|The Ants of Africa
First Edition - 1998
The past forty years have seen greatly increased interest in tropical forests and in the crops grown in once-forested areas. Among the most abundant animals in the tropics are the ants (Hymenoptera: Formicidae). For instance, Entwistle (1972) related findings from Ghana (by Leston & Gibbs) and from Nigeria (by Booker) which showed that as many as 87% of the total number of individual insects found on cocoa are ants. This, coupled with the colonial lifestyle of many species and their activity as predators, surely makes them major components of both the terrestrial and arboreal ecosystems.
Tree crops, especially cocoa, are of great economic significance in the forest zone of Nigeria and the ants which can be found on all the trees profoundly affect both pests and diseases of the crops. As members of a team studying Black Pod disease of cocoa, caused by one of two fungi, Phytophthora palmivora (Butler) Butler, or the closely related Phytophthora megakarya Brasier & Griffin, my staff and I made extensive collections and surveys of ants at the Cocoa Research Institute of Nigeria (CRIN) and elsewhere in the cocoa growing areas of western Nigeria. Our findings on the distribution and inter-specific relations of the ants and their role in the annual Black Pod epidemic appeared in the Final Report of the Project (Gregory & Maddison, 1981) and in specific research papers (Taylor, 1977; Taylor & Adedoyin, 1978). Although the number of ant species which were both abundant and involved in Black Pod disease was found to be quite small, we encountered some 100 species foraging or nesting on cocoa trees and another 50 species in cocoa plantations or in the surrounding forest. This contrasted greatly to the report by Booker (1968), who collected only 19 ant species in repeated surveys of the tree canopy of two blocks of cocoa at CRIN, despite using regular pyrethrum knockdown (pkd) for nearly two years, and the list of 27 species given by Eguagie (1971).
The problem of identifying tropical insects is widely recognised and ants are no exception. Strickland (1948), whose bionomic studies in Ghana were among the earliest on the ants of West Africa, described how -
"Nearly 50 species of ant have so far been taken attending P. njalensis colonies in the field" but "Only nine of the species have so far been identified".
In Nigeria, during 1974-76, the output from our daily surveys and monitoring work included a bewildering array of small brown-black moving objects, ants, some or all of which might be playing a role in the transmission of cocoa black pod disease. In order to satisfy the crucial need to rapidly separate the numerous ants, I started to make small annotated illustrations of the worker and soldier castes of each species. Two slices of good fortune combined, however, to ease the task of naming the species. First, CRIN at that time had a moderate collection of ants which had been curated by Barry Bolton, now one of the world's great ant taxonomists. Secondly, Bolton, by then at the Natural History Museum, also had just published keys to the genera of West African ants (Bolton 1973a). The combination enabled me to separate out all the specimens to genus level. The labels on CRIN species and the first of Bolton's revisionary studies (Bolton, 1969, 1970-71, 1971a, 1971b, 1972, 1973, 1974a, 1974b, 1975a, 1975b) enabled me to assign specific names to a good proportion of the taxa we collected. In many instances, separating them to species level still meant either using the code letters or numbers of specimens in the CRIN collection, some of which we did not encounter, or devising further code designations. The major problem at that time, however, was the poor state of taxonomy, especially at the species level, and over 50 of the species could be labelled only as "species 1, species 2, species A, species B, etc.", my additional species were labelled "species T¹, species T², and so-on". The literature on research in Ghana, which is referred to extensively later in the text, shows similar code numbers, etc., for many species, probably reflecting the fact that CRIN originally was an outstation of the main institute in Ghana (first known as the West African Cocoa Research Institute; now The Cocoa Research Institute of Ghana, CRIG). The source of some of the codes, moreover, may have lain elsewhere, especially the British Museum (Natural History). For instance, Kemp (1951), reporting work on Pheidole in (now) Tanzania, listed two species as Pheidole species A and Q, with the identifications being attributed to H. St. J. Donisthorpe of that Museum. The latter also had written two short papers which dealt with ants collected by workers based in Ghana (see Chapter 2). The abundance of such coded "forms" can be seen by a quick glance at the two Taxonomic Name Indices which deal with Forms A to N and Forms O to Z.
The initiative, in 1975, to begin publication of a CRIN
Bulletin encouraged me to turn my illustrations into a proper field
guide which could be used by anyone needing to identify ants of the
area. The original monograph, Ants of the Nigerian Forest Zone,
was written in early 1976, and appeared in five parts (Taylor 1976,
1978, 1979, 1980a, 1980b). Advances in ant taxonomy since that time
have been considerable and an update has become both desirable and
possible. Extensive revisionary work, mostly by Barry Bolton, has given
proper names to many of the species and synonymized a number of others.
[November 2015 - the monograph in pdf form, together
with an up-to-date list of the species can be seen on the linked pages
from Ant Guide update.]
Even with all the taxonomic work, however, several important, perhaps the most important, genera remain inadequately classified. The most urgent needs are for revisions of Camponotus, Crematogaster and Pheidole. Wilson (1976) had speculated that the three genera had 641, 274 and 391 members respectively. The three genera form what Hölldobler & Wilson (1990) termed the crux myrmecologorum, but they noted that the extreme example, Pheidole, is thought to have more than 1,000 members, two and a half times as many as was thought only 14 years earlier. Thus, even very recent reports of ant-related research in West Africa, such as Campbell (1994), have had to resort to using the name Pheidole megacephala to cover several, perhaps as many as eighteen, morphologically and behaviourally similar taxa, and Belshaw & Bolton (1994b) listed eleven undetermined species. The magnitude of the taxonomists' task was illustrated by Bolton (1981b), when he described the then state of knowledge of the subfamily Myrmicinae. There were some 43 genera recognised from sub-Saharan Africa; of which 29 genera had 1-10 species, 9 had 10-40 species and 5 had more than 40 species. Of the last, Tetramorium, Pheidole and Crematogasterwere thought to have well over 100 species each in the region. His own revisionary studies had enabled him to recognise 176 species of Tetramorium (Bolton, 1980) and 33 species of Triglyphothrix (Bolton 1976, the genus was synonymized subsequently with Tetramorium, Bolton, 1985). Recently, Wilson, together with Bill Brown of Cornell University, announced that a start is being made on the task of preparing a monograph on Pheidole (Hölldobler & Wilson, 1994). [In the event, sadly, Brown died in 1997 and Wilson's Pheidole revsion covered only species from The Americas (apart from two tramp species)].
Taking the foregoing, I decided to adopt a two-pronged approach. This was to compile the fullest possible taxonomic catalogue, illustrated by my numerous drawings of species from Nigeria, and to pull together all useful knowledge of the biology and strategic importance of ants in the Forest Zone and the adjoining Guinea Savannah Zone of West Africa. Although my illustrations cover only some 40% of the known, described species, they do show almost all the relatively common species and at least one representative of most of the genera. Perusing the rather battered copy of Wheeler (1922) held by the Royal Entomological Society led me to decide to incorporate relevant drawings (originally by Mrs Helen von Ziska, and denoted by W '22) and illustrations both to complement this text and to further enhance their value for posterity. To aid those who wish to look further into the literature and to identify the many other species; wherever possible, I have drawn on modern publications to provide Keys to genus members. Then I have added brief notes on the particular distinguishing features of each of the listed species, including indicating illustrations in the reference literature. If available, information on habitats, nest sites, etc. is provided. Unless indicated, the information comes from the collections by my staff and myself. The emphasis remains on ants in cocoa farms simply because of the paucity of other research.
A final justification, if any is needed, for this book is to bridge the gap between anglophone and francophone entomologists. While in Nigeria, I met entomologists from Ivory Coast and Cameroun who were working on cocoa problems but I have to admit to being ignorant of the work under way at Lamto in the Ivory Coast, although that was in its earlier days. My contemporary compatriots, such as Mike Bigger and Colin Campbell, who were studying aspects of cocoa entomology in Ghana and published a little after my main papers, make no reference to work published in French, excusable perhaps as they were not dealing specifically with ant mosaics or general biology. In converse, moreover, Lévieux (1983) wrote how little or nothing had been written on the biology of Myrmicaria, whereas Myrmicaria striata featured in my papers (notably Taylor & Griffin, 1981).
Receipt (in mid-1999) of a copy of Francis Bernard's treatise on the ants of Mt. Nimba, Guinea, over 100 pages of fascinating text, showed again how much the English-speaking world has failed to appreciate texts written in other languages. I have endeavoured to do justice to Bernard's work and, thus, have included (my) translations of his descriptions of previously new species and of many of his commentaries. He examined several of the major museum collections of ants and I have taken his naming of species as valid, especially as, obviously, that is better than the code designations I have given for many unassigned forms. Given my own difficulty in obtaining a copy of his treatise, I have also incorporated Bernard's illustrations (distinguishing them by "B'52").
For those interested in ants there is no better starting point than the magnificent book The Ants by Bert Hölldobler and Edward O. Wilson (Hölldobler & Wilson, 1990). An easier read but no less fascinating is its companion book Journey to the Ants (Hölldobler & Wilson, 1994). Contemporaneously, the narrower but fascinating aspect of ant morphology and taxonomy has been well served by Barry Bolton (1994), whose great book Key to the Ant Genera of the World has 522 superb photographs, taken with a Scanning Electron Microscope by Laraine Ficken, and showing almost all recognised genera. Complementing that work now is his comprehensive A New General Catalogue of the Ants of the World, in which one finds a full list of what, at 31 December 1993, were 16 subfamilies, 296 genera, and 9536 species (Bolton, 1995). More than that, however, he gives extensive coverage of subspecies, junior synonyms, junior homonyms and unavailable names. The whole is completed by references to all the source papers for the names and many ancillary papers. All three have proved very useful to me in writing this text.
Coming across the website of the Japanese Ant Color Image Database filled me with admiration for the outstanding quality of the colour images. Importantly, the "Database" includes several "tramp species" found in West Africa, plus some genera not pictured on my site. Thus, I have added links to my site which, writing this on 1 January 2000, show how the internet really can open up the guarded world of the museum specialists.
This second version of the text rises from my plan to update it as further source publications become available. Presently the main gaps lie in the difficulty of access to work published from Universities in francophone Africa, but the amount of information embodied in this version justifies making it available as it stands. With the generous gift from Professor Xavier Espadaler Gelabert, from Barcelona University, of the extensive report by Francis Bernard (1952), I have been able now (mid-1999) to move forward to this second version. In addition to illustrations from Bernard's work, I have replaced the ".jpg" format species drawings by ".gif" versions, thus, giving clearer white areas. The "Frontispiece" and "Many Faces...." sections were added after the original web posting and the analyses in "Biodiversity and Niches" are updated. New also are the separate chapter sections on Guinea and the ecological findings by Bernard & Lamotte.
Third edition - 2001
With the receipt of numerous specimens from the Doyle McKey team (see below), I have been able to add a considerable amount of information and scanned images of the species. The specimens include several species previously recorded only from the Congo Basin and, so, I have decided to expand the overall list by adding records of species known from that area.
Note From the 2001 edition, the reader will find a number of "missing" illustrations - these are not lost relative to the earlier editions but indicate images in published works not available for copyright purposes. This availability, I hope, will be improved before too long. I am also continuing the process of gathering original publications from nearly a century ago and so adding to the text and images.
Fourth edition - 2002
The catalogue now includes species known from the Congo Basin, including information from the writings of F. Santschi. Other fresh material is from a paper by Cedric Collingwood and a very interesting set of specimens of Driver Ants sent to me by Tatyana Humle, of the University of Stirling, who is studying chimpanzee behaviour, especially the use of tools, such as for "ant-dipping".
Fifth edition - 2003
For each species a link has been provided to the appropriate page in the Hymenoptera Name Server. Continuing the policy of ongoing development the addition of historical and bionomic material continues. The spreadsheet catalogue (Microsoft Excel 2000 format) now has links between all species and the appropriate page in the main catalogue.
Sixth Edition - 2004
More by accident than design but made possible through the efforts led by Donat Agosti to get all the source publications on-line, I have embarked on an expansion to bring as much as possible of, at least, the taxonomic knowledge of ants from all of sub-Saharan Africa into this sixth edition. The "publication" will be progressive due to the vast amount of available information but I will place more and more on the site as time goes by. I began to translate original texts from French, Spanish or even Italian but I have no knowledge of German. Because that exercise will be very demanding on time, I have decided to create "file cards", e.g. , each of which contain edited versions from scans of the original texts and to link these from the summary descriptions. I have also created edited versions of illustrations from original papers, so that nearly all are in a uniform style.
Ragge (1980) expressed his preference for the term 'Afrotropical Region', rather than the long-held and, in my view, now misunderstood term 'Ethiopian Region', thus, supporting the view of Crosskey & White (1977) - a concept with which I fully agree.
Seventh Edition - 2005
The extensive work to collate information on all the described species from sub-Saharan Africa (inclusive of Sudan and Ethiopia) has been largely completed.
I have started to add cards summarising the distribution, as known from published sources, plus new material incorporated on this website. Thus, for each species, there will be a "distribution card" - . In due course, my intention is to add similar cards for, at least, summaries, of biological information, again using "file cards", e.g. .
Eighth Edition - June 2005
The near completion of the ant literature collection available
the Hymenoptera Name Server means that very little remians unaccessed
of the original descriptions and distribution records.
Further, it now has become possible to incorporate illustrations and other material from the numerous taxonomic papers written by Barry Bolton. For that kind permission was given by Prof. Quentin D. Wheeler, Keeper and Head of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD U.K.
Thus, there are now almost 3600 description "cards" covering all but 20 of 1819 species. For 721 species there are no known illustrations and for 34 species original illustrations remain unaccessed. Illustrations are on the website for 1064 species.
Ninth Edition - November 2007
This is a wholesale revision of the basic structure of the website. The central feature is that each species has its own webpage. The former comparative format with related species on a shared page has been changed but not lost as an attempt has been made to create useable keys for all subfamilies and genera with more than a small number of members. A core element of each key is the incorporation of thumbnail images. Each of the species pages is linked from the related key.
For all but two of the now recognised 1901 species, the original or translated descriptions are given either in the main text or on linked "TAXONOMY" cards; of which there are around 5000, covering the subspecies and synonyms as well as the type species. In constructing the keys, many of the original descriptions have had to be translated from non-English sources, most commonly French but also German and Italian, with a few from Latin.
The main caution that needs to be understood by the user is that the author has not sought out or accessed type material, other than where photographs of types have been made available on the internet. Thus, there may be instances where the identification of fresh specimens sent to the author ultimately turns out to be wrong. For the present the identifications should be viewed as "best guess" determinations, although in each case the original descriptions of the species and its parent species where the status has been changed from subspecies, variety or stirps, etc. have been read.
Tenth Edition - December 2008
This stems primarily from cataloguing the 500 or so species that I now have and, thanks to the generosity of Dr George McGavin and the Oxford Museum of Natural History, am able to arrange in proper insect cabinets.
The publication of revisions, e.g. Fernandez (2004), synonymizing Afroxyidris, Oligomyrmex and Paedalgus under Carebara; Bolton (2007) on Technomyrmex; and, Bolton & Fisher (2008) on Asphinctopone, are incorporated, at least, in part.
Additional and better images of, for instance, the Cameroun McKey material, and fresh images of specimens from Congo (Yves Braet & Eric Nzassi), the Central African Republic (Philippe Annoyer), Niger, etc (David King), bring the total number of species for which illustrations are available to around 1500 of the total of 1925 named and as yet unnamed species. Mike Lush also has contributed some photos from his http://sifolinia.blogspot.com/2007/10/gambian-ant-list.html web blog. There are photographs of about 862 species, over 480 being my originals. A result is the further use of images in keys. The number of species for which no illustrations are available is down to around 390. The known but as yet unavailable drawings is down to five but three of these have photographs. All of the published descriptions all have been accessed and almost all are linked from the appropriate species page.
Eleventh edition - 2013
The withdrawal of hosting by the
the need to
incorporate a clearer listing of all the many specimens sent to me, led
me to add tables of collection details for all that I have
identified and that will be deposited in the Oxford University Museum
of Natural History. The tables incorporate links to albums,
that show my diagnostic photographs of specimens,in effect providing a "Virtual Museum".
After much effort and hope for an alternative institutional host, I decided in August 2013 to personally meet the costs of hosting, initially on a two year contract. Once fully opened, I will ask the British Library Web Archive to update their holding.
Twelth Edition - 2015
After considerable time and a lot of work, I have completed the addition of photomontages of images of type specimens put on-line by the tremendous efforts of the Antweb.org programme, led by Dr Brian Fisher.
Of a total of some 2132 species, about 70 of which are yet to
be formally published in print, only 148 have no photographs but there
are drawn images of 90 of those, thus only 58 have no illustrations at
There are images of 1455 holotypes and of paratype or syntypes of the others.
For 840 species there are my images of fresh specimens, in many cases with examples from two or more locations.
From now on the main changes will be addition of new species,
incorporation of taxonomic revisions, or notes on such revisions if I
have doubts about them.
As they come my way, images of fresh specimens will be
incorporated, either on the species page or in the albums of my
Archived by The British Library
Guidance for users
Ultimately, my hope is for a full printed or DVD edition but use of a Web version, despite some limitations in the presentation of the many illustrations, should offer an accessible text for those in remoter global areas and with limited funding. It will also allow very rapid updating as further information, taxonomic revisions and the like come to hand.
The use of an electronic format gives certain advantages in the use of the text - by permitting cross-access through "clickable links", these have particular advantages in moving from the "Contents" page to individual sections, from sections to illustrations and so-on.
When using the "Keys to Species" the "clickable
links" enable rapid movement from couplet to couplet or on
to the individual taxon and, from either, a
The "Taxonomic Name Indices", which can be accessed from the "Contents" page and from all sections of the narrative Chapters, represent my attempt to alphabetically list all the published names which relate to the presently definitive species. Each name has a "clickable link" to the definitive species - and thus provides a quick access from published names - be they subspecies, junior synonyms, outdated names, or whatever. The indices include many names which probably have not been used in the geographical context of West Africa. The supreme example of a multiplicity of names under a single species can be found with Camponotus maculatus which, bearing in mind the poor state of all Camponotine taxonomy, has some 10 subspecies, 31 junior synonyms and 11 other "unavailable names" etc. - it has been found throughout Africa, right across Asia and into Australasia. When genus revisions and synonymization relate to West African and Congo Basin species I have listed the "original" genus name, but have not done so for ants from elsewhere.
In addition to the indexing of published names, I have attempted also to list all "forms" - a term which I use for the ants listed in many bionomics and survey reports simply by the Genus name followed by "species X", where X is a code letter or number or combinations of letter and number(s). In a number of cases the form is linked to a subsequently determined definitive species. Sadly, the taxonomists have not always indicated the "form" used by the original collector.
©1998-9, 2001, 2002, 2003, 2004, 2005, 2006, 2007,
2008, 2009, 2012, 2013, 2015 - Brian Taylor CBiol FRSB FRES
11, Grazingfield, Wilford, Nottingham, NG11 7FN, U.K.