The Ants of Africa
Chapter 2 - History - Introduction

See also Listing of known ant collectors and The Exploration of Africa by Europeans and Colonial Development - up to ca 1907

Historic Collections

Initially, I sought to examine and cite work published since 1945 but the new catalogue by Bolton (1995) and other modern taxonomic revisionary studies show how much we owe to the assiduous collectors and expeditions of the late 19th and early 20th Centuries. Throughout the text the date suffixes for taxonomic papers are those given in Bolton (1995).

{P A Latreille}As a Briton, I have to start by mentioning the Rev. T.S. Savage, a medical missionary in West Africa who described the marching habits of "drivers" or "visiting ants" as long ago as 1847 (Savage, 1847, 1849; Westwood, 1847). His contemporary, W.E. Shuckard wrote a series of four papers in a review of the "Dorylidae" which included a number of West African species of Dorylus (Shuckard, 1840 a-d).

The names of many of the collectors can be found attached to the species named during the era and a potted biography of a number of the taxonomists can be found in Morley (1953). The pioneer taxonomist Pierre Latreille, describer of Oecophylla longinoda in 1802, was the subject of a short biography in Biologist, 44, 276 (February 1997, O.A.E. Sparagagno), the profile (right) is from the frontispiece of the Centenary volume of the journal of the Entomological Society of France (Santschi, 1932a).

{H W Bates} In a paper on the specimens of ant held in the Zoological Museums in Munich, Forel (1911), has notes on a good number of species collected by H. W. Bates, one of the great collectors of the Victorian era and author of The Naturalist on the Amazons (1863), who was the first to describe the leaf-cutter ants of that area.
H(enry) W(alter) Bates, born 8 Feb 1825, died 16 Feb 1892, was a naturalist and explorer whose demonstration of the operation of natural selection in animal mimicry (the imitation by a species of other life forms or inanimate objects), published in 1861, gave firm support to Charles Darwin's theory of evolution. He and Alfred Russel Wallace left England in 1842 to explore and collect insects in the Amazon basin. Bates spent 11 years in Amazonia amassing large collections of insects that were sent back to museums and collectors in Europe. Bates was quick to embrace Darwin's and Wallace's theory of evolution by natural selection. Bates' own theory of mimicry, which now bears his name (Batesian mimicry), provided evidence for evolution by natural selection. From 1864-1892, he was Assistant Secretary at The Royal Geographical Society in London. The RGS doled out more than £20,000 (a substantial sum in Victorian monies) specifically for African exploration during the years 1864-1892, a period that matched H.W. Bates’s twenty-eight-year tenure as assistant secretary and during which Britain maintained an unusually strong interest in Africa. And as the society’s only permanent officer, Bates, a naturalist who had returned to London in the late 1850s after extensive travels in South America, single-handedly prepared virtually all of its correspondence. It proved to be a pivotal position. Bates did more than simply administer the RGS during one of the most active periods in its history. Rare was the explorer seeking British support for expeditions in Africa (or anywhere else in the world) who did not go through him. Furthermore, as a biographer said of his remarkable career at the RGS, "[H]e gave understanding help to little-known men setting out on their journeys, and when they returned famous, he was usually the first to greet them." [partial sources of brief biography - Today in Science and Mercator's World]. Not mentioned in the biographies I have seen, however, seem to be later journeys (or a single journey) taking in countries in South Asia (Cambodia, Celebes, Ceram, Sulawesi, New Guinea). Perhaps on that journey, he collected ants in Old Calabar (eastern Nigeria) and Natal, specimens of which were among those Forel described, notably Crematogaster batesi and Aenctus batesi (a junior synonym of Aenictus decolor). An alternative that seem possible is that the ants came from a collection originally belonging to Bates, with the specimens collected by someone else - perhaps Wallace? In his paper Forel does not give any collection dates.

The first taxonomic work in what might be called a proper scientific style was that of Gustav L. Mayr (publishing from 1853 to 1907), who identified and reported several collections. The earliest (Mayr, 1862) was a study of ants in the University Museum in Vienna, among which was a small collection of ants from the Gold Coast (Ghana), apparently collected or sent to Professor Rudolf Kner, possibly the collector was Pirazzoli, among them Camponotus acvapimenis and Camponotus flavomarginatus, both from the Akwapim Hills. Carebara sicheli was described by Mayr, from a specimen collected in Senegal by Dr Sichel. In the same paper he described species from South Africa, including Lepisiota capensis - from the Cape of Good Hope, hence the name capensis. Most of the material, at as with his later description of Anochetus africanus was collected on a circumnavigation voyage by the Austrian frigate "Novara" under the command of Commodore B. von Wüllerstorf-Urbair (Mayr, 1865) - described at A small amount of material also had come from the collecting efforts of Herr Ritter van Frauenfeld in the Sinai Pensinsula of Egypt. Writing in 1874, Forel commented (my translation) - "Without doubt the most important author is Mayr. His remarkable perspicacity in the creation of genera, and in general in the recognition of the value of zoological characters, the minute exactitude of all his writings which represent a very considerable sum of work has raised myrmecology to the highest level of entomological understanding".

Later, there were the collections by Professor Dr Yngve Sjöstedt and R. Buchholz from Cameroun (Mayr, 1896, 1901a), and A. Mocquerys from Sierra Leone (pre-1889). Sjöstedt personally described his collections of Orthoptera in Cameroun and later led an expedition to the Mount Kilimanjaro area of Tanzania; Mocquerys also collected in Senegal. Others were Professor R. Buchholz, collecting in upper Guinea and Accra, Ghana (Gold Coast), and Dr H Brauns in the "Los-Inseln" of Senegambia. In Mayr's 1862 paper there is what must be one of the earliest uses of a dichotomous key, "eine analytische Arbeitertabelle", in this case to the genera of Formicidae and Ponerinae ("Poneridae") and to the species of the Australasian genus Myrmecia. In most cases, his individual descriptions of new species are in the format and detail common to any modern paper.

A number of species, notably Ponerines, were described by Julius Roger in a short series of papers (1859, 1860, 1862a, 1862b). He was based in the Museum in Berlin but his 1860 paper gives an interesting list of the museum taxonomists active across Europe at that time.

{Felix Santschi}Perhaps the most outstanding collector was Professor Filippo Silvestri, whose 1913 "voyage" to west and south Africa (he was at Conakry, Guinea, on 8.viii.1913; a date of 24.viii.1912, see Cataulacus tardus, seems likely to be wrong) yielded many new species. The species were described by the very prolific taxonomist Felix Santschi (1872-1940), who was publishing between 1906 and 1941 (see Santschi 1914b & d). Professor Silvestri earlier collected extensively in southern South America (Emery, 1906c) and later collected ants in South-East Asia (Indochina, Wheeler, 1927) and himself studied Chalcidid wasps, also writing on South American termites.

{Santschi signature ?}Santschi himself was described by Forel (1904c) as "M. le Dr. Santschi, de Lausanne, établi comme médecin dans la ville sainte des Arabes de Tunisie" and seems to have lived mainly in Kairouan, Tunisia, around where he made personal studies and collections of primarily desert ants. Kairouan, now known as Al Qayrawân, is also called the City of 100 Mosques. The earliest of his own reports of personal collections in Kairouan is recorded as January 1906 (Santschi, 1907), although Forel (1905b) described Monomorium santschii [as Wheeleria santschii] as collected by Santschi at Kairouan on 19 August 1903, and Solenopsis santschii as collected in a Kairouan street on 1 August 1903. In his 1907 paper, Santschi dedicated the newly named species Leptothorax foreli to Professor A. Forel, "who initiated my interest into the so interesting study of ants" [my translation]. What appears to be his signature (right) is written on the copy of his 1932a paper that has been scanned for the Antbase website.

Others whose ant collections were described by Santschi are Ch. Alluaud, in the "Territory of Assinie" (Ivory Coast, close to Ghana border) in July-August 1886 (Plectroctena minor, Santschi, 1914b, Emery, 1892d), he collected ants in East Africa in June 1903 to May 1904, and in 1911, together with René Jeannel (1879-1965), he had been to Madagascar in 1893 and to the Seychelles in 1899. Dr. F. Zumpt, on a voyage to Cameroun, in October 1935 (Santschi, 1937b); and R.P.E. Wassman, at Gr. Batanga, Cameroun (Santschi, 1926b). A. Weiss collected several Polyrhachis species from Congo Brazzaville, including Polyrhachis weissi (Santschi, 1909). Santschi (1910c) describes Weiss as being a member of an expedition to study sleeping sickness ("maladie du sommeil") in French Congo, spending a year in the Brazzaville region (Mindouli, M'Bomou), from where he found a good number of new species and numerous species not previously known from that part of the Congo Basin. Another collector whose work Santschi described was Ivar Trågärdh who was active in South Africa in 1913-1914. Trågärdh had earlier collected in Sudan (Mayr, 1904b).

{short description of image}Photo: Collecting Microdipnus jeanneli All. on north-east slope of the Mt. Kenya, 2.750m. Ch. Alluaud (in center) and R. Jeannel (on right), February 1912 (from Jeannel, 1963). On his website at, Vasily Grebennikov writes - "I clearly remember the day when for the first time I understood that I want to go to East African mountains. It happened in mid-October 1992, in the library of the Zoological Institute in St. Petersburg. Then an undergraduate, I was reading Rene Jeannel's (1963) monograph on Anillina (Coleoptera: Carabidae: Trechitae: Tachyini); a group of carabid beetles which is totally absent from the former USSR. There is a picture in that book: Jeannel and Alluaud turning a large rock on the Mt. Kenya searching for annilines. Well, this was it and since then I started to plan the expedition".

Carlo Emery, who was based first in the University of Bologna, where he was Professor of Zoology, and later in Geneva, published extensively between 1869 and 1926, including collections by L. Conradt at Mundame in Cameroun (Polyrhachis platyomma, 9.xi.1895, in Emery, 1921e) (others of Conradt's collections were described by Stitz).

Among the taxonomists who named species in that golden era was E. André (publishing between 1874 and 1905), who described much material from Sierra Leone (André, 1889, 1890), collected by Albert Mocquerys, who also collected in Ogooué, Congo (André, 1895a); and from Cameroun, collected by L. Conradt; others from whom he had material were Wilverth (from Congo) and Professor Aurivillius. Staudinger & Bang-Haas also forwarded specimens to him from South Africa, Angola and Nigeria (Emery, 1915c, referred to them as "the house of Staudinger & Bang-Haas", so it seems they were dealers in insect specimens).

{Forel 1902 personal postscript}Auguste Forel (publishing between 1870 and 1922, based in Geneva, see his personal postscript, right, to his 231st paper on his "little old friends", when the moment came for him to stop because of glaucoma affecting his sight) had among his works the names of the collectors Ingenieur Alfred Ing, Dr. Liengme, Missionary Paul Berthoud and Dr. Arthur Müller, whose collections included some from Senegal (e.g. Pheidole mayri, in Forel, 1894b). Forel (1875), himself then medical associate at the hospital for aliens in Munich, described Missionary Paul Berthoud and his wife, otherwise Mlle Exchaquet, as sent by the free church of the canton of Vaud to Lesotho. Others were J. Bequaert (Monomorium bequaerti, from Lumbumbashi [Elizabethville], Zaïre, 20.iii.1912, in Forel, 1913b), Father Hermann Kohl (see Crematogaster kohli, in Forel, 1909b), who appears to have lived at the Roman Catholic Mission station at Kisangani [Stanleyville], Zaïre, in 1901-1905, and was at St Gabriel Mission in 1909 or thereabouts (from Wasmann, 1918b). Both Bequaert, a botanist, and Kohl were particularly interested in ants associated with plants. To them one can add E. Luja (see Bondroitia lujae, he also collected in Mozambique) and R. Mayné, who collected in Zaïre (then the Belgian Congo, e.g. Serrastruma maynei, Forel 1913b, 1913c, 1916, and see below); collections by all of them were published by the other taxonomists (Emery and Santschi). In the 1909b paper, other collectors whose names appear also include - Lemaire (Katanga), Duchesne (Haut Congo), Kinsbergen (Ikelemba), Dr A Jullien (Luki), Lamarche (Léopoldville), Leboutte (Boma), Lindemans (Iringui), Deleval (Congo), Cabra (Mayumbe, Chikai, La Lukula); R. Rhode (Mukonje Farm, Cameroun), don Lopez (Zambi), Tosquinet (Vivi, Congo), Lemarinel (Congo), De Pauw (Léopoldville), Waelbroek (Kinshasa), Petit (Landaéna and Condé), Dr Neave (Kalumba and Katumba), Weyns (Leo-Stanleyville), Rollin (Boma Sundi); and Solon (who sent him specimens from in the stomach of a pangolin from Bas Congo

H. Stitz (papers from 1909 to 1939) described ants collected from West Africa by Tessmann, who was from Lübeck (Stitz, 1910), and by the German Central Africa Expedition 1910-11 (Stitz, 1916, Pachycondyla sennaarensis).

{Menozzi signature}C. Menozzi (publishing from 1918 to 1952; see 1933a) was a more recent taxonomist. He identified material collected by Professor E. Zavattari, of the University of Pavia, at locations around the Gulf of Guinea, including in Gambia (Bathurst), Nigeria (Port Harcourt, Koko, Sapele), and Gabon (Port Gentil) (Menozzi, 1926a).

Another who made a relatively limited contribution to the knowledge of African ants was H. Wiehmeyer of Dresden. He examined specimens mainly from (Anglo-Egyptian) Sudan collected by Austrian and Swedish expeditions of the early 1900's; and, earlier, specimens from Tanzania and Cameroun (both then German colonies).

H. Wasmann, a Jesuit, of Valkenburg, Holland, also published a few texts on Congo ants, with some from Father Hermann Kohl (see above) (Pheidole neokohli) and more from von Rothkirch who collected at an altitude of 730-800 m on Mount Cameroun in December 1912. Wasmann's studies seem primarily to have been on insect guests or parasites of ants.

Although his primary interest was in the ants of southern Africa, George Arnold, Curator of the Rhodesia Museum, Bulawayo, produced an extensive Monograph of the Formicidae of South Africa, published in parts between 1915 and 1924, with an appendix in 1926. In addition to his own descriptions, Arnold provided translations of papers originally written, for instance, in German, French or Italian. As a fair number of the species he examined had been described from the Congo Basin, if not from West Africa, his catalogue is a useful source of descriptions. His later papers, the last being published in 1962, include some relevant observations.

There were others but one who must be remembered for one major piece of work on Africa, among many publications from other areas between 1900 and 1942, is William Morton Wheeler, who made a massive study of the ants collected by the American Museum Belgian Congo expedition of Herbert O. Lang and James P. Chapin in 1910 (-1913, see Ragge, 1980), together with a smaller collection by Dr. J. Bequaert (Wheeler, 1922). Wheeler was aided especially in separating out the various Crematogaster by Dr. F. Santschi. The volume of reports from the expedition includes Parts by Bequaert, on the predacious enemies of ants (III), and on the diverse relations of ants to the plant world (IV). Wheeler himself provided Parts covering the distribution of ants in the Ethiopian and Malagasy Regions (I), the ants collected by the expedition (II), keys to the genera and subgenera of ants (VII) and synonymic lists of the ants of the Ethiopian Region (VIII) and the Malagasy region (IX). In Part VIII, he provided a map of the collection locations and a list of localities in Africa, with latitude and longitude where possible.

Although I have extracted much of the information on the ants, the full text of the expedition reports now (March 2002) has been placed on the web at and is well worth a visit.

Modern Collections

More recent expeditionary studies are given, together with the entomologists who have worked on agricultural and ecological projects, in the subchapters which follow but include:
The work in Guinea where Maxime Lamotte and A. Villiers collected ants from the Mt. Nimba area, reported by Francis Bernard (1952);
Between June 1949 and January 1950, Albert Raignier and Jozef van Boven of the Instiute de Zoologie, Louvain, Belgium, carried out intensive studies of Dorylines in Yangambi, Belgian Congo, now Zaïre. It seems that Father J van Boven subsequently was at Roermond, Netherlands.
{W L Brown signature}William L. Brown, of Cornell University and a major ant taxonomist of the modern era, with D.E. Brown, in Ivory Coast, in (January) 1963;
William H. Gotwald Jr., a specialist in Army Ants, of Utica College of Syracuse University, in several countries in the early 1970s, sometimes accompanied by R. Schaefer;
V. Mahnert & J.-L. Perret (a mammalologist) in 1980 in Ivory Coast, having collected in Kenya in 1977;
E.S. Ross (of the California Academy of Sciences) and R.E. Leech in several countries, mostly in East Africa in 1957-58;
E.S. Ross and K. Lorenzen made collections in several countries in 1966;
I. Löbl collected in Ivory Coast, in 1977;
Neal A. Weber, of Swarthmore College and the American Museum of Natural History, himself an ant taxonomist, collected in Chad, at Hout Mbomu, around 1950. Brown (1952) described Weber as having made "two extensive tours in Africa". Weber (1942a, 1943c) published his own findings from a study of the Imatong (or Lolibai) Mountains of south-west Sudan. Much of the ant fauna he described is of the Congo Basin faunal group.

The early 1990's saw a resurgence in survey and collecting activities. The results, however, have been slow to appear taking until the past two years (2000-onwards) to appear in formal publications). The primary teams are -
R. Belshaw in Ghana, in 1993, in a collaborative project with Barry Bolton of the (British) Natural History Museum;
Allan Watt, Nigel Stork and colleagues studying the impact of tree plantation establishment in southern Cameroun;
the superb series of studies centered around the Campo Forest of southwest Cameroun, led by Alain Dejean.

Since I resumed active work on ants, initially with this website, I have been asked to help with identification of specimens collected in diverse research studies - the chimpanzee projects in the Mt Nimba area of Guinea/Ivory Coast; the Wolbachia and ant-mutualism studies led by Doyle McKey in Cameroun; and others.  Now (2012) this list is considerable and can be accessed from the Contents page

A final thought on the diversity of collectors and taxonomists has to be that their frenetic activity appears to reflect the patriotic fervour of the heady days of Imperial competition which ultimately climaxed in the Great War of 1914-1918. This can be seen from Bolton's new catalogue, where the extensive lists of junior synonyms and "subspecies" show a vast proliferation of "species", suggesting that the taxonomists acted with little regard to whatever their contemporaries in the museums of other Empires or loyalties had done or were doing. Presumably like the modern pre-occupation with publication of small papers to ensure a flow of grants, the kudos of expeditions yielding many "new species" proved irresistible. Something of this proliferation of names can be seen in the "taxonomic name indices" which I have prepared as part of this guide.

© 1998-99, 2002, 2003, 2009, 2012 - Brian Taylor CBiol FSB FRES
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