Aenictus Shuckard 1840
Creators: name: Gómez, Kiko
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hymenoptera
Formicidae
Aenictus

description: Genus <i>Aenictus</i> Shuckard, 1840 <i>Aenictus</i> SHUCKARD, 1840: 266. Type-species: <i>Aenictus ambiguus</i>, by original designation. <i>Enictus</i> WALKER, 1860: 306; SMITH, 1865: 79; incorrect subsequent spellings of <i>Aenictus</i>: BOLTON, 1995: 19. [unavailable name]. <i>Paraenictus</i> WHEELER, 1929: 27 [as subgenus of <i>Aenictus</i>]. Type-species: <i>Aenictus (Paraenictus) silvestrii</i>, by monotypy. [Junior synonym of <i>Aenictus</i>: WILSON, 1964: 444]. <i>Typhlatta</i> SMITH, 1857: 79. Type-species: <i>Typhlatta laeviceps</i>, by monotypy. [Junior synonym of <i>Aenictus</i>: FOREL, 1890: ciii.; revived from synonymy as subgenus of <i>Aenictus</i>: WHEELER, 1930: 198; junior synonym of <i>Aenictus</i>: WILSON, 1964: 444]. An excellent account on morphology, phylogeny, taxonomy and biology of the genus is given in BOROWIEC (2016), including a detailed description for worker and male castes. FISHER &amp; BOLTON (2016) also diagnoses the worker caste for the Afrotropical species. The following notes complement these two main sources and apply solely to the worker caste of the Afrotropical species. Most species are monomorphic, with some having a wide range of sizes (e.g. CSR up to 144 for <i>A. eugenii</i>) but without marked allometric morphological differences among the maxima and minima workers other than size, colour or sculpturation. However, some species in the <i>rotundatus</i> group represent the extreme of this variation with noticeable allometry in some indexes (e.g. <i>A. hitai</i> sp. nov. 0.28&lt;HW&lt;0.60, CSR 178, 74&lt;CI&lt;100) which led EMERY (1895) to comment about <i>A. mariae</i> “it is hardly correct to say that workers of this species are monomorphic”. As a rule of thumb, minima workers present smaller SIL and CI than maxima workers, meaning that they have more elongated heads with relatively shorter scapes. Some Asian <i>Aenictus</i> species possess clearer cuticular patches laterally at the upper half of the head (“ <i>Typhlatta</i> spot”), but this character is always absent in the Afrotropical species described to date. In the Afrotropical species antennae are 10 segmented, with subcylindrical scapes, bent outwards in frontal view and widening in its apical half to almost twice its width, usually the basal third narrow and the rest widening to the apex. Its length range from short to medium (30&lt;SIL&lt;100) not reaching the occipital margin and never as long as in some Asian species groups where scapes reach and even surpass the occipital margin (<i>A</i>. <i>hottai</i>, <i>A</i>. <i>inflatus</i>, <i>A</i>. <i>laeviceps</i>, <i>A</i>. <i>pachycerus</i>, <i>A</i>. <i>wroughtonii</i> (part) groups) (JAITRONG &amp; YAMANE, 2011). Mandible shape is remarkably diverse among the Afrotropical species and relatively stable within species groups. As a consequence, mandible shape is the main character used to define such groups. Most species display linear or triangular shapes, but may have highly specialized morphology (as in the <i>decolor</i> or <i>popeyei</i> groups which are not present in any other <i>Aenictus</i> group worldwide). Mandibular dentition is also variable and linked to the mandibular shape. In triangular mandibles it consists of a long apical tooth followed or not by a preapical, and up to 8–12 smaller denticles, but in the <i>rixator</i> group with 3–4 defined teeth, in the <i>asantei</i> group just an apical tooth and in the <i>decolor</i> and <i>popeyei</i> groups apical and preapical tooth. No Afrotropical species has a row of basal teeth or denticles as seen in the Asian <i>minutulus</i> (formerly <i>piercei</i>) group (JAITRONG &amp; HASHIMOTO, 2012) nor a large basal tooth as in the Asian <i>javanus</i> group (JAITRONG &amp; YAMANE, 2012). Sometimes a sharp, differentiated edge can be present basally (e.g. <i>A. popeyei</i>), but in most species the basal mandibular margin is flat and undifferentiated. Usually pro- and mesonotum are convex and the propodeum almost flat, both meeting at an angle, except for some minute species with almost flat mesosoma in lateral view (e.g. <i>rixator</i> species group or minima workers in <i>rotundatus</i> species group). Mesonotum-mesopleuron ridge is always absent (present in the Asian <i>philippinensis group</i>) (JAITRONG &amp; YAMANE, 2012). Petiole is always sessile or weakly subsessile. Subpetiolar process is always present, highly specific and interspecifically variable, with a long range of shapes and sizes and presenting or not a flat lamella. Petiolar carinae are usually present and developed: anterolateral and anterodorsal carinae are often present, while dorsolateral carinae, when present, are weak and not surpassing the petiolar spiracle (except for the heavily sculptured <i>A. susanae</i>). A few heavily sculptured species may present petiolar posterolateral and/or posterodorsal carinae. Femora and tibiae with their distal halves swollen (maximum width/minimum width~3–4 typically), except for the <i>koloi</i> species group, in which femora and tibiae have cylindrical shapes. Setae are always present, usually unequal, with very long and shorter setae, from semierect to erect in the whole body. A few species may have reclined and similar in length setae (e.g. <i>A. mvuvii</i>, <i>A. jacki</i>). Funiculus is always coated with short, dense pubescence and semierect short setae, slightly longer than funiculus width. Pubescence is absent in the rest of the body, except for <i>A. steindachneri</i> Mayr, 1901, which presents short, dense pubescence on lateropropodeum and petiolar sides. SPECIES GROUPS: AFROTROPICAL VS. PALAEARCTIC, ASIAN AND AUSTRALASIAN FAUNA <i>West Palaearctic</i> The Western Palaearctic <i>Aenictus</i> fauna lacks a global taxonomic revision and comprises four species: <i>A. arabicus</i> Sharaf &amp; Aldawood, 2012; <i>A. dlusskyi</i> Arnol’di, 1968; <i>A. rhodiensis</i> Menozzi, 1936 and <i>A. vaucheri</i> Emery, 1915. JAITRONG &amp; RUANGSITTICHAI (2018) have modified the initial definition of the <i>wroughtonii</i> group given in JAITRONG &amp; YAMANE (2011) to include species with short scapes and listing <i>A. arabicus</i> and <i>A. rhodiensis</i> into that group. <i>Asia, Indomalaya and Australasia</i> WILSON (1964) defined five species groups for the Indo-Australian area based on morphological characters. These groups were modified and refined for the Eastern Oriental, Indo-Australian and Australasian regions (JAITRONG &amp; YAMANE, 2011; JAITRONG &amp; HASHIMOTO, 2012; JAITRONG &amp; RUANGSITTICHAI, 2018). The current definition includes twelve groups and comprises the whole distribution range for the genus excluding the four Palaearctic species and the Afrotropical region. Twelve Afrotropical species were included in WILSON (1964), and though these species were not included in the final groups, a brief statement was given on the African fauna: “On examining the numerical data on African <i>Aenictus</i> […], it can be seen that the following form a closely knit group: <i>asperivalvus</i>, <i>mariae</i>, <i>mentu</i>, <i>rotundatus</i>, <i>steindachneri</i> and <i>weissi</i>. These, in turn are allied to the <i>ceylonicus</i> group of species in Asia, especially <i>peguensis</i> of Burma” [WILSON, 1964: 436]. The inclusion of <i>asperivalvus</i> in the analysis must be considered as an error, as it’s only known from males. GOTWALD &amp; BARR (1988) follow Wilson’s definition and include all the Afrotropical species studied into this group. Further redefinition of the group (JAITRONG &amp; YAMANE, 2011) leave aside almost all of the Afrotropical species. Also note that all the mentioned species but <i>mentu</i> are included in this revision under the <i>rotundatus</i> species group. Lacking molecular and/or genetic evidence, the morphological information suggests that the relation of Afrotropical and Asian groups appears to be weak. This is suggested from the following characters: <b>•</b> the absence of the “ <i>Typhlatta</i> spot” in any Afrotropical species excludes the <i>currax</i>, <i>inflatus</i>, <i>laeviceps</i> and <i>leptotyphlatta</i> groups; <b>•</b> scapes relatively short-medium sized (usually SIL&lt;85) and never reaching posterolateral corner of head excludes <i>hottai</i> and <i>pachycerus</i> groups. This is true except for the Afrotropical group <i>decolor</i> with relatively long scapes (SIL 85-100), but their characteristic mandibular morphology is not found in any Asian species; <b>•</b> 10 segmented antennae excludes <i>silvestrii</i> group; <b>•</b> absence of a conspicuous row of denticles at the basal margin of mandibles excludes <i>minutulus</i> (formerly <i>piercei</i>) group (JAITRONG &amp; HASHIMOTO, 2012); <b>•</b> the absence of a mesonotum-mesopleuron ridge excludes the <i>philippinensis</i> group; <b>•</b> the lack of a masticatory margin with a large basal tooth excludes <i>javanus</i> group. The remaining two groups (<i>ceylonicus</i> and <i>wroughtonii</i>) deserve a more detailed analysis. The <i>ceylonicus</i> group comprises the species, among other characteristics <i>“with mandibles closed, a gap present between mandibles and anterior margin of clypeus; anterior clypeal margin weakly concave or almost straight, lacking denticles”</i> (JAITRONG &amp; YAMANE, 2010, see also LIU <i>et al</i>., 2015). Two Afrotropical species have linear mandibles that do not close against the clypeus (<i>A. eugenii</i> and <i>A. mvuvii</i>) but both species present two small clypeal denticles between (<i>eugenii)</i> or just below (<i>mvuvii)</i> the antennal insertions. The <i>wroughtonii</i> group (clypeus with a row of triangular teeth and poorly developed subpetiolar process) has been recently redefined (JAITRONG &amp; RUANGSITTICHAI, 2018) to include the species with short scapes. Under this new definition, several Afrotropical species could belong to this group except for the fact that very similar species may or may not present or not developed subpetiolar processes, even in series from the same nest in some species. The above data strongly suggests that the similarities of the Afrotropical groups with their Asian counterparts is more of convergent nature, and there is substantial morphological differentiation between the groups in both biogeographical regions. Therefore, instead of expanding the definitions of <i>ceylonicus</i> and <i>wroughtonii</i> groups, and along with the other clearly differentiated Afrotropical five groups, I prefer to define the new <i>eugenii</i> and <i>rotundatus</i> groups for the Afrotropical species until genetic or molecular studies are available and the real affinities can be properly established. Despite this decision, these two pairs of species groups seem to be the best candidates for a link between Asian and Afrotropical faunas. AFROTROPICAL SPECIES GROUPS: DEFINITION AND IDENTIFICATION TIPS Unfortunately, genetic data for the reviewed species were not available for this study and building a phylogeny to define species groups based on it was not possible. From this point of view, the species groups defined in this paper should be considered as informal and preliminary, waiting for complementary studies. This revision proposes organizing the Afrotropical species in seven groups, is based mainly on mandibular and clypeal shape. These two characters alone seem to be good enough to separate the species groups in a robust manner. The <i>A</i>. <i>koloi</i> group is defined based on its cylindrical legs and sculptured heads (which are swollen and glassy smooth respectively in the other groups). The <i>rotundatus</i> group is also separated into two species complexes based on SIL index. Besides the cited clypeal and mandibular shape, the most helpful characters to differentiate species are subpetiolar process, propodeal shape, scape length relative to head length (SIL), sculpture and setation. When mounting specimens it is important to lift the abdomen upwards and push the third pair of legs forward, so that the subpetiolar process is visible. SYNOPSIS OF AFROTROPICAL SPECIES Only species with described worker caste. [Q] ‘Queen also described’ <i>asantei</i> group <i>asantei</i> Campione, Novak &amp; Gotwald, 1983 [Q] <i>decolor</i> group <i>bidentatus</i> Donisthorpe, 1942 <b>stat. rev.</b> <i>decolor</i> (Mayr, 1879) [Q] = <i>batesi</i> Forel, 1911 <i>villiersi</i> Bernard, 1953 <i>eugenii</i> group <i>eugenii</i> Emery, 1895 [Q] <i>=brazzai</i> Santschi, 1910 <b>syn. nov.</b> <i>= eugenii caroli</i> Forel, 1910 <b>syn. nov.</b> = <i>eugeniae v. kenyensis</i> Santschi, 1933 <i>= eugenii henrii</i> Santschi, 1924 <b>syn. nov.</b> <i>mvuvii</i> <b>sp. nov.</b> <b> <i>koloi</i> group</b> <i>koloi</i> <b>sp. nov.</b> <i>susanae</i> <b>sp. nov.</b> <i>xegi</i> <b>sp. nov.</b> <i>popeyei</i> group <i>popeyei</i>sp. nov. <i>rixator</i> group <i>mentu</i> Weber, 1942 <i>rixator</i> Emery, 1901 <i>rotundatus</i> group <i>mariae species complex</i> <i>boltoni</i> <b>sp. nov.</b> <i>hitai</i> <b>sp. nov.</b> <i>mariae</i> Emery, 1895 <i>= mariae natalensis</i> Emery, 1901 <b>syn. nov.</b> <i>steindachneri</i> Mayr, 1901 <i>rotundatus</i> species complex <i>congolensis</i> Santschi, 1911 [Q] <i>guineensis</i> Santschi, 1924 <i>jacki</i> <b>sp. nov.</b> <i>nyuyi</i> <b>sp. nov.</b> <i>rotundatus</i> Mayr, 1901 <i>=furibundus</i> Arnold, 1959 <b>syn. nov.</b> = <i>rotundatus merwei</i> Santschi, 1932 <b>syn. nov.</b> <i>ugaduwensis</i> <b>sp. nov.</b> <i>weissi</i> Santschi, 1910 [Q cited, not described] KEY TO SPECIES (WORKERS) 1 Head glassy smooth (Fig. 2 A); femora with their distal halves expanded (Fig. 2 C) (Maximum width / minimum width ~ 3–4).............................................................................. 2 - Head alutaceus to heavily reticulated, matt (Fig. 2 B); femora subcylindrical (Fig. 2 D) (Maximum width / minimum width &lt;2)........................................................... 7 (<i>koloi</i> group) 2 (1) Mandibles either linear or triangular, never with either a large blunt, rounded basal tooth nor with its basal half expanded and semi-spherical (Fig. 1 A, C, D, G, H)............................ 3 - Mandibles massive, neither linear nor triangular, either with a large, blunt, rounded basal tooth separated from the rest by a depression (mitten-shaped) or with its basal half expanded and semi-spherical (Fig. 1 E, F)...................................................................................................... 6 3 (2) Mandibles linear, leaving a gap between themselves and the clypeus when closed. Clypeus reduced to two triangular minute teeth between or beneath the antennal sockets (Fig. 1 C, D)........................................................................................................................ 11 (<i>eugenii</i> group) - Mandibles linear to triangular with no gap between themselves and the clypeus when mandibles closed. Clypeus usually with a row of minute triangular denticles, and never reduced to two triangular minute teeth between or beneath the antennal sockets (Fig. 1 A, G, H)............................................................................................................................................................ 4 4(3) Mandibles very long, crossing over at the mid-length when closed, each mandibular apex clearly reaching the opposite antennal insertion; apical tooth massive, usually without preapical tooth or denticles (Fig. 1 G)............................................. (<i>asantei</i> group), one species, <i>asantei</i> - Mandibles linear to triangular, neither crossing over at the mid-length nor with each mandibular apex; preapical tooth and/or denticles may be present (Fig. 1 A, H).................... 5 5 Mandibles linear, with three or four defined teeth. Clypeus reduced to a thin line, virtually not visible in frontal view (Fig. 1 H)............................................................. 12 (<i>rixator</i> group) - Mandibles triangular, always with more than 4 teeth or denticles in total. Dentition variable, usually with one apical tooth followed by a preapical tooth and/or a series of smaller denticles, but some species with an apical tooth followed by smaller well defined triangular teeth. Clypeus present as a narrow lamella or a row of triangular denticles, sometimes small and difficult to see when mandibles closed (Fig. 1 A).................................................... 13 (<i>rotundatus</i> group) 6 (2) Mandibles with basal half expanded and semi-spherical, surrounded by a thin cutting edge (Fig. 1 E)........................................................... <i>popeyei</i> group (one species, <b> <i>popeyei</i> sp. nov.</b> ) - Mandibles with a conspicuous, blunt, large proximal tooth clearly separated from the rest of the mandible, mitten shaped (Fig. 1 F)........................................................... 9 (<i>decolor</i> group) KEY FOR <i>koloi</i> GROUP 7 (5) Numerous long, white, semierect to erect setae present on scapes, dorsum of mesosoma, petiole, postpetiole and femora. Petiole with clearly more than 10 setae (Fig. 22 C).......................................................................................................................................... <b> <i>susanae</i> sp. nov.</b> - Scattered white, erect setae present on scapes, dorsum of mesosoma, petiole, postpetiole and femora. Petiole with 4 to 8 setae.............................................................................................. 8 8 (7) Subpetiolar process small, size similar to the prora, shaped as a quarter of ellipse with the vertical face facing forward, followed by an almost horizontal ovoid lamella (Fig. 20 C); smaller size (HW&lt;0.60)...................................................................................... <b> <i>koloi</i> sp. nov.</b> - Subpetiolar process very developed, subrectangular and followed by a rectangular lamella usually inclined 45 degrees forward, the whole process clearly larger than the prora (Fig. 24 C); larger size (HW&gt;0.65)........................................................................................... <b> <i>xegi</i> sp. nov.</b> KEY FOR <i>decolor</i> GROUP 9 (8) Scapes relatively shorter, slightly surpassing three quarters of head length when laid back (SIL&lt;80), smaller size (HL&lt;0.70, WL&lt;1.15)........................................................... <i>bidentatus</i> - Scapes relatively longer, distinctly surpassing three quarters of head length when laid back (SIL&gt;80), larger size (HL&gt;0.70, WL&gt;1.15)........................................................................... 10 10 (9) Scapes reaching three quarters of the head length when laid back (80&lt;SIL&lt;95). Subpetiolar process subrectangular, volume slightly smaller than petiolar dome and of similar shape, displaced forward (Fig. 7 C)....................................................
description: Published as part of <i>Gómez, Kiko, 2022, A revision of the Afrotropical species of the Dorylinae ant genus Aenictus (Hymenoptera: Formicidae) based on the worker caste, pp. 1-86 in Belgian Journal of Entomology 124</i> on pages 6-16, DOI: 10.5281/zenodo.5898821
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