either side of body. Dorsal setae very small, spiniform ; ventral ones much longer and slender. Anterior and posterior dorsal ostioles not prominent, lips membranous with a few trilocular pores and small setae. Circulus transversely elongate, membranous. Legs well developed, rather stout, with a few translucent pores on hind tibia ; claw with a small denticle ; ungual digitules slightly knobbed, tarsal ones finely pointed. Antennae with nine joints.

Pseudococcus bechuanae Brain (Text-fig. 5) Pseudococcus bechuanae, 1915, Trans. roy. Soc. S. Afr. 5: 108. Two sides were studied, each containing a single specimen, partly distorted, labelled: “‘ Pseudococcus bechuanae Br.; on Geranium; Vryburg, Bechuanaland, aeen.tor4; ©. Kh. Brain; B. 53; C.K.B."*

1 In the original paper the date is 3.xii.1914. Evidently there has been a mistake on labelling the slides, as all other collecting data are identical.

88 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

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Fig. 5. Pseudococcus bechuanae Brain

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 89

“The specimens had been badly shaken in the post, but it was clear that large patches of the stem had been completely covered with ovisacs. The white secretion appeared powdery rather than cottony. The females embedded in this were pinkish in colour when young, but purplish in older specimens. There were no lateral filaments, and only two short, blunt, caudal ones. The female when adult reaches approximately 3 mm. in length.” (Brain, /.c.). Mounted specimens elongate elliptical, membranous. Margin of body provided with anal and preanal cerarii only. Each of them built up with two conical spines surrounded by a small group of trilocular pores and a few slender auxiliary setae ; area about spines not chitinized. Ventral side of each anal lobe without chitinized bar ; apical seta robust, somewhat shorter than those of anal ring; subapical seta much shorter. 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) 6; (vi) 18; (vii) 21; (vili) 29; (ix + x) 28. A few multilocular disc pores occur on dorsal side of three abdominal segments anterior to anal one. Tubular ducts with oral rim fairly numerous on dorsum and a few scattered along marginal and submarginal areas of abdomen. Tubular ducts with oral collar arranged in four or five groups on ventral marginal area of last abdominal segments anterior to anal lobes ; others distributed in irregular rows in association with dorsal and ventral multilocular disc pores. Trilocular pores rather numerous and uniformly distributed on both sides of body. Circular disc pores apparently absent. Anterior and posterior dorsal ostioles inconspicuous, lips membranous with a small grouping of trilocular pores and a few small setae. Circulus absent. Legs well developed, without translucent pores. Ventral and dorsal body setae very small and slender. Antennae 8-jointed with a pseudoarticulation on apical joint.?

Pseudococcus burnerae Brain (Text-fig. 6)

Pseudococcus burnerae Brain, 1915, Trans. voy. Soc. S. Afy. 5: 111. Pseudococcus simulator James, 1933, Bull. ent. Res, 24: 434 (SYN. NOV.).

A fairly long series of specimens from the typical material originally studied by Brain was available. The slides were as follows: four with altogether twelve speci- mens labelled: “‘ Pseudococcus burnerae Brain ; on Sida longipes; Pretoria: Oct. II, 1914; Char. K. Brain; B. 45, C.K.B.” Two more slides of this series contained only eggs, larvae and adult males. Two slides each with a single specimen marked only with the serial number “ B. 50, C.K.B.” Three slides with altogether twelve specimens also marked with the serial number only ‘‘ 66, C.K.B.’’. Three slides each with a single specimen labelled : ‘‘ Pseudococcus burnerae Brain ; on granadilla ;

1 Pseudococcus bechuanae is so close to lounsburyi that a careful and detailed study of a long series of specimens of both species might lead to the conclusion that they are actually synonyms or at most that they represent only forms of the same systematic species. As far as it can be presumed from the structures of the paratypes of both species and from supplementary material of Jounsburyi examined up to present, bechuanae differs from lounsburyi by having the tubular ducts of oral collar type confined to the abdominal segments, while in /ownsburyi they extend as far as the head.

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THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

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Fic. 6, Pseudococcus burnevae Brain

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA og1

Pretoria: Dec. 1914; paratypes; 66a, C.K.B.” Seven more slides with eighteen specimens labelled : ‘‘ Pseudococcus burnerae Brain ; Foxslave; Pretoria: 10.1.1915 ; Char. K. Brain; 47a, C.K.B.” were also seen, although they do not belong to the typical series.

“ Adult females with ovisacs usually aggregated in compact masses on the under- side of the leaves of the various host plants. The ovisacs are white, cottony, generally more or less spherical when complete, but the exact contour often obscured by the crowding of many together. The adult female is generally visible on top of the ovisac, the sides of which are raised around the insect. It therefore looks as though it had been pushed into the soft cottony mass. In other cases the insect is found at one side of the ovisac, which protrudes behind and below it, giving a decided Pulvinaria effect ... The adult female is so densely covered with white, powdery secretion that the general body colour is entirely obscured. The colour impression conveyed, however, is grey—perhaps slightly purplish or brown. There is no median, dorsal, clear patch as in citvi. All filaments are slender except the two caudal ones, which are more delicate than those of citvi. The caudal filaments may attain one-third the length of the body. The lateral ones are similar to those of capensis in that they are shortest at the anterior end, and gradually increase in length towards the posterior extremity. When mounted the adult female is about 2-5 mm. long and 1-5 mm. broad.”’ (Brain, /.c.).

The following redescription is based on a series of twelve specimens collected in Pretoria on 16.vii.1954 on Caralluma caudata N. E. Br. by Dr. H. K. Munro, which were compared with the Brain’s paratypes.

Outline of mounted specimens elliptical. Margin of body with seventeen pairs of cerarii. Anal lobe cerarii each built up with two conical spines beset with a small group of trilocular pores and four or five short auxiliary setae ; area about cerarian spines not chitinized. Each of remaining cerarii also with two spines somewhat shorter, except frontal (xvii) and ocular (xvi) each of which have three—occasionally four—spines ; each cerarius is surrounded by five to eight trilocular pores without auxiliary setae. Ventral side of anal lobes each with an ill-bordered slightly chitinized bar; apical seta robust, distinctly longer than those of anal ring; subapical one much shorter. Multilocular disc pores distributed in five groups on ventral side of last abdominal segments as follows: (v) 12-19; (vi) 15-21; (vii) 15-20; (viii) 22-26; (ix + x) 15-24. On segments anterior to genital opening they are arranged in transverse rows along distal margin of segment involved. Dorsal tubular ducts with oral rim few ; one duct is normally associated with each abdominal cerarius anterior to anal lobe and one duct is always present near each frontal cerarius; one duct on median area of first to third abdominal segments anterior to anal one ; occasional- ly a few ducts are interpolated on median or submedian areas of thorax. Ventral tubular ducts with oral collar crowded all along marginal area ; a few are associated with ventral multilocular disc pores. Trilocular pores not numerous and evenly distributed. Circular disc pores smaller than trilocular pores, very few on both sides of body. Dorsal setae few, all short and slender; ventral ones much longer and more abundant. Anterior and posterior dorsal ostioles rather inconspicuous with lips membranous. Circulus rather variable in size, normally small, roundish or

ENTOM. 7, 3. 4§§

92 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

elongate, with surface smooth. Legs well developed with some translucent pores on hind coxa, femur and tibia. Antennae with eight joints.

In our paper dealing with the Pseudococcidae described by H. C. James from East Africa (De Lotto, 1957) we sunk Pseudococcus simulator James as a synonym of muraltiae Brain. In that action we assumed the occurrence of a circulus in Brain’s paratype of muraltiae studied in London, but examination of further specimens showed that it is not present in the species. After examining one more paratype of muraltiae and the long series of slides of burnerae above listed, in the collection of the Department of Agriculture in Pretoria, we came to the conclusion that in these two species, which are structurally very similar, the only reliable character on which they depart is the circulus, which in muraltiae is entirely missing, whilst in burnerae is always present, as is the case in simulator. Therefore Pseudococcus simulator James is now synonymized with burnerae instead of muraltiae. In simulator the dorsal tubular ducts with oral rim tend to be fewer than in typical burnerae, but in all other respects they are identical.

Pseudococcus caffra Brain (Text-fig. 7) Pseudococcus caffra Brain, 1915, Trans. voy. Soc. S. Afr. 5: 138.

One single slide marked with the serial number of Brain’s collection “ 41, C.K.B.” was seen. It contained two specimens originally mounted without having been boiled in KOH or cleaned of the body content. In one specimen both antennae were partly broken away.

Adult female ‘‘ appeared to have made elongate ovisacs which had been crushed or broken in transit. There was no cottony material present, but rather a dense, powdery secretion. The adult female is orange-brown in colour except at the extre- mities, which are decidedly pinkish. The largest specimen seen measured 2:4 mm. in length. Mounted specimens average 1-8 mm. in length and 0-8 mm. broad.” (Brain, /.c.). Mounted specimens very elongate elliptical, membranous. Marginal and anal lobe cerarii entirely missing. Ventral side of anal lobes each with a robust apical seta, somewhat longer than those of anal ring; chitinized bar absent. Multi- locular disc pores of usual type very numerous on ventral side of last four abdominal segments ; on three segments anterior to genital opening they are arranged in trans- verse rows near distal margin; other pores are scattered on either side of body. | Other larger multilocular disc pores having twenty or more small loculi are fairly abundant on dorsum and on marginal area of abdomen. Tubular ducts with oral collar few on ventral side of last five abdominal segments. Trilocular pores absent. Circular disc pores small, few and scattered on both sides of body. Dorsal setae, few, very short, spiniform ; ventral setae more abundant and much longer. Anterior dorsal ostioles not recognizable ; posterior ones small, with lips membranous. Circulus absent. Legs long and slender with ungual digitules knobbed ; tarsal ones finely pointed ; tibiae of hind legs with a few translucent pores ; claw without denticle. In the specimen in which the antennae were intact, both were 8-jointed. ©

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 93

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Fic. 7. Pseudococcus caffra Brain

94 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Pseudococcus capensis Brain (= Pseudococcus maritimus (Ehrhorn)) Pseudococcus capensis Brain, 1912, Ann. ent. Soc. Amer. 5: 182.

This species was synonymized with Pseudococcus maritimus (Ehrhorn) by Joubert in 1928. A re-examination of some of Brain’s types fully supports this action.

Pseudococcus citri phenacocciformis Brain (= Planococcus citri (Risso)) Pseudococeus citvi phenacocciformis Brain, 1915, Trans. voy. Soc. S. Afr. 5: 116.

Recently Ezzat & McConnell (1956) recognized this variety as identical with Plano- coccus citri (Risso) with which they synonymized it. Two of Brain’s types were seen by us and we agree with Ezzat & McConnell’s opinion.

Pseudococcus elisabethae Brain (= Pseudococcus quaesitus Brain) Pseudococcus elisabethae Brain, 1915, Tvans. voy. Soc. S. Afr. 5: 126. .

One slide was seen containing four young adult females, three of which in rather poor condition. It was labelled: “‘ Pseudococcus rhenosterbosi Brain ; on rhenoster- bosch ; Newlands, C.P.: Jan. 17, 1915; paratype; B. 58, C.K.B.” Although the specific name on the label is different from that published by Brain, there are no doubts that the specimens actually belong to elisabethae inasmuch as the collection data and serial number are exactly the same as those reported in his original paper.

Pseudococcus elisabethae is structurally identical wih quaesitus described by Brain in the same paper and with which it is here synonymized, guaesitus having page precedence. Brain’s diagnosis of elisabethae was based on a series of very young adult females,

Pseudococcus flagrans Brain (Text-fig. 8) Pseudococcus flagrans Brain, 1915, Trans. voy. Soc. S. Afy. 5: 140.

Three slides containing altogether five specimens in rather good condition were seen. All slides were marked with the serial number of Brain’s collection: ‘69, C.K.B.”

“ Adult female: elongate, parallel-sided, about 4 mm. long, bright pink to flesh- coloured. No lateral filaments were observed, but in two cases short caudal ones were present, while one insect indicated that there may, in certain cases, be four caudal filaments instead of two ... Females ... up to 4 mm. in length.” (Brain, /.c.). Mounted specimens elongate elliptical, membranous. Margin of body with fifteen to seventeen pairs of cerarii. Anal lobe and preanal lobe cerarii each provided with two slender slightly lanceolate spines. Antepenultimate cerarii each with one or two spines smaller than those of anal lobes. Remaining cerarii each with a single spine ; the spines tending to be progressively smaller anteriorly. Area about cerarian spines not chitinized ; auxiliary setae absent. Apical seta of

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA _ 95

Fic. 8. Pseudococcus flagrans Brain

96 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

each anal lobe long and robust; dorsal chitinized bar absent. Multilocular disc pores of usual type abundantly distributed all over body and particularly numerous on ventral side of last four abdominal segments. Larger multilocular disc pores represented by a type with five loculi, seldom with four. They are numerous on both sides of body, particularly so on dorsum and marginal area of venter. Tubular ducts very small with inner portion somewhat chitinized, rather numerous on either side of body. Circular disc pores small, not abundant and scattered on venter and dorsum. Trilocular pores entirely absent. Ventral and dorsal setae long, robust and fairly numerous. Anterior and posterior dorsal ostioles rather inconspicuous, lips membranous. Circuli four, circular or nearly so in shape, with surface membranous, rim thick and heavily chitinized. First two circuli about same diameter, third one smaller ; the posterior one smaller still and about half the size of anteriors. Legs all well developed with a dentical on claw ; tarsal digitules finely pointed, ungual ones knobbed at apex. No translucent pores were seen on hind legs of specimens examined. Antennae with nine joints. Anal ring entire with six robust setae distinctly shorter than apical seta of anal lobes.

Pseudococcus fragilis Brain

Pseudococcus fragilis Brain, 1912, Ann. ent. Soc. Amer. 5: 186. Pseudococcus gahani Green, 1915, Ent. mon. Mag. 51: 179.

At page 351 of his book College Entomology, Essig (1942) recorded Pseudococcus gahant Green, I915 as a.synonym of P. fragilis Brain, 1912, without giving any indication how he came to this conclusion or on whose authority it was based. In recent correspondence received from Mr. C. J. Joubert of the Stellenbosch-Elsenburg Agricultural College, Stellenbosch, South Africa, we have been informed that several years ago he studied the identity of fragilis and gahamt which he recognized structurally identical. According to him, in 1933 he communicated his finding to Essig who apparently took for granted that by the time his text-book was issued Joubert would have published the result of his investigation, which he has not done.

Independently we came to the same conclusion when at the British Museum (Natural History), London, we compared types or paratypes of both species.

Pseudococcus lounsburyi Brain Pseudococcus lounsburyi Brain, 1912, Amn. ent. Soc. Amer. 5: 179.

Two slides were available. One with a single specimen, the other with five, all rather young adult females with many setae and cerarian spines broken away and partly attacked by fungi. Both slides bore the following label: ‘‘ Pseudococcus lounsburyi Brain; paratype; but setae of anal lobes lacking; on Agapanthus sp.; Kenilworth, S.A.: 14.ix.1910.”

Two excellent redescriptions of this species were recently published by Ferris (in Zimmerman, 1948; Ferris, 1950). On some remarks on the species contained in the Atlas of the Scale Insects of North America, it is pointed out that according to Morrison’s opinion the specimens from Hawaii determined by Ferris as lounsburyi

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA _ 097

and used for diagnosis and figure, although very close to it, would not belong to that species in view of a noticeable difference in the number of multilocular disc pores occurring on the ventral side of the abdominal segments. Besides the paratypes above mentioned, at the British Museum (Natural History), London, we examined another slide of the same typical series, and supplementary specimens are at hand from South Africa on Agapanthus sp. and on bulbs of Narcissus sp. as well as from Kenya also on Agapanthus sp. We are prepared to accept the insect redescribed by Ferris as lounsburyi, since the number of multilocular disc pores in the species in question is variable and occasionally a few occur even on the dorsum of the last two or three segments anterior to the anal one. Even the number of tubular ducts with oral rim is rather variable.

Pseudococcus mallyi Brain (Text-fig. 9) Pseudococcus mallyi Brain, 1915, Trans. voy. Soc. S. Afr. 5: 120.

The material examined was represented by two slides containing altogether three specimens in fairly good condition, labelled: ‘‘ Pseudococcus mallii! Brain ; on grass ; Rosebank, C.P.; Nov. 25,1914; paratype; Char. K. Brain ; 32, C.K.B.” Four other slides not belonging to the typical material, although marked with the same serial number of Brain’s collection, were seen. They bore the following label : “ Pseudococcus mallyi Brain; on grass; Rosebank Exp. Sta.: May 10, 1915; Bet CRs 3a,C.KB,”’

“No definite ovisac was seen, but adult females and young were clustered in a white powdery secretion in the leaf sheaths ... + The adult is bright rose-pink in colour, some specimens being uniformly covered with white powder. There were no traces of filaments of any kind. Legs and antennae colourless. The insect is exceptionally long and narrow, mounted specimens averaging 2:16 mm. long by 0-75 mm. broad.” (Brain, /.c.). Body very elongate elliptical, membranous. Cerarii recognizable only on last three or four abdominal segments. Anal lobe cerarii each with two rather long slender spines of different size. On each of preceding cerarii spines tend to be more slender and set apart each other. Cerarii on third segment anterior to anal lobes at times represented by a single spine. All cerarii without auxiliary setae or grouping of trilocular pores; area about spines not chitinized. Ventral side of each anal lobe without chitinized bar ; apical seta robust and longer than those of anal ring ; subapical one much shorter. Multilocular disc pores rather numerous and distributed on either side of body ; on ventral side of two segments anterior to genital opening several pores are crowded in transverse rows along distal margin. Trilocular and quinquelocular pores entirely absent. Tubular ducts of oral collar type very short, not numerous and distributed on venter and dorsum without any peculiar arrangement. Circular disc pores few and scattered on both sides of body. Dorsal setae very short, spiniform, rather few ; ventral setae more numerous and much longer. Anterior dorsal ostioles absent ; posterior ones incon-

1 Evidently a pen slip for mailyi,

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

98

Pseudococcus mallyi Brain

9.

Fic.

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. Afy. 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!; 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, l.c.). 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 in 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

too THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 10. Pseudococcus mirabilis Brain

i ——

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA to1

Pseudococcus muraltiae Brain (Text-fig. 11) 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 I-g 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, /.c.). 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; (vil) 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. Anteriorand posterior dorsal ostiolesinconspicuous withlipsmembranous 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, Tvans. 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 Nzpaecoccus

1io2 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

sr -<

e}

Fic. 11. Pseudococcus muraltiae Brain

—

THE PSEUDOCOCCIDAE (HOM. 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, /.c.). 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.

10o4 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 12. Pseudococcus nitidus Brain

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 105

Pseudococcus quaesitus Brain (Text-fig. 13)

Pseudococcus quaesitus Brain, 1915, Trans. voy. Soc. S. Afr. 5 : 123. Pseudococcus elisabethae Brain, 1915, Tvans. voy. Soc. S. Afr. 5: 126 (SYN. NOV.). Pseudococcus trichiliae Brain, 1915, Tvans. 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: “ of Pseudococcus quaesitus Brain; Pretoria; 3 paratype; gf 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, /.c.). Body of mounted specimens elliptical. Margin of body with a series of seventeen pairs of cerarli. 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; (vil) 21; (vili) 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 (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 13. Pseudococcus quaesitus Brain

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 107

Pseudococcus segnis Brain (Text-fig. 14) Pseudococcus segnis Brain, 1915, Tvans. voy 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: 17.xii.1914; B. 55, C.K.B.” Although the collecting data is not exactly the same recorded in Brain’s paper, that is 13th 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, /.c.). Body elliptical with eighteen pairs of cerarii. Anal lobe cerarii each apparently formed by two spines! 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.; 52 B.”’ 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 soczalis 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.

108 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 14. Pseudococcus segnis Brain

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 109

Fic. 15. Pseudococcus socialis Brain

110 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

long and o-8 mm. broad.” (Brain, /.c.). 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; (vili) 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.

“Qvisac: the ovisacs are rounded masses of cottony material ... In form they appear almost spherical ... The greatest diameter averages approximately 25 mm. ... The adult female is pale canary-yellow in colour, about 2 mm,

THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA 111

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Fic. 16. Pseudococcus stelli Brain

112 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, /.c.). 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. voy. Soc. S. Afr. 5: 149.

This variety was described on a few specimens collected on the same twigs as those attacked by P. stellt 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 st¢el/i with which it is here synonymized.

Pseudococcus transvaalensis Brain

Pseudococcus transvaalensis Brain, 1915, Trans. voy. 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.1914; 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. Afy. 5: 131.

Three slides containing altogether six specimens labelled: ‘‘ Pseudococcus trichliae Brain ; on Trichila sp.; Durban, Natal; 27.x.1I914; 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 tvichiliae 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.19g10; 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, /.c.).

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; (vili) 34; (ix+ x) 26. On

1144 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 17. Pseudococcus wachendorfiae 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. voy. Soc. S. Afy. 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, D.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, /.c.). 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 setat 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 dorsumonly. 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.

116 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Fic. 18. Puto africanus Brain

THE PSEUDOCOCCIDAE (HOM. 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 (= Rhizoecus 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 tenutfolia ; 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 2mm. to 2:5 mm. long.” (Brain, /.c.). 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. Maultilocular 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.

118 THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

4)

md

Fic. 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. Amn. ent. Soc. Amer. 5 : 177-189.

1915. The Coccidae of South Africa, I. Tvrans. 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.

Essic, E.O. 1942. College Entomology, pp. vii + 900, McMillan Co., New York.

Ezzat, Y. M. & McConneELL, H. S. 1956. A classification of the Mealybug Tribe Plano- coccini (Pseudococcidae, Homoptera). Bull. agric. exp. Stn., Univ. Maryland, A—84,

. 108. Beas, 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.

1zo THE PSEUDOCOCCIDAE (HOM. COCCOIDEA) FROM SOUTH AFRICA

Hatt, W. J. 1937. Observations on the Coccidae of Southern Rhodesia. VIII. Trams. 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. Tvamns. 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). J. Wash. Acad. Sci. 35 : 38-55.

Witurams, 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 ‘

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

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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 Degeertella-complex.

THE Degeertella-cOMPLEX. It is difficult to delimit this group exactly but the following genera should probably be included : Degeeriella Neumann (= Kélerinirmus Eichler), Acutifrons Guimaraes, Austrophilopterus Ewing, Capraiella Conci, Cotin- gacola Carriker, Cuculicola Clay & Meinertzhagen, Lagopecus Waterston, (= Colini- cola Carriker), Picicola Clay & Meinertzhagen (= Tyrannicola Carriker), Tvogoni- niyrmus 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 Briielia (Clay, 1951) ; pulvinus usually in the form of a single sac-like structure. 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 Degeerielia- and Briielia-complexes, however the species of Degeeriella from Chelictina and Elanoides have the characters of these structures as in some species of Briielia (see Clay, 1958).

ENTOM. 7, 4. 5

124 REVISIONS OF MALLOPHAGA GENERA

setae elongated. Prothorax with one marginal seta each side (except in Lagopoecus meinerizhagent Clay); third episternum fully sclerotized laterally. Abdomen with postspiracular setae on segments III-VII' (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. mookenjeei 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. 1) 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 Ausiro- gonotdes, Osculotes, Chelopistes and members of the Heptapsogaster-complex. The Otidoecus-complex (Otidoecus, Rhynonirmus and Cuclotogaster) have an unpaired diverticulum arising from the ductus ejaculatorius?. 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. Brielia) ; genital region without lateral spine-like setae (cf. Rallicola) or clump of setae on tubercle-like area (cf. Briielia). 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 Degeertella-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. 1, fig. 1). 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 vierai 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 Galliformes 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 (nec Hermann, 1804).

Degeeriella. Neumann, 1906, Bull. Soc. zool. Fr. 20:60. Nomen novum for Nirmus Nitzsch nec Hermann. Type species by subsequent designation, Johnston & Harrison, 1911, Proc. Linn. Soc. N.S.W. 36 : 326: “‘ D. discocephalus N.”’

Kélerinirmus. Eichler, 1940, Zool. Anz. 130: 101. Type species: ‘‘ Nirymus fuscus Nitzsch in Denny.”

Ischnocera not exceeding 3 mm. in length; usually without marked sexual dimorphism, but the females average