Introduction The Ants of Africa
Chapter 2 - Geography and History - Ghana

Summary of known collectors
W Belfield (Bolton, 1980); R Belshaw (Belshaw, 1994b; Bolton, 2002); M Bigger (Bigger, 1981a); B Bolton (Bolton, 1971a, 1974b, 1975, 1976, 1980, 1981b, 1982, 1983, 1991; Brown, 1978c); H E Box (Donisthorpe, 1945b); H Brauns (Mayr, 1895); R Buchholz (Mayr, 1902); C A M Campbell (Bolton, 1981b, Shattuck, 1991); C A Collingwood (Bolton, 1973b, 1974a, 1980, 1982, 1983, 1987); G F Cotterell (Bolton, 1973b, 1981b; Donisthorpe, 1942e); D J Cross (Bolton, 1973b, 1981b); Fisch (Forel, 1913h; Santschi, 1919b); Ganger (Forel, 1894b); D Gibbs (Bolton, 1974a); W H Gotwald (Bolton, 1983; Gotwald, 1973); W J Hall (Donisthorpe, 1945b); Hollingsworth (Hollingsworth, 1960); Imhoff (Emery, 1902c); A B S King (Bolton, 1971a); D Leston (Bolton, 1973b, 1974a, 1975a, 1975b, 1980, 1981a, 1983, 1987, 2002 ; Leston, 1973); J D Majer (Bolton, 1980, 1982; Majer, 1975); J Paine (Donisthorpe, 1945b); H Pirazzoli (Mayr, 1862); J Plisko (Bolton, 2002); Edmund Reitter ((Mayr, 1895); O W Richards (Bolton, 1974a); P M Room (Bolton, 1980, 1981a, 1981b, 1987; Room, 1971, 1975); E S Ross & R E Leech (Brown, 1978c); E S Ross & K Lorensen (Bolton,1983); F Silvestri (Santschi, 1914d); A H Strickland (Donisthorpe, 1948f; Strickland1951a); Unger (Wheeler, 1922 list).

Note: Marcus Stüben has told me (September 2009) that Isert (Paul Erdmann Isert), who often is listed from Fabricius (1793) as collecting in "Guinea" (and was by me also until now) actually collected in "Danish Guinea" = Gold Coast = modern Ghana. Marcus wrote:-

"I found out that the type specimen of "Formica foetens Fabricius, 1793" [Pachcondyla (Megaponera) analis] collected by "Isert in Guinea" has actually been collected in "Danish Guinea" = "Gold Coast" = present-day Ghana. Isert collected mainly plants, but also fishes, insects (including ants), and probably also other animals. He mainly collected in Ghana, but sailed also along the coast of Dahomey (Togo) and collected also in southern Benin. For Togo and Benin I have so far no evidence that he collected ants there.

Paul Erdmann Isert, in Danish service as surgeon/physician (and naturalist): born 20.10.1756 in Brandenburg, Germany, died 21.1.1789 in Ghana. He published in German. Apart from West Africa he collected also in the Caribbean: St. Croix, etc.
Isert in Ghana: 11.1783 to 10.1786, 11.1788 to 21.1.1789 (death of Isert).
See also here: Isert, P. E. (1788). Reise nach Guinea und den Caribaischen inseln in Columbien, in Briefen an seine Freunde beschrieben, Verlag J.F. Morthorst, 376pp.
Wilks, I. (1994). "Letters on West Africa and the Slave Trade. Paul Erdmann Isert’s ‘Journey to Guinea and the Caribbean Islands in Columbia’ (translator and editor Selena Axelrod Winsnes)." Sudanic Africa 5: 274-276. and On 30 March 1850 all Danish Gold Coast Settlements were sold to Britain and incorporated into the British Gold Coast. See also my Chapter 2 - Geography - West Africa & the Congo Basin - introduction.

The early collections include Missionary Widmann, whom Dr Ludwig Imhoff reported (1852) as observing ants at the Akropong Mission. Mayr described Camponotus flavomarginatus (Mayr, 1862) and Phasmomyrmex aberrans (Mayr, 1895) from Ghana. Dorylus (Anomma) funereus and Dorylus (Anomma) gerstaeckeri have type localities in Ghana (Emery, 1895j). H. Brauns collected at Kitta (modern name Keta, 5°55'N, 0°55'E) and Accra (5°31'N, 0°15'W), before 1895, he also collected in Liberia, and after (?) was in Angola and South Africa. An unknown collector was responsible for discovering Camponotus acvapimensis from "Akwapim Mountains" (5°50'N, 0°20'W), a hilly area a short distance inland from Accra (Mayr, 1862). R. Buchholz also collected at Accra (Lepisiota capensis var. guineensis, in Mayr, 1902). Professor F. Silvestri, who collected at Aburi, in 1913 (Santschi, 1914d). Most of these locations are shown in Map 2.

Several ant species were described by H.St.J. Donisthorpe, then the British Museum myrmecologist, who seems to have been responsible for many of the older labels on ants in the CRIG and CRIN collections (Donisthorpe, 1942, 1945a & 1945d). He also wrote a note on ants associated with coccids (Donisthorpe, 1945b). Three or four field entomologists were involved. Harold E. Box, was a somewhat peripatetic figure, reporting on coffee pests in Kenya in 1921, sending insects from British Guiana to the Imperial Institute of Entomology in 1923, and at the Imperial College of Agriculture in Trinidad in the 1930s. In Ghana, he studied capsids (a cocoa mirid is Boxia khayae) and swollen-shoot virus vectors; making several ant collections, including Crematogaster wellmanni (as Crematogaster boxi). He also reported on visits to sight the cocoa pest situation in West Africa; travelling to Togo (British Togo, see Crematogaster togoensis), Dahomey (French Togo), Nigeria (including Owena, see Oecophylla longinoda) and the Cameroun, between December 1943 and March 1944 (WACRI Technical Bulletin, 1, 69) and perhaps was active in the area until at least January 1945. In November 1944 - January 1945, J. Paine appears to have worked or liaised with Box (see Crematogaster painei). W.J. Hall was a coccid specialist and another of the much travelled entomologists. In 1923 he was Entomologist with the Agriculture Department in Cairo, then in the early 1930s was in Zimbabwe (Southern Rhodesia). From Ghana, he described a mealybug vector of cocoa swollen shoot virus (information given and reports cited in Entwistle, 1972) and was denoted by the naming of Crematogaster buchneri junior synonym halli. G. F. Cotterell, who was in Ghana (Gold Coast) from before 1926 to at least 1942), being in the Department of Agriculture and finally at the "Central Cocoa Research Station, Tafo", also worked on capsids, collecting Monomorium rosae, in December 1940 (as Diplomorium cotterelli, in Donisthorpe, 1942b), and Polyrhachis weissi.

The research station at New Tafo, was established in 1938 to investigate the rapid die-back of cocoa. It became the West African Cocoa Research Institute in 1944, and, after independence, was re-named the Cocoa Research Institute of Ghana (CRIG) (CRIG in 1975) . The Institute was described by Gibbs & Leston (1970) as located at the eastern end of the Ghana-Guinea bloc of moist forest at low and medium altitudes, with most of the cocoa under "Ghana farm conditions", where many forest trees are left to give shade. Under the canopy, this "cocoa forest" was characterised by a low intensity of light comparable to that in primary forest. CRIG itself lies at 140 m a.s.l., Kade is at 200 m, and Mampong Akwapim at around 460 m; cocoa farms are found up to around 600 m (Akotoye, 1975). Map 3, has a general map of southern Ghana, showing many of the collection areas (adapted from Belshaw & Bolton, 1994b); and Map 4 is a larger scale map of the area in which much of the survey work was carried out (adapted from Room, 1971).

The earliest bionomic study of ants in West African cocoa, or at least the recognised starting point for modern research, was that of A.H. Strickland, in 1945. Not long before it had been discovered that mealy-bugs (Pseudococcidae) were vectors of swollen shoot disease of cocoa and Strickland's main task was an investigation of their association with ants (Strickland, 1948, 1951a, b). He described finding at least 75 ant species associated with cocoa in Ghana (as the Gold Coast) but his lists grouped ten Pheidole spp., and various Crematogaster species (into the then and now-accepted subgenera of Wheeler, 1922). Strickland's detailed investigations all appear to have been made at CRIG, although he gave some information on the wider occurrence of species. One of Strickland's colleagues appears to have been H.M. Amoah (see Paedalgus distinctus).

P.B. Cornwell, followed Strickland's line of study with a closer examination of the habitats and environment on populations of mealybugs and ants in the "same square mile of cocoa at Tafo as that surveyed by Strickland". His work was carried out from 5.i.1954 to 11.ii.1955 (Cornwell, 1957).

The work of the International Capsid Research Team, which was based at CRIG between 1965 and 1971, with two members (R.H. Booker and Barry Bolton) spending some time also in Nigeria, included extensive investigations into the question of ant-capsid relations (Collingwood, 1971). The main studies on ants were by Dennis Leston, in 1966-68 (see below); Cedric Collingwood, in 1968-71, as Team Leader; and, Barry Bolton, in 1968 and 1970-71, who concentrated on collecting and starting to identify the many ant species which were being encountered (see, for instance, Bolton, 1970-71). Room (1971) stated that Bolton et al. (unpublished) had found some 250 species of ant in Ghanaian cocoa farms but no comprehensive list seems ever to have been made available. D.J. Cross, another member of the team, mainly worked on the capsids but did make a study of Oecophylla and predation (Cross, 1969, cited in Entwistle, 1972). H. Marchart used radiotracers to study mirid predators and found Platythyrea conradti to be among them (Marchart, 1968, cited in Entwistle, 1972).

Also based at CRIG and contemporary with the Capsid Team, Harry Evans investigated various aspects of cocoa black pod disease, including the involvement of several species of ant (Evans, 1971, 1973; Evans & Leston, 1971). A further effort at CRIG was the further investigation of Swollen Shoot Virus by Mike Bigger, who started in 1969, and Colin Campbell, who joined the SSV team in 1973. Both focused on ant-Homoptera associations. Their work was confined, however, to restricted areas - four plots each of around 225 trees by Bigger, and ten sampling areas from within a 13 ha plantation with some 14,000 trees by Campbell (Bigger, 1981,1993a, b; Campbell, 1984, 1994). Perhaps less glamorous but useful work on insecticides, insect control and minor pests, including those attended by ants, was done by Miss B. Adjei and Ed. Owuso-Mano.

A second focus of research was the University of Ghana at Legon, near Accra; notably by Dennis Leston, who moved there from working at CRIG, Peter Room and Jonathan Majer. Much of this research concentrated on cocoa in southern Ghana. An MSc thesis by a student at the University, E.A. Aryeetey, reported a study of Tetramorium aculeatum (as Macromischoides aculeatus). Although the thesis has been cited several times by contemporary workers, the results were not published and cannot be described as readily available. A few ant collections were made by S.B.V. Mkhize, who studied the lepidoptera of cocoa farms in Ghana for a Legon MSc. S.K. Firempong examined the biology of the aphid, Toxoptera aurantii, on Ghana cocoa and noted that it was tended by certain ants.

Between October 1968 and March 1970, Peter Room made a detailed survey of a small cocoa farm, the Mampong Cemetery site, beside the Mampong-Mamfe road, at the southern edge of the cocoa belt, bordering on the coastal savannah (Room, 1971). He also surveyed the cocoa canopy and mistletoe (hand sampling from a ladder) on thirty farms over a wider area of southern Ghana cocoa, shown in a map in his paper (see Map 4). Room later reported specific collections of the insects found on cocoa mistletoe, Tapinanthus bangwensis (Room, 1975). A final aspect of his work seems to have been a survey of northern savannah areas, locations including Tumu and Navrongo, a northern border town; work which he extended into the Sahel in neighbouring countries (e.g. Ouagadougou, in Burkina Faso; Gao, Bourem, Tessalit, Anefis and Labezanga, in Mali; Ayorou, in Niger; all, I surmise, while on a trans-Saharan road journey).

Jonathan Majer did all his research into the ant mosaic at the Agricultural Research Station of the University of Ghana. This is near Kade, which is in the Celtis-Triplochiton moist semi-deciduous forest. Most of the results were obtained from two adjacent plots of cocoa, both under relatively light shading, and with a total area of 2.25 ha. His list of the insects on cocoa was compiled from the use of pyrethrum knock-down (pkd) insecticide sprays, to collect 144 twenty-five m² samples over a 16 month period, January 1971 to April 1972, and from an undefined number of hand sampling collections (Majer, 1972, 1975, 1976a, b, c).

The work by Dennis Leston, who initially worked with the Capsid team at CRIG (being there in 1966), and was at Legon in 1969-72, is widely cited but, curiously, only limited details can be found in easily available, published, sources. One such source is that on the "Ectatommini" of Ghana (Leston, 1971); however, the ants concerned were Proceratium species, which now are placed in the Tribe Proceratiini. For instance, in his generic revisions, Bolton often lists findings such as, "1966, ant ecology sample (D. Leston)" - among the locations were CRIG; Kade (see below); Mt. Atewa, a primary forest area close to Kibi in the Eastern region, from which there are numerous unique findings; Legon (a coastal savannah location); Wiawso, Western Region (Evans & Leston, 1971); and, a number of locations most of which actually are on the map given by Room (1972), or, it seems safe to assume, are in the area covered by the map (Aburi, Asikaisu, Bosusu, Nkwantanan, Sajimasi). Leston's collection methods include pkd sprays, simple hand collections, and the undefined 'ant ecology samples'. At CRIG, between March 1966 and September 1967, he and D.G. Gibbs made regular pkd collections in three blocks of cocoa, but the detailed publication from that work (Gibbs & Leston, 1970) has information on only two species of ants, Oecophylla longinoda and Odontomachus troglodytes (as Odontomachus haematodus).

{Coastal scrub near Accra}Other than the cocoa entomologists, there is a small number of other findings of ants. These include - Serrastruma lujae, by E.S. Ross & K. Lorenzen, at Bolgatanga (Bolton, 1983); Serrastruma ludovici, by W.H. Gotwald & R. Schaefer; Tetramorium angulinode, by W. Belfield at the Agricultural Station, Pokoase (he described West African scorpions in 1956; location misspelt by Bolton, 1980; 5°41'37"N 0°16'52"E); and, those of J.C. Grieg at Mole Game Reserve (Polyrhachis viscosa, Monomorium rosae). Cataulacus huberi was collected by O.W. Richards at Mt. Atewa and Mampong; plus Cataulacus traegaordhi at Accra; the latter species was also found, at Pokoase and Koforidua, by N.L.H. Krauss.

The study of driver ants in Kenya and Ghana (Dorylus nigricans and Dorylus gerstaeckeri) by William H. Gotwald Jr. has added interest because he included photographs and descriptions of his study areas (Gotwald, 1974). The coastal scrub and grassland of the Accra Plains near Legon has relatively few and scattered small trees; the Guinea savannah-woodland of the Mole Game Reserve near Damongo has larger and more dense tree cover; and, the third area, the dense moist semi-deciduous forest at Kade, is secondary forest with a good number of climax trees still present. He cited the area of Guinea savannah-woodland in Ghana as being some 168,000 km², and the high forest as about 65,000 km². At Legon, he studied the driver ants on the University campus and in the cultivated fields of the University farm system. In addition to the forest at Kade, he also studied colonies foraging in and around cocoa plots at CRIG.

{Guinea savannah} The most recent research has been by Robert Belshaw, who studied the leaf litter fauna of various forest areas between December 1991 and November 1992, under the guidance of Barry Bolton (Belshaw & Bolton, 1993, 1994b). With the results of their survey, they provided a map of the 34 sites, which were scattered across the moist semi-deciduous tropical forest zone (see Map 1). The south-eastern sites are in the same localities as the most northern sites surveyed by Room (1971); and the map shows Room's other sites to lie in the dry semi-deciduous forest. Interestingly, no significant difference was found between the sites in primary forest, secondary forest and cocoa farms, either in species composition or in species richness (Belshaw & Bolton, 1994b). All told, they collected a total of 197 species and 47 genera of ants. Of the total at least 17 species were definitely new to science, plus two new genera (Afroxyidris and Loboponera). A further 57 species were in the groups which remain lacking modern keys or sound taxonomic studies. The leaf litter inhabitants numbered 176 species, which they calculated to be some 81% of the actual species present. Three species were subterranean, being found only in soil samples, up to about 5 cm deep. The remaining 18 species were regarded as 'tourists', most being arboreal, plus four Camponotus species which are known to forage on the ground.

By extrapolation from Belshaw & Bolton (1994b), one can get an idea of how many other species there could be in Ghana. First, they believed that they probably missed a number of species which nest in rotting logs, then there are the purely arboreal species, fast moving terrestrial species (especially Camponotus species), and the known abundant Dorylines (of which they found only one undetermined species). They enumerated some 120 species from well-studied genera, and were unable to even hint at how many Pheidologetonini, Pheidolini and Crematogasterini there may be. Thus, in all it is easy to suggest a figure of well over 350 species of ant which could be found in the forest zone of Ghana.

©1998, 2001, 2003, 2006, 2009, 2012, 2014 - Brian Taylor CBiol FSB FRES
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