The Ants of Africa
Contents - Pseudomyrmecinae

Genus Tetraponera F Smith (1852: 44) (includes previous Pachysima and Viticicola).

Diagnostic Features - Slender elongate ants with relatively short legs. Clypeus sometimes produced into a spine or armed with a row of teeth or with a crenulate anterior margin. Either petiole alone with a ventral process or both segments without ventral processes.

F Smith's (1852) genus definition is at {original description}. [Smith, F. 1852. Descriptions of some hymenopterous insects captured in India, with notes on their oeconomy, by Ezra T. Downes, Esq., who presented them to the Honourable the East India Company. Ann. Mag. Nat. Hist., (2) 9, 44-50]. Roger's (1863a) definition of Sima is at {original description}.

All are arboreal species and very active, with characteristic rapid, jerky movements, and abrupt changes of direction. The previous Pachysima and Viticicola, were characterised because of their habit of living inside plant stems, sometimes as commensals, Pachysima in plants of the genus Barteria and tending large coccids on the plants. Hölldobler & Wilson (1990, page 535) describe the protection provided by Tetraponera (as Viticicola) tessmanni, which live only in stems of the liana, Vitex staudtii Guerkel; and by Tetraponera (as Pachysima) aethiops for Barteria fistulosa in Zaïre; they also list the incidence of Tetraponera ledouxi as a temporary parasite of Tetraponera anthracina (page 438, and Terron, 1969); and how one species, Tetraponera unidens (as nasuta), uniquely has a major caste with unknown functions (Terron (1971).

Earlier, Bernard (1952) remarked how all genus members inhabit hollow stems or dead wood and some support specific plants. From Africa, he reckoned there were some 30 species, with many still to be found.

Ward (1990: 449, 470) covered Tetraponera and confirmed earlier suggestions of the synonymy of Pachysima Emery and Viticicola Wheeler, also the non-African Parasima and Sima. The synonymy within Tetraponera is recognised also by Bolton (1994, 1995) but now Ward & Downie (2005), using DNA analysis appear to find differences that might substantiate the early genera as having merit.

{short description of image}Ward & Downie (2004) made several analyses of the phylogenetic relations within the Pseudomyrmecinae, of which Fig. 2 is an example. Morphological features that they considered useful were (w = worker, q = queen) - the angulate surface of the mandible above the trulleum (w), reduction in mandibular teeth (w, q), and the narrow notchlike cleft on the distal margin of the labrum (w, q). Non-African species listed are rufonigra (India), pilosa (Borneo), grandidieri (Madagascar), allaborans (Sri Lanka), morondaviensis (Madagascar), nigra (India) and punctulata (Australia). The possible separation might be into Pachysima for the rufonigra group; Tetraponera for the T. nigra group (represented by the bottom couplet); and, Sima Roger "for some fraction of the remaining species". I note that Ward & Downie have separated caffra and natalensis, whereas Bolton (1995) has caffra as a junior synonym of natalensis.

Ward (2006) continued his publication of studies of the genus mambers. In this latest part, he has established five monophyletic species groups covering all the Afrotropical and Malagasy species. These groups, for which he gave a key (adapted below), are rufonigra-group, ambigua-group, allaborans-group, grandidieri-group (Malagasy only] and natalensis-group. He also offered a revision of the T. ambigua-group. The paper can be seen in full - WARD, P.S. 2006: The ant genus Tetraponera in the Afrotropical region: synopsis of species groups and revision of the T. ambigua-group (Hymenoptera: Formicidae). Myrmecologische Nachrichten, 8, 119-130

NOTE - I am not convinced that Ward's interpretation of the T. ambigua-group is correct. My species key, below, therefore, does not wholly support his revisions. The availability of type images now (April 2015) appears to show I am correct.

Ward designated one new species, Tetraponera parops Ward, from East Africa. He was hesitant to treat it as a new species but I think his designation is correct. However, I have specimens of Tetraponera unidens (Santschi) new status that have the lateral rough patches on the occiput used by Ward as a primary distinguishing character for parops. Ironically, Tetraponera parops actually seems close to the original description of ambigua. The further specimens from Tanzania support the separation of parops, which is distinctly smaller than the related species.

With fresh specimens from Gabon, I regard Tetraponera angolensis (Santschi) as a clearly distinct species.

Under the blanket species Tetraponera ambigua (Emery), Ward states "mesonotum weakly convex and not separated from anterior margin of propodeum by prominently raised metanotal spiracles". He appears unaware that the drawing of occidentalis by Stitz (1917) and the description of angolensis by Santschi (1930b) have these spiracles as prominent.

Tentatively, I have separated occidentalis (ambigua race erythraea var. occidentalis Stitz 1917: 336; Menozzi, 1934: 154, worker) from Algeria to subspecies of Tetraponera erythraea (Emery), as having a wider, oval postpetiole. Moreover, if occidentalis is a genuine subspecies of ambigua, the distribution in North Africa is strange; Menozzi (1934) wrote of the finding in southwest Algeria, plus his record of its collection in Libya (Cirenaica, by Professor L di Caporiacco) as being of the form typical of southern Africa. My co-author on Ants of Egypt, Mostafa Sharaf, collected a specimen that clearly shows the distinctiveness of the North African species.

Ward's drawing of ambigua shows roughened patches on the lateral occipitum and prominent erect hairs on the sides of the head. Santschi's description of angolensis has no roughened patches on the occipitum, no lateral erect hairs on the head, indeed very few erect hairs at all, a more rounded propodeum amd a lower petiole. It is distinctly smaller than the specimen in Ward's illustration, for which he gave no origin, and has the darkened posterior to the gaster mentioned by Santschi, plus the narrow frontal area shown by Santschi. There appear to be five distinct teeth on the mandible as opposed to four for Ward's ambigua (also in Arnold's note on rhodesiana). I feel justified in elevating angolensis to full species status - Tetraponera angolensis (Santschi).

As Ward states he examined syntype workers of angolensis and erythraea (but not occidentalis) I surmise that he followed the style of Bolton, in "lumping" the ancient "varieties, subspecies and stirps", mostly attributable to Forel and Santschi. The converse, however, is that, at least by Bolton, other species have been erected on quite small characteristics. My experience from reading all the source papers and with a wide range of fresh material is that Santschi was remarkably accurate in separating his "stirps" but was influenced by his mentor, Forel, and so was reluctant to designate new species.

I have mentioned Tetraponera unidens above, Ward (1990: 489) gave the "current nominal combination" T. ophthalmica Emery (1912: 98; from Cameroun), Tetraponera ophthalmica tenebrosa Santschi (1928: 61; from Zaïre) and Tetraponera ophthalmica unidens Santschi (1928: 60; from Zaïre); with the annnotation that he had examined specimens. I am confident from the original descriptions, my own drawings of a Nigerian specimen and fresh specimens from Cameroun and Congo, that it is correct to separate Tetraponera unidens and Tetraponera tenebrosa off from Tetraponera ophthalmica (Emery).

Fresh specimens from Tanzania, have revealed Tetraponera bifoveolata (Mayr) to be like unidens in having a major worker. In both species this form has a curiously elongated head.

Grouping defined by Ward, with additions by myself (BT)
1 {Tetraponera aethiops}Large species, HW > 1.70, distance between frontal carinae [MFC] > 0.35; head with 3 distinct ocelli; petiole with prominent recurved antero-ventral tooth
aethiops & latifrons
T. rufonigra-group
-- Smaller, HW < 1.70, MFC < 0.25; head with 0-2 distinct ocelli, or, petiole with small simple antero-ventral tooth, or, both 2
2 {Tetraponera prelli}Pronotum with sharp lateral margination, continuing backwards to propodeum and petiole; overall appearance matt; mesonotum 2-3 times wider than long and semicircular in dorsal view; petiole with notched posteroventral margin
andrei, angusta, anthracina, caffra [Ward list; ssp of natalensis], capensis, emacerata, ledouxi, lepida [BT], mocquerysi, monardi, natalensis, poultoni, prelli, schulthessi & triangularis
[former Sima] T. natalensis-group
-- Pronotum, propodeum and petiole usually with rounded lateral margins; if margin distinct then shiny; mesonotum about as long as wide and ovoid or subrectangular in dorsal view; petiole without a posteroventral notch 3
3 Tetraponera ophthalmicaIn full face view, large eyes with long axis angled inwards anteriorly; upper half of mesosternum sparsely pubescent; workers dimorphic with distinct soldier caste
ambigua, bifoveolata [BT], encephala [BT], erythraea [BT], maculifrons [BT], ophthalmica, parops and unidens [BT]
T. ambigua-group
-- {Tetraponera braunsi}In full face view, eyes with long axis vertical; upper half of mesosternum densely pubescent; workers monomorphic; all but zavattarii have a distinctive face shape with the occciput arcuate commencing from just posterior to the eyes
braunsi, claveaui, clypeata, emeryi, gerdae, liengmei, mayri, penzigi, scotti, tenebrosa [BT], tessmanni & zavattarii
T. allaborans-group

Provisional key to workers of the Tetraponera of Africa, derived in part from Arnold (1916: 170), who gave a key to the then known species (as Sima); this is at {original description}. OI = (eye length/head width) X 100

¤ Tetraponera encephalaKnown only from the queen; TL 5.5 mm, testaceous red, alitrunk brown; quite shiny; possibly synonymous with bifoveolata.
Senegal - encephala
¤ Tetraponera gerdae queenKnown only from the queen; wholly black except for lighter appendages; TL 4.5 mm; the profile, narrow gaster and overall size suggest this may be the queen of bifoveolata Tanzania - gerdae
¤ Tetraponera lemoultiKnown only from the queen; TL 10 mm, close to T. mocquerysi; mostly testaceous red
Congo - lemoulti
¤ Known only from the queen; TL 6.3 mm; red brown; no images
Cameroun - mayri
1 {Tetraponera aethiops face}Thickset species; frontal carinae set far apart; 3 ocelli; eyes relatively small 2
-- Smaller, more slender, most with frontal carinae close together 4
2 {Tetraponera tessmanni}Small species, TL 3.0-3.5 mm, eyes set forward of midlength of head; amber yellow
. {Tetraponera tessmanni} (former subgenus Viticicola) - Congo Basin - tessmanni
-- Large stout species, eye set towards rear of head (former subgenus Pachysima) - 3
3 {Tetraponera aethiops pedicel}TL 9-10 mm, shining jet-black; petiole and postpetiole in dorsal view ovoid; noticeable ventral processes on both the petiole and postpetiole but these are simple triangles .
. {Tetraponera aethiops} Panafrican (in Barteria) - aethiops
-- {Tetraponera latifrons}TL 7-8.5 mm, generally more slender; clypeus with a conspicuous fringe of yellow ciliary bristles; petiole and postpetiole in dorsal view more angular (rather than ovoid); petiole and postpetiole with rearward hooked processes, quite pronounced on petiole; black and shining Cameroun & Congo Basin - latifrons
-- Smaller, more slender, most with frontal carinae close together --
4 {Tetraponera ledouxi}Pronotum with distinct sharply margined anterior angles; 2 or 3 distinct ocelli (former subgenus Sima) 5
-- {Tetraponera zavattarii}Pronotum with anterior angles rounded (may be laterally margined); ocelli rudimentary or absent .
. {short description of image} (former subgenus Tetraponera s.s.) 14
-- --Former Sima - usually with alitrunk and sometimes petiole dorsum with distinct lateral margins --
5 General colour dark brown to  black 6
-- General colour orange brown or paler 12
-- Black to blackish species --
6 Tetraponera andreiTL 4.5 mm; smallest of the group; similar to but smaller than mocquerysi with rectangular head longer than wide; colour primarily black but dull; mandibles coarsely striate; almost no erect pilosity and very fine sparse pubescence
Mozambique - andrei
-- TL > 5 mm 7
7 Head with near straight occiput 8
-- Head with rounded occiput, at least the lateral angles; TL at least 7 mm 11
8 TL > 5 mm < 7 mm 9
-- TL at least 7 mm 10
9 {Tetraponera emacerata}TL 6.0 mm; head longer, about a quarter longer than wide, parallel sided; posterior border more strongly impressed (prelli, mocquerysi and anthracina being enlarged in the posterior one-third and with convex sides). .
.9 {Tetraponera emacerata} Panafrican - emacerata
-- {Tetraponera mocquerysi}TL 6.0 mm; head rectangular (type apparently straighter sided than biozellata) HL > HW; anterior margin of clypeus weakly arcuate with only fine crenellation; body with yellowish pubescence, almost no erect hairs; head and thorax densely and finely reticulate, semi-matt; entirely black expect extremities; OI 58 Panafrican - mocquerysi
10 {Tetraponera triangularis}TL 7.5 mm; black; erect whitish pilosity fine, absent on thorax and base of gaster; subopaque, very finely and densely reticulate, serrated puncturation on head and thorax; head as mocquerysi but eyes larger and with three ocelli; anterior of clypeus a short rectangular lobe, margin weakly concave (lobe weakly indented in mocquerysi); OI 52

This and T. anthracina (below) appear to be identical. If so, the latter, published 23.ii.1910, takes precedence over this, published (31).viii.1910 (dates in Bolton, 1995)

Central African Republic, Equatorial Guinea & Nigeria - triangularis
-- {Tetraponera anthracina}TL 6.5-7.0 mm; colour deep red-brown; shiny but with dense pubescence; head near rectangular, OI ca 48, 2 ocelli West Africa & Congo Basin - anthracina
-- Head with rounded occiput, at least the lateral angles; TL at least 7 mm --
11 {Tetraponera ledouxi}TL 7.1-7.9 mm; colour black, with extremities brown; cuticle smooth and not finely puncturate as in anthracina; whitish pubescence on all the body; on the head and thorax less dense but longer than on anthracina; OI 53 Cameroun & Central African Republic - ledouxi

{Tetraponera lepida}TL 6.5-7.0 mm (9.3 mm for CRIN form); anterior margin of clypeus near straight with 3-5 strong crenellations (teeth); abundant erect hairs all over; colour dark red-brown to black, shiny; OI 60 Congo & Nigeria - lepida stat. nov

Tetraponera poultoniTL 7.5 mm; generally as triangularis but anterior margin of clypeus with five small teeth, the median tooth broader and blunt; only two small ocelli; petiole with sharp dorsal borders and ventral processes of a small anterior forward-directed tooth and a larger posterior backward-directed tooth; margination of alitrunk "fading out" on the propodeum declivity; black, apex of gaster segments yellowish brown; OI = 52 Uganda & Kenya - poultoni
return to key-- General colour orange brown or paler --
12 Tetraponera n sp fohlenHW < HL; TL ca 4.9 mm; colour yellow-orange; small near globular postpetiole, with a well defined metanotal groove Namibia - n. sp. fohlen
-- Alitrunk without an impressed metanotal groove 13
13 {short description of image}Head sub-square; TL 5.6 mm; alitrunk dorsum feebly convex without any impression of the sutures; head and alitrunk densely puncturate and subopaque; brown but with many areas lighter, notably the yellow-rust dorsum of the first gaster segment Tanzania - prelli

{Tetraponera natalensis}TL 6-8 mm; colour yellow or brownish yellow
Note: the type form is a queen
. {Tetraponera natalensis}bulawayensis worker Tanzania southwards to the Cape of Good Hope - natalensis

{Tetraponera capensis}TL 7.0 mm; dark reddish yellow South Africa - capensis
-- {Tetraponera caffra}TL ca 7.5 mm; narrow head, broad posterior to petiole in dorsal view; anterior margin of clypeus dentate; yellow-brown apex of gaster darker
South Africa, Angola - caffra
-- Tetraponera schultessi queenTL 9.0 mm; more robust than natalensis; head wider; alitrunk dorsum flatter; declivity of propodeum twice as long as dorsum; colour yellow-rust
Queen TL ca 11 mm
Mozambique & Zimbabwe - schulthessi
-- {Tetraponera angusta}TL ca 7.0 mm; narrow head, narrow petiole in dorsal view; anterior margin of clypeus dentate; reddish ochreous Zimbabwe - angusta
return to key-- Former subgenus Tetraponera - no more than anterior of pronotum with any lateral margination -
14 {Tetraponera nasuta}Dimorphic, major TL 4.6-4.9 mm; minor TL 4.1-4.7 mm; head cylindrical, clypeus of major (and queen) with distinctive elongated median anterior process; both petiole and to a lesser extent, postpetiole distinctively low and elongated; generally golden-yellow with at least part of the gaster darker .
. {Tetraponera unidens morphs} West Africa & Congo Basin - (nasuta new status) unidens
-- Most known only as monomorphic - note 2007 finding of a major for bifoveolata, anterior clypeus without prominent median process, although smaller processes may be present 15
15 {Tetraponera clypeatus}TL 3.6-4.0 mm; anterior margin of clypeus with two pairs of sharp processes or teeth; black South Africa - clypeata
-- Anterior margin with no or less than four teeth 16
16 {Tetraponera braunsi}Tetraponera braunsi TL 4-4.4 mm; clypeus with three teeth on anterior margin; black with appendages reddish or lighter
queen with the median tooth very small or absent, TL 7 mm
South Africa - braunsi
-- Tetraponera braunsi durbanensis TL 3.8-4.0 mm; more slender; clypeus with three teeth on anterior margin; yellow with head and alitrunk South Africa - (braunsi) durbanensis
-- Anterior clypeal margin with no or less than three teeth 17
17 {Tetraponera monardi pedicel}Colour black or brown group - pear-shaped petiole 18
-- Colour yellow-brown 22
-- Dark brown to black --
18 {Tetraponera monardi}TL 11 mm; head rectangular
Angola - monardi
-- TL < 11 mm 19
19 Head elongated-rectangular 20
-- Head elongated ovoid 21
20 {short description of image}TL 3.4 mm; elongated head but eyes less than one-third of head length; postpetiole rounded in dorsal view; postpetiole similar to the petiole but more spherical in profile and dorsal views (latter is oval in penzigi). Separable from liengmei and penzigi by its elongated head and the postpetiole shape. Senegal & Gambia - claveaui
-- Tetraponera penzigiTL 3.1-4.7 mm (continua smaller); clypeus angular in the middle, postpetiole ovoid in dorsal view; black South and East Africa savannah - penzigi

{short description of image}TL 3.4-3.7 mm; head elongate-rectangular 1.5 X longer than wide; clypeus with two longitudinal carinae ending in minute teeth; the eyes are larger than claveaui but still relatively small; very shiny brown-black Mozambique - liengmei
-- head elongated ovoid --
21 Tetraponera scotti worker headHead oblong-oval HW 0.53 HL 0.69 shining black; queen TL 6 mm. Anterior margin of clypeus on the worker recessed arcuate.
Ethiopia - scotti
-- {Tetraponera zavattarii}TL 4.0-4.5 mm; petiole distinctly domed; black Ethiopia - zavattarii
. {Tetraponera tenebrosa}TL ca 3.8 mm; HL 0.9 HW 0.56 SL 0.35 EL 0.40 PW 0.30; CI 60 SI 67 OI 68
Colour dark brown; pronotum but not rest of alitrunk or petiole margined and without distinct ocelli, characteristic of former genus Tetraponera members. Anterior margin of clypeus weakly concave without any dentition; mandibles dull but not striate. Over shape narrowly elongate; pedicel segments in profile low and weakly domed, dorsum of petiole narrow and of postpetiole trapezoidal, anterior about half width of posterior.  Distinctive quite dense layer of decumbent golden pubescence on the postpetiole and gaster; overall sculpturation of dense fine spiculation.
Congo Basin - tenebrosa
return to key-- Yellow or reddish yellow --
22 Tetraponera emeryiTL 4.5 mm; pronotum not margined, clypeus with five teeth on the anterior margin, the outer two smallest; eyes near 1/3 of head length; reddish-yellow
South Africa - emeryi
-- Pronotum at least laterally margined; yellow or reddish yellow, head and thorax not smooth and shining 23
23 Clypeus emarginate medianly, evenly convex 24
-- Clypeus not emarginate medianly; rectangular and/or with a feebly tridentate margin 27
24 At least gaster with moderate to abundant pubescence 25
-- Pubescence absent or very sparse 26
25 {short description of image}TL 3.8-4.2 mm; with abundant pubescence, especially so on the gaster; reddish yellow East Africa north from Mozambique - bifoveolata
. {short description of image} major of bifoveolata - first record
-- {Tetraponera ophthalmica}TL 4.8 mm; colour yellow, shiny almost transparent cuticle. Fine, dense pubescence restricted to gaster. Eye length 0.39 mm (0.45 of HL). No or only small ocelli.

TL 4.08 mm, HL 0.86, HW 0.53, SL 0.39, PW 0.36; CI 60  SI 80 EI 46
Western Africa - Cameroun to Angola - ophthalmica
26 {short description of image}TL ca 6 mm; mesonotum with strong sutures all around; mesonotum shallowly domed; petiole relatively more domed; apparently without significant pubescence Djibouti & Egypt - maculifrons
-- Tetraponera angolensisTL ca 4 mm; without significant pubescence; posterior of gaster darkened Angola & Gabon - angolensis
-- Clypeus not emarginate medianly; rectangular and/or with a feebly tridentate margin --
27 {Tetraponera ambigua}TL 4.5 mm; anterior margin of clypeus produced medianly into a rectangular lobe; frontal carinae set quite far apart; relatively short, sparse erect pilosity on alitrunk dorsum and head; transition from dorsum to declivity of propodeum near right-angled
Southern & Eastern Africa - ambigua.
. Anterior margin of clypeus not lobed, feebly tridentate in the minor and distinctly so in the major ambigua ssp rhodesiana
-- TL < 4.5 mm; frontal carinae set quite close together; postpetiole ovoid; erect pilosity restricted; 28
28 Tetraponera erythraeaTL 4.0 mm; mesonotum with a raised angular profile; eyes closer to posterior border of head; piligerous punctures on head stronger giving a coarse uneven appearance; femora with a brownish dorsal edge; abundant erect hairs distinctive on head and dorsal alitrunk
Yemen (Aden) & Algeria - erythraea
-- Tetraponera paropsTL ca 3 mm; mesonotum with concave profile; occiput with lateral patches of short bristly pilosity, pronotum with paired erect hairs East Africa - parops


Tetraponera species A

Collected in Ghana by Bigger (1981a) at CRIG, 30 workers by pkd from the canopy of four of five plots of Amelonado cocoa and 1 from the ground under the cocoa.

Tetraponera species B

Given in Room's (1971) report as being found on the cocoa canopy on a single farm in Ghana.

© 2007, 2008, 2009, 2010, 2011, 2013, 2014, 2015, 2019 - Brian Taylor CBiol FRSB FRES
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