Cephalotes Latreille, 1802

Oliveira, Aline Machado, Powell, Scott & Feitosa, Rodrigo Machado, 2021, A taxonomic study of the Brazilian turtle ants (Formicidae: Myrmicinae: Cephalotes), Revista Brasileira de Entomologia 65 (3), No. e 20210028, pp. 1-52 : 3-7

publication ID	https://doi.org/ 10.1590/1806-9665-RBENT-2021-0028
DOI	https://doi.org/10.5281/zenodo.5512228
persistent identifier	https://treatment.plazi.org/id/038587E5-FFF6-FFC6-FCB3-FB84D197FC43
treatment provided by	Carolina
scientific name	Cephalotes Latreille, 1802
status

Treatment

Cephalotes Latreille, 1802 View in CoL

Type-species: Formica View in CoL atrata, by monotypy.

[Type-species not Formica View in CoL cephalotes, unjustified subsequent designation by Wheeler, 1911: 160; corrected by Wheeler, 1913: 78].

Cephalotes View in CoL as junior synonymofCryptocerus: Fabricius, 1804: 419.

Cephalotes View in CoL 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 View Figure 1 ): 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 View Figure 1 ): 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 View Figure 1 ): 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 View Figure 1 ):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 View in CoL based on workers

( Figs. 2 View Figure 2 a-r)

1 In lateralview, vertexal corners with a pairof spines (pd, pv) ( Fig.2a View Figure 2 ). Comparatively larger ants............................................................ atratusgroup View in CoL

1’ Vertexal corners can have projections, but in a different pattern than above, usually as lamellar expansions (vle) ( Figs. 2b, c View Figure 2 ). Comparatively smaller ants...................................................................................................................2

2 Indorsalview, mesonotumandpropodeumwithoutlateralprojections ( Fig. 2e View Figure 2 ).............................................................................................. solidusgroup View in CoL

2’ Mesonotumandpropodeum with lateral projections.These projections can be denticles, spines (ps) ( Figs. 2 View Figure 2 f-h) or lamellar expansions (ple) ( Figs. 2i View Figure 2 , k-l) sometimes restricted to the declivous face of propodeum ( Fig. 42c View Figure 42 )....................................................................................................................3

3 In dorsal view, lateral margins of propodeum with lamellar expansions (ple) ( Figs. 2i View Figure 2 , k-l) sometimes restricted to the declivousface of propodeum ( Fig. 42c View Figure 42 )............................................................................................4

3’ Lateral margins of propodeum with denticles or spines (ps) ( Figs. 2 View Figure 2 f-h).............................................................................................................7

4 In dorsal view, gaster completely surrounded by a lamella ( Fig. 2o View Figure 2 )....... ........................................................................................................... clypeatus View in CoL group

4’ In dorsal view, gaster with lamellar expansions restricted to the anterior portion (ge) ( Figs. 2 View Figure 2 k-l, p-r)...............................................................5

5 Infrontal view, vertexal corner extending laterally, overhanging the eyes ( Fig. 2d View Figure 2 - dashed). Body color reddish brown ( Fig. 2d, i View Figure 2 ) ................ ............................................................................................................. pallens View in CoL group

5’ In frontal view, vertexal corners not extending laterally ( Fig. 2c View Figure 2 - 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 View Figure 2 ).................................................................................................. grandinosus View in CoL group

6’ Metafemur without lamellarexpansions ( Fig. 2l View Figure 2 ).......... pineliigroup View in CoL

7 In dorsal view, propodeum with two pairs of spines, the anterior one longer than the posterior and curved backwards ( Fig. 2g View Figure 2 ). Spines sometimes with denticles.................................................... depressusgroup View in CoL

7’ Indorsal view, propodeumwithvariablenumberof spines ( Fig. 2h View Figure 2 ); if there are two pairs, theanterior oneisnever longer than theposterior ( Fig. 2f View Figure 2 )....................................................................................................................8

8 In dorsal view, first gastral tergite yellowish, with a cross-shaped dark macula ( Fig. 2p View Figure 2 )..................................................... umbraculatusgroup View in CoL

8’ Firstgastral tergite variablein color; maculaabsent ( Fig. 2q View Figure 2 ) or never cross-shaped ( Fig. 2r View Figure 2 )............................................................................................9

9 Posterior femora witha medianprojection ( Fig.2j View Figure 2 ). Metatibiae usually marginated ( Fig. 2j View Figure 2 ).......................................................................... basalisgroup View in CoL

9’ Posterior femora without projections ( Fig. 2l View Figure 2 ). Metatibiae never marginated ( Fig. 2l View Figure 2 )............................................................................................10

10 In dorsal view, anterior gastral expansions (ge) withouta translucent lamella ( Fig. 2q View Figure 2 ).....................................................................................................11

10’ In dorsal view, anterior gastral expansions (ge) translucent ( Fig. 2r View Figure 2 ).. ...................................................................................................................................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 View in CoL

11’ Dorsal and declivous faces of propodeum continuous, not meeting at a distinct propodeal angle; lateral margins of declivous face without lamellarexpansions....................................................................... fiebrigigroup View in CoL

In dorsal view, propodeum with two pairs of spines, the posterior one longer than the declivous face of propodeum ( Fig. 2f View Figure 2 )...................... ....................................................................................................... pusillus View in CoL group

12’ Indorsal view, propodeumwithvariablenumberof spines, none of them longer than the declivous face of propodeum ( Fig. 2h View Figure 2 ) .............. ..................................................................................................................................13

13 Indorsal view, margins of declivous face of propodeum withlamellar expansions ( Fig. 2m View Figure 2 ). 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 View Figure 2 ). In dorsal view, margins of pronotum with denticlesor spines...................................................................... angustus View in CoL group Key to the identification of Brazilian species groups of Cephalotes View in CoL based on soldiers

( Figs. 3 View Figure 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 View Figure 1 ), 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. View Figure 1

1 In frontal view, cephalic dorsum completely covering the mandibles ( Fig. 3a View Figure 3 ).............................................................................................. pallens View in CoL group

1’ In frontal view, mandibles visible, at least partially ( Figs. 3 View Figure 3 b-e)..........2

2 In frontal view, dorsum of head completely marginated by a carina (blue dotted), forming a disc ( Fig. 3b View Figure 3 ). Vertexal corners (pink dotted) separated from the dorsal cephalic disc ( Fig. 3b View Figure 3 ).......................................3

2’ In frontal view, dorsum of head square shaped ( Fig. 3c View Figure 3 ) or dome shaped ( Fig. 3d, e View Figure 3 ), 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 View Figure 3 c-e).........................................................................8

3 In dorsalview, petiole andpostpetiole with lateral lamellar expansions ( Fig. 3h View Figure 3 )....................................................................................................................4

3’ Petiole and postpetiole with lateral acute projections, forming spines ordenticles ( Figs. 3i, j, m, o, p View Figure 3 ).........................................................................5

4 Posterior femora with lamellar expansions in dorsal and/ or ventral face ( Fig. 3k View Figure 3 ), which frequently are crenulate and narrow....................................................................................... grandinosus View in CoL group

4’ Posterior femora without lamellar expansions ( Fig. 3l View Figure 3 )........................... ................................................................................................................... pineliigroup View in CoL

5 In dorsal view, propodeum with two pairs of spines, the anterior one longer than the posterior one and slightly curved backwards ( Fig. 3j View Figure 3 )...................................................................... depressusgroup View in CoL (in part)

5’Indorsalview,propodeumwithvariablenumberof spines,iftherearetwo pairs, the anterior one is never longer than the posterior ( Figs. 3f, g, i View Figure 3 )........6

6 Body yellowish; first gastral tergite witha cross-shaped dark macula ( Fig. 3p View Figure 3 ). In dorsal view, pronotum crest with pointed edges ( Fig. 3f View Figure 3 )...... ................................................................................................ umbraculatusgroup View in CoL

6’ Body black to darkbrown; gastermacula absent or in ashape different thana cross ( Fig.3o View Figure 3 ), if cross-shaped, thenpronotumcrest with rounded or subrectangular edges ( Fig. 3g View Figure 3 )....................................................................7

7 In dorsal view, anterior expansions of gaster without a translucent lamella ( Fig. 3m View Figure 3 )......................................................... fiebrigigroup View in CoL (in part)

7’ In dorsalview, anterior gasterexpansions with atranslucent lamella, usually very thin ( Fig. 3o View Figure 3 )........................................................ angustus View in CoL group

8 In frontal view, frontal carinae continuous withthe vertexal corners, cephalic dorsum square shaped ( Fig. 3c View Figure 3 )......................................................9

8’ In frontal view, frontal carinae ends before reaching the vertexal corners, cephalic dorsum dome shaped ( Fig. 3d, e View Figure 3 )................................11

9 Gaster completely surrounded by lamellar expansions ( Fig. 3n View Figure 3 )....... ........................................................................................................ clypeatus View in CoL group

9’ Lamellar expansions, if any, present only in the anterior portion of the gaster ( Fig. 3o, p View Figure 3 )...........................................................................................10

10 Posterior femora with a median projection, metatibiae marginated......................................................................................... basalisgroup View in CoL

10’ Posterior femora without projections, metatibiae never marginated................................................................................................ mannigroup View in CoL

11 In frontal view, head longer than wide, with contiguous foveae and suberect hairs ( Fig. 3e View Figure 3 ).............................................. fiebrigigroup View in CoL (in part)

11’ Infrontalview, headsubquadrate, with foveae separatebyinterspaces, and short appressed hairs ( Fig. 3d View Figure 3 )..............................................................12

12 Propodeum withtwo pairs of lateral projections, the posterior one longer than the anterior one ( Fig. 3i View Figure 3 ).................................... pusillus View in CoL group

12’ Propodeum with two pairs of lateral projections, the anterior one longer than the posterior one ( Fig. 3j View Figure 3 )............ depressusgroup View in CoL (in part)