taxonID	type	description	language	source
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	description	Figs 1 – 33, distribution map: Fig. 32; Table 1	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	type_taxon	Type species Eutermes (Cubitermes) minitabundus Sjöstedt, 1913.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	diagnosis	Diagnosis As in the other Cubitermitinae, the worker’s, soldier’s and imago’s guts are characterized by a mixed segment between mid and hind gut; four Malpighian tubules fused at base and by a blind diverticulum, or caecum, on the paunch of the hind gut (Noirot & Kovoor 1958; Noirot 2001) (Figs 2 – 3). The genus Isognathotermes is best and fully characterized by its worker’s EVA which takes two main forms and an intermediate form (workers of no other Cubitermitinae genera show such EVAs.): - in the fungifaber pattern (Figs 4, 6), the odd-numbered primary cushions (slightly longer than the even primary cushions) are frequently narrowed and crested in their downstream parts (the crests, however, are only visible if the PCs are twisted and viewed in profile), in some species, they are only slightly protruding or even not at all in I. acristatus sp. nov., the crested and distal parts of the PCs bear long curved or hooked bristles; the transition from the part with supporting bristles to the bulbous part is relatively gradual; see also Noirot & Kovoor (1958: 458). - in the finitimus pattern (Figs 5 – 6), the odd primary cushions are more robust, less or not at all narrowed; a rather swollen part (viewed in profile) bears numerous short, stiff, non-curved bristles and the distal part bears some long curved to hooked bristles; the transition from the part with supporting bristles to the bulbous part is relatively abrupt. - in the intermediate pattern (Fig. 6) the odd-numbered primary cushions have relatively small crests, are not swollen in their downstream part, and bear numerous slightly curved bristles; the distal parts bear some long curved to hooked bristles (it is closer to the fungifaber pattern than the finitimus pattern and occurs only in species of the fungifaber pattern, mainly I. severus and I. ugandensis). The secondary cushions are wide with a homogeneous spine scattering. The worker is also characterized by the shape and spinosity of its front legs: the fore coxa is carinated and bears two to seven spines on the ventral carina (Fig. 7); the fore trochanter bears three to eight spines. Mandibles as in the imagines. The soldier is dark-gutted (its gut contains some soil); it has a well-developed frontal gland and a frontal pore (fontanelle) on the frons surrounded by a tuft of setae; it has a deeply bifurcated labrum with straight, convex or, more generally sinuous sides (lyre-shaped labrum), almost always with subtruncated lobes ending in fine, whitish or translucent tips, and bearing three to ten large setae on each lobe; its EVA, however, is not as well characterized as that of worker, especially in the fungifaber pattern where it can be confused with a bilobatodes or a muneris pattern; in the finitimus pattern, the soldier’s EVA is generally recognizable (Fig. 6). Its fore trochanter and carinated fore coxa often bear some spines (but less than the worker’s). It has a long head (SHdL = 2.18 – 3.77 mm), a high head elongation index (SHdL / SHdW = 1.22 – 1.67), and the sabre-like mandibles are always shorter than the head (SMlL / SHdL = 0.66 – 0.92); antennae of 14 – 15 articles. All these characters taken together make it possible to recognise the Isognathotermes soldiers. The imago cannot be distinguished easily from other, related Cubitermitinae genera; it is always rather dark (C 6 – C 8), its head bears some large setae overhanging a generally dense mat of short bristles. Its meso- and metanotum generally bear some very fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 – 80 × (without any visible small setae in 6.5 % of the imagines examined), and frequently with three to four larger setae. The fontanelle is always small; antennae of 16 articles (but shortened in functional queens and kings). Mandibles with apical tooth longer than marginal teeth, three on the left mandible, two on the right. Legs: tibial spurs 3: 2: 2, middle tibia with two extra subapical spines. The enteric valve is weakly developed and can hardly be used for identification (Fig. 6).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	etymology	Etymology From Greek ΙΣΟΣ (isos, equal, even), γναΘΟΣ (gnatho s, mandible), and Latin termes (termite), the name Isognathotermes probably refers to the type species (see the chapter I. minitabundus), either to the resemblance between both mandibles or to the fact that their diameter varies little between both ends. Historical review The first mention in the literature of a termite that could be an Isognathotermes probably dates back to Smeathman (1781) under the name Termes atrox: see the historical review of I. severus. This genus was created by Sjöstedt (1926) for a species previously described (soldier only) as Eutermes (Cubitermes) minitabundus by himself in 1913. Indeed, Sjöstedt (1926: 214) considered, in his identification key, that this monotypic genus was significantly different from all other Cubitermes species by their soldiers’ mandibles which are straight with slightly curved tips. It shared this feature with Megagnathotermes but in the latter genus, soldiers have longer mandibles and a less deeply emarginated labrum. In the same work, Sjöstedt (1926: 216) described the imago and worker of I. minitabundus which was then the only species of the genus, so these descriptions applied also to the genus itself. According to Sands (1965: 98) and to Krishna et al. (2013: 1858, 1927), “ fide Emerson card catalog (unpublished) ”, the imago of I. minitabundus had already been described erroneously under the name Eutermes (Trinervitermes) carbo by Sjöstedt (1924 a: 42). The species was later reverted to the genus Cubitermes by Snyder (1949: 161). The genus Isognathotermes was recently restored as valid by Hellemans et al. (2021: 233) based on a phylogenetic analysis. It now gathers several previous Cubitermes species sharing the same kinds of Enteric Valve Architectures (EVAs), i. e., of the fungifaber or finitimus patterns as defined by Josens & Deligne (2019). Among the genera derived from Cubitermes (sensu Wasmann), Isognathotermes is probably the most confusing with the most numerous synonymies and misidentifications in museum collections. The mitogenomes of 12 out of the 15 valid species have been sequenced. Species included The genus Isognathotermes clusters 22 taxa previously housed in the genus Cubitermes (sensu Wasmann), namely C. antennalis Sjöstedt, 1924; C. banksi (Emerson, 1928); C. bulbifrons Sjöstedt, 1924; C. comstocki (Emerson, 1928); C. finitimus Schmitz, 1916; C. fungifaber (Sjöstedt, 1896); C. fungifaber var. elongata Sjöstedt, 1924; C. gaigei (Emerson, 1928); C. gibbifrons Sjöstedt, 1924; C. heghi Sjöstedt, 1924; C. kemneri (Emerson, 1928); C. loubetsiensis Sjöstedt, 1924; C. minitabundus (Sjöstedt, 1913); C. modestior Silvestri, 1914; C. planifrons Sjöstedt, 1924; C. schmidti (Emerson, 1928); C. severus Silvestri, 1914; C. silvestrii Sjöstedt, 1925; C. speciosus Sjöstedt, 1924; C. subarquatus Sjöstedt, 1926; C. ugandensis Fuller, 1923; and C. zenkeri (Desneux, 1904). However, 13 of these 22 species are now considered to be junior synonyms of other species, as explained below. In addition, one species previously housed in the genus Cubitermes (sensu Wasmann), namely C. congoensis (Emerson 1928) and three possibly new species could not be classified for sure; they are housed in the Incertae sedis chapter. The genus Isognathotermes also includes seven new species: I. acristatus sp. nov., I. magniplanifrons sp. nov., I. modicus sp. nov., I. phallicaecalis sp. nov., I. phalloides sp. nov., I. rectimalatus sp. nov., and I. similifinitimus sp. nov.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	description	Redescription of the genus Imago COLOUR. Head capsule well sclerotised, dark, C 6 – C 8, paler (up to C 4) in long preserved samples. Fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 7 concolorous or somewhat paler (up to two levels) than head capsule. Antennae C 4 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 5 – C 7, somewhat paler than head capsule; meso- and metanotum C 5 – C 7, as pronotum. Legs C 3 – C 5, tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 2 – Cf 4), anterior veins darker. Abdomen: tergites C 5 – C 7, as pronotum. Sternites appreciably paler in middle (C 3 – C 5) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule with some prominent setae, mainly near the eyes, overhanging a generally dense mat of short bristles. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and many shorter ones in middle; meso- and metanotum with some fine, pale setae arranged in a medio-longitudinal strip, mainly on mesonotum and the anterior lobe of metanotum, generally visible at 20 – 80 ×, but sometimes without any visible small setae; frequently 1 – 4 larger setae on metanotum. Legs very pilose, furnished (among numerous fine setae) with some large setae becoming sometimes spine-like: 0 – 18 on the carina of fore coxa and 0 – 8 on the anterior side of fore coxa; tibia pilose and furnished with 15 – 30 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 1; Figs 8 – 12, 23). Size: the imagines of the genus Isognathotermes (with those of the genus Polyspathotermes) are among the largest of those derived from the former genus Cubitermes (sensu Wasmann); however, with large overlapping with Ternicubitermes and Nitiditermes. This can be seen in most raw data, exemplified by the head width (Fig. 23); in contrast, indices differ little between genera. Head capsule large; fontanelle generally a tiny round or elongate marking (Fig. 8). Compound eyes round to shortly oval (IEy D / IEy d = 1.00 – 1.25, Fig. 9). Ocelli round to oval (IOc D / IOc d = 1.00 – 1.80). Antennae: 16 articles on alate individuals (one out of 278 alate imagines with 17 articles), always shortened by amputation of 1 – 4 articles in queens and kings. Labrum: cupola shaped, generally wider than long. Left mandible with apical tooth longer and more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth (disappearing in worn mandibles); only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth longer and more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel (Fig. 10). Thorax: pronotum appreciably wider than long (Fig. 11) and narrower than head width, straight to very weakly sellate with anterior lobe short and very slightly elevated. Legs: the fore coxa is flanged ventrally resulting in a more or less sharp edge called here ‘ carina’. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 6 and 7 – 15 branches respectively (Fig. 12). Abdomen of imagines shows 10 fully visible undivided tergites in both sexes. The first sternite is absent or vestigial; fully visible undivided sternites are 8 in male and 6 in female. In the male, the 8 visible undivided sternites are followed by a reduced one divided into two paraprocts; in female the sixth visible sternite (genital plate) is elongated and often covers the three following ones which are divided and reduced, the third having become the two paraprocts. In both male and female each paraproct bears a two-jointed small cercus. Gut: The gut of the imago is similar to that of workers with notably a well-defined mixed segment and a caecum (Fig. 3), but the caecum is proportionally less developed. Soldier COLOUR. Head capsule generally Cd 4 – Cd 6 becoming fader and darker (e. g., Cf 5 – Cf 7) in long preserved samples; frons frequently 1 – 2 levels darker than back (e. g., C 6 – C 7) sometimes with a sharp contrast giving the impression of a bicolour head; this two-tone colouring has no specific value. Gulamentum concolorous with head or somewhat darker. Antennae and labrum one level paler than head capsule. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (two levels) which are generally of the same colour as frons. Thorax, nota and legs paler than head capsule (C 3 – C 5) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow tinge on tergites. SETATION. Head capsule with few scattered fine setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 4 – 9 large setae on each lobe. Thorax: pro- and mesonotum with some setae mainly located on fore and hind margins. Legs: fore coxa furnished with 0 – 3 spines on carina and not any (or rarely one) spine on ventral side; trochanter generally with six or seven long lined-up strong setae or spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 6 – 15 spines. Abdomen: tergites with few large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 1; Figs 13 – 17, 24). Size: the soldiers of Isognathotermes (with those of Polyspathotermes) are among the largest of those derived from the former genus Cubitermes (sensu Wasmann); however, with some overlapping with Ternicubitermes and Nitiditermes. This can be seen in most raw data that vary with genus, exemplified by the head length (Fig. 24); in contrast, most indices differ little between genera. Head capsule always clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a more or less strong narrowing near posterior third or fourth (Fig. 13); however, in the soldiers of incipient colonies, lateral sides are slightly convex without any narrowing; posterior side variable: regularly rounded or sometimes with a short straight or even concave part in the middle, or with three short straight or even concave parts. In profile: upper profile always more or less concave, also in the case of soldiers from incipient colonies; angle between extended mandibles and frons varies from almost right to 130 °; frons with or without any small frontal hump overhanging the fontanelle (Fig. 14). Gulamentum: profile generally with a flat posterior part; in ventral view always more or less constricted in its posterior half, with sides of anterior part variable, from smoothly rounded (Fig. 15) to extended into a kind of angular ears. Antennae generally of 15 articles (14.5 articles in 9 %, 14 articles in 6 % and only 13 articles in one soldier from an incipient colony). Labrum: always deeply bifurcate and generally wider than long, generally (in 82 %) with sinuous sides (lyre-shaped labrum) (with convex sides in 11 %, or with straight sides in 7 %) and almost always with subtruncated lobes ending in fine, whitish or translucent tips; anterior margin concave (Fig. 16). Mandibles: sabre-like to almost hooked distally; inner edges smooth with one distinct but generally very small marginal tooth near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, narrower than head width, with generally entire anterior and posterior margins (Fig. 17). Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions generally showing trilateral symmetry: the odd primary cushions are on average 16 % longer than the even cushions; the odd cushions generally without any crest or with crests weakly developed; in the species with finitimus EVAs the pilosity becomes abruptly very dense showing the place where a crest would be expected (in comparison with the worker’s EVA); secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, either shortly lobed (in most species) or developed as a finger-like process that is sometimes capped or swollen distally. Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always 1 – 2 levels darker (C 4 – C 5). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust, scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 7 spines on carina (only 2 spines in two out of 400 samples) and 0 – 4 spines on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs, and 0, 2, 0 subapical spurs respectively and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 1; Figs 18 – 19, 25). Size: the workers of Isognathotermes (with those of Polyspathotermes) are among the largest of those derived from the former genus Cubitermes (sensu Wasmann); however, with large overlapping with Ternicubitermes and Nitiditermes. This can be seen in most raw data that vary with genus, exemplified by the hind tibia lengths (Fig. 25); in contrast, indices differ very little between genera. Head capsule weakly sclerotised (except mandibles) (Fig. 18). Antennae 15 articles, more rarely 14.5 or 14, and only 13 in a worker from an incipient colony. Labrum: cupola shaped, generally wider than long. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed, second marginal faint (visible as an undulated edge but disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth with its proximal end partly generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis (Fig. 19). Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel (Fig. 19). Thorax: pronotum sellate. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. The enteric valve with either a fungifaber or a finitimus EVA or an intermediate EVA (but in one species, I. acristatus sp. nov., with a muneris EVA), always with a triradial symmetry; the odd PCs, in their downstream part, either bear crests that are as high as or higher than they are wide, with long and strong bristles (fungifaber EVA, Fig. 4) or bear at that place a higher density of rather short bristles on a globular bulge (finitimus EVA, Fig. 5); supporting bristles are generally numerous: 13 – 38 on each side of the odd PCs (with an exception in I. acristatus); secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, frequently with two or three small lobes or developed as a finger-like process that is sometimes capped or swollen distally. Morphologic analysis The worker’s valve remains the best criterion to recognise the genus Isognathotermes, with the exception of I. acristatus sp. nov. whose valve is similar to the Ternicubitermes pattern. Additionally, the spinosity of workers’ fore coxae characterises well the genus Isognathotermes (including I. acristatus), although this feature is shared with Polyspathotermes and some species of Nitiditermes. While the soldier caste allows the best separation between species, several soldier characteristics need to be brought together to delimit the Isognathotermes genus (see diagnosis). The imagines provide few discriminating criteria and are sometimes difficult to distinguish from other Cubitermitinae imagines. As a notable exception, however, four subspecies of I. ugandensis, practically indistinguishable from the soldiers, could be evidenced from the imagines’ overall dimensions, as well both eyes and ocelli relative dimensions. Ideally, therefore, all three castes should be available for proper identification, bearing in mind that Isognathotermes termite mounds, especially older ones, often contain several species and can lead to misidentification. An initial distinction between finitimus – type and fungifaber – type EVAs is essential: a Principal Component Analysis (PCA) on the soldiers’ morphology involving all the species seems indeed confusing (Fig. 20). However, applying a PCA separately to species with a fungifaber – type EVA (Fig. 21) and those with a finitimus – type EVA (Fig. 22) results in better (though not yet complete) separation of the species. Nevertheless, it is important to note that creating sub – genera based on EVA types is not feasible because they do not delineate two distinct clades. Colour does not provide any diagnostic criterion: it depends on both the age of the individual at the time of capture and the time spent in the alcohol before being studied. Chorology-ecology This genus has been collected in forests and savannahs of West, Central, and East Africa, from Gambia to Kenya and from Angola to Malawi (Fig. 32). The species with a finitimus EVA are almost restricted to forested areas of central Africa while the species with a fungifaber EVA or an intermediate EVA can be found in forests and savannahs and is therefore present in a much larger area, from west to east equatorial Africa. There are only a few localities where species with finitimus and fungifaber EVAs coexist (Fig. 32). All species are soil feeders. Phylogenetic analyses Our phylogenetic tree integrates the mitogenomes of 101 samples from the genus Isognathotermes (Fig. 33). Mitogenome similarities between all members of Isognathotermes are larger than 91.23 % (Table ST 2). These analyses revealed that the analysed samples belonged to 12 lineages (out of the 15 morphological valid species reported herein), excluding the unidentified species represented by the samples DJ P 141 and DJ 0937. All species were monophyletic, with the exception of the intricate paraphylies of I. similifinitimus sp. nov. and I. phallicaecalis sp. nov. Note that our exclusive use of (maternally-inherited) mitogenomes may have limited our ability to fully resolve the evolutionary relationships among species of Isognathotermes. The 12 species could further be subdivided into five main clades. The first four clades are monospecific, all supported by SHalrt> 99 and UFB = 100, and respectively composed of: the species (i) I. acristatus sp. nov.; (ii) I. rectimalatus sp. nov.; (iii) I. bulbifrons; and (iv) I. fungifaber. The fifth clade combines the eight remaining species: I. severus, I. minitabundus, I. ugandensis, I. finitimus, I. phallicaecalis sp. nov., I. phalloides sp. nov., I. planifrons, and I. similifinitimus sp. nov., While the relationships among species of the first four clades are highly supported (SHalrt> 99 and UFB = 100), the relationships between species of the fifth clade are unresolved. Interestingly, the clades evidenced from the mitogenomic tree did not perfectly match EVA patterns. While clades I and IV are limited to the fungifaber EVA, and clades II and III to the finitimus EVA; the last unresolved clade V groups species exhibiting both EVAs. Additionally, we reconstructed a phylogenetic tree based on the COII gene (Supp. file 2) to study the placements of samples from Roy et al. (2006) and Hellemans et al. (2021) in the herein presented larger sampling of Isognathotermes. The COII tree included 120 samples of Isognathotermes, and recovered similar patterns as the mitogenome-based tree including the paraphyly of I. similifinitimus sp. nov. and I. phallicaecalis sp. nov. In addition, both I. phalloides (DJ 0459) and I. aff. planifrons (DJ P 141) were inserted within I. severus. We specifically detail our reanalyses of samples within I. fungifaber and I. planifrons in their respective sections.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768241FFFFAFDF4FAD8FDC1FE34.taxon	description	14. Worker’s left mandible with a larger apico-marginal distance (WMlAmD = 0.20 – 0.21 mm), soldier’s mandibles, especially the right one, very slightly curved (SMrc = 0.13 – 0.19 mm, SMrC = 0.20 – 0.25 mm); sample from Angola ........................................ I. rectimalatus Josens & Deligne sp. nov. – Worker’s left mandible generally with a smaller apico-marginal distance (WMlAmD = 0.15 – 0.22 mm); soldier’s mandibles generally more curved (SMrc = 0.16 – 0.38 mm, SMrC = 0.24 – 0.49 mm); sample from elsewhere .................................................................................................. 15 15. Larger species (SHdL = 2.98 – 3.77 mm) from Haut-Uele or Kivu, DRC ............................................ .................................................................................... I. magniplanifrons Josens & Deligne sp. nov. – Generally smaller species (SHdL = 2.60 – 3.76 mm) from elsewhere ............................................. 16 16. Soldier’s head generally more heavily humped (SHdC = 0.01 – 0.12 mm); worker’s valve with longer secondary cushions (WVS-AvL = 0.47 – 0.62 mm) ............................... I. bulbifrons (Sjöstedt, 1924) – Soldier’s head generally more slightly humped (SHdC = 0.01 – 0.08 mm); worker’s valve with shorter secondary cushions (WVS-AvL = 0.38 – 0.56 mm) ......................................................................... 17 17. Soldier’s mandibles somewhat more curved (SMlc = 0.19 – 0.35 mm); indistinguishable workers and imagines; sample from forests of western-central Africa (Cameroon, CAR, Gabon, Congo and Kongo-Central, DRC, Fig. 47) ............................................................. I. planifrons (Sjöstedt, 1924) – Soldier’s mandibles somewhat less curved (SMlc = 0.16 – 0.31 mm); indistinguishable workers and imagines; sample from continental forests of central Africa .......................................................... 18 18. Two species indistinguishable without sequencing; sample from the continental forests of Central Africa (CAR, DRC, Uganda, Fig. 38) ................................................... I. finitimus (Schmitz, 1915) – Cryptic species; only one site known to date, south of Kisangani, DRC (Fig. 47) .............................. ........................................................................................ I. similifinitimus Josens & Deligne sp. nov. Valid species and subspecies	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682433FFF6FE1AFE1FFCD4F9E5.taxon	description	Figs 26 – 34, distribution map: Fig. 47; Table 2	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682433FFF6FE1AFE1FFCD4F9E5.taxon	diagnosis	Diagnosis This species is rather atypical in the genus Isognathotermes; the worker is small sized and characterised by its odd PCs without crests but furnished with strong setae at the place where crests would be expected (Fig. 34); it could therefore be confused with a species of Ternicubitermes (bilobatodes EVA). The PCs carry a low number (5 – 10) of supporting bristles; it could therefore also be confused with a species of Cubitermes (sensu Hellemans et al. 2021). However, it has carinated fore coxae bearing 3 – 4 strong spines (the workers of Ternicubitermes and Cubitermes have unarmed conical coxae). It is also characterised by a high PC elongation index: WVPOdd-Elg = 9.79 – 13.6. The soldier is small sized in the genus Isognathotermes and can be recognised by its very weakly curved mandibles (even less curved than those of I. rectimalatus sp. nov.): SMlc = 0.11 – 0.16 mm. The imago is medium sized in the genus Isognathotermes and can hardly be distinguished from I. severus or I. ugandensis ugandensis. It has larger ocelli than I. ugandensis (IOc D = 0.20 – 0.22 mm). To date, this species is only known from Angola.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682433FFF6FE1AFE1FFCD4F9E5.taxon	etymology	Etymology The epithet acristatus, from Greek α (without) and Latin cristatus (crested) refers to the quasi absence of crests on the odd PCs of the enteric valve, even in the worker caste.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682433FFF6FE1AFE1FFCD4F9E5.taxon	materials_examined	Material examined Four samples from two locations. Holotype ANGOLA • soldier; Mussende; 10 ° 33.17 ′ S, 15 ° 39.492 ′ E; 25 Oct. 2021; M. Hasson leg.; study code: DJ 0811; GenBank no PQ 679177 (mitogenome); BE RMCA INS. Iso. 059148. Paratypes ANGOLA • soldier, worker, ♂ (alate), ♀ (queen); same data as for holotype; BE RMCA INS. Iso. 059149. Other material examined ANGOLA • soldier, worker; Cangandala N. P.; 9 ° 49.33 ′ S, 16 ° 40 ′ E; 1 Mar. 2019; M. Viage leg.; study code: DJ 0865; GenBank no PQ 679228 (mitogenome); BE RMCA INS. Iso. 059145 • soldier, worker; Cangandala N. P.; 9 ° 49.33 ′ S, 16 ° 40 ′ E; 1 Mar. 2019; M. Viage leg.; study code: DJ 0866; GenBank no PQ 679192 (mitogenome); BE RMCA INS. Iso. 059147 • soldier, worker; Cangandala N. P.; 9 ° 49.33 ′ S, 16 ° 40 ′ E; 1 Mar. 2019; M. Viage leg.; study code: DJ 0868; BE RMCA INS. Iso. 059146. Historical review Specimens of this species were recently collected by two of us (MV & MiH) in Angola; the imago, the soldier, and the worker are described here.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682433FFF6FE1AFE1FFCD4F9E5.taxon	description	Description Imago COLOUR. Head capsule: well sclerotised and dark (C 6); fontanelle slightly (one level) paler. Postclypeus concolorous with head capsule. Antennae C 4 – C 5. Thorax: pronotum concolorous with head capsule; meso- and metanotum C 4 slightly paler than head. Legs tibia C 5 slightly paler than femur C 4. Abdomen: tergites C 5. Sternites appreciably paler in middle (C 3) with both sides darker (C 5) posterior sternites darker (C 4 – C 5) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, visible at 80 ×, with some large setae on posterior lobe. Legs very pilose, furnished (among numerous fine setae) with 9 – 10 stronger setae on the carina of fore coxa and 3 – 4 on the ventral side of fore coxa and 8 – 9 on trochanter; tibia pilose, fore, mid, and hind tibia furnished with 15 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 2, Fig. 34). Size: the imagines of I. acristatus sp. nov. are medium sized in the genus Isognathotermes (Fig. 26). Head capsule: compound eyes shortly oval; ocelli rather large (IOcD> 0.20 mm) oval, removed from eyes by a distance equal to 0.8 – 0.9 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: 16 articles, shortened to 13 by amputation in queen. Labrum: cupola shaped, wider than long. Left mandible with apical tooth small sized in the genus (Fig. 27) and more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimen; premolar tooth unobstructed and clearly visible in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; second marginal tooth with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a sharp carina. Gut not studied. Soldier COLOUR. Head capsule C 4 – C 6. Antennae and labrum concolorous or somewhat paler than head capsule. Mandibles dark (C 6 – C 8) with an abrupt clearing on their bases (two levels) which is generally the same colour as frons. Thorax and legs paler than head capsule (C 2 – C 4). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum with 4 – 5 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa with at least one fine seta and 2 – 6 spines or strong setae on carina and not any spine on ventral side; trochanter generally with some strong lined-up setae; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 15 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 2, Fig. 34). Size: the soldiers of I. acristatus sp. nov. is among the small soldiers in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a very slight narrowing near posterior third or fourth; from antennal sockets sides converge towards bases of mandibles; posterior side regularly convex. Slightly to strongly concave upper profile; angle between extended mandibles and frons obtuse; frons with a small bump that overhangs the fontanelle. Gulamentum in ventral view constricted in its posterior half, with sides of anterior part smoothly rounded to extended into a kind of angular ears. Antennae: of 14 – 15 articles. Labrum: deeply bifurcate and wider than long, with almost straight sides; rounded lobes without translucent tips; anterior margin concave. Mandibles: sabre-like; inner edges smooth with one distinct but small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Mandibles very slightly curved, among the least curved in the genus. Right mandible slightly more curved than left. Thorax: pronotum sellate, as wide as 52 to 56 % of head width, with straight anterior and posterior margins. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum rather small, best seen in ventral view, near centre of abdomen, three- to four-lobed. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions narrow and longer than the even ones (as in the muneris EVA). Worker COLOUR. Head capsule pale (C 2 – C 3). Antennae: proximal articles pale (C 2), distal articles two levels darker (C 4). Thorax, nota, and legs pale (C 2 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 3 – 4 spines on carina and 1 – 3 on ventral side; fore trochanter with 6 lined up spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 7 – 15 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 2, Fig. 34). Size: the workers of I. acristatus sp. nov. are medium sized in the genus Isognathotermes. (Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae: 14 – 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed but shorter than in most other species; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge and disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth unobstructed and clearly visible in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as about 63 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the muneris pattern with triradial symmetry: the odd PCs, in their downstream part, are narrow and bear strong, bent setae; supporting bristles are few: 5 – 10 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, shortly lobed (3 – 4 small lobes). Chorology-ecology This species is only known from Angola, its geographic distribution (Fig. 47) is linked with the Angolan Miombo woodland ecoregion. Molecular data Mitogenomes of I. acristatus sp. nov. (three samples) exhibit less than 1 % dissimilarity with each other (Supp. file 4: GenBank accessions: see supplementary Supp. file 3 and ‘ Material examined’), and much more with other species. The species acristatus is found sister – group to all other Isognathotermes species (Fig. 33); its atypical EVA - very simple and comparable to the EVA of Cubitermes or Ternicubitermes - may represent the ancestral state of this genus’ EVA.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768243FFFEBFDD1F9BFFBA4FB16.taxon	description	Figs 26 – 31, 33, 35 – 37, distribution map: Fig. 38; Table 3	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768243FFFEBFDD1F9BFFBA4FB16.taxon	diagnosis	Diagnosis The individuals of this species come in two forms, one medium sized (previously called I. gibbifrons) and a large one (previously called I. bulbifrons) (Fig. 72); these two species are here synonymised on the basis of chorological-ecological data and the sequencing of their mitogenomes. The worker has a finitimus EVA (Fig. 5) and is among the medium sized to the largest workers of the genus Isognathotermes (Fig. 30; WHdW = 1.11 – 1.43 mm) it can hardly be distinguished from the workers of other species which have the same kind of EVA. The soldier has generally a recognizable finitimus EVA; it is among the medium to the largest sized soldiers of the Isognathotermes species with a finitimus EVA (SHdL = 2.65 – 3.76 mm; Figs 28 – 29), with less curved mandibles (SMlc = 0.14 – 0.31 mm) than I. magniplanifrons sp. nov. As it is suggested by its name, its head is bulbous: the upper profile is moderately to strongly concave (SHdC = 0.01 – 0.12 mm), on average, clearly more than that of I. magniplanifrons (but this is shared with I. severus, I. minitabundus, and I. ugandensis which have fungifaber EVAs). The imago is among the medium to the largest sized imagines in the genus Isognathotermes with a finitimus EVA, (IHdW = 1.44 – 1.73 mm); the larger ones can be confused with the imago of I. magniplanifrons sp. nov. Its mandibular apico-marginal distance is clearly larger (IMlAmD = 0.20 – 0.30 mm) than in any other Isognathotermes species with a finitimus EVA (Fig. 27). Some samples could be confused with I. finitimus (same size, mandibles of I. bulbifrons less curved than in I. finitimus) or with I. phallicaecalis sp. nov. that has a different caecum. This species is also defined by its ecology: all known samples come from forested regions up to about 300 km from the Atlantic coast (Fig. 38).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768243FFFEBFDD1F9BFFBA4FB16.taxon	etymology	Etymology The name bulbifrons from the Latin bulbus (onion) and frons (forehead) refers to the fact that the frons of the soldier viewed from the side is distinctly bulbous, as Sjöstedt pointed out in the original description.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768243FFFEBFDD1F9BFFBA4FB16.taxon	materials_examined	Material examined Sixty-five samples from 23 locations (and four incipient or young colonies from one location). Of the 55 old museum samples examined, three were initially labelled as Cubitermes bulbifrons, five as C. gibbifrons, 14 as C. heghi, one as C. kemneri, two as C. subarquatus, one as C fungifaber, 17 as Cubitermes sp., and five as Cubitermes 2, and seven were not identified. Of the four incipient or young colonies, two were labelled as C. gaigei and two as C. heghi. This species also includes six samples of C. aff. subarquatus “ spC ” (in Roy et al. 2006). Paratypes of Cubitermes bulbifrons Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Mukimbungu; 5 ° 8.5 ′ S, 14 ° 2.3 ′ E; ca 1904; K. E. Laman leg.; study code: DJ 0298; initially C. bulbifrons; AMNH. Syntypes of Cubitermes gibbifrons Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Mukimbungu; 5 ° 8.5 ′ S, 14 ° 2.3 ′ E; ca 1904; K. E. Laman leg.; study code: DJ 0280; initially C. gibbifrons; NHMM. Paratypes from type colony of Cubitermes heghi Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Mukimbungu; 5 ° 8.5 ′ S, 14 ° 2.3 ′ E; ca 1904; K. E. Laman leg.; study code: DJ 0290; initially C. heghi → C. bulbifrons; AMNH. Paratypes from type colony of Mirotermes (Cubitermes) kemneri Emerson, 1928 CAMEROON • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0297; initially C. kemneri; AMNH. Other material examined ANGOLA • soldier, worker; Quimucanda; 8 ° 5.14 ′ S, 15 ° 7.43 ′ E; 27 Nov. 2024; J. Šobotnik leg.; study code: DJ 0970; BE RMCA INS. Iso. 059892. CAMEROON • soldier, worker, ♀ (queen); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; [no date, around 1912]; from J. Bequaert collection; study code: DJ 0610; initially C. bulbifrons; AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0109; initially C. heghi in RMCA; BE RMCA INS. Iso. 059152 • soldier, worker, ♀ (alate); Mbalmayo; 3 ° 31 ′ N, 11 ° 30 ′ E; 17 Oct. 1962; G. Becker leg.; study code: DJ 0723; initially C. heghi; NHMUK • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 120; initially C. heghi; NHMUK 13671911 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 122; initially C. heghi; NHMUK 13671907 • soldier, worker, ♀ (alate); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ 0727; initially C. heghi; NHMUK • soldier, worker, ♀ (queen); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ 0725; initially C. heghi; NHMUK • soldier, worker, ♂ (king), ♀ (queen); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ 0726; initially C. heghi; NHMUK • soldier, ♀ (queen); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 104; initially C. gaigei; NHMUK 13671901 • soldier, ♀ (queen); Edea; 3 ° 48 ′ N, 10 ° 8 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 109; initially C. gaigei; NHMUK 13671899 • soldier, worker; Kala; 3 ° 51 ′ N, 11 ° 22 ′ E; 27 Apr. 1992; A. Dejean leg.; study code: DJ 0063; initially C. subarquatus in RMCA; BE RMCA INS. Iso. 059154 • soldier, worker; Nkolbisson (Yaoundé); 3 ° 52.25 ′ N, 11 ° 27.25 ′ E; 20 Jun. 1992; A. Dejean leg.; study code: DJ 0062; GenBank no PQ 679245 (mitogenome); initially C. subarquatus in RMCA; BE RMCA INS. Iso. 059153 • soldier, worker; Akok; 2 ° 47 ′ N, 10 ° 17 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0714; initially C. bulbifrons; NHMUK 13671835 • soldier, worker; Bilik; 3 ° 31 ′ N, 11 ° 17 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0706; initially C. heghi; NHMUK 13671839 • soldier; Ebogo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 094; initially C. heghi; NHMUK 13671838 • soldier; Ebogo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 095; initially C. heghi; NHMUK 13671842 • soldier; Eboufek; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 098; initially C. heghi; NHMUK 13671843 • soldier, worker, ♀ (queen); Eboufek; 3 ° 30 ′ N, 11 ° 53 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0715; GenBank no PQ 679207 (mitogenome); initially C. heghi; NHMUK 13671836 • soldier, worker; Mbalmayo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0713; initially C. heghi; NHMUK 13671837 • soldier, worker; Mbalmayo; 3 ° 31 ′ N, 11 ° 30 ′ E; Apr. 1996; P. Eggleton et al. leg.; study code: DJ 0712; initially C. heghi; NHMUK 13671840 • soldier, worker; Mbalmayo; 3 ° 25.1 ′ N, 11 ° 27.1 ′ E; 17 Apr. 2015; J. Šobotnik leg.; study code: DJ 0253; BE RMCA INS. Iso. 059151 • soldier; Ejokou; 3 ° 20 ′ N, 11 ° 27 ′ E; 12 Mar. 2018; J. Šobotnik leg.; study code: DJ 0968; BE _ RMCA _ INS. Iso. 059947. DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Luali; 5 ° 6 ′ S, 12 ° 29 ′ E; 26 Aug. 1912; J. Bequaert leg.; study code: DJ 0058; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059234 • soldier, worker; Luali; 5 ° 6 ′ S, 12 ° 29 ′ E; 26 Aug. 1913; J. Bequaert leg.; study code: DJ 0614; initially C. gibbifrons; AMNH • soldier; Sona Pangu; 5 ° 30 ′ S, 13 ° 55 ′ E; 9 Apr. 1948; A. Emerson leg.; study code: DJ U 117; initially C. gibbifrons; NHMUK 13671905 • soldier, worker, ♂ (king), ♀ (queen); Sona Pangu; 5 ° 30 ′ S, 13 ° 55 ′ E; 9 Apr. 1948; A. Emerson leg.; study code: DJ 0615; initially C. gibbifrons; AMNH • soldier, worker; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 19 Apr. 1965; A. Bouillon leg.; study code: DJ 0890; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059229 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ B 280; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059218 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ B 281; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059219 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ B 284; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059222 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ B 288; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059224 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ B 289; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059216 • soldier, worker; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; A. Bouillon (?) leg.; study code: DJ 0176; BE RMCA INS. Iso. 059225 • soldier, worker; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ 0888; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059226 • soldier, worker; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ 0889; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059227 • soldier, worker, ♀ (queen); Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ 0915; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059231 • soldier, worker, ♀ (queen); Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 20 Sep. 1965; G. Mathot leg.; study code: DJ 0916; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059232 • soldier; Mbamba Kilenda; 4 ° 55 ′ S, 15 ° 30 ′ E; 19 Oct. 1966; C. Nkakala leg.; study code: DJ B 311; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059150 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 278; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059214 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 279; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059215 • soldier, worker, ♂ (king); Kai Tshianga; 5 ° 51.5 ′ S, 12 ° 42 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ 0917; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059233 • soldier; Kai-Tshianga; 5 ° 52 ′ S, 12 ° 42 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 305; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059217 • soldier; Kai-Tshianga; 5 ° 52 ′ S, 12 ° 42 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 306; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059220 • soldier; Kai-Tshianga; 5 ° 52 ′ S, 12 ° 42 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 307; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059223 • soldier, worker; Kai-Tshianga; 5 ° 52 ′ S, 12 ° 42 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ 0892; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059228 • soldier; Kiemi; 5 ° 32 ′ S, 12 ° 49 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 310; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059221 • soldier, worker; Kiemi; 5 ° 16 ′ S, 12 ° 34 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ 0893; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059230. GABON • soldier; Mekambo; 1 ° 0.4 ′ N, 13 ° 56 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 158; GenBank no PQ 679204 (mitogenome); MNHN EP 9829 • soldier; Piste du Bouéni; 0 ° 20 ′ N, 13 ° 11 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 170; MNHN EP 9831 • soldier; route Makokou – Booué; 0 ° 35 ′ N, 12 ° 45 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 161; GenBank no PQ 679219 (mitogenome); MNHN EP 9830 • soldier; Piste du Bouéni; 0 ° 44 ′ N, 13 ° 12 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 160; MNHN EP 9881 • soldier; Piste du Bouéni, km 53; 0 ° 44 ′ N, 13 ° 12 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 165; MNHN EP 9882 • soldier, worker; Route Makokou – Mekambo; 0 ° 41 ′ N, 13 ° 2 ′ E; 24 Mar. 1962; J. Pasteels leg.; study code: DJ 0722; initially C. heghi; NHMUK • soldier, worker; Simintang; 0 ° 34 ′ N, 12 ° 42 ′ E; 16 Apr. 1962; J. Deligne leg.; study code: DJ 0721; initially C. heghi; NHMUK • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 215; MNHN EP 9828 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 7 Mar. 1998; M. Harry leg.; study code: DJ P 246; GenBank no PV 564650 (mitogenome); MNHN EP 9883 • soldier, worker; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0621; GenBank nos DQ 246543 (COII) DQ 246533 (ITS 2) PQ 679175 (mitogenome); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9834 • worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0627; GenBank nos DQ 127304 (COII), DQ 246537 (ITS 2), PQ 679232 (mitogenome); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9835 • soldier, worker; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0628; GenBank nos DQ 127303 (COII), DQ 246531 (ITS 2), PQ 679186 (mitogenome); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9836 • soldier, worker; La Lopé N. P.; 0 ° 16.7 ′ S, 11 ° 33 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0922; MNHN EP 9832 • soldier, worker; La Lopé N. P.; 0 ° 16.7 ′ S, 11 ° 33 ′ E; 11 Mar. 1998; M. Harry leg.; study code: DJ 0629; GenBank nos DQ 127307 (COII), DQ 246534 (ITS 2), PQ 679180 (mitogenome); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9837 • soldier, worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ 0630; GenBank nos DQ 246544 (COII), DQ 246535 (ITS 2), PQ 679240 (mitogenome); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9838 • soldier, worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ 0631; GenBank nos DQ 127306 (COII), DQ 246536 (ITS 2); initially C. aff. subarquatus “ spC ” (cf. Roy et al. 2006); MNHN EP 9839 • soldier, ♂ (king); La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 16 Mar. 1998; M. Harry leg.; study code: DJ P 239; MNHN EP 9833. Historical review The soldier alone was first described shortly as Cubitermes bulbifrons by Sjöstedt (1924 b: 256) on a sample from Mukimbungu, DRC; all castes were subsequently described in Sjöstedt (1926: 247 – 248) on a sample from Gabon. In the same papers, Sjöstedt (1924 b: 255) described shortly the soldier of Cubitermes heghi, whose soldier and worker he described with more details (1926: 249) on a sample from Mukimbungu, DRC. He considered both species as very similar, C. bulbifrons being somewhat larger than C. heghi. Sjöstedt (1926: 221 – 226) included both species in a key of the Cubitermes soldiers and inserted them in a “ bulbifrons - Gruppe ” of species with an upwards humped frons. He also included C. bulbifrons in a key of the imagines (Sjöstedt 1926: 218 – 220). Snyder (1949: 157) catalogued Cubitermes bulbifrons and C. heghi in the sub-family Termitinae. Ruelle (1975: 7) established C. heghi as a junior synonym of C. bulbifrons; however, Ruelle (1992: 500), changed his mind and established C. bulbifrons as a junior synonym of C. heghi for a reason of page priority. Krishna et al. (2013: 1916) housed Cubitermes bulbifrons in the sub-family Cubitermitinae; espousing the view of Ruelle (1975) but contesting the reversal of precedence proposed by Ruelle (1992), they therefore considered this species as the senior synonym of C. heghi. Synonymy of the two species is compatible with their enteric valve morphology, as both species belong to the same finitimus valve group (Josens & Deligne 2019: 42 – 44). Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Cubitermes aff. subarquatus “ spC ”, mentioned by Roy et al. (2006) in La Lopé National Parc, Gabon was considered a cryptic species on a genetical base. Hellemans et al. (2021: 231) placed it in the restored genus Isognathotermes and maintained its cryptic status. But with the recent sequencing of several samples of I. bulbifrons (not available in 2021), it turned out that C. aff. subarquatus “ spC ” must become a junior synonym of I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768243FFFEBFDD1F9BFFBA4FB16.taxon	description	The NHMUK collection contains two samples from incipient colonies (DJ 0725 and DJ 0726) and two samples of very young colonies (DJ U 104 and DJ U 109) that were collected at Edea, Cameroon, by N. M. Collins; they are identified as I. bulbifrons based on their queens (not at all physogastric but wingless and with shortened antennae). If the soldiers had been used instead, the resulting identification would have be ‘ I. congoensis’. It is indeed well known (Noirot 1955; Mathot 1964; and personal (G. J., J. D.) observations) that the first soldiers and workers produced by young colonies are significantly smaller than the soldiers and workers of mature colonies. In the PCA (Fig. 22), the ellipse labelled “ c + b ” pools I. congoensis and the young societies of I. bulbifrons: it shows a strong difference of size (x-axis) from mature colonies (labelled ‘ bulb’); moreover, it is stretched because it is made up on the one hand of the two incipient colonies plus the two I. congoensis samples (the four points furthest to the left = smallest individuals) and, on the other hand, of the two very young colonies (slightly larger individuals). Therefore, the samples identified as I. congoensis (the imagines of them being unknown), might also be incipient colonies of I. bulbifrons, hence their location in the chapter Incertae sedis. Redescription Imago The description provided by Emerson (1928: 511 – 513) under the name Mirotermes (Cubitermes) heghi concerned in fact Cubitermes (now Polyspathotermes) sulcifrons. The imago of P. heghi has indeed a much smaller fontanelle and, among the species of the former Cubitermes genus of West Africa, only imagines of P. sulcifrons show a fontanelle as large as that described and figured by Emerson (Josens & Deligne 2019: 62). Further, the imago of Cubitermes kemneri was never described, so the imago of I. bulbifrons was only described to-date by Sjöstedt (1926: 247 – 248) based on alates and queen from Gabon (not examined). COLOUR. Head capsule: well sclerotised, dark, C 6 – C 7; fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 6 slightly paler (one level) than head capsule. Antennae C 5 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 7, as head capsule; meso- and metanotum C 6 – C 7, as pronotum. Legs C 4 – C 5, tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 3 – Cf 4), anterior veins darker. Abdomen: tergites C 6 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule with some prominent setae, mainly near the eyes, and a higher density of shorter, finer setae everywhere. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at magnification 50 × or more). Thorax: pronotum with prominent setae mainly on margins and many shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 20 – 40 ×; sometimes three to four larger setae on metanotum. Legs very pilose, furnished (among numerous fine setae) with some fine spines: 6 – 11 on the carina of fore coxa and 1 – 3 on the ventral side; tibia pilose; fore, mid, and hind tibia furnished with 30 – 40 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 3; Figs 26 – 27, 35). Size: the imagines of I. bulbifrons are, on average, rather variable (Fig. 26); some individuals (with I. zenkeri), have the largest apical teeth and the largest apico-marginal distances (IMlAmD = 0.20 – 0.30 mm (Fig. 27). Head capsule: compound eyes nearly round; fontanelle generally a tiny elongate marking; ocelli nearly round to oval removed from eyes by a distance about equal to 0.7 to 1.3 ocellus small diameter. Antennae: 16 articles on alate individuals, always shortened by amputation by one to three articles in queens and kings. Labrum: cupola shaped. Left mandible with apical tooth, on average, longer than in most other species of the genus and always longer and more prominent than first marginal; marginal teeth three in number but second one only suggested by a slight undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always longer and more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and slightly narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a sharp carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 3 and 10 – 11 branches, respectively. Soldier COLOUR. Head capsule generally Cd 4 – Cd 6 becoming fader and darker (e. g., Cf 5 – Cf 7) in long preserved samples; there is frequently a gradient from a darker frons to a paler back (e. g., C 6 → C 4). Gulamentum concolorous with or somewhat darker than head. Antennae and labrum somewhat paler (one to two levels) than head capsule. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons. Thorax, nota and legs paler than head capsule (C 3 – C 5) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of hair-like setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 5 – 9 large setae on each lobe. Thorax: pro- and mesonotum with some setae located mainly on margins. Legs: fore coxa furnished with 0 – 3 spines on carina and not any one on ventral side; trochanter generally with some long lined-up setae, sometimes with four to seven spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 6 – 15 spines. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 3; Figs 28 – 29, 36). Size: the soldiers of I. bulbifrons are rather variable, the largest of them are (with those of I. magniplanifrons) the largest of the genus Isognathotermes; this is best seen in the length of the soldiers’ heads (Fig. 28). Head capsule: always clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a narrowing near posterior third or fourth (however, in the soldiers of incipient or very young colonies, lateral sides are slightly convex without any narrowing); from antennal sockets sides converge more or less clearly towards bases of mandibles; posterior side variable. In profile: upper profile always clearly concave, even in soldiers from incipient colonies; frons with a small frontal hump (rarely with no hump) overhanging the fontanelle, even in soldiers from incipient colonies. Gulamentum in ventral view always more or less constricted in its posterior half, with sides of anterior part softly curved or forming an acute widening or even a kind of ear on each side; generally, with a flat posterior part from side view. Antennae: of 15, sometimes 14.5, articles (only 13 articles in one soldier from an incipient colony). Labrum: always deeply bifurcate and wider than long, with sides generally lyre-shaped, rarely straight; lobes angular, with fine, whitish or translucent tips; anterior margin concave. Mandibles: sabre-like, almost hooked distally; inner edges smooth with one distinct but generally very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, as wide as 52 – 68 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions are on average 17 % longer than the even cushions, generally without any hump or with humps weakly developed, however, the pilosity becoming abruptly very dense shows the place where a hump is expected (Fig. 6); secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed (two or three small lobes). Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 4 spines on carina and 1 – 3 on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 3; Figs 30 – 31, 37). Size: the workers of I. bulbifrons are among the medium sized to the largest of the genus Isognathotermes (with large overlapping on several other species: Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae: generally 14.5 articles, 14 in workers from incipient colonies. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but Fig. 36 (continued on next page). Isognathotermes bulbifrons (Sjöstedt, 1924) and junior synonym species: soldiers. Lateral and dorsal views of a soldier’s head capsule, ventral view of its mandibles, and dorsal view of its labrum. A. From type colony of Cubitermes bulbifrons Sjöstedt, 1924 (DJ 0298). B. From type colony of Cubitermes heghi Sjöstedt, 1924 (DJ 0290). C. From type colony of Mirotermes (Cubitermes) kemneri Emerson, 1928 (DJ 0297). D. From type colony of Cubitermes gibbifrons Sjöstedt, 1924 (DJ 0280). with a blunt tip even when fresh, second marginal faint (visible as an undulated edge but disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 64 to 78 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a high density of rather short bristles on a globular bulge; supporting bristles are generally numerous: 22 – 38 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, shortly lobed (three, rarely two or four, small lobes, sometimes only sketched). Fig. 36 (continued). E. From “ C. aff. subarquatus spC ” (DJ 0621). F. From an incipient colony of I. bulbifrons (DJ 0725). G. From type colony of Mirotermes (Cubitermes) congoensis Emerson, 1928 (DJ 0299), probably also an incipient colony of I. bulbifrons (Sjöstedt, 1924) (labrum somewhat distorted by dessication). Fig. 37 (continued on next page). Isognathotermes bulbifrons (Sjöstedt, 1924) and junior synonym species: soldiers and workers. PC 1 s of soldier’s and worker’s enteric valves, worker’s caecum and foreleg. A. From type colony of Cubitermes bulbifrons Sjöstedt, 1924 (DJ 0298). B. From type colony of Cubitermes heghi Sjöstedt, 1924 (DJ 0290), the downstream end of worker’s PC 1 is twisted. C. From type colony of Mirotermes (Cubitermes) kemneri Emerson, 1928 (DJ 0299). D. From type colony of Cubitermes gibbifrons Sjöstedt, 1924 (DJ 0280). Chorology-ecology This species has been collected in the northwestern Congolian lowland forest and the Atlantic equatorial coastal forest ecoregions (Cameroon, Gabon, DRC and Angola) up to about 300 km from the Atlantic coast (Fig. 38). Molecular data The species I. bulbifrons corresponds to the Cubitermes “ spC ” in Roy et al. 2006 and Hellemans et al. 2021, proposed on basis of the COII gene. Our present sampling includes full mitogenomes from Gabonese samples previously published under the appellation “ Cubitermes aff. subarquatus spC ” by Roy et al. (2006), which we herein formally assign to I. bulbifrons. Ten mitogenomes of I. bulbifrons collected from Gabon and Cameroon are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). Overall, mitogenomes exhibit less than Fig. 37 (continued). E. From “ Cubitermes aff. subarquatus spC ” (DJ 0621). F. From an incipient colony of I. bulbifrons (Sjöstedt, 1924) (DJ 0726). G. From type colony of Mirotermes (Cubitermes) congoensis Emerson, 1928 (DJ 0299 & 0484), which is, probably, also an incipient colony of I. bulbifrons. 2 % dissimilarity with each other (Supp. file 4 with the samples DJ 0621 and DJ P 158 being the most dissimilar). Of the ten samples sequenced, nine belonged to the large form of I. bulbifrons and only one (DJ P 246) to the former I. gibbifrons (the medium-sized form of I. bulbifrons).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	description	Figs 5, 13 – 20, 22, 26 – 33, 39 – 41, distribution map: Fig. 38; Table 4	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	diagnosis	Diagnosis As already pointed out by H. Schmitz (1916), a soldier or an imago alone does not allow a confident identification. Indeed, this species is difficult to identify from its morphology since it is in a medium position in the genus Isognathotermes (blue ellipse in the middle of Fig. 22). It is obviously easier to identify this species from its mitogenome: I. finitimus makes a clade well apart from the planifrons clade (Hellemans et al. 2021: 231). If molecular data are not available, the best recognition is reached by a combination of worker’s EVA, soldiers’ mandibles, and geographical origin. The worker has a finitimus EVA and is among the medium-sized workers of the genus Isognathotermes (Fig. 30). The soldier is also medium sized in the genus Isognathotermes (Fig. 28; SHdL = 2.70 – 3.45 mm), and its EVA can generally be recognized as a finitimus EVA. The forehead has little, or no prominence, and the upper profile is only slightly concave. In comparison with I. planifrons, it has, on average, a higher acceleration of the mandible curvature: ln (SMlpR) - ln (SMldR) = 1.28 – 2.42 (0.89 – 2.28 in I. planifrons), that means: mandibles less curved at the base and more so at the tip. In I. planifrons, the soldier has on average slightly more curved mandibles. The imago is also medium sized (Fig. 26) in the genus Isognathotermes (IHdW = 1.47 – 1.71 mm). This medium-sized species can be confused with several other species with a finitimus EVA: I. planifrons and I. bulbifrons (I. phalloides sp. nov. and I. phallicaecalis sp. nov. have also similar sizes and morphologies but they can be distinguished easily by their caecum). Isognathotermes finitimus can also be characterized by its ecology and chorology: it has always been found in continental forests east of longitude 15 ° E (but the I. finitimus and I. planifrons areas overlap in the south-west of CAR (Figs 38, 47); other species with a finitimus EVA can be found in this area: I. magniplanifrons sp. nov. with distinctly larger soldiers or I. phallicaecalis sp. nov. and I. phalloides sp. nov. with a characteristic, different caecum.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	etymology	Etymology The epithet finitimus from the Latin finitimus (bordering, close to) doubtless refers to the proximity of C. finitimus with both C. fungifaber and C. zenkeri, discussed at length by Schmitz in the original description.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	materials_examined	Material examined Sixty samples from 24 locations. Of the 35 old museum samples examined, nine samples were initially labelled as Cubitermes finitimus, four as C. fungifaber, one as C. fungifaber var. elongata, one as C. loubetsiensis, six as C. minitabundus, 11 as C. speciosus, two as C. subarquatus, and one as Cubitermes sp. Syntype of Cubitermes finitimus Schmitz, 1916 DEMOCRATIC REPUBLIC OF THE CONGO • ♀ (queen); Stanleyville [= Kisangani]; 0 ° 31 ′ N, 25 ° 12 ′ E; before 1904; H. Kohl leg.; study code: DJ 0291; initially C. finitimus → C. fungifaber; AMNH. Syntypes of Cubitermes speciosus Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (queen); Moto; 2 ° 54.47 ′ N, 29 ° 24.85 ′ E; 1923; L. Burgeon leg.; study code: DJ 0061; initially C. speciosus in RMCA; BE RMCA INS. Iso. 059178. Krishna et al. (2013: 1937) mention that other syntype samples are deposited in AMNH and PPRI, not examined. Syntypes of Cubitermes subarquatus Sjöstedt, 1926 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, ♂ (alate); Stanleyville [= Kisangani]; 0 ° 31 ′ N, 25 ° 12 ′ E; before 1904; H. Kohl leg.; study code: DJ 0803; initially C. subarquatus → C. finitimus; AMNH, IZC 00321262. This latter label (DJ 0803) has been amended: on the one hand Emerson (1928: 510 – 511) mentioned for this sample: “ Lubila River, Belgian Congo: Lang-Chapin Coll., Sept. 20, 1919 ” but in his introduction he wrote: “ the collection was made by Mr. Herbert Lang and Mr. James P. Chapin during the American Museum Congo expedition, 1909 – 1915, … ”: this period does not include the collecting date of September 1919. On the other hand, the photo of the nest (plate XXIX) is dated “ September 20, 1909 ”, which is consistent with the dates of the Lang-Chapin expedition. In agreement with Sjöstedt (1926: 244), we changed the label to: “ Stanleyville [= Kisangani], H. Kohl coll. [Date unknown, probably before 1904] ”. Krishna et al. (2013: 1938) mention that another syntype sample is deposited in PPRI, not examined. Other material examined CENTRAL AFRICAN REPUBLIC • soldier, worker; Lobaye rain forest; 4 ° 5 ′ N, 17 ° 0 ′ E; 1975; G. Becker leg.; study code: DJ 0680; initially C. speciosus; NHMUK 13671988 • soldier; near Bangui; 4 ° 16 ′ N, 18 ° 31 ′ E; 1975; G. Becker leg.; study code: DJ 0679; initially C. speciosus; NHMUK 13671989. DEMOCRATIC REPUBLIC OF THE CONGO • soldier, ♀ (alate); Sankuru; 3 ° 57 ′ S, 21 ° 48 ′ E; [no date]; unknown leg.; study code: DJ 0279; initially C. fungifaber var. elongata; AMNH • soldier, worker, ♀ (queen); unclear data: “ Lubila R. ”; 1 ° 0 ′ N, 27 ° 10 ′ E; 20 Sept. 1919 or Stanleyville [= Kisangani]; 0 ° 31 ′ N, 25 ° 12 ′ E; before 1904; H. Kohl leg.; study code: DJ 0295; initially C. subarquatus → C. finitimus; AMNH • soldier, worker; Irumu – Avakubi road; 1 ° 21 ′ N, 29 ° 6 ′ E; Oct. 1906; unknown leg.; study code: DJ 0677; initially C. subarquatus; NHMUK 13671992 • soldier, worker; Medje; 2 ° 25 ′ N, 27 ° 18 ′ E; Jun. 1914; Lang-Chapin leg.; study code: DJ 0122; initially C. loubetsiensis in RMCA; BE RMCA INS. Iso. 059177 • soldier, ♀ (queen); Medje; 2 ° 25 ′ N, 27 ° 18 ′ E; Jun. 1914; Lang Chapin leg.; study code: DJ N 125; initially C. fungifaber; AMNH • soldier, worker; Medje; 2 ° 25 ′ N, 27 ° 18 ′ E; Jul. 1914; Lang-Chapin leg.; study code: DJ 0612; initially C. finitimus; AMNH • soldier, worker; unclear location “ Lubila R. ”; [1 ° 0 ′ N, 27 ° 10 ′ E?]; 20 Sep. 1919; Lang-Chapin leg.; study code: DJ 0064; initially C. subarquatus in RMCA; BE RMCA INS. Iso. 059182 • soldier, worker; Moto; 3 ° 2 ′ N, 29 ° 30 ′ E; 1922; L. Burgeon leg.; study code: DJ 0166; initially C. speciosus in RMCA; BE RMCA INS. Iso. 059179 • soldier, ♀ (alate), ♂ (alate), ♀ (queen); Moto; 3 ° 2 ′ N, 29 ° 30 ′ E; 1923; L. Burgeon leg.; study code: DJ 0071; initially C. speciosus in RMCA; BE RMCA INS. Iso. 059180 • soldier, worker, ♀ (alate), ♂ (alate); Moto; 3 ° 2 ′ N, 29 ° 30 ′ E; 1923; L. Burgeon leg.; study code: DJ 0072; initially C. speciosus in RMCA; BE RMCA INS. Iso. 059181 • soldier, worker, ♀ (alate), ♀ (queen); Epulu; 1 ° 24 ′ N, 28 ° 36 ′ E; 2 May 1948; A. Emerson leg.; study code: DJ 0613; initially C. finitimus; AMNH • soldier, worker, ♀ (queen); Epulu; 1 ° 24 ′ N, 28 ° 36 ′ E; 12 May 1948; A. Emerson leg.; study code: DJ 0618; initially C. finitimus; AMNH • soldier, worker; Epulu; 1 ° 24 ′ N, 28 ° 36 ′ E; 12 May 1948; A. Emerson leg.; study code: DJ 0619; initially C. finitimus; AMNH • soldier, worker, ♀ (queen); Epulu; 1 ° 24 ′ N, 28 ° 36 ′ E; 13 May 1948; A. Emerson leg.; study code: DJ 0617; initially C. finitimus; AMNH • soldier, worker, ♀ (queen); Epulu; 1 ° 24 ′ N, 28 ° 36 ′ E; 15 May 1948; A. Emerson leg.; study code: DJ 0286; initially C. finitimus; AMNH • soldier, ♀ (queen); Yangambi; 0 ° 48 ′ N, 24 ° 29.3 ′ E; 29 May 1948; A. Emerson leg.; study code: DJ N 127; initially C. fungifaber; AMNH • soldier, ♂ (king); Stanleyville [= Kisangani]; 0 ° 31 ′ N, 25 ° 11 ′ E; 1 Jun. 1948; A. Emerson leg.; study code: DJ N 128; initially C. fungifaber; AMNH • soldier, ♀ (queen); Stanleyville [= Kisangani]; 0 ° 31 ′ N, 25 ° 12 ′ E; 1 Jun. 1948; A. Emerson leg.; study code: DJ 0616; initially C. finitimus; AMNH • soldier, ♂ (king); Mondongo; 2 ° 16.5 ′ N, 19 ° 9.4 ′ E; 13 Sep. 1966; J. Ruelle leg.; study code: DJ B 318; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059173 • soldier, worker; near Mongolo; 2 ° 14 ′ N, 19 ° 12 ′ E; 14 Sep. 1966; J. Ruelle leg.; study code: DJ 0874; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059170 • soldier, worker; Beronge; 1 ° 30 ′ S, 19 ° 0 ′ E; 18 Sep. 1966; P. Cappelle leg.; study code: DJ 0897; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059176 • soldier, worker; Yalosemba; 2 ° 34.5 ′ N, 21 ° 47.5 ′ E; 24 Sep. 1966; J. Ruelle leg.; study code: DJ 0876; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059174 • soldier, worker, ♂ (king); Yalosemba; 2 ° 34.5 ′ N, 21 ° 47.5 ′ E; 24 Sep. 1966; J. Ruelle leg.; study code: DJ 0877; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059175 • soldier, worker, ♀ (queen); Mondongo; 2 ° 16.5 ′ N, 19 ° 9.4 ′ E; 27 Sep. 1966; unknown leg.; study code: DJ 0875; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059171 • soldier, worker; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 1 Apr. 1972; A. Bouillon leg.; study code: DJ 0878; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059172 • soldier, worker, ♀ (queen); Kisangani, Masako reserve; 0 ° 38 ′ N, 25 ° 15 ′ E; 1990; K. Soki leg.; study code: DJ 0012; BE RMCA INS. Iso. 059169 • soldier, worker, ♀ (queen); Yaekela; 0 ° 49.12 ′ N, 24 ° 16.67 ′ E; 2 May 2010; B. Le Ru leg.; study code: DJ 0739; GenBank no PQ 679218 (mitogenome); BE RMCA INS. Iso. 059166 • soldier, worker, ♂ (king); Bomane; 1 ° 15.74 ′ N, 23 ° 42.4 ′ E; 19 May 2010; B. Le Ru leg.; study code: DJ 0741; GenBank no PQ 679248 (mitogenome); BE RMCA INS. Iso. 059168 • soldier, worker; Lokulu; 1 ° 14.12 ′ N, 23 ° 43.19 ′ E; 22 May 2010; B. Le Ru leg.; study code: DJ 0742; GenBank no PQ 679241 (mitogenome); BE RMCA INS. Iso. 059167 • soldier, worker; Yangambi; 0 ° 52.9 ′ N, 24 ° 19.5 ′ E; 3 Jul. 2013; Y. Roisin leg.; study code: DJ 0086; GenBank nos MN 646708 (COI), MN 685910 (COII), MN 685971 (28 S); BE RMCA INS. Iso. 059183 • soldier, worker; Yangambi; 0 ° 47.369 ′ N, 24 ° 31.08 ′ E; 5 Jul. 2013; Y. Roisin leg.; study code: DJ 0652; GenBank nos MN 646709 (COI), MN 685911 (COII), MN 685972 (28 S); BE RMCA INS. Iso. 059184 • soldier; Yangambi; 0 ° 52.957 ′ N, 24 ° 19.475 ′ E; 8 Jul. 2013; Y. Roisin leg.; study code: DJ Y 143; BE RMCA INS. Iso. 059163 • soldier, worker; Yangambi; 0 ° 47.369 ′ N, 24 ° 31.08 ′ E; 9 Jul. 2013; Y. Roisin leg.; study code: DJ 0654; BE RMCA INS. Iso. 059186 • soldier, worker; Yangambi; 0 ° 47.151 ′ N, 24 ° 31.437 ′ E; 10 Jul. 2013; Y. Roisin leg.; study code: DJ 0655; GenBank no PQ 679174 (mitogenome); BE RMCA INS. Iso. 059187 • soldier; Yangambi; 0 ° 46.851 ′ N, 24 ° 31.255 ′ E; 11 Jul. 2013; Y. Roisin leg.; study code: DJ Y 147; BE RMCA INS. Iso. 059164 • soldier; Yangambi; 0 ° 46.851 ′ N, 24 ° 31.255 ′ E; 11 Jul. 2013; Y. Roisin leg.; study code: DJ Y 148; BE RMCA INS. Iso. 059165 • soldier, worker; Yangambi; 0 ° 46.851 ′ N, 24 ° 31.255 ′ E; 11 Jul. 2013; Y. Roisin leg.; study code: DJ 0656; GenBank no PQ 679205 (mitogenome); BE RMCA INS. Iso. 059188 • soldier, worker, ♀ (alate); Yangambi; 0 ° 46.851 ′ N, 24 ° 31.255 ′ E; 11 Jul. 2013; Y. Roisin leg.; study code: DJ 0657; GenBank no PQ 679173 (mitogenome); BE RMCA INS. Iso. 059189 • soldier; Yangambi; 0 ° 48.152 ′ N, 24 ° 29.258 ′ E; 15 Jul. 2013; Y. Roisin leg.; study code: DJ Y 149; BE RMCA INS. Iso. 059156 • soldier, worker; Yangambi; 0 ° 48.152 ′ N, 24 ° 29.268 ′ E; 15 Jul. 2013; Y. Roisin leg.; study code: DJ 0653; GenBank nos MN 646710 (COI), MN 685912 (COII), MN 685973 (28 S), PQ 679167 (mitogenome); BE RMCA INS. Iso. 059185 • soldier; Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 18 Jul. 2013; Y. Roisin leg.; study code: DJ Y 144; BE RMCA INS. Iso. 059157 • soldier, worker, ♂ (king); Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 18 Jul. 2013; Y. Roisin leg.; study code: DJ 0757; BE RMCA INS. Iso. 059190 • soldier; Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 19 Jul. 2013; Y. Roisin leg.; study code: DJ Y 145; BE RMCA INS. Iso. 059158 • soldier; Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 19 Jul. 2013; Y. Roisin leg.; study code: DJ Y 150; BE RMCA INS. Iso. 059159 • soldier; Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 19 Jul. 2013; Y. Roisin leg.; study code: DJ Y 151; BE RMCA INS. Iso. 059160 • soldier; Yangambi; 0 ° 47.598 ′ N, 24 ° 29.498 ′ E; 19 Jul. 2013; Y. Roisin leg.; study code: DJ Y 152; BE RMCA INS. Iso. 059161 • soldier; Yangambi; 0 ° 48.51 ′ N, 24 ° 31.647 ′ E; 20 Jul. 2013; Y. Roisin leg.; study code: DJ Y 146; BE RMCA INS. Iso. 059162 • soldier, worker; Yangambi; 0 ° 48.51 ′ N, 24 ° 31.647 ′ E; 20 Jul. 2013; Y. Roisin leg.; study code: DJ 0620; GenBank no KY 224569 (mitogenome); BE RMCA INS. Iso. 059191 • soldier, worker, ♂ (king); Yangambi; 0 ° 48.51 ′ N, 24 ° 31.647 ′ E; 20 Jul. 2013; Y. Roisin leg.; study code: DJ 0758; BE RMCA INS. Iso. 059192 • soldier, worker, ♂ (king); Yangambi; 0 ° 48.51 ′ N, 24 ° 31.647 ′ E; 20 Jul. 2013; Y. Roisin leg.; study code: DJ 0758; BE RMCA INS. Iso. 059192. UGANDA • soldier, ♀ (queen); Bugondo forest; 1 ° 46 ′ N, 31 ° 35 ′ E; 1944; C. J. Eggding and W. V. Harris leg.; study code: DJ U 086; initially C. speciosus; NHMUK 13671980 • soldier, worker; Kawanda; 0 ° 25.23 ′ S, 32 ° 32.43 ′ E; 6 Dec. 1949; W. V. Harris leg.; study code: DJ 0684; initially C. speciosus; NHMUK 13671982 • soldier; Kampala – Fort Portal road; 0 ° 22 ′ N, 32 ° 16.5 ′ E; 3 Feb. 1955; R. M. C. Williams leg.; study code: DJ U 088; initially C. speciosus; NHMUK 13671981 • soldier, worker, ♀ (alate); Entebbe – Kampala road; 0 ° 7.5 ′ N, 32 ° 31.5 ′ E; 2 Jul. 1970; Kistner leg.; study code: DJ 0681; initially C. speciosus; NHMUK 13671985 • soldier; Kisubi forest; 0 ° 7 ′ N, 32 ° 30 ′ E; 12 Jul. 1970; Kistner leg.; study code: DJ U 087; initially C. speciosus; NHMUK 13671987. Historical review Schmitz (1916: 120 – 124) described this species under the name Cubitermes finitimus. He provided a description and measurements of king, queen, and soldier, mentioning that imagines are like those of C. zenkeri while the soldier closely resembles the soldier of C. fungifaber. Sjöstedt (1926: 221 – 226) included C. finitimus in his soldiers’ key of Cubitermes species and inserted it in a “ zenkeri - Gruppe ” of Cubitermes species with relatively shorter soldier’s mandibles. He clarified some morphological differences between C. finitimus, C. zenkeri and C. fungifaber. Snyder (1949: 159) catalogued this species under the name Cubitermes finitimus in the sub-family Termitinae and reported its known geographical distribution. Emerson, in his unpublished “ card catalog ” considered C. subarquatus as a synonym of C. finitimus (quoted by Krishna et al. 2013: 1938): this is confirmed in this work. Bouillon & Vincke (1971: 269) described the enteric valve of C. finitimus as belonging to the first of three types, the “ simple type ” without any spatula. Krishna et al. (2013: 1920) housed Cubitermes finitimus in the sub-family Cubitermitinae. Josens & Deligne (2019: 42 – 44) choose this species as type of the finitimus valve pattern group. Hellemans et al. (2021: 231 – 233) placed this species in the restored genus Isognathotermes and within a “ finitimus clade ”.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, dark when fresh, C 6 – C 7, lighter in old, preserved samples (C 4 – C 6); fontanelle concolorous with or somewhat paler than head capsule. Postclypeus C 6 – C 7 slightly paler (one level) than head capsule. Antennae C 5 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 7, concolorous with or somewhat paler than head capsule; meso- and metanotum C 6 – C 7, as pronotum. Legs C 4 – C 5. Wings hyaline with brown to grey tinge (Cf 3 – Cf 4), anterior veins darker. Abdomen: tergites C 6 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule with some prominent setae, mainly near the eyes, and a higher density of shorter, finer setae everywhere. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and many shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 20 to 80 ×, with frequently one to four larger setae on metanotum. Legs very pilose, frequently furnished (among numerous fine setae) with some strong setae: 6 – 10 on the carina of fore coxa and 0 – 2 on the ventral side of fore coxa; tibia pilose; fore, mid, and hind tibia furnished with 10 – 30 strong setae and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 4; Figs 26, 39). Size: the imagines of I. finitimus are, on average, medium sized in the genus Isognathotermes; as shown in Fig. 26. Head capsule (Fig. 39): compound eyes nearly round; fontanelle generally a tiny round to elongate marking; ocelli nearly round to oval, removed from eyes by a distance about equal to 0.7 – 1.5 ocellus small diameter. Antennae: 16 articles on alate individuals, always shortened by amputation by one-three articles in queens and kings. Labrum: cupola shaped. Left mandible with apical tooth always longer and more prominent than first marginal; marginal teeth three in number but second one only suggested by a slight undulation of edge between first and third marginal teeth (this undulation disappears in worn mandibles); only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always longer and more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a more or less sharp carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 4 and 10 – 15 branches respectively. Gut (only studied in some cases): enteric valve weakly developed and hardly comparable with workers’ and soldiers’ valves; odd PCs are slightly longer than even PCs (Fig. 6). Caecum present as a small amorphous button. Soldier COLOUR. Head capsule generally Cd 4 – Cd 6 becoming fader and darker (e. g., Cf 5 – Cf 7) in long preserved samples; there is frequently a gradient from a darker frons to a paler back (e. g., from C 6 – C 4) sometimes giving the impression that the head capsule is bicolorous (Fig. 13). Gulamentum generally concolorous with head. Antennae and labrum slightly paler (half a level) than head capsule. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons. Thorax, nota and legs paler than head capsule (C 3 – C 4). Abdomen grey to red-brown owing to digestive bolus, rarely with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 3 – 6 large setae on each lobe. Thorax: pro- and mesonotum with some setae mainly located on margins. Legs: fore coxa furnished with 2 – 4 (rarely 1 or 5) spines on carina and not any one on ventral side; trochanter generally with some long lined-up setae, sometimes with 4 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 7 – 15 spines. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 4; Figs 28, 40). Size: the soldiers of I. finitimus are medium sized in the genus Isognathotermes; this can be seen in the soldiers’ head lengths (Fig. 28). Head capsule (Fig. 40): always clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a slight to very slight narrowing near posterior third or fourth; from antennal sockets sides converge clearly towards bases of mandibles; posterior side variable. In profile: upper profile slightly to clearly concave; frons: fontanelle overhanged by a very small frontal hump (or sometimes no hump at all). Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either forming an acute widening or even a kind of ear on each side. Antennae: of 15 articles, rarely 14.5 or 14. Labrum: always deeply bifurcate and wider than long, with sides always more or less lyre-shaped; lobes angular, with fine, whitish, or translucent tips; anterior margin concave (Fig. 40). Mandibles: sabre-like, generally regularly curved but sometimes almost hooked distally; inner edges smooth with one distinct but generally very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, clearly narrower than head, generally with a slight notch in anterior margin and entire posterior margin. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions are on average 18 % longer than the even cushions, generally without any hump or with humps weakly developed, however, the pilosity becoming abruptly very dense shows the place where a hump is expected (Fig. 41); secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed (two to five small lobes). Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 7 spines on carina and 0 – 2 on ventral side; fore trochanter with 5 – 7 spines or strong setae; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 4; Figs 30, 41). Size: the workers of I. finitimus are, on average, medium sized in the genus Isognathotermes (but with large overlapping on several other species: Fig. 30). Head capsule weakly sclerotised (except mandibles). Antennae 14.5 articles, rarely 14. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge and disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 64 – 74 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a very high density of rather short bristles on a globular bulge (Fig. 41); supporting bristles are generally numerous: 22 – 29 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, shortly lobed (two to five small lobes, sometimes only sketched). Chorology-ecology The geographic distribution of I. finitimus is linked with the continental forests of central Africa; it includes the central and northeastern Congolian lowland forest and the Victoria Basin forest-savanna mosaic ecoregions (Fig. 38). Molecular data Seven mitogenomes of I. finitimus are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). They are characterized by less than 1.05 % dissimilarity with each other (Supp. file 4), with the sample DJ 0741 being the most dissimilar. Phylogenetic reconstructions based on full mitogenomes placed them in a sister relationship to the species I. ugandensis and I. minitabundus, to which they exhibit less than 2 % dissimilarity.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682422FF9DFDD9FAC9FAC3FC05.taxon	discussion	Remarks In the absence of genetic information, I. finitimus is often difficult to ascertain as its morphology places it in the middle of the genus Isognathotermes; for the species with a finitimus EVA alone, there are partial overlaps with I. bulbifrons and I. planifrons, and also with I. phallicaecalis sp. nov., and I. phalloides sp. nov. but these last two are distinguished by their caecum. Integrative taxonomy comes into its own here: in particular, chorology is a valuable aid: two samples – DJ 0655 and DJ 0656 – collected at Yangambi (Tshopo Province, DRC), were initially identified as I. gibbifrons on the basis of their morphology (although in disagreement with the chorology of this species). Mitogenome-based phylogenetic reconstructions subsequently placed these samples in I. finitimus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	description	Figs 1, 3 – 4, 6, 26 – 31, 33, 42 – 44, distribution map: Fig. 45; Tables 5 – 6	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	diagnosis	Diagnosis The worker has a fungifaber EVA (Fig. 4) but in 2 % of the samples examined, it has an EVA intermediate between the fungifaber and finitimus patterns. The worker itself and its enteric valve are among the smallest in the genus Isognathotermes (Fig. 31). The soldier is among the smallest in the genus Isognathotermes; (SHdL = 2.37 – 3.11 mm, Fig. 28); its EVA generally looks like a muneris EVA and is among the smallest in the genus Isognathotermes EVAs. In some samples, however, the soldier is as large as the smallest soldiers of I. severus. The soldier has, along with I. planifrons, the most evenly curved mandibles from base to tip: ln (SMlpR) - ln (SMldR) = 0.85 – 1.88. The imago is, on average and with that of I. phalloides sp. nov., the smallest in the genus Isognathotermes; (IHdW = 1.40 – 1.61 mm, Fig. 26). This species is also defined by its ecology and chorology: it has always been found in evergreen forests from RCI to Gabon at less than 300 km from the Atlantic coast (Fig. 45 but see later, ‘ Supplementary material of dubious I. fungifaber ’).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	etymology	Etymology The name fungifaber from the Latin fungus (mushroom) and faber (artisan) doubtless refers to the building of mushroom-shaped epigeous nests.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	materials_examined	Material examined One hundred twenty-four samples from 44 locations (plus six samples of “ dubious I. fungifaber ” from five locations). Of the 64 old museum samples examined, 49 were initially labelled as C. fungifaber, one as C. fungifaber var. elongata, four as C. banksi, one as C. comstocki, three as C. gaigei, three as C. kemneri, two as C. modestior, and one as C. schmidti. Of the six samples of “ dubious I. fungifaber ”, four were initially labelled as C. fungifaber and two not identified. This species also includes C. aff. subarquatus “ spA ” and “ spD ” (in Roy et al. 2006). Syntypes of Eutermes fungifaber Sjöstedt, 1896 CAMEROON • soldier, worker, ♀ (alate), ♀ (queen); Mbongé; 4 ° 32.18 ′ N, 9 ° 6.67 ′ E; Oct. 1891; Y. Sjöstedt leg.; study code: DJ 0271; initially Eutermes fungifaber; NHMM • soldier, worker, ♀ (queen); Mbongé; 4 ° 32.18 ′ N, 9 ° 6.67 ′ E; 27 Oct. 1891; Y. Sjöstedt leg.; study code: DJ 0272; initially Eutermes fungifaber; AMNH • worker, ♀ (alate); Mbongé (?); 4 ° 32.18 ′ N, 9 ° 6.67 ′ E; 1891; Y. Sjöstedt leg.; study code: DJ 0965; initially Eutermes fungifaber ex typis; RBINS. Krishna et al. (2013: 1921) mention that other syntype samples are deposited in NHRM, not examined. Paratypes of Mirotermes (Cubitermes) banksi Emerson, 1928 CAMEROON • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0287; initially Mirotermes (C.) banksi → C. fungifaber; AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0097; initially C. banksi in RMCA; BE RMCA INS. Iso. 059210. Krishna et al. (2013: 1913) mention that the holotype and paratypes are deposited in UMMZ, not examined. Paratypes of Mirotermes (Cubitermes) comstocki Emerson, 1928 CAMEROON • soldier, worker, ♀ (queen); Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0273; initially Mirotermes (C.) comstocki → C. fungifaber; AMNH • soldier, worker, ♀ (alate); Bipindi?; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0278; initially C. comstocki → C. fungifaber; AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0121; initially C. comstocki in RMCA; BE RMCA INS. Iso. 059212. Krishna et al. (2013: 1918) mention that the holotype (soldier) and paratypes are deposited in UMMZ, not examined, and that other paratypes are deposited in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, not examined. Paratypes of Mirotermes (Cubitermes) schmidti Emerson, 1928 CAMEROON • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0275; initially C. schmidti → C. fungifaber in AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0124; initially Mirotermes (C.) schmidti in RMCA; BE RMCA INS. Iso. 059213 Other material examined CAMEROON • soldier; Abonando [= Ossidinge]; 5 ° 54 ′ N, 9 ° 8 ′ E; no date; G. Mansfeld leg.; study code: DJ N 123; initially C. fungifaber; AMNH • soldier, worker, ♀ (alate); Mungo; 5 ° 12 ′ N, 9 ° 31 ′ E; 1903; Rohde leg.; study code: DJ 0269; initially C. fungifaber var. elongata; AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0963; initially Eutermes fungifaber; RBINS • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0964; initially Eutermes fungifaber; RBINS • soldier; Réserve du Nyong; 3 ° 50 ′ N, 10 ° 50 ′ E; 6 Dec. 1949; J. Birkett-Smith and J. Dahl leg.; study code: DJ U 036; initially C. fungifaber; NHMUK 13671854 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 033; initially C. fungifaber; NHMUK 13671851 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 039; initially C. fungifaber; NHMUK 13671847 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 040; initially C. fungifaber; NHMUK 13671850 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 041; initially C. fungifaber; NHMUK 13671848 • soldier; Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ U 042; initially C. fungifaber; NHMUK 13671846 • soldier; Mbalmayo; 3 ° 30 ′ N, 11 ° 31 ′ E; 1 Dec. 1962; G. Becker leg.; study code: DJ U 065; initially C. fungifaber; NHMUK 13671852 • soldier, worker; Mamfe – Ikom road; 5 ° 47 ′ N, 8 ° 58 ′ E; 18 Sep. 1966; W. Wilkinson leg.; study code: DJ 0718; initially C. gaigei; NHMUK 13671886 • worker, ♂ (king); Edea; 3 ° 29 ′ N, 9 ° 50 ′ E; Jul. – Sep. 1973; N. M. Collins leg.; study code: DJ 0649; initially C. fungifaber; NHMUK 13671846 • soldier, worker, ♀ (alate); Ebodjié; 2 ° 38 ′ N, 9 ° 53 ′ E; Nov. 1991; A. Dejean leg.; study code: DJ 0096; initially C. banksi in RMCA; BE RMCA INS. Iso. 059211 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 049; initially C. fungifaber; NHMUK 13671877 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 050; initially C. fungifaber; NHMUK 13671876 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 052; initially C. fungifaber; NHMUK 13671879 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 053; initially C. fungifaber; NHMUK 13671880 • soldier, ♀ (alate); Ebogo; 3 ° 23 ′ N, 11 ° 28 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 045; initially C. fungifaber; NHMUK 13671874 • soldier; Ebogo; 3 ° 23.5 ′ N, 11 ° 28 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 068; initially C. kemneri; NHMUK 13672024 • soldier; Ebogo; 3 ° 23.5 ′ N, 11 ° 28 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 069; initially C. kemneri; NHMUK 13672025 • soldier; Ebogo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 092; initially C. banksi; NHMUK 13671831 • soldier, ♂ (king), ♀ (queen); Eboufek; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 051; initially C. fungifaber; NHMUK 13671875 • soldier, worker, ♂ (king); Eboufek; 3 ° 30 ′ N, 11 ° 53 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0651; initially C. kemneri; NHMUK 13672023 • soldier, worker, ♂ (alate); Eboufek; 3 ° 30 ′ N, 11 ° 53 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0707; initially C. banksi; NHMUK 13671828 • soldier, ♂ (king); Ekombitié; 3 ° 28 ′ N, 11 ° 31 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 048; initially C. fungifaber; NHMUK 13671872 • soldier; Mbalmayo; 3 ° 27 ′ N, 11 ° 29 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 047; initially C. fungifaber; NHMUK 13671871 • soldier; Mbalmayo; 3 ° 27 ′ N, 11 ° 29 ′ E; Feb. 1996; P. Eggleton et al. leg.; study code: DJ U 044; initially C. fungifaber; NHMUK 13671870 • soldier; Mbalmayo; 3 ° 27 ′ N, 11 ° 29 ′ E; Mar. 1996; P. Eggleton et al. leg.; study code: DJ U 046; initially C. fungifaber; NHMUK 13671859 • soldier; Mbalmayo; 3 ° 27 ′ N, 11 ° 29 ′ E; Aug. 1996; P. Eggleton et al. leg.; study code: DJ U 043; initially C. fungifaber; NHMUK 13671873 • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 136; GenBank no PQ 679252 (mitogenome); MNHN EP 9869 • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 137; MNHN EP 9870 • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 138; MNHN EP 9871 • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 139; MNHN EP 9872 • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 140; MNHN EP 9873 • soldier, worker, ♂ (alate); Makak; 3 ° 38 ′ N, 11 ° 4 ′ E; 27 Nov. 2013; J. Birkett-Smith and J. Dahl leg.; study code: DJ 0650; initially C. fungifaber; NHMUK 13671853 • soldier, worker, ♂ (alate); Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0452; GenBank nos MN 646713 (COI), MN 685915 (COII), MN 685976 (28 S), PQ 679181 (mitogenome); BE RMCA INS. Iso. 059200 • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0468; GenBank no PQ 679188 (mitogenome); BE RMCA INS. Iso. 059196 • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 23 Nov. 2016; P. Akama leg.; study code: DJ 0469; GenBank no PQ 679195 (mitogenome); BE RMCA INS. Iso. 059195 • soldier, worker, ♀ (alate); Mbongé; 4 ° 32.2 ′ N, 9 ° 6.7 ′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0450; GenBank nos MN 646712 (COI), MN 685914 (COII), MN 685975 (28 S); BE RMCA INS. Iso. 059194 • soldier, worker; Mbongé; 4 ° 32.2 ′ N, 9 ° 6.7 ′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0451; GenBank no PQ 679193 (mitogenome); BE RMCA INS. Iso. 059202 • soldier, worker, ♂ (alate); Mbongé; 4 ° 32.2 ′ N, 9 ° 6.7 ′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0466; GenBank no PQ 679246 (mitogenome); BE RMCA INS. Iso. 059198 • soldier, worker, ♂ (alate); Mbongé; 4 ° 32.2 ′ N, 9 ° 6.7 ′ E; 30 Nov. 2016; P. Akama leg.; study code: DJ 0467; GenBank no PQ 679226 (mitogenome); BE RMCA INS. Iso. 059203 • soldier, worker, ♀ (queen); Ebogo; 3 ° 22.894 ′ N, 11 ° 27.862 ′ E; 23 May 2017; Y. Roisin leg.; study code: DJ 0506; GenBank nos MN 646714 (COI), MN 685916 (COII), MN 685977 (28 S); BE RMCA INS. Iso. 059206 • soldier, worker; Ebogo; 3 ° 22.872 ′ N, 11 ° 27.796 ′ E; 6 Jun. 2017; Y. Roisin leg.; study code: DJ 0521; GenBank no PQ 679222 (mitogenome); BE RMCA INS. Iso. 059205 • soldier, worker; Mbalmayo; 3 ° 28.841 ′ N, 11 ° 34.646 ′ E; 6 Jun. 2017; Y. Roisin leg.; study code: DJ 0520; GenBank no PQ 679185 (mitogenome); BE RMCA INS. Iso. 059207 CÔTE D’IVOIRE • soldier; Banco N. P.; 5 ° 23 ′ N, 4 ° 3 ′ W; 1 Jan. 1934; P. - P. Grassé leg.; study code: DJ P 199; MNHN EP 9849 • soldier; Banco N. P.; 5 ° 23 ′ N, 4 ° 3 ′ W; 8 Jan. 1934; P. - P. Grassé leg.; study code: DJ P 153; MNHN EP 9844 • soldier, worker; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 9 Jan. 1934; P. - P. Grassé leg.; study code: DJ 0203; initially C. fungifaber in MNHN; MNHN EP 9840 • soldier; Banco N. P.; 5 ° 23 ′ N, 4 ° 3 ′ W; Jan. 1939; P. - P. Grassé leg.; study code: DJ P 157; MNHN EP 9845 • soldier, ♀ (queen); Banco N. P.; 5 ° 23 ′ N, 4 ° 3 ′ W; 6 Feb. 1947; C. Noirot leg.; study code: DJ P 194; MNHN EP 9848 • soldier, ♂ (king); Adiopodoumé; 5 ° 19.75 ′ N, 4 ° 8 ′ W; 22 May 1947; C. Noirot leg.; study code: DJ P 193; MNHN EP 9847 • soldier; Banco N. P.; 5 ° 19 ′ N, 4 ° 10 ′ W; 26 Jul. 1947; C. Noirot leg.; study code: DJ P 325; MNHN EP 9863 • soldier; Mambo; 5 ° 46 ′ N, 4 ° 3 ′ W; 19 Oct. 1947; C. Noirot leg.; study code: DJ P 324; MNHN EP 9862 • soldier; Mambo; 5 ° 46 ′ N, 4 ° 3 ′ W; 19 Oct. 1947; C. Noirot leg.; study code: DJ P 333; MNHN EP 9948 • soldier; Mambo; 5 ° 46 ′ N, 4 ° 3 ′ W; 20 Oct. 1947; C. Noirot leg.; study code: DJ P 334; MNHN EP 9949 • soldier; Yapo; 5 ° 44 ′ N, 4 ° 5 ′ W; 10 Oct. 1947; C. Noirot leg.; study code: DJ P 326; MNHN EP 9864 • soldier, worker, ♀ (alate); Petit Yapo réserve forestière; 5 ° 46 ′ N, 4 ° 8 ′ W; 22 Aug. 1953; M. Lüscher leg.; study code: DJ 0277; initially C. fungifaber; AMNH • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 6 Dec. 1959; C. Noirot leg.; study code: DJ P 211; MNHN EP 9860 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 25 Dec. 1959; C. Noirot leg.; study code: DJ P 208; MNHN EP 9857 • soldier, worker, ♂ (alate); near Abidjan; 5 ° 23 ′ N, 4 ° 3 ′ W; ca 1960; C. Noirot leg.; study code: DJ 0268; initially C. fungifaber; AMNH • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P 203; MNHN EP 9852 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P 205; MNHN EP 9854 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jan. 1960; C. Noirot (?) leg.; study code: DJ P 207; MNHN EP 9856 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jan. 1960; C. Noirot leg.; study code: DJ P 209; MNHN EP 9858 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jan. 1960; C. Noirot leg.; study code: DJ P 210; MNHN EP 9859 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 30 Jan. 1960; C. Noirot (?) leg.; study code: DJ P 201; MNHN EP 9850 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 30 Jan. 1960; C. Noirot (?) leg.; study code: DJ P 204; MNHN EP 9853 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 20 Mar. 1960; C. Noirot leg.; study code: DJ P 329; MNHN EP 9866 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 7 May 1960; C. Noirot (?) leg.; study code: DJ P 202; MNHN EP 9851 • soldier, ♂ (alate); Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 7 May 1960; C. Noirot (?) leg.; study code: DJ P 206; MNHN EP 9855 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 12 Jun. 1960; C. Noirot leg.; study code: DJ P 330; MNHN EP 9867 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 17 Jun. 1960; C. Noirot leg.; study code: DJ P 336; MNHN EP 9951 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 9 Oct. 1960; C. Noirot leg.; study code: DJ P 327; MNHN EP 9865 • soldier; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 9 Oct. 1960; C. Noirot leg.; study code: DJ P 331; MNHN EP 9868 • soldier, ♂ (king); Abidjan; 5 ° 20 ′ N, 3 ° 59 ′ W; 29 Dec. 1960; C. Noirot leg.; study code: DJ P 347; MNHN • soldier, ♂ (alate); Adiaké; 5 ° 22 ′ N, 3 ° 19.7 ′ W; 15 Apr. 1963; C. Noirot leg.; study code: DJ P 348; MNHN • soldier; Banco N. P.; 5 ° 23 ′ N, 4 ° 3 ′ W; 21 Aug. 1968; G. Josens leg.; study code: DJ U 001; initially C. modestior; NHMUK 13671927 • soldier, worker; Banco N. P.; 5 ° 24 ′ N, 4 ° 3 ′ W; 21 Aug. 1968; G. Josens leg.; study code: DJ 0202; initially C. modestior in RMCA; BE RMCA INS. Iso. 059204 • soldier, worker; Ahoutoué; 5 ° 28 ′ N, 3 ° 45 ′ W; 9 Aug. 1969; Vincent leg.; study code: DJ 0229; BE RMCA INS. Iso. 059209 • soldier, worker; Banco N. P.; 5 ° 23.005 ′ N, 4 ° 3.341 ′ W; 17 Feb. 2015; G. Josens leg.; study code: DJ 0412; GenBank nos MN 646711 (COI), MN 685913 (COII), MN 685974 (28 S), PQ 679172 (mitogenome); BE RMCA INS. Iso. 059208. EQUATORIAL GUINEA • soldier; Punta Frailes; 3 ° 47 ′ N, 8 ° 43 ′ E; ca 1894; R. M. Downes leg.; study code: DJ U 110; initially C. gaigei; NHMUK 13671887 • soldier, worker, ♀ (alate); Punta Frailes; 3 ° 47 ′ N, 8 ° 43 ′ E; Oct. 1901; L. Fea leg.; study code: DJ 0311; initially C. fungifaber; MCGD. GABON • soldier; Mekambo; 1 ° 1 ′ N, 13 ° 56 ′ E; Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 167; MNHN EP 9846 • soldier, worker; Piste du Bouéni; 0 ° 13 ′ N, 11 ° 48 ′ E; 1 Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ 0855; MNHN EP 9843 • soldier, worker; Piste du Bouéni, km 53; 0 ° 44 ′ N, 13 ° 12 ′ E; 1 Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ 0853; MNHN EP 9841 • soldier, worker, ♂ (king), ♀ (queen); forêt dense de Djidji; 0 ° 13 ′ N, 11 ° 48 ′ E; 25 Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ 0854; MNHN EP 9842 • soldier, worker; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ P 242; MNHN EP 9875 • soldier, worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0623; GenBank nos DQ 127302 (COII), DQ 246527 (ITS 2), PQ 679189 (mitogenome); initially C. aff. subarquatus “ spA ” (cf. Roy et al. 2006); MNHN EP 9878 • soldier, worker; Doda (north of La Lopé N. P.); 0 ° 4.5 ′ S, 11 ° 25.5 ′ E; 10 Mar. 1998; M. Harry leg.; study code: DJ 0625; GenBank nos DQ 246541 (COII), DQ 246526 (ITS 2) initially C. aff. subarquatus “ spD ” (cf. Roy et al. 2006); MNHN EP 9879 • soldier, ♀ (queen); La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ P 243; MNHN EP 9876 • soldier, worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 13 Mar. 1998; M. Harry leg.; study code: DJ 0626; GenBank nos DQ 127299 (COII), DQ 246529 (ITS 2), PQ 679214 (mitogenome); initially C. aff. subarquatus “ spA ” (cf. Roy et al. 2006); MNHN EP 9880 • soldier; La Lopé N. P.; 0 ° 9 ′ S, 11 ° 36 ′ E; 6 Mar. 1998; M. Harry leg.; study code: DJ P 232; MNHN EP 9874 • soldier, worker; Bitam; 2 ° 5 ′ N, 11 ° 30 ′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0456; BE RMCA INS. Iso. 059197 • soldier, worker; Bitam; 2 ° 5 ′ N, 11 ° 30 ′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0457; BE RMCA INS. Iso. 059201 • soldier; Bitam; 2 ° 5 ′ N, 11 ° 30 ′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0458; BE RMCA INS. Iso. 059199 • soldier, worker; Nkobissimo [= Nkolmengboua?]; 2 ° 14.3 ′ N, 11 ° 29.25 ′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0453; GenBank no PQ 679216 (mitogenome); BE RMCA INS. Iso. 059193 • soldier, worker; Ipassa; 0 ° 30.66 ′ N, 12 ° 48 ′ E; 27 Jan. 2025; J. Šobotnik leg.; study code: DJ 0974; BE RMCA INS. Iso. 059949. GHANA • soldier, ♀ (queen); Bobiri Forest Reserve; 6 ° 40 ′ N, 1 ° 21 ′ W; 21 Feb. 1959; W. A. Sands leg.; study code: DJ U 114; initially C. gaigei; NHMUK 13671881. NIGERIA • soldier; Ijebu – Ode road; 6 ° 50 ′ N, 3 ° 54 ′ E; 2 May 1950; G. C. Webb leg.; study code: DJ N 126; initially C. fungifaber; AMNH • soldier; North of Lagos; 6 ° 49 ′ N, 3 ° 13 ′ E; 11 Feb. 1955; W. V. Harris leg.; study code: DJ U 063; initially C. fungifaber; NHMUK 13671865 • soldier; Mofu River forest reserve; 6 ° 10 ′ N, 7 ° 10 ′ E; 15 Feb. 1955; W. V. Harris leg.; study code: DJ U 062; initially C. fungifaber; NHMUK 13671864 • soldier; Port Harcourt; 4 ° 46 ′ N, 7 ° 1 ′ E; 5 Dec. 1955; D. Kay leg.; study code: DJ U 057; initially C. fungifaber; NHMUK 13671868 • soldier; Sapele; 5 ° 53 ′ N, 5 ° 41 ′ E; 31 Jan. 1956; D. Kay leg.; study code: DJ U 056; initially C. fungifaber; NHMUK 13671866 • soldier; Obanikoro; 6 ° 33 ′ N, 3 ° 22 ′ E; 8 Jan. 1957; W. Wilkinson leg.; study code: DJ U 034; initially C. fungifaber; NHMUK 13671861 • soldier; Ugo; 6 ° 5 ′ N, 6 ° 0 ′ E; 11 Jan. 1957; W. Wilkinson leg.; study code: DJ U 061; initially C. fungifaber; NHMUK 13671860 • soldier, worker; Ilaro (near)?; 7 ° 26 ′ N, 3 ° 5 ′ E; 6 Apr. 1957; W. Wilkinson leg.; study code: DJ 0549; initially C. fungifaber; NHMUK • soldier, ♀ (queen); Calabar-Mamfe road; 5 ° 17 ′ N, 8 ° 34 ′ E; 22 Apr. 1957; W. Wilkinson leg.; study code: DJ U 035; initially C. fungifaber; NHMUK 13671863 • soldier; Port Harcourt – Owerri road; 5 ° 11 ′ N, 6 ° 52 ′ E; 19 Jun. 1957; W. Wilkinson leg.; study code: DJ U 064; initially C. fungifaber; NHMUK 13671867 • soldier, worker; Port Harcourt – Owerri road; 5 ° 9.5 ′ N, 6 ° 51 ′ E; 19 Jun. 1957; W. Wilkinson leg.; study code: DJ 0548; initially C. fungifaber; NHMUK • soldier; Mamu forest reserve; 6 ° 11 ′ N, 7 ° 10 ′ E; 28 Feb. 1958; W. A. Sands leg.; study code: DJ U 058; initially C. fungifaber; NHMUK 13671862 • soldier; Enugu – Otukpo road; 6 ° 45 ′ N, 7 ° 28 ′ E; 5 Mar. 1958; W. A. Sands leg.; study code: DJ U 055; initially C. fungifaber; NHMUK 13671858 • soldier; Ayangba; 7 ° 30 ′ N, 7 ° 10 ′ E; 6 Mar. 1958; W. A. Sands leg.; study code: DJ U 059; initially C. fungifaber; NHMUK 13671857 • soldier; Ijebu – Ode road; 6 ° 49 ′ N, 3 ° 55 ′ E; 12 Dec. 1959; W. A. Sands leg.; study code: DJ U 060; initially C. fungifaber; NHMUK 13671859. Supplementary material of dubious I. fungifaber Four samples from the CAR, near Bangui, and two from eastern DRC (Fig. 45) show a soldier’s morphology entirely consistent with that of I. fungifaber. As these samples come from semi-deciduous forest and forest galleries (which is not characteristic of I. fungifaber) and as it was not possible to sequence them, these six samples, are presented here as “ dubious, possibly cryptic I. fungifaber ”. The distance between Bangui and the nearest harvesting point in Gabon is around 500 km, and between Bangui and eastern DRC, the distance is around 1300 km. Of the six old museum samples examined, four samples were initially labelled as C. fungifaber and two not identified. CENTRAL AFRICAN REPUBLIC • soldier; route Mbaïki – Bagandou; 3 ° 52 ′ N, 17 ° 52 ′ E; 21 Jun. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 188; MNHN EP 9825 • soldier; Boukoko; 3 ° 54 ′ N, 17 ° 55 ′ E; 2 Jul. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 189; MNHN EP 9826 • soldier; Bangui; 4 ° 27 ′ N, 18 ° 32 ′ E; 1975; Becker leg.; study code: DJ U 037; initially C. fungifaber; NHMUK 13671856 • soldier; Bangui; 4 ° 27 ′ N, 18 ° 32 ′ E; 1975; Becker leg.; study code: DJ U 038; initially C. fungifaber; NHMUK 13671855. DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (queen); “ Alto Uele, foresta ”; 2 ° 30 ′ N, 29 ° 30 ′ E; [no date]; S. Patrizi leg.; study code: DJ 0312; initially C. fungifaber in MCGD • soldier, ♀ (alate); Epulu; 1 ° 24 ′ N, 28 ° 34 ′ E; 15 May 1948; A. Emerson leg.; study code: DJ N 129; initially C. fungifaber in AMNH. Historical review The alate imago from Bonge [= Mbongé], Cameroon, was briefly described by Sjöstedt (1896: 297) under the name Eutermes fungifaber, mentioning it as very close to Eutermes mordax (Smeathman, 1781). Four years later, Sjöstedt (1900: 143 – 149) provided more detailed descriptions (text and figures) of all castes and nest. Sjöstedt (1926: 218 – 225) included Cubitermes fungifaber in imagines' and soldiers' keys of Cubitermes species and inserted it in a “ fungifaber - Gruppe ” of species with a not forward humped soldier’s frons. Emerson (1928: 517 – 519) referred to this species as Mirotermes (Cubitermes) fungifaber; he provided further descriptions and measurements of all castes, based on samples collected by G. Zenker in Cameroon. Snyder (1949: 159) catalogued this species under the name Cubitermes fungifaber in the sub-family Termitinae. Bouillon & Vincke (1971: 269) described the enteric valve of C. fungifaber as belonging to the first of three types, the “ simple type ” without any spatula. Cubitermes fungifaber was considered as a senior synonym of (a) C. banksi, (b) C. comstocki and (c) C. schmidti (Emerson unpublished “ Card catalog ”; Ruelle 1992: 501; Krishna et al. 2013: 1913, 1918, 1935). These three species are somewhat smaller than C. fungifaber, but synonymies are compatible with their enteric valve patterns (Josens & Deligne 2019: 39 – 42). To obtain genetic sequences of I. fungifaber, one of us (PA) went and collected samples at Mbongé, type location of I. fungifaber, and at Bipindi, type location of I. banksi, I. comstocki and I. schmidti; some of the soldiers in those samples were as small as the soldiers of C. comstocki and Hellemans et al. (2021: 233) confirmed their synonymies and placed the species in the restored genus Isognathotermes. Cubitermes fungifaber was considered a senior synonym of C. fungifaber var. elongata (Krishna et al. 2013: 1923) and a possible senior synonym of C. planifrons (Emerson in his unpublished “ card catalog ”, quoted by Krishna et al. 2013: 1931) but these synonymies are rejected because the two taxa proposed as junior synonyms do not have enteric valves of the fungifaber pattern, but well of the finitimus one (Josens & Deligne 2019: 42 – 44). Cubitermes aff. subarquatus “ spA ” and Cubitermes aff. subarquatus “ spD ” were mentioned by Roy et al. (2006) in La Lopé National Parc, Gabon; on a genetical base they were considered as cryptic species. Hellemans et al. (2021: 231) placed these species in the restored genus Isognathotermes as synonyms of I. fungifaber. As already mentioned, the geographic distribution of I. fungifaber is clearly dependant on humid, forest environments. In at least one documented case, its geographic distribution has been changing during the second half of the 20 th century, possibly in relation with global and local climate change. Isognathotermes fungifaber used to be common in the Banco National Park near Abidjan, RCI: see the publications by Noirot et al. (1986), Han & Lepage (1991), and personal observation (GJ, in 1970); however, nowadays, this national park is almost totally included within the city of Abidjan, which presumably influences the local climate. Moreover, the annual rainfall in Côte d’Ivoire has been declining by about 20 % during the last decades, mainly during the late sixties and early seventies (Servat et al. 1999), possibly linked with huge deforestation. This might explain that two of us (GJ and YR) could hardly find a single nest of this species in 2015 in the Banco National Park.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, fresh samples very dark, C 7 – C 8 (C 6 – C 7 in long preserved imagines); fontanelle concolorous or almost so with head capsule. Postclypeus C 6 – C 7 slightly paler (one level) than head capsule. Antennae C 5 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 8 (C 5 – C 7 in long preserved imagines), as postclypeus, generally one level paler than head capsule; meso- and metanotum C 5 – C 7, sometimes somewhat paler than pronotum. Legs C 3 – C 5; tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 3 – Cf 4), anterior veins darker. Abdomen: tergites C 5 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 ×, sometimes only at 80 ×. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 5 – 12 on the carina of fore coxa and 1 – 8 on the ventral side of fore coxa and trochanter; tibia pilose fore, mid, and hind tibia furnished with 30 – 40 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 5; Fig. 42). Size: the imagines of I. fungifaber are, on average, among the smallest of the genus Isognathotermes (Fig. 26). Head capsule: compound eyes shortly oval; ocelli shortly oval to oval, removed from eyes by a distance equal to 0.7 – 1.4 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: generally 16 articles on alate individuals, rarely 17 (one individual out of 16), always shortened by amputation by three to four articles in queens and kings. Labrum: cupola shaped, wider than long. Left mandible with apical tooth on average longer than in most other species of the genus (Fig. 27) and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a more or less sharp carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 3 – 5 and 7 – 11 branches respectively. Gut (only studied in some cases): enteric valve weakly developed and hardly comparable with workers’ and soldiers’ valves; odd PCs are slightly longer than even PCs (Fig. 6). Caecum present as a small amorphous button. Soldier COLOUR. Head capsule generally tending towards deep palette (Cd 4 – Cd 6) becoming fader and darker (e. g., Cf 5 – Cf 7) in long preserved samples; there is always a gradient from a darker frons to a paler back (e. g., from C 6 – C 4), smooth and extended in 55 % of the samples examined but more abrupt in the remaining 45 % giving in some cases the impression that the head capsule is bicolorous (as in Fig. 13). Antennae and labrum sometimes concolorous or one level paler than head capsule. Mandibles dark (C 6 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons; in long preserved samples, this clearing tends to disappear. Thorax and legs paler than head capsule (C 2 – C 4) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow to reddish-brown tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with some large setae on lobes. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta but not any spines (in 68 % of the samples examined) or furnished with 1 – 3 spines on carina and 0 – 2 on ventral side; trochanter generally with some long lined-up setae, sometimes with six or seven spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 15 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 5; Fig. 43). Size: the soldiers of I. fungifaber are among the smallest of the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with (in 86 % of the samples examined) a narrowing near posterior third or fourth; from antennal sockets sides converge more or less clearly towards bases of mandibles; posterior side regularly rounded or sometimes (in 29 % of the samples examined) with a short straight or even concave part in the middle. In profile: upper profile almost always slightly concave; angle between extended mandibles and frons varies from right to a little obtuse; frons with a sketched or without any anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either roundly convex or forming an acute widening or even a kind of ear on each side. Antennae: of 14.5 – 15 articles. Labrum: always deeply bifurcate and wider than long, with sides varying from lyre-shaped (in 84 % of the samples examined) to slightly convex; lobes angular, frequently with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like strongly curved (on average, I. fungifaber, and I. planifrons, have the highest left mandible apical curvature index in the genus Isognathotermes); inner edges generally smooth with one distinct but small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, as wide as 55 – 70 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed, sometimes (in 18 % of the samples examined) with a lobe expanded somewhat forward. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 20 % longer than the even cushions, generally without any crest or with crests very weakly developed. Fig. 43 (continued on next page). Isognathotermes fungifaber (Sjöstedt, 1896) and junior synonym species: soldiers. Lateral and dorsal views of a soldier’s head capsule, ventral view of its mandibles, and dorsal view of its labrum. A. From type colony of Eutermes fungifaber Sjöstedt, 1896 (DJ 0271). B. From type colony of Mirotermes (Cubitermes) banksi Emerson, 1928 (DJ 0287). C. From type colony of Mirotermes (Cubitermes) comstocki Emerson, 1928 (DJ 0273). Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and 1 – 4 on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 5, Fig. 44). Size: the workers of I. fungifaber are, on average, among the smallest of the genus Isognathotermes (but with large overlapping on several other species: Fig. 31). Head capsule: weakly sclerotised (except mandibles). Antennae of 14.5 (rarely 14) articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with Fig. 43 (continued). D. From type colony of Mirotermes (Cubitermes) schmidti Emerson, 1928 (DJ 0124). E. From a non-type colony of I. fungifaber (Sjöstedt, 1896) (DJ 0268). F. From a colony suspected to be a cryptic species of “ Cubitermes aff. subarquatus spA ” (DJ 0626). a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 57 – 70 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are as high as or higher than they are wide, with long and strong bristles; supporting bristles are generally numerous: Fig. 44 (continued on next page). Isognathotermes fungifaber (Sjöstedt, 1896) and junior synonym species: soldiers and workers. PC 1 s of soldier’s and worker’s enteric valves, worker’s caecum and foreleg. A. From type colony of Eutermes fungifaber Sjöstedt, 1896 (DJ 0271). B. From type colony of Mirotermes (Cubitermes) banksi Emerson, 1928 (DJ 0287). C. From type colony of Mirotermes (Cubitermes) comstocki Emerson, 1928 (DJ 0273). 14 – 38 on each side of the odd PCs; only three samples (out of 54) show an intermediate EVA between the fungifaber and finitimus patterns; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, generally with two or three (up to five) short lobes, sometimes (in 14 % of the samples examined) with a lobe expanded forward. Chorology-ecology The geographic distribution of I. fungifaber is clearly linked with humid, evergreen forests generally at less than 300 km from the Atlantic coast (Fig. 45): it includes from west to east the eastern Guinean forests, the Cross-Sanaga-Bioko coastal forests, the Atlantic Equatorial coastal forests, and Fig. 44 (continued). D. From type colony of Mirotermes (Cubitermes) schmidti Emerson, 1928 (DJ 0124). E. From a non-type colony of Isognathotermes fungifaber (Sjöstedt, 1896) (DJ 0268). F. From a colony suspected to be a cryptic species of “ Cubitermes aff. subarquatus spA ” (DJ 0626). the Northwestern Congolian lowland forests ecoregions. This distribution partly resembles that of I. planifrons (that has a different EVA). However, some samples of “ dubious I. fungifaber ” come from the northeastern Congolian lowland forests ecoregion without it being possible to guarantee the true species of these samples. Molecular data Thirteen mitogenomes of I. fungifaber are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). These were collected from RCI, Cameroon, and Gabon. Most samples exhibit less than 1 % dissimilarity with each other (Supp. file 4); except for sample DJ 0451 being the most dissimilar (1.50 %) collected in one of the type localities of the species, and for sample DJ 0412, collected at the westernmost part of its distribution (1.32 %). Our phylogenetic reconstructions based on the COII gene (Figure SF 2) includes samples identified as “ Cubitermes aff. subarquatus spA ” (DJ 0623 and DJ 0626) and “ spD ” (DQ 246541, DJ 0625) of Roy et al. (2006). The two mitogenomes of “ spA ” samples were successfully re-sequenced from the samples having produced the COII sequences published under accession DQ 127302 and DQ 127299 but not that of the “ spD ” sample. Therefore, the cryptic species status of the “ spD ” proposed by Roy et al. (2006) cannot be discussed here.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682454FF8CFDCFFBD8FCEBFE0B.taxon	discussion	Remarks Isognathotermes fungifaber is quite variable in size and much more in soldiers and workers than in imagines: we can deduct from Table 5 that variability in head width for I. fungifaber is 14 % in imagines, but 31 % in soldiers and 28 % in workers The soldiers of several samples have an intermediate morphology between I. fungifaber and I. severus (having the same kind of EVA): the easiest way to distinguish them is their origin since they seem to be mutually exclusive. This is linked with their preferred habitat: on the one hand, I. fungifaber seems to be restricted to evergreen forests with, however, one known exception: DJ 0625, that was proposed as a cryptic species (“ spD ”) by Roy et al. 2006: it was collected in the Doda gallery forest, surrounded with savannahs, north of La Lopé National Park, Gabon. On the other hand, I. severus is common in savanna landscapes of West and Central Africa (from Gambia to CAR) but also in orchards, secondary forests and, in some regions, it is also found in evergreen forest but only in places where I. fungifaber is absent as for example in RCI near the border with Liberia.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682445FF8AFE70FDCDFD46FDBB.taxon	description	Figs 26 – 31, 46, distribution map: Fig. 47; Table 7	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682445FF8AFE70FDCDFD46FDBB.taxon	diagnosis	Diagnosis The worker has a finitimus EVA and is among the large workers of the genus Isognathotermes (WHdW = 1.30 – 1.42 mm, Fig. 30) but rather difficult to distinguish from that of other species. The soldier has generally a recognizable finitimus EVA; it is, on average, and with the soldier of I. bulbifrons, the largest of the Isognathotermes species with a finitimus EVA (SHdL = 2.98 – 3.77 mm, Fig. 28), with more curved mandibles (SMlc = 0.24 – 0.36 mm, Fig. 29), as in I. planifrons; its head is slightly bulbous (SHdC = 0.0 – 0.07 mm), less than that of I. bulbifrons and similar to I. planifrons. The imago is, on average, and with the imago of I. bulbifrons, the largest in the genus Isognathotermes with a finitimus EVA (IHdW = 1.62 – 1.78 mm, Fig. 26); but its mandibular apico-marginal distance is generally smaller (IMlAmD = 0.20 – 0.24 mm, Fig. 27) than in I. bulbifrons (IMlAmD = 0.20 – 0.30 mm in I. bulbifrons). Isognathotermes magniplanifrons sp. nov., is therefore morphologically close to I. bulbifrons but their chorology is different: to date, I. magniplanifrons is only known from the north-east of the DRC (Haut-Uele and Kivu provinces).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682445FF8AFE70FDCDFD46FDBB.taxon	etymology	Etymology The name magniplanifrons is from the Latin magnus (large) and the species name planifrons, referring to the resemblance with I. planifrons but with larger individuals in all castes.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682445FF8AFE70FDCDFD46FDBB.taxon	materials_examined	Material examined Twenty-four samples from two locations. Of the 24 old museum samples, one was initially labelled as C. minitabundus, one as C. speciosus, and 22 as Cubitermes sp. Holotype DEMOCRATIC REPUBLIC OF THE CONGO • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 7 Sep. 1970; A. Wabo leg.; study code: DJ 0920; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059253. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♂ (alate), ♀ (queen); same data as for holotype; BE RMCA INS. Iso. 059935. Other material examined DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Moto; 3 ° 2 ′ N, 29 ° 30 ′ E; 1923; L. Burgeon leg.; study code: DJ 0070; initially C. speciosus in RMCA; BE RMCA INS. Iso. 059258 • soldier, worker; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 10 Apr. 1952; A. Wabo leg.; study code: DJ 0896; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059256 • soldier, worker, ♂ (alate); Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 4 Sep. 1970; A. Wabo leg.; study code: DJ 0894; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059252 • soldier, worker, ♂ (king); Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 4 Sep. 1970; A. Wabo leg.; study code: DJ 0919; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059251 • soldier, worker, ♂ (king), ♀ (queen); Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 6 Sep. 1970; A. Wabo leg.; study code: DJ 0929; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059254 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 21 Sep. 1970; A. Wabo leg.; study code: DJ B 262; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059238 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 21 Sep. 1970; A. Wabo leg.; study code: DJ B 263; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059239 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 21 Sep. 1970; A. Wabo leg.; study code: DJ B 264; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059240 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 21 Sep. 1970; A. Wabo leg.; study code: DJ B 265; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059241 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 20 Sep. 1970; A. Wabo leg.; study code: DJ B 267; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059236 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 20 Sep. 1970; A. Wabo leg.; study code: DJ B 268; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059237 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 20 Sep. 1970; A. Wabo leg.; study code: DJ B 269; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059243 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 25 Jan. 1972; A. Wabo leg.; study code: DJ B 272; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059246 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 28 Feb. 1972; A. Wabo leg.; study code: DJ B 303; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059249 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 13 Mar. 1972; A. Wabo leg.; study code: DJ B 304; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059250 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 31 Mar. 1972; A. Wabo leg.; study code: DJ B 261; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059235 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 13 Mar. 1972; A. Wabo leg.; study code: DJ B 274; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059248 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 1 Apr. 1972; A. Wabo leg.; study code: DJ B 273; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059247 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 9 May 1972; A. Wabo leg.; study code: DJ B 270; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059244 • soldier, ♀ (alate); Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 1972?; A. Wabo leg.; study code: DJ B 316; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059255 • soldier, ♀ (alate); Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 1972?; A. Wabo leg.; study code: DJ B 317; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059257 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 11 Sep. 1972; A. Wabo leg.; study code: DJ B 266; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059242 • soldier; Irangi; 1 ° 54 ′ S, 28 ° 27 ′ E; 15 Sep. 1972; A. Wabo leg.; study code: DJ B 271; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059245. Historical review This species is described here. It was collected in 1923 in the Haut-Uele Province, DRC (and misidentified), and in 1970 in Kivu, DRC. This species is known from 24 samples, 23 of which come from the same locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682445FF8AFE70FDCDFD46FDBB.taxon	description	Description Imago COLOUR. Head capsule: well sclerotised, dark, C 6 – C 7; fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 7 concolorous or slightly paler (one level) than head capsule. Antennae C 4 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 5 – C 6, as head capsule or slightly paler (one level); meso- and metanotum C 4 – C 6, as pronotum or slightly paler. Legs C 4 – C 5. Wings hyaline with brown to grey tinge (Cf 3), anterior veins darker. Abdomen: tergites C 4 – C 6. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule with some prominent setae, mainly near the eyes, and a higher density of shorter, finer setae everywhere. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at magnification 50 × or more). Thorax: pronotum with prominent setae mainly on margins and many shorter ones in middle; meso- and metanotum without any visible setae or with some fine, pale setae, arranged in a medio-longitudinal strip, visible at 40 ×; generally three to four larger setae on metanotum. Legs very pilose, furnished (among numerous fine setae) with some fine spines: 6 – 10 on the carina of fore coxa and 0 – 2 on the ventral side; tibia pilose; fore, mid, and hind tibia furnished with 20 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 7, Fig. 46). Size: the imagines of I. magniplanifrons sp. nov. are, on average, rather large (but not the largest of the genus Isognathotermes) (IHdW = 1.62 – 1.78 mm). Head capsule: compound eyes nearly round; fontanelle generally a tiny elongate marking; ocelli nearly round to oval removed from eyes by a distance about equal to 0.7 – 1.1 ocellus small diameter. Antennae: 16 articles on alate individuals, shortened by amputation by five to six articles in queens and kings. Labrum: cupola shaped. Left mandible with mandibular apico-marginal distance clearly smaller (IMlAmD = 0.20 – 0.24 mm) than in I. bulbifrons (Fig. 27); apical tooth longer and more prominent than first marginal; marginal teeth three in number but second one only suggested by a slight undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth longer and more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and slightly narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a sharp carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 3 and 12 – 13 branches, respectively. Soldier COLOUR. Head capsule generally C 6 – C 7, paler (C 5) in a long-preserved sample; with a gradient from a darker frons to a paler back (e. g., C 6 → C 4). Gulamentum concolorous with head. Antennae and labrum somewhat paler (1 – 2 levels) than head capsule. Mandibles dark (C 8) with or without a slight clearing on their bases (one palette level) which is generally the same colour as frons. Thorax, nota and legs paler than head capsule (C 4 – C 5). Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of hair-like setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 5 – 7 large setae on each lobe. Thorax: pro- and mesonotum with some setae located mainly on margins. Legs: fore coxa furnished with 1 – 2 spines on carina and not any one on ventral side; trochanter generally with some long lined-up setae, sometimes with two spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 6 – 15 spines. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 7; Fig. 46). Size: the soldiers of I. magniplanifrons sp. nov. are, on average, the largest of the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a narrowing near posterior third or fourth; from antennal sockets sides converge more or less clearly towards bases of mandibles. In profile: upper profile slightly concave (as in I. planifrons, much less than in I. bulbifrons); frons with no hump overhanging the fontanelle. Gulamentum in ventral view always more or less constricted in its posterior half, with sides of anterior part either forming an acute widening or even a kind of ear on each side; generally, with a flat posterior part from side view. Antennae: of 15 articles. Labrum: always deeply bifurcate and wider than long, with lyre-shaped sides; lobes angular, with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like; inner edges smooth with one small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, as wide as 57 – 63 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions are on average 14 % longer than the even cushions, generally without any hump or with humps weakly developed; however, the pilosity becoming abruptly very dense shows the place where a hump is expected (Fig. 46); secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed (2 – 4 small lobes). Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and 1 – 2 on ventral side; fore trochanter with 6 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 7, Fig. 46). Size: the workers of I. magniplanifrons sp. nov. are, on average, among the largest of the genus Isognathotermes (but with large overlapping on several other species: Fig. 30). Fore coxa flanged ventrally resulting in a sharp carina. Head capsule: weakly sclerotised (except mandibles). Antennae: 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge), third marginal with a blunt tip; premolar tooth with its proximal end hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 70 % of head width. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a high density of rather short bristles on a globular bulge; supporting bristles are generally numerous: 20 – 38 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, shortly lobed (3 – 4 small lobes). Chorology-ecology The geographic distribution of I. magniplanifrons sp. nov. is linked with the northeastern Congolian lowland forests ecoregion (Fig. 47). Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	description	Figs 2, 8 – 12, 26 – 31, 33, 48, distribution map: Fig. 49; Table 8	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	description	Josens & Deligne 2019: 60 – 61.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	diagnosis	Diagnosis This species is very close to I. ugandensis (and I. severus). The worker has a fungifaber EVA (Fig. 4) but in 3 % of the samples examined, it has an EVA intermediate between the fungifaber and finitimus patterns. It is among the medium-sized workers of the genus Isognathotermes (Fig. 30). As in I. ugandensis, the premolar tooth on left mandible is more apparent than in most other species. The soldier is among the large-sized soldiers in the genus Isognathotermes (Fig. 28), on average somewhat larger than that of I. ugandensis; it has a left mandible very slightly curved to almost straight at the base and generally less curved than the right mandible (SMlc / SMrc = 0.51 – 1.03, with a weaker asymmetry than in I. ugandensis: SMlc / SMrc = 0.92 – 1.14), resulting in a low soldiers’ left mandible apical curvature index (Fig. 29); its head is generally clearly bulbous (but this is shared with I. ugandensis and I. severus, and also with I. bulbifrons which has a finitimus EVA). The imago is among the medium- to large-sized imagines in the genus Isognathotermes (Figs 26, 65); the eyes and ocelli are relatively small, as a consequence, the eye to ocellus minimal distance index is high: IEyOcD / IOc d = 1.16 – 1.61 (vs 0.54 – 1.58 in I. ugandensis). The premolar tooth on left mandible is better seen than in most other species (but this is shared with I. severus and I. ugandensis). This species is also defined by its ecology and chorology: it is the southernmost species of the genus Isognathotermes, it has been found in forest-savannah mosaics and woodlands (miombo) to the south and east of the Congolian forests (Fig. 49).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	etymology	Etymology The epithet minitabundus from the Latin minitabundus (threatening) probably refers to a supposed threat.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	materials_examined	Material examined Thirty-six samples from 15 locations. Of the 24 old museum samples, 18 were initially labelled as C. minitabundus, four as C. ugandensis, one as Eutermes fungifaber and one as Cubitermes sp. Syntype of Eutermes (Cubitermes) minitabundus (Sjöstedt, 1913) DEMOCRATIC REPUBLIC OF THE CONGO • soldier; Lumata; 12 ° 0.05 ′ S, 27 ° 31 ′ E; 6 Jun. 1912; J. Bequaert leg.; study code: DJ 0129; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059280. Other material examined DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Hemptine Saint Benoît [= Bunkonde]; 6 ° 18 ′ S, 22 ° 31 ′ E; 1912; P. Callewaert leg.; study code: DJ 0140; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059284 • soldier, worker, ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 21 Jan. 1912; P. Callewaert leg.; study code: DJ 0094; initially Eutermes fungifaber in RMCA; BE RMCA INS. Iso. 059274 • soldier, worker; Hemptine Saint Benoît [= Bunkonde]; 6 ° 18 ′ S, 22 ° 32 ′ E; 26 Jan. 1912; P. Callewaert leg.; study code: DJ 0015; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059286 • soldier, worker, ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 26 Oct. 1921; P. Callewaert leg.; study code: DJ 0139; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059283 • soldier, worker, ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 26 Oct. 1921; P. Callewaert leg.; study code: DJ 0157; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059276 • soldier, worker, ♀ (alate), ♂ (alate), ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 27 Oct. 1921; P. Callewaert leg.; study code: DJ 0144; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059285 • soldier, worker, ♀ (alate), ♂ (alate), ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 27 Oct. 1921; P. Callewaert leg.; study code: DJ 0155; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059278 • soldier, worker, ♀ (alate), ♂ (alate), ♀ (queen); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 27 Oct. 1921; P. Callewaert leg.; study code: DJ 0156; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059275 • soldier, worker, ♀ (alate); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 27 Oct. 1921; P. Callewaert leg.; study code: DJ 0158; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059277 • soldier, worker, ♀ (alate); Luluabourg [= Kananga]; 5 ° 53 ′ S, 22 ° 26 ′ E; 27 Oct. 1921; P. Callewaert leg.; study code: DJ 0728; initially C. minitabundus; AMNH • soldier, worker; Upemba N. P., Lusinga; 8 ° 56.4 ′ S, 27 ° 8 ′ E; 17 Jul. 1947; G. F. De Witte et al. leg.; study code: DJ 0137; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059282 • soldier, worker, ♂ (king), ♀ (queen); Upemba N. P., Lusinga; 8 ° 56.4 ′ S, 27 ° 8 ′ E; 17 Jul. 1947; G. F. De Witte et al. leg.; study code: DJ 0138; initially C. minitabundus in RMCA; BE RMCA INS. Iso. 059281 • soldier, worker, ♂ (king); Keyberg (Lubumbashi); 11 ° 47 ′ S, 27 ° 25 ′ E; 21 Apr. 1948; A. Emerson leg.; study code: DJ 0729; initially C. minitabundus in AMNH • soldier, worker, ♂ (king); Likasi; 10 ° 59.3 ′ S, 26 ° 4.27 ′ E; 1 Dec. 1971; P. Vincke leg.; study code: DJ 0898; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059271 • soldier, worker, ♂ (alate); Kundelungu P. N.; 10 ° 27 ′ S, 27 ° 42 ′ E; 15 Oct. 1972; P. Vincke leg.; study code: DJ 0899; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059272 • soldier, worker; Kipopo; 11 ° 33 ′ S, 27 ° 21 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0908; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059273 • soldier, worker; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; 11 Feb. 2011; G. Josens leg.; study code: DJ 0010; BE RMCA INS. Iso. 059279 • soldier, worker; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; Jun. 2016; P. Kasangij leg.; study code: DJ 0348; GenBank nos MN 646704 (COI), MN 685906 (COII), MN 685967 (28 S), PQ 679238 (mitogenome); BE RMCA INS. Iso. 059270 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; 3 Jun. 2016; P. Kasangij leg.; study code: DJ B 008; BE RMCA INS. Iso. 059259 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; 3 Jun. 2016; P. Kasangij leg.; study code: DJ B 009; GenBank no PQ 679225 (mitogenome); BE RMCA INS. Iso. 059260 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; 3 Jun. 2016; P. Kasangij leg.; study code: DJ B 010; BE RMCA INS. Iso. 059261 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.3 ′ E; 3 Jun. 2016; P. Kasangij leg.; study code: DJ B 011; BE RMCA INS. Iso. 059262 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.18 ′ E; 14 Jun. 2016; P. Kasangij leg.; study code: DJ B 012; BE RMCA INS. Iso. 059263 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.18 ′ E; 14 Jun. 2016; P. Kasangij leg.; study code: DJ B 014; BE RMCA INS. Iso. 059264 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.2 ′ E; 14 Jun. 2016; P. Kasangij leg.; study code: DJ B 017; BE RMCA INS. Iso. 059265 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.2 ′ E; 14 Jun. 2016; P. Kasangij leg.; study code: DJ B 018; BE RMCA INS. Iso. 059266 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.2 ′ E; 22 Jun. 2016; P. Kasangij leg.; study code: DJ B 019; BE RMCA INS. Iso. 059267 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.2 ′ E; 22 Jun. 2016; P. Kasangij leg.; study code: DJ B 020; BE RMCA INS. Iso. 059268 • soldier; Kisangwe, Mikembo sanctuary; 11 ° 29.1 ′ S, 27 ° 39.2 ′ E; 2 Jul. 2016; P. Kasangij leg.; study code: DJ B 021; BE RMCA INS. Iso. 059269. ZAMBIA • soldier, worker; Kitwe, Mwekera Forest reserve; 12 ° 52 ′ S, 28 ° 21 ′ E; 25 Jan. 1956; W. G. H. Coaton leg.; study code: DJ 0696; initially C. minitabundus; NHMUK 13671922 • soldier, worker; Samfya; 11 ° 22 ′ S, 29 ° 33 ′ E; 27 Jan. 1957; W. G. H. Coaton leg.; study code: DJ 0665; initially C. ugandensis; NHMUK 13672010 • soldier, worker; Chembe; 11 ° 57.5 ′ S, 28 ° 44 ′ E; 28 Jan. 1957; W. G. H. Coaton leg.; study code: DJ 0685; initially C. minitabundus; NHMUK 13671920 • soldier; Ndola – Kitwe road; 12 ° 59 ′ S, 28 ° 24.5 ′ E; 30 Jan. 1957; W. G. H. Coaton leg.; study code: DJ 0694; initially C. minitabundus; NHMUK 13671921 • soldier, worker; Kasama – Mpika road; 11 ° 0 ′ S, 31 ° 0 ′ E; 19 Dec. 1969; M. G. Bingham leg.; study code: DJ 0695; initially C. minitabundus; NHMUK 13671925 • soldier, worker; Chililabombwe; 12 ° 23 ′ S, 27 ° 50 ′ E; 22 Dec. 1970; M. G. Bingham leg.; study code: DJ 0693; initially C. minitabundus; NHMUK 13671926. Historical review Sjöstedt (1913: 368 – 369) described the soldier of his species under the name Eutermes (Cubitermes) minitabundus. Schmitz (1916: 119) referred to this species under the name Cubitermes minitabundus. Sjöstedt (1924 a: 42) described the imago of Eutermes (Trinervitermes) carbo that was in fact the imago of Cubitermes minitabundus (see Sands 1965: 98 and Krishna et al. 2013: 1858). Sjöstedt (1926: 216 – 217) transferred the species in a new, monotypic genus (Isognathotermes). He provided figures of the soldier and a description of imago and worker. Snyder (1949: 161) catalogued this species under the name Cubitermes minitabundus and housed it in the sub-family Termitinae. Harris (1958: 13 – 14) provided measurements and figures of imago and soldier on basis of specimens collected in the Upemba National Park (Haut-Katanga Province, DRC). Williams (1966: 90 – 92) provided a detailed redescription of imago and soldier along with measurements and figures. Bouillon & Vincke (1971: 269) described the enteric valve of C. minitabundus as belonging to the first of three types, the “ simple type ” without any spatula. Krishna et al. (2013: 1927) referred to this species under the name Cubitermes minitabundus and housed it in the sub-family Cubitermitinae. Josens & Deligne (2019: 39 – 42) placed this species within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682443FF81FE3CFD9DFACCFC1D.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, fresh samples very dark, C 6 – C 8 (C 5 – C 7 in long preserved samples); fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 7 concolorous or slightly paler (one level) than head capsule. Antennae C 4 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 5 – C 7, as postclypeus, concolorous or one level paler than head capsule; meso- and metanotum C 4 – C 7, sometimes somewhat paler than pronotum. Legs C 3 – C 5; tibia usually concolorous with femur. Wings hyaline with brown to grey tinge (Cf 2 – Cf 3), anterior veins darker. Abdomen: tergites C 4 – C 6. Sternites appreciably paler in middle (C 2 – C 3) with both sides darker (C 3 – C 4); posterior sternites darker (C 3 – C 4) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 20 – 80 ×, but rarely even not visible at 80 ×; when present, these fine setae do not extend over the posterior margin; sometimes a small number of large setae are present on metanotum. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 8 – 12 on the carina of fore coxa and 0 – 6 on the ventral side of fore coxa and trochanter; tibia pilose fore, mid, and hind tibia furnished with 20 – 40 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 8; Fig. 48). Size: the imagines of I. minitabundus are among the medium- to large-sized species of the genus Isognathotermes (as I. ugandensis) (Fig. 26). Head capsule: compound eyes nearly round to oval; ocelli shortly to moderately oval, removed from eyes with a higher eye to ocellus minimal distance index: IEyOcD / IOc d = 1.16 – 1.61 (0.54 – 1.58 in I. ugandensis); fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on alate individuals, always shortened by amputation by two to five articles in queens and kings. Labrum: cupola shaped, wider than long. Left mandible with apical tooth medium to large sized (IMlAmD = 0.21 – 0.27 mm vs 0.19 – 0.27 in I. ugandensis, Fig. 27) and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end hardly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 4 and 10 – 12 branches, respectively. Gut not studied. Soldier COLOUR. Head capsule generally tending towards deep palette (Cd 4 – Cd 6) becoming fader (Cf 4 – Cf 5) in long preserved samples; there is generally a smooth gradient from a darker frons to a paler back. Antennae and labrum generally one to two palette levels paler than head capsule. Mandibles dark (C 6 – C 8) generally with an abrupt clearing on their bases (two levels) which is generally the same colour as frons; in long preserved samples, this clearing tends to disappear. Thorax and legs paler than head capsule (C 3 – C 5) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum with 5 – 8 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta and 1 – 3 spines (exceptionally 5) on carina and not any spine on ventral side; trochanter generally with some strong lined-up setae, or with 3 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 7 – 18 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 8; Fig. 48). Size: the soldiers of I. minitabundus are part of the medium to larger soldiers in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a clear or slight narrowing near posterior third or fourth; from antennal sockets sides converge towards bases of mandibles; posterior side regularly convex or sometimes with short straight or even concave parts. In profile: upper profile always slightly to strongly concave; angle between extended mandibles and frons obtuse; frons with an outlined or clear anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either roundly convex or more frequently forming an acute widening or even a kind of ear on each side. Antennae: of 14 – 15 articles. Labrum: always deeply bifurcate and wider than long, with sides varying from lyre-shaped (in 87 % of the samples examined) to almost straight; lobes angular, frequently with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like and rather clearly asymmetrical, the left mandible is slightly curved to almost straight and hooked distally and obviously less curved than the right mandible; inner edges generally smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible always clearly more curved than left. Thorax: pronotum sellate, as wide as 52 – 61 % of head width, with straight or indented anterior margin and posterior straight margins. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, three-lobed. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 18 % longer than the even cushions, generally without any crest or with crests very weakly developed. Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 4 – 6 spines on carina and 0 – 2 on ventral side; fore trochanter with 6 – 8 spines, all except one lined up; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 18 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 8, Fig. 48). Size: the workers of I. minitabundus are part of the medium-sized workers in the genus Isognathotermes (with large overlapping on several other species: Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14.5 (sometimes 14) articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal appearing as an undulated edge, reduced but still present in worn mandibles (more visible than in other species), third marginal tooth with a blunt tip; premolar tooth generally well visible (more than in other species); molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh and frequently with a wavy inner edge near the base; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 65 – 70 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are as high as or higher than they are wide, with long and strong bristles; 15 – 26 supporting bristles on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, generally with three to five short lobes. Chorology-ecology The geographic distribution of I. minitabundus is linked with the southern Congolian forest-savanna mosaic and the Central Zambezian Miombo woodland ecoregions; this is, with I. ugandensis malawii subsp. nov., the southernmost species of the genus Isognathotermes (Fig. 49). Molecular data The species I. minitabundus was previously referred to as “ I. afug ” (DJ 0348) by Hellemans et al. (2021), which was identified as a close species to I. ugandensis. Herein, we recognise this sample to belong to its own species, due to distinct morphologies and geographical distributions. Two mitogenomes of this species (both collected from Kisangwe, DRC, Haut-Katanga Province) are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). The difficulty of morphological delimitation between I. minitabundus and I. ugandensis is also reflected from analyses of their mitogenomes, with I. minitabundus placed as sister to I. ugandensis in a phylogenetic reconstruction (Fig. 33), and with whom they exhibit on average less than 2 % dissimilarity (Supp. file 4). However, they also exhibit less than 2 % dissimilarity with I. finitimus (Supp. file 4).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768244BFFBEFE23FEEBFD46FB60.taxon	description	Figs 26 – 31, 50, distribution map: Fig. 53; Table 9	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768244BFFBEFE23FEEBFD46FB60.taxon	diagnosis	Diagnosis This is a very small species (the smallest with a finitimus EVA), so small that the first sample was initially thought to come from an incipient colony, but a physogastric queen ruled out this possibility. The worker has a finitimus EVA and is among the smallest in the genus Isognathotermes: (WT 3 L = 1.12 – 1.20 mm), and its enteric valve is also the smallest (Fig. 31). The soldier is among the smallest in the genus Isognathotermes (SHdL = 2.42 – 2.66 mm), as small as the smallest I. fungifaber but with a finitimus EVA. It has evenly curved mandibles with a very low acceleration index of the mandible curvature between proximal and distal parts (ln (SMlpR) - ln (SMldR) = 0.96 – 1.44) as it is the case for some I. planifrons but with different sizes. The imago, however, is among the medium-sized imagines in the genus Isognathotermes (Fig. 26) and rather like those of I. finitimus, I. rectimalatus sp. nov., and I. planifrons. To date, this species is only known from the “ forêt des Abeilles ”, Gabon.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768244BFFBEFE23FEEBFD46FB60.taxon	etymology	Etymology The epithet modicus means ‘ modest’ in Latin. It refers to the small size of this species.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768244BFFBEFE23FEEBFD46FB60.taxon	materials_examined	Material examined Five samples from one location. Holotype GABON • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau and A. Robert leg.; study code: DJ 0862; MNHN EP 9886. Paratypes GABON • worker, ♂ (king), ♀ (queen); same data as for holotype; MNHN EP 9886. Other material examined GABON • soldier, ♀ (queen); Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 214; MNHN EP 9887 • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 216; MNHN EP 9888 • soldier, worker; Forêt des Abeilles; 0 ° 35 ′ S, 12 ° 25 ′ E; 1995; C. Bordereau and A. Robert leg.; study code: DJ 0858; MNHN EP 9884 • soldier, worker, ♂ (king), ♀ (queen); Forêt des Abeilles; 0 ° 35 ′ S, 12 ° 25 ′ E; 1995; C. Bordereau and A. Robert leg.; study code: DJ 0859; MNHN EP 9885. Historical review This species is described here. It was discovered by C. Bordereau and A. Robert in the so-called “ forêt des Abeilles ”, Gabon.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768244BFFBEFE23FEEBFD46FB60.taxon	description	Description Imago COLOUR. Head capsule: well sclerotised and dark C 6 – C 7; fontanelle concolorous with head. Postclypeus one level paler than head capsule. Antennae C 5 – C 6. Thorax: pronotum concolorous or almost so with head capsule; meso- and metanotum C 6 slightly paler than head. Legs C 4. Abdomen: tergites C 6. Sternites appreciably paler in the middle (C 3 – C 5) with both sides darker (C 5 – C 6) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 20 – 40 ×; sometimes with one or two large setae on posterior lobe. Legs very pilose, furnished (among numerous fine setae) with 7 – 11 stronger setae on the carina of fore coxa and 1 – 3 on the ventral side of fore coxa and trochanter; tibia pilose, fore, mid, and hind tibia furnished with 30 – 40 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 9, Fig. 50). Size: the imagines of I. modicus sp. nov. are, on average, medium sized in the genus Isognathotermes (Fig. 26). Head capsule: compound eyes nearly round; ocelli shortly oval, removed from eyes by a distance equal to 0.9 – 1.3 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: shortened to 12 articles by amputation of probably four articles in queens and kings. Labrum: cupola shaped, wider than long. Left mandible with apical tooth medium sized in the genus (Fig. 27) and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth almost totally obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a sharp carina. Gut not studied. Soldier COLOUR. Head capsule uniformly C 5. Antennae and labrum one to two palette levels paler than head capsule. Mandibles dark (C 6) with an abrupt clearing on their bases (one levels) which is generally the same colour as frons. Thorax and legs paler than head capsule (C 4). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum with 4 – 6 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa with at least one fine seta and 2 – 3 spines on carina and not any spine on ventral side; trochanter generally with some strong lined-up setae, including 3 – 4 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 12 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 9, Fig. 50). Size: the soldiers of I. modicus sp. nov. are part of the smallest soldiers in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a very slight narrowing near posterior third or fourth; from antennal sockets sides converge towards bases of mandibles; posterior side regularly convex. Slightly concave or straight upper profile; angle between extended mandibles and frons obtuse; frons flat or with a small anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part roundly convex. Antennae: of 14.5 – 15 articles. Labrum: always deeply bifurcate and wider than long, with lyre-shaped sides; lobes angular, with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like; inner edges smooth with one distinct but small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible slightly more curved than left. Thorax: pronotum sellate, as wide as 56 – 60 % of head width, with straight anterior and posterior margins. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum rather small, best seen in ventral view, near centre of abdomen, three-lobed. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions recognizable as a finitimus EVA. Worker COLOUR. Head capsule pale (C 2 – C 3). Antennae: proximal articles pale (C 2), distal articles two levels darker (C 4). Thorax, nota, and legs pale (C 2 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and not any one on ventral side; fore trochanter with 4 – 6 lined up spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 6 – 12 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 9, Fig. 50). Size: the workers of I. modicus sp. nov. are the smallest workers with a finitimus EVA, of the same size as the smallest workers of I. fungifaber (which have a fungifaber EVA). Head capsule: weakly sclerotised (except mandibles). Antennae: 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge and disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as about 64 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a high density of rather short bristles on a globular bulge (Fig. 50); supporting bristles are rather numerous: 20 – 28 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, shortly lobed (three or four small lobes). Chorology-ecology To date, this species is only known only from the “ Forêt des Abeilles ”, Gabon, at the limit of the northwestern Congolian lowland forests and the Atlantic equatorial coastal forest ecoregions (Fig. 53). Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682477FFBBFE02FB3BFEA8FA0A.taxon	description	Figs 26 – 31, 33, 51 – 52, distribution map: Fig. 53; Table 10	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682477FFBBFE02FB3BFEA8FA0A.taxon	diagnosis	Diagnosis Soldiers and workers of Isognathotermes phallicaecalis sp. nov. and I. phalloides sp. nov. share the same kind of phalloid cecum, extended forward in a finger-like process that can be swollen distally or capped or crowned (Figs 52, 56), which distinguishes them from all other species of Isognathotermes; they have morphologically similar soldiers, with a slight difference in the curvature of the mandibles (Fig. 22); however, they come from different ecosystems: continental evergreen forest in the case of I. phallicaecalis, forest galleries in the case of I. phalloides. The worker of I. phallicaecalis sp. nov. has a finitimus EVA and is among the small workers in the genus Isognathotermes. Its head is, on average, somewhat wider than that of I. phalloides sp. nov. (Fig. 30). The soldier’s EVA can generally be recognized as belonging to the finitimus pattern; on average it has longer mandibles than I. phalloides sp. nov.: SMlL = 2.17 – 2.53 mm (vs 1.99 – 2.36 mm in I. phalloides). The imago is unknown. This species is also defined by its ecology and chorology: to date it has been found only in forested environments of the northern Congo Republic and Cameroon (Fig. 53).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682477FFBBFE02FB3BFEA8FA0A.taxon	etymology	Etymology The epithet phallicaecalis from the Greek φαλλός (phallos, phallus) and the Latin caecalis (of the caecum) refers to the caecum phalloid morphology in workers and soldiers.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682477FFBBFE02FB3BFEA8FA0A.taxon	materials_examined	Material examined Twelve samples from three locations. Holotype CONGO REPUBLIC • soldier; Loundoungou; 2 ° 22.827 ′ N, 17 ° 4.226 ′ E; 4 Dec. 2017; Y. Roisin leg.; study code: DJ 0529; GenBank nos MN 646722 (COI) MN 685925 (COII) MN 685986 (28 S) PQ 679196 (mitogenome); BE RMCA INS. Iso. 059288. Paratypes CONGO REPUBLIC • soldier, worker; same data as for holotype; BE RMCA INS. Iso. 059936. Other material examined CAMEROON • soldier; Dja Rock; 3 ° 20.73 ′ N, 12 ° 42.84 ′ E; 30 Mar. 2015; J. Šobotnik leg.; study code: DJ B 320; BE RMCA INS. Iso. 059287. CONGO REPUBLIC • soldier, worker; Mokabi; 3 ° 8.796 ′ N, 16 ° 57.869 ′ E; 8 Dec. 2017; Y. Roisin leg.; study code: DJ 0527; GenBank no PQ 679190 (mitogenome); BE RMCA INS. Iso. 059290 • soldier, worker; Mokabi; 3 ° 8.795 ′ N, 16 ° 57.826 ′ E; 8 Dec. 2017; Y. Roisin leg.; study code: DJ 0528; BE RMCA INS. Iso. 059289 • soldier, worker; Loundoungou; 2 ° 22.929 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0759; GenBank no PV 564652 (mitogenome); BE RMCA INS. Iso. 059296 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0760; GenBank no PV 564657 (mitogenome); BE RMCA INS. Iso. 059291 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0761; GenBank no PQ 679202 (mitogenome); BE RMCA INS. Iso. 059297 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0762; BE RMCA INS. Iso. 059292 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0763; BE RMCA INS. Iso. 059295 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0764; BE RMCA INS. Iso. 059293 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0765; BE RMCA INS. Iso. 059298 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0766; GenBank no PV 564655 (mitogenome); BE RMCA INS. Iso. 059294. Historical review This species is described here; the imago is still unknown. It was recently discovered by three of us (JŠ, YR & SL) in Cameroon and Northern Congo.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682477FFBBFE02FB3BFEA8FA0A.taxon	description	Description Imago The imago is unknown. Soldier COLOUR. Head capsule tending towards faded palette (Cf 4 – Cf 5); there is a gradient from a darker frons to a paler back. Antennae and labrum concolorous with or somewhat paler than frons. Mandibles dark (C 7 – C 8) with an abrupt clearing on their bases (C 5 – C 6) which is concolorous with frons. Thorax and legs generally paler (C 3 – C 4) than head capsule. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellowish tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 3 – 6 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bear 1 – 3 spines on carina and none (rarely one) on ventral side; trochanter with 2 – 8 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 15 spines along their shaft. Abdomen: tergites with some large setae, mainly on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 10; Figs 51 – 52). Size: the soldiers of I. phallicaecalis sp. nov. are medium sized among of the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised, appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a clear narrowing near posterior fourth (Fig. 51), from antennal sockets sides converge more or less clearly towards bases of mandibles; posterior side regularly rounded or with a short straight middle part; upper profile concave. Angle between extended mandibles and frons a little obtuse; frons without any or with a week anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part forming an acute widening or a kind of ear on each side. Antennae: of 14.5 – 15 articles. Labrum: always deeply bifurcate and wider than long, with lyre-shaped sides; lobes angular, with fine, translucent tips; anterior margin concave. Mandibles: sabre-like with a middle-sized curvature in the genus Isognathotermes; inner edges generally smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible slightly more curved than left. Thorax: pronotum sellate, as wide as 59 – 64 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather well developed, best seen in ventral view, near centre of abdomen, as a typical finger-like process, extended forward and generally swollen distally or seeming to be capped (Figs 52, 56). Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being 20 – 25 % longer than the even cushions, with a pilosity becoming very dense distally showing the place where a hump would be expected (in comparison with the worker’s EVA). Worker COLOUR. Head capsule pale (C 2 – C 3). Antennae: proximal articles pale (C 2), distal articles two levels darker (C 4). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 3 – 6 spines on carina and 1 – 2 on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 6 – 15 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 10; Figs 51 – 52). Size: the workers of I. phallicaecalis sp. nov. are, on average, medium sized among the genus Isognathotermes (Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14.5 (rarely 14) articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with a blunt tip; premolar tooth with its proximal end not hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 66 – 71 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern (Fig. 5) with triradial symmetry: the odd PCs, in their downstream part, bear a higher density of rather short bristles on a globular bulge, the latter sometimes weakly developed; supporting bristles are numerous: 18 – 33 on each side of the odd PCs (Fig. 51); secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always well developed, best seen in ventral view, near centre of abdomen, as a finger-like process, extended forward and often swollen distally or seeming to be capped (Figs 52, 56). Chorology-ecology To date, this species is only known from two sites in the northern Congo Republic and one in Cameroon (in the northwestern Congolian lowland forest ecoregion: Fig. 53). Molecular data This species was previously published under the label “ pha 2 ” (DJ 0529), established as sister to I. planifrons (Hellemans et al. 2021). Six mitogenomes of I. phallicaecalis sp. nov. are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). The mitogenomes exhibit less than 1.21 % dissimilarity with each other (Supp. file 4); and a dissimilarity of up to 2.26 % with I. phalloides sp. nov. – the other samples exhibiting phalloid caeca – which justifies the creation of two different (new) species. Of note, I. phallicaecalis sp. nov. is paraphyletic with respect to I. similifinitimus sp. nov. (Fig. 33), to which mitogenomes are less than 1.50 % dissimilar. Even though these two species were found non-monophyletic, the clear morphological and anatomical differences (e. g., caecum) support their recognition as distinct species. Our use of (maternally inherited) mitogenomes only may have limited our ability to resolve their evolutionary relationships. Thus, we posit that their respective paraphylies may either stem from incomplete lineage sorting, or mitogenome introgression following hybridization. Future analyses leveraging nuclear loci will be crucial to shed further light on their relationships.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682472FFB3FE17F9CDFE16F9CD.taxon	description	Figs 26 – 31, 33, 54 – 56, distribution map: Fig. 53; Table 11	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682472FFB3FE17F9CDFE16F9CD.taxon	diagnosis	Diagnosis This species is close to I. phallicaecalis sp. nov.; like the latter, its soldier and worker are characterised by a phalloid caecum, extended forward in a finger-like process that can be swollen distally or capped or crowned (Figs 52, 56). The imago’s caecum, without being phalloid, is also more developed than in imagines of other species and takes the appearance of a forward-pointing comma. Isognathotermes phallicaecalis sp. nov. and I. phalloides sp. nov. have morphologically similar soldiers, with a slight difference in the curvature of the mandibles (Fig. 22); however, they come from different ecosystems: continental evergreen forest in the case of I. phallicaecalis, forest galleries in the case of I. phalloides. The worker has a finitimus EVA and is among the smallest workers in the genus Isognathotermes; its head is, on average, narrower than that of I. phallicaecalis sp. nov. (Fig. 30). The soldier can be distinguished from I. phallicaecalis sp. nov. by its shorter mandibles, SMlL = 1.99 – 2.36 mm (vs SMlL = 2.17 – 2.53 mm in I. phallicaecalis); its EVA can generally be recognized as belonging to the finitimus pattern. The imago is, on average and with I. fungifaber, the smallest in the genus Isognathotermes: IHdW = 1.38 – 1.54 mm (Fig. 26). This species is also defined by its ecology and chorology: all samples come from forest galleries south-east of the Congolian forests (Fig. 53).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682472FFB3FE17F9CDFE16F9CD.taxon	etymology	Etymology The epithet phalloides from the Greek φαλλός (phallos, phallus) and εἶδος (eidos, like) refers to the caecum morphology in workers and soldiers.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682472FFB3FE17F9CDFE16F9CD.taxon	materials_examined	Material examined Fifty-two samples from 17 locations. Of the 46 old museum samples examined, six were initially labelled as C. fungifaber, three as C. fungifaber var. elongata, five as C. gibbifrons, and 32 as Cubitermes sp. Holotype DEMOCRATIC REPUBLIC OF THE CONGO • soldier; Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0087; initially C. gibbifrons in RMCA; BE RMCA INS. Iso. 059341. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (alate), ♂ (alate); same data as for holotype; BE RMCA INS. Iso. 059937. Other material examined ANGOLA • soldier, worker; Quimucanda; 8 ° 5.14 ′ S, 15 ° 7.43 ′ E; 27 Nov. 2024; J. Šobotnik leg.; study code: DJ 0971; BE RMCA INS. Iso. 059427 • soldier, worker; Quimucanda; 8 ° 5.14 ′ S, 15 ° 7.43 ′ E; 27 Nov. 2024; J. Šobotnik leg.; study code: DJ 0972; BE RMCA INS. Iso. 059428. DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (alate), ♂ (alate); Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0088; initially C. gibbifrons in RMCA; BE RMCA INS. Iso. 059342 • worker; Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0089; initially C. gibbifrons in RMCA; BE RMCA INS. Iso. 059343 • soldier, worker, ♀ (alate), ♂ (alate); Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0090; initially C. gibbifrons in RMCA; BE RMCA INS. Iso. 059344 • soldier, worker, ♀ (queen); Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; 1905; E. Luja leg.; study code: DJ 0095; initially Eutermes fungifaber in RMCA; BE RMCA INS. Iso. 059338 • worker, ♂ (alate); Kondué; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0098; initially C. fungifaber var. elongata in RMCA; BE RMCA INS. Iso. 059339 • worker, ♀ (alate), ♂ (alate) ;; 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0099; initially C. fungifaber var. elongata in RMCA; BE RMCA INS. Iso. 059340 • worker, ♂ (alate); Kondué (?); 4 ° 58 ′ S, 23 ° 19 ′ E; ca 1905; E. Luja leg.; study code: DJ 0606; initially C. gibbifrons; IEAP • soldier, worker, ♀ (alate); Bandundu; 3 ° 19 ′ S, 17 ° 20 ′ E; 28 Aug. 1939; J. Ruelle leg.; study code: DJ 0879; initially C. fungifaber var. elongata in RMCA; BE RMCA INS. Iso. 059326 • soldier, worker, ♂ (king); Kinshasa Campus Lovanium) [= Unikin]; 4 ° 25 ′ S, 15 ° 18 ′ E; 25 Feb. 1960; A. Bouillon leg.; study code: DJ 0913; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059336 • soldier; Kinshasa Campus Lovanium) [= Unikin]; 4 ° 25.16 ′ S, 15 ° 18.55 ′ E; 5 Mar. 1960; unknown leg.; study code: DJ B 293; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059300 • soldier, ♀ (queen); Kenje; 4 ° 45 ′ S, 16 ° 57 ′ E; 14 Apr. 1964; G. Mathot leg.; study code: DJ B 299; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059332 • soldier; Route vers Sanda; 4 ° 33.75 ′ S, 12 ° 24.5 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ B 314; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059311 • soldier; Route vers Sanda; 4 ° 33.75 ′ S, 12 ° 24.5 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ B 315; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059312 • soldier, worker; Takundi; 4 ° 45 ′ S, 16 ° 33 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ 0880; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059328 • soldier, worker; Takundi; 4 ° 45 ′ S, 16 ° 33 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ 0883; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059321 • soldier, worker; Takundi; 4 ° 45 ′ S, 16 ° 33 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ 0884; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059322 • soldier, worker; Takundi; 4 ° 45 ′ S, 16 ° 33 ′ E; 19 Apr. 1964; G. Mathot leg.; study code: DJ 0885; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059324 • soldier; Kimwenza; 4 ° 28.5 ′ S, 15 ° 18.25 ′ E; 17 Mar. 1965; S. Kikomba leg.; study code: DJ B 294; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059299 • soldier; Kinshasa Campus Lovanium) [= Unikin]; 4 ° 25.16 ′ S, 15 ° 18.55 ′ E; 13 Mat. 1965; C. Nkakala leg.; study code: DJ B 292; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059307 • soldier, worker, ♂ (king); Maluku; 4 ° 21 ′ S, 15 ° 48 ′ E; 25 May 1967; C. Nkakala leg.; study code: DJ 0912; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059335 • soldier, worker, ♀ (queen); Ndjili; 4 ° 28 ′ S, 15 ° 21 ′ E; 1 Feb. 1968; A. Bouillon leg.; study code: DJ 0914; initially C. fungifaber in RMCA; BE RMCA INS. Iso. 059337 • soldier; Route vers Sanda; 4 ° 33.5 ′ S, 15 ° 24.5 ′ E; 19 Apr. 1969; G. Mathot leg.; study code: DJ B 302; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059306 • soldier; Kinshasa Campus Lovanium) [= Unikin]; 4 ° 25.16 ′ S, 15 ° 18.55 ′ E; 7 Aug. 1969; C. Nkakala leg.; study code: DJ B 290; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059305 • soldier; Kinshasa Campus Lovanium) [= Unikin]; 4 ° 25.16 ′ S, 15 ° 18.55 ′ E; 7 Aug. 1969; C. Nkakala leg.; study code: DJ B 291; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059310 • soldier, ♂ (alate); Ndjili; 4 ° 24 ′ S, 15 ° 25 ′ E; 5 Oct. 1970; A. Bouillon leg.; study code: DJ B 296; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059327 • soldier, worker, ♀ (alate); Ndjili; 4 ° 27.22 ′ S, 15 ° 20.6 ′ E; 5 Nov. 1970; A. Bouillon leg.; study code: DJ 0909; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059325 • soldier; Ndjili; 4 ° 24 ′ S, 15 ° 25 ′ E; 12 Jan. 1971; A. Wabo leg.; study code: DJ B 297; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059144 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 275; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059303 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 276; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059308 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 277; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059309 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 282; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059313 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 283; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059315 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 285; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059318 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 286; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059319 • soldier; Luki; 5 ° 39 ′ S, 13 ° 4 ′ E; 1 Apr. 1971; A. Bouillon leg.; study code: DJ B 287; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059314 • soldier; Kiemi; 5 ° 32 ′ S, 12 ° 49 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 308; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059316 • soldier; Kiemi; 5 ° 32 ′ S, 12 ° 49 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ B 309; initially Cubitermes 2 in RMCA; BE RMCA INS. Iso. 059317 • soldier, worker; Kiemi; 5 ° 16 ′ S, 12 ° 34 ′ E; 2 Apr. 1971; A. Bouillon leg.; study code: DJ 0891; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059329 • soldier; Kenje; 4 ° 12 ′ S, 15 ° 43 ′ E; 14 Juil. 1971; P. Vincke leg.; study code: DJ B 300; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059301 • soldier, ♂ (king); Menkao; 4 ° 12 ′ S, 15 ° 39 ′ E; 14 Jul. 1971; P. Vincke leg.; study code: DJ B 298; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059323 • soldier; Menkao; 4 ° 12 ′ S, 15 ° 43 ′ E; 14 Juil. 1971; P. Vincke leg.; study code: DJ B 301; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059302 • soldier, worker, ♂ (king); Menkao; 4 ° 10 ′ S, 15 ° 42 ′ E; 14 Jul. 1971; P. Vincke leg.; study code: DJ 0910; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059333 • soldier, worker, ♀ (queen); Menkao; 4 ° 10 ′ S, 15 ° 42 ′ E; 14 Jul. 1971; P. Vincke leg.; study code: DJ 0911; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059334 • soldier; Kimwenza; 4 ° 29 ′ S, 15 ° 18 ′ E; 20 Apr. 1973; P. Vincke leg.; study code: DJ B 295; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059304 • soldier, worker; Itaanda (Popokabaka); 5 ° 41 ′ S, 16 ° 40 ′ E; 22 Apr. 1973; P. Vincke leg.; study code: DJ 0881; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059330 • soldier, worker, ♂ (king), ♀ (queen); Itaanda (Popokabaka); 5 ° 41 ′ S, 16 ° 40 ′ E; 4 May 1973; P. Vincke leg.; study code: DJ 0882; BE RMCA INS. Iso. 059331 • soldier, worker; Duale; 4 ° 20 ′ S, 16 ° 5 ′ E; Jan. 2011; C. Kifukieto leg.; study code: DJ 0083; BE RMCA INS. Iso. 059345 • soldier, worker; Duale; 4 ° 20 ′ S, 16 ° 5 ′ E; Jan. 2011; C. Kifukieto leg.; study code: DJ 0141; BE RMCA INS. Iso. 059346 • soldier, worker; Bombo-Luméné; 4 ° 25.94 ′ S, 16 ° 2.83 ′ E; 22 Apr. 2017; C. Kifukieto leg.; study code: DJ 0459; GenBank nos MN 646723 (COI), MN 685926 (COII), MN 685987 (28 S), PQ 679184 (mitogenome); BE RMCA INS. Iso. 059320. Historical review This species is described here. Although several samples were present in museum collections under the names of C. gibbifrons, C. fungifaber, or C. fungifaber var. elongata, the species was not recognized until Josens & Deligne (2019: 42 – 44) placed it within the finitimus valve pattern group.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682472FFB3FE17F9CDFE16F9CD.taxon	description	Description Imago The description of the imagines is based on 14 Isognathotermes phalloides sp. nov. imagines (five female alates, six male alates, two queens, and one king) from nine colonies; only nine individuals concerning the wings. COLOUR. Head capsule: well sclerotised, very dark (C 6 – C 8), paler in long preserved samples; fontanelle concolorous with or slightly paler than head capsule. Postclypeus C 5 – C 7, often paler (one level) than head capsule. Antennae C 4 – C 6. Thorax: pronotum C 5 – C 7 as postclypeus, generally one level paler than head capsule; meso- and metanotum C 5 – C 7. Legs C 4 – C 5; tibia concolorous with or slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 2 – Cf 4), anterior veins darker. Abdomen: tergites C 5 – C 6. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 5); posterior sternites darker (C 4 – C 5) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 × or 80 ×; frequently some larger setae on the posterior lobe of metanotum. Legs very pilose, furnished (among numerous fine setae) with 8 – 10 stronger setae on fore coxa and 5 – 6 on trochanter; tibia pilose fore, mid, and hind tibia furnished with 20 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 11, Fig. 54). Size: the imagines of I. phalloides sp. nov. are among the smallest of the genus Isognathotermes (Fig. 26). Head capsule: compound eyes nearly round; ocelli shortly oval to oval, removed from eyes by a distance about equal to 0.9 – 1.5 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on alate individuals, shortened by amputation by three to four articles in queens and kings. Labrum: cupola shaped. Left mandible with apical tooth on average shorter than in most other species of the genus (Fig. 27) but always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 4 and 9 – 12 branches, respectively. Soldier COLOUR. Head capsule generally tending towards faded palette (Cf 4 – Cf 5) becoming darker (e. g., Cf 5 – Cf 6) in long preserved samples; there is always a gradient from a darker frons to a paler back, frequently abrupt giving in some cases the impression that the head capsule is bicolorous (as in Fig. 13). Antennae and labrum one or two levels paler than frons. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally paler than frons. Thorax and legs generally concolorous with head capsule (C 3 – C 5) or somewhat paler. Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellowish tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 4 – 5 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears 4 – 7 spines on carina but none on ventral side; trochanter generally with 1 – 4 spines and some lined-up setae; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 12 spines along their shaft. Abdomen: tergites with some large setae, mainly on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 11, Fig. 54). Size: the soldiers of I. phalloides sp. nov. are medium sized in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised, appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a narrowing (sometimes strong) near posterior fourth; from antennal sockets sides converge towards bases of mandibles; posterior side regularly rounded or sometimes with three short straight or even concave parts; upper profile concave. Angle between extended mandibles and frons always a little obtuse; frons generally without any anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part forming an acute widening or a kind of ear on each side. Antennae: of 15, rarely 14, articles. Labrum: always deeply bifurcate and wider than long, with sides generally lyre-shaped; lobes angular, with fine, whitish or translucent tips; anterior margin concave. Mandibles: sabre-like with a medium-sized curvature in the genus Isognathotermes; inner edges generally smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally more curved than left. Thorax: pronotum sellate, as wide as 57 – 63 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather well developed (as in the worker), best seen in ventral view, near centre of abdomen, as a typical finger-like process, extended forward and sometimes swollen distally or capped (Figs 52, 56). Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 20 % longer than the even cushions, with a pilosity becoming abruptly very dense showing the place where a crest would be expected (in comparison with the worker’s EVA). Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 6 spines on carina and 0 – 3 on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 15 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 11, Fig. 55). Size: the workers of I. phalloides sp. nov. are small sized in the genus Isognathotermes (Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14 – 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with a blunt tip; premolar tooth with its proximal end partly hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 66 – 73 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, bear a higher density of rather short bristles on a globular bulge (finitimus EVA, Fig. 5); supporting bristles are numerous: 19 – 32 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather well developed, best seen in ventral view, near centre of abdomen, as a typical finger-like process, extended forward and sometimes swollen distally or seeming to be capped (Fig. 56). Chorology-ecology This species is linked with the forest galleries of the south-western Congolian forest savanna mosaic ecoregion (Fig. 53). Molecular data Herein, one mitogenome of I. phalloides sp. nov. is published alongside this work (GenBank accession: see Supp. file 3 and ‘ Material examined’). The representative of I. phalloides sp. nov. exhibited a dissimilarity of up to 2.26 % with samples of I. phallicaecalis sp. nov. – the other species exhibiting a phalloid caecum –, and up to 2.43 % with samples of I. similifinitimus sp. nov. (see Supp. file 4). This sample was previously published under the label “ phal ” (DJ 0459), inserted within the species I. severus based on a phylogeny reconstructed from the COII gene (Hellemans et al. 2021). Herein, the same sample was re-sequenced for its mitogenome. The phylogeny based on COII extracted from mitogenomes again placed this sample within the species I. severus (Supp. file 2). However, the phylogenetic reconstruction from full mitogenomes places this species as sister to a clade containing finitimus + minitabundus + ugandensis + severus (Fig. 33), with low support values (SH-aLRT = 70.7; UFB = 82). Further analyses leveraging more samples and nuclear loci will be required to better ascertain its phylogenetic position.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768247AFFA5FDD1F990FDA5FC2E.taxon	description	Figs 26 – 31, 33, 57 – 59, distribution map: Fig. 47; Table 12	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768247AFFA5FDD1F990FDA5FC2E.taxon	diagnosis	Diagnosis This species is difficult to identify from its morphology since it is located near I. finitimus and I. phalloides sp. nov. It is obviously easier to identify from its COII sequence: I. planifrons makes a clade well apart from the finitimus clade (Hellemans et al. 2021: 231). Best recognition, if molecular data are not available, by a combination of worker’s EVA, soldier’s mandibles curvature, and geographic origin. The worker has a finitimus EVA (Fig. 5) and is among the medium-sized workers of the genus Isognathotermes (Fig. 30) but with a rather small-sized enteric valve (Fig. 31). The soldier has, on average and along with that of I. fungifaber, the highest left mandible apical curvature index; it is among the medium-sized soldiers of Isognathotermes (Fig. 28); on the odd PCs of its enteric valve, the pilosity becomes very dense showing the place where a bulge exists in the worker’s valve, its finitimus pattern is therefore recognizable. The imago is, on average, medium sized in the genus Isognathotermes (Fig. 26) with, on average, larger eyes and ocelli than I. finitimus. This species is also defined by its ecology and chorology: it has always been found in evergreen forests from Gabon to CAR, up to 1000 km from the Atlantic coast (Fig. 47). Some samples of I. planifrons are like I. fungifaber, but they have different EVAs. Some samples of I. planifrons are like I. phalloides sp. nov.; (Fig. 22) they have similar EVAs, but they have different caeca and geographical distributions. Many samples of I. planifrons are like I. finitimus, (Fig. 22) they have similar EVAs and caeca; but their geographical distributions are different, but in CAR (Figs 38, 47).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768247AFFA5FDD1F990FDA5FC2E.taxon	etymology	Etymology The epithet planifrons from the Latin planus (flat) and frons (forehead) refers to the soldier’s head morphology (not protruding frons).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768247AFFA5FDD1F990FDA5FC2E.taxon	materials_examined	Material examined Seventy-seven samples from 29 locations. Of the 41 old museum samples examined, only one, the type, was identified as C. planifrons (and subsequently renamed “ C. fungifaber ”); six samples were labelled as C. banksi, two as C. bulbifrons, one as C. finitimus, three as C. fungifaber, one as C. fungifaber var. elongata, six as C. gaigei, one as C. loubetsiensis, one as C. subarquatus, six as Cubitermes sp. and 13 not identified. This species also includes C. aff. subarquatus “ spB ” (in Roy et al. 2006). Syntypes of Cubitermes planifrons Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Mukimbungu; 5 ° 8.5 ′ S, 14 ° 2.3 ′ E; 25 Oct. 1904; K. E. Laman leg.; study code: DJ 0296; initially C. planifrons → C. fungifaber; AMNH. Krishna et al. (2013: 1931) report imagines in the AMNH syntype but the sample that was examined did not contain any. They also mention that another syntype subsample is deposited in NHRM, not examined. Paratypes from type colony of Cubitermes fungifaber var. elongata Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Mukimbungu; 5 ° 8.5 ′ S, 14 ° 2.3 ′ E; ca 1904; K. E. Laman leg.; study code: DJ 0270; initially C. fungifaber var. elongata; AMNH Krishna et al. (2013: 1931) mention that another syntype subsample is deposited in NHRM (not examined). Paratypes of Mirotermes (Cubitermes) gaigei Emerson, 1928 CAMEROON • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0274; initially C. gaigei → C. fungifaber; AMNH • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0288; initially C. gaigei → C. fungifaber; AMNH. Paratypes of Cubitermes loubetsiensis Sjöstedt, 1924 CONGO REPUBLIC • soldier; Loubetsi; 3 ° 12 ′ S, 12 ° 10 ′ E; 1924; Nilsson leg.; study code: DJ 0611; initially C. loubetsiensis → C finitimus; AMNH • soldier; Loubetsi; 3 ° 12 ′ S, 12 ° 10 ′ E; [no date]; Kranz leg.; study code: DJ 0281; initially C. loubetsiensis; NHMM. Other material examined ANGOLA • soldier, worker, ♀ (alate); Quimucanda; 8 ° 5.14 ′ S, 15 ° 7.43 ′ E; 28 Nov. 2024; J. Šobotnik leg.; study code: DJ 0973; BE _ RMCA _ INS. Iso. 059898. CAMEROON • soldier, worker; Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1903; G. Zenker leg.; study code: DJ 0961; initially Eutermes zenkeri; RBINS • soldier, worker, ♀ (queen); Bipindi; 3 ° 5 ′ N, 10 ° 24.5 ′ E; 1920; G. Zenker leg.; study code: DJ 0276; initially C. fungifaber; AMNH • soldier; 45 km from Abong-Mbang; 5 ° 16 ′ N, 17 ° 38 ′ E; 1 Jul. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 195; MNHN EP 9900 • soldier, worker, ♂ (alate); Réserve forestière de Nyong; 3 ° 33 ′ N, 11 ° 37 ′ E; 5 Dec. 1949; J. Birkett-Smith and J. Dahl leg.; study code: DJ 0678; initially C. subarquatus; NHMUK 13671991 • soldier; Eboufek; 3 ° 30 ′ N, 11 ° 53 ′ E; 1 Dec. 1962; G. Becker leg.; study code: DJ U 066; initially C. banksi; NHMUK 13671829 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 096; initially C. bulbifrons; NHMUK 13671844 • soldier; Akok; 3 ° 53 ′ N, 11 ° 57 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 097; initially C. bulbifrons; NHMUK 13671841 • soldier; Bilik; 3 ° 31 ′ N, 11 ° 17 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 067; initially C. banksi; NHMUK 13671832 • soldier, ♂ (king); Bilik; 3 ° 31 ′ N, 11 ° 17 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 091; initially C. banksi; NHMUK 13671827 • soldier, worker; Bilik; 3 ° 31 ′ N, 11 ° 17 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0705; initially C. banksi; NHMUK 13671830 • soldier; Ebogo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 093; initially C. banksi; NHMUK 13671833 • soldier; Ebogo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 119; initially C. gaigei; NHMUK 13671904 • soldier, worker; Ebogo; 3 ° 23 ′ N, 11 ° 28 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0703; initially C. banksi; NHMUK 13671834 • soldier; Mbalmayo; 3 ° 28 ′ N, 11 ° 31 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ U 118; initially C. gaigei; NHMUK 13671903 • ♀ (queen); Mbalmayo; 3 ° 31 ′ N, 11 ° 30 ′ E; ca 1995; P. Eggleton et al. leg.; study code: DJ 0724; initially C. gaigei; NHMUK • soldier, worker; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ 0754; GenBank no PQ 679239 (mitogenome); MNHN EP 9901 • soldier, worker; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ 0755; GenBank no PQ 679178 (mitogenome); MNHN EP 9902 • soldier; Dja forest; 3 ° 20.63 ′ N, 12 ° 43.03 ′ E; 30 Mar. 2015; J. Šobotnik leg.; study code: DJ B 321; BE RMCA INS. Iso. 059347 • soldier; Dja Rock; 3 ° 20.73 ′ N, 12 ° 42.84 ′ E; 30 Mar. 2015; J. Šobotnik leg.; study code: DJ B 319; BE RMCA INS. Iso. 059348 • soldier, worker; near Somalomo; 3 ° 20.7 ′ N, 12 ° 42.8 ′ E; 30 Mar. 2015; J. Šobotnik leg.; study code: DJ 0259; BE RMCA INS. Iso. 059356 • soldier, worker; Ebogo; 3 ° 22.95 ′ N, 11 ° 27.718 ′ E; 22 May 2017; Y. Roisin leg.; study code: DJ 0509; BE RMCA INS. Iso. 059361 • soldier, worker; Ebogo; 3 ° 22.772 ′ N, 11 ° 27.899 ′ E; 24 May 2017; Y. Roisin leg.; study code: DJ 0518; BE RMCA INS. Iso. 059362 • soldier, worker, ♂ (king); Ebogo; 3 ° 22.914 ′ N, 11 ° 27.8 ′ E; 26 May 2017; Y. Roisin leg.; study code: DJ 0519; GenBank no PQ 679176 (mitogenome); BE RMCA INS. Iso. 059363 • soldier, worker; Yolo-Chimpa; 5 ° 35.136 ′ N, 10 ° 53.301 ′ E; 30 May 2017; Y. Roisin leg.; study code: DJ 0508; GenBank nos MN 646726 (COI), MN 685929 (COII), MN 685990 (28 S); BE RMCA INS. Iso. 059364 • soldier, worker; Akometa; 3 ° 28.194 ′ N, 11 ° 32.814 ′ E; 6 Jun. 2017; Y. Roisin leg.; study code: DJ 0507; GenBank nos MN 646725 (COI), MN 685928 (COII), MN 685989 (28 S), PQ 679210 (mitogenome); BE RMCA INS. Iso. 059365 • soldier, worker, ♂ (king), ♀ (queen); Ebogo; 3 ° 22.935 ′ N, 11 ° 27.793 ′ E; 7 Jun. 2017; Y. Roisin leg.; study code: DJ 0511; BE RMCA INS. Iso. 059366 • soldier, worker; Nguinda; 4 ° 15 ′ N, 12 ° 11 ′ E; 23 Nov. 2023; J. Šobotnik leg.; study code: DJ 0954; GenBank no PV 564654 (mitogenome); BE RMCA INS. Iso. 059489 • soldier, worker; Nguinda; 4 ° 15 ′ N, 12 ° 11 ′ E; 25 Nov. 2023; J. Šobotnik leg.; study code: DJ 0951; GenBank no PV 564644 (mitogenome); BE RMCA INS. Iso. 059372 • soldier, worker; Nguinda; 4 ° 15 ′ N, 12 ° 11 ′ E; 25 Nov. 2023; J. Šobotnik leg.; study code: DJ 0953; GenBank no PV 564645 (mitogenome); BE RMCA INS. Iso. 059488 • soldier, worker; Nguinda; 4 ° 15 ′ N, 12 ° 11 ′ E; 3 Dec. 2023; J. Šobotnik leg.; study code: DJ 0950; GenBank no PV 564653 (mitogenome); BE RMCA INS. Iso. 059373 • soldier; Nguinda; 4 ° 15 ′ N, 12 ° 11 ′ E; 3 Dec. 2023; J. Šobotnik leg.; study code: DJ 0957; GenBank no PV 564659 (mitogenome); BE RMCA INS. Iso. 059155. CENTRAL AFRICAN REPUBLIC • soldier; Boukoko; 3 ° 54 ′ N, 17 ° 55 ′ E; 21 Jun. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 185; MNHN EP 9898 • soldier; Boukoko; 3 ° 54 ′ N, 17 ° 55 ′ E; 21 Jun. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 186; MNHN EP 9899 • soldier, worker, ♀ (alate); Bounguélé; 3 ° 57 ′ N, 16 ° 45 ′ E; 12 May 2007; N. Sion leg.; study code: DJ 0931; BE RMCA INS. Iso. 059355. CONGO REPUBLIC • soldier; Ouesso; 1 ° 37 ′ N, 16 ° 1 ′ E; 27 Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 168; MNHN EP 9896 • soldier, worker; Dimonika; 4 ° 12 ′ S, 12 ° 20 ′ E; 21 Apr. 1983; C. Noirot leg.; study code: DJ 0930; MNHN EP 9894 • soldier, worker, ♀ (queen); Loundoungou; 2 ° 22.973 ′ N, 17 ° 4.298 ′ E; 3 Dec. 2017; Y. Roisin leg.; study code: DJ 0526; GenBank nos MN 646728 (COI), MN 685931 (COII), MN 685992 (28 S); BE RMCA INS. Iso. 059357 • soldier, worker; Loundoungou; 2 ° 22.929 ′ N, 17 ° 4.301 ′ E; 4 Dec. 2017; Y. Roisin leg.; study code: DJ 0524; GenBank no PQ 679227 (mitogenome); BE RMCA INS. Iso. 059358 • soldier, worker; Mokabi; 3 ° 8.795 ′ N, 16 ° 57.826 ′ E; 6 Dec. 2017; Y. Roisin leg.; study code: DJ 0523; GenBank nos MN 646727 (COI), MN 685930 (COII), MN 685991 (28 S); BE RMCA INS. Iso. 059359 • soldier, worker; Mokabi; 3 ° 9.022 ′ N, 16 ° 58.214 ′ E; 8 Dec. 2017; Y. Roisin leg.; study code: DJ 0525; GenBank no PQ 679194 (mitogenome); BE RMCA INS. Iso. 059360 • soldier, worker; Loundoungou; 2 ° 22.93 ′ N, 17 ° 4.301 ′ E; Dec. 2018; S. Lenz leg.; study code: DJ 0767; GenBank no PV 564648 (mitogenome); BE RMCA INS. Iso. 059371. DEMOCRATIC REPUBLIC OF THE CONGO • soldier; Mekambo; 1 ° 1 ′ N, 13 ° 56 ′ E; 8 Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 171; MNHN EP 9897 • soldier; Forêt de Djidji; 0 ° 13 ′ N, 11 ° 48 ′ E; 26 Jan. 1957; P. - P. Grassé and C. Noirot leg.; study code: DJ P 162; GenBank no PQ 679182 (mitogenome); MNHN EP 9895 • soldier; Route Makokou-Mekambo; 0 ° 41 ′ N, 13 ° 2 ′ E; 6 Jun. 1962; J. Deligne leg.; study code: DJ U 099; initially C. gaigei; NHMUK 13671888 • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 218; MNHN EP 9891 • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 219; MNHN EP 9892 • soldier; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau (?) leg.; study code: DJ P 221; MNHN EP 9893 • soldier, worker; Forêt des Abeilles; 0 ° 35 ′ S, 12 ° 25 ′ E; 1995; C. Bordereau and A. Robert leg.; study code: DJ 0861; MNHN EP 9889 • soldier, worker; Forêt des Abeilles; 0 ° 19 ′ S, 12 ° 7 ′ E; 1995; C. Bordereau and A. Robert leg.; study code: DJ 0863; MNHN EP 9890 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 6 Mar. 1998; M. Harry leg.; study code: DJ P 236; MNHN EP 9910 • soldier; La Lopé N. P.; 0 ° 9 ′ S, 11 ° 36 ′ E; 7 Mar. 1998; M. Harry leg.; study code: DJ P 227; MNHN EP 9904 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 7 Mar. 1998; M. Harry leg.; study code: DJ P 247; MNHN EP 9915 • worker; La Lopé N. P.; 0 ° 12.7 ′ S, 11 ° 33.5 ′ E; 7 Mar. 1998; M. Harry leg.; study code: DJ 0632; GenBank nos DQ 246542 (COII), DQ 246524 (ITS 2), PQ 679220 (mitogenome); initially C. aff. subarquatus “ spB ” (cf. Roy et al. 2006); MNHN EP 9919 • soldier; La Lopé N. P.; 0 ° 9 ′ S, 11 ° 36 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ P 229; MNHN EP 9905 • soldier, worker; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 9 Mar. 1998; M. Harry leg.; study code: DJ 0624; GenBank nos DQ 127312 (COII), DQ 246520 (ITS 2), PQ 679183 (mitogenome); initially C. aff. subarquatus “ spB ” (cf. Roy et al. 2006); MNHN EP 9918 • soldier, worker; Doda (north of La Lopé N. P.); 0 ° 4.5 ′ S, 11 ° 25.5 ′ E; 10 Mar. 1998; M. Harry leg.; study code: DJ 0923; MNHN EP 9903 • soldier; La Lopé N. P.; 0 ° 12 ′ S, 11 ° 33 ′ E; 11 Mar. 1998; M. Harry leg.; study code: DJ P 233; MNHN EP 9908 • soldier; La Lopé N. P.; 0 ° 12 ′ S, 11 ° 33 ′ E; 11 Mar. 1998; M. Harry leg.; study code: DJ P 234; MNHN EP 9909 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 11 Mar. 1998; M. Harry leg.; study code: DJ P 248; MNHN EP 9916 • soldier; La Lopé N. P.; 0 ° 9 ′ S, 11 ° 41 ′ E; 15 Mar. 1998; M. Harry leg.; study code: DJ P 230; MNJN EP 9906 • soldier; La Lopé N. P.; 0 ° 9 ′ S, 11 ° 41 ′ E; 15 Mar. 1998; M. Harry leg.; study code: DJ P 231; MNHN EP 9907 • soldier, ♀ (queen); La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 15 Mar. 1998; M. Harry leg.; study code: DJ P 245; MNHN EP 9914 • worker, ♂ (king); La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 15 Mar. 1998; M. Harry leg.; study code: DJ P 250; MNHN EP 9917 • soldier, ♀ (queen); La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 16 Mar. 1998; M. Harry leg.; study code: DJ P 238; MNHN EP 9912 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 16 Mar. 1998; M. Harry leg.; study code: DJ P 240; MNHN EP 9913 • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 23 Mar. 1998; M. Harry leg.; study code: DJ P 237; MNHN EP 9911 • soldier, worker; Nkobissimo [= Nkolmengboua?]; 2 ° 14.3 ′ N, 11 ° 29.25 ′ E; 15 Apr. 2017; G. Trembleau leg.; study code: DJ 0455; GenBank nos MN 646724 (COI), MN 685927 (COII), MN 685988 (28 S); BE RMCA INS. Iso. 059354. Historical review Sjöstedt (1924 b: 255 – 256) briefly described the soldier of this species under the name Cubitermes planifrons. Two years later, Sjöstedt (1926: 243 – 244) provided more detailed descriptions of all three castes, housing the species in the sub-family Termitinae. He included (1926: 218 – 226) C. planifrons in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ fungifaber - Gruppe ” of species with a not protruding soldier’s frons. Snyder (1949: 162) catalogued this species under the name Cubitermes planifrons and housed it in the sub-family Termitinae. This is one of the species that Ruelle (1992: 500) called “ forgotten species ”. Krishna et al. (2013: 1931) housed Cubitermes planifrons in the sub-family Cubitermitinae. Following Emerson’s “ card catalog ”, they mentioned that this species might become a junior synonym of C. fungifaber; however, this synonymy could not be accepted because C. planifrons and C. fungifaber belong to different valve pattern groups: Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Cubitermes aff. subarquatus “ spB ” was mentioned by Roy et al. (2006) in La Lopé National Parc, Gabon. On a genetical base, it was considered a cryptic species. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes as synonym of I. planifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768247AFFA5FDD1F990FDA5FC2E.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, fresh samples very dark, C 7 – C 8 (C 6 – C 7 in long preserved imagines); fontanelle concolorous with or slightly paler than head capsule. Postclypeus C 6 – C 7 slightly paler (one level) than head capsule. Antennae C 5 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 7 as postclypeus, generally one level paler than head capsule; meso- and metanotum C 5 – C 6, concolorous with or somewhat paler than pronotum. Legs C 3 – C 5; tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 3 – Cf 4), anterior veins darker. Abdomen: tergites C 6 – C 7. Sternites appreciably paler in middle (C 2 – C 5) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 20 – 40 ×. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 6 – 10 on the carina of fore coxa and 1 – 4 on the ventral side of fore coxa and sometimes also some on trochanter; tibia pilose fore, mid, and hind tibia furnished with 15 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 12; Figs 26 – 27, 57). Size: the imagines of I. planifrons are, on average, among the medium-sized of the genus Isognathotermes (Fig. 26). Head capsule: compound eyes nearly round; ocelli nearly round to oval, removed from eyes by a distance about equal to 0.7 – 1.2 ocellus small diameter; fontanelle generally a tiny round marking. Antennae: 16 articles on alate individuals, always shortened by amputation in queens and kings. Labrum: cupola shaped. Left mandible with apical tooth on average rather small in comparison with other species of the genus (Fig. 27) and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a more or less sharp carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 3 – 4 and 10 – 11 branches respectively. Gut: enteric valve not studied. Soldier COLOUR. Head capsule generally tending towards deep palette (Cd 4 – Cd 5) becoming fader in long preserved samples; there is always a gradient from a darker frons to a paler back (e. g., from C 5 to C 4), smooth and extended in 60 % of the samples examined but more abrupt in the remaining 40 % giving the impression that the head capsule is bicolorous (as in Fig. 13). Antennae and labrum sometimes concolorous or one level paler than head capsule. Mandibles dark (C 6 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons. Thorax and legs paler than head capsule (C 2 – C 4) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with some large setae on lobes. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta and not any spines in some (10 %) samples examined but generally furnished with 1 – 4 spines on carina and no spines on ventral side; trochanter generally with some long lined-up setae, sometimes with five or six spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 15 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 12; Figs 28, 58). Size: the soldiers of I. planifrons are medium sized in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides subparallel with a narrowing near posterior third or fourth; from antennal sockets sides converge more or less clearly towards bases of mandibles; posterior side regularly rounded or (in 56 % of the samples examined) with a short straight part in the middle. Upper profile almost always slightly concave; angle between extended mandibles and frons always a little obtuse; frons generally (88 %) without any anterior hump, rarely with a slight hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either roundly convex or forming an acute widening or even (in 36 % of the samples studied) a kind of ear on each side. Antennae: of 15 articles. Labrum: always deeply bifurcate and wider than long, with sides varying from lyre shaped (in 92 % of the samples examined) to straight; lobes angular, frequently with fine, whitish or translucent tips; anterior margin concave. Mandibles: sabre-like, strongly curved (on average, I. fungifaber, and I. planifrons, have the highest left mandible apical curvature index in the genus Isognathotermes); inner edges generally smooth with one distinct but generally very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible generally slightly more curved than left. Thorax: pronotum sellate, as wide as 56 – 64 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a more or less sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 20 % longer than the even cushions, generally without any crest or with crests very weakly developed. Caecum always rather small, best seen in ventral view, near centre of abdomen, shortly lobed. Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and generally 1, rarely 2 or 3 spines on ventral side; fore trochanter with 5 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 12; Figs 30, 59). Size: the workers of I. planifrons are, on average, medium sized in the genus Isognathotermes with large overlapping on several other species (Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 63 – 76 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a high density of rather short bristles on a globular bulge; supporting bristles are generally numerous: 17 – 31 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, shortly lobed (generally three small lobes). Chorology-ecology The geographic distribution of I. planifrons is linked with humid, evergreen forest environments up to 800 km from the Atlantic coast: it includes the Atlantic equatorial coastal forests and the northwestern Congolian lowland forests ecoregions (Fig. 47). This distribution is partly similar to that of I. fungifaber (but with another EVA). Molecular data Sixteen mitogenomes of the morphological species I. planifrons are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). The morphological species I. planifrons appears to be genetically heterogeneous, with sample DJ P 141 differing from the others by a dissimilarity of over 2 %, which probably makes it a cryptic species, but as it is currently unique, it has not yet been described and was moved as I. aff. planifrons in the Incertae sedis chapter. Surprisingly, two samples of I. planifrons (DJ 0754 and DJ 0755) as well as I. aff. planifrons (DJ P 141) were collected almost in the same place (between Nsimi and Zoétélé, Cameroon). After having discarded DJ P 141 (which is slightly more similar to I. phallicaecalis sp. nov. and I. phalloides sp. nov.), the fifteen remaining mitogenomes exhibited a dissimilarity of up to 2.1 % with each other (Supp. file 4). Our present sampling includes full mitogenomes from Gabonese samples previously published under the appellation Cubitermes aff. subarquatus “ spB ” by Roy et al. (2006). This species “ spB ” was already reassigned to I. planifrons by Hellemans et al. (2021) based on COII phylogeny (accession DQ 127312).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246CFFA2FE0AFBF2FD9AFDB1.taxon	description	Figs 26 – 31, 33, 60, distribution map: Fig. 53; Table 13	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246CFFA2FE0AFBF2FD9AFDB1.taxon	diagnosis	Diagnosis The worker has a finitimus EVA (Fig. 5) and is among the large-sized workers of the genus Isognathotermes (Fig. 30) with large-sized enteric valve (Fig. 31); in particular, the odd PCs elongation index is very high (WVPOdd-Elg = 8.60 – 11.6). The soldier is medium sized in the genus Isognathotermes (Fig. 28) and has almost straight mandibles with a low left mandible apical curvature index (SMlc / SMlL = 0.05 – 0.07), as in I. minitabundus and I. ugandensis (Fig. 29) but the latter have a fungifaber EVA. The only known imago is medium sized, its compound eyes are also medium sized, but they are located far from the head margin and close to the ocelli. This species is also defined by its ecology and chorology: it has been found only in Angolan woodlands (Fig. 53).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246CFFA2FE0AFBF2FD9AFDB1.taxon	etymology	Etymology The epithet rectimalatus from the Latin rectus (straight) and malatus (mandible) refers to the straight mandibles of the soldier.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246CFFA2FE0AFBF2FD9AFDB1.taxon	materials_examined	Material examined Six samples from two locations. Holotype ANGOLA • soldier; Mussende; 10 ° 33.17 ′ S, 15 ° 39.492 ′ E; 21 Sep. 2019; M. Hasson leg.; study code: DJ 0719; GenBank no PQ 679198 (mitogenome); BE RMCA INS. Iso. 059376. Paratypes ANGOLA • soldier, worker; same data as for holotype; BE RMCA INS. Iso. 059938. Other material examined ANGOLA • soldier, worker; Mussende; 10 ° 33.17 ′ S, 15 ° 39.492 ′ E; Dec. 2018; M. Hasson leg.; study code: DJ 0771; GenBank no PQ 679237 (mitogenome); BE RMCA INS. Iso. 059379 • soldier, worker; Cangandala N. P.; 9 ° 49.33 ′ S, 16 ° 40 ′ E; 1 Mar. 2019; M. Viage leg.; study code: DJ 0864; GenBank no PQ 679197 (mitogenome); BE RMCA INS. Iso. 059374 • soldier, worker; Cangandala N. P.; 9 ° 49.33 ′ S, 16 ° 40 ′ E; 1 Mar. 2019; M. Viage leg.; study code: DJ 0867; GenBank no PQ 679203 (mitogenome); BE RMCA INS. Iso. 059375 • worker; Mussende; 10 ° 33.17 ′ S, 15 ° 39.492 ′ E; 13 Aug. 2019; M. Hasson leg.; study code: DJ 0710; GenBank no PQ 679233 (mitogenome); BE RMCA INS. Iso. 059377 • soldier, ♂ (alate); Mussende; 10 ° 33.17 ′ S, 15 ° 39.492 ′ E; 25 Oct. 2021; M. Hasson leg.; study code: DJ 0812; BE RMCA INS. Iso. 059378. Historical review The species is described here. It was discovered by two of us (MV & MiH) in the Angolan Miombo.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246CFFA2FE0AFBF2FD9AFDB1.taxon	description	Description Imago The imagines are very similar to those of I. finitimus. COLOUR. Head capsule: well sclerotised, very dark (C 7); fontanelle paler than head capsule. Postclypeus C 5, paler than head capsule. Antennae C 5. Thorax: pronotum C 7 as head capsule; meso- and metanotum C 6. Legs C 4 – C 5; tibia slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 3), anterior veins darker. Abdomen: tergites C 6. Sternites appreciably paler in middle (C 3) with both sides darker (C 5); posterior sternites darker (C 5) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum without any visible setae, even at 100 ×. Legs very pilose, furnished (among numerous fine setae) with 6 stronger setae on fore coxa and 4 – 5 on trochanter; tibia pilose fore, mid, and hind tibia furnished with 20 – 35 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs, respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 13; Figs 26, 60). Size: the only known imago of I. rectimalatus sp. nov. is among the medium sized of the genus Isognathotermes (Fig. 26). Head capsule: compound eyes shortly oval; ocelli shortly oval, removed from eyes by a distance about equal to 0.7 – 0.8 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on the single known alate. Labrum: cupola shaped. Left mandible apical tooth always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa weakly flanged ventrally resulting in an unclear carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 3 and 9 branches, respectively. Soldier COLOUR. Head capsule C 4 – C 5; frons C 5 – C 6. Gulamentum somewhat darker than head. Antennae and labrum concolorous with head capsule. Mandibles dark (C 7 – C 8) with an abrupt clearing on their bases (two palette levels) which is the same colour as frons. Thorax, nota and legs somewhat paler than head capsule (C 3 – C 4). Abdomen grey to red-brown owing to digestive bolus, rarely with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum with some large setae on lobes. Thorax: pro- and mesonotum with some setae mainly located on margins. Legs: fore coxa furnished with 1 – 3 spines on carina and 0 – 1 on ventral side; trochanter with 3 – 6 long lined-up spines; fore, mid-, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 9 – 15 spines. Abdomen: tergites with some large setae, mainly on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 13, Figs 28, 60). Size: the soldiers of I. rectimalatus sp. nov. are medium sized in the genus Isognathotermes (Fig. 28). Head capsule: clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a slight narrowing near posterior third; from antennal sockets sides converge clearly towards bases of mandibles; posterior side regularly convex. In profile: upper profile concave; frons: fontanelle overhanged by a sketched frontal hump. Gulamentum in ventral view constricted in its posterior half, with sides of anterior part forming an acute widening with a kind of ear on each side. Antennae of 15 articles. Labrum: deeply bifurcate and wider than long, with sides lyre-shaped; lobes angular or truncated, with fine, whitish or translucent tips; anterior margin concave. Mandibles: sabre-like, almost straight at the base and slightly curved distally; left mandible apical curvature index very low (SMlc / SMlL <0.075); inner edges smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Thorax: pronotum sellate, clearly narrower than head, sometimes with a slight notch in anterior margin and entire posterior margin. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions are 19 % longer than the even cushions, with humps weakly developed; secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, best seen in ventral view, near centre of abdomen, lobed (three small lobes). Worker COLOUR. Head capsule pale (C 1 – C 3). Antennae: proximal articles pale (C 2), distal articles two levels darker (C 4). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 2 – 4 spines on carina and 0 – 2 on ventral side; fore trochanter with 7 – 9 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 16 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 13, Figs 30 – 31, 60). Size: the workers of I. rectimalatus sp. nov. are medium to large sized in the genus Isognathotermes (Fig. 30). Head capsule weakly sclerotised (except mandibles). Antennae 14 – 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed (WMlAmD> 0.19 mm) with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge), third marginal with a blunt tip; the premolar tooth is best seen than in most other species; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 63 – 70 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a very high density of rather short bristles on a globular bulge; supporting bristles are generally numerous: 25 – 33 on each side of the odd PCs; worker’s valve very large (WVP-AvL> 0.72 mm, Fig. 31) odd PCs elongation index very high (WVPOdd-Elg> 8.5); secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, shortly lobed (three small lobes). Chorology-ecology This species is only known from Angola; its geographic distribution is linked with the Angolan Miombo woodland ecoregion (Fig. 53). Molecular data The five mitogenomes of I. rectimalatus sp. nov. exhibited less than 2 % dissimilarity with each other (Supp. file 4: GenBank accessions: see supplementary Supp. file 3 and ‘ Material examined’). After I. acristatus sp. nov., the species I. rectimalatus sp. nov. was established as sister to all remaining Isognathotermes members (Fig. 33).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246BFF56FDE1FD64FB4AFD6D.taxon	description	Figs 26 – 31, 33, 61 – 63, distribution maps: Figs 45, 49; Table 14	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246BFF56FDE1FD64FB4AFD6D.taxon	diagnosis	Diagnosis This is the Isognathotermes species with the largest geographic distribution: from Gambia to CAR (Figs 45, 49). The worker has a fungifaber EVA but in 12 % of the samples examined, it has an EVA intermediate between the fungifaber and finitimus patterns; it is among the medium-sized workers of the genus Isognathotermes (Fig. 30). Its premolar tooth on left mandible is generally better seen than in most other species. The soldier is, on average, among the medium-sized soldiers in the genus Isognathotermes (Figs 21, 28) and its EVA, generally without any crest, looks like a muneris EVA; its head is generally bulbous but this is shared with I. minitabundus and I. ugandensis (and with I. bulbifrons that has a finitimus EVA). The imago is, on average, among the middle- to large-sized imagines in the genus Isognathotermes (Fig. 26). Confusion may occur mainly with I. fungifaber in West Africa and with I. ugandensis in central Africa. (see discussion about the ambiguities between I. severus and I. ugandensis ugandensis). This species has been found in various ecosystems (see chorology).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246BFF56FDE1FD64FB4AFD6D.taxon	etymology	Etymology The epithet severus means ‘ serious, earnest’ in Latin.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246BFF56FDE1FD64FB4AFD6D.taxon	materials_examined	Material examined One hundred fifty-seven samples from 88 locations. Of the 101 old museum samples examined, 42 were initially labelled as Cubitermes severus, two as C. severus var. modestior, one as C. fungifaber, 15 as C. gaigei, six as C. silvestrii, four as C. speciosus, six as C. ugandensis, one as Cubitermes sp. and 24 not identified. Syntypes of Cubitermes severus Silvestri, 1914 GUINEA • soldier, worker, ♀ (queen); Kakoulima; 9 ° 46 ′ N, 13 ° 27 ′ W; Aug. 1912; F. Silvestri leg.; study code: DJ 0608; initially C. severus; IEAP • soldier, worker, ♀ (queen); Camayenne; 9 ° 31.83 ′ N, 3 ° 41.17 ′ W; 16 Oct. 1912; F. Silvestri leg.; study code: DJ 0609; initially C. severus; IEAP • soldier, worker; Kakoulima; 9 ° 46 ′ N, 13 ° 27 ′ W; before 1904; F. Silvestri leg.; study code: DJ 0292; initially C. severus; AMNH • soldier, worker; Camayenne; 9 ° 31.8 ′ N, 13 ° 41.5 ′ W; 16 Oct. 1912; F. Silvestri leg.; study code: DJ 0293; initially C. severus; AMNH Syntypes of Cubitermes severus var. modestior Silvestri, 1914 GUINEA • soldier, worker, ♂ (king); Kindia; 10 ° 3 ′ N, 12 ° 52 ′ W; 20 Aug. 1912; F. Silvestri leg.; study code: DJ 0294; initially C. severus var. modestior; AMNH • soldier, worker, ♀ (queen); Kindia; 10 ° 3 ′ N, 12 ° 52 ′ W; 20 Aug. 1912; F. Silvestri leg.; study code: DJ 0341; initially C. severus var. modestior; IEAP. Krishna et al. (2013: 1927) mention that another subsample is deposited in PPRI (not examined). Syntypes of Cubitermes silvestrii Sjöstedt, 1925 GUINEA • soldier, worker; Kakoulima; 9 ° 46 ′ N; 13 ° 27 ′ W), Fr. Guinea [= Guinea]; ca 1912; F. Silvestri leg.; study code: DJ 0302; initially C. silvestrii; AMNH. Krishna et al. (2013: 1937) mention that another subsample is deposited in NHRM (not examined). Other material examined BENIN • soldier, worker; Birni (forêt classée); 10 ° 1 ′ N, 1 ° 32 ′ E; Nov. 2016; L. E. Loko leg.; study code: DJ 0402; GenBank no PQ 679191 (mitogenome); BE RMCA INS. Iso. 059446 • soldier, worker, ♀ (queen); Birni (forêt classée); 10 ° 1 ′ N, 1 ° 32 ′ E; Nov. 2016; L. E. Loko leg.; study code: DJ 0403; GenBank nos MN 646735 (COI), MN 685940 (COII), MN 686001 (28 S); BE RMCA INS. Iso. 059448 • soldier, worker; Birni (forêt classée); 10 ° 1 ′ N, 1 ° 32 ′ E; Nov. 2016; L. E. Loko leg.; study code: DJ 0404; GenBank no PQ 679230 (mitogenome); BE RMCA INS. Iso. 059445 • soldier, worker; Birni (forêt classée); 10 ° 1 ′ N, 1 ° 32 ′ E; Nov. 2016; L. E. Loko leg.; study code: DJ 0405; GenBank no PQ 679200 (mitogenome); BE RMCA INS. Iso. 059449 • soldier, worker; Birni (forêt classée); 10 ° 1 ′ N, 1 ° 32 ′ E; Nov. 2016; L. E. Loko leg.; study code: DJ 0406; GenBank no PQ 679221 (mitogenome); BE RMCA INS. Iso. 059447. BURKINA FASO • soldier, worker; Bama; 11 ° 21 ′ N, 4 ° 23 ′ W; Feb. 2012; S. Traoré leg.; study code: DJ 0052; BE RMCA INS. Iso. 059412. CAMEROON • soldier, worker, ♂ (alate); Mont Guendérou; 7 ° 10 ′ N, 14 ° 14 ′ E; before 1914?; Riggenbach leg.; study code: DJ 0324; initially C. silvestrii; AMNH • soldier, worker, ♀ (alate), ♂ (alate); Ekoua; 3 ° 52.6 ′ N, 12 ° 15.5 ′ E; 28 Mar. 2015; J. Šobotnik leg.; study code: DJ 0249; GenBank no PQ 679208 (mitogenome); BE RMCA INS. Iso. 059401 • soldier, worker, ♀ (alate), ♂ (alate); Batié; 5 ° 17 ′ N, 10 ° 14.7 ′ E; 7 Apr. 2015; J. Šobotnik leg.; study code: DJ 0250; BE RMCA INS. Iso. 059402 • soldier, worker, ♀ (alate), ♂ (alate); Koutaba; 5 ° 39.3 ′ N, 10 ° 46.4 ′ E; 8 Apr. 2015; J. Šobotnik leg.; study code: DJ 0251; GenBank no PQ 679250 (mitogenome); BE RMCA INS. Iso. 059403 • soldier, worker; Koutaba; 5 ° 39.3 ′ N, 10 ° 46.4 ′ E; 8 Apr. 2015; J. Šobotnik leg.; study code: DJ 0258; GenBank no PQ 679229 (mitogenome); BE RMCA INS. Iso. 059407 • soldier, worker, ♀ (alate), ♂ (alate); Boulourou; 4 ° 48.8 ′ N, 11 ° 0.1 ′ E; 10 Apr. 2015; J. Šobotnik leg.; study code: DJ 0252; BE RMCA INS. Iso. 059404 • soldier, worker; Koutaba; 5 ° 35.988 ′ N, 10 ° 44.973 ′ E; 31 May 2017; Y. Roisin leg.; study code: DJ 0530; GenBank no PQ 679166 (mitogenome); BE RMCA INS. Iso. 059408 • soldier, worker; Mancha; 5 ° 33.534 ′ N, 10 ° 52.836 ′ E; 1 Jun. 2017; Y. Roisin leg.; study code: DJ 0510; BE RMCA INS. Iso. 059409 • soldier, worker; Koutaba; 5 ° 35.816 ′ N, 10 ° 45.856 ′ E; 2 Jun. 2017; Y. Roisin leg.; study code: DJ 0531; GenBank no PQ 679168 (mitogenome); BE RMCA INS. Iso. 059410 • soldier, worker; Kimi; 6 ° 5 ′ N, 11 ° 29 ′ E; 30 Nov. 2023; J. Šobotnik leg.; study code: DJ 0949; GenBank no PV 564658 (mitogenome); BE RMCA INS. Iso. 059405 • soldier, worker; Kimi; 6 ° 5 ′ N, 11 ° 29 ′ E; 30 Nov. 2023; J. Šobotnik leg.; study code: DJ 0952; GenBank no PV 564651 (mitogenome); BE RMCA INS. Iso. 059406. CENTRAL AFRICAN REPUBLIC • soldier; Bouala; 6 ° 22.8 ′ N, 15 ° 37 ′ E; 8 Jan. 1914; Elbert leg.; study code: DJ U 006; initially C. severus; NHMUK 13671956 • soldier, ♀ (alate); Bosoum; 6 ° 19 ′ N, 16 ° 22 ′ E; 1 May 1914; G. Tessmann leg.; study code: DJ U 016; initially C. severus; NHMUK 13671955 • soldier; Boukoko; 3 ° 54 ′ N, 17 ° 55 ′ E; 2 Jul. 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ P 187; MNHN EP 9939 • soldier; 20 km N of Bangui; 4 ° 37 ′ N, 18 ° 33 ′ E; Nov. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 174; MNHN EP 9928 • soldier; 20 km N of Bangui; 4 ° 36 ′ N, 18 ° 31 ′ E; Nov. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 177; MNHN EP 9931 • soldier; Bossembélé; 5 ° 16 ′ N, 17 ° 38 ′ E; 21 Nov. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 175; MNHN EP 9929 • soldier; Bossembélé; 5 ° 16 ′ N, 17 ° 38 ′ E; 21 Nov. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 181; MNHN EP 9935 • soldier; 70 km from Alindao; 5 ° 30 ′ N, 20 ° 49 ′ E; Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 180; MNHN EP 9934 • soldier; 70 km from Alindao; 5 ° 30 ′ N, 20 ° 49 ′ E; Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 184; MNHN EP 9938 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 178; MNHN EP 9932 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 179; MNHN EP 9933 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 182; MNHN EP 9936 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; 4 Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 183; MNHN EP 9937 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; 13 Dec. 1956; P. - P. Grassé and C. Noirot leg.; study code: DJ P 176; MNHN EP 9930 • soldier; Bambari; 5 ° 46 ′ N, 20 ° 40 ′ E; 1975; P. Eggleton et al. leg.; study code: DJ U 070; initially C. ugandensis; NHMUK 13671994 • soldier; near Bangui; 4 ° 23 ′ N, 18 ° 34.5 ′ E; 1975; Becker leg.; study code: DJ U 111; initially C. gaigei; NHMUK 13671894 • soldier, worker, ♂ (alate); Boika (Mbaïki); 3 ° 52 ′ N, 17 ° 58 ′ E; 11 Apr. 2007; N. Sion leg.; study code: DJ 0934; GenBank no PV 564656 (mitogenome); BE RMCA INS. Iso. 059399 • soldier, worker; Mopiti (Mbaïki); 3 ° 52 ′ N, 17 ° 58 ′ E; 14 Apr. 2007; N. Sion leg.; study code: DJ 0932; GenBank no PV 564660 (mitogenome); BE RMCA INS. Iso. 059418 • soldier, worker; Mopiti (Mbaïki); 3 ° 52 ′ N, 17 ° 58 ′ E; 14 Apr. 2007; N. Sion leg.; study code: DJ 0933; BE RMCA INS. Iso. 059417 • soldier, worker; Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 23 Apr. 2007; N. Sion leg.; study code: DJ 0004; BE RMCA INS. Iso. 059400 • soldier, worker; Gbasabangui (Bangui); 4 ° 22.5 ′ N, 18 ° 34.75 ′ E; 3 May 2007; N. Sion leg.; study code: DJ 0935; GenBank no PV 564646 (mitogenome); BE RMCA INS. Iso. 059452 • soldier, worker; Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 27 May 2007; N. Sion leg.; study code: DJ 0001; BE RMCA INS. Iso. 059450 • soldier, worker; near Bangui; 4 ° 23 ′ N, 18 ° 34.5 ′ E; 30 Oct. 2010; G. Josens leg.; study code: DJ 0367; BE RMCA INS. Iso. 059414 • soldier, worker, ♀ (alate); Mbaïki; 3 ° 55 ′ N, 18 ° 0 ′ E; 24 Apr. 2018; S. P. Wango leg.; study code: DJ 0586; BE RMCA INS. Iso. 059395 • soldier, worker, ♀ (alate); Mbaïki; 3 ° 55 ′ N, 18 ° 0 ′ E; 24 Apr. 2018; S. P. Wango leg.; study code: DJ 0587; GenBank no PQ 679223 (mitogenome); BE RMCA INS. Iso. 059388 • soldier, worker; Mbaïki; 3 ° 55 ′ N, 18 ° 0 ′ E; 24 Apr. 2018; S. P. Wango leg.; study code: DJ 0588; BE RMCA INS. Iso. 059394 • soldier, worker; Mbaïki; 3 ° 55 ′ N, 18 ° 0 ′ E; 24 Apr. 2018; S. P. Wango leg.; study code: DJ 0595; BE RMCA INS. Iso. 059389 • soldier, worker; Mbaïki; 3 ° 55 ′ N, 18 ° 0 ′ E; 24 Apr. 2018; S. P. Wango leg.; study code: DJ 0596; GenBank no PQ 679247 (mitogenome); BE RMCA INS. Iso. 059396 • soldier, worker, ♀ (alate); Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 26 Apr. 2018; S. P. Wango leg.; study code: DJ 0589; GenBank nos MN 646738 (COI), MN 685943 (COII), MN 686004 (28 S); BE RMCA INS. Iso. 059393 • soldier, worker, ♀ (alate); Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 26 Apr. 2018; S. P. Wango leg.; study code: DJ 0590; GenBank no PQ 679244 (mitogenome); BE RMCA INS. Iso. 059390 • soldier, worker, ♂ (alate); Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 26 Apr. 2018; S. P. Wango leg.; study code: DJ 0591; GenBank no PQ 679206 (mitogenome); BE RMCA INS. Iso. 059392 • soldier, worker; Bondoé; 5 ° 10.3 ′ N, 17 ° 44.2 ′ E; 26 Apr. 2018; S. P. Wango leg.; study code: DJ 0594; GenBank no PQ 679201 (mitogenome); BE RMCA INS. Iso. 059391 • soldier, worker, ♂ (king); Bangui?; 4 ° 21 ′ N, 18 ° 30 ′ E; 3 May 2007; N. Sion leg.; study code: DJ 0936; GenBank no PV 564647 (mitogenome); BE RMCA INS. Iso. 059398. CHAD • soldier, worker; Moundou; 8 ° 24 ′ N, 16 ° 24 ′ E; 23 May 1948; P. - P. Grassé and C. Noirot leg.; study code: DJ 0431; MNHN EP 9923. CÔTE D’IVOIRE • soldier; 6 km E of Odienné; 9 ° 30 ′ N, 7 ° 30 ′ W; Dec. 1938; P. - P. Grassé leg.; study code: DJ P 154; MNHN EP 9925 • soldier; 6 km E of Odienné; 9 ° 30 ′ N, 7 ° 30 ′ W; Dec. 1938; P. - P. Grassé leg.; study code: DJ P 155; MNHN EP 9926 • soldier; Odienné; 7 ° 15 ′ N, 7 ° 14 ′ W; Dec. 1938; P. - P. Grassé leg.; study code: DJ P 156; MNHN EP 9927 • soldier; Mont Tonkui; 7 ° 24 ′ N, 7 ° 9 ′ W; 18 Mar. 1947; C. Noirot leg.; study code: DJ P 338; MNHN EP 9953 • soldier; Ferkessédougou; 9 ° 35 ′ N, 5 ° 12 ′ W; 2 Oct. 1947; C. Noirot leg.; study code: DJ P 192; MNHN EP 9940 • soldier; Bouaké – Mbayakro road; 7 ° 40 ′ N, 4 ° 54 ′ W; 8 Oct. 1947; C. Noirot leg.; study code: DJ P 332; MNHN EP 9947 • soldier, worker; Youhouli; 5 ° 25 ′ N, 4 ° 29 ′ W; 7 Sep. 1953; M. Lüscher leg.; study code: DJ 0330; initially C. severus; AMNH • soldier; Yapo; 5 ° 44 ′ N, 4 ° 5 ′ W; 26 May 1960; C. Noirot leg.; study code: DJ P 323; MNHN EP 9861 • soldier, ♀ (alate); Kpouébo (Orumbo Boka); 6 ° 21 ′ N, 4 ° 53.7 ′ W; 30 Oct. 1960; C. Noirot leg.; study code: DJ P 346; MNHN • soldier, ♂ (alate); Adiaké; 5 ° 27 ′ N, 3 ° 15.5 ′ W; 15 Apr. 1963; C. Noirot leg.; study code: DJ P 349; MNHN • soldier; Dabou; 5 ° 19 ′ N, 4 ° 20 ′ W; 10 Jun. 1964; P. Bodot leg.; study code: DJ U 014; initially C. severus; NHMUK 13671954 • soldier; Dabou; 5 ° 19 ′ N, 4 ° 20 ′ W; 10 Jun. 1964; P. Bodot leg.; study code: DJ U 015; initially C. severus; NHMUK 13671953 • worker; Taï; 5 ° 52 ′ N, 7 ° 20 ′ W; 1970; C. Kerdelhué leg.; study code: DJ 0749; GenBank no PQ 679224 (mitogenome); MNHN EP 9942 • soldier, worker; Taï; 5 ° 52 ′ N, 7 ° 20 ′ W; 1970; C. Kerdelhué leg.; study code: DJ 0750; GenBank no PQ 679242 (mitogenome); MNHN EP 9943 • soldier, worker, ♂ (alate); Pakobo (Lamto research centre); 6 ° 12 ′ N, 4 ° 59 ′ W; 8 Mar. 2010; K. Dosso leg.; study code: DJ 0017; BE RMCA INS. Iso. 059385 • soldier, worker; Pakobo (Lamto research centre); 6 ° 12 ′ N, 4 ° 59 ′ W; 8 Mar. 2010; K. Dosso leg.; study code: DJ 0066; BE RMCA INS. Iso. 059386 • soldier, worker; Pakobo (Lamto research centre); 6 ° 12 ′ N, 4 ° 59 ′ W; 8 Mar. 2010; K. Dosso leg.; study code: DJ 0069; BE RMCA INS. Iso. 059387 • soldier, worker, ♀ (alate), ♂ (alate); Korhogo; 9 ° 24 ′ N, 5 ° 37 ′ W; 2013; T. Coulibaly leg.; study code: DJ 0044; BE RMCA INS. Iso. 059411 • soldier; Korhogo; 9 ° 24 ′ N, 5 ° 37 ′ W; 2013; T. Coulibaly leg.; study code: DJ 0056; BE RMCA INS. Iso. 059441 • soldier, worker; Youhouli; 5 ° 26.433 ′ N, 4 ° 29.351 ′ W; 18 Feb. 2015; G. Josens leg.; study code: DJ 0408; BE RMCA INS. Iso. 059439 • soldier, worker; Youhouli; 5 ° 26.417 ′ N, 4 ° 29.333 ′ W; 18 Feb. 2015; G. Josens leg.; study code: DJ 0410; GenBank no PQ 679209 (mitogenome); BE RMCA INS. Iso. 059440 • soldier, worker; Pakobo (Lamto research centre); 6 ° 12.56 ′ N, 4 ° 58.584 ′ W; 21 Feb. 2015; G. Josens leg.; study code: DJ 0411; GenBank no PQ 679169 (mitogenome); BE RMCA INS. Iso. 059415 • soldier, worker; Taabo; 6 ° 17.87 ′ N, 5 ° 0.129 ′ W; 21 Feb. 2015; G. Josens leg.; study code: DJ 0409; GenBank nos MN 646736 (COI), MN 685941 (COII), MN 686002 (28 S); BE RMCA INS. Iso. 059455 • soldier, worker; Katiola; 7 ° 58.6 ′ N, 5 ° 4 ′ W; 26 Feb. 2015; G. Josens leg.; study code: DJ 0260; BE RMCA INS. Iso. 059413 • soldier, worker; Odienné; 9 ° 26.252 ′ N, 7 ° 34.935 ′ W; 26 Feb. 2015; G. Josens leg.; study code: DJ 0414; BE RMCA INS. Iso. 059444 • soldier, worker, ♀ (queen); Man; 7 ° 24.7 ′ N, 7 ° 35.1 ′ W; 27 Feb. 2015; G. Josens leg.; study code: DJ 0261; BE RMCA INS. Iso. 059416 • soldier, worker, ♂ (king), ♀ (queen); Zéalé; 5 ° 57.9 ′ N, 8 ° 11.4 ′ W; 28 Feb. 2015; G. Josens leg.; study code: DJ 0262; BE RMCA INS. Iso. 059442 • soldier, worker; Zéalé; 6 ° 57.879 ′ N, 8 ° 11.386 ′ W; 28 Feb. 2015; G. Josens leg.; study code: DJ 0415; GenBank no PQ 679171 (mitogenome); BE RMCA INS. Iso. 059443 • soldier, worker; Route Man – Duékoué; 6 ° 50.051 ′ N, 7 ° 24.53 ′ W; 1 Mar. 2015; G. Josens leg.; study code: DJ 0413; GenBank nos MN 646737 (COI), MN 685942 (COII), MN 686003 (28 S), PQ 679236 (mitogenome); BE RMCA INS. Iso. 059451. GABON • soldier; La Lopé N. P.; 0 ° 15 ′ S, 11 ° 35 ′ E; 15 Mar. 1998; M. Harry leg.; study code: DJ P 251; MNHN EP 9945. GAMBIA • soldier, ♂ (king); Tabo Kotor; 13 ° 24 ′ N, 16 ° 40 ′ W; 10 Sep. 1966; W. A. Sands leg.; study code: DJ U 100; initially C. gaigei; NHMUK 13671897 • soldier, ♂ (king); Bathurst [= Banjul] – Yundum road; 13 ° 28 ′ N, 16 ° 38 ′ W; 12 Sep. 1966; W. A. Sands leg.; study code: DJ U 103; initially C. gaigei; NHMUK 13671898 • soldier; Sanyangh – Gunjur road; 13 ° 15 ′ N, 16 ° 46 ′ W; 16 Sep. 1966; W. A. Sands leg.; study code: DJ U 108; initially C. gaigei; NHMUK 13671885 • soldier, worker, ♀ (queen); Bathurst [= Banjul]; 13 ° 13 ′ N, 16 ° 19 ′ W; 18 Sep. 1966; W. A Sands leg.; study code: DJ 0717; initially C. gaigei; NHMUK 13671896 • soldier; Bakau; 13 ° 28 ′ N, 16 ° 40 ′ W; 17 Nov. 1973; R. M. C. Williams and R. W. Lamb leg.; study code: DJ U 007; initially C. severus; NHMUK 13671950 • soldier; Jambanjeli; 13 ° 17 ′ N, 16 ° 44 ′ W; 25 Nov. 1973; R. W. Lamb leg.; study code: DJ U 018; initially C. severus; NHMUK 13671952 • soldier; Abuko game reserve; 13 ° 24 ′ N, 16 ° 39 ′ W; 5 Dec. 1973; R. M. C. Williams and R. W. Lamb leg.; study code: DJ U 008; initially C. severus; NHMUK 13671951 • soldier, ♂ (king); Abuko game reserve; 13 ° 24 ′ N, 16 ° 39 ′ W; 5 Dec. 1973; R. M. C. Williams and R. W. Lamb leg.; study code: DJ U 017; initially C. severus; NHMUK 13671949 • soldier, worker; Brikama (Nyambai Forest?); 13 ° 18 ′ N, 16 ° 40 ′ W; 1 Nov. 2013; A. Ndiaye leg.; study code: DJ 0220; BE RMCA INS. Iso. 059397. GHANA • soldier, ♀ (queen); Larabanga – Sawla road; 9 ° 13 ′ N, 1 ° 55 ′ W; 22 Feb. 1959; W. A. Sands leg.; study code: DJ U 115; initially C. gaigei; NHMUK 13671882 • soldier; Bolgatanga; 10 ° 34 ′ N, 0 ° 50 ′ W; 9 Mar. 1959; W. A. Sands leg.; study code: DJ U 030; initially C. severus; NHMUK 13671966 • soldier; Tumu; 11 ° 5 ′ N, 2 ° 17 ′ W; 16 Mar. 1959; W. A. Sands leg.; study code: DJ U 027; initially C. severus; NHMUK 13671970 • soldier; Lawra; 10 ° 39 ′ N, 2 ° 52 ′ W; 18 Mar. 1959; W. A. Sands leg.; study code: DJ U 031; initially C. severus; NHMUK 13671967 • soldier; Tamale – Yeji road; 8 ° 55 ′ N, 0 ° 30 ′ W; 27 Mar. 1959; W. A. Sands leg.; study code: DJ U 024; initially C. severus; NHMUK 13671969 • soldier, ♀ (queen); Akumadan on Wenchi – Kumasi road; 7 ° 28 ′ N, 1 ° 58 ′ W; 1 Apr. 1959; W. A. Sands leg.; study code: DJ U 101; initially C. gaigei; NHMUK 13671883 • soldier, ♀ (queen); Akumadan on Wenchi – Kumasi road; 7 ° 28 ′ N, 1 ° 58 ′ W; 1 Apr. 1959; W. A. Sands leg.; study code: DJ U 102; initially C. gaigei; NHMUK 13671885 • soldier; Nakpanduri; 10 ° 34 ′ N, 0 ° 17 ′ W; 6 Jun. 1959; W. A. Sands leg.; study code: DJ U 023; initially C. severus; NHMUK 13671968 • soldier; Ejura; 7 ° 23 ′ N, 1 ° 22 ′ W; 20 Oct. 1966; W. A. Sands leg.; study code: DJ U 020; initially C. severus; NHMUK 13671971 • soldier, ♀ (alate); Kumasi – Sunyani; 6 ° 53 ′ N, 1 ° 52 ′ W; 12 Jul 1975; R. M. C. Williams leg.; study code: DJ U 116; initially C. gaigei; NHMUK 13671895. GUINEA • ♀ (alate), ♂ (alate); “ La Mé forest ”; 9 ° 31.5 ′ N, 13 ° 41.1 ′ W; 31 Jan. 1931; P. - P. Grassé leg.; study code: DJ 0429; initially C. fungifaber in MNHN; MNHN EP 9921 • soldier; Camayenne; 9 ° 31.5 ′ N, 13 ° 41.1 ′ W; 1934; P. - P. Grassé leg.; study code: DJ 0427; MNHN EP 9920 • soldier, worker, ♀ (queen); Camayenne; 9 ° 31.5 ′ N, 13 ° 41.1 ′ W; 1934; P. - P. Grassé leg.; study code: DJ 0430; MNHN EP 9922 • soldier, ♂ (alate); Nzo; 7 ° 41 ′ N, 8 ° 19 ′ W; 25 Mar. 1947; C. Noirot leg.; study code: DJ P 322; MNHN EP 9924 • soldier, ♀ (alate); Nzo; 7 ° 41 ′ N, 8 ° 19 ′ W; 26 Mar. 1947; C. Noirot leg.; study code: DJ P 343; MNHN • soldier, ♀ (queen); Nzo; 7 ° 41 ′ N, 8 ° 19 ′ W; 27 Mar. 1947; C. Noirot leg.; study code: DJ P 342; MNHN • soldier; Nimba mountain; 10 ° 0 ′ N, 13 ° 0 ′ W; Apr. 1996; M. Harry leg.; study code: DJ 0751; MNHN EP 9944 • soldier; Foulaya; 10 ° 0 ′ N, 12 ° 50 ′ W; 20 Jun. 1996; F. L. Guilavogui leg.; study code: DJ 0068; initially C. severus in RMCA; BE RMCA INS. Iso. 059421 • soldier; Forécariah; 9 ° 28 ′ N, 13 ° 6 ′ W; 21 Jun. 1996; O. Diallo leg.; study code: DJ 0067; initially C. severus in RMCA; BE RMCA INS. Iso. 059420 • soldier, worker; Haut-Niger N. P.; 10 ° 14.43 ′ N, 10 ° 27.67 ′ W; 22 Dec. 1996; M. Mei leg.; study code: DJ 0314; initially C. severus; MURS. LIBERIA • soldier, ♀ (alate); unknown location; 7 ° 0 ′ N, 10 ° 0 ′ W; 12 Feb. 1975; G. Becker leg.; study code: DJ U 009; initially C. severus; NHMUK 13671957. NIGERIA • soldier, ♂ (alate); Enugu Township; 6 ° 27 ′ N, 7 ° 32 ′ E; 13 Feb. 1955; W. V. Harris leg.; study code: DJ U 106; initially C. gaigei; NHMUK 13671893 • soldier; Kaduna; 10 ° 15 ′ N, 7 ° 50 ′ E; 15 Feb. 1957; W. A. Sands leg.; study code: DJ U 025; initially C. severus; NHMUK 13671974 • soldier, ♂ (alate); Kwei Hill, Heipang; 9 ° 39 ′ N, 8 ° 50 ′ E; 29 Mar. 1957; W. A. Sands leg.; study code: DJ U 026; initially C. severus; NHMUK 13671977 • soldier, worker, ♂ (alate); Ropp; 9 ° 30 ′ N, 8 ° 55.7 ′ E; 3 Apr. 1957; W. A. Sands leg.; study code: DJ 0545; initially C. severus; NHMUK • soldier, ♂ (alate); Bauchi – Gombe road; 10 ° 17 ′ N, 9 ° 56 ′ E; 7 May 1957; W. A. Sands leg.; study code: DJ U 105; initially C. gaigei; NHMUK 13671890 • soldier; Donkin; 8 ° 37 ′ N, 11 ° 44 ′ E; 24 May 1957; W. A. Sands leg.; study code: DJ U 028; initially C. severus; NHMUK 13671972 • soldier; Wamba; 9 ° 31 ′ N, 8 ° 37 ′ E; 23 Feb. 1958; W. A. Sands leg.; study code: DJ U 022; initially C. severus; NHMUK 13671973 • soldier, ♂ (alate); Gboko – Otupko road; 7 ° 17 ′ N, 8 ° 54 ′ E; 26 Feb. 1958; W. A. Sands leg.; study code: DJ U 032; initially C. severus; NHMUK 13671976 • soldier, ♀ (queen); Kabba – Ondo road; 7 ° 34.75 ′ N, 5 ° 48 ′ E; 2 Mar. 1958; W. A. Sands leg.; study code: DJ U 107; initially C. gaigei; NHMUK 13671889 • soldier, ♂ (alate); Enugu – Onitsha road; 6 ° 25 ′ N, 7 ° 24 ′ E; 3 Mar. 1958; W. A. Sands leg.; study code: DJ U 113; initially C. gaigei; NHMUK 13671892 • soldier, ♂ (alate); Enugu – Otukpo road; 6 ° 45 ′ N, 7 ° 28 ′ E; 5 Mar. 1958; W. A. Sands leg.; study code: DJ U 112; initially C. gaigei; NHMUK 13671891 • soldier, worker; Dekina; 7 ° 40.25 ′ N, 6 ° 56 ′ E; 7 Mar. 1958; W. A. Sands leg.; study code: DJ 0659; initially C. gaigei; NHMUK 13671993 • soldier, ♀ (alate); Lokoja; 7 ° 50 ′ N, 6 ° 33 ′ E; 10 May 1958; W. A. Sands leg.; study code: DJ U 029; initially C. severus; NHMUK 13671975 • soldier; Mayo Ndaga; 6 ° 55 ′ N, 11 ° 27 ′ E; ca 1975; J. T. Medler leg.; study code: DJ U 021; initially C. severus; NHMUK 13671978. SIERRA LEONE • soldier, worker, ♀ (alate); unknown location; 8 ° 0 ′ N, 12 ° 0 ′ W; 14 Jun. 1917; P. A. Buxton leg.; study code: DJ 0323; initially C. silvestrii; AMNH • soldier; Njala; 8 ° 6 ′ N, 12 ° 4 ′ W; 24 Aug. 1930; E. Hargreaves leg.; study code: DJ U 019; initially C. severus; NHMUK 13671979 • soldier, worker; Njala; 8 ° 10 ′ N, 12 ° 24 ′ W; 24 Aug. 1930; E. Hargreaves leg.; study code: DJ 0325; initially C. silvestrii; AMNH • soldier, worker; Njala; 8 ° 10 ′ N, 12 ° 24 ′ W; 24 Aug. 1930; E. Hargreaves leg.; study code: DJ 0540; initially C. silvestrii; NHMUK • soldier, worker; Njala; 8 ° 10 ′ N, 12 ° 24 ′ W; Apr. 1936; E. Hargreaves leg.; study code: DJ 0546; initially C. severus; NHMUK • soldier, worker, ♂ (alate), ♀ (queen); Njala; 8 ° 10 ′ N, 12 ° 24 ′ W; 29 May 1947; F. A. Squire leg.; study code: DJ 0326; initially C. silvestrii; AMNH • soldier; Freetown airport; 8 ° 37 ′ N, 13 ° 11 ′ W; Jan. 1955; W. V. Harris leg.; study code: DJ U 011; initially C. severus; NHMUK 13671960 • soldier; Njala; 8 ° 6 ′ N, 12 ° 4 ′ W; 27 Jan. 1955; W. V. Harris leg.; study code: DJ U 013; initially C. severus; NHMUK 13671964 • soldier; Freetown; 8 ° 25 ′ N, 13 ° 13 ′ W; Feb. 1955; W. V. Harris leg.; study code: DJ U 002; initially C. severus; NHMUK 13671959 • soldier; Freetown; 8 ° 25 ′ N, 13 ° 13 ′ W; Feb. 1955; W. V. Harris leg.; study code: DJ U 003; initially C. severus; NHMUK 13671961 • soldier, worker, ♀ (queen); Freetown; 8 ° 28 ′ N, 13 ° 15 ′ W; 27 May 1956; J. K. Ocloo and R. M. C. Williams leg.; study code: DJ 0544; initially C. severus; NHMUK • soldier; Freetown; 8 ° 28 ′ N, 13 ° 15 ′ W; 8 Jan. 1958; W. Wilkinson leg.; study code: DJ U 005; initially C. severus; NHMUK 13671965 • soldier; Freetown; 8 ° 28 ′ N, 13 ° 15 ′ W; 8 Jan. 1958; W. Wilkinson leg.; study code: DJ U 012; initially C. severus; NHMUK 13671960 • soldier; “ Bo residential ”; 7 ° 57 ′ N, 11 ° 44 ′ W; 10 Jan. 1958; W. Wilkinson leg.; study code: DJ U 010; initially C. severus; NHMUK 13671963 • soldier; Makeni; 8 ° 53 ′ N, 12 ° 2 ′ W; 24 Sep. 1966; W. A. Sands leg.; study code: DJ U 004; initially C. severus; NHMUK 13671958. TOGO • soldier; Kloto; 6 ° 57.17 ′ N, 0 ° 34.21 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ BB 01; BE RMCA INS. Iso. 059380 • soldier; Kloto; 6 ° 57.17 ′ N, 0 ° 34.21 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ BB 02; BE RMCA INS. Iso. 059381 • soldier; Kloto; 6 ° 57.17 ′ N, 0 ° 34.21 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ BB 03; BE RMCA INS. Iso. 059383 • soldier; Kloto; 6 ° 57.17 ′ N, 0 ° 34.21 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ BB 04; BE RMCA INS. Iso. 059384 • soldier; Kloto; 6 ° 57.17 ′ N, 0 ° 34.21 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ BB 05; BE RMCA INS. Iso. 059382 • soldier, worker; Kloto; 6 ° 57.15 ′ N, 0 ° 34.2 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ 0635; BE RMCA INS. Iso. 059454 • soldier, worker; Kloto; 6 ° 57.15 ′ N, 0 ° 34.2 ′ E; 8 Dec. 2018; G. Goergen leg.; study code: DJ 0636; BE RMCA INS. Iso. 059453 • soldier; PN Fazao-Malfakassa, Bonako; 9 ° 10.08 ′ N, 0 ° 57 ′ E; 17 Feb 2025; Y. Roisin leg.; study code: DJ Y 383; BE RMCA INS. Iso. 059952 • soldier; Route Atakpamé – Badou; 7 ° 32.4 ′ N, 1 ° 0 ′ E; 19 Feb 2025; Y. Roisin leg.; study code: DJ Y 381; BE RMCA INS. Iso. 059950 • soldier; Route Atakpamé – Badou; 7 ° 33.24 ′ N, 1 ° 0.54 ′ E; 19 Feb 2025; Y. Roisin leg.; study code: DJ Y 382; BE RMCA INS. Iso. 059951. Historical review The first mention of an Isognathotermes species in the literature probably dates back to Smeathman (1781): of two mushroom-shaped nest-building species, “ The largest species, the Termes atrox, in its perfect state measures one inch and three-tenths from the extremities of the wings on the one side to the extremities to the other ”. As Smeathman collected in Sierra Leone, this dimension (~ 33 mm) is entirely compatible with the wing dimensions of I. severus found in West Africa. However, Smeathman’s material could not be traced even in the early 20 th century (Sjöstedt 1926: 218) and is therefore considered lost and unavailable for a systematic revision. Silvestri (1914: 92 – 93) described this species under the name Cubitermes severus. He provided a description of queen, soldier, and worker together with measurements and figures. In the same paper, he described Cubitermes severus var. modestior (1914: 93 – 94); he mentioned that the soldiers and imagines are like the typical form (= Cubitermes severus) but smaller. Sjöstedt (1926: 250 – 251) housed C. severus in the sub-family Termitinae and provided a first description of alates. Sjöstedt (1926: 238 – 239) considered that C. severus var. modestior deserved to be elevated to the species level. He referred to it under the name C. modestior, housed it in the sub-family Termitinae, and included it in a “ glebae - Gruppe ” of small Cubitermes species. He also included C. severus in his soldier’s key of Cubitermes species and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped frons. Snyder (1949: 163) catalogued C. severus and C. modestior and housed them in the sub-family Termitinae. Krishna et al. (2013: 1936) housed C. severus and C. modestior in the sub-family Cubitermitinae. Josens & Deligne (2019: 39 – 42) placed these species within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed these species in the restored genus Isognathotermes. The small size of I. modestior might suggest a very young colony, but this is excluded with a physogastric queen; this small size could also be the result of environmental conditions. As I. modestior is known from a single nest, without the aid of sequencing (old sample), there is still some doubt as to its status (abnormal sample or distinct species?). On the basis of morphology alone, since the type material of I. modestior (Silvestri, 1914) is now included among the smallest specimens of I. severus, it is now considered a junior synonym of Isognathotermes severus (Silvestri, 1914).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768246BFF56FDE1FD64FB4AFD6D.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, fresh samples very dark, C 6 – C 8 (but C 4 – C 6 in long preserved imagines); fontanelle concolorous with head capsule or paler (up to two levels). Postclypeus C 6 – C 7 slightly paler than head capsule. Antennae C 4 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 7 (paler in long preserved samples), as postclypeus, generally somewhat paler than head capsule; meso- and metanotum C 4 – C 7, sometimes somewhat paler than pronotum. Legs C 3 – C 5; tibia usually slightly (one level) darker than femur. Wings hyaline with brown to grey tinge (Cf 2 – Cf 4), anterior veins darker. Abdomen: tergites C 5 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 ×, sometimes only at 80 ×. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 7 – 12 on the carina of fore coxa and 0 – 8 on the ventral side of fore coxa; fore trochanter with some strong setae; tibia pilose fore, mid, and hind tibia furnished with 10 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 14; Figs 26, 61). Size: the imagines of I. severus are, on average, medium to large sized in the genus Isognathotermes (Fig. 26). Head capsule: compound eyes round to shortly oval; ocelli shortly to strongly oval, removed from eyes by a distance equal to 0.4 – 1.3 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on alate individuals, always shortened by amputation by two to four articles in queens and kings. Labrum: cupola shaped, wider than long. Left mandible with apical tooth on average medium sized in the genus and always more prominent than first marginal; marginal teeth three in number but second one only suggested by an undulation of edge between first and third marginal teeth; only the apical tooth is acute in unworn specimens; premolar tooth with proximal end obscured or partly obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly * Abbreviations: see definitions in Material and methods. elevated. Fore coxa flanged ventrally resulting in a carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 4 and 8 – 13 branches, respectively. Gut (only studied in some cases): enteric valve weakly developed and hardly comparable with workers’ and soldiers’ valves; odd PCs are slightly longer than even PCs. Caecum present as a small amorphous button. Soldier COLOUR. Head capsule generally tending towards deep palette (Cd 3 – Cd 6); there is always a gradient from a darker frons to a paler back, smooth and extended in 72 % of the samples examined but more abrupt in the remaining 28 % giving in some cases the impression that the head capsule is bicolorous (as in Fig. 13). Antennae and labrum concolorous with or one level paler than head capsule. Mandibles dark (C 6 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons. Thorax and legs paler than head capsule (C 2 – C 4). Abdomen grey to red-brown owing to digestive bolus, sometimes with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 5 – 8 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta and generally 0 – 3 spines on carina and generally not any spine (rarely 1 – 2) on ventral side; trochanter generally with some long lined-up setae, sometimes with 6 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 17 strong setae or spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 14; Fig. 62). Size: the soldiers of I. severus are medium sized in the genus Isognathotermes (Figs 21, 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel always with a narrowing near posterior third or fourth, from antennal sockets sides converge clearly towards bases of mandibles; posterior side regularly rounded or sometimes with a short straight or even concave part in the middle. Upper profile almost always slightly to strongly concave; angle between extended mandibles and frons varies from right to obtuse; frons generally with a sketched or small anterior hump (without any hump in 10 % of the samples examined). Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part either roundly convex or forming an acute widening or even a kind of ear on each side. Antennae: 14.5 – 15 articles). Labrum: always deeply bifurcate and wider than long, with sides varying from lyre-shaped (in 85 % of the samples examined) to convex; lobes angular, frequently with fine, whitish or translucent tips; anterior margin concave. Mandibles: sabre-like generally strongly curved; inner edges generally smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible equally or slightly more curved than left. Thorax: pronotum sellate, as wide as 54 – 64 % of head width, with generally entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, lobed, rarely (in 6 % of the samples examined) with a lobe expanded somewhat forward. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 16 % longer than the even cushions, generally without any crest (muneris EVA pattern) or with crests very weakly developed. Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust, scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and 0 – 4 on ventral side; fore trochanter with 5 – 8 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 6 – 15 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 14, Fig. 63). Size: the workers of I. severus are, on average, medium sized in the genus Isognathotermes (with large overlapping on several other species: Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14 or 14.5 (rarely 15) articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal tooth faint (visible as an undulated edge and disappearing in worn mandibles), third marginal tooth with a blunt tip; premolar tooth with its proximal end generally (in 70 % of the samples examined) well visible; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 57 – 73 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are generally as high as or higher than they are wide, with long and strong bristles; however, rather frequently (in 11 cases out of 92), including the syntype from Kakoulima (but not the syntype from Camayenne), the EVA is intermediate between the fungifaber and the finitimus patterns; supporting bristles are generally numerous: 15 – 25 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, generally with two or three (up to five) short lobes, rarely (in 5 % of the samples examined) with a lobe expanded forward. Chorology-ecology The geographic distribution of I. severus is linked with the Sudanian savanna, the Guinean forest-savanna mosaic, and the northern Congolian forest-savanna mosaic ecoregions. Some samples were also found in evergreen forests in RCI and in southwest CAR and were certified by their mitogenomes. This species shows an extended repartition from Gambia to CAR. It seems to be in competition with I. fungifaber (Figs 45, 49, see the general discussion). Molecular data Twenty mitogenomes of I. severus are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). Despite the disparity in the origin of the samples (from RCI to CAR), mitogenomes exhibit less than 1.2 % dissimilarity with each other (Supp. file 4).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249FFF52FE7EFD33FC7CF86E.taxon	description	Figs 26 – 31, 33, 64, distribution map: Fig. 47; Table 15	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249FFF52FE7EFD33FC7CF86E.taxon	diagnosis	Diagnosis The worker has a finitimus EVA and looks a little more like I. finitimus than I. planifrons. The soldier also looks a little more like I. finitimus than I. planifrons. The imago, however, clearly looks more like I. planifrons. The specimens now assigned to I. similifinitimus sp. nov. were first considered to be close to I. planifrons; besides their mixed morphology, they were found outside the geographic range of I. planifrons. Isognathotermes similifinitimus sp. nov. is a cryptic species that can be distinguished for sure only with molecular tools: it is therefore possible that some of the samples previously identified as I. finitimus (that could not be sequenced) belong in fact to I. similifinitimus. In terms of its ecology and chorology, this species is up to now only known from the Congolese forest south of Kisangani, DRC.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249FFF52FE7EFD33FC7CF86E.taxon	etymology	Etymology The name similifinitimus (from the Latin similis, ‘ similar’ and finitimus from the species I. finitimus) refers to the morphology that is like that of I. finitimus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249FFF52FE7EFD33FC7CF86E.taxon	materials_examined	Material examined Two complete samples from one location. Holotype DEMOCRATIC REPUBLIC OF THE CONGO • soldier; Yoko; 0 ° 17.62 ′ N, 25 ° 17.77 ′ E; 8 Jun. 2010; B. Le Ru leg.; study code: DJ 0744; GenBank no PQ 679199 (mitogenome); BE RMCA INS. Iso. 059457. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (queen); same data as for holotype; BE RMCA INS. Iso. 059939. Other material examined DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♂ (king), ♀ (queen); Yoko; 0 ° 17.62 ′ N, 25 ° 17.98 ′ E; 8 Jun. 2010; B. Le Ru leg.; study code: DJ 0745; GenBank no PQ 679249 (mitogenome); BE RMCA INS. Iso. 059456. Historical review This species is described here; only two samples are available which may seem too little for describing a new species but both samples are complete (imago, soldier, worker, and mitogenome). It was discovered by one of us (BLR) in an evergreen forest south of Kisangani, DRC.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249FFF52FE7EFD33FC7CF86E.taxon	description	Description Imago COLOUR. Head capsule: well sclerotised, dark, C 6 – C 7; fontanelle concolorous with head capsule. Postclypeus concolorous with head capsule. Antennae C 5 – C 6 without any difference between proximal and distal articles. Thorax: pronotum C 6 – C 7, concolorous with head capsule; meso- and metanotum C 6 – C 7, as pronotum. Legs C 3 – C 4. Abdomen: tergites C 6 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule with some prominent setae, mainly near the eyes, and a higher density of shorter, finer setae everywhere. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and many shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, visible at 20 ×, with or without two larger setae on metanotum. Legs very pilose, furnished (among numerous fine setae) with some strong setae: 10 – 15 on the carina of fore coxa and 2 on the ventral side of fore coxa; tibia pilose; fore, mid, and hind tibia furnished with 10 – 30 strong setae and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 15; Figs 26, 64). Size: the imagines of I. similifinitimus sp. nov. are medium sized in the genus Isognathotermes; as shown in Fig. 26. Head capsule: compound eyes nearly round; fontanelle a tiny round to elongate marking; ocelli nearly round to oval, removed from eyes by a distance about equal to 0.8 – 0.9 ocellus small diameter. Labrum: cupola shaped. Left mandible with apical tooth always longer and more prominent than first marginal; marginal teeth three in number but second one only suggested by a slight undulation of edge between first and third marginal teeth; premolar tooth with proximal end obscured by molar prominence in dorsal view; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth longer and more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip; second marginal tooth smaller and with a blunt tip; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a sharp carina. Soldier COLOUR. Head capsule C 4 – C 6; there is, in one of the two samples, a strong gradient from a darker frons to a paler back (e. g., from C 6 to C 4) giving the impression that the head capsule is bicolorous (as in Fig. 13). Gulamentum, antennae, and labrum concolorous with head. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (two palette levels) which is generally the same colour as frons. Thorax, nota C 4 and legs paler (C 2 – C 3). Abdomen grey owing to digestive bolus, with a yellow tinge on tergites. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 5 – 6 large setae on each lobe. Thorax: pro- and mesonotum with some setae mainly located on margins. Legs: fore coxa furnished with 1 – 2 spines on carina and not any one on ventral side; trochanter with some long setae, and 4 – 5 lined-up spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 7 – 15 spines. Abdomen: tergites with some large setae, only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 15; Figs 28, 64). Size: the soldiers of I. similifinitimus sp. nov. are medium sized in the genus Isognathotermes; (Fig. 28). Head capsule (Fig. 64): always clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a slight narrowing near posterior third or fourth; from antennal sockets sides converge clearly towards bases of mandibles. Upper profile clearly concave; frons: flat without any hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part forming a kind of ear on each side. Antennae: of 15 articles. Labrum: deeply bifurcate and wider than long, with sides lyre-shaped; lobes angular, with translucent to whitish tips; anterior margin concave. Mandibles: sabre-like, regularly curved; inner edges smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Thorax: pronotum sellate, clearly narrower than head, with a slight notch in anterior margin and entire posterior margin. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions are 14 to 18 % longer than the even cushions, with humps hardly developed, moreover, the pilosity becoming abruptly very dense shows the place where a hump is expected; secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, best seen in ventral view, near centre of abdomen, lobed (three small lobes). Worker COLOUR. Head capsule pale (C 2 – C 3). Antennae: proximal articles pale (C 2), distal articles two levels darker (C 4). Thorax, nota and legs pale (C 1 – C 3). Abdomen grey owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 5 spines on carina and 2 on ventral side; fore trochanter with 6 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 15; Figs 30, 64). Size: the workers of I. similifinitimus sp. nov. are, on average, medium sized in the genus Isognathotermes (but with large overlapping with several other species: Fig. 30). Head capsule weakly sclerotised (except mandibles). Antennae: 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge), third marginal with a blunt tip; premolar tooth with its proximal end hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 72 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a very high density of rather short bristles on a globular bulge; supporting bristles are numerous: 25 – 27 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, shortly lobed (three or four small lobes). Chorology-ecology To date, this species is only known from two samples collected in an evergreen forest near Kisangani, DRC, in the northeastern Congolian lowland forests ecoregion (Fig. 47). Molecular data Two mitogenomes of I. similifinitimus sp. nov. are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’), with a dissimilarity of 1.48 % between them, even though they come from the same locality (Supp. file 4). The phylogenetic reconstructions based on full mitogenomes indicated that this species is paraphyletic with respect to I. phallicaecalis sp. nov. (Fig. 33), to which mitogenomes were up to 1.50 % dissimilar. In this case, the separation of the two species is not justified by molecular dissimilarity but by the different types of caeca.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	description	Figs 26 – 31, 33, 65 – 69, distribution map: Fig. 49; Tables 16 – 17	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	description	Isognathotermes ugandensis burundii Josens & Deligne subsp. nov. Isognathotermes ugandensis kenyae Josens & Deligne subsp. nov. Isognathotermes ugandensis malawii Josens & Deligne subsp. nov. Isognathotermes ugandensis ugandensis Josens & Deligne subsp. nov.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	diagnosis	Diagnosis This species is very close to I. minitabundus and I. severus. The worker (all four subspecies) has a fungifaber EVA but in 6 % of the samples examined, it has an EVA intermediate between the fungifaber and finitimus patterns. Its valve is, on average, somewhat larger than that of I. minitabundus (Fig. 31). It is among the medium- to large-sized workers of the genus Isognathotermes (Fig. 30). As in I. minitabundus, the premolar tooth of the left mandible is more apparent than in most other species. The soldier (all four subspecies) is among the medium- to large-sized soldiers in the genus Isognathotermes (Fig. 28), on average somewhat larger than I. severus. It has the left mandible slightly curved to almost straight and obviously less curved than the right mandible as in I. minitabundus but more asymmetrical than in I. severus (SMlc / SMrc = 0.68 – 1.02, vs 0.51 – 1.03 in I. minitabundus and 0.79 – 1.24 in I. severus). Its head is generally clearly bulbous, but this is shared with I. severus (and also with I. bulbifrons which has a finitimus EVA). The imago is among the medium- to large-sized imagines of the genus Isognathotermes (Figs 26, 65). The premolar tooth on left mandible is more apparent than in most other species (but this is shared with I. severus and I. minitabundus). Contrary to usual, the morphology of imagines is more heterogeneous than that of soldiers and workers (Figs 65 – 69), and this heterogeneity is linked to origin and altitude (Fig. 66), revealing clear subgroups that are hardly detectable in soldiers or workers. Pending more comprehensive molecular data (most of the available samples being too old to be sequenced), we propose to distinguish four subspecies based on overall size (first PCA factor) and eye and ocelli dimensions (second PCA factor).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	etymology	Etymology The name ugandensis (from Uganda and Latin desinence - ensis, from) clearly refers to the country (Uganda) of the type locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	materials_examined	Material examined See hereafter the material examined of Isognathotermes ugandensis ugandensis and other subspecies. Historical review Fuller (1923: 191 – 192) described this species under the name Cubitermes ugandensis. He provided descriptions of alate, imago, and soldier. Sjöstedt (1926: 251 – 252) provided a redescription of alate imago and soldier, together with figures of the soldier’s head, and a short first description of the worker. Sjöstedt (1926: 249 – 250) included Cubitermes ugandensis in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped soldier’s frons. Snyder (1949: 165) catalogued Cubitermes ugandensis in the sub-family Termitinae. Williams (1966: 107 – 108) provided a detailed redescription of imago and soldier along with measurements and figures. He stated that the alate imago described by Fuller (1923) and Sjöstedt (1926) belonged to Ophiotermes ugandaensis. Williams (1966: 108) considered C. ugandensis as a senior synonym of C. antennalis. Emerson, in his unpublished “ card catalog ”, named six samples (from three nests) “ Cubitermes basilewskyi ” in honour of Pierre Basilewsky who collected this material in Rwanda, but it was neither described nor published (nomen nudum). Two such subsamples preserved in RMCA were examined by Williams in 1968 and considered to be Cubitermes ugandensis; confirmed in this study as I. u. burundii. Bouillon & Vincke (1971: 269) described the enteric valve of C. ugandensis as belonging to the first of three types, the “ simple type ” without any spatula. Krishna et al. (2013: 1942) housed Cubitermes ugandensis in the sub-family Cubitermitinae. Josens & Deligne (2019: 39 – 42) placed this species within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Here, we divide the species I. ugandensis into four subspecies that could later become true species. The problem at this stage of our knowledge is that molecular data is still scarce; these taxa can therefore only be recognised by the morphology of the imagines (which are missing from almost half the samples) and by their geographical origin.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	description	Redescription The redescription concerns the species; the distinction between subspecies is based mainly on the imagines’ sizes and origins: see diagnosis. Imago COLOUR. Head capsule: well sclerotised, fresh samples very dark, C 6 – C 8 (C 6 – C 7 in long preserved imagines); fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 7 concolorous or slightly paler (one level) than head capsule. Antennae C 4 – C 7 without any difference between proximal and distal articles. Thorax: pronotum C 5 – C 7, as postclypeus, concolorous or slightly (one level) paler than head capsule; meso- and metanotum C 5 – C 6, sometimes somewhat paler than pronotum. Legs C 3 – C 5; tibia concolorous with femur or somewhat (one level) darker. Wings hyaline with brown to grey tinge (Cf 2 – Cf 3), anterior veins darker. Abdomen: tergites C 6 – C 7. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 5); posterior sternites darker (C 4 – C 5) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 – 80 ×, these fine setae do not extend over the posterior margin. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 7 – 12 on the carina of fore coxa and 1 – 6 on the ventral side of fore coxa and on trochanter; tibia pilose, fore, mid, and hind tibia furnished with 15 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Tables 16 – 17; Fig. 67). Size: the imagines of I. ugandensis are among the medium- to large-sized species of the genus Isognathotermes (as I. minitabundus) (Fig. 26). The imagines’ size is one of the criteria that distinguish the subspecies (see diagnosis). Head capsule: compound eyes oval; ocelli shortly to moderately oval, removed from eyes by a short distance with an eye to ocellus minimal distance index: IEyOcD / IOc d = 0.54 – 1.58 somewhat larger than in I. severus (0.39 – 1.29) and smaller than in I. minitabundus (1.10 – 1.70); fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on alate individuals, shortened by amputation by some articles in king and queen. Labrum: cupola shaped, wider than long. Left mandible with apical tooth medium to small sized (IMlAmD = 0.19 – 0.27 mm: Fig. 27) and always more prominent than first marginal; marginal teeth three in number; second marginal appearing as an undulated edge, reduced but still present in worn mandibles, third marginal tooth with a blunt tip; premolar tooth generally well visible (best seen than in most other species); only the apical tooth is acute in unworn specimens; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 2 – 5 and 9 – 12 branches, respectively. Gut not studied. Soldier COLOUR. Head capsule generally tending towards deep palette (Cd 4 – Cd 6) becoming fader and somewhat darker in long preserved samples; there is generally a smooth gradient from a darker frons to a paler back. Antennae and labrum generally one to two palette levels paler than head capsule. Mandibles dark (C 6 – C 8) generally with an abrupt clearing on their bases (one to two levels) which is generally the same colour as frons; in long preserved samples, this clearing tends to fade. Thorax and legs paler than head capsule (C 3 – C 4) somewhat darker in long preserved samples. Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 4 – 8 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta and 1 – 3 spines (exceptionally 6) on carina and not any spine on ventral side (rarely one); trochanter generally with some strong lined-up setae, or with 3 – 7 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 7 – 18 spines along their shaft. Abdomen: tergites with some large setae, mainly or only on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 16; Figs 28, 67). Size: the soldiers of I. ugandensis are part of the medium to larger soldiers in the genus Isognathotermes (Fig. 28). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a clear narrowing near posterior third or fourth, from antennal sockets sides converge towards bases of mandibles; posterior side regularly convex or sometimes with short straight parts. Upper profile always rather strongly concave; angle between extended mandibles and frons obtuse; frons with sketched to clear anterior hump. Gulamentum in ventral view always constricted in its posterior half, with sides of anterior part generally forming an acute widening or even a kind of ear on each side. Antennae: of 14 – 15 articles. Labrum: always deeply bifurcate and wider than long, with sides varying from lyre-shaped (in 90 % of the samples examined) to convex; lobes angular, generally with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like and rather clearly asymmetrical (but a little less than in I. minitabundus), the left mandible is slightly curved and obviously less curved than the right mandible; inner edges generally smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible always clearly more curved than left. Thorax: pronotum sellate, as wide as 54 – 61 % of head width, with straight or indented anterior margin and posterior straight margin. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, generally three-lobed. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 16 % longer than the even cushions, generally without any crest or with crests very weakly developed. Worker COLOUR. Head capsule pale (C 1 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale (C 1 – C 3). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and 0 – 2 on ventral side; fore trochanter with 6 – 7 spines, all except one lined up; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 7 – 16 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 16, Figs 30, 67). Size: the workers of I. ugandensis are part of the medium to larger workers in the genus Isognathotermes (but with large overlapping on several other species: Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14 or more often 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal appearing as an undulated edge, reduced but still present in worn mandibles (better seen than in other species), third marginal tooth with a blunt tip; premolar tooth generally well visible (also better seen than in other species); molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh and sometimes with a wavy inner edge near the base; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 60 – 68 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are as high as or higher than they are wide, with long and strong bristles (but in 6 % of the samples examined, the EVA is intermediate between the fungifaber and finitimus patterns); 16 – 27 supporting bristles on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum always rather small, visible in ventral view, near centre of abdomen, generally with two short lobes. Chorology-ecology Isognathotermes ugandensis is the easternmost and southernmost species of the genus Isognathotermes (Fig. 49) its geographic distribution is linked with the Victoria basin forest-savanna mosaic, the east Sudanian savanna, the central Zambezian miombo woodlands, and the northern Acacia-Commiphora bushlands and thickets ecoregions. It settles at high altitudes: Williams (1966) mentioned that all his C. ugandensis samples had been collected between 1100 and 2400 m a. s. l. Here, we confirm and extend this range to 625 – 2500 m with a range of four subspecies. Molecular data Of the four considered subspecies, only two (I. ugandensis burundii subsp. nov., and I. u. kenyae subsp. nov.) were successfully sequenced. Six mitogenomes of I. ugandensis are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). Overall, mitogenomes exhibited less than 1.3 % dissimilarity with each other (see Supp. file 4), with I. u. kenyae (DJ 0599) being the most dissimilar. Our phylogenetic reconstructions placed I. ugandensis + I. minitabundus as sister to I. finitimus (Fig. 33; SHalrt / UFB = 99.3 / 93), with whom they are less than 2 % dissimilar. Of note, I. ugandensis and I. minitabundus exhibit a fungifaber EVA which is distinctive from I. finitimus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768249AFF4BFDD8FEEBFCD8FBCD.taxon	discussion	Remarks The subdivision of I. ugandensis into four subspecies and the difficulty of incorporating I. antennalis into one of these subspecies betrays the vagueness that surrounds the Isognathotermes of East Africa. The morphological diversity of the imagines may result from genuine genetic differences, in which case the subspecies could become species, but it could also reflect phenotypic plasticity linked to altitude: only further molecular studies could resolve this question.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682482FF47FEB4FB90FD46F9AD.taxon	description	Figs 65 – 69	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682482FF47FEB4FB90FD46F9AD.taxon	diagnosis	Diagnosis Soldiers and workers do not allow this subspecies to be distinguished from the other three. The imago is close to that of I. u. burundii subsp. nov.; it is, on average, the smallest of the four subspecies, with, relatively speaking, the widest eyes (“ ugug ” in Fig. 65, Table 17). Isognathotermes u. ugandensis is also the subspecies that comes from the lowest average altitude (625 – 1500 m, average = 1015 m); the samples come from Uganda, South Sudan, eastern CAR and Haut-Uele province, DRC). To date, no samples have been sequenced.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682482FF47FEB4FB90FD46F9AD.taxon	etymology	Etymology The name refers to the country (Uganda) of the type locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682482FF47FEB4FB90FD46F9AD.taxon	materials_examined	Material examined Sixty one old samples from 28 locations. Of the 61 old museum samples examined, 20 were initially labelled as Cubitermes ugandensis, 18 as C. antennalis, five as C. speciosus, and 18 as Cubitermes sp. Syntypes of Cubitermes ugandensis Fuller, 1923 = type of Isognathotermes ugandensis ugandensis subsp. nov. UGANDA • soldier, worker; near Entebbe?; 2 ° 0 ′ N, 33 ° 0 ′ E; 1921; H. L. Duke leg.; study code: DJ 0289; initially C. ugandensis; AMNH. Syntypes of Cubitermes antennalis Sjöstedt, 1924 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (alate), ♀ (queen); Moto; 2 ° 54.47 ′ N, 29 ° 24.85 ′ E; 1923; L. Burgeon leg.; study code: DJ 0013; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059422 • worker, ♀ (alate), ♂ (alate); Moto; 2 ° 54.47 ′ N, 29 ° 24.85 ′ E; 1923; L. Burgeon leg.; study code: DJ 0031; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059430. Krishna et al. (2013: 1936) mention that another subsample is deposited in NHRM (not examined). Other material examined CENTRAL AFRICAN REPUBLIC • soldier, worker; Zemio; 5 ° 2 ′ N, 25 ° 7 ′ E; Mar. 1948; N. A Weber leg.; study code: DJ 0808; initially C. antennalis in AMNH; IZC 00321222 • soldier, worker, ♂ (alate); Djema; 6 ° 5 ′ N, 25 ° 12 ′ E; 6 Mar. 1948; N. A. Weber leg.; study code: DJ 0805; initially C. antennalis in AMNH; IZC 00321221. DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker, ♀ (alate); Garamba N. P.; 4 ° 10 ′ N, 29 ° 30 ′ E; 19 Mar. 1912; Lang-Chapin leg.; study code: DJ 0806; initially C. antennalis; AMNH, IZC 00321228 • soldier, worker, ♀ (queen); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; Jul. 1912; Lang-Chapin leg.; study code: DJ 0736; initially C. speciosus; AMNH • soldier, worker; Garamba N. P.; 4 ° 10 ′ N, 29 ° 30 ′ E; Sep. 1912; Lang-Chapin leg.; study code: DJ 0809; initially C. antennalis; AMNH, IZC 00321227 • soldier, worker; Manda; 4 ° 0 ′ N, 24 ° 54 ′ E; 2 Mar. 1925; H. Schouteden leg.; study code: DJ 0019; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059434 • soldier; Manda; 4 ° 0 ′ N, 24 ° 54 ′ E; 27 Nov. 1925; H. Schouteden leg.; study code: DJ 0025; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059435 • soldier, worker, ♀ (alate); Asa; 4 ° 34 ′ N, 25 ° 49 ′ E; 4 Mar. 1948; N. A. Weber leg.; study code: DJ 0804; initially C. antennalis; AMNH, IZC 00321224 • worker; Garamba N. P.; 4 ° 20 ′ N, 29 ° 15 ′ E; 25 Jan. 1950; G. Demoulin leg.; study code: DJ 0039; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059433 • soldier, worker, ♂ (king); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0026; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059437 • soldier, ♂ (king); Garamba N. P.; 3 ° 45 ′ N, 29 ° 16 ′ E; ca 1951; H. De Saeger et al. leg.; study code: DJ U 078; initially C. ugandensis; NHMUK 13672006 • soldier; Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0028; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059438 • worker, ♂ (alate); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0029; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059425 • ♂ (alate); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0033; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059423 • ♀ (alate); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0035; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059424 • ♂ (king), ♀ (queen); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. de Saeger et al. leg.; study code: DJ 0037; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059431 • ♂ (alate); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; ca 1951; H. De Saeger et al. leg.; study code: DJ 0672; initially C. ugandensis; NHMUK 13672005 • soldier, worker; Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; 3 Feb. 1951; H. de Saeger et al. leg.; study code: DJ 0040; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059432 • soldier, worker; Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; 1 Oct. 1951; H. de Saeger et al. leg.; study code: DJ 0023; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059436 • soldier, worker, ♀ (queen); Garamba N. P.; 3 ° 55 ′ N, 29 ° 25 ′ E; 10 Nov. 1951; H. de Saeger et al. leg.; study code: DJ 0024; initially C. antennalis in RMCA; BE RMCA INS. Iso. 059426. “ EAST AFRICA ” (UGANDA?) • soldier; Mabira?; 0 ° 37 ′ N, 33 ° 9 ′ E; no date; Oscar John leg., other label “ collector almost certainly J. Sokolov in 1914 R. M. C. W. ”; study code: DJ U 090; initially C. speciosus; NHMUK 13671990. SOUTH SUDAN • soldier, worker, ♀ (alate), ♂ (alate); Madebe; 4 ° 47 ′ N, 29 ° 3 ′ E; 14 May 1937; J. G. Myers leg.; study code: DJ 0737; initially C. speciosus; AMNH. UGANDA • soldier; Mityara, [= Mityana?]; 0 ° 24 ′ N, 32 ° 2 ′ E; Oct. 1926; H. Hargreaves leg.; study code: DJ N 134; initially C. ugandensis; AMNH • soldier, ♀ (queen); Bugishu [Bugisu]; 1 ° 6 ′ N, 34 ° 11 ′ E; Feb. 1930; H. Hargreaves leg.; study code: DJ N 133; initially C. ugandensis; AMNH • soldier; Kampala – Entebbe road; 0 ° 11 ′ N, 32 ° 32 ′ E; 2 Dec. 1934; H. Kirby leg.; study code: DJ N 131; initially C. ugandensis; AMNH • soldier, worker, ♂ (king); Aduma; 3 ° 0 ′ N, 31 ° 0 ′ E; 1 Sep. 1934; Buxton and H. Kirby leg.; study code: DJ 0807; initially C. antennalis; AMNH, IZC 00321218 • soldier; Bugondo forest; 1 ° 42 ′ N, 31 ° 27 ′ E; 15 Dec. 1934; H. Kirby leg.; study code: DJ N 130; initially C. ugandensis; AMNH • soldier, ♀ (alate); Kambrega Kiyezi; 1 ° 13 ′ S, 29 ° 53 ′ E; 1947; W. V. Harris leg.; study code: DJ U 082; initially C. ugandensis; NHMUK 13671996 • soldier, ♂ (alate); Hoima; 1 ° 26 ′ N, 31 ° 21 ′ E; 4 Dec. 1948; W. V. Harris leg.; study code: DJ 0674; initially C. ugandensis; NHMUK 13671999 • soldier, ♀ (alate); Kiziba Bulemezi; 0 ° 50 ′ N, 32 ° 35 ′ E; 15 Apr. 1949; W. V. Harris leg.; study code: DJ U 074; initially C. ugandensis; NHMUK 13672020 • soldier, worker, ♂ (alate); Kampala – Masake road; 0 ° 3 ′ N, 32 ° 5 ′ E; 4 Feb. 1955; W. Wilkinson leg.; study code: DJ 0570; initially C. ugandensis; NHMUK • soldier, ♀ (alate); Fort Portal – Masindi road; 0 ° 44 ′ N, 30 ° 41 ′ E; 12 Feb. 1955; R. M. C. Williams leg.; study code: DJ U 083; initially C. ugandensis; NHMUK 13671995 • soldier, worker, ♂ (alate), ♂ (king); Fort Portal – Masindi road; 0 ° 58 ′ N, 30 ° 51 ′ E; 12 Feb. 1955; R. M. C. Williams leg.; study code: DJ 0571; initially C. ugandensis; NHMUK • soldier, worker, ♀ (alate); Kyenjojo – Mubende road; 0 ° 33 ′ N, 30 ° 54 ′ E; 14 Feb. 1955; R. M. C. Williams leg.; study code: DJ 0663; initially C. ugandensis; NHMUK 13671997 • soldier, ♀ (alate); Hoima – Fort Portal road; 1 ° 11 ′ N, 31 ° 2 ′ E; 17 Feb. 1955; R. M. C. Williams leg.; study code: DJ U 071; initially C. ugandensis; NHMUK 13671998 • soldier, worker, ♂ (king); Hoima – Fort Portal road; 1 ° 11 ′ N, 31 ° 2 ′ E; 17 Feb. 1955; unknown leg.; study code: DJ 0683; initially C. speciosus; NHMUK 13671984 • soldier, ♂ (alate); Paidha – Pakwash road; 2 ° 24 ′ N, 31 ° 2 ′ E; 24 Feb. 1955; R. M. C. Williams leg.; study code: DJ U 077; initially C. ugandensis; NHMUK 13672021 • soldier, ♀ (alate); Masindi – Kampala road; 1 ° 37 ′ N, 31 ° 45 ′ E; 27 May 1955; R. M. C. Williams leg.; study code: DJ U 075; initially C. ugandensis; NHMUK 13672022 • soldier, worker, ♂ (alate); Mpigi – Masaka road; 0 ° 10 ′ N, 32 ° 16 ′ E; 25 Apr. 1955; R. M. C. Williams leg.; study code: DJ 0682; initially C. speciosus; NHMUK 13671983 • soldier, ♀ (alate); Kampala, Kawanda; 0 ° 25 ′ N, 32 ° 32 ′ E; 25 Jul. 1955; R. M. C. Williams leg.; study code: DJ U 076; initially C. ugandensis; NHMUK 13672019 • soldier, ♂ (king), ♀ (queen); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 253; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059474 • soldier, ♀ (alate), ♂ (king); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 254; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059475 • soldier, ♂ (king), ♀ (queen); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 255; initially Cubitermes sp. (+ Ophiotermes) in RMCA; BE RMCA INS. Iso. 059479 • soldier; Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 256; initially “ Big Cubi ” in RMCA; BE RMCA INS. Iso. 059471 • soldier, ♀ (queen); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 257; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059481 • soldier; Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 258; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059472 • soldier, ♂ (king); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 259; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059482 • soldier, ♂ (king); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 27 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 260; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059484 • soldier; Luweero; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Sep. 1972; P. Vincke leg.; study code: DJ B 252; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059473 • soldier, worker, ♀ (queen); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 15 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0905; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059419 • soldier, worker, ♂ (king); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 15 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0906; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059485 • soldier, worker, ♂ (king); Kampala – Murchinson Falls road; 1 ° 10 ′ N, 32 ° 15 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0907; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059487 • soldier, worker, ♀ (queen); Luweero; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0928; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059476 • soldier, worker; near Luweero; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0900; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059486 • soldier, worker, ♀ (queen); near Luweero; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0903; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059480 • soldier, worker, ♂ (king); near Luweero; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0904; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059483 • soldier, worker; near Murchison Falls N. P.; 1 ° 35 ′ N, 32 ° 0 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0901; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059478 • soldier, worker, ♀ (queen); near Murchison Falls N. P.; 0 ° 51 ′ N, 32 ° 28 ′ E; 2 Nov. 1972; P. Vincke leg.; study code: DJ 0902; initially Cubitermes sp. in RMCA; BE RMCA INS. Iso. 059477.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682482FF47FEB4FB90FD46F9AD.taxon	description	Description Imago See the description of the imago of I. ugandensis (Fuller, 1923) with the particularity that the imago of I. u. ugandensis subsp. nov. is clearly the smallest of the four subspecies, with, relatively speaking, the widest eyes, best seen in a PCA (“ ugug ” in Fig. 65, Table 17), hence the highest IEyD / IHdW ratio (0.25 – 0.34 vs 0.23 – 0.28 for the other three subspecies). Samples previously known as I. antennalis and synonymised with I. ugandensis (Williams 1966) are here assimilated to the subspecies I. u. ugandensis subsp. nov., because of the proximity of their geographical areas and morphological similarity of their imagines. However, not all individuals of I. antennalis should be considered synonyms of I. u. ugandensis as without molecular data, there is no reason to attribute certain samples to I. ugandensis rather than I. severus. And indeed the ‘ unusually’ large size of the “ ugxx ” ellipse in Fig. 65 (I. antennalis samples) suggests a high “ intrasubspecific ” variability, probably linked with genetic heterogeneity. Soldier See the description of the soldier of Isognathotermes ugandensis (Fuller, 1923). Worker See the description of the worker of Isognathotermes ugandensis (Fuller, 1923). Chorology-ecology Isognathotermes ugandensis ugandensis subsp. nov. (including the samples previously identified as I. antennalis) is the subspecies that comes from the lowest altitude (625 – 1500 m, average = 1015 m); it was found in Uganda, south-east CAR, South-Sudan, and north-east DRC. Its geographic distribution is linked with the northern Congolian forest-savanna mosaic, the Victoria basin forest-savanna mosaic, and the east Sudanian savanna ecoregions. Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248EFF41FE44F977FBEAFADB.taxon	description	Figs 33, 65 – 66, 69	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248EFF41FE44F977FBEAFADB.taxon	diagnosis	Diagnosis This subspecies is best characterised by its mitogenome sequence. Soldiers and workers do not allow this subspecies to be distinguished from the other three. The imago of I. u. burundii subsp. nov. is close to that of I. u. ugandensis subsp. nov.; on average, it is somewhat larger with generally smaller eyes (“ ugbu ” in Fig. 65, Table 17); it can be found in Burundi, Rwanda, and Tanzania at a relatively low altitude (1200 – 1700 m, average = 1350 m).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248EFF41FE44F977FBEAFADB.taxon	etymology	Etymology The name burundii (from Burundi) refers to the country of the type locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248EFF41FE44F977FBEAFADB.taxon	materials_examined	Material examined Twenty one samples from 14 locations; of the 13 old museum samples examined, nine were initially labelled as Cubitermes ugandensis and four as C. basilewskyi (nomen nudum). Holotype BURUNDI • soldier; Ruvubu N. P.; 2 ° 56.7 ′ S, 30 ° 26.3 ′ E; Sep. 2014; B. Host leg.; study code: DJ 0396; GenBank no PQ 679187 (mitogenome); BE RMCA INS. Iso. 059458. Paratypes BURUNDI • soldier, worker, ♂ (alate); same data as for holotype; BE RMCA INS. Iso. 059940. Other material examined BURUNDI • soldier, worker; Moso; 4 ° 2 ′ S, 30 ° 8 ′ E; 4 Jul. 1989; J. Ruelle (?) leg.; study code: DJ 0021; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059467 • soldier, worker; Kibago; 4 ° 18.152 ′ S, 29 ° 55.73 ′ E; 25 Nov. 2013; Y. Roisin leg.; study code: DJ 0662; GenBank no PQ 679212 (mitogenome) BE RMCA INS. Iso. 059464 • soldier, worker; Gihofi – Giharo road pk 75; 3 ° 59.124 ′ S, 30 ° 8.826 ′ E; 27 Nov. 2013; Y. Roisin leg.; study code: DJ 0661; GenBank nos MN 646742 (COI), MN 685948 (COII), MN 686009 (28 S), PQ 679235 (mitogenome); BE RMCA INS. Iso. 059465 • soldier, worker; Mahanga; 4 ° 14.5 ′ S, 29 ° 57.6 ′ E; 29 Nov. 2013; Y. Roisin leg.; study code: DJ 0091; GenBank no KY 224661 (mitogenome) BE RMCA INS. Iso. 059462 • soldier, worker; Ruvubu N. P.; 2 ° 56.93 ′ S, 30 ° 25.777 ′ E; 29 Nov. 2013; Y. Roisin leg.; study code: DJ 0660; GenBank nos MN 685947 (COII), MN 686008 (28 S), PQ 679215 (mitogenome); BE RMCA INS. Iso. 059463 • soldier, worker; Rusigabangazi; 3 ° 14.5 ′ S, 30 ° 46.9 ′ E; 30 Nov. 2013; Y. Roisin leg.; study code: DJ 0107; BE RMCA INS. Iso. 059461 • soldier, worker; Ruvubu N. P.; 2 ° 56.7 ′ S, 30 ° 26.3 ′ E; Sep. 2014; B. Host leg.; study code: DJ 0267; GenBank no PQ 679217 (mitogenome) BE RMCA INS. Iso. 059459 • soldier, worker, ♀ (alate); Ruvubu N. P.; 3 ° 3.209 ′ S, 30 ° 30.226 ′ E; Sep. 2014; B. Host leg.; study code: DJ 0368; BE RMCA INS. Iso. 059460. RWANDA • soldier, worker; Gabiro; 1 ° 30.7 ′ S, 30 ° 28 ′ E; 27 Dec. 1934; H. Kirby leg.; study code: DJ 0793; initially C. basilewskyi (unpublished); AMNH, IZC 00321237 • soldier, worker, ♀ (queen); Butare (← Astrida); 2.36 ° 36 ′ S, 29 ° 45 ′ E; 10 Jun. 1952; W. V. Harris leg.; study code: DJ 0675; initially C. ugandensis; NHMUK 13672007 • soldier; Gabiro; 1 ° 30.5 ′ S, 30 ° 28 ′ E; 8 Jul. 1952; W. V. Harris leg.; study code: DJ U 079; initially C. ugandensis; NHMUK 13672008 • soldier, worker; Kinazi; 2 ° 22 ′ S, 30 ° 6 ′ E; 5 – 8 Jan. 1953; P. Basilewsky leg.; study code: DJ 0022; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059468 • soldier, worker; Kinazi; 2 ° 8.5 ′ S, 30 ° 6 ′ E; 7 Jan. 1953; P. Basilewsky leg.; study code: DJ 0792; initially C. basilewskyi (unpublished); AMNH, IZC 00321235 • soldier, worker; Kinazi; 2 ° 8.5 ′ S, 30 ° 6 ′ E; 7 Jan. 1953; P. Basilewsly leg.; study code: DJ 0794; initially C. basilewskyi (unpublished); AMNH, IZC 00321234 • soldier, worker; Rukara; 1 ° 50 ′ S, 30 ° 28 ′ E; 4 Feb. 1953; P. Basilewsky leg.; study code: DJ 0791; initially C. basilewskyi (unpublished); AMNH, IZC 00321236 • soldier, worker; Rukara, near Mohasi lake; 1 ° 50 ′ S, 30 ° 27 ′ E; 5 Nov. 1953; P. Basilewsky leg.; study code: DJ 0020; initially C. ugandensis in RMCA; BE RMCA INS. Iso. 059466. TANZANIA • soldier, ♀ (alate); Bukoba; 1 ° 20 ′ S, 31 ° 48 ′ E; 1937; W. V. Harris leg.; study code: DJ U 081; initially C. ugandensis; NHMUK 13672012 • soldier, worker, ♂ (alate); Geita – Uzinu road; 2 ° 52 ′ S, 32 ° 14 ′ E; Sep. 1937; W. V. Harris leg.; study code: DJ 0664; initially C. ugandensis; NHMUK 13672014 • soldier, ♀ (alate); Katoro; 3 ° 0 ′ S, 31 ° 55 ′ E; Sep. 1937; W. V. Harris leg.; study code: DJ U 084; initially C. ugandensis; NHMUK 13672011 • soldier, worker, ♀ (alate); Mkwese; 5 ° 38 ′ S, 34 ° 49 ′ E; before 1959; P. B. Kemp leg.; study code: DJ 0667; initially C. ugandensis; NHMUK 13672013.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248EFF41FE44F977FBEAFADB.taxon	description	Description Isognathotermes ugandensis burundii subsp. nov. is best characterised by its mitogenome sequence. Imago See the description of the imago of I. ugandensis (Fuller, 1923) with the particularity that, on average, the imago of I. u. burundii subsp. nov. is the second smallest of the four subspecies, best seen in a PCA (“ ugbu ” in Fig. 65, Table 17). Soldier See the description of the soldier of Isognathotermes ugandensis (Fuller, 1923). Worker See the description of the worker of Isognathotermes ugandensis (Fuller, 1923). Chorology – ecology Isognathotermes ugandensis burundii subsp. nov. was found in Burundi, Rwanda, and Tanzania at altitudes between 1200 and 1700 m (average = 1350 m). Its geographic distribution is linked with the central Zambezian miombo woodlands and the Victoria basin forest-savanna mosaic ecoregions. Molecular data Six mitogenomes of I. ugandensis burundii subsp. nov. are published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). Overall, I. ugandensis burundii subsp. nov. mitogenomes exhibited less than 1 % dissimilarity with each other (see Supp. file 4), but 1.2 % with I. u. kenyae subsp. nov. (DJ 0599).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682488FF42FE4EFABEFEADF838.taxon	description	Figs 33, 65 – 66, 69	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682488FF42FE4EFABEFEADF838.taxon	diagnosis	Diagnosis This subspecies is best characterised by its mitogenome sequence. Soldiers and workers do not allow this subspecies to be distinguished from the other three. The imago is close to that of I. u. malawii subsp. nov. (but their geographical distributions are disjunct); it is, on average, the largest of the four subspecies (ugke, Fig. 65, Table 17) with the longest wings (IWgL = 18.9 – 19.5 mm (vs 13.6 – 18.1 mm for two other subspecies, I. u. malawii is not yet available). It is also the subspecies that comes from the highest altitude (1500 – 2500 m, average = 2200 m); to date, all the samples have come from Kenya.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682488FF42FE4EFABEFEADF838.taxon	etymology	Etymology The name kenyae (from Kenya) refers to the country of the type locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682488FF42FE4EFABEFEADF838.taxon	materials_examined	Material examined Seven samples from five locations; all six old museum samples examined were initially labelled as Cubitermes ugandensis. Holotype KENYA • soldier; Mukuyuni; 0 ° 1.612 ′ N, 36 ° 0.303 ′ E; 13 Aug. 2015; J. Šobotnik leg.; study code: DJ 0599; GenBank no PQ 679170 (mitogenome); BE RMCA INS. Iso. 059469. Paratypes KENYA • soldier, worker; same data as for holotype; BE RMCA INS. Iso. 059941. Other material examined “ EAST AFRICA ” (KENYA?) • worker, ♀ (alate); Molo?; 0 ° 15 ′ S, 35 ° 42 ′ E; 5 Jul. 1914; J. Sokolov (?) leg.; study code: DJ 0658; initially C. ugandensis; NHMUK. KENYA • soldier; Thompson falls [= Nyahururu]; 0 ° 2.75 ′ N, 36 ° 22.25 ′ E; Mar. 1934; T. Anderson leg.; study code: DJ N 132; initially C. ugandensis; AMNH • soldier, worker, ♀ (alate); Maji-Mazuri; 1 ° 14 ′ S, 36 ° 56 ′ E; 31 Mar. 1952; W. V. Harris leg.; study code: DJ 0666; initially C. ugandensis; NHMUK 13672002 • soldier, worker, ♂ (alate); Londiani – Makutano (road?); 0 ° 2 ′ S, 36 ° 20 ′ E; 18 Feb. 1954; R. M. C. Williams leg.; study code: DJ 0669; initially C. ugandensis; NHMUK 13672003 • soldier, worker, ♀ (alate); Londiani – Kisumu road; 0 ° 15 ′ S, 35 ° 30 ′ E; 19 Feb. 1954; R. M. C. Williams leg.; study code: DJ 0668; initially C. ugandensis; NHMUK 13672001 • soldier, ♀ (alate); Timboroa – Eldoret road; 0 ° 9 ′ N, 35 ° 28 ′ E; 12 May 1954; W. V. Harris leg.; study code: DJ U 085; initially C. ugandensis; NHMUK 13672000.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B97682488FF42FE4EFABEFEADF838.taxon	description	Description Isognathotermes ugandensis kenyae subsp. nov. is best characterised by its mitogenome sequence. Imago See the description of the imago of I. ugandensis (Fuller, 1923) with the particularity that the imago of I. u. kenyae subsp. nov. is clearly the largest of the four subspecies, best seen in a PCA (“ ugke ” in Fig. 65, Table 17). On average, the imago of this subspecies has the longest wings (IWgL = 18.9 – 19.5 mm (vs 13.6 – 18.1 mm for two other subspecies, I. u. malawii subsp. nov. is not yet available). Soldier See the description of the soldier of Isognathotermes ugandensis (Fuller, 1923). Worker See the description of the worker of Isognathotermes ugandensis (Fuller, 1923). Chorology-ecology Isognathotermes ugandensis kenyae subsp. nov. is the subspecies found at the highest altitude (1500 to 2500 m, average = 2200 m) in the East African montane forest ecoregion of Kenya. Molecular data One sample of I. ugandensis kenyae subsp. nov. has recently been collected by one of us (JŠ) in Kenya, enabling its sequencing; it is published alongside this work (GenBank accessions: see Supp. file 3 and ‘ Material examined’). Its mitogenome exhibits 1.2 % dissimilarity with I. u. burundii subsp. nov. samples (Supp. file 4).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248AFF7CFE48FEEBFD46FC51.taxon	description	Figs 65 – 66, 69	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248AFF7CFE48FEEBFD46FC51.taxon	diagnosis	Diagnosis Soldiers and workers do not allow this subspecies to be distinguished from the other three. The imago is close to that of I. u. kenyae subsp. nov. (but their geographical distributions are disjunct); it is, on average, the second largest of the four subspecies, best seen in a PCA (“ ugma ”, Fig. 65); it is the subspecies with the greatest distance between the apical and the first marginal teeth (IMlAmD = 0.23 – 0.27 mm vs 0.20 – 0.25 mm for all three other subspecies, Table 17). It comes from a relatively high altitude (1460 – 1770 m, average = 1560 m), from Malawi and northern Zambia.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248AFF7CFE48FEEBFD46FC51.taxon	etymology	Etymology The name malawii (from Malawi) refers to the country of the type locality.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248AFF7CFE48FEEBFD46FC51.taxon	materials_examined	Material examined Eleven samples from eight locations; all old museum samples initially identified either as Cubitermes minitabundus (six samples) or C. ugandensis (five samples). Holotype MALAWI • soldier; Dowa-Lilongwe road; 13 ° 39 ′ S, 33 ° 51 ′ E; 13 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0699; initially C. minitabundus; NHMUK 13671919. Paratypes MALAWI • soldier, worker, ♀ (queen); same data as for holotype. Other material examined MALAWI • soldier, worker, ♀ (queen); Dedza; 14 ° 22 ′ S, 34 ° 19.5 ′ E; 4 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0688; initially C. minitabundus; NHMUK 13671917 • soldier, worker; Dedza; 14 ° 23.5 ′ S, 34 ° 18.5 ′ E; 2 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0698; initially C. minitabundus; NHMUK 13671918 • soldier, ♀ (queen); Dowa – Lilongwe road; 13 ° 39 ′ S, 33 ° 51 ′ E; 11 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0697; initially C. minitabundus; NHMUK 13671916 • soldier; Mzimba – Kasungu road; 12 ° 16 ′ S, 33 ° 38 ′ E; 18 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ U 073; initially C. ugandensis; NHMUK 13672015 • soldier, worker; Nyika Plateau; 10 ° 54 ′ S, 33 ° 26 ′ E; 29 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0670; initially C. ugandensis; NHMUK 13672018 • soldier, worker; Nyika Plateau; 10 ° 45 ′ S, 33 ° 57 ′ E; 30 Sep. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ 0671; initially C. ugandensis; NHMUK 13672016 • soldier; Vipya Plateau, Ekwendeni; 11 ° 23 ′ S, 33 ° 54 ′ E; 20 Oct. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ U 072; initially C. ugandensis; NHMUK 13672017. ZAMBIA • soldier; Tunduma; 9 ° 26 ′ S, 32 ° 51 ′ E; 5 Aug. 1953; W. A. Sands and W. Wilkinson leg.; study code: DJ U 080; initially C. ugandensis; NHMUK 13672009 • soldier, worker; Kasama – Mbala road; 10 ° 2 ′ S, 31 ° 15 ′ E; 23 Dec. 1969; M. G. Bingham leg.; study code: DJ 0690; initially C. minitabundus; NHMUK 13671923 • soldier, worker; Mbala – Mpulungu road; 8 ° 48 ′ S, 31 ° 8 ′ E; 24 May 1970; J. R. Clover and Kistner leg.; study code: DJ 0689; initially C. minitabundus; NHMUK 13671924.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B9768248AFF7CFE48FEEBFD46FC51.taxon	description	Description Imago See the description of the imago of Isognathotermes ugandensis (Fuller, 1923) with the particularity that the imago of I. u. malawii subsp. nov. is the second largest of the four subspecies, best seen in a PCA (“ ugma ” in Fig. 65, Table 17). This is the subspecies with the greatest distance between the apical and the first marginal teeth (IMlAmD = 0.23 – 0.27 mm vs 0.20 – 0.25 mm for all three other subspecies, Table 17). Soldier See the description of the soldier of Isognathotermes ugandensis (Fuller, 1923). Worker See the description of the worker of Isognathotermes ugandensis (Fuller, 1923). Chorology-ecology Isognathotermes ugandensis malawii subsp. nov. was found between altitudes of 1460 and 1770 m (average = 1560 m), in the central Zambezian miombo woodlands in Malawi and northern Zambia. Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B5FF7AFDE0FC0BFB2BFC25.taxon	description	Figs 26 – 31, 70, distribution map: Fig. 49; Table 18	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B5FF7AFDE0FC0BFB2BFC25.taxon	diagnosis	Diagnosis The workers and soldiers of I. zenkeri are larger than those of the other species with a fungifaber EVA, but the imagines are almost indistinguishable from those of I. severus and I. ugandensis. The worker has a fungifaber EVA; it is among the large-sized workers of the genus Isognathotermes (Fig. 30) with the largest apico-marginal distances in the genus (WMlAmD = 0.23 – 0.25 mm). The soldier is among the large-sized soldiers of the genus Isognathotermes (Fig. 28); the mandibles are shorter than the head but longer than in other species of the genus (SMlL = 2.75 – 2.99 mm; SMlL / SHdL = 0.86 – 0.92), some soldiers of I. bulbifrons can have equally long mandibles but they have another EVA. Gulamentum with almost straight, parallel lateral edges, only slightly curved. The imago is among the middle- to large-sized imagines of the genus Isognathotermes (Fig. 26) with the largest left and right apico-marginal distances; IMlAmD = 0.26 – 0.28 mm (Fig. 27); ocellus large, protruding and removed from eye by a distance shorter than 0.6 ocellus small diameter.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B5FF7AFDE0FC0BFB2BFC25.taxon	etymology	Etymology This species was named in honour of G. A. Zenker who discovered this species and collected the type material.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B5FF7AFDE0FC0BFB2BFC25.taxon	materials_examined	Material examined Five samples from one (?) location, all labelled as Termes, or Eutermes, or Cubitermes zenkeri. Syntypess of Termes zenkeri Desneux, 1904 CAMEROON • soldier, worker, ♀ (alate); Bipindi? (3 ° 5 ′ N, 10 ° 24.5 ′ E); before 1904; G. Zenker leg.; study code: DJ 0283; initially Termes zenkeri; NHMM • soldier, worker, ♀ (alate); Cameroon, from Hill collection; [no date, no location]; G. Zenker (?) leg.; study code: DJ 0730; initially Termes zenkeri; AMNH • soldier, worker, ♀ (alate); Kamerun, from Holmgren collection; [no date, no location]; G. Zenker (?) leg.; study code: DJ 0731; initially Termes zenkeri; AMNH • soldier, worker, ♂ (alate); Bipindi (3 ° 5 ′ N, 10 ° 24.5 ′ E); 1903; G. Zenker leg.; study code: DJ 0962; initially Eutermes zenkeri; RBINS. Krishna et al. (2013: 1945) mention that other type (sub) samples are deposited in NHRM, and in the Zoologisches Museum der Natur, Hamburg, Germany, not examined. Other material examined CAMEROON • soldier, worker, ♀ (alate), ♀ (queen); Bipindi? (3 ° 5 ′ N, 10 ° 24.5 ′ E); [no date]; G. Zenker leg.; study code: DJ 0321; initially C. zenkeri; AMNH. Five samples from one (?) location, all labelled as Termes, or Eutermes, or Cubitermes zenkeri. Historical review Desneux (1904 a: 148 – 149) described this species under the name Termes Zenkeri (species name incorrectly published with a capital letter and only corrected in 1949). He provided a short description of alate imago, soldier, and worker along with a few measurements, mentioning some differences with C. fungifaber. The species was later referred to under the name of Termes (Eutermes) Zenkeri by Desneux 1904 b, Cubitermes (Termes) Zenkeri by Wasmann (1911: 158), Mirotermes (Cubitermes) Zenkeri by Holmgren (1912), and Cubitermes Zenkeri by Sjöstedt (1926: 236). Sjöstedt (1926: 218 – 226) included the species in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ Zenkeri - Gruppe ” of large Cubitermes species. Snyder (1949: 166) catalogued Cubitermes zenkeri in the subfamily Termitinae while Krishna et al. (2013: 1945) housed it in the subfamily Cubitermitinae. Espousing the view of Ruelle (1975: 8) they considered Cubitermes kemneri as a junior synonym of C. zenkeri. Bouillon & Vincke (1971: 269) described the enteric valve of C. zenkeri as belonging to the first of three types, the “ simple type ” without any spatula. Josens & Deligne (2019: 39 – 42) placed this species within the fungifaber valve pattern group. Therefore, C. zenkeri cannot be the senior synonym of C. kemneri as the types of both species belong to different valve pattern groups, i. e., fungifaber - and finitimus - group respectively. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. This species is confusing: all the known samples seem to come from the same site (Bipindi, Cameroon) and the same collector (G. Zenker), and it has not been collected since, despite the numerous collecting campaigns that have been organised in Cameroon.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B5FF7AFDE0FC0BFB2BFC25.taxon	description	Redescription Imago COLOUR. Head capsule: well sclerotised, and very dark, C 6 – C 7; fontanelle concolorous or almost so with head capsule. Postclypeus C 5 – C 6, slightly paler than head capsule. Antennae C 5, slightly paler than head capsule without any difference between proximal and distal articles. Thorax: pronotum C 5 – C 6, as postclypeus, slightly paler than head capsule; meso- and metanotum C 5 – C 6, concolorous with pronotum. Legs C 4 – C 5; tibia concolorous with femur. Wings hyaline with brown to grey tinge (Cf 3 – Cf 4), anterior veins darker. Abdomen: tergites C 6. Sternites appreciably paler in middle (C 3 – C 4) with both sides darker (C 4 – C 6); posterior sternites darker (C 4 – C 6) than anterior. SETATION. Head capsule, with some prominent setae set amongst a high density of short, fine setae forming a dense mat. Labrum and postclypeus with some prominent setae mixed with shorter ones. Antennae with some prominent setae, some more numerous smaller setae and, mainly distally on most articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Thorax: pronotum with prominent setae mainly on margins and shorter ones in middle; meso- and metanotum with some fine, pale setae, arranged in a medio-longitudinal strip, generally visible at 40 – 80 ×; some large setae sometimes present on metanotum. Legs very pilose, furnished (among numerous fine setae) with some stronger setae: 4 – 10 on the carina of fore coxa and 1 – 4 on the ventral side of fore coxa; tibia pilose fore, mid, and hind tibia furnished with 15 – 30 spines and bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively. Abdomen: tergites with many large and small setae. Sternites with long setae, erect or directed slightly forward, and many long and smaller setae directed backwards. STRUCTURE (measurements in Table 18, Figs 26, 70). Size: the imagines of I. zenkeri are among the medium- to large-sized species of the genus Isognathotermes (Fig. 26) with the largest left and right apico-marginal distances (Fig. 27). Head capsule: compound eyes large and shortly oval; ocelli large, protruding and shortly to moderately oval, removed from eyes by a shorter distance than 0.6 ocellus small diameter; fontanelle generally a tiny round or elongate marking. Antennae: 16 articles on alate individuals, shortened by amputation by three articles in the queen. Labrum: cupola shaped, wider than long. Left mandible with apical tooth large sized in the genus and always more prominent than first marginal; marginal teeth three in number; second marginal appearing as an undulated edge, third marginal tooth with a blunt tip; only the apical tooth is acute in unworn specimens; premolar tooth partly hidden; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible with apical tooth large sized in the genus and always more prominent than first marginal; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum appreciably wider than long and narrower than head width (including the eyes), straight to very weakly sellate with anterior lobe short and very slightly elevated. Fore coxa flanged ventrally resulting in a carina. Wings: R 1 fused entirely with costal margin, sclerotised; Rs simple, sclerotised; M and Cu not or weakly sclerotised with 3 – 4 and 10 – 13 branches, respectively. Gut not studied. Soldier COLOUR. Head capsule C 4 – C 6, darkened by age, with a smooth gradient from a darker frons to a paler back. Antennae and labrum approximately concolorous with head capsule. Mandibles dark (C 7 – C 8) generally with an abrupt clearing on their bases (one palette level) which is generally the same colour as frons. Thorax and legs paler than head capsule (C 3 – C 5), darkened by age. Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule with few scattered setae; on frons a dense bunch of setae surrounds and overhangs fontanelle. Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of very fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with 7 – 10 large setae on each lobe. Thorax: pro- and mesonotum with a small number of setae mainly located on margins. Legs: fore coxa bears at least one fine seta and 2 – 3 spines on carina and 0 – 1 spine on ventral side; trochanter with some strong lined-up setae, or with 5 – 6 spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively (the latter sometimes weakly developed) and a row of 9 – 14 spines along their shaft. Abdomen: tergites with some large setae, mainly on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 18; Figs 28, 70). Size: the soldiers of I. zenkeri are among the large-sized soldiers in the genus Isognathotermes (Fig. 29). Head capsule: always clearly sclerotised; appreciably longer than wide. Dorsal view: lateral sides mostly subparallel with a slight narrowing near posterior third or fourth, from antennal sockets sides converge towards bases of mandibles; posterior side regularly convex or with a short straight part. Upper profile slightly concave; angle between extended mandibles and frons obtuse; frons with a slight anterior hump (Fig. 70). Gulamentum in ventral view with almost parallel sides, very weakly constricted in its posterior half; with sides of anterior part generally forming an acute widening or even a kind of ear on each side. Antennae: of 15 articles. Labrum: always deeply bifurcate and wider than long, with lyre-shaped sides; lobes angular, generally with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like; inner edges smooth with one distinct but very small marginal tooth, near molar tooth on each mandible; mandibles shorter than head but the longest in relative value in the genus (85 – 90 % of head length); entire surface of both mandibles smooth and glossy. Right mandible equally or slightly more curved than left. Thorax: pronotum sellate, as wide as about 60 % of head width, with straight or indented anterior margin and posterior straight margin. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Caecum always rather small, best seen in ventral view, near centre of abdomen, three-lobed. Arrangement of enteric valve cushions showing trilateral symmetry, the odd cushions being about 20 % longer than the even cushions, generally without any crest. Worker COLOUR. Head capsule pale but darkened by age. Antennae: proximal articles pale (C 2 – C 3), distal articles always one to two levels darker (C 4 – C 5). Thorax, nota, and legs pale but darkened by age. Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect, scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa always carinated, bearing one fine seta and furnished with 3 – 4 spines on carina and one on ventral side; fore trochanter with 6 – 7 spines, all except one lined up; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of 8 – 15 spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 18, Figs 30, 70). Size: the workers of I. zenkeri are part of the large-sized workers in the genus Isognathotermes (but with large overlapping on several other species: Fig. 30). Head capsule: weakly sclerotised (except mandibles). Antennae of 14 or 14.5 articles. Labrum: cupola shaped. Left mandible: apical tooth well developed, the largest in the genus, with a sharp tip when fresh; marginal teeth three in number, first marginal tooth well developed but with a blunt tip even when fresh, second marginal appearing as an undulated edge, third marginal tooth with a blunt tip; premolar tooth partly hidden underneath molar tooth; this latter bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed, the largest in the genus, with a sharp tip when fresh and sometimes with a wavy inner edge near the base; marginal teeth two in number; first marginal tooth well developed with a sharp tip when fresh; second marginal tooth smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate, as wide as 70 – 75 % of head width. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in second half of abdomen. Arrangement of enteric valve cushions of the fungifaber pattern with triradial symmetry: the odd PCs, in their downstream part, are rather narrow and bear crests that are as high as or higher than they are wide, with long and strong bristles; 19 – 27 supporting bristles on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum with three lobes, either small or with one lobe somewhat extended forwards, visible in ventral view, near centre of abdomen, Chorology-ecology Only one collecting site is suspected in the Atlantic Equatorial coastal forest ecoregion of Cameroon (Fig. 49). Its precise origin is uncertain: in the original description, Desneux (1904 a) only mentions that it came from “ Kamerun ” and that it was discovered by G. Zenker in a large nest (1 m high) at the base of a tall tree. Georg August Zenker was working at Jaunde [= Yaoundé] from 1889 to 1895 and was later domiciled at Bipindi from 1896 to 1922 (Wikipedia 2024 a, German version), it is therefore believed that Termes zenkeri was found at Bipindi. However, this large and very characteristic species has never been found again despite numerous collections that were made in Cameroon, especially around Mbalmayo and in the region from Yaoundé to Kribi (including Bipindi). As Zenker collected in a vast region of Cameroon, it is quite possible that I. zenkeri, a species with a supposedly limited area, did not originate from Bipindi. Molecular data No genetic sequence is currently available. Incertae sedis – species inquirenda This chapter contains a species already described (I. congoensis (Emerson, 1928 )), but of uncertain matches; one potential species already named (I. bredoi) but not described (nomen nudum) and two potential species with unclear identity (a possible cryptic species of I. planifrons and an unknown species from Malawi). All these samples are classified “ Incertae sedis ” and “ Species inquirenda ”.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B3FF7BFDE9FBFFFD46FC56.taxon	diagnosis	Diagnosis The worker has a fungifaber EVA. The PCA on soldiers’ morphology places this sample at the intersection of the severus and ugandensis ellipses, and near the acristatus ellipse. However, the PCAs on workers’ and soldiers’ valves project this species onto the fungifaber ellipse. The enteric valve has poorly developed crests, which brings it closer to I. acristatus sp. nov. Therefore, this sample seems to be an intermediate between I. ugandensis, I. fungifaber and I. acristatus. This sample therefore deserves to be elevated to the rank of species, provided that other samples with the same characteristics can be found. Hence their provisional classification as Incertae sedis. The imago is unknown.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B3FF7BFDE9FBFFFD46FC56.taxon	etymology	Etymology The name Cubitermes bredoi was given (not published) by A. E. Emerson in honour of Hans Bredo who collected the sample.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B3FF7BFDE9FBFFFD46FC56.taxon	materials_examined	Material examined Two subsamples from the same colony. ZAMBIA (?) • soldier, worker; Swali; 8 ° 45 ′ S, 29 ° 20 ′ E; 25 May 1941; H. Bredo leg.; study code: DJ 0778; initially C. bredoi (unpublished); AMNH, IZC 00321243 • soldier, worker; Swali; 8 ° 45 ′ S, 20 ° 20 ′ E; 25 May 1941; H. Bredo leg.; study code: DJ 0779; initially C. bredoi (unpublished); AMNH, IZC 00321242. Historical review This putative species is known from a single nest sampled in 1941. The sample has been named “ Cubitermes bredoi ” by A. Emerson in his unpublished “ card catalog ” but was not described. It was nevertheless quoted by Mathur & Thapa (1962), and was therefore mentioned as nomen nudum by Krishna et al. (1913: 1915) with an origin (Zimbabwe) which is likely incorrect. This sample is not described here, its status remaining uncertain. We don’t wish to describe a new species of which only one nest is known, and which could be an anomaly. Emerson got away with creating a new name, but he too did not describe a new species. Chorology-ecology The location (Swali or Suali) could not be found; it is assumed to be in Zambia, near the border with Katanga, DRC, where H. Bredo used to work and where there is a river bearing that name. Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B2FF76FDC8FC0BFD46F95C.taxon	description	Figs 22, 28, 30 – 31, 36 – 37, distribution map: Fig. 38; Table 19	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B2FF76FDC8FC0BFD46F95C.taxon	diagnosis	Diagnosis The worker has a finitimus EVA and is among the smallest workers of the genus Isognathotermes (Fig. 30). The soldier is among the smallest in the genus Isognathotermes (Fig. 28); the soldiers of I. fungifaber are sometimes as small but have a fungifaber EVA; the soldier of I. phalloides sp. nov. (also with a finitimus EVA) is sometime as small but has another, characteristic caecum. The imago is unknown. This species is also defined by its chorology: the two known samples come from forested regions up to about 200 km from the Atlantic coast (Fig. 38).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B2FF76FDC8FC0BFD46F95C.taxon	etymology	Etymology The name congoensis (from Congo and the Latin desinence - ensis, ‘ from’) clearly refers to the country of type locality (“ Belgian Congo ”, now DRC).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B2FF76FDC8FC0BFD46F95C.taxon	materials_examined	Material examined Three samples from two locations (probably from two colonies), all identified as C. congoensis. Paratypes of Cubitermes congoensis Emerson, 1928 DEMOCRATIC REPUBLIC OF THE CONGO • soldier, worker; Banana; 5 ° 58 ′ S, 12 ° 26 ′ E; Aug. 1915; Lang-Chapin leg.; study code: DJ 0299; initially C. congoensis; AMNH • soldier, worker; Banana; 6 ° 1 ′ S, 12 ° 25 ′ E; Aug. 1915; Lang-Chapin leg.; study code: DJ 0484; initially C. congoensis; PPRI, ISTM 5690. Krishna et al. (2013: 1918) mention that the holotype is deposited in AMNH, not examined. Other material examined CAMEROON • soldier, worker; Mbalmayo; 3 ° 31 ′ N, 11 ° 30 ′ E; Mar. 1996; P. Eggleton et al. leg.; study code: DJ 0648; initially C. congoensis; NHMUK 13671845. Historical review Emerson (1928: 519 – 520) described this taxon under the name Mirotermes (Cubitermes) congoensis. He provided a description (text, measurements, and figure) of the soldier, mentioning differences with other species. Snyder (1949: 158) catalogued this taxon under the name Cubitermes congoensis in the sub-family Termitinae and reported its known geographical distribution. Krishna et al. (2013: 1918) housed C. congoensis in the sub-family Cubitermitinae. Josens & Deligne (2019: 42 – 44) placed this taxon in the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this taxon in the restored genus Isognathotermes. The sample, however, could not be sequenced. The validity of Isognathotermes congoensis, is puzzling: it has been found in two remote places: the type colony (DJ 0299 & DJ 0484) near the mouth of the Congo River, DRC, and another sample near Mbalmayo, Cameroon (DJ 0648). As explained in the chapter bulbifrons, four samples from incipient or very young colonies collected by N. M. Collins near Douala, Cameroon, were identified on the basis of their young queens as being I. bulbifrons; if the soldiers had been used instead, the resulting identification would have been I. congoensis. In the PCA on soldier morphology, those soldiers join the soldiers from I. congoensis colonies (ellipse labelled “ c + b ” in Fig. 22). Therefore, the two samples identified as I. congoensis might also be incipient colonies of I. bulbifrons. This hypothesis is supported by the absence of imagines. This name might become a synonym if it turns out that it was assigned to an incipient colony of I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B2FF76FDC8FC0BFD46F95C.taxon	description	Redescription Imago The imago is unknown. Soldier COLOUR. Head capsule C 4 – C 5 becoming fader and darker (Cf 5 – Cf 6) in long preserved samples. Gulamentum concolorous with head. Antennae and labrum paler (C 3 – C 4) than head capsule, darker in long preserved samples. Mandibles dark (C 7) with an abrupt clearing on their bases (C 5 – C 6) which is same colour as frons. Thorax, nota and legs paler than head capsule (C 2 – C 4), darker in long preserved samples. Abdomen grey owing to digestive bolus. SETATION. Head capsule with few scattered fine setae; on frons a scattered bunch of setae surrounds and overhangs fontanelle (but one sample with very few setae around fontanelle). Antennae with some prominent setae, more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Labrum always with some large setae on lobes. Thorax: pro- and mesonotum with some setae mainly located on margins. Legs: fore coxa furnished with only one large seta on carina; fore trochanter generally with five or six aligned setae; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively; all tibiae furnished with a row of 4 – 6 spines. Abdomen: tergites with some large setae, mainly on their posterior margins. Sternites with long setae, erect or slightly directed forward, often coloured, and smaller setae directed backwards. STRUCTURE (measurements in Table 19; Fig. 28). Size: the soldiers of I. congoensis are, on average, clearly among the smallest of the genus Isognathotermes; this is better seen in the soldiers’ head lengths (Fig. 29). Head capsule: clearly sclerotised and appreciably longer than wide. Dorsal view: lateral sides subparallel, slightly convex to narrowed near posterior third or fourth, converging slightly towards anterior until antennal sockets; from antennal sockets sides converge towards bases of mandibles; posterior side rounded. In profile: clearly concave; frons with a faint frontal hump overhanging the fontanelle or no hump at all. Gulamentum in ventral view constricted in its posterior half, with sides of anterior part convex; rounded or with a slightly concave, posterior part from side view. Antennae: of 14.5 or 15 articles. Labrum: deeply bifurcate, with lyre-shaped sides; lobes angular or subtruncated, with fine, whitish, or translucent tips; anterior margin concave. Mandibles: sabre-like; inner edges smooth with one distinct but small marginal tooth, near molar tooth on each mandible; mandibles clearly shorter than head; entire surface of both mandibles smooth and glossy. Right mandible slightly more curved than left. Thorax: pronotum sellate, with entire anterior and posterior margins. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions showing trilateral symmetry: the odd cushions being somewhat longer and wider than the even cushions; the odd cushions with crests weakly developed; secondary cushions wide at the upstream end narrowing noticeably downstream with a homogeneous spine scattering. Caecum best seen in ventral view, near centre of abdomen, with three faint lobes. Worker The description of the worker is given here for the first time. COLOUR. Head capsule pale (C 2 – C 3) turning grey in long preserved samples. Antennae: proximal articles pale (C 2), distal articles darker (C 3). Thorax, nota and legs pale (C 1 – C 2). Abdomen grey to red-brown owing to digestive bolus. SETATION. Head capsule and postclypeus with few, erect scattered setae. Labrum with few, robust scattered setae. Antennae with some prominent setae, some more numerous smaller setae and at distal extremity of distal articles, a bunch of fine, bent setae (visible only at high magnification, 50 × or more). Thorax: nota with some scattered setae. Legs: fore coxa carinated, bearing one fine seta and furnished with 3 – 5 spines on carina and 0 – 2 on ventral side; fore trochanter with six spines; fore, mid, and hind tibia bearing 3, 2, 2 apical spurs and 0, 2, 0 subapical spurs respectively and a row of five to six spines. Abdomen: tergites with scattered setae. Sternites with long setae, erect or slightly directed forward, often coloured and smaller setae directed backwards. STRUCTURE (measurements in Table 19, Fig. 30). Size: the workers of I. congoensis are, on average, among the smallest of the genus Isognathotermes: Fig. 30. Head capsule weakly sclerotised (except mandibles). Antennae 14 – 14.5 articles. Labrum: cupola shaped, wider than long. Left mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth three in number, first marginal well developed but with a blunt tip even when fresh, second marginal faint (visible as an undulated edge and disappearing in worn mandibles), third marginal with a blunt tip; premolar tooth with its proximal end generally hidden under molar prominence; molar tooth bearing a rounded molar prominence dorsally and ending posteriorly in a tiny acute apophysis. Right mandible: apical tooth well developed with a sharp tip when fresh; marginal teeth two in number; first marginal well developed with a sharp tip when fresh; second marginal smaller and with a blunt tip even when fresh; molar tooth bearing a ventral rounded flange and ending posteriorly in a kind of heel. Thorax: pronotum sellate. Fore coxa flanged ventrally resulting in a sharp carina. Gut: enteric valve seating on left side, best seen in ventral view, situated in posterior half of abdomen. Arrangement of enteric valve cushions of the finitimus pattern with triradial symmetry: the odd PCs, in their downstream part, are wide and bear at that place a very high density of rather short bristles on a globular bulge; supporting bristles are numerous: 22 – 33 on each side of the odd PCs; secondary cushions are wide at the upstream end, narrowing noticeably downstream with a homogeneous spine scattering. Caecum rather small, visible in ventral view, near centre of abdomen, shortly lobed (small lobes, sometimes only sketched). Chorology-ecology The type of this species was collected near the mouth of the Congo River (Fig. 38) and despite several collecting sessions in the Kongo-Central Province by A. Bouillon and his collaborators and by other entomologists, no other samples of this species have been found there. However, a sample from Cameroon matches I. congoensis morphology. Molecular data No genetic sequence is currently available.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BFFF77FE3AF92FFEADFD2C.taxon	description	Fig. 33	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BFFF77FE3AF92FFEADFD2C.taxon	diagnosis	Diagnosis This species has a soldier of planifrons morphology; nothing distinguishes it from other planifrons but the mitogenome has a dissimilarity> 2 % with the other samples of I. planifrons. It is therefore considered a possible cryptic species. However, there is only one sample that has this genetic signature, so it has not yet been described.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BFFF77FE3AF92FFEADFD2C.taxon	materials_examined	Material examined One sample. CAMEROON • soldier; Nsimi; 3 ° 10 ′ N, 11 ° 58 ′ E; Nov. 1996; M. Harry leg.; study code: DJ P 141; GenBank no PQ 679213 (mitogenome); MNHN EP 9827. Historical review This putative cryptic species is known from a single nest, sampled in 1996, by one of us (MyH) between Nsimi and Zoétélé, Cameroon. Molecular data The mitogenome of DJ P 141 exhibits up to 2.70 % dissimilarity with the other samples of I. planifrons (Supp. file 4).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BEFF70FDE6FD1DFB57FA19.taxon	description	Fig. 33 (as ‘ I. sp. ’)	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BEFF70FDE6FD1DFB57FA19.taxon	diagnosis	Diagnosis The worker has a fungifaber EVA and is difficult to distinguish from other species. The soldier’ morphology is similar to that of I. ugandensis malawii and I. minitabundus. The imago is unknown. However, this sample (DJ 0937) has a peculiar molecular signature, and as there is only one sample that has this genetic signature, it has therefore not yet been described.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BEFF70FDE6FD1DFB57FA19.taxon	materials_examined	Material examined One sample. MALAWI • soldier, worker; Nyika N. P.; 10 ° 48.642 ′ S, 33 ° 35.384 ′ E; 20 Jun. 2019; P. Stiblik leg.; study code: DJ 0937; GenBank no PV 564649 (mitogenome); BE RMCA INS. Iso. 059470. Historical review This unknown species was collected by one of us (PK) in Malawi. Molecular data This unidentified sample from Malawi was placed as sister to all species of the clade V (containing similifinitimus + phallicaecalis + planifrons + phalloides + finitimus + minitabundus + ugandensis + severus), and exhibited its highest similarity with I. minitabundus (96.5 %). Junior synonyms Isognathotermes antennalis (Sjöstedt, 1924) Figs 67 – 68, distribution map: Fig. 49 Revised herein as junior synonym of I. ugandensis ugandensis.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BEFF70FDE6FD1DFB57FA19.taxon	etymology	Etymology The name antennalis probably refers to the 14 - segmented antennae of the soldier (instead of 15 in other species), underlined by Sjöstedt (1924 c: 493) in the original description. Historical review The soldier of Cubitermes antennalis was briefly described by Sjöstedt (1924 c: 493), mentioning that it is close to C ubitermes severus Silvestri, 1914, but with antennae of 14 articles. Two years later he provided more detailed descriptions of all castes (Sjöstedt 1926: 252 – 253), he included C. antennalis in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped frons (Sjöstedt 1926: 218 – 226). Emerson (1928: 513 – 515) referred to this species under the name Mirotermes (Cubitermes) antennalis and provided further descriptions of all castes on material collected by Lang-Chapin in the Garamba National Park (Haut-Uele province, DRC) but mentioned “ antennae with 14 or 15 segments ”. Snyder (1949: 156) catalogued Cubitermes antennalis in the sub-family Termitinae. Williams (1966: 107 – 108) considered that Cubitermes antennalis Sjöstedt, 1924 was a synonym of Cubitermes ugandensis Fuller, 1923, since the syntype soldiers of the former fell well within the range of variation shown by the very large amount of material of the latter. Krishna et al. (2013: 1943) housed Cubitermes antennalis in the sub-family Cubitermitinae, as a junior synonym of C. ugandensis. Josens & Deligne (2019: 62) placed this species within the fungifaber valve pattern group and echoed the opinion of Williams, considering C. antennalis as a junior synonym of C. ugandensis. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. In this study, we relate I. antennalis to the subspecies I. u. ugandensis. Two syntype subsamples have been examined (see I. ugandensis, DJ 0013 and DJ 0031). Krishna et al. (2013: 1943) mention that another subsample is deposited in NHRM (not examined).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B9FF71FDE3F9FCFD37FA5E.taxon	description	Revised herein as junior synonym of I. fungifaber.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B9FF71FDE3F9FCFD37FA5E.taxon	etymology	Etymology In the original description, Emerson (1928: 522 – 523) stated: “ I have named the species in honor of Dr. Nathan Banks, the authority on the termites of the United States and the Neotropical region. ” Historical review Emerson (1928) described this species under the name Mirotermes (Cubitermes) banksi in the family Termitidae. He provided a description, biometrical data and figures of imago and soldier, together with a much shorter description of the worker. He noted several morphological differences between imago and soldier of the new species and those already described among which C. fungifaber. However, he did not compare with C. comstocki or C. schmidti. Snyder (1949: 156) catalogued Cubitermes banksi in the sub-family Termitinae. This is one of the species that Ruelle (1992: 501) called “ forgotten species ”. Krishna et al. (2013: 1913) housed this species in the sub-family Cubitermitinae. According to them, Emerson wrote in his unpublished “ card catalog ” that C. banksi should be considered as a junior synonym of C. fungifaber. This latter opinion of Emerson had already been reported by Ruelle (1992: 501). Synonymy of the two species is compatible with their enteric valve morphology, as both species belong to the same fungifaber valve pattern group (Josens & Deligne 2019: 39 – 42). However, I. banksi, along with I. comstocki and I. schmidti are rather small in comparison with most I. fungifaber samples (see Table 6). To clarify this possible synonymy, one of us (PA) went and collected new samples at Bipindi, Cameroon (the type locality of Cubitermes banksi as well as of C. comstocki and C. schmidti), and at Mbongé, Cameroon (one of the type localities of C. fungifaber). Indeed, some of the samples he collected, especially at Bipindi, were very small and might previously have been identified as I. banksi, I. comstocki, or I. schmidti. Subsequent COII and 28 S sequences placed all these samples in one single “ I. fungifaber clade ” (Hellemans et al. 2021). Therefore, I. banksi (Emerson, 1928) is confirmed as a junior synonym of I. fungifaber (Sjöstedt, 1896). Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Two subsamples have been examined (see Isognathotermes fungifaber, DJ 0097 and DJ 0287). Krishna et al. (2013: 1943) mention that other subsamples are deposited in UMMZ: holotype (soldier) and morphotype (winged imago), not examined.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B8FF72FDCEFA02FDE9FB4E.taxon	description	Revised herein as junior synonym of I. fungifaber.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824B8FF72FDCEFA02FDE9FB4E.taxon	etymology	Etymology In the original description, Emerson (1928: 525) stated: “ I have named the species in honor of Prof. J. H. Comstock, to whom I owe a great deal of my enthusiasm for the study of insects. ” Historical review Emerson (1928: 523 – 525) described this species under the name Mirotermes (Cubitermes) comstocki. He provided a description (with measurements and figures) of queen and soldier, mentioning differences with other species, among which M. (C.) fungifaber. Snyder (1949: 158) catalogued C. comstocki in the sub-family Termitinae and reported its known geographical distribution. This is one of the species that Ruelle (1992: 501) called “ forgotten species ”. Krishna et al. (2013: 1918) housed C. comstocki in the subfamily Cubitermitinae. They reported that Emerson (in his unpublished “ card catalog ”) considered C. comstocki as a junior synonym of C. fungifaber. This latter opinion of Emerson was also reported by Ruelle (1992: 501). Synonymy of the two species is compatible with their enteric valve morphology, as both species belong to the same fungifaber valve pattern group (Josens & Deligne 2019: 39 – 42). Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Three subsamples have been examined (see Isognathotermes fungifaber, DJ 0121, DJ 0273, and DJ 0278). Krishna et al. (2013: 1943) mention that other subsamples are deposited in UMMZ: holotype (soldier) and morphotype (queen), not examined. Isognathotermes comstocki, along with I. banksi and I. schmidti are rather small in comparison with I. fungifaber (see Table 6). As already explained for I. banksi, new material was collected by one of us (PA) and sequenced (Hellemans et al. 2021: 233) with the conclusion that the four above species belong to the same clade. Therefore, I. comstocki (Emerson, 1928) is confirmed as a junior synonym of I. fungifaber (Sjöstedt, 1896).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BBFF73FE7EFB11FE0CFB8E.taxon	description	Figs 58 – 59 Revised herein as junior synonym of I. planifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BBFF73FE7EFB11FE0CFB8E.taxon	etymology	Etymology The epithet elongata from the Latin elongatus (elongated) refers to the soldier’s head, longer than in I. fungifaber. Historical review Sjöstedt (1924 b: 256) very briefly described the soldier of this taxon under the name Cubitermes fungifaber var. elongata, mentioning that it had a longer head than the soldier of the main form. Two years later he included this taxon in his soldier’s key of Cubitermes species (Sjöstedt 1926: 221 – 225). Snyder (1949: 159) catalogued this taxon under the name Cubitermes fungifaber var. elongata in the sub-family Termitinae and reported its known geographical distribution. According to Krishna et al. (2013: 1923), Emerson wrote in his unpublished “ card catalog ” that the subspecies C. fungifaber elongata was invalid and proposed that this subspecies became a junior synonym of C. fungifaber. However, this synonymy can no longer be accepted because the types of both taxa belong to different valve pattern groups, i. e., finitimus and fungifaber groups respectively (Josens & Deligne 2019: 42 – 44). Among the museum material identified as Cubitermes fungifaber var. elongata, the syntype morphology and EVA agree well with Cubitermes planifrons described in the same paper (Sjöstedt 1924 b: 255 – 256). Other samples from the AMNH and bearing the same name are also misidentified and belong to either I. fungifaber or I. severus. Other samples from the RMCA belong to I. phalloides sp. nov. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. One subsample has been examined (see Isognathotermes fungifaber, DJ 0270). Krishna et al. (2013: 1943) mention that other subsamples are deposited in UMMZ: holotype (soldier) and morphotype (queen), not examined. Given its EVA and morphology and since it was collected in the same type of ecosystem, Cubitermes fungifaber var. elongata (Sjöstedt, 1924) is from now on considered a junior synonym of Isognathotermes planifrons (Sjöstedt, 1924).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BAFF6CFDE6FB50FCAFFCC3.taxon	description	Figs 58 – 59 Revised herein as junior synonym of I. planifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824BAFF6CFDE6FB50FCAFFCC3.taxon	etymology	Etymology In the original description, Emerson (1928: 517) stated: “ I have named the species in honor of Dr F. M. Gaige, through whose courtesy I was allowed to study the Cameroon collections included in this report. ” Historical review Emerson (1928: 516 – 517) described this species under the name Mirotermes (Cubitermes) gaigei. He provided a description (with measurements and figures) of soldier together with a few measurements of the worker. He considered that the species was very close to C. loubetsiensis, and a little larger than C. fungifaber; he mentioned some differences with other species. Snyder (1949: 159) catalogued C. gaigei in the sub-family Termitinae. This is one of the species that Ruelle (1992: 501) called “ forgotten species ”. Krishna et al. (2013: 1923) housed C. gaigei in the sub-family Cubitermitinae. Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Two subsamples have been examined (see Isognathotermes planifrons, DJ 0274 and DJ 0288). Krishna et al. (2013: 1923) mention that another subsample is deposited in UMMZ: holotype (soldier), not examined. Isonathotermes gaigei has a morphology that matches well that of I. planifrons and was collected in the same type of ecosystem; therefore, I. gaigei (Emerson, 1928) is from now on considered a junior synonym of Isognathotermes planifrons (Sjöstedt, 1924).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A5FF6DFDD2FC96FCC6FDD3.taxon	description	Figs 35 – 37 Revised herein as junior synonym of I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A5FF6DFDD2FC96FCC6FDD3.taxon	description	Junior synonym of Isognathotermes bulbifrons (Sjöstedt, 1924). Syn. nov.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A5FF6DFDD2FC96FCC6FDD3.taxon	etymology	Etymology The epithet gibbifrons from the Latin gibber (humped) and frons (forehead) refers to the upwards humped frons of the soldier. Historical review Sjöstedt (1924 b: 255) briefly described the soldier of this species under the name Cubitermes gibbifrons, mentioning that its soldier is close to the soldier of C. heghi but smaller. Two years later Sjöstedt (1926: 249 – 250) provided a first description of the alate imago and worker together with additional information on soldier. He included (1926: 218 – 225) Cubitermes gibbifrons in the imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped soldier’s frons. Snyder (1949: 160) catalogued this species under the name Cubitermes gibbifrons in the sub-family Termitinae and reported its known geographical distribution. Bouillon & Vincke (1971: 269) described the enteric valve of C. gibbifrons as belonging to the first of three types, the “ simple type ” without any spatula. This is one of the species that Ruelle (1992: 500) called “ forgotten species ”. Krishna et al. (2013: 1923) housed Cubitermes gibbifrons in the sub-family Cubitermitinae. Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. However, sequencing of the mitogenomes, despite the difference in size, confirmed synonymy with I. bulbifrons, a synonymy that had been suspected by the fact that samples of I. bulbifrons and I. gibbifrons had been collected in the same locations.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A4FF6EFDEDFD87FCAFFDAD.taxon	description	Revised herein as junior synonym of I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A4FF6EFDEDFD87FCAFFDAD.taxon	etymology	Etymology In the original description, Sjöstedt (1924 b: 255) stated: “ This species is named in honour of Mr E. Hegh, attached to the Belgian Ministry of Colonies and author of the beautiful and well-researched summary work ‘ Les Termites’ ”. Historical review Sjöstedt (1924 b: 255) briefly described the soldier of this species under the name Cubitermes heghi. Two years later he provided a more detailed description of soldier and a first description of worker (Sjöstedt 1926: 249). He included Cubitermes heghi in his soldiers’ key of Cubitermes species (1926: 221 – 226) and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped soldier’s frons. Emerson (1928: 511 – 513) referred to this species under the name Mirotermes (Cubitermes) heghi and provided a redescription (with figures) of the soldier; he noted that the soldiers of M. (C.) heghi and of M (C) bulbifrons were similar to each other but that the imagines were very different. However, the imago described by Emerson was misidentified and belonged to Cubitermes (now Polyspathotermes) sulcifrons, characterized by a large fontanelle. Snyder (1949: 160) catalogued this species under the name Cubitermes heghi and housed it in the sub-family Termitinae. Ruelle (1975: 7) considered C. bulbifrons as senior synonym of C. heghi; however, later, Ruelle (1992: 500) gave precedence to C. heghi over C. bulbifrons. Krishna et al. (2013: 1916) housed C. bulbifrons in the sub-family Cubitermitinae. Espousing the view of Ruelle 1975 but contesting the reversal of precedence proposed by Ruelle 1992, they considered this species as the senior synonym of C. heghi. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. One subsample has been examined (see Isognathotermes bulbifrons, DJ 0290). Krishna et al. (2013: 1917) mention that other paratype subsamples are deposited in PPRI (not examined) and IEAP; however, the material that was lent by this latter museum did not contain any Cubitermes heghi sample. Synonymy of both species is compatible with their enteric valve morphology, as both species belong to the same finitimus valve pattern group (Josens & Deligne 2019: 42 – 44). Their morphology also supports synonymy. Following Ruelle (1975) and Krishna et al. (2013), I. heghi is therefore considered a junior synonym of Isognathotermes bulbifrons (Sjöstedt, 1924).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A7FF6FFDD9FD70FC59FE76.taxon	description	Revised herein as junior synonym of I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A7FF6FFDD9FD70FC59FE76.taxon	etymology	Etymology In the original description, Emerson (1928: 509) stated: “ I take pleasure in naming this species in honor of Dr. N. Kemner, who has done such valuable work on oriental termites and termitophiles. ” Historical review Emerson (1928: 509) described this species under the name Mirotermes (Cubitermes) kemneri. He provided description and measurements of the soldier, noting that it is very close to Cubitermes heghi. Snyder (1949: 161) catalogued this species under the name C. kemneri and housed it in the sub-family Termitinae. Ruelle (1975: 8), on the advice of Emerson (in litteris), proposed C. kemneri as a junior synonym of C. zenkeri. This is one of the species that Ruelle (1992: 501) called “ forgotten species ”. Krishna et al. (2013: 1945) housed C. kemneri in the sub-family Cubitermitinae. Espousing the view of Ruelle 1975 they considered this species as a junior synonym of C. zenkeri. However, C. kemneri cannot be synonymized with C. zenkeri because the types of both species have different EVAs, i. e., of the finitimus and fungifaber groups respectively (Josens & Deligne 2019: 42 – 44). Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. One type subsample has been examined (see Isognathotermes bulbifrons, DJ 0297). Krishna et al. (2013: 1945) mention that other subsamples are deposited in UMMZ: holotype (soldier), and a paratype subsample in PPRI (not examined). Only two samples in the museum collections bore the name “ C. kemneri ”: the type sample and a misidentified sample (presently assigned to I. fungifaber). Our morphology study (this article) shows that the type matches well I. bulbifrons. Therefore, I. kemneri (Emerson, 1928) is from now on considered a junior synonym of Isognathotermes bulbifrons (Sjöstedt, 1924).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A6FF6FFDC2FE29FE0CF825.taxon	description	Fig. 58 Revised herein as junior synonym of I. planifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A6FF6FFDC2FE29FE0CF825.taxon	etymology	Etymology The epithet loubetsiensis (from Loubetsi and Latin suffix - ensis, from) refers to the type locality (Loubetsi, Republic of Congo). Historical review Sjöstedt (1924 c: 493) briefly described the soldier of this species under the name Cubitermes loubetsiensis, mentioning that it is close to the soldier of C. subarquatus. Two years later, he provided a first description and some measurements of imago and worker (Sjöstedt 1926: 246 – 247). He included (1926: 218 – 226) this species in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ fungifaber - Gruppe ” of species with a not humped soldier’s frons. Emerson (1928: 515 – 516) re-described the species under the name Mirotermes (Cubitermes) loubetsiensis based on specimens collected at Medje (DRC, Haut-Uele Province). He provided a redescription of queen and soldier together with some measurements of worker. However, these specimens were misidentified (Krishna et al. 2013: 1926); they are presently assigned to I. finitimus. Snyder (1949: 161) catalogued this species under the name Cubitermes loubetsiensis and housed it in the sub-family Termitinae. Krishna et al. (2013: 1926) housed Cubitermes loubetsiensis in the sub-family Cubitermitinae. They reported that Emerson (in his unpublished “ card catalog ”) considered C. loubetsiensis as a junior synonym of C. finitimus. Synonymy of the two species is compatible with their EVAs, as Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Two type subsamples have been examined, both limited to the soldier caste (see Isognathotermes planifrons, DJ 0281 and DJ 0611). Only four samples in the museum collections bore the name “ C. loubetsiensis ”: the type samples and two misidentified samples (presently assigned to I. finitimus). Our morphology study (this article) shows that the type matches well I. planifrons. Therefore, I. loubetsiensis (Sjöstedt, 1924) is from now on considered a junior synonym of Isognathotermes planifrons (Sjöstedt, 1924).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A1FF68FDD3FEEBFD5EF8FA.taxon	description	Figs 28, 30 – 31, 61 – 63, distribution map: Fig. 45 Revised herein as junior synonym of I. severus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A1FF68FDD3FEEBFD5EF8FA.taxon	etymology	Etymology The epithet modestior means ‘ smaller’ in Latin. It referred to the smaller size of the variety C. severus modestior when compared to the main form, as underlined by Silvestri in the original description. Historical review Silvestri (1914: 93 – 94) described this taxon under the name Cubitermes severus var. modestior; it is known from a single nest. He provided a short description of imago, soldier, and worker together with measurements and figures, mentioning that the imagines are like the typical form (= Cubitermes severus), and indeed they are among the smallest individuals of the other I. severus imagines but also that the soldiers and workers are smaller than the main form. Sjöstedt (1926: 238 – 239) considered that this taxon deserved to be elevated to the species level. He referred to it under the name Cubitermes modestior, housed it in the sub-family Termitinae, and included it in the “ glebae - Gruppe ” of small Cubitermes species. Snyder (1949: 161) catalogued this species under the name C. modestior and housed it in the sub-family Termitinae. Krishna et al. (2013: 1927) referred to this species under the name C. modestior and housed it in the sub-family Cubitermitinae. Josens & Deligne (2019: 39 – 42) placed this species within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Two type subsamples have been examined (see Isognathotermes severus, DJ 0294 and DJ 0341). Krishna et al. (2013: 1927) mention that another subsample is deposited in PPRI (not examined). Since the type material of I. modestior (Silvestri, 1914) comes from a single nest and is included in the smaller specimens of I. severus (from other regions), it is from now on considered a junior synonym of Isognathotermes severus (Silvestri, 1914).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A1FF69FDD7F8DEFE73FA2E.taxon	description	Revised herein as junior synonym of I. fungifaber.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A1FF69FDD7F8DEFE73FA2E.taxon	etymology	Etymology In the original description, Emerson (1928: 521) stated: “ I have named the species in honor of Mr. Karl P. Schmidt, to whom I owe my first introduction to termites. ” Historical review Emerson (1928: 520 – 521) described this species under the name Cubitermes (Mirotermes) schmidti in the family Termitidae. He provided a description, biometrical data and a figure of imago and soldier, together with a few measurements of worker. Snyder (1949: 163) catalogued C. schmidti in the sub-family Termitinae. This is one of the species that Ruelle (1992: 501) called “ forgotten species ”. Krishna et al. (2013: 1935) referred to this species under the name C. schmidti and housed it in the sub-family Cubitermitinae. They reported that Emerson (in his unpublished “ card catalog ”) considered C. schmidti a junior synonym of C. fungifaber. This latter opinion of Emerson is also reported by Ruelle (1992: 501). Synonymy of the two species is compatible with their enteric valve morphology, as Josens & Deligne (2019: 39 – 42) placed C. schmidti within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. Two type samples have been examined (see Isognathotermes fungifaber, DJ 0124 and DJ 0275). Isognathotermes schmidti, along with I. banksi and I. comstocki are rather small in comparison with I. fungifaber (see Table 6). As already explained for I. banksi, new material was collected by one of us (PA) and sequenced (Hellemans et al. 2021) with the conclusion that the four above species belong to the same clade. Therefore, I. schmidti (Emerson, 1928) is confirmed as a junior synonym of I. fungifaber (Sjöstedt, 1896).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A0FF6AFDDDF9F1FD5EFB22.taxon	description	Figs 62 – 63 Revised herein as junior synonym of I. severus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A0FF6AFDDDF9F1FD5EFB22.taxon	etymology	Etymology The species was named in honour of F. Silvestri who collected the type material. Historical review Sjöstedt (1925: 54) briefly described the soldier of this species under the name Cubitermes Silvestrii. One year later Sjöstedt (1926: 251) provided more detailed descriptions together with figures of the soldier, specifying that they were based on specimens from Kakoulima (Guinea) first identified as Cubitermes severus by Silvestri. He housed the species in the sub-family Termitinae, included it (1926: 218 – 226) in a soldier’s key of Cubitermes species and inserted it in a “ bulbifrons - Gruppe ” of species with an upwards humped frons. Snyder (1949: 164) catalogued Cubitermes silvestrii in the sub-family Termitinae. This is one of the species that Ruelle (1992: 500) called “ forgotten species ”. Krishna et al. (2013: 1937) housed C. silvestrii in the sub-family Cubitermitinae. Josens & Deligne (2019: 39 – 42) placed this species within the fungifaber valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. One type sample has been examined (see Isognathotermes severus, DJ 0302). Krishna et al. (2013: 1937) mention that another subsample is deposited in NHRM (not examined). The type of Cubitermes silvestrii came from the same locality as one of the syntypes of Cubitermes severus and our morphology study shows that it matches well I. severus. Therefore, Cubitermes silvestrii (Sjöstedt, 1925) is from now on considered a junior synonym of Isognathotermes severus (Silvestri, 1914).	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A3FF6BFDD7FAF6FA91FC0B.taxon	description	Figs 39 – 41 Revised herein as junior synonym of I. finitimus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A3FF6BFDD7FAF6FA91FC0B.taxon	etymology	Etymology The epithet speciosus means ‘ beautiful, superb’ in Latin. Historical review Sjöstedt (1924 b: 256) briefly described the soldier of this species under the name Cubitermes speciosus, mentioning that it is close to the soldier of C. fungifaber but larger. Two years later Sjöstedt (1926: 242 – 243) provided a first description of alate imago and worker together with additional information on soldier. He included (1926: 218 – 225) C. speciosus in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ fungifaber - Gruppe ” of species with a not protruding soldier’s frons. Snyder (1949: 164) catalogued C. speciosus in the sub-family Termitinae. Williams (1966: 102 – 104) provided a detailed redescription of imago and soldier along with measurements and numerous figures. Bouillon & Vincke (1971: 269) described the enteric valve of C. speciosus as belonging to the “ simple type ” without any spatula. Krishna et al. (2013: 1937) housed C. speciosus in the sub-family Cubitermitinae. Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes. One type sample has been examined (see Isognathotermes finitimus, DJ 0061). Krishna et al. (2013: 1937) mention that other syntype samples are deposited in AMNH and PPRI (not examined). As the type material of Isognathotermes speciosus (Sjöstedt, 1924) matches well Isognathotermes finitimus (Schmitz, 1916) and come from the same forested environment, they are from now on considered as synonyms.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A2FF64FDC5FBCFFEA7FBF2.taxon	description	Figs 39 – 41 Revised herein as junior synonym of I. finitimus.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A2FF64FDC5FBCFFEA7FBF2.taxon	description	Isognathotermes aff. subarquatus “ spA ” junior synonym of Isognathotermes fungifaber. Isognathotermes aff. subarquatus “ spB ” junior synonym of Isognathotermes planifrons. Isognathotermes aff. subarquatus “ spC ” junior synonym of Isognathotermes bulbifrons. Syn. nov. Isognathotermes aff. subarquatus “ spD ” junior synonym of Isognathotermes fungifaber.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
038B976824A2FF64FDC5FBCFFEA7FBF2.taxon	etymology	Etymology The epithet subarquatus from the Latin sub- (to a lesser degree) and arquatus (yellowish) can refer to the red-yellowish colour of the soldier’s head mentioned in the original description, but arcuatus (or arquatus) also means ‘ curved, in the shape of a bow’. Historical review Sjöstedt (1926: 244 – 245) described alate imago, soldier, and worker, along with a few measurements and provided figures of the soldier’s head. He included (1926: 218 – 226) Cubitermes subarquatus in imagines’ and soldiers’ keys of Cubitermes species and inserted it in a “ fungifaber - Gruppe ” of species with a not protruding soldier’s frons. Emerson (1928: 510 – 511) referred to this species under the name Mirotermes (Cubitermes) subarquatus. He provided further descriptions (with measurements and figures) of imago and soldier together with photographs of the nest. Snyder (1949: 164) catalogued C. subarquatus in the sub-family Termitinae. Krishna et al. (2013: 1938) housed C. subarquatus in the sub-family Cubitermitinae. They reported that Emerson (in his unpublished “ card catalog ”) considered C. subarquatus a junior synonym of C. finitimus. Synonymy of the two species is compatible with their EVAs, as Josens & Deligne (2019: 42 – 44) placed this species within the finitimus valve pattern group. Hellemans et al. (2021: 233) placed this species in the restored genus Isognathotermes; and COII and 28 S sequences placed it in the “ I. finitimus clade ”. Our morphology study (this article) also shows that the type matches well I. finitimus. Roy et al. (2006) pointed to the possible existence of cryptic species on base of nuclear and mitochondrial markers. Among the four cryptic species, two of them (“ spA ” and “ spD ”) had fungifaber - EVAs and their DNA sequences also classified them in I. fungifaber (Hellemans et al. 2021). The two other cryptic species (“ spB ” and “ spC ”) had finitimus - EVAs: “ spB ” was recognized as I. planifrons and “ spC ” as I. bulbifrons.	en	Josens, Guy, Deligne, Jean, Harry, Myriam, Roy, Virginie, Akama, Pierre D., Coulibaly, Tenon, Dosso, Kanvaly, Goergen, Georg, Hasson, Michel, Kasangij, Patrick Kasangij A, Kifukieto, Carmel, Ru, Bruno Le, Loko, Laura Estelle Yêyinou, Ndiaye, Abdoulaye Baila, Roisin, Yves, Sion, Noémie, Šobotnik, Jan, Stiblik, Petr, Kuenda, Soki Kue Di, Traoré, Saran, Viage, Manuela, Wango, Solange Patricia, Kaymak, Esra, Bourguignon, Thomas, Hellemans, Simon (2025): An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies. European Journal of Taxonomy 1024: 1-197, DOI: 10.5852/ejt.2025.1024.3099, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3099/13837
