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The genus Epeolus Latreille fromsubsaharan Africa (Hymenoptera:Anthophoridae)C.D. Eardley aa National Collection of Insects , Plant Protection ResearchInstitute , Private Bag X134, Pretoria, 0001, South AfricaPublished online: 17 Feb 2007.
To cite this article: C.D. Eardley (1991) The genus Epeolus Latreille from subsaharanAfrica (Hymenoptera: Anthophoridae), Journal of Natural History, 25:3, 711-731, DOI:10.1080/00222939100770471
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JOURNAL OF NATURAL HISTORY, 1991, 25, 7 1 1 - 7 3 1
The genus Epeolus Latreille from subsaharan Africa (Hymenoptera: Anthophoridae)
C. D. EARDLEY
National Collection of Insects, Plant Protection Research Institute, Private Bag X134, Pretoria, 0001, South Africa
(Accepted 27 November 1990)
The genus Epeolus from subsaharan Africa is revised. Eleven species are recognized, of which E. cestus is described as new. Ten species and one subspecies names have been placed in synonymy, and E. caffer (Lepeletier), is transferred to the genus from Crocisa Jurine. A checklist and a key for the identification of the subsaharan species are given.
KEYWORDS: Anthophoridae, Epeolus, subsaharan Africa, taxonomy.
Introduction The genus Epeolus Latreille belongs to the Epeolini, a tribe of the large
cosmopolitan subfamily Nomadinae, family Anthophoridae. All the nomadine bees are cleptoparasitic and an outstanding discussion on cleptoparasitism and the phylogeny of the Nomadinae has been given by Alexander (1990).
The Epeolini apparently occur on all continents except Australia (Michener, 1965). The tribe is commonly recognized as comprising six genera. Five of these are confined to the New World and Epeolus occurs in the Old World and North America (Rozen, 1989). They are known to be cleptoparasitic in the nests of Colletes Latreille (Friese, 1915; Rozen, 1966, 1977) and Tetralonia Spinola (Medler, 1980).
In subsaharan Africa the genus is represented by 11 species, eight of which appear to be endemic to southern Africa, two are widespread and one is known only from Ethiopia.
This is the first detailed study of the subsaharan species of Epeolus and it necessitated the examination of all the type material as well as the redescription of all the previously known species from the area. Bischoff (1923) provided a key to the species, in which he described several new species, but this study is outdated and cannot be used for reliable identifications.
Methods Sexual dimorphism in Epeolus is slight and, apart from the number of antennal
segments, is largely limited to the distal region of the metasoma. Detailed descriptions of only the female of each species have therefore been included, and the diagnostic sex- limited characters of the males have been added in parentheses in the female descriptions. The seventh and eighth metasomal sterna and the genitalia of the males do not differ significantly between species and have not been dealt with.
The abbreviations T and S are used in the descriptions for the metasomal terga and sterna, respectively. The legs are described as though extended laterally, with the axis of
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the limb perpendicular to the axis of the body. I have generally followed the terminology used by Michener (1944), but have referred to the posterior region of the dorsal surface of the pronotum as the pronotal collar, and the line of dense pubescence that extends from the middle of each half of the anterior margin of the scutum backwards to a point in about the same plane as the anterior ends of the tegulae as the anterior mediolateral scutal stripe. The distribution maps include both the localities of the material examined during this study and published distribution records.
Institutions and collections The material on which this study was based is in the institutions recorded below,
and these are referred to by the accompanying abbreviations.
BLCU DMSA LACM
MNHN MRAC NCSA
NHML SAMC
SC SMWH TMSA
ZMHB
Utah State University, Bee Biology and Systematics Laboratory, Logan Durban Natural Science Museum, Durban Los Angeles County Museum of Natural History, Los Angeles Musrum National d'Histoire Naturelle, Paris Musre Royal d'Afrique Centrale, Tervuren National Collection of Insects, Plant Protection Research Institute, Pretoria The Natural History Museum, London South African Museum, Cape Town Schwarz private collection, Ansfelden, Austria State Museum, Windhoek Transvaal Museum, Pretoria Museum fiir Naturkunde der Humboldt-Universit/it, Berlin
Genus Epeolus Latreille
Epeolus Latreille, 1802: 427; Sandhouse 1943: 548. Type species: Nomada varieoata Fabricius, 1775 (monobasic).
With the exception of E.friesei Brauns, in which the axilla is simple, both sexes of all the species of Epeolus can be easily recognized by the presence of a well-developed, posteriorly projecting, spine on the axilla. The genus is characterized in subsaharan Africa in that it is the only anthophorid genus with patches of short, pale, appressed, squamiform pubescence, in which the female has a transverse pseudopygidial area on the fifth metasomal tergum that is curved gently downwards and clothed with fine silvery pubescence. The male also has patches of short, pale, appressed, squamiform pubescence, but it can be identified by the well-developed subapical fringes on the fourth and fifth metasomal sterna.
The subsaharan species of Epeolus are divided into three species groups, namely the E. natalensis, E. pyomaeorum and the E. caffer species groups. The first group comprises E. natalensis Smith, E. kristenseni Friese and E. rufothoracicus Bischoff; the second group includes E. pyomaeorum Cockerell, E. alatus Friese, E. amabilis Gerstaecker, E. pubescence Cockerell, E. fulviventris Bischoff and E. cestus and the third group contains E. caffer (Lepeletier) and E. friesei. The placement of species in the first two groups is based on the structure of the axilla. In the E. caffer group the axillae are dissimilar and unlike the remainder of the subsaharan fauna. The axilla is largely fiat dorsally and carinate laterally in the E. natalensis species group and strongly carinate
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The genus Epeolus Latreille f rom subsaharan Africa 713
dorsolateral ly in the E. pygmaeorum species group. Al though E. caffer and E. fr iesei
cannot be grouped by the s tructure of their axillae, they do show a r emarkab le overall similarity.
K e y t o t h e s p e c i e s
1 Axilla without a posteriorly projecting spine . . E. friesei Brauns - Axilla with a well-developed, posteriorly projecting spine 2
2 Labrum without a distinct mediolongitudinal concavity; spine on axilla not carinate, but cylindrical and curved dorsolaterally (Fig. 15); scutellum with mediolongitudinal groove either absent or very weakly developed, and with each half of scutellum not distinctly raised medially . . . . . . . . E. caffer (Lepeletier)
- Distal half of mediolongitudinal region of labrum with a shallow, distinct concavity; axilla carinate and projecting more or less directly backwards, distal end sometimes curved laterally (Figs 3, 8); scutellum with mediolongitudinal groove moderately well developed to very well developed, and with each half strongly rounded medially. 3
3 Pronotal collar very densely pubescent 4 - Pronotal collar sparsely pubescent 7
4 Scutellum and axilla black. 5 - Scutellum and axilla orange 6
5 Labial palp relatively long, subequal in length to antennal flagellar segment II (known from central, eastern and southern Africa) . . . E. natalensis Smith
- Labial palp short, about half as long as antennal segment II (known only from Ethiopia) E. kristenseni Friese
6 Axilla with a well-developed lateral carina (cf. Fig. 3) . E. rufothoracicus Bischoff - AxiUa with a well-developed dorsolateral carina (cf. Fig. 8i E. fulviventris Bischoff
7 Pallid bands on T3-T4 (T3-T5 of males) either continuous of narrowly interrupted medially (Figs 12, 13) . . . . 8
- Pallid bands on T3-T4 (T3-T5 of males) t~roadiy interrupted medially (Fig. 9) i 9
8 Scutum and propodeum completely black (occurring on west coast of Namibia) E. pubescence Cockerell
- Scutum and propodeum each with at least lateral regions reddish-orange (occurring in Zimbabwe and the Transvaal province in South Africa) E. cestus sp. nov.
9 Integument of head, mesosoma and metasoma completely black . 10 - Head, mesosoma and metasoma black with many orange or orangish areas
E. amabilis Gerstaecker
10 Face moderately densely clothed with long white pubescence (Fig. 6); anterior mediolateral scutal stripes well developed (Fig. 8); mesopleuron moderately densely clothed with long white vestiture . . . . . . E. pygmaeorum Cockerell
- Face with upper third of clypeus, supraclypeus, supra-antennal area and paraocular area very densely clothed with short white vestiture (cf. Fig. 1); scutum without anterior mediolateral stripes; mesopleuron densely clothed with short white vestiture
E. alatus Friese
The Epeo lus natalensis species g roup
This species g roup comprises E. natalensis, E. kristenseni and E. rufothoracicus. I t is character ized by the distinctly spinose axilla tha t is largely fiat dorsal ly and carinate laterally.
Apar t f rom the similarity in the structure of their axillae, these three species all have densely pubescent p rono ta l collars and are devoid of anter ior mediola tera l scutal
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714 C.D. Eardley
stripes. But these characters are not unique to the group. Epeolus alatus and E.fulviventris, which belong to the E. pyomaeorum species group, resemble this group in that they do not have anterior mediolateral scutal stripes, and E.fulviventris also has a very densely pubescent pronotal collar.
Epeolus natalensis Smith
(Figs 1-4, 19)
Epeolus natalensis Smith, 1879:101 ($ holotype, NHML). Epeolus coelostoma Bischoff, 1923:601 (~ holotype, ZMHB), syn. nov. Epeolus bidens Bischoff, 1923:601 (6 holotype, ZMHB), syn. nov. Epeolusfoveilabris Bischoff, 1923:601 (~ holotype, TMSA), syn. nov. Epeolus 91yptochilus Bischoff, 1923:601 (~ leetotype, TMSA), syn. nov.
Bischoff(1923) was presumably not aware of E. natalensis, as he did not include it in his key to the subsaharan species of the genus. Epeolus coelostoma, E. bidens, E.foveilabris and E. 91yptochilus were separated by Bischoff(1923) on the sculpture and the number of denticles on the labrum. Careful examination of the labrum of each specimen studied revealed that the structures that Bischoff (1923) considered to be diagnostic characters for separating the species mentioned above have numerous intermediate forms, and these differences can be more realistically attributed to intraspecific variation. I, therefore, regard them as being conspecific. Epeolus
FIGS 1-5. Epeolus spp. 1-4. E. natalensis, female: 1, frontal view of head; 2, external view of labrum; 3, dorsal view ofmesosoma; 4, dorsolateral view of metasoma; 5, E. rufothoracicus, external view of female labrum.
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The genus Epeolus Latreille from subsaharan Africa 715
glyptochilus was described from three male and four female specimens, a male from Rikatla in Mozambique is here designated as the lectotype of this species.
Description. Lengths. Head 2-1-2.4 mm; scutum 1-7-2-1 mm; fore wing 6.1-7.9 mm; body 8-4-9.9 mm.
Colour of integument. Black. Vestiture. Face as in Fig. 1, with upper third of clypeus, paraocular area,
supraclypeus and supra-antennal area very densely clothed with white, plumose vestiture; remainder of clypeus and labrum short, simple and black; vestiture on vertex sparse, long, white and simple, and on gena short, white and plumose; pronotum with short, white, sparse, plumose vestitute, except collar very densely pubescent, as in Fig. 3, and pronotal lobe surrounded by a dense white fringe; mesosomal dorsum, as illustrated in Fig. 3, mostly sparsely pubescent, vestiture mostly long, white and simple; anterior, anterolateral and posterolateral regions of scutum more densely pubescent; posteromesal region of tegula densely clothed with white vestiture; mesopleuron with dorsal half of lateral surface very densely clothed with white pubescence, anterior surface and ventral half of lateral surface sparsely clothed with very short black vestiture; metapleuron clothed with white pubescence, that on anterior region more dense than posterior region; posterior, subvertical surface of scutellum, entire metanotum and posterolateral region of propodeum densely clothed with white vestiture; remainder of propodeum largely naked; fore and middle legs with lateral regions of coxae, posterior surfaces of femora and dorsal surfaces of tibiae densely clothed with white pubescence; fore and middle basitarsi with simple white vestiture above and orangish setation below; outer surface of hind coxa clothed with white vestiture; hind femur sparsely clothed with white vestiture above and black anteriorly; dorsal surfaces, especially posterodorsal surfaces, of hind tibia and basitarsus fairly densely clothed with white vestiture; hind basitarsus brownish-orange below; meta- soma with following areas densely clothed with short white vestiture (Fig. 4): subvertical anterior, dorsolateral, laterally and posterolateral regions of T1 white; T2 with a medially, broadly interrupted distal band, and sometimes with lateral regions white; T3 with a medially, broadly interrupted distal band, and distal margin ofT4 with either a continuous or medially interrupted distal band; T5 with posterolateral region white and pseudopygidium silvery-white (male T5-T6 with a continuous, sparsely pubescent, white bands on distal margins); posterolateral regions of $4-$5 often with a little white vestiture (male with a little white vestiture on posterolateral regions of $2- $3, $4-$5 with subapical fringes of long black hairs, $6 black); remaining areas either naked or sparsely clothed with short simple black vestiture.
Structure. Head: frontal carina well developed, rising gradually from a point about half way between antennal socket and median ocellus to end abruptly in same plane as lower level of antennal sockets; upper region of carina with lateral sides steeply raised and lower region of carina only slightly raised above swollen inter-antennal socket area; labrum with median region mostly gently concave (Fig. 2); distal edge of labrum usually concave medially, with one apicomedial tubercle and with one to two subapicolateral teeth (Fig. 2); mandible with inner, subapical tooth small and tuberculate; maxillary palp two-segmented, and about as long as segment II of antennal flagellum. Mesosoma: dorsum moderately densely punctate; axilla spinose and with a well-developed carina on lateral surface (Fig. 3); scutellum strongly concave mediolongitudinally and each half distinctly convex (Fig. 3). Metasoma: (male with $4-$5 strongly concave posteromesally and with well-developed subapical fringes of long black hairs that curve towards mediolongitudinal axis of metasoma).
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716 C.D. Eardley
Distribution (Fig. 19). This species is known from numerous localities from Zaire, Uganda and Kenya, in the north, to Namibia and the Orange Free State and Natal, South Africa, in the south.
Discussion. Within the E. natalensis group this species is remarkably similar to E. kristenseni, differing mainly in the relative length of the maxillary palp. The structure of the maxillary palp of this species is unique among the subsaharan fauna. Epeolus natalensis can be easily separated from E. rufothoracicus by the colour of the integument and the vestiture, and by the structure of the labrum.
Material examined. Type material. Epeolus natalensis, 3` holotype: 'Pt. Natal, 57 3, Epeolus natalensis Type Sm., B.M. Type Hym. 17B 510', NHML.
Epeolus coelostoma, ~ holotype: 'Rikatla, Delagoa, (Junod), Epeolus coelostoma Bisch. ~ Type', ZMHB.
Epeolus bidens, 3` holotype: 'Nguelo, Usambara, H. Rolle, Berlin SW 1, Epeolus bidens Bisch. Type 3`', ZMHB.
Epeolus foveilabris, 3` holotype: 'Delarey, W. Transvaal, Dr Brauns, 15.1.1917, Epeolus foveilabris 3 ̀Bisch. Type', TMSA.
Epeolus glyptochilus, 3` lectotype: 'Port Natal, Ostafrica, Dr Brauns, 24.4.93, Epeolus glyptochilus Bisch. 3 ̀ Type', TMSA; ~ paralectotype E. glyptochilus: 'Johannesburg, Transvaal, G. Kobrow, February 1906, Epeolus glyptochilus Bisch.
Type', TMSA. Additional material. 16~ 243`: ZAIRE: Kivu, Mulungu, Tshibinda, xi.1951, P. C.
Lefrvre, 2~ MRAC; Kivu, Bwito, 26.vi.1934, Lt Marlier, 1~ MRAC; W. Kivu, Loashi, viii.1937, J. Ghesquirre, lc~ MRAC; Kivu, Lubero, Mulo, vi-vii.1953, R. P. M. J. Crlis, 13 ̀MRAC; Elisabethville [ = Lubumbashi], 30.v.1933, Dr M. Bequaert, 1~ 13` MRAC. KENYA: Laragei Springs, 12 km SW Maralal, Samburu district, 17-22.i.1973, J. P. Donahue, 1~ LACM. ZIMBABWE: Vumba mountains, 20 km SE Mutare, 19.iv.1985, J. Gusenleitner, 1~ SC; Selukwe, 19.i.1922, 13` SAMC; Umtali district, 4.i.1931, P. A. Sheppard, 13 ̀TMSA; Salisbury I-=HaraeJ, i.1900, G. A. K. Marshall, 1~ NHML. SOUTH AFRICA: TRANSVAAL: Entabeni Forest Reserve, Soutpansberg, 23.00S 30.16E, 3-7.xi.1980, R. S. du Plessis, 1~ NCSA; Woodbush, ii.1977, G. L. Prinsloo, 13 ̀NCSA; Mogoto Nature Reserve, Zebediela, 24.15S 29.13E, 22-25.x.1979, C. D. Eardley, lc~ NCSA; Haenertsburg, 9.xii.1909, C. J. Swierstra, 13` TMSA; Rustenburg, Nature Reserve, 25.40S 27.12E, 23-26.ii.1981, C. D. Eardley, 1~ lc~ NCSA; Renosterkop, Pretoria, 25.46S 28.17E, 30.iii.1990, M. Schwarz, 1~ SC; Pretoria, Fountains, ii.1901, R. v. Jutr, 13' TMSA; Rietvlei Dam, 13.viii.1980, H. Empey, 13` NCSA; Johannesburg, ii.1906, G. Kobrow, 17 TMSA; Florida, xii.1918, R. W. Tucker, 13` SAMC; Strubens Valley, Florida, different dates in December to February, H. Empey, 3~ 43` NCSA; NATAL: Pinetown, 29.xii.1916, H. W. Bell-Marley, 1~ DMSA; Weenen, i.1924, H. P. Thomasset, 13` NCSA; ORANGE FREE STATE: Adullan farm, near Clarens, 28.34S 28.28E, different dates in January and February, different collectors, 2~ 33` NCSA. NAMIBIA: Kaoko Otavi, iii.1926, 13' SAMC; Gobabis, 21.xii.1974, H. Empey, 13 ̀NCSA; Otjiwarongo, ii.1978, S. J. van Tonder, 23` NCSA.
Epeolus kristenseni Friese
(Fig. 19)
Epeolus kristenseni Friese, 1915: 265, 290-291 (~ lectotype, TMSA).
The female syntype of E. kristenseni is here designated as the lectotype of this species. As I have neither been able to locate the male paralectotype nor had the
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The genus Epeolus Latreille from subsaharan Africa 717
opportunity to study other males of this species, I have not included any features of the male in the description below. These can be obtained from the original description of this species.
Description. Female. Similar to E. natalensis except as follows: mesosomal venter sparsely clothed with fine, simple, white vestiture; T2 without any white vestiture on lateral region; posterior regions of $3-$5 mostly clothed with white pubescence; maxillary palp two-segmented and about half as long as segment II of antennal flagellum.
Distribution (Fig. 19). This species is only known from Ethiopia. Discussion. The relationships between this species and the remainder of the group,
including its closest relative, are dealt with in the discussion on E. natalensis. Epeolus kristenseni was recorded, in the original description, as being a parasite of
Colletes abessinicus. Material examined. Type material. Epeolus kristenseni, c~ lectotype: 'NO.-Afrika,
Abessinien, Harrar 11, Kristensen, Epeolus kristenseni 1912 Friese det. ~', TMSA. Additional material. 1~: ETHIOPIA. Le Kempti ( = Nak'amet), 2200 m, 25.v. 1946,
K. M. Guichard, 1~ NHML.
Epeolus rufothora¢icus Bischoff
(Figs 5, 20)
Epeolus rufothoracicus Bischoff, 1923:600 (~ lectotype, TMSA). Epeolus haemataspis Cockerell, 1937:160 (9 holotype, NHML), syn nov. Epeolus roseatus Cockerell, 1937:160 (~ holotype, NHML), syn. nov.
Cockerell (1937) separated E. haemataspis and E. roseatus on the basis of several small differences in the width of the median interruptions of the yellowish distal bands on the metasomal dorsum, and the colour of the clypeus and the second metasomal tergite. Detailed study of the types of E. rufothoracicus, E. haemataspis and E. roseatus, together with several additional specimens ofE. rufothoracicus, clearly revealed that the differences described by Coekerell are within the range of intraspecific variation of this species. Cockerell made no comment on the similarity between E. rufothoracicus, E. haemataspis and E. roseatus, which suggests that he was not aware of their similarity.
Description. Lengths. Head l-6-2.0mm; scutum 1.5-1.8 mm; fore wing 5"5-6-6 mm; body 5-9-8-32 mm.
Colour of inteoument. Black to orangish-black with clypeus, labrum, mandible, antenna, pronotal lobe, anterolateral region of scutum, tegula, axiUa, scutellum, mesopleuron and legs usually mostly orange.
Vestiture. Head moderately densely clothed with white vestiture, except area between upper limit of frontal line and upper margin of clypeus densely pubescent, cf. Fig. 1; pronotum sparsely clothed with very short, plumose, yellowish vestitute; except pronotal collar very densely clothed with yellow pubescence (cf. Fig. 3), and with a fairly dense white fringe surrounding pronotal lobe, similar to E. natalensis; mesosomal dorsum sparsely clothed with a mixture of both simple and plumose yellowish vestiture, mesal region sometimes a little more densely pubescent, and anterolateral and posterolateral regions white and generally more densely pubescent; metanotum densely clothed with white vestiture; remainder of mesosoma sparsely clothed with short, white, plumose vestiture, except region near propodeal spiracle with long vestiture and posterior surface of propodeum naked; legs sparsely to moderately densely clothed with pallid vestiture, except ventral surfaces of tarsi orangish; T1
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718 C.D. Eardley
similar to E. natalensis, except pallid vestitute yellowish-white; T2-T4 black with medially interrupted, yellowish-white bands on distal margins; T5 sparsely clothed with very short yellowish-white vestiture and with a silvery-white pseudopygidium (male T5-T6 with continuous white bands on distal margins), remaining areas of metasomal dorsum either naked or sparsely clothed with short, simple, black hairs; metasomal venter generally white to yellowish-white (male with $4-$6 black to orangish-black, $4-$5 with subapical fringes of long black hairs, similar to E. natalensis).
Structure. Similar to E. natalensis, except distal edge of labrum always devoid of an apicomedian tubercle and with only one subapicolateral tooth, as in Fig. 5; maxillary palp two-segmented and about half as long as segment II of antennal flagellum.
Distribution (Fig. 20). Epeolus rufothoracicus is apparently endemic to South Africa. Discussion. Epeolus rufothoracicus is most closely related to E. natalensis and
E. kristenseni. Within this group it can be easily recognized by the colour of the integument and the vestiture, and by the structure of the labrum.
Material examined. Type material. Epeolus rufothoracicus, ~ lectotype: 'Algoa Bay, Capland, Dr Brauns, 22.11.96, Epeolus rufothoracicus Bisch. ~ Type', TMSA; paralectotype E. rufothoracicus as in lectotype except '26.4.96', TMSA.
Epeolus haemataspis, ~ holotype: 'Ceres, Cape Province, 1300 ft., Dec. 1920, Epeolus haemataspic Ckll. TYPE, B.M. TYPE HYM. 17B 512', NHML.
Epeolus roseatus, J; holotype: 'Port St John, Pondoland, Jan. 1924, Epeolus roseatus Ckll. TYPE, B.M. TYPE HYM. 17B 509', NHML.
Additional material. 39, 4c~: SOUTH AFRICA: NATAL: Port Edward, 21.iii. 1969, L. C. Starke, 19, 1~ NCSA; CAPE PROVINCE: Kenton-on-Sea, 21.xii.1979, H. Empey, 1~ NCSA; Pearly Beach, Bredasdorp, xii.1958, 19 SAMC; Upper Sources, Olifants River, near Ceres, xii.1949, 2~ SAMC; Ceres, xii.1920, R. E. Turner, 19 NHML.
The Epeolus pygmaeorum species group
The following species are included in this group: E. pygmaeorum, E. alatus, E. amabilis, E. pubescence, E.fulviventris and E. cestus. In these six species the axilla is spinose and strongly carinate dorsolaterally.
Most of these species also have a sparsely pubescent pronotal collar and distinct anterior mediolateral scutal stripes. In E. fulviventris the pronotal collar is densely pubescent, and neither E. fulviventris nor E. alatus have anterior mediolateral scutal stripes. In these characters they resemble the E. natalensis species group more closely.
El~olus pygmaeorum CockereU
(Figs 6-9, 19)
Epeolus pygmaeorum Cockerell, 1932:115-116 (~ holotype, NHML).
Description. Lengths. Head 2.3-2-6 mm; scutum 2-0-2.4 mm; fore wing 6.8-7.6 ram; body 8.1-9.9 mm.
Colour of integument. Mostly black, labrum, mandible and legs partly orange. Vestiture. Head moderately densely clothed with long white vestiture, upper third of
clypeus, paraocular area, supraclypeus and supra-antennal area a little more dense and comprising plumose hairs (Fig. 6); pronotum sparsely clothed with white vestiture; scutum sparsely clothed with simple yellowish vestiture, except anterior mediolateral scutal stripe, anterolateral and lateral scutal margin adjacent to anterior half of tegula
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The genus Epeolus Latreille from subsaharan Africa
, . . ! , .
719
11
FIGS 6-11. Epeolus spp. 6-9. E. pygmaeorum, female: 6, frontal view of head; 7, external view of labrum; 8, dorsal view of mesosoma; 9, dorsolateral view of metasoma; 10, E. amabilis, frontal view of female head; 11, E. fulviventris, external view of female labrum.
and posterolateral region of scutum densely clothed with white plumose vestiture; tegula with anterior and anterolateral regions clothed with white pubescence and with a little white vestiture posteriorly; mesosomal dorsum (Fig. 8) with dorsal half of mesopleuron and posterolateral region of propodeum densely clothed with long white vestiture; remainder of mesosoma sparsely clothed with short white pubescence, except anteroventral region of lateral surface of propodeum black and posterior surface of propodeum largely naked; fore, middle and hind legs similar to E. natalensis, except middle and hind coxae and trochanters and hind femur largely clothed with short white pubescence, and dorsal surfaces of all tibiae with short, stout black setae intermixed with white vestiture; metasoma similar to E. natalensis, except lateral region of T2 anterior to distal band always white and posterior bands on T3-T4 interrupted medially by black and with each half divided in half by a little black vestiture (Fig. 9); T5
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with a dense patch of white posterolaterally and with a silvery-yellowish pseudopy- gidium (male T6 with a more or less continuous white band on distal margin); metasomal venter largely clothed with short white vestiture (male with a little white vestiture on posterolateral regions of $2-$3, $4-$6 black).
Structure. Head: frontal carina similar to E. natalensis, except dorsal region rises more steeply; outer surface of labrum with distal half of median region gently concave, concavity widens progressively towards distal edge, and with one tooth on either side of concavity in about middle of labrum and a subapicolateral tooth on either side of concavity; distal edge of labrum concave medially (Fig. 7); mandible with inner, subapical tooth moderately well developed; maxillary palp with two free segments and about half as long as antennal flagellar segment II. Mesosoma: mesosomal dorsum densely and deeply punctate; axilla spinose and strongly carinae dorsolaterally; (male $4-$5 strongly concave posteromesally and with well-developed subapical fringes of long black hairs that curve forwards mediolongitudinally).
Distribution (Fig. 19). This species is known from Zaire, Malawi, Zimbabwe and the Transvaal and Natal in South Africa.
Discussion. Within this species group E. pyomaeorum can be identified by the black integument, the relatively long, sparse vestiture on the face and pronotal collar, the presence of anterior mediolateral scutal stripes, and lateral interruptions of the distal bands on metasomal terga III-IV.
Material examined. Type material. E. p ygmaeorum, 9 holotype: 'Tshihinda Aug. 26. J.O., Epeolus pyomaeorum Ckll. TYPE, B.M. TYPE HYM. 17B 504', NHML.
Additional material. 109, 6c~: ZAIRE: Secteur Tshiaberimu, Hintumo (lieu-dit), P. Vanschuytbroeck H. Synave, 2c~ MRAC. MALAWI: Mlanje mountain, 11.xi.1982, T. and R. Griswold, 39 BLCU. ZIMBABWE: Chimanimani mountains, 4.iv.1972, R. H. Watmough, 19 NCSA; Troutbeck Inyanga mountains, 23.iv.1985, J. Gusenleit- ner, 39 SC; Vumba mountains, different dates in October, November, January and February, 29, 2c~ SAMC. SOUTH AFRICA: TRANSVAAL: Heidelburg, 23.x. 1966, H. Empey, lc~ NCSA; NATAL: Cathedral Peak Forestry area, 28.55S 29.14E, 10.xi.1981, S. J. van Tonder, C. Kok, lc~ NCSA. SWAZILAND: Mbabane, ii.1954, Osborn, 19 TMSA.
Epeolus alatus Friese
(Fig. 20)
Epeolus alatus Friese, 1922:36 (~ lectotype, TMSA).
The type series of E. alatus comprises two male specimens, one of which is here designated as the lectotype of this species (in TMSA). As this species is only known from the type material, I am unable to describe the female.
Description. Female. Unknown. Male. Similar to E. pygmaeorum in size and colour of integument. Vestiture. Head and mesosoma mostly similar to E. natalensis except as follows:
pronotal collar a little less densely pubescent, similar to E. pyomaeorum, cf. Fig. 8; anterior region of scutum a little more densely pubescent, but without anterior mediolateral scutal stripe; mesopleuron with white vestiture more extensive and with anterior, anteroventral and ventral regions sparsely clothed with short white to yellowish hair; propodeum clothed with short white hair; legs similar to those of E. pygmaeorum; metasoma similar to E. pygmaeorum.
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The genus Epeolus Latreille from subsaharan Africa 721
Structure. Similar to E. pygmaeorum. Distribution (Fig. 20). This species is only known from Harare in Zimbabwe. Discussion. Epeolus alatus is remarkably similar to E. pygmaeorum. These two
species differ mainly in the absence of mediolateral scutal stripes in E. alatus. In this respect E. alatus resembles the E. natalensis species group more closely.
Material examined. Type material. Epeolus alatus, ~ lectotype: 'Salisbury, Mashonaland, April 1906, G. A. K. Marshall, Epeolus alatus Fr. ~', TMSA; paralectotypes: 'Salisbury, Capland, iii.02, Epeolus alatus, 1904 Friese det. ~', ZMHB.
Epeolus amabilis Gerstaecker
(Figs 10, 19)
Epeolus amabilis Gerstaecker, 1869:159 (~ lectotype, ZMHB). Epeolus variegatus (Linnaeus): Smith, 1854:255 [misidentification]; Day, 1979: 75. Epeolus incrassatus Meade-Waldo, 1913: 97; Bischoff, 1923: 599. (~ holotype, NHML). Epeolus amabilis rhodesicus Cockerell, 1921:203 (~ holotype, SAMC), syn. nov.
Smith (1854) recorded E. variegatus from numerous European countries and from the Cape Province. The material from the Cape was studied by Gerstaecker (1869) who found that it had been misidentified by Smith, and that it belonged to a new species which he described as E. amabilis. The original description of E. amabilis included both sexes and the male syntype is here designated as the lectotype of this species.
Bischoff (1923) synonymized amabilis and incrassatus. I have studied the type material of these two species and I agree with the synonymy. Further, the study of the holotype of Cockerell's (1921) E. amabilis rhodesicus together with a large amount of material ofamabilis revealed that the colour in this species shows a continuous range of variation and subspecies cannot be maintained on the basis of these differences. I therefore synonymize rhodesicus and amabilis.
Description. Lengths. Head 1-6-2"3 mm; scutum 1.2-2.2 mm; fore wing 4-7~5-8 mm; body 5"6-9.2 mm.
Colour of integument. Black and usually with ventral and ventrolateral regions of clypeus, entire labrum and mandible orange; antenna blackish-orange; pronotal collar, pronotal lobe, tegula, axilla, scutellum and mesopleuron usually partly orange; anterolateral region of scutum occasionally reddish-orange; legs mostly orange and blackish-orange; lateral regions of T1-T2 partly orange and metasomal venter black, orange and/or blackish-orange.
Vestiture. Face mostly densely pubescent, consisting of long, plumose, white hair, that on lower region of face short and plumose, and on vertex sparse, long, simple and yellowish, as in Fig. 10; gena densely clothed with short, plumose, white vestiture; pronotum sparsely clothed with short, white pubescence; scutum with narrow anterior margin, anterior mediolateral scutal stripe, anterolateral and posterolateral regions densely clothed with long, plumose, white vestiture; remainder of scutum simple and black; tegula with anterior and posterior regions clothed with plumose white vestiture; axilla mostly with simple, black hair, region below spine with plumose, white hair; mesosomal dorsum similar to that of E. pygmaeorum, cf. Fig. 8; scutellum black with anterior margin and posterolateral region white; metanotum mostly white; median region black; mesopleuron, metapleuron and entire ventral surface of mesosoma densely clothed with short, plumose, white vestiture; lateral region of propodeum with short, plumose, white vestiture and that on posterolateral region long and plumose; posterior surface of propodeum largely naked; legs and metasoma similar to
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722 C.D. Eardley
E. pygmaeorum, except short, stout setae on dorsal surface of tibiae orangish; T5 largely clothed with dense white vestitute and with a silvery-yellowish pseudopygidium.
Structure. Similar to E. pygmaeorum. Distribution (Fig. 19). This species occurs in eastern and southern Africa. Discussion. This species is most closely related to E. pygmaeorum and E. alatus, but
can be easily identified by the characters given in the key. The only known host plant is Grielum, but as E. amabilis is a cleptoparasite it is
unlikely to be an important pollinator. A male and a female specimen from Paleisheuwel in the western Cape Province
(detailed below) are here assigned to E. amabilis with some reservation. These two specimens do not differ in structure from the type and other material studied, but they are more extensively marked with orange as follows: entire clypeus and part of lateral region of pronotum of female orange; both sexes with entire lateral region of scutum, and almost entire axilla, scutellum and mesopleuron orangish; posterolateral regions of propodeum, almost entire legs and most of metasomal dorsum orange, entire median region of T3 and part of median regions T4-T5 black. The taxonomic significance of this colour difference is not clear, and more material from that region will have to be studied before the true identity of this material can be ascertained.
Material examined. Type material. Epeolus amabilis, 3 lectotype: 'Capland, Krebs S., amabilis Gerst. 3 Ep. variegatus var. Smith', ZMHB.
E. incrassatus, 3 holotype: 'Basutoland, R. Crowshay, 1902~a4, Epeolus incrassatus Type 3 G. Meade-Waldo, det. B.M. Type Hym 17B 511', NHML.
holotype E. amabilis rhodesicus: 'S. Rhodesia, Salisbury, D. Dodds, Epeolus amabilis rhodesicus Ckll. TYPE', SAMC.
Additional material. 92~, 343: ZIMBABWE: Inyanga, 7.ii.1940, 15 ̀ SAMC. NAMIBIA: Barby 26, Bethanie, 2516DC, 2-7.x.1972, 1 ~ SMWH; Uguchab River, near Aurusberg, 27.32S 16.11E, 22.iv.1988, C. D. Eardley, 2~ NCSA. SOUTH AFRICA: TRANSVAAL: Wolkberg, 21 km SW Tzaneen, 7.iii.1976, R. H. Watmough, 19 NCSA; Graskop, 6.v.1964, H. Empey, 13 NCSA; Johannesburg, iv.1916, G. Kobrow, 13 TMSA; Florida, xii.1918, R. W. Tucker, 1~ SAMC; ORANGE FREE STATE: Harrismith, i.1978, C. Eardley, 1~ NCSA; NATAL: Garden Castle State Forest, 7.ii.1986, V. B. Whitehead, 1~ SAMC; National Park (= Royal Natal National Park), 3-15.iii.1932, L. Ogilvie, 13 NHML; CAPE PROVINCE: Katberg, xii.1932 and i-ii.1933, R. E. Turner, 7~, 75 ̀NHML 15 ̀SAMC; Cookhouse, iii.1954, 1~ SAMC; Middelburg, xi. 1935, 1 ~, 15 ̀SAMC; Grahamstown, Dr Tenther, 1 ~ TMSA; Resolution, Grahamstown, i-iv.1928, Miss Walton, 1~ SAMC; Sunday River, 27.xii.1897, Dr Brauns, 1~ TMSA; Algoa Bay, 11.iii.1896, Dr Brauns, 15 ̀ TMSA; Graaff-Reinet, 13.iii.1969, L. C. Starke, 15 ̀NCSA; WiUowmore, different dates from December to March, Dr Brauns, 24~, 48' TMSA; Fort Beaufort, Umdala, iii.1954, 1~, 43 SAMC; Rust en Vrede, Oudtshoorn district, x.1951, 13 SAMC; Great Brak River, Mossel Bay, ii.1960, 13 SAMC; Mossel Bay, xii.1921 and i.1922, R. E. Turner, 2~, 13 NHML; Richmond district, xii.1939 and iii.1931, 4~, 13 SAMC; Murraysburg, x-xi.1935, 5~, 33 SAMC; Merweville, i.1959, H. Zinn, 1~ SAMC; Merweville district, i-ii.1947, H. Zinn, 1~ SAMC; Coup, Lainsburg district, ix.1937, 2~ SAMC; Oukloof, Beaufort West, i.1949, H. Zinn A. Hesse, 1~ SAMC; Molteno Pass, 32.16S 22.34E, 14.xi.1988, C. D. Eardley, 1~ NCSA; Goedehoop, Heidelberg district, x.1951, 19 SAMC; Theekloof, Fraserburg district, xi.1935, 2~ SAMC; 30 km E Touwsrivier, on road to Hondewater, xii.1962, 3~ SAMC; near Touwsrivier, between Ladismith and Montagu, x.1937, 3~ SAMC; Matjesfontein, 14-27.xi.1928, R. E. Turner, 13 NHML; Pearly Beach,
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The genus Epeolus Latreille from subsaharan Africa 723
Bredasdorp, xii.1958, lq SAMC; Stellenbosch, xi.1925, H. Brauns, 13 ̀ TMSA; Kommetjie, 3418AB, 14.xi. 1976, lq SAMC; Gydo Pass, 15 km N Prince Albert Hamlet, 33.08S 19.20E, 19.xi.1984, C. D. Eardley, 13 ̀NCSA; Paleisheuwel, xi.1948, lq, 13' SAMC; Bot River, 11.xi.1933, L. Ogilvie, lq NHML; Ceres, xii.1920 and xi.1924, R. E. Turner, 2~, 13 ̀NHML; Upper Sources, Olifants river, near Ceres, xii.1949, 1~ SAMC; Calvinia, 13.xii, J. Ogilvie, 13 ̀SAMC; Kamieskroon, 26.v.1978, V. B. Whitehead, on Grielum sp., lq SAMC; Oorlogskloof, Nieuwoudtville, 27.x.1"978, V. B. Whitehead, lq SAMC; Hester Malan Nature Reserve, near Springbok, 2917DB, 18.x.1985, M. Struck, 13 ̀NCSA; Henkries, Bushmanland, Lightfoot, 1~ SAMC; Between Springbok and Pella, Aggenys or Bushmanland, x.1939, 3~ SAMC; Bowesdorp, ix.1941, 1~ SAMC; 10km N Copperton, 29.52S 22.15E, 6.v.1985, M. L. Penrith, V. B. Whitehead, M. Macpherson, 2~ SAMC; Swartkops, 30.08S 20.59E, 1.v.1985, M. L. Penrith, V. B. Whitehead, M. Macpherson, 2~ SAMC.
Epeolus pubescence Cockerell
(Figs 12, 20)
Epeolus pubescence Cockerell, 1937:159 (c~ holotype, NHML).
Description. Female. Unknown. Male. Similar to E. amabilis except as follows: integument of T1-T2 mostly orange;
entire distal margins of T1-T6 clothed with short white, appressed, vestiture, except for that on T3 which is narrowly interrupted with a little black vestiture medially (Fig. 12); metasomal dorsum otherwise similar to that of E. cestus; distal margin of $4-$5 strongly concave posteromesally and with well-developed subapical fringes of long blackish-orange hairs that curve forwards mediolongitudinaUy.
Distribution (Fig. 20). Known from Swakopmund in Namibia. Discussion. This species is more closely related to E. pyomaeorum, E. amabilis and
E. cestus than to the other species in this group. Its most conspicuous diagnostic features within the group are the colour of the integument in combination with the continuous or very narrowly interrupted distal bands on the metasomal terga.
Material examined. Type material. Epeolus pubescence, 3 ̀holotype: 'S.W. Africa, R. E. Turner, Brit. Mus. 1928-249, Swakopmund, 2-4.iv.1928, Epeolus pubescence Ckll. TYPE, B.M. TYPE HYM. 17B 508', NHML.
Epeolus fulviventris Bischoff
(Figs 11, 20)
Epeolusfulviventris Bischoff, 1923: 600; Friese, 1925: 503-504, (~ holotype, ZMHB).
This species was described as new by both Bischoff(1923) and Friese (1925), and it appears that they based their descriptions on the same material.
Description. Size, colour and relative density of vestiture on head, mesosoma and metasoma similar to E. rufothoracicus except as follows: integument of mesopleuron mostly black, with a little reddish-orange medially; vestiture on T1 sparse and pale yellowish-white on subvertical, anterior and extreme lateral surfaces, distal margin with a densely pubescent pale yellowish-white band that is broadly interrupted medially; T2 with a broadly interrupted distal band of pale yellowish vestiture; T3-T4 (T3-T5 in males) with more or less continuous pale yellowish-white distal bands; T5 sparsely clothed with pale yellowish vestiture, pseudopygidium silvery-yellowish;
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I R
C. D. Eardley
17
/
FIGS 12-18. Epeolus spp. 12, E. pubescence, dorsolateral view of male metasoma; 13, E. cestus, dorsolateral view of female metasoma. 14-16. E. caffer, female: 14, external view oflabrum; 15, dorsal view of mesosoma; 16, dorsolateral view of metasoma. 17, 18. E.friesei, female: 17, external view of labrum; 18, dorsal view of mesosoma.
S1-$5 sparsely clothed with short orangish-white hairs (male with $4-$5 with orange subapical fringes).
Structure. Similar to that of E. pygmaeorum except as follows: labrum with a weakly developed posteromedian concavity, and with three median tubercles, one sub- apicomedian tubercle and two apicolateral tubercles on each side of apicomedian tubercle, as in Fig. 11.
Distribution (Fig. 20). Epeolus fulviventris is known only from Natal and the Transkei, South Africa.
Discussion. This species is placed in the E. pygmaeorum species group because it resembles the species of that group in the structure of the axilla. But it differs from the other species in the group in that the pronotal collar is very densely pubescent and the scutum does not have any anterior mediolateral stripes.
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The genus Epeolus Latreille from subsaharan Africa 725
Material examined. Type material. Epeolus fulviventris, 3 ̀ holotype: 'Mfongosi, Zulu L., WE Jones, Epeolusfulviventris 1019 Friese det. 3 ,̀ Epeolusfulviventris 3 ̀Fr. i.1. Bisch. Type', ZMHB.
Additional material. 1 ~: SOUTH AFRICA: NATAL: Braemar, 23.iii. 1927, L. Bevis, DMSA.
Epeolus cestus sp. nov.
(Figs 13, 20)
This name is taken from the Greek for girdle, which refers to the bands of white vestiture on the metasomal dorsum.
Description. Lengths. Head 1.9-2-2 mm; scutum 1-7-2.1 mm; fore wing 5.7-6.5 mm; body 6.6-9.2 mm.
Colour of integument. Mostly reddish-orange, vertex and paraocular area of head and mediolongitudinal region of scutum black (male with scutum largely black, narrow lateral margins reddish); posterior surface of propodeum sometimes partly black and dorsal surface of metasoma sometimes with a blackish tinge.
Vestiture. Head and mesosoma, including legs, generally similar to E. pygmaeorum, most of scutal vestiture has a yellowish tinge, except densely pubescent anterior mediolateral scutal stripe, lateral and posterolateral regions which are white, and short, stout setae on dorsal surface of tibiae which are orangish; T1 with subvertical anterior, lateral and posterior regions mostly white to pale yellowish, posterior band narrowly interrupted medially (males sometimes with this band continuous); T2-T4 with white to pale yellowish distal bands and sometimes with a little white vestiture laterally, T2 with distal band occasionally narrowly interrupted medially and T3-T4 with distal bands very occasionally narrowly interrupted medially, but sometimes interrupted in about middle of each half and continuous medially; T5 sparsely clothed with very short white vestiture and with a silvery-yellowish pseudopygidium (male T5-T6 each with a more or less continuous pale yellowish band on distal margin); remainder ofmetasomal terga clothed with short reddish hairs, sometimes reddish-black or black medially; metasomal terga as in Fig. 13; metasomal venter largely white (male with $4-$6 orange, including subapical fringes on $4-$5).
Structure. Similar to that of E. pygmaeorum. Distribution (Fig. 20). Epeolus cestus is only known from Zimbabwe and the
Transvaal in South Africa. Discussion. This species is unusual in that the female scutum is largely red with a
black mediolongitudinal stripe, in strong contrast to the male in which the scutum is mostly black, and the distal bands on terga three and four are sometimes interrupted laterally and usually continuous medially.
Material examined. Type material. 7~, 63`: ZIMBABWE: Bulawayo, x-xi.1923, R. Stevenson, ~ holotype 2~, 13 ̀ paratypes TMSA 2~, 13 ̀ paratypes NCSA 2~, 13' paratypes SAMC; Hope Fountain, 8.x.1923, N. Jones, 13 ̀paratype TMSA; Druid Mine, Filabusi, 20.ix.1923, 13 ̀paratype SAMC. SOUTH AFRICA: TRANSVAAL: Pretoria, 2.x.1933, G. van Son, 13 ̀paratype NCSA.
The Epeolus caller species group
Epeolus caffer and E. friesei constitute this species group. In contrast to the other two groups the structure of the axilla is diagnostic of the species and not the group. The
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726 C.D. Eardley
remarkable similarity in cotour, vestiture and structure, and in particular the flat to weakly tuberculate labrum, evenly rounded scuteUum and long black subapical fringes on the fourth and fifth metasomal sterna of the male were used to formulate the group.
I am of the opinion that the general similarity between these two species is indicative of close relationship and that the loss of the spine on the axilla of E.friesei is a derived character.
Epeolus cafl'er (Lepeletier), comb. nov.
(Figs 14-16, 19)
Crocisa caffra Lepeletier, 1841: 455; Friese, 1905: 173; Meyer, 1921: 132-133; Cockerell, 1935:89 (~ holotype, MNHN).
Epeolus militaris Gerstaecker, 1869:160 (~ holotype, ZMHB), syn. nov. Epeolus pectinatus Friese, 1922: 35; Bischoff, 1923: 598. (~ lectotype, ZMHB), syn. nov. Epeolus albohirtus Friese, 1922:35-36 (~' holotype, ZMHB), syn. nov.
Lepeletier (1841) described this species in the genus Crocisa (= Thyreus) and I here transfer it to Epeolus. Epeolus caffer was synonymized with Thyreus albomaculatus, by Friese (1905), and with Thyreus plumifer, by Meyer (1921), neither of which are congeneric with this species (Eardley, 199l). The holotype of E. caffer is fragile and was, therefore, not dissected. Nevertheless, I am confident of the identity of this species. The head of the holotype has been lost and what appears to be the head of a wasp has been glued in its place.
"Epeolus militaris was described by Gerstaecker (1869), who was apparently not aware that this species had been described. His description of E. militaris dealt primarily with colour. Friese (1922), in the descriptions of E. pectinatus and E. albohirtus, separated these two species primarily on size and the colour of the vestiture on the metasomal renter. I have studied much material of E. caffer and found it to be variable in many respects. The differences between the types of E. miIitaris, E. pectinatus and E. albohirtus, which I have also studied, fall well within the variation of this species. Epeolus militaris, E. pectinatus and E. albohirtus have therefore been synonymized with E. caffer.
Description. Lengths. Head 1.7-2-1 mm; scutum 1.4-2-1 mm; fore wing 5.1-6-4 mm; body 6.3-8"1 mm.
Colour of integument. Head black, except antennal flagellum and bulla of labrum orangish; mesosoma black, except tegula and all tibiae and tarsi orange, trochanters and femora sometimes orangish; metasoma generally with first two to three segments orange or orangish and with distal region black.
Vestiture. Head either completely clothed with sparse, long, black, simple vestiture, or with lower region of face partly or completely clothed with short, white, plumose pubescence (male with head generally sparsely pubescent above and densely pubescent below antennal sockets, upper region of head ranges from mostly black to mostly white and region of face below antennal sockets always mostly white); mesoma sparsely clothed with long vestiture, ranging from completely black to mostly white, usually predominantly white anteriorly and black posteriorly; legs with basal three segments mostly black and distal segments largely white; dorsal surfaces of tibiae and tarsi sparsely clothed with short, white vestiture which is intermixed with long, thick, black to orangish setae, except hind tibia has very few thick setae and distal end of dorsal
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The genus Epeolus Latreille from subsaharan Africa 727
surface of middle tibia has a tuft of orange to blackish-orange hair (male with legs clothed with fine white vestiture, dorsal surfaces of tibiae with a few long, thick, orange spines intermixed with white vestiture); ventral surfaces of tarsi orange; metasomal dorsum occasionally completely clothed with short, sparse, black hairs, usually as in Fig. 16 with a little white pubescence posterolaterally on T1, T2-T4 with broad, white distal margins that are broadly interrupted medially and T5 black with a silvery-white to silvery-yellowish pseudopygidium (male with T3-T5 similar to T3-T4 of female, except distal bands shorter, they do not extend to posterolateral region of each tergum); S1-$4 with several long, black setae and with a little white posterolaterally, $5 completely black (male with $4-$6 blackish).
Structure. Head: frontal carina moderately developed, lateral sides gradually raised so that region between antennal sockets together with virtually entire supra-antennal area strongly raised; labrum flattish, sometimes with a small to very small tubercle on each side of median axis near distal end (Fig. 14), or with a weakly developed mediolongitudinal carina on distal half; mandible without an inner tooth; maxillary palp two-segmented and about one-third as long as antennal flagellar segment II. Mesosoma: mesosomal dorsum densely punctate; axilla spinose, spine not carinate and projects posteriorly in a more or less dorsolateral direction (Fig. 15); scutellum evenly rounded and not concave mediolongitudinally (male $4-$5 with long, black subapical fringes).
Distribution (Fig. 20). This species is apparently endemic to the Cape Province, South Africa. Its range incorporates several different biotypes, such as the Karoo, Namaqualand, the south-western Cape Province and the southern Cape coast.
Discussion. Epeolus caffer can be easily identified by the peculiar shape of the axilla. There is a single record of a male visiting flowers of Hermannia sp. Material examined. Type material. Epeolus caffer, ~ holotype: 'Afrique, Cafreria,
Delalande, Cafforue [? hand writing illegible], Crocisa caffra Lepeletier, Type!', MNHN.
Epeolus militaris, ~ holotype: 'Capland, S. Windt., mititaris Gerst. c~', ZMHB. Epeolus pectinatus, ~ lectotype: 'Saldanha B., Sept. 1912, L.P., Epeolus pectinatus
1904 Friese det. ~', ZMHB; ~ paralectotype E. pectinatus: 'Capland, Epeolus pectinatus 1904 Friese det. ~', ZMHB.
Epeolus albohirtus, ~ holotype: 'Capland, Cradock, 1902, Epeolus albohirtus, 1910 Friese det. c~', ZMHB.
Additional material. 36~, 13~: SOUTH AFRICA: CAPE PROVINCE: Algoa Bay, 1.xi.1897, Dr Brauns, 2~, lz~ TMSA; Mossel Bay, viii.1932, R. E. Turner, 1~ SAMC 3d' NHML; Willowmore, 1.xi.1919, Dr Brauns, ld' TMSA; Rust en Vrede, Oudtshoorn district, x.1951, 1~ SAMC; Skurfkop, 3118AB, 18.viii.1983, V. B. Whitehead, on Hermannia sp. white, ld' SAMC; Stellenbosch, Joosterbergkloof, 3318DD, 7.x.1989, V. B. Whitehead, 1~ SAMC; Malmesburg, 14.x.1923, lq) SAMC; Lambert's Bay, 32.04S 18.20E, 4.x.1974, R. H. Watmough, 1~ NCSA; Elands Bay, 3218AD, 26.ix.1978, V. B. Whitehead, 1~ SAMC; Augustfontein, ix.1947, 1~, ld' SAMC; Biedouw Valley, Clanwilliam district, 32.08S 19.14E, 5-7.ix.1987, C. D. Eardley, 1~ NCSA; Ramskop, Clanwilliam, 24.viii.1985, V. B. Whitehead, 25 SAMC; Ramskop cam_p, Clanwilliam, 3218BB, 20.viii.1984, V. B. Whitehead, 4~ SAMC; 10km N Clanwilliam, 3218BB, 30.vii.1983, V. B. Whitehead, 1~ SAMC; Vanrhynsdorp, viii.1927, Dr Brauns, 7~, lc~ TMSA 2~, 1~ NCSA; Vanrhynsdorp, vii-viii.1927, G. van Son, 5~, 1~ TMSA 2~, 1~ NCSA; near Kamieskroon on Wallekraal road, 3017BA, 3.viii.1988, V. B. Whitehead, 1~ SAMC.
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728 C .D. Eardley
t~-_ I , i ~'- i I 1 1 ~ / ..... \_ t
o ~ ~ - ~ ' ::--' :. ,,,_ .... :~ ........
%. ....... °
E. n a t a l e n s i s . """n'"" l E. p y g m a e o r u m •
E. a m a b i l u s +
FIG. 19. Distribution of E. natalensis, E. kristenseni, E. pygmaeorum and E. amabilus.
Epeolus friesei Brauns
(Figs 17, 18, 20)
Epeolusfriesei Brauns, 1903:362-366 (3' iectotype, TMSA). Epeolus karroensis Brauns, 1909:10 (~' holotype, TMSA), syn. nov.
The holotype of E. karroensis is in my opinion a melanized specimen of E.friesei. This specimen differs from the other material of E.friesei that I have studied mainly in that the first two segments of the metasoma are reddish-black and not orange.
Description. Similar to E. caffer except as follows: metasoma generally more variable, colour of integument of T l -T2 occasionally reddish-black; posterior margin of T1 may be devoid of any white vestiture, have a little white posterolaterally or have a medially interrupted distal band of white vestiture; white distal bands on T2-T3 range from continuous to broadly interrupted medially; T4 with distal band ranging from continuous to completely absent; labrum flattish and devoid of carina and tubercles (Fig. 17); axilla simple in structure (Fig. 18).
Distribution (Fig. 20). This species is apparently endemic to the Karoo and Namaqualand in the Cape Province of South Africa.
Discussion. Epeolusfriesei is unique among the subsaharan species of this genus in that it does not have a spinose axilla. The general structure and colour of this species, however, suggest that it is most closely related to E. caffer.
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The genus Epeolus Latreille from subsaharan Africa 729
r ~
%
i
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3i
FIG. 20. Distribution of E. rufothoracicus, E. fulviventris, E. alatus, E. pubescence, E. cestus, E; caffer and E. friesei.
Material examined. Type material. Epeolus friesei, ~ lectotype: 'Willowmore, Capland, Dr Brauns, 15.9.1902, Type ~ Epeolusfriesei Brauns', TMSA; 1~ paralecto- type E. friesei: data as in holotype except ~ specimen, TMSA; 33' paralectotypes E. friesei: data as in holotype except dates which are as follows, 20.8.1906, 18.8.1906 and 20.8.1906, TMSA.
Epeolus karroensis, ~ holotype: 'Capland, Willowmore, 18.8.1906, Dr Brauns, Type Epeolus karroensis Brauns', TMSA.
Additional material. 4~, 1~: SOUTH AFRICA: CAPE PROVINCE: Lambert's Bay, 32.04S 18.20E, 4.x.1974, R. H. Watmough, from nest to sandy soil, 2~? NCSA; Augustfontein, ix.1947, 1~ SAMC; Mecrhofskasteel, Nuwerus, 30.viii.1984, M. Macpherson, lc~ SAMC.
Acknowledgements I thank Dr G. L. Prinsloo of the Plant Protection Research Institute, Pretoria, for
reading the manuscript, and Mrs M. Verster, of the same institute, for her illustrations (Figs 1-18). The Department of Environment Affairs and the Transvaal Provincial Administration, Nature Conservation Division are thanked for permission to collect in areas under their control. I express my gratitude to the following persons and institutions for the loan of their material: Dr E. De Coninck, MRAC; Mr G. Else, NHML; Dr T. L. Griswold, BLCU; Mr J. Irish, SMWH; Dr F. Koch, ZMHB; Mr C. D. Quickelberg, DMSA; Mr M. Schwarz, SC; Mr R. Snelling, LACM; Dr R. Toms, TMSA; Dr J. C. Weulisse, MNHN; Dr V. B. Whitehead, SAMC.
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Checklist of species The arrangement under each entry is as follows: species or subspecies name, author,
year in which described, original generic placement if different from current one or species name in case of subspecies in square brackets and synonyms in round parentheses. Invalid names are in italics.
alatus Friese, 1922. albohirtus Friese, 1922 (see caffer). amabilis Gerstaecker, 1869 (variegatus, incrassatus, rhodesicus). bidens Bischoff, 1923 (see natalensis). caffer (Lepeletier), 1841 [Crocisa] (militaris, pectinatus, albohirtus). cestus sp. nov. coelostoma Bischoff, 1923 (see natalensis). foveilabris Bischoff, 1923 (see natalensis). friesei Brauns, 1903 (karroensis). fulviventris Bischoff, 1923. glyptochilus Bischoff, 1923 (see natalensis). haemataspis Cockerell, 1937 (see rufothoracicus). incrassatus Meade-Waldo, 1913 (see amabilis). karroensis Brauns, 1909 (see friesei). kristenseni Friese, 1915. militaris Gerstaecker, 1869 (see caffer). natalensis Smith, 1879 (coelostoma, bidens, foveilabris, glyptochilus). pectinatus Friese, 1922 (see caffer). pubescence Cockerell, 1937. pygmaeorum Cockerell, 1932. rhodesicus Cockerell, 1921 [amabilis] (see amabilis). roseatus Cockerell, 1937 (see rufothoracicus). rufothoracicus Bischoff, 1923 (haemataspis, roseatus). variegatus (Linnaeus), 1758 [Apis] (see amabilis, misidentification).
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