Cephalotes Latreille 1802
Creators: name: Oliveira, Aline Machado
name: Powell, Scott
name: Feitosa, Rodrigo Machado
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hymenoptera
Formicidae
Cephalotes

description: Cephalotes Latreille, 1802
Type-species: Formica atrata, by monotypy.
[Type-species not Formica cephalotes, unjustified subsequent designation by Wheeler, 1911: 160; corrected by Wheeler, 1913: 78].
Cephalotes as junior synonymofCryptocerus: Fabricius, 1804: 419.
Cephalotes as senior synonym of Cryptocerus: Wheeler, 1913: 78.
Cephalotes in Myrmicinae, Cryptocerini: Emery, 1914: 42; Forel, 1917: 246; Wheeler, 1922: 665; Emery, 1924: 303; all subsequent authors to 1949, and Dlussky and Fedoseeva, 1988: 79 (anachronism).
Cephalotes in Myrmicinae, Cephalotini: Smith, 1949: 19; Kempf, 1951: 105; all subsequent authors except the above.
Cephalotes in Myrmicinae, Attini: Ward et al., 2015: 17.
General diagnosis: Females: Frontal carinae covering the genae in frontal view. Mandibles small and thick, with two apical teeth. Eyes produced laterally, very close to the vertex. Antennae short, with 11 articles. Deep antennal scrobes. Humeral angles with spines or lamellae. Hard integument. Male: Size smaller than or subequal to gyne. Frontal carinae short to obsolete. Antennae with 13 articles, remarkably longer than in the females. Antennal scrobes vestigial to absent. Propodeum unarmed or armed with a pair of obtuse angles ordenticles.
Worker (Fig. 1A): Mandibles small and thick, with twoapical teeth. Palpal formula 5:3. Clypeus narrow. Frontal lobesgreatly expanded and covering the genae. Frontal carinae divergent posteriorly andreaching the vertex.Antennalscrobes deeplyexcavated.Antennae with 11 articles. Lateral margins of pronotum withspines, denticles or lamellae. Dorsum of propodeum without a transversal crest. Midand hind tibiae without apical spurs. The first gastral tergite comprises almost the total length of gaster. Sting reduced or missing.
Soldier (Figs. 1C, D): Presentin most species. Differsfrom the worker by the presence of a variably developed transverse crest or carina on dorsum of pronotum and, in most species, by the head specialization, which variesindegree of modification infour morphotypes (Powellet al., 2020). Of the species presentin Brazil, the squaremorphotype occurs in the atratus, basalis, clypeatusandmanni groups.The dome morphotype occurs in thebasalis, pusillus, anddepressus groups.The discmorphotype occursin the angustus, depressus, grandinosus, pinelii, umbraculatus, and fiebrigi groups. Finally, the dish morphotype occurs only in the pallens group. Thesoldier for the Brazilianrepresentatives of thecoffeae species group (Cephalotes trichophorus) and thesolidus species group (Cephalotes solidus) are currently unknown.
Gyne (Figs. 1E, G): Head generally like thesoldier; whenthe soldier caste is absent, gyne’s head is similar to the workers but usually larger and moreconvex.Eyescomparatively large; ocellialwayspresent.Scutum andscutellum weaklyconvex to flat. Forewingswith Sc+R meeting the strongly pigmented stigma; Rextendsbeyond the stigma, not reaching the external margin of wing; Rs meeting Rposteriorly and forming a marginalcell with 2r-rs; M+Cu diverging in Mand Cu, forming a discal cell with m-cu; Mand Cu extends towards margin after discal cell, but spectral for most of its length, not reaching the wing margin; vein A extends beyond the cu-a, which is interrupted, not connecting M+Cu and A; claval furrow marked by a conspicuous notch; length of R, M, Cu, and A can be variable between specimens, but never reaching the margin of wing. Hindwing with R+Rs diverging in R1 and Rs; M+Cu diverging in Cu and rs-m+M; vein Aextends beyond the cu-a, which is interrupted, not connecting M+Cu and A; length of R1, Rs, Cu, and Acan be variable between specimens, but never reaching the wing margin; hamuli variable innumber between species. Gasterlonger than mesosoma, with four tergites visible, the first one occupying almost the total length of gaster.
Male (Fig.1B):Size smaller thanorsubequaltogyne.Headsubcircular indorsal view.Mandibleslike workers, butmasticatory margins usually withasingle apical tooth, oftenfollowed by aseries of denticles. Clypeus subrectangular. Frontal carinae short to obsolete. Antennal scrobes vestigial to absent. Antennae with 13 articles, remarkablylonger than in thefemales;scape shorterthansecond funicularsegment; pedicel shorter thanscape, antennal condyleexposed. Mesosomawithnotauli Y-shaped, deeplyimpressed. Propodeum unarmed orarmed with a pair of obtuse angles or denticles. Hind tibiae usually without apical spurs. Petiole cylindrical or nodiform. Wings as in gyne. Gaster usually shorter than mesosoma, with five visible tergites; tergite Ialwayslarger than others.
Taxonomic synopsis of Cephalotes for Brazil
angustus speciesgroup
Cephalotes adolphi (Emery, 1906)
Cephalotes angustus (Mayr, 1862)
Cephalotes conspersus (Smith, 1867)
Cephalotes frigidus (Kempf, 1960)
Cephalotes gabicamacho newspecies
Cephalotes goeldii (Forel, 1912)
Cephalotes marycorn newspecies
Cephalotes monicaulyssea newspecies
Cephalotes notatus (Mayr, 1866)
Cephalotes pallidicephalus (Smith, 1876)
Cephalotes targionii (Emery, 1894)
atratus species group
Cephalotes atratus (Linnaeus, 1758)
= Cephalotes marginatus (Fabricius, 1804) new synonym Cephalotes oculatus (Spinola, 1851)
Cephalotes opacus Santschi, 1920
Cephalotes placidus (Smith, 1860)
Cephalotes serraticeps (Smith, 1858)
basalis speciesgroup
Cephalotes basalis (Smith, 1876)
Cephalotes complanatus (Guérin-Méneville, 1844)
Cephalotes cordiae (Stitz, 1913)
Cephalotes ramiphilus (Forel, 1904)
clypeatus speciesgroup
Cephalotes clypeatus (Fabricius, 1804)
Cephalotes membranaceus (Klug, 1824)
Cephalotes ustus (Kempf, 1973)
coffeae speciesgroup
Cephalotes trichophorus De Andrade, 1999
depressus species group
Cephalotes betoi De Andrade, 1999
Cephalotes borgmeieri (Kempf, 1951)
Cephalotes cordatus (Smith, 1853)
Cephalotes depressus (Klug, 1824)
Cephalotes eduarduli (Forel, 1921)
Cephalotes palustris De Andrade, 1999
Cephalotes pavonii (Latreille, 1809)
fiebrigi speciesgroup
= bruchi species group new synonym
Cephalotes bruchi (Forel, 1912)
Cephalotes fiebrigi (Forel, 1906)
Cephalotes guayaki De Andrade, 1999
Cephalotes jheringi (Emery, 1894)
Cephalotes liviaprado newspecies
Cephalotes pilosus (Emery, 1896)
Cephalotes quadratus (Mayr, 1868)
Cephalotes specularis Brandão, Feitosa, Powelland Del-Claro, 2014
grandinosus speciesgroup
Cephalotes grandinosus (Smith, 1860)
Cephalotes klugi (Emery, 1894)
Cephalotes persimilis De Andrade, 1999
Cephalotes persimplex De Andrade, 1999
manni speciesgroup (here proposed)
Cephalotes manni (Kempf, 1951)
pallens speciesgroup
Cephalotes pallens (Klug, 1824)
Cephalotes pallidoides De Andrade, 1999
Cephalotes pallidus De Andrade, 1999
Cephalotes patellaris (Mayr, 1866)
Cephalotes pellans De Andrade, 1999
pinelii speciesgroup
Cephalotes incertus (Emery, 1906)
Cephalotes liepini De Andrade, 1999
Cephalotes maculatus (Smith, 1876)
Cephalotes mariadeandrade newspecies
Cephalotes nilpiei De Andrade, 1999
Cephalotes pinelii (Guérin-Méneville, 1844)
pusillus species group
= laminatus species group new synonym
Cephalotes duckei (Forel, 1906)
Cephalotes inaequalis (Mann, 1916)
Cephalotes laminatus (Smith, 1860)
Cephalotes minutus (Fabricius, 1804)
Cephalotes pusillus (Klug, 1824)
Cephalotes simillimus (Kempf, 1951)
Cephalotes spinosus (Mayr, 1862)
solidus speciesgroup
Cephalotes solidus (Kempf, 1974)
umbraculatus speciesgroup
Cephalotes umbraculatus (Fabricius, 1804)
Key to the identification of Brazilian species groups of Cephalotes based on workers
(Figs. 2 a-r)
1 In lateralview, vertexal corners with a pairof spines (pd, pv) (Fig.2a). Comparatively larger ants............................................................ atratusgroup
1’ Vertexal corners can have projections, but in a different pattern than above, usually as lamellar expansions (vle) (Figs. 2b, c). Comparatively smaller ants...................................................................................................................2
2 Indorsalview, mesonotumandpropodeumwithoutlateralprojections (Fig. 2e).............................................................................................. solidusgroup
2’ Mesonotumandpropodeum with lateral projections.These projections can be denticles, spines (ps) (Figs. 2 f-h) or lamellar expansions (ple) (Figs. 2i, k-l) sometimes restricted to the declivous face of propodeum (Fig. 42c)....................................................................................................................3
3 In dorsal view, lateral margins of propodeum with lamellar expansions (ple) (Figs. 2i, k-l) sometimes restricted to the declivousface of propodeum (Fig. 42c)............................................................................................4
3’ Lateral margins of propodeum with denticles or spines (ps) (Figs. 2 f-h).............................................................................................................7
4 In dorsal view, gaster completely surrounded by a lamella (Fig. 2o)....... ........................................................................................................... clypeatus group
4’ In dorsal view, gaster with lamellar expansions restricted to the anterior portion (ge) (Figs. 2 k-l, p-r)...............................................................5
5 Infrontal view, vertexal corner extending laterally, overhanging the eyes (Fig. 2d - dashed). Body color reddish brown (Fig. 2d, i) ................ ............................................................................................................. pallens group
5’ In frontal view, vertexal corners not extending laterally (Fig. 2c - dashed). Body variable in color.........................................................................6
6 Metafemur with lamellar expansion (fle) on dorsal and/ or ventral face, which frequently are crenulate and narrow (Fig. 2k).................................................................................................. grandinosus group
6’ Metafemur without lamellarexpansions (Fig. 2l).......... pineliigroup
7 In dorsal view, propodeum with two pairs of spines, the anterior one longer than the posterior and curved backwards (Fig. 2g). Spines sometimes with denticles.................................................... depressusgroup
7’ Indorsal view, propodeumwithvariablenumberof spines (Fig. 2h); if there are two pairs, theanterior oneisnever longer than theposterior (Fig. 2f)....................................................................................................................8
8 In dorsal view, first gastral tergite yellowish, with a cross-shaped dark macula (Fig. 2p)..................................................... umbraculatusgroup
8’ Firstgastral tergite variablein color; maculaabsent (Fig. 2q) or never cross-shaped (Fig. 2r)............................................................................................9
9 Posterior femora witha medianprojection (Fig.2j). Metatibiae usually marginated (Fig. 2j).......................................................................... basalisgroup
9’ Posterior femora without projections (Fig. 2l). Metatibiae never marginated (Fig. 2l)............................................................................................10
10 In dorsal view, anterior gastral expansions (ge) withouta translucent lamella (Fig. 2q).....................................................................................................11
10’ In dorsal view, anterior gastral expansions (ge) translucent (Fig. 2r).. ...................................................................................................................................12
11 Dorsal and declivous faces of propodeum meeting at a distinct propodeal angle; lateral margins of the declivous face with broad lamellar expansion from the posterior dorsal spines of propodeum to the petiolar insertion..................................................................... mannigroup
11’ Dorsal and declivous faces of propodeum continuous, not meeting at a distinct propodeal angle; lateral margins of declivous face without lamellarexpansions....................................................................... fiebrigigroup
12
In dorsal view, propodeum with two pairs of spines, the posterior one longer than the declivous face of propodeum (Fig. 2f)...................... ....................................................................................................... pusillus group
12’ Indorsal view, propodeumwithvariablenumberof spines, none of them longer than the declivous face of propodeum (Fig. 2h) .............. ..................................................................................................................................13
13 Indorsal view, margins of declivous face of propodeum withlamellar expansions (Fig. 2m). In dorsal view, margins of pronotum with lamellar expansion, which can be crenulate, but not forming denticles or spines............................................................................................ coffeaegroup
13’ In dorsal view, margins of declivous face of propodeum without lamellarexpansions (Fig.2n). In dorsal view, margins of pronotum with denticlesor spines...................................................................... angustus group Key to the identification of Brazilian species groups of Cephalotes based on soldiers
(Figs. 3 a-p)
Note 1: soldiers are not present in the Brazilian species of coffeae and solidus groups.
Note 2: soldiers have a transversal crest or carina on the dorsum of pronotum (Fig. 1D), except in atratusgroup, which can be identified with the worker’s key.
Note 3: for a better comprehension of the morphology of soldier`s head see Fig. 1C.
1 In frontal view, cephalic dorsum completely covering the mandibles (Fig. 3a).............................................................................................. pallens group
1’ In frontal view, mandibles visible, at least partially (Figs. 3 b-e)..........2
2 In frontal view, dorsum of head completely marginated by a carina (blue dotted), forming a disc (Fig. 3b). Vertexal corners (pink dotted) separated from the dorsal cephalic disc (Fig. 3b).......................................3
2’ In frontal view, dorsum of head square shaped (Fig. 3c) or dome shaped (Fig. 3d, e), but never disc shaped. In both cases, the vertexal corners (pink dotted) are not separated from the dorsum of head by a carina (blue dotted) (Fig. 3 c-e).........................................................................8
3 In dorsalview, petiole andpostpetiole with lateral lamellar expansions (Fig. 3h)....................................................................................................................4
3’ Petiole and postpetiole with lateral acute projections, forming spines ordenticles (Figs. 3i, j, m, o, p).........................................................................5
4 Posterior femora with lamellar expansions in dorsal and/ or ventral face (Fig. 3k), which frequently are crenulate and narrow....................................................................................... grandinosus group
4’ Posterior femora without lamellar expansions (Fig. 3l)........................... ................................................................................................................... pineliigroup
5 In dorsal view, propodeum with two pairs of spines, the anterior one longer than the posterior one and slightly curved backwards (Fig. 3j)...................................................................... depressusgroup (in part)
5’Indorsalview,propodeumwithvariablenumberof spines,iftherearetwo pairs, the anterior one is never longer than the posterior (Figs. 3f, g, i)........6
6 Body yellowish; first gastral tergite witha cross-shaped dark macula (Fig. 3p). In dorsal view, pronotum crest with pointed edges (Fig. 3f)...... ................................................................................................ umbraculatusgroup
6’ Body black to darkbrown; gastermacula absent or in ashape different thana cross (Fig.3o), if cross-shaped, thenpronotumcrest with rounded or subrectangular edges (Fig. 3g)....................................................................7
7 In dorsal view, anterior expansions of gaster without a translucent lamella (Fig. 3m)......................................................... fiebrigigroup (in part)
7’ In dorsalview, anterior gasterexpansions with atranslucent lamella, usually very thin (Fig. 3o)........................................................ angustus group
8 In frontal view, frontal carinae continuous withthe vertexal corners, cephalic dorsum square shaped (Fig. 3c)......................................................9
8’ In frontal view, frontal carinae ends before reaching the vertexal corners, cephalic dorsum dome shaped (Fig. 3d, e)................................11
9 Gaster completely surrounded by lamellar expansions (Fig. 3n)....... ........................................................................................................ clypeatus group
9’ Lamellar expansions, if any, present only in the anterior portion of the gaster (Fig. 3o, p)...........................................................................................10
10 Posterior femora with a median projection, metatibiae marginated......................................................................................... basalisgroup
10’ Posterior femora without projections, metatibiae never marginated................................................................................................ mannigroup
11 In frontal view, head longer than wide, with contiguous foveae and suberect hairs (Fig. 3e).............................................. fiebrigigroup (in part)
11’ Infrontalview, headsubquadrate, with foveae separatebyinterspaces, and short appressed hairs (Fig. 3d)..............................................................12
12 Propodeum withtwo pairs of lateral projections, the posterior one longer than the anterior one (Fig. 3i).................................... pusillus group
12’ Propodeum with two pairs of lateral projections, the anterior one longer than the posterior one (Fig. 3j)............ depressusgroup (in part)
description: Published as part of Oliveira, Aline Machado, Powell, Scott & Feitosa, Rodrigo Machado, 2021, A taxonomic study of the Brazilian turtle ants (Formicidae: Myrmicinae: Cephalotes), pp. 1-52 in Revista Brasileira de Entomologia 65 (3) on pages 3-7, DOI: 10.1590/1806-9665-RBENT-2021-0028, http://zenodo.org/record/5512220