Taxonomy of Afrotropical and West Palaearctic ants of the ponerine genus Hypoponera Santschi (Hymenoptera: Formicidae).
Author
Bolton, B.
Author
Fisher, B. L.
text
Zootaxa
2011
2843
1
118
http://antbase.org/ants/publications/23490/23490.pdf
journal article
23490
Hypoponera punctatissima (Roger)
(Figs 97
-
102)
Ponera punctatissima Roger
, 1859: 246, pl. 7, fig. 7. Syntype workers and queen, POLAND: Rauden (now Rudy, Opole Prov.) and GERMANY: Berlin, in hothouses (Roger) (MNHN) [examined]. [Combination in
Hypoponera
: Taylor, 1967: 12.] (See note 1.)
Ponera androgyna Roger
, 1859: 246 (footnote). Syntype ergatoid males (not workers), POLAND: Rauden (now Rudy, Opole Prov.) (Roger) (ZMHB) [not seen]. [Junior synonym of
punctatissima
: Emery & Forel, 1879: 455; Seifert, 2003: 69.]
Ponera tarda Charsley
, 1877: 162. Syntype workers and queen, GREAT BRITAIN: Oxford, 1877 (R.S. Charsley) (OXUM) [not seen]. [Junior synonym of
punctatissima
: Dalla Torre, 1893: 41; Seifert, 2003: 69.]
Ponera punctatissima r. jugata Forel
, 1892: 251. Holotype queen, MADAGASCAR: Prov.
d'Imerina
(Sikora) (MHNG) [not seen]. [Raised to species: Dalla Torre, 1893: 39. Reverted to subspecies of
punctatissima
: Emery, 1899: 268. Combination in
Hypoponera
: Bolton, 1995: 215. Junior synonym of
punctatissima
: Seifert, 2003: 69.]
Ponera ergatandria Forel
, 1893: 365. Syntype workers, queens and ergatoid male, SAINT VINCENT I. (Antilles): 41b (H.H. Smith). (MHNG, BMNH) [BMNH ergatoid male examined]. [Junior synonym of
punctatissima
: Taylor, 1968: 65. Revived status as species and combination in
Hypoponera
: Kempf, 1972: 122. Reverted to junior synonym of
punctatissima
: Smith, D.R. 1979: 1343, here confirmed.] (See note 2).
Ponera kalakauae Forel
, 1899: 115. Syntype worker-queen intercaste (not worker) and queen, HAWAIIAN IS: Kauai I., Lahue, 2000 ft, vii.1896 (R.C.L. Perkins) (intercaste), and Honolulu (Oahu I.), xi.1896 (R.C.L. Perkins) (queen) (BMNH) [examined]. [Junior synonym of
gleadowi
: Wilson, 1958: 328. Junior synonym of
punctatissima
: Wilson & Taylor, 1967: 29 (in text).] (See note 3.)
Ponera
punctatissima subsp. schauinslandi Emery
, 1899: 439. Syntype queens, HAWAIIAN IS: Laysan I. (now Laycan I.) (Schauinsland) (MSNG) [not seen] Syn. rev. [Combination in
Hypoponera
and junior synonym of
punctatissima
: Wilson & Taylor, 1967: 28. Revived from synonymy and raised to species: Seifert, 2003: 69.]
Ponera ergatandria subsp. bondroiti Forel
, 1911: 285. Syntype workers, queen and ergatoid male, BELGIUM: Bruxelles, Jardin Bot., xi.1909, 5.xi.1909, 15.xi.1909 (Bondroit) (MHNG) [not seen]
Syn. n.
[Raised to species: Santschi, 1937b: 364. Combination in
Hypoponera
: Onoyama, 1989: 2. Previously junior synonym of
schauinslandi
: Seifert, 2003: 69.]
Ponera dulcis var. aemula Santschi
, 1911: 351. Lectotype and paralectotype workers (designated by Seifert, 2003: 68), TANZANIA: Kilimandjaro, zone des cultures, Kiboscho, 1400 m, 1904 (Ch. Alluaud) (NHMB) [examined]
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 213. Previously junior synonym of
schauinslandi
: Seifert, 2003: 69.] (See note 4.)
Ponera ergatandria st. cognata Santschi
, 1912: 153. Syntype workers, ANGOLA: Benguela, Cucula (J. Cruchet) (NHMB) [examined].
Syn. n.
[Unresolved junior primary homonym of
Ponera cognata Emery
, 1896: 56 (now in
Pachycondyla
). Raised to species: Bernard, 1953: 202. Combination in
Hypoponera
: Bolton, 1995: 213.] (See note 5.)
Ponera ragusai var. sordida Santschi
, 1914a: 54. Holotype worker, KENYA:
region
cotiere
, Shimoni, st. no. 9, xi.1911 (Alluaud& Jeannel) (NHMB) [examined].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 216.]
Ponera incisa Santschi
, 1914b: 320, fig. 8. Holotype ergatoid male (not worker), NIGERIA: Lagos (F. Silvestri) (not in NHMB, presumably in DEUN) [not seen].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 214.] (See note 6).
Ponera sulcatinasis r. durbanensis Forel
, 1914: 213. Syntype workers, SOUTH AFRICA: Natal, Durban, 16.i.1914, no. 323 (G. Arnold) (MHNG) [examined].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 214.]
Ponera ergatandria r. petri Forel
, 1916: 397. Holotype worker-queen intercaste (not worker), DEMOCRATIC REPUBLIC OF CONGO: St Gabriel (Kohl) (MHNG) [examined].
Syn. n.
[Raised to species: Santschi, 1938: 78. Combination in Hypoponera: Bolton, 1995: 216.]
Ponera brevis Santschi
, 1921a: 113, fig. 1a-c. Holotype worker, BENIN (Dahomey on label): Porto Novo (Le Moult) (NHMB) [examined].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 213.] (See note 7.)
Ponera punctatissima var. exacta Santschi
, 1923: 134. Syntype workers, TUNISIA: Hammamet (Santschi) (NHMB, BMNH) [BMNH syntype examined]. [Junior synonym of
punctatissima
: Atanassov & Dlussky, 1992: 71; Seifert, 2003: 69.]
Ponera mina Wheeler
, W.M. 1927b: 131, fig. 2. Syntype workers, queens and ergatoid male, AUSTRALIA: Norfolk I., 1915 (A.M. Lea) (MCZC and possibly SAMA) [not seen]. [Junior synonym of
gleadowi
: Wilson, 1958: 329. Revived from synonymy and combination in
Hypoponera
: Taylor & Brown, D.R. 1985: 31. Junior synonym of
punctatissima
: Wilson & Taylor, 1967: 29 (in text); Taylor, 1987: 30.]
Ponera argonautorum Arnol'di
, 1932: 66, figs. 6, 7. Holotype queen, RUSSIA: Black Sea Coast, Anapa, 26.viii.1925, Nr. 422 (K.V.
Arnol'di
) (probably in ZMUM) [not seen].
Syn. n.
(provisional). [Combination in
Hypoponera
: Bolton, 1995b: 213.] (See note 8.)
Ponera mumfordi Wheeler
, W.M. 1933: 141. Syntype workers, FRENCH POLYNESIA: Marquesas Is, Uapou; Kohepu (Tekohepu) summit, alt. 3000 ft, 3.xi.1931 (LeBronnec) (probably in MCZC) [not seen]. [Junior synonym of
gleadowi
: Wilson, 1958: 328. Junior synonym of
punctatissima
: Wilson & Taylor, 1967: 29 (in text).]
Ponera mesoepinotalis Weber
, 1942: 44, fig. 4. Holotype worker, SUDAN: Imatong Mts, 2.viii.1939, 6400 ft, no. 1395 (N.A. We b e r) (not in MCZC, presumed lost)
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 215.] (See note 9.)
Ponera breviceps Bernard
, 1953: 202, fig. 3A. LECTOTYPE dealate queen (by present designation) and two paralectotype dealate queens, GUINEA: Nimba (Lamotte) (MNHN) [examined].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 213.] (See note 10.)
Ponera ursoidea Bernard
, 1953: 203, fig. 3C. LECTOTYPE worker-queen intercaste (not worker) (by present designation), GUINEA:
Crete
de Nion, 1300 m. (no
collector's
name, probably Lamotte) (MNHN) [examined].
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 216.] (See note 11.)
Ponera sulcitana Stefani
, 1970: 1, figs. 1
-
12. Holotype worker; paratype workers, queens and ergatoid male, ITALY: Sardinia, Grotta dei Fiori presso Carbonia (A. Serra) (IZUC) [not seen]
Syn. n.
[Combination in
Hypoponera
: Bolton, 1995: 216.] (See note 12.)
NOTES.
1 Seifert (2003) refers to each of the
punctatissima
type-specimens he examined as a paralectotype. While there seems no doubt that the material originated in
Roger's
collection and is type-material, no trace of a lectotype designation can be found. Therefore the two MNHN specimens examined here, and those recorded by Seifert as present in ZMHB and DEIC, revert to syntype status.
2 The BMNH ergatoid male has the data:
"41b"
and
"
Ponera ergatandria Forel
, male type, St. Vincent
"
. This is certainly a minor ergatoid male of
punctatissima
(yellow, eyeless and with 12 antennal segments). In the publication, Forel (1893) gives the following localities for
ergatandria
, all of which are syntypes. 41a: Bowwood Valley, nr Kingston, 800 ft, 15.x.; 41b: Islet fronting
Chateaubelais
Bay, 31.x.; 41c: Richmond Estate, 31.x.; 41d: Villa Estate, south end of island, 14.x.
3 Wilson (1958: 328-329) recorded the names
decipiens
,
kalakauae
,
mina
and
mumfordi
as junior synonyms of
gleadowi
. In the text of Wilson & Taylor (1967: 29) all these names were transferred to the synonymy of punctatissima
,
but
decipiens
was later reverted to the synonymy of
gleadowi
by Onoyama (1989) following his examination of its holotype. These transfers of synonymy were inadvertently omitted from the Bolton (1995) catalogue.
4 The short history of the name
aemula
amply illustrates the confused nature of the early taxonomy of Afrotropical
Hypoponera
. When introduced,
aemula
was described as an infraspecific form of
dulcis
, now in
H. dulcis
group. At its next appearance (Santschi, 1914a: 53)
aemula
had become an infrasubspecific name attached to
coarctata subsp. boerorum
. The first of these names is now in a separate genus (
Ponera
) and
boerorum
is a member of the
H. boerorum
group. The first critical examination of the syntypes of
aemula
, by Seifert (2003), placed it firmly in what is now the
H. punctatissima
group, which is confirmed here.
5 In the original description of
cognata, Santschi
mentions three workers, with the data recorded above. Of these, two are normally mounted and the head of the third is mounted on a small perspex slide, without any remnants of its body. In NHMB are two more workers, also labelled as
"types"
but from
"
Benguela, Caconda (Cruchet)
"
. These are identical to the Cucula specimens, but because they were not mentioned in the original description they have no type-status.
6 The holotype of
incisa
is not among
Santschi's
collection in NHMB. If it still exists it is probably in the Silvestri collection at Portici (DEUN), which is not currently available. The original description gives the holotype as a worker, but it seems most likely that it is really an ergatoid male of
punctatissima
. Characters noted or figured by Santschi that point towards this identification include: extremely short scapes (from the drawing SI ca 65; measured ergatoid males SI 68
-
72), coupled with worker-like mandibles; large, rectangular, worker-like head; metanotal groove sharp and deep; gastral segments somewhat enlarged. Unfortunately the apical gastral segments are missing from
Santschi's
drawing, which may imply that they are absent from the specimen. But the description and figure suggest most strongly that the holotype is an ergatoid male of
punctatissima
. It is also interesting to note that on the same page Santschi (1914b: 320) recorded a queen of
punctatissima
, collected from the same locality as
incisa
.
7 The holotype of
brevis
is in reasonable condition but is flat-mounted on card and much of the lower and posterior petiole, and the base of the gaster, are obscured by glue. Several characters noted by Santschi in the original description and figures are misleading. For instance, the metanotal groove is not shown in his fig. 1c, but in the text it is stated as present and it is conspicuous on the holotype. The propodeum is not marginate between the declivity and the sides. The lower petiole is obscured not only by glue but by the position of the legs. However, the node in profile appears to taper slightly towards the apex and is not
"
as thick at the summit as at the base
"
.
8
Ponera argonautorum
was provisionally described as new by
Arnol'di
, based on a single alate queen discovered flying in a garden at Anapa, on the Russian north-east coast of the Black Sea. Its holotype has not been examined but it seems obvious, from the description and figures, that this is a queen of
punctatissima
. It is provisionally synonymised here to draw attention to our opinion, until the holotype can be examined in detail.
9 Stefan Cover (MCZC) informs us that although the holotype of
mesoepinotalis
is noted in the MCZC typeregister, the specimen has disappeared from the collection and cannot be found.
Weber's
description and figure are sufficient to place the name with considerable confidence as a junior synonym of
punctatissima
.
10
Bernard's
syntype series of
breviceps
consisted of four specimens mounted on two pins. The upper specimen on the first pin is a dealate queen of
punctatissima
, with its gaster detached and mounted separately. The lower specimen on this pin is a headless male of a small species of
Leptogenys
; its pectinate pretarsal claw is clearly visible on the right foreleg. This pin bears three additional labels, as follows:
"
Queen lectotype
Ponera breviceps Ber.
Designated by R.W. Taylor, 10 Aug. 63
"
;
"
Leptogenys
sp. male det. R.W. Taylor, 10 Aug. 63
"
;
"
Synonym of Ponera
gleadowi Forel
. Det. R.W. Taylor, 10 Aug. 63
"
. On the second pin both specimens are dealate queens of punctatissima and bear a secondary label:
"
Paralectotypes
Ponera breviceps Bern
. Designated by R.W. Taylor, 10 Aug. 63
"
.
Taylor's
designations were never published and the lectotype designation is formalised here. The
Leptogenys
specimen is excluded from further consideration.
11
Bernard's
syntype series of
ursoidea
included three workers and a queen from Guinea:
Crete
de Nion (all on one pin) and a single worker from Guinea: Nimba (Lamotte) on a second pin. On the first pin the top specimen belongs to
punctatissima
but the two workers and the alate queen mounted below it are all
dulcis
. The worker on the Nimba pin is a specimen of
dulcis
that has the right side of its petiole scale damaged by indentation so that it is deformed.
Bernard's
description contains a mixture of characters from both species. The first pin bears two secondary labels:
"
Lectotype (top specimen with marked pin) and paralectotypes of
Ponera ursoidea Ber. Designated
by R.W. Taylor
"
, and
"
A synonym of
Ponera gleadowi Forel
. Det. R.W. Taylor, 10 Aug. 63
"
. The lectotype designation
was
never published by Taylor and is now formally established here. The remaining specimens, all
dulcis
, are hereby excluded from the type-series. One small point remains: the lectotype is not a worker but an intercaste, as indicated by its larger than worker-sized eye and the presence of a mesonotal-mesopleural suture, a character absent from genuine workers of this species.
12 There is nothing in
Stefani's
descriptions and figures of the various sexes and castes of
sulcitana
that separates it from
punctatissima
. The worker description and drawings (figs. 1, 6, 11) show nothing that would detract from this conclusion, and the worker LPeI ca 50, estimated from his fig. 1, is within the range of
punctatissima
.
Stefani's
“apterous queen
"
(his fig. 3) appears to be a normal dealate gyne, not an intercaste, as the mesosoma seems to have a full complement of flight sclerites. His description and drawing of the ergatoid male makes the synonymy confident as the specimen is said to have 12 antennal segments, worker-like mandibles and minute eyes. Some years ago, Prof. Cesare Baroni Urbani (pers. com.) said that he had tried to obtain specimens from Stefani several times, but without success.
FIGURES 97
-
99. Lateral, full face and dorsal view of body.
Hypoponera punctatissima
worker CASENT0226650.
FIGURES
100
-
102. Lateral, full face and dorsal view of body.
Hypoponera punctatissima
ergatoid male CASENT0055828.
WORKER. Measurements: HL 0.56
-
0.72, HW 0.46
-
0.60, HS 0.515
-
0.660, SL 0.35
-
0.48, PrW 0.33
-
0.43, WL 0.70
-
0.90, HFL 0.36
-
0.48, PeNL 0.14
-
0.20, PeH 0.30
-
0.39, PeNW 0.22
-
0.29, PeS 0.223
-
0.290 (60 measured). Indices: CI 79
-
87, SI 75
-
84, PeNI 63
-
74, LPeI 43
-
53, DPeI 140
-
165.
Eyes small but conspicuous, of 1
-
6 often poorly defined or partially fused ommatidia, located far forward on the side of the head. Impression that extends back along midline of head from the frontal lobes reaches at least the midlength of the vertex and usually further. Apex of scape, when laid straight back from its insertion, usually fails to reach or less commonly just touches the midpoint of the posterior margin in full-face view; SL/HL 0.62
-
0.70. Minute punctulate sculpture of cephalic dorsum extremely fine and superficial. Mesonotal-mesopleural suture absent. Metanotal groove deeply incised across dorsum of mesosoma; mesonotum with a well-defined posterior margin. Propodeal declivity rounds into the sides, without sharp marginations or carinae. In profile the anterior margination of the mesopleuron rounded to very bluntly angular behind base of anterior coxa; without a prominent sharp angle or tooth. Mesopleuron smooth and shining, at most with a few scattered punctures; usually with a small reticulate patch in the extreme posteroventral corner above the mesocoxa. Petiole in profile with the anterior and posterior faces of the node weakly convergent dorsally; node longer just above the anterior tubercle than at the dorsum
.
Sternite of petiole in profile a rounded lobe, without angles anteriorly or posteriorly. Maximum width of first gastral tergite in dorsal view less than the width of the second tergite at its midlength. Base of cinctus of second gastral tergite smooth and shining in dorsal view, without cross-ribs. Posttergite of second gastral segment, from posterior margin of cinctus to apex, much broader than long. Disc of second gastral tergite minutely punctulate, appearing weakly microreticulate in places. Full adult colour varies from dull brownish yellow to very dark brown or almost black. In general, workers at the bottom end of the size range given above tend to be lighter and the colour becomes darker with increasing size, but this is by no means an entirely consistent rule.
Apart from variations in size and colour, all the worker material of this widely distributed species is morphologically uniform and answers the description above.
H. punctatissima
is without doubt the
world's
most accomplished ponerine tramp-species. Its range incorporates all tropical and subtropical zoogeographical regions, including most oceanic islands, and it also penetrates well into the temperate zones of both hemispheres where it is frequently synanthropic; for a brief synopsis of world distribution see Delabie & Blard (2002); European distribution is summarised in Seifert (2003). As well as the usual workers and alate queens, it also produces worker-queen intercastes (ergatoids) and dimorphic large and small ergatoid males, but never the usual alate male form. Because of its tramping ability, oddly polymorphic sex/caste system and the considerable confusion about the taxonomy within the genus,
punctatissima
has amassed an inordinately large synonymy. A glance at the synonymic synopsis above will illustrate how often intercastes and ergatoid males have been misinterpreted as workers of separate taxa by authors dating back to Roger (1859) and how often supposed infraspecific forms were incorrectly associated with other species.
In a recent analysis, Seifert (2003) reached the conclusion that
punctatissima
actually consisted of two species,
punctatissima
and
schauinslandi
, that were not distinguishable by standard morphology but could be separated by micro-morphometrics subjected to discriminant analysis. He concluded that
punctatissima
had the junior synonyms
androgyna
,
exacta
,
jugata
and
tarda
(of which only
exacta
had not been previously proposed as a synonym), and that
schauinslandi
was the senior synonym of
aemula
and
bondroiti
. He stated that he had examined type-material of all the available names that could probably represent
punctatissima
or
schauinslandi
, to ensure that no possible senior name remained unconsidered. Unfortunately, a quick glance at the synopsis above shows that many names associated with
punctatissima
remained unexamined for the Afrotropical region, that several synonyms proposed by Wilson & Taylor (1967) had been omitted, and that one name from the latter publication has the same date as
schauinslandi
. It is not surprising that Seifert failed to detect the Afrotropical relations as their identities could not possibly be assessed based solely on their often inadequate and frequently misleading original descriptions. Taken together, the Afrotropical type-material and the mass of other material examined here did not fall neatly into the two presumed species, or into any one of the two, which raises the possibility that either there are more than two extremely cryptic and virtually inseparable species currently confused here, or that there is only one somewhat variable genuine species present. The possibility remains strong that the discriminant functions applied by Seifert do not isolate discrete species, but rather indicate allopatric populations of the same species, or even different ecomorphs of a single species. The problem could not be completely resolved here, though the ecomorph possibility is favoured, so it was decided that the most parsimonious approach would be to regard all as a single species, under the earliest available name, until a complete analysis of all material, world-wide, becomes feasible. Whether this can be accomplished by morphometrics or must await a DNA analysis of various populations remains to be seen, but a dedicated study is certainly essential.
Therefore, at present,
H. punctatissima
workers are regarded as any specimen that possesses the characters of the
punctatissima
group (see introduction) and fits the description above. Its worker-queen intercastes are very worker-like but have larger eyes (usually 10 to about 20 ommatidia) and a developed mesonotal-mesopleural suture that is absent in the worker. Ergatoid males fall into major (larger, brown and with small eyes present, Figs 100
-
102) and minor (smaller, yellow and eyeless) categories (Yamauchi, et al. 1996) and are again very worker-like, especially in head structure, but have shorter scapes (SI 68
-
72), only 12-segmented antennae (as do workers, intercastes and queens), and of course fully developed but retractile male genitalia. Alate males have never been seen and it is probably now safe to say are never produced. True queens are alate, with much larger eyes (obviously>50 ommatidia) that have short setae projecting between the ommatidia, and have a full complement of flight sclerites on the mesosoma.
A closely related species,
ragusai
, has been confused (under the name of its junior synonym,
gleadowi
) with
punctatissima
in the past (e.g. Wilson, 1958).
H. ragusai
has much the same sex/caste system as
punctatissima
,
including
monomorphic ergatoid males which have 13-segmented antennae, as is most frequently the case in male ants. This species is very uncommon in the Afrotropical region and indeed seems rare everywhere. In general the worker of
ragusai
is small and yellowish. Its petiole in profile is lower and longer than in
punctatissima
and in dorsal view its petiole node is longer in relation to its width than in
punctatissima
. For detailed comparative notes see under
ragusai
.
One other Afrotropical species-rank name may also be a synonym of
punctatissima
,
H. lea (Santschi)
. Unfortunately, no type-material of this name seems to have survived and its identity has been rendered obscure. For the circumstances of this see under lea in the species inquirendae.
Finally, the small species described from Guinea as
sinuosa
by Bernard (discussed below) is also related to
punctatissima
and its allies. The single specimen of this species seems distinct from
ragusai
in the form of its petiole, but any revision of those two species should also reconsider
sinuosa
.
H. punctatissima
is very widely distributed in sub-Saharan Africa and appears relatively common. It is often retrieved from litter and topsoil samples in forests, and also from rotten wood, but it seems particularly frequent in areas disturbed by human activities, especially in gardens, plantations and crop fields.
Material examined. Guinea: Crete de Nion (Lamotte); Nimba (Lamotte). Ivory Coast: Bingerville (Mahnert & Perret); Banco (Delamare-Debouttville); Abidjan, Banco Nat. Pk (I.
Loebl
); Monogaga (Mahnert & Perret). Ghana: Legon (D. Leston); Poano (R. Belshaw). To g o:
Palime
, Klouto For. (Vi t). Benin: Porto Novo (Le Moult). Cameroun: Nkoemvon (D. Jackson); Mbalmayo (N. Stork); Mt Cameroun, Buea (Ross & Lorenzen); Victoria Div., Muyuka (B. Malkin); Prov. Sud, PN Campo (B.L. Fisher); Res. de Campo (D.M. Olson); Res. de Faune de Campo,
Ebodje
(B.L. Fisher). Gabon: Makokou (I. Lieberburg); Prov. Woleu-Ntem, Minvoul (B.L. Fisher); Prov. Ogooue-Maritime, Res. Monts Doudou, Doussala (B.L. Fisher); Doudou, Doussala (S. van Noort); For. de Mondah, Cap Esterias (Bartolozzi & Taiti).
Sao
Tome
& Principe:
Sao
Tome
I., Obo N.P. (J.M. Ledford). Central African Republic: Dzanga-Sangha, Bayanga (B.L. Fisher); Dzanga-Ndoki, Lidjombo (B.L. Fisher); Dzanga-Ndoki,
Mabea
Bai (B.L. Fisher). Democratic Republic of Congo: Elisabethville (Ross & Leech); Matadi (Ross & Leech); Lwiro Riv., Bukavu (Ross & Leech); St Gabriel (Kohl). Angola: Bruco (P. Hammond); Villa Arriaga (Ross & Leech); Benguela, Cucula (J. Cruchet); Benguela, Caconda (J. Cruchet). Ethiopia: Ilubador Reg., Buno Bedele (Sforzi & Bartolozzi). Rwanda: Rangiro (P. W e r n e r). Kenya: Shimoni (Alluaud & Jeannel); Likoni (Alluaud & Jeannel); Western Prov., Kakamega For.,
Udo's
camp (S. Maurer); Ekero (G. F i s c h e r); Western Prov., Mwanza (F. Hita Garcia); Kakamega For., Colobus (M. Peters); Kakamega For., Isecheno (F. H e r c h e n); Isecheno (R.R. Snelling); Isecheno Nat. Res., Kalunya Glade (R.R. Snelling); Western Prov., Bungoma, Nzoia (F. H e r c h e n). Uganda: Jinja (Ross & Leech); Kalinzu For., Bushenyi Distr. (S. Yamane). Ta n za n i a: Amani (Ross & Leech); Kilimandjaro, Kiboscho (C. Alluaud); Morogoro Reg., Mkungwe For. Res. (Hawkes, Bhoke & Richard). Zambia: Lusaka (W. H. Lange). Botswana: Serowe (P. Forchhammer). Zimbabwe: Bulawayo (G. Arnold); Umtali, Melsetter (R. Mussard). South Africa: Natal, Durban (F. B. M a r l e y); Durban (G. A r n o l d); Zululand, Eshowe (R.E. Turner); E. Cape, Mkambati Game Res. (B.L. Fisher); Cape Devon, Stellenbosch (
O'Brien
& Marshall); Cape Prov., Oudtshoorn (B. Malkin); Cape Prov., Tsitsikamma N.P. (R.B. Kimsey). Comoros: Anjouan (Fisher, et al.); Anjouan, Ntringui (Fisher, et al.); Anjouan, Lac
Dzialandee
(Fisher, et al.); Moheli (Fisher, et al.); Grand Comore, Karthala (Fisher, et al.); Grand Comore,
Grille
(Fisher, et al.). Mayotte I.: Dapani (Fisher, et al.); Mont Chongui (Fisher, et al.); Sohoa For. Res. (Fisher, et al.); Mont Benara (Fisher, et al.).
Reunion
I.: Le Port (F. B la r d); Mare Longue (Fisher, et al.); Les Makes (Fisher, et al.). Mauritius: Le Pouce Mt, Moka Range (Fisher, et al.); Cocotte Mt (Fisher, et al.); Pieter Both Mt, Moka Range (Fisher, et al.); Basin Blanc (Fisher, et al.). Seychelles: Alphonse I. (J. Gerlach). Poland: Opole Prov., Rudy (Roger). Germany: Berlin (Roger). Netherlands: Mijdrecht (P. v. d. W i e l). Italy: Sardinia, Villamassargia, Grotta dei Pipistrelli (S. Puddu); Sicily, Siracusa Prov., Pachino, Grotta Calafarina (B. Poldi). United Kingdom: England, Southsea (H. Donisthorpe); Oxfordshire, Oxford (E. Saunders); Oxford (P. Harwood); Middlesex, Old Ford (Bedwell); Hertfordshire, Waltham Cross (W. H. T u r n e r); Dorsetshire, Holt (P. Harwood); Kent, colliery below ground (no
collector's
name); Yorkshire, Barnsley, Darton (no
collector's
name); Scotland, Edinburgh (Godfrey); Glasgow (Min. of Agiculture); Wales, Cardiff (no
collector's
name). Ireland: Co. Cavan, Coote Hill (J.H. Cole). Madeira I.: Praia Formosa pr. Funchal (Lindberg). St Helena I.: (Wo l l a s t o n). Canada:
Quebec
(A. Francoeur). Hawaiian Is.: Kauai I., Lahue (R.C.L. Perkins); Oahu I., Honolulu (R.C.L. Perkins). New Zealand: Wellington, intercepted in quarantine, from Canada (no
collector's
name). St Vincent I. (Antilles):
Chateaubelais
Bay (H.H. Smith).