taxonID	type	description	language	source
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	type_taxon	Type-species: Formica atrata, by monotypy. [Type-species not Formica cephalotes, unjustified subsequent designation by Wheeler, 1911: 160; corrected by Wheeler, 1913: 78].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	discussion	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.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	diagnosis	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.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	description	Worker (Fig. 1 A): 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. 1 C, 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. 1 E, 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 2 r-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 R 1 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 R 1, 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. 1 B): 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.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	discussion	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)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	description	(Figs. 2 a-r)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	description	(Figs. 3 a-p)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF6FFC6FCB3FB84D197FC43.taxon	discussion	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. 1 D), 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. 1 C.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF3FFC8FFDFFF0BD196F86B.taxon	description	(Figs. 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF3FFC8FFDFFF0BD196F86B.taxon	discussion	The angustus group was proposed by Kempf (1958 a) based on the subgenus Harnedia of the genus Paracryptocerus, with 26 species divided into seven subgroups. De Andrade and Baroni Urbani (1999) reorganized this scheme, reducing the angustus group to Kempf’s angustus subgroup, which was composed by C. adolphi, C. angustus, C. goeldii, C. notatus (= P. (Harnedia) fleddermanni in Kempf, 1958 a), C. pallidicephalus (= P. (Harnedia) striativentris in Kempf, 1958 a), and C. targionii. These authors also addedC. conspersus in the group, which was considered by Kempf (1951) as a taxonomically isolated species, andtwo species described bythem, C. dentidorsum and C. palta. These last two speciesare known only from Colombia and werenot included in this study. All species in this group are exclusively South American. In the morphological phylogeny by De Andrade and Baroni Urbani (1999, see their Fig. 24), this group was considered the sister group to the clade formed by the bruchi, fiebrigi and prodigiosus groups. The current phylogeny confirms the relationshipsbetweenthese groups (Fig. 54). The bruchi, fiebrigi and prodigiosus groups are recovered as monophyletic, but they render the angustus group paraphyletic, suggesting that all thesegroupsmight represent asinglelineage (Fig. 54). Morphologically, these groups share the declivous face of the propodeum continuous with its dorsal face, not meeting at a distinct propodeal angle, and with a variable number of spines, but differing by the anterior expansions of the gaster, which are a thin translucent lamella in the angustus group and a thick opaque lamella in bruchi, fiebrigi and prodigiosus groups. Although the phylogenetic data (Fig. 54) suggests theangustus, bruchi, fiebrigi andprodigiosus groups couldbe synonymized, we decidedto keepangustus asa valid groupuntil abroader phylogeny, including additional species from all these groups, isavailable. The Argentinianprodigiosus group was not included in the present study, due to itsabsence in Brazil. The discussion about bruchi and fiebrigi groups is presented in the fiebrigi group section below.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF3FFC8FFDFFF0BD196F86B.taxon	diagnosis	Diagnosis: In workers, body with appressedhairs (Fig. 2 h). Declivous face of propodeum continuous with dorsal face, not meeting at a distinct propodealangle, withavariablenumberofspines (Fig. 2 h). If onlytwopairs of spinesarepresenton propodeum, the anteriorone is neverthelongest, andthe posterioroneisnever longerthanthedeclivousface (Fig. 2 h). First tergite of gaster with anterior lamellar expansions translucent (Fig. 2 r). Insoldiers, propodeumwithvariablenumberof spines, iftherearetwo pairs, the anterior one is never longer than the posterior. Anterior gastral expansions with a translucent lamella, usually very thin (Fig. 3 o).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF3FFC8FFDFFF0BD196F86B.taxon	discussion	Brazilian species of angustus group Cephalotes adolphi (Emery, 1906) Obsolete combination: Cryptocerus adolphi Cephalotes angustus (Mayr, 1862) Obsolete combinantion: Cryptocerus angustus = Cephalotes angustus clarior (Forel, 1912) Obsolete combination: Cryptocerus angustus clarior = Cephalotes cristatus ruficeps (Santschi, 1929) Obsolete combination: Cryptocerus cristatus ruficeps Cephalotes conspersus (Smith, 1867) Obsolete combinantion: Cryptocerus conspersus = Cephalotes denticulatus variegata (Forel, 1911) Obsolete combination: Cryptocerus denticulatus var. variegata Cephalotes frigidus (Kempf, 1960) Obsolete combination: Paracryptocerus frigidus Cephalotes gabicamacho newspecies Cephalotes goeldii (Forel, 1912) Obsolete combination: Cryptocerus goeldii Cephalotes marycornnewspecies Cephalotes monicaulyssea new species Cephalotes notatus (Mayr, 1866) Obsolete combination: Cryptocerus notatus = Cephalotes fleddermanni (Kempf, 1958 a) Obsoletecombination: Paracryptocerus (Harnedia) fleddermanni Cephalotes pallidicephalus (Smith, 1876) Obsolete combination: Cryptocerus pallidicephalus = Cephalotes striativentris (Emery, 1894) Obsolete combination: Cryptocerus striativentris Cephalotes targionii (Emery, 1894) Obsolete combination: Cryptocerus targionii = Cephalotes denticulatus (Emery, 1894) Obsolete combination: Cryptocerus denticulatus	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF3FFC8FFDFFF0BD196F86B.taxon	description	(Figs. 5 a-p)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFDFFCAFFDFFB4BD619F86B.taxon	description	Figs. 6 a-c, 7 a-c, 15	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFDFFCAFFDFFB4BD619F86B.taxon	materials_examined	Holotype: BRAZIL, MT, Coxipó, ix. 1900, typus (worker), - 15.614986, - 56.030674 [MSNG] [examined byimages - AntWeb. org CASENT 0904906]. Additional material examined: Brazil, MG, Uberlândia, Clube de caça e pesca Itororó, 20. vii. 2007, - 19.004404 - 48.312329, S. Powell cols, Cerrado, / C 05 - 192 (1 soldier) [DZUP], Panga Ecological Reserve, 10. vii. 2015, - 19.174260 - 48.400940, S. Powell cols, Cerrado, / C 15 - 45 (2 soldiers) [DZUP], Panga Ecological Reserve, 10. vii. 2015, - 19.174260 - 48.400940, S. Powellcol, Cerrado, / C 15 - 45 (1 gyne) [DZUP]. Worker descriptioncan be found in De Andrade and Baroni Urbani, 1999, page 733. Soldiermeasurements (N = 3): HL 1.95 - 2.12; HW 1.72 - 1.88; EL 0.33 - 0.35; PW 1.52 - 1.76; WL 1.60 - 1.69; PTL 0.64 - 0.72; PTW 0.20 - 0.24; PPL 0.70 - 0.76; PPW 0.20 - 0.26; GL 1.88 - 1.94; HBL 0.46 - 0.48; HBW 0.11 - 0.13; TL 6.85 - 7.16; CI 87.1 - 89.7; OI 18.6 - 20; PI 266 - 327; HBI 23.1 - 27.2. Soldier (first description): Head, dorsum of mesosoma and legs yellowish to ferruginous. Propleura predominantly black with the upper surface yellowish; meso- and metapleura predominantly yellowish with dark spots. First gastral tergite yellowish with a transverse dark macula. First gastral sternite brownish (Fig. 6). Mandiblesalveolate; ventralfaceofheadweaklyfoveate, spacebetween foveaemicroalveolate; dorsumofheadirregularlyfoveate (Fig. 6 a). Dorsum of mesosomafoveate-microalveolate; lateralof mesosomafinelyrugosewith sparsefoveae. Petioleandpostpetiolesmooth. Gastermicroalveolate (Fig. 6 c). Mandibles with appressed canaliculate hairs and erect simple hairs. Head, mesosoma, petiole, postpetiole and legs with appressed canaliculate hairs; frontal carinae with erect simple hairs. Gaster with appressed simple hairs; posterior portion of first sternite and edges of each tergite and sternite of gaster with erect simple hairs (Fig. 6 b). Headlongerthanwide (CI 87.1 - 89.7). Mandibleswithastronglongitudinal lateralangle. Clypeuswithapairofdenticles (Fig. 6 a). Dorsumofheaddisc shaped, slightlyconvexanteriorly (Fig. 6 b). Frontalcarinaecrenulateanteriorly. Antennae with a three-segmented club. Roof of antennal scrobes with a lateral carinae and a posterior rounded projection. Lateroventral margins of head without carinae. Vertexal corners forming pointed projections separated of the dorsum cephalic disc (Fig. 6 a, b). In lateral view, pronotum ascending, with a transversal carina raised ina crest; pronotalcrestnotcrenulate, witha mediandepression (Fig. 6 c). Indorsalview, anteriormarginof pronotumgentlyrounded, lateralmargins with twopairs of denticles, the anterior oneacute, thesecond oneobtuse. Mesonotumandpropodeumdiscontinuousandflat (Fig. 6 b); mesonotum with a pair of blunt rounded denticles; propodeal groove impressed; dorsal and declivous faces of propodeum meeting in a distinct propodeal angle; indorsalview, lateralmarginsof propodeum witha pairof median spinesandobtuse denticlesstrongly bent dorsally, with theapices curved anteriorly (Fig. 6 b). Femoranotangulateddorsally, mid, andhindbasitarsi not flattened, with subparallel dorsal and ventral faces. Indorsalview, petiolecompressedanteroposteriorly, anteriormargin with a discrete median concavity, lateral spines curved backwards, dorsum witha pairof denticles (Fig. 6 b), subpetiolarprocess broader and rounded anteriorly (Fig. 6 c). Postpetioleslightly wider andlonger than petiole, spines narrow andcurved forward, dorsum witha “ v ” sharped elevation (Fig. 6 b), subpostpetiolar process pronounced ventrally and compressed anteroposteriorly (Fig. 6 c). Gaster elongate, with narrowanterior lamellar expansions (Fig. 6 c). Gyne measurements (N = 1): HL 1.80; HW 1.64; EL 0.40; PW 1.60; WL 2.05; PTL 0.60; PTW 0.28; PPL 0.70; PPW 0.30; GL 2.56; HBL 0.56; HBW 0.14; TL 7.71; CI 91.1; OI 24.4; PI 214; HBI 25. Gyne (first description): Body black; dorsumandlateralofhead, lateral marginsof pronotum and propodeum, centraldisc of anepisternum, and legs yellowish to ferruginous. First gastral tergite withan anterior and a posterior pair of yellowish spots, each one occupying more than one third of the first tergite length; posterior margins of each tergite and sternite yellowish (Fig. 7). Sculpture of head, petiole, postpetiole, legs and gaster as in the soldiers. Dorsum of mesosoma with sparse foveae, space between foveae microalveolate. Propleura deeply striate. Anepisternum deeply and denselyfoveate, withoutspacebetween foveae; katepisternum and metapleura finely rugose with some sparse foveae (Fig. 7 b). Declivous face of propodeum finely rugose and microalveolate. Pilosity of head as in thesoldier. Mesoscutum, axillaeand legs with some short erecthairs. First gastraltergitewith sparse appressed simple hairs and some short erect hairs; first gastral sternite with short and long appressed simple hairs, and short erect hairs (Fig. 7 c) Head as in the soldier, but shorter (HL 1.80) (Fig. 7 a). Indorsalview, anteriormarginofpronotumslightlyrounded, lateral margins with a pair of denticles pointed anteriorly, pronotal carina weakly developed, not crenulate, with a median depression (Fig. 7 c). Dorsally, mesoscutum subtriangular, anterior margins rounded; notauli absent; parapsidial lines feebly visible and parallel; transscutal line impressed, reaching the lateral margins of mesosoma; scutoscutellar groove impressed, extending laterally, separating scutellum and axillae; axillae rounded posteriorly (Fig. 7 c). Laterally, mesopleural groove dividing anepisternum and katepisternum; metapleura divided in upper and lower metapleura by a deep groove; metapleuropropodeal groove not impressed (Fig. 7 b). In dorsal view, posterior margin of propodeum concave with a pair of short, blunt denticles (Fig. 7 c). Wings unknown. Indorsal view, petiole subquadrate, anteriormargin concave and lateral margins convex, without lateral or dorsalprojections, subpetiolar process narrow. Postpetiole wider than petiole, with a “ v ” sharped dorsal elevation and lateral blunt projections, subpostpetiolar process pronounced ventrally and compressed anteroposteriorly (Fig. 7 b). Gaster with anteriorly protruding lobes (Fig. 7 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	conservation	urn: lsid: zoobank. org: act: F 19 B 756 C- 3 FA 8 - 45 BB-AA 3 F- 9 D 9480071 D 8 F Figs. 8 a-c, 9 a-c, 10 a-c, 15	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	materials_examined	Holotype: BRAZIL: MG, Uberlândia, Clube de Caça e Pesca Itororó, - 19.00473 - 48.31253, S. Powellcol. / C 06 - 94, DZUP 550148 (worker) [DZUP]. Paratypes: samedataas holotype: DZUP 550149 (2 workers), DZUP 550150 (1 worker, 1 soldier), DZUP 550151 (1 soldier, 1 gyne), DZUP 550152 (1 gyne) [DZUP], DZUP 550159 (1 worker, 1 soldier) [MZSP], DZUP 550157 (1 worker, 1 soldier) [USNM]; - 19.00124 - 48.31239, S. Powellcol. / C 05 - 134, DZUP 550153 (3 workers), DZUP 550154 (2 soldiers) [DZUP], DZUP 550158 (1 worker, 1 soldier) [MZSP], DZUP 550156 (1 soldier) [USNM], DZUP 550163 (2 workers), DZUP 550162 (1 soldier) [INPA], DZUP 550161 (2 workers) DZUP 550160 (1 soldier) [MPEG], Paraopeba, iii. 2011, C. R. Ribas, Cerrado, pitfall arbóreo [unique specimenidentifier UFV-LABECOL- 004439] (1 worker) [DZUP].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	diagnosis	Diagnosis: Amemberoftheangustus speciesgroup. In workers, propleura striate. Firstgastral sternite microalveolate, without striae (Fig. 4 m). First gastraltergiteanteriorlywithoutstriae, withabundantappressedcanaliculate hairs, thedistance betweeneachhairshorterthantheir length, appressed simplehairspresentonmiddleof tergite (Fig. 8 c). Insoldiers, indorsalview, lateral margins of pronotum straight and subparallel (Fig. 9 c). Propodeal groove strongly impressed, forming a depression (Fig. 9 c). Worker measurements (N = 15): HL 0.95 - 1.20; HW 0.98 - 1.28; EL 0.30 - 0.34; PW 0.82 - 1.03; WL 1.10 - 1.24; PTL 0.18 - 0.23; PTW 0.53 - 0.60; PPL 0.22 - 0.25; PPW 0.54 - 0.63; GL 1.40 - 1.64; HBL 0.35 - 0.40; HBW 0.08 - 0.10; TL 3.85 - 4.46; CI 95.1 - 120; OI 24.9 - 31.6; PI 30 - 38.3; HBI 20 - 26.3.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	description	Worker description: Body black; frontal lobes, anterior spots on gastral tergite, apices of each segment of legs, and external face of tibiae yellowish (Fig. 8). Mandibles, legs andgaster microalveolate. Head, mesosoma, petiole and postpetiole foveate-microalveolate. Propleura totally striate. Declivous face of propodeum sparsely striate. Body with appressed canaliculate hairs (Fig. 8 c), more concentrated on meso- and metapleura. Mandibles and anterior margin of clypeus with suberect clavate and simple hairs (Fig. 8 a). Declivous face of propodeum glabrous. First sternite of gaster with long erect simple hairs (Fig. 8 b). Posterior edge of the gastral tergites with erect hairs. Head slightly wider than long (CI 95.1 - 120), dorsum slight convex (Fig. 8 b). Mandibleswithweaklydevelopedlateral angle. Anteriormargin of clypeus concave, without lateral denticle. Frontal carinae notched anteriorly to eyes, not bent dorsally over eyes (Fig. 8 c). Antennae with three-segmented club. Lateroventral margins of head with posterior carinae extending posteriorly to eyes until vertexal corners. Vertexal corners with narrow, irregular lamellar expansion (Fig. 8 a). Mesosomaweaklyconvexinlateralview (Fig. 8 b). Indorsalview, lateral margins of pronotum with three denticles, anterior two acute, posterior broadandsometimesbifid, almostformingfourthdenticle; promesonotal grooveabsent (Fig. 8 c). Mesonotumwithapairofshortdenticles. Propodeal grooveimpressed only laterally. Dorsal anddeclivous facesof propodeum continuous, not meeting in a distinct propodeal angle; lateral margins of propodeum with variable number of denticles (Fig. 8 c). Femora not angulated dorsally, mid and hind basitarsi not flattened, with subparallel dorsal and ventral faces. Indorsalview, anteriormarginof petioleconcave, laterallywithpairof spines (Fig. 8 c), dorsum with a pair of tiny denticles (Fig. 8 b), subpetiolar process broader anteriorly with narrow translucent lamellae (Fig. 9 b). Postpetiole wider and longer than petiole (Fig. 8 c), with a pair of spines curvedbackwards broader than spines of petiole, dorsum without carina ordenticles (Fig. 8 c), subpostpetiolarprocesspronouncedandcompressed anteroposteriorly (Fig. 8 b). Gaster suboval, deeply concave anteromedially, with broad anterior lamellar expansions, not extended posteriorly in a carina (Fig. 8 c). Soldiermeasurements (N = 10): HL 1.43 - 1.72; HW 1.15 - 1.58; EL 0.33 - 0.36; PW 1.18 - 1.48; WL 1.40 - 2.00; PTL 0.20 - 0.33; PTW 0.53 - 0.68; PPL 0.28 - 0.32; PPW 0.63 - 0.73; GL 1.72 - 2.06; HBL 0.36 - 0.42; HBW 0.10 - 0.12; TL 5.17 - 6.09; CI 77.1 - 98.2; OI 20.1 - 28.3; PI 34.4 - 50.8; HBI 23.8 - 31.6. Soldierdescription: Bodyblack; dorsumandlateralofhead, apices of femora andexternalface of tibiaeferruginous. Firstgastral tergite with an anterior and a posterior pair of yellowish spots, each one occupying less than one third of the firsttergite length; posterior margins of eachtergite and sternite yellowish (Fig. 9). Sculpturingasintheworkers, exceptbypropleurawithstriaenotfully occupying the propleura, often on the lower part of the surface (Fig. 9 b). Pilosity as in the workers, expect by gaster, which has appressed simple hairs (Fig. 9 b, c). Headlongerthanwide (CI 77.1 - 98.2). Mandibleswithastronglongitudinal lateral angle. Clypeusslightlyconcave without a pair of denticles (Fig. 9 a). Dorsumofheaddiscshapedandslightlyconcaveanteriorly. Frontalcarinae crenulateanteriorly. Antennalclubill-defined. Roof ofantennalscrobeswith a lateral carinae and a posterior denticle (Fig. 9 b). Lateroventral margins of head without carinae. Vertexal corners forming pointed projections separated of the dorsum cephalic disc (Fig. 9 a, b). In lateral view, pronotum ascending, with a transversal carina not raisedinacrest; pronotalcarinaweaklydeveloped, notcrenulate, medially interrupted (Fig. 9 c). In dorsal view, anterior margin of pronotum gently rounded, lateralmargins subparallel, with a pair of anterior denticles. Mesonotumandpropodeumdiscontinuousandflat (Fig. 9 b); mesonotum withapairofbluntroundedprojections; propodealgroovewellimpressed; dorsal and declivous faces of propodeum meeting in a distinct propodeal angle (Fig. 9 b); indorsalview, lateralmarginsof propodeumwithtwo pairs of projections, the anterior obtuse, the posterior acute, short, not curved anteriorly (Fig. 9 b). Legs as in the worker. In dorsal view, anterior margin of petiole slightly concave, laterally with a pair of spines curved backwards (Fig. 9 c), dorsum with a pair of denticles (Fig. 9 b), subpetiolar process broader anteriorly with an acute projection. Postpetiole wider andlonger than petiole (Fig. 9 c), with a pair of spines broader than spines of petiole and curved anteriorly, dorsum of postpetiole with a transversal elevation (Fig. 9 c), subpostpetiolar process pronounced and compressed anteroposteriorly (Fig. 9 b). Gaster elongate, protruding anteriorly, without anterior lamellae or carinae (Fig. 9 c). Gyne measurements (N = 2): HL 1.32 - 1.40; HW 1.28 - 1.38; EL 0.32 - 0.35; PW 1.38 - 1.43; WL 1.85 - 1.88; PTL 0.33 - 0.34; PTW 0.48 - 0.50; PPL 0.35 - 0.38; PPW 0.63; GL 2.31 - 2.40; HBL 0.46 - 0.50; HBW 0.12; TL 6.20 - 6.36; CI 96.6 - 98.2; OI 25.1 - 25.4; PI 65 - 7.58; HBI 24.0 - 26.1. Gyne description: Body black; head predominantly black, with the edges of dorsum and part of lateral face yellowish. Apices of femora, tarsi and external face of tibiae yellowish. First gastral tergite with an anterior and a posterior pair of yellowish spots, each one occupying about one fifth of the first tergite length; posterior margins of each tergite and sternite yellowish (Fig. 10). Sculpturing and pilosity as in the soldiers, except by some erect hairs on mesoscutum and first tergite of gaster. Head as in the soldier (CI 96.6 - 98.2), but with frontal carinae convergingposteriorly (Fig. 10 a). Indorsalview, anteriormarginofpronotumslightlyrounded, narrower than in the soldiers, lateral margins with a pair of denticles pointed forwards, pronotal carina weakly developed (Fig. 10 b). Dorsally, mesoscutumsubtriangular, anteriormargin rounded; notauli absent; parapsidiallinesfeeblyvisibleand parallel; transscutal line weakly impressed, reaching the lateral margins of mesosoma; scutoscutellar groove weakly impressed, arched; scutellum well delimited and broader anteriorly; axillae rounded posteriorly (Fig. 10 c). Laterally, mesopleural groove dividing anepisternum and katepisternum; metapleura divided in upper metapleura and lower metapleura by a groove; metapleuropropodeal groove not impressed (Fig. 10 b). In dorsal view, posterior margin of propodeum concave, with a pair of denticles (Fig. 10 b). Wings as in genus description. Petiole and postpetiole as in the soldier, but longer (Fig. 10 b, c). Gaster elongated, protruding anteriorly, without anterior lamellae or carinae (Fig. 10 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	discussion	Comments: This species differs from C. marycorn new species and C. monicaulyssea new species by the anterior portion of the first tergite of gaster, near of the postpetiolar insertion, not striate, while the two first species present short striae in this portion. Also, the first gastral tergite in C. gabicamacho is covered by appressed canaliculate hairs anteriorly and posteriorly, with a narrow central portion with appressed simple hairs, while C. marycorn newspecies has appressed canaliculatehairs only laterally onanterior portion and posterioredges of tergite, the central portion has sparse appressed simple hairs, and C. monicaulyssea new species has abundant appressed canaliculate hairs evenly distributedover the tergite. Cephalotes gabicamacho is the species C. sp. 3 in the molecular phylogeniesof Price et al. (2014, 2016), where it is placed as sister to the clade containing all other members of the angustus, fiebrigi and prodigious groups and thus contributes to the paraphyly of theangustus group (Fig. 54). Natural history: This specieswas collected in two cities of state of Minas Gerais, both in the core region of the Brazilian biome Cerrado, a savanna physiognomy. In Paraopeba, asingle worker was collected in an arboreal pitfall trap, in a Cerrado area, at 750 m a. s. l. In Uberlândia, the specimens werecollected in twopoints at the Private Natural Heritage Reserve of Clube Caça e Pesca Itororó. This reserve is a 640 ha area, with Cerrado sensu strictu as predominant vegetation. This Cerrado physiognomy is characterized by tree canopies approximately 3 - 6 m high, annual average temperature between 18 and 24 ° C, and altitude 850 m a. s. l. Annual average precipitation is 1,700 mm, with two welldefined seasons, rainy summer (October to March) and dry winter (April to September) (Pinese et al., 2015). The coloniesfrom the Reserve of Clube Caça e Pesca Itororó were discovered in April 2005 (paratype C 05 - 134) and July 2006 (holotype C 06 - 94), respectively, as part of a larger survey of Cephalotes diversity at the reserve (see Powell, 2008, 2016). In both cases, discovery occurred during a transect baiting approach, designed to maximize the discovery of Cephalotes forager recruitment (See Powell, 2008 for explanation). For each colony, all occupied nests were located using dense baiting within the home tree and visually tracking recruits back to their nest (following methods of Powell, 2009). Collections were made by sealing each nest in the early morning, when foragers are not active, and then removing the intact branches for dissection in the laboratory (following methods of Powell, 2009). The colonies occupiedsix and eleven nests, respectively. The two colonies occupied different tree species, indicating no tree species preference. Nevertheless, nest entrance size was limited to sizes that closely fit a single soldier head (see data for C. sp 3 in Powell, 2016), characteristic of species with adisc-headed soldier morphology (Powell etal., 2020).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	distribution	Distribution: Minas Gerais, Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFFEFFCCFCB3FCB9D130FD98.taxon	etymology	Etymology: This speciesis named, inapposition, after the Brazilian myrmecologist Gabriela Procópio Camacho, for herimportant contribution to our current knowledge on Hymenoptera evolution, especially Ectatommine ants. Gabriela is a close friend of the three authors and her work and career trajectory have provided personal inspiration, as well as inspiring many other female myrmecologists.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	description	Figs. 11 a-c, 16	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	materials_examined	Holotype: BRAZIL: MG, Manga, Parque Estadual da Mata Seca, ix. 2011, - 14.84833 - 43.96555, R. Garro & R. Antoniazzi cols. Dossel 15.5 m, in Myracrodruon urundeuva, DZUP 550164 (worker) [DZUP]. Paratypes: samedataas holotype, - 14.84833 - 43.96694, dossel 14 m, DZUP 550169 (1 worker) [DZUP]; - 14.84833 - 43.98805, dossel 10.3 m, DZUP 550165 (1 worker) [DZUP]; - 14.84833 - 43.98861, dossel 10.3 m, DZUP 550168 (1 worker) [MZSP]; - 14.84833 - 43.987777, dossel 23 m, DZUP 550167 (1 worker) [DZUP]; - 14.848333 - 43.965555, dossel 11.8, in Handroanthus chrysotrichus, DZUP 550166 (1 worker) [DZUP]; - 14.848333 - 43.98805, dossel 15.8, in Handroanthus chrysotrichus dossel 15.8 m, DZUP 550170 (1 worker) [USNM].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	diagnosis	Diagnosis: Amember of angustus species group. Workers with incomplete striae on propleura. First sternite of gaster laterally striate. Anteriorportion offirsttergite of gaster, near tothe postpetiolar insertion, withshort striaeand sparseappressedsimple hairs, the distancebetween each hair longerthan their length; appressed canaliculate hairs present only laterally on gastral anterior portion (Fig. 11). Worker measurements (N = 7): HL 0.98 - 1.08; HW 1.13 - 1.33; EL 0.28 - 0.32; PW 0.92 - 1.10; WL 0.90 - 1.16; PTL 0.18 - 0.21; PTW 0.50 - 0.57; PPL 021 - 0.25; PPW 0.53 - 0.60; GL 1.35 - 1.68; HBL 0.34 - 0.40; HBW 0.10; TL 4.10 - 4.34; CI 111 - 123; OI 23.8 - 25.5; PI 17.6 - 19.0; HBI 24.0 - 28.0. Worker description: Body black; mandibles, frontal lobes, apices of femora, dorsal face of tibiae and tarsi yellowish tobrownish (Fig. 11). Mandibles, legs, gasteranddeclivousface of propodeummicroalveolate. Head, mesosoma, petioleandpostpetiolefoveate-microalveolate. Propleura striate-microalveolate, striae not fullyoccupying thepropleura. First tergite of gaster microalveolate with some anterior striae near of the postpetiole insertion; first sternite medially smooth and shiny, laterally striate-microalveolate. Body with appressed canaliculate hairs (Fig. 11 c). Mandibles and anterior margin of clypeus with suberect clavate and simple hairs (Fig. 11 a). Declivous face of propodeum glabrous. First tergite of gaster with sparse appressed simple hairs, first sternite with short erect simple hairs (Fig. 11 b). Some erect hairs present on the posterior edge of the gastral tergites. Head widerthan long (CI 111 - 123), dorsum slight convex (Fig. 11 b). Mandibles with a weakly developed lateral angle. Anterior margin of clypeus concave without a pair of denticles. Frontal carinae sinuous anterior tothe eyes, not bent dorsally over the eyes (Fig. 11 c). Antennae with a three-segmented club. Lateroventral margins of head with posterior carinae extending beyond the eyes until vertexal corners. Vertexal corners with anarrow, irregular lamellar expansion (Fig. 11 a). Mesosoma convex in lateral view (Fig. 11 b). Indorsal view, lateral margins of pronotum with three denticles, the anterior two acute, the posterior broad and sometimes bifid, almost forming a fourth denticle; promesonotalgroove absent (Fig. 11 c). Mesonotum with a pair of short denticles. Propodeal groove impressed only laterally. Dorsal and declivous faces of propodeum continuous, not meeting in a distinct propodeal angle; lateral margins of propodeum with variable number of denticles (Fig. 11 c). Femora notangulated dorsally, midand hind basitarsi not flattened, with subparallel dorsal and ventral faces. In dorsal view, anterior margin of petiole concave, laterally with a pair of spines (Fig. 11 c), petiolar dorsum with a pair of tiny denticles (Fig. 11 b), subpetiolar process broader anteriorly (Fig. 11 b). Postpetiole wider and longer than petiole (Fig. 11 c), with a pair of spines curved backwards, broader than the spines of petiole. Dorsum of postpetiole without carinaeordenticles (Fig. 11 c), subpostpetiolarprocesspronounced and compressed anteroposteriorly (Fig. 11 b). Gaster suboval, deeply concave anteromedially, with broad anterior lamellar expansions, not extending posteriorly in a carina (Fig. 11 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	discussion	Comments: This species differs from C. gabicamacho and C. monicaulyssea newspecies by the anterior portionof thefirst tergite of gaster, near the postpetiolar insertion, which presents short striae andsparse appressed simplehairs, while inC. gabicamacho this portion is not striate and covered by abundant, appressed canaliculate hairs, and in C. monicaulyssea new species this portion has short striae, but the hairs are canaliculate and evenly distributed. Natural history: The only series of this species known so far was collected at the Parque Estadual da Mata Seca, an area of successional stageswherehadnoanthropogenic interventionforat least 60 years. There are trees exceeding 20 m inheight andfewer newtrees and lianas when compared with other adjacent areas at different levels of regeneration. That area is a transition zone between three Brazilian biomes: Cerrado and Caatinga (Brazilian savanna), and Mata Atlântica (Brazilian Atlantic Forest). That locality is at almost 500 m a. s. l., with average annual temperature of 25 ºC, and average annual precipitation of 818 mm, the rainiest months are November and April (Antoniazzi et al., 2019). Workers were sampled in arboreal pitfalls which contained water and soap, on the canopy of twotree species, between 10 and 23 meters. Handroanthus chrysotrichus (Mart. Ex DC.) (Bignoniaceae), known as Golden trumpet tree (in Brazil, ipê), is found in open formation of Atlantic Forest, in dry forests, on topof hills, disturbed areas, associated with sandy soils, and is widely used as ornamental tree in urban areas (Bittencourt Junior and Moraes, 2010). This species presents extrafloral nectaries located on leaves (Gonzalez, 2013), which is highly attractive for ants, and Cephalotes genus are often sampled on these trees. Myracrodruon urundeuva Allemão (Anacardiaceae), known as Aroeira, has an ethnobotany role, as an important species for communities in northeastern Brazil, because of its use for medicinal, construction, fuel and forage purposes (Barros et al., 2016). However, precisely because of its importance for the community, this species was evaluated as one of the ten most priority for conservation, in comparation with almost 150 species with medicinal use endemic from Brazil (Campos and Albuquerque, 2020).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	distribution	Distribution: Minas Gerais, Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF8FFCDFCB3FDE5D103FBA1.taxon	etymology	Etymology: This speciesis named, in apposition, after Mary Lynne Corn, an early pioneer in the study of Cephalotes ants. Her dissertation on Cephalotes atratus from 1976 was groundbreaking in its detailed examination ofCephalotes biology. Her work representsas anessential contribution to our knowledge of this remarkable group of ants and stands as an inspiration to all students of Cephalotes biology.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	description	urn: lsid: zoobank. org: act: 2 D 8 A 0 B 94 - E 099 - 4 E 2 E- 9 BBB-A 6 F 8 B 34408 DF Figs. 12 a-c, 13 a-c, 14 a-c, 16	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	materials_examined	Holotype: BRAZIL: MG, Santana do Riacho, Serra do Cipó, 1267 m, - 19.294090 - 43.587950, 25. vii. 2018, S. Powell col., Campo rupestre (Vellozia) / C 18 - 32, DZUP 550180 (worker) [DZUP]. Paratypes: samedataas holotype: DZUP 550181 (2 workers, 1 soldier), DZUP 550182 (1 worker, 1 soldier, 1 gyne) [DZUP]; - 19.29370 - 43.588160, 13. viii. 2017, DZUP 550186 (1 worker, 1 soldier) [MZSP]; 1258 m, - 19.294040 - 43.587760, 17. vii. 2019, S. Powellcol. Humiria balsamifera / C 19 - 27, DZUP 550183 (1 worker, 1 gyne), DZUP 550184 (1 worker) [DZUP], DZUP 550185 (1 worker, 1 gyne) [USNM].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	diagnosis	Diagnosis: Amember of angustus species group. Workers with incomplete striae onthe propleura. First gastral tergite withshort striae near of postpetiole insertion, and with evenly distributed appressed canaliculate hairs, without simple hairs (Fig. 12 c). In soldiers, in dorsal view, lateral margins of pronotum convex and converging posteriorly (Fig. 13 c). Propodeal groove weakly impressed, marked on the sides but absent medially (Fig. 13 c). First sternite of the gaster with striae laterally (Fig. 5 n). Worker measurements (N = 8): HL 0.85 - 1.20; HW 1.00 - 1.38; EL 0.30 - 0.38; PW 0.79 - 1.25; WL 0.93 - 1.35; PTL 0.17 - 0.24; PTW 0.46 - 0.60; PPL 0.17 - 0.28; PPW 0.49 - 0.66; GL 1.05 - 1.76; HBL 0.28 - 0.41; HBW 0.10 - 0.11; TL 3.43 - 4.77; CI 105 - 117; OI 25.2 - 36.0; PI 37.9 - 43.3; HBI 26.3 - 34.6. Worker description: Body black; frontal lobe, vertexal lamellar expansions, apices of femora, tarsi, dorsal face of tibiae, and gastral lamellar expansions yellowish (Fig. 12). Head, mesosoma, legs, petioleandpostpetiolefoveate-microalveolate. Propleura striate-microalveolate, striae not fully covering propleura. Propodeal declivitywith some striae. Firsttergiteof gastermicroalveolate, with some short striae in the anterior portion, near to the postpetiolar insertion; first sternite medially smooth and shiny, laterally striatemicroalveolate (Fig. 12 b, c). Bodywith appressedcanaliculate hairs (Fig. 12 c), moreconcentrated on meso-, metapleura and anterior lamellar expansions of the gaster. Mandibles andanterior margin of clypeus witherect clavate and simple hairs (Fig. 12 a). Declivous face of propodeumglabrous. First sternite of gaster with long erect simple hairs (Fig. 12 b). Head wider than long (CI 105 - 117), dorsum slightlyconvex (Fig. 12 b). Mandibles with a weakly developed lateral angle. Anterior margin of clypeusconcavewitha pair of denticles. Frontalcarinae sinuous tonotched anteriorly the eyes, in frontal view (Fig. 12 a), gently bent dorsally over the eyes, in posterior view (Fig. 12 c). Antennae with three-segmented club. Lateroventral margins of head with posterior carinae extending posteriorly to vertexal corners, in lateral view. Vertexal corners with a narrow irregular lamellar expansion (Fig. 12 a). Mesosoma convex in lateral view (Fig. 12 b). In dorsal view, lateral margins of pronotum with variable number of denticles, often three, the posterior one bifurcate or trifurcate; promesonotal groove absent (Fig. 12 c). Mesonotumwithapairofshortdenticles. Propodealgroove impressed only laterally. Dorsal and declivous faces of propodeum continuous, notmeetingin adistinct propodealangle; lateral marginsof propodeum witha pair of anterior denticles and rowof minordenticles posteriorly near petiolar insertion (Fig. 12 c). Femora not angulated dorsally, mid and hind basitarsi not flattened, with subparallel dorsal and ventral faces. Indorsal view, anterior marginof petiole concave; lateralmargin with posteriorly curved spine, dorsum with a pair of denticles, subpetiolar process in lateral view with an anterior angle (Fig. 12 b). Postpetiole wider and longer than petiole (Fig. 12 c), with lateral spines broader than petiolar spines and curved backwards, dorsum without carinae or denticles, subpostpetiolar process in lateral view pronounced and compressed anteroposteriorly (Fig. 12 b). Gaster suboval, deeplyconcave anteromedially withbroad anterior anterolateral lamellar expansions, not extendingposteriorly as acarina (Fig. 12 c). Soldiermeasurements (N = 3): HL 1.80 - 2.00; HW 2.00 - 2.16; EL 0.40 - 0.43; PW 1.88 - 2.03; WL 1.73 - 2.00; PTL 0.30 - 0.35; PTW 0.73 - 0.80; PPL 0.33 - 0.38; PPW 0.80 - 0.88; GL 2.30 - 2.55; HBL 0.44 - 0.48; HBW 0.14 - 0.20; TL 5.76 - 7.28; CI 109 - 111; OI 19.7 - 20.0; PI 41.4 - 43.7; HBI 32.0 - 43.4. Soldier description: Body mostly black; anterolateral faceand edges of the dorsum of headferruginous, only central portion of the cephalic disc black. Lateral margins of pronotum with ferruginous spots. Apices of femora, tarsi and dorsal face of tibiae ferruginous. First gastral tergite with an anterior and a posterior pair of weakly yellowish spots, each one occupying less than one fourth of the first tergite length (Fig. 13). Sculpturing and pilosity as in workers, except by the presence of appressed simple hairson the first tergite of gaster, andthe erect short hairs on the posterior edge of the tergites and sternites (Fig. 13 b). Head slightlywider thanlonger (CI 106 - 111). Mandibles with strong longitudinal lateral angle. Clypeus slightlyconcave with apair of denticles (Fig. 13 a). Dorsum of head disc shaped and concave anteriorly. Frontal carinae crenulate anteriorly. Antennalclub ill-defined. Roof of antennal scrobe with lateralcarinae and posterior denticle. Lateroventral margin of head without carina. Vertexal corners forming pointed projections separated of the dorsum cephalic disc (Fig. 13 a, b). In lateral view, pronotum ascending, with a transversal carina not raised in a crest; pronotal carinae weakly developed, not crenulate (Fig. 13 c). In dorsal view, anteriormargin of pronotum gently rounded, lateral margins slightly concave and converging posteriorly, with anterolateral denticle. Mesonotum and propodeum continuous andflat (Fig. 13 b); mesonotum with blunt rounded posterolateral projection; propodeal grooveimpressed only laterally; dorsal and declivous faces of propodeum meeting in a distinct propodeal angle; in dorsal view, lateral margin of propodeum with three projections, the anterior one obtuse, the median one broad and acute, the posterior one acute and long, with the apices curved anteriorly. Legs as in worker. Petiole and postpetiole as in worker, except for the presence of a transverse elevation of dorsum of postpetiole. Gaster elongated, with narrow anterior lamellae, not extending posteriorly as a carina (Fig. 13 c). Gyne measurements (N = 3): HL 1.88 - 2.16; HW 1.88 - 1.96; EL 0.43 - 0.46; PW 1.80 - 1.88; WL 2.52 - 2.60; PTL 0.32 - 0.44; PTW 0.71 - 0.78; PPL 0.45 - 0.51; PPW 0.86 - 0.93; GL 0.33 - 0.36; HBL 0.56 - 0.58; HBW 0.15 - 0.17; TL 5.59 - 5.97; CI 90.7 - 101; OI 22.4 - 24.5; PI 44.1 - 61.6; HBI 26.1 - 31.2. Gyne description: Color, sculpturing and pilosity as in soldiers, except for some erecthairs on mesoscutum (Fig. 14). Head as in the soldier (CI 90.7 - 101), but longer (Fig. 14 a). Indorsalview, anteriormarginofpronotumslightlyrounded, narrower than in the soldiers, lateral margin with a pair of denticles pointed forward, pronotal carina weakly developed (Fig. 14 c). Dorsally, mesoscutum subtriangular, anteriormarginrounded; notauli absent; parapsidial line welldeveloped and parallel; transscutal line impressed, reaching lateral margin of mesosoma; scutoscutellar groove deeply impressed, arched, and scrobiculate; scutellum well delimited, broadest anteriorly; axillaeroundedposteriorly (Fig. 14 c). Laterally, mesopleural groove dividing anepisternum and katepisternum; metapleura divided in upper metapleura and lower metapleura by groove; metapleuropropodeal groove notimpressed (Fig. 14 b). In dorsal view, posterior margin of propodeum slightly concave with a pair of short denticles (Fig. 14 b). Wings unknown. Petiole and postpetiole as in the soldier, but longer (Figs. 14 b, c). Gaster elongated, anteriorly with narrow carinae, not extending posteriorly (Fig. 14 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	discussion	Comments: This species is very similar to C. angustus, differing by the presence of striae on the first sternite of gaster and the gastral anterior lamellae not extending posteriorly as a carina. Cephalotes monicaulyssea differs from new species C. marycorn and C. gabicamacho by the first tergite of gaster with abundant appressed canaliculate hairs evenly distributed. While C. gabicamacho has appressed canaliculate hairs anteriorly and posteriorly, with a narrow central portion with appressed simple hairs, and C. marycorn has appressed canaliculate hairs only laterally of anterior portion and posterior edge of tergite, the central portion has sparse appressed simple hairs. Natural history: This species was collected at the same locality in three different years (2017, 2018 and 2019), at Serra do Cipó, the southern part of the mountainous system of Serra do Espinhaço, Minas Gerais state. The local climate is well-defined, with fresh and rainy summers and dry season, the annual average temperature is 18 ° C, and the annual average precipitation is 1,600 mm. The dominant vegetation of the region is the Cerrado, a Neotropical savanna (Callisto et al., 2001). The specimens were only collected in altitudes above 1200 m, in a phytophysiognomy know as Campo Rupestre, despite targeted Cephalotes sampling at nearby locations at 1000 m and 800 m. Campo Rupestre is characterized by occurring exclusively on top of mountains, above 900 m, with rocky outcrops, shallow soils, and vegetation herbaceous-shrubs (Vasconcelos, 2011). It has been inferred that 30 % of the taxa occurring in campos rupestres are restricted to this formation (Lousada et al., 2011). The larger colony collection (C 18 - 32) was taken from four nests in the thicker basal stems of a Vellozia Vand. plant, a shrub typical of Campo Rupestres, occurring between 1000 and 2000 m a. s. l. (Lousada et al., 2011). This colony was discovered initially with baiting, and subsequent baiting across Vellozia individuals showed that Cephalotes monicaulyssea occupied most plants of this species within the vegetation patch. Viable populations of Cephalotes monicaulyssea may then be highly dependent on Vellozia at this altitude. Nevertheless, the second collection, a small incipient colony (C 19 - 27), was made inadvertently when opening dead stems of a Humiria balsamifera Aubl. (Humiriaceae) tree within the same vegetation patch as C 18 - 32. Additionally, a large reproductively mature colony of Cephalotes monicaulyssea was also collected from dead stems of a Vochysia thyrsoidea Pohl. (Vochysiaceae) tree in the same patch. This indicates that despite the high density of this Cephalotes monicaulyssea in Vellozia plants, it is not indicative of an obligate ant-plant relationship. Nevertheless, the discovery of this high-density population at 1200 m a. s. l. and not at lower elevations does suggest that C. monicaulyssea may be restricted to high altitude in Campos Rupestres.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	distribution	Distribution: Minas Gerais, Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFF9FFCFFCADFBDED7BDF8FA.taxon	etymology	Etymology: This species is named, in apposition, after Mônica Antunes Ulysséa, a Brazilian myrmecologist, poetess, activist, and agroecologist, for her contribution to the knowledge of Neotropical ants, especially the taxonomy of Myrmicinae. Besides being a great friend, Mônica’s protagonism in academic feminism has been an inspiration for contemporary and future generations of female myrmecologists.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE5FFD2FFDFF89FD197FAEB.taxon	description	(Figs. 17, 18)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE5FFD2FFDFF89FD197FAEB.taxon	materials_examined	The atratus group was proposed for the first time by De Andrade and Baroni Urbani (1999). Before that, the species C. alfaroi, C. atratus and C. serraticeps belonged tothe genus Cephalotes, while the species C. oculatus, C. opacusandC. placidus belongedto thegenusEucryptocerus. The species C. alfaroi was not included in this study since it does not occur in Brazil. In the morphological phylogeny by De Andrade and Baroni Urbani (1999, see their Fig. 24), the Central American hamulus group is the sister group of all other groups in Cephalotes, followed by theatratus group. In the recent molecular phylogenies (Price et al., 2014, 2016, see their Fig. S 3) this relationship isinverse, and theatratus group is the sister-mostgroup in the topology (Fig. 54). Theatratus and hamulus groups share the absenceof the soldier caste in some species of atratus and all species of hamulus. The species C. alfaroi, C. opacus, and C. serraticeps have known soldiers, while the species C. atratus, C. oculatus, and C. placidus do not have soldiers, as far as we know. Of these, large colony series ofC. atratus have revealed that while this species certainly lacks morphologically differentiated soldiers and any pronounced allometric morphological scaling, the worker caste is highly variable insize within mature colonies (Corn, 1980). This species is broadly distributed, and frequently collected (Fig. 18), and is the second most common species of the genus in Brazil, after C. pusillus. The other species without soldiers are not highly variable in size, and their distribution is more restricted (Amazon / Atlantic Forest (Fig. 18 )).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE5FFD2FFDFF89FD197FAEB.taxon	diagnosis	Diagnosis: In workers and soldiers vertexal corners of head, in lateral view, with a pair of spines (Fig. 2 a). Pronotum always with a pair of long dorsolateral spines; a pair of short median spines can be present in some species, sometimes weakly developed. Postoccipital carinae with ventral expansions (Fig. 17 a-b).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE5FFD2FFDFF89FD197FAEB.taxon	discussion	Brazilian species of atratus group Cephalotes atratus (Linnaeus, 1758) Obsolete combinations: Cryptocerus atratus, Formica atrata = Cephalotes atratus crassispina Santschi, 1920 = Cephalotes atratus erectus Kempf, 1951 = Cephalotes atratus nitidiventris Santschi, 1920 = Cephalotes atratus quadridens (De Geer, 1773) Obsolete combinations: Cephalotes quadridens, Formica quadridens = Cephalotes atratus rufiventris (Emery, 1894) Obsolete combination: Cryptocerus atratus rufiventris = Cephalotes dubitatus (Smith, 1853) Obsolete combination: Cryptocerus dubitatus = Cephalotes marginatus (Fabricius, 1804) new synonym Obsolete combination: Cryptocerus marginatus Cephalotes oculatus (Spinola, 1851) Obsolete combination: Cryptocerus oculatus = Cephalotes aethiops (Smith, 1853) Obsolete combination: Cryptocerus aethiops Cephalotes opacus Santschi, 1920 = Cephalotes abdominalis Santschi, 1929 Cephalotes placidus (Smith, 1860) Obsolete combination: Cryptocerus placidus = Cephalotes fenestralis (Smith, 1876) Obsoletecombination: Cryptocerus fenestralis. Cephalotes serraticeps (Smith, 1858) Obsolete combination: Cryptocerus serraticeps Note: The synapomorphiesso far accepted for C. marginatus include abundant suberect pilosity onthefirst gastral sternite, instead of sparse as in C. atratus, and median pronotal spines at least with ¼ of the size of the dorsolateral ones, instead of minute or absent as in C. atratus (De Andradeand Baroni Urbani, 1999). Cephalotes marginatus was described by Fabricius (1804), synonymized under C. atratus by Klug (1824), and then revived by De Andrade and Baroni Urbani (1999). In the latter study, the species C. decemspinosus Santschi, 1920 was synonymizedunder C. marginatus. Kempf (1951) has already argued that the status of this species should be changed, as there were individuals in series of C. atratus with the same characteristics of C. decemspinosus. After a careful examination of both species based on specimens from a wide range of distribution, we found great variation of the putative synapomorphic characters of C. marginatus among samples of C. atratus. Cephalotes marginatus is known only for the Amazonian arch in South America, while C. atratus is widely distributed from Mexico to north of Argentina and isone of the most commonly sampled species of Cephalotes. Therefore, consideringmorphological and geographic evidence, we here propose the synonymy of C. marginatus under C. atratus.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE5FFD2FFDFF89FD197FAEB.taxon	description	(Figs. 17 a-g)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	description	(Figs. 19, 20, 21) The basalis group of De Andradeand Baroni Urbani (1999) includes the complanatus group of Paracryptoceus by Kempf (1951) (C. complanatus, C. cordiae, C. ramiphilus), and two isolated species, C. basalis and C. manni. Additionally, there are five other species that do not occur in Brazil (C. brevispineus, C. cordiventris, C. femoralis, C. inca and C. mompox). Cephalotes manni is morphologically very distinct and can be easilyseparated from all other species in the group, by the combination of dentiform lamellar expansions on pronotum, declivous face of propodeum and anterior face of petiole concave, apices of anterior lamellar expansion of gaster near to postpetiole insertion, and median dorsal projection on hind femora absent. Thus, we transfer C. manni to the monotypicmanni species group, proposed here as discussed in its respective section.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	discussion	According to the molecular phylogeny (Price et al., 2016, see their Fig. S 3), basalis group forms a clade with the exclusively North and Central Americanmultispinosus, wheeleri andtexanus species groups. All species of basalis group recorded for Brazil occur only in the North and Midwest regions (Fig. 21).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	diagnosis	Diagnosis: In workers and soldiers, margin of pronotum with lamellar expansions, without denticles or spines. Declivous face of propodeum and anterior face of petiole truncate (Fig. 19 c-d). Hind femora with a median projection dorsally (Fig. 2 j). In dorsal view, the anterior lamellarexpansions of the gaster with apices distally directed in relation to the postpetiole insertion. Brazilian species of basalis group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	discussion	Cephalotes basalis (Smith, 1876) Obsolete combination: Cryptocerus basalis = Cephalotes multispinus (Emery, 1890) Obsolete combinations: Cryptocerus cordatus multispinus, Cryptocerus multispinus Cephalotes complanatus (Guérin-Méneville, 1844) Obsolete combination: Cryptocerus complanatus = Cephalotes angulatus (Smith, 1858) Obsolete combination: Cryptocerus angulatus = Cephalotes multispinus amazonensis (Forel, 1911) Obsolete combination: Cryptocerusmultispinus amazonensis Cephalotes cordiae (Stitz, 1913) Obsolete combination: Cryptocerus cordiae Cephalotes ramiphilus (Forel, 1904) Obsolete combination: Cryptoceruscomplanatus ramiphilus	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	description	(Figs. 19 a-f)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE0FFD5FFFDFF0BD1B4FEF7.taxon	description	(Figs. 20 a-g)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE2FFD6FFFDFF0BD196FDFD.taxon	description	(Figs. 22, 23)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE2FFD6FFFDFF0BD196FDFD.taxon	discussion	The clypeatus groupwas proposed byDe Andrade and Baroni Urbani (1999) to include three species from the former genus Zacryptocerus. The authorsstatedthat the diagnostic charactersof this group (triangular vertexal angles, and pronotal and propodeal spines fused by lamella) could indeed be enough to keep them in a separated genus from Cephalotes. However, thiswould render Cephalotes paraphyletic since the phylogeny shows this group as a clade within the genus Cephalotes (Fig. 54). All species in this group are exclusively South American.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE2FFD6FFFDFF0BD196FDFD.taxon	diagnosis	Diagnosis: In workers andsoldiers, pronotal andpropodeal spines fused by lamellae (Fig. 2 o). Gaster surrounded by a lamella (Fig. 22). Brazilian species of clypeatus group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE2FFD6FFFDFF0BD196FDFD.taxon	discussion	Cephalotes clypeatus (Fabricius, 1804) Obsolete combination: Cryptocerus clypeatus Cephalotes membranaceus (Klug, 1824) Obsolete combination: Cryptocerus membranaceus = Cephalotes fervidus (Smith, 1876) Obsolete combination: Cryptocerus fervidus Cephalotes ustus (Kempf, 1973) Obsolete combination: Zacryptocerus ustus	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE2FFD6FFFDFF0BD196FDFD.taxon	description	(Figs. 22 a-c)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE3FFD7FFDFF893D1B4F822.taxon	description	(Figs. 25, 26)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE3FFD7FFDFF893D1B4F822.taxon	discussion	The depressus group is composed by the former pavonii group of Paracryptocerus byKempf (1951), andthespeciesC. betoi andC. palustris, described by De Andrade in De Andrade and Baroni Urbani (1999). In the recentmolecularphylogeny (Priceetal., 2016, seetheirFig. S 3) thisgroupis sistertotheclade formedbycoffeae, patei, emeryi andcrenaticeps groups. The majority of speciesin the depressus group occur in Brazil. Cephalotes cristatus is the only member that does not occur in Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE3FFD7FFDFF893D1B4F822.taxon	diagnosis	Diagnosis: In workers and soldiers, dorsal and declivous face of propodeumcontinuous, notmeetinginadistinct propodealangle, withtwo pairs of spines, the anterior one longer than the posterior one (Fig. 25 d-i). Brazilian species of depressus group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE3FFD7FFDFF893D1B4F822.taxon	discussion	Cephalotes betoi De Andrade, 1999 Cephalotes borgmeieri (Kempf, 1951) Obsolete combination: Paracryptocerus borgmeieri Cephalotes cordatus (Smith, 1853) Obsolete combination: Cryptocerus cordatus = Cephalotes cordatus boliviensis (Santschi, 1921) Obsolete combination: Cryptocerus cordatus boliviensis Cephalotes depressus (Klug, 1824) Obsolete combination: Cryptocerus depressus = Cephalotes depressus sorocabensis (Forel, 1912) Obsolete combination: Cryptocerus depressus sorocabensis Cephalotes eduarduli (Forel, 1921) Obsolete combination: Cryptocerus eduarduli Cephalotes palustris De Andrade, 1999 Cephalotes pavonii (Latreille, 1809) Obsolete combination: Cryptocerus pavonii	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE3FFD7FFDFF893D1B4F822.taxon	description	(Figs. 25 a-o)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	description	(Figs. 27, 28, 29, 30, 31, 32) Kempf (1958 a) created the jheringi subgroup of the angustus group in Paracryptocerus (Harnedia) with seven species (C. bivestitus, C. bohlsi, C. bruchi, C. fossithorax, C. jheringi, C. prodigious, andC. quadratus). Later, De Andrade and Baroni Urbani (1999) separated it into three groups of Cephalotes. The first is the monotypic bruchi group, characterized by cephalicdiscincompleteinsoldiersandgynes, asignificantcharacterintheir analysis. Thesecondistheexclusively Argentinianprodigiosus groupwith the species C. bivestitus and C. prodigiosus, characterized by the concave cephalic disc insoldiers. Finally, the fourremaining species of thejheringi subgroup were joined to the Kempf’s pilosus group, originally formed by C. fiebrigi, C. liogaster, andC. pilosus; resultinginthecurrentfiebrigi group, with the addition of C. guayaki, C. lanuginosus, and C. supercilii described by De Andrade in De Andrade and Baroni Urbani, 1999.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	discussion	Inthemorphological phylogeny (De Andradeand Baroni-Urbani, 1999, see their Fig. 24) the bruchi group was recovered as sister to the fiebrigi group. The authors argued that the incomplete cephalic disc could be a secondary loss in bruchi group, and the unknown ancestor of C. bruchi should have had both soldiers and gynes with a complete disc, but they keptthespeciesinaseparatedgroup. However, themolecularphylogenies (Fig. 54) recovered C. bruchi within the fiebrigi group, as sister to the groupingformed byC. jheringi, C. bohlsi andC. specularis. Therefore, based onmorphologicalandmolecularevidence, weheretransfer C. bruchi tothe fiebrigi group, extinguishingthe bruchi speciesgroup (Fig. 54).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	diagnosis	Diagnosis: In workers, dorsal and declivous faces of propodeum continuous, not meeting in a distinct propodeal angle, and converging posteriorly towards the petiolar insertion, the lateral marginswithout lamellar expansions (Fig. 2 n). Anterior portion of gaster with opaque expansion, never extending posteriorly in the form of translucent lateral lamellae (Fig. 2 q). In soldiers, propodeum with variable number of spines, if there are two pairs, the anterior one is never longer than the posterior. Anterior gastral expansions not translucent (Fig. 2 q). Brazilian species of fiebrigi group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	discussion	Cephalotes bruchi (Forel, 1912) Obsolete combination: Cryptocerus bruchi = Cephalotes jheringi pampaensis (Santschi, 1931) = Cephalotes ridiculus (Santschi, 1915) Obsolete combination: Cryptocerus ridiculus Cephalotes fiebrigi (Forel, 1906) Obsolete combination: Cryptocerus pilosus fiebrigi = Cephalotes guttifer (Santschi, 1919) Obsoletecombination: Cryptocerus (Paracryptocerus) guttifer Cephalotes guayaki De Andrade, 1999 Cephalotes jheringi (Emery, 1894) Obsolete combination: Cryptocerus jheringi = Cephalotes peltatus (Emery, 1896) Obsolete combination: Cryptocerus peltatus = Cephalotes peltatus ellenriederi (Forel, 1911) Obsolete combination: Cryptocerus peltatus ellenriederi Cephalotes liviaprado new species Cephalotes pilosus (Emery, 1896) Obsolete combination: Cryptocerus pilosus Cephalotes quadratus (Mayr, 1868) Obsolete combination: Cryptocerus quadratus = Cephalotes convexus (Santschi, 1916) Obsolete combination: Cryptocerus convexus Cephalotes specularis Brandão, Feitosa, Powell and Del-Claro, 2014	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	description	(Figs. 27 a-l)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEEFFDBFFFDFF0BD1B4FD52.taxon	description	(Figs. 28 a-l)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE8FFDDFFFDFF0BD035FBC6.taxon	materials_examined	Holotype: BRAZIL, MS, Porto Murtinho, 28. i. 2015 (dique, espinho P. ruscifolia), P. R. Souza, DZUP 550171 (worker) [DZUP]. Paratypes: samedataas holotype: DZUP 550173 (2 workers, 1 soldier), DZUP 550172 (1 worker, 1 soldier) [DZUP], DZUP 550174 (2 workers, 1 soldier) [MZSP], DZUP 550175 (1 worker, 1 soldier) [INPA].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE8FFDDFFFDFF0BD035FBC6.taxon	diagnosis	Diagnosis: Amember offiebrigi species group. Workers withlong flexuous hairs (Fig. 29 b). In frontal view, frontal carinae with a lateral projection anteriorly to the eyes (Fig. 27 d). In soldier, first gastraltergite, with erect hairs, few and sparse appressedhairs can be present (Fig. 30). Worker measurements (N = 7): HL 1.08 - 1.23; HW 1.18 - 1.33; EL 0.32 - 0.38; PW 1.10 - 1.25; WL 1.21 - 1.34; PTL 0.26 - 0.29; PTW 0.62 - 0.64; PPL 0.26 - 0.31; PPW 0.66 - 0.75; GL 1.50 - 1.80; HBL 0.41 - 0.48; HBW 0.13 - 0.14; TL 4.36 - 4.92; CI 104 - 109; OI 26.4 - 29.8; PI 37.5 - 44.3; HBI 27.2 - 30.8. Worker description: Body black; frontal lobes, mandibles, and apices of each segment of legs brownish to ferruginous (Fig. 29). Mandibles, dorsum of head, mesosoma, petiole and postpetiole foveate-rugose, space between foveae microalveolate (Fig. 29 a, c). Frontal lobes weakly striate (Fig. 29 a); ventral face of head and lateral of mesosoma areolate-rugose; rugosities of dorsum of mesosoma extending to the middle of propodeum declivous face, lower portion of propodeum microalveolate. Legs microalveolate, except by tibiae, which external face is areolate-rugose. Gaster microalveolate, the anterior half of first tergite with short longitudinal striae originating near the postpetiole insertion (Fig. 29 c); first sternite laterally with some irregular weak rugosities. Body with abundant flexuous hairs, and some sparse appressed canaliculate hairs. Anterior margin of clypeus with suberect canaliculate hairs (Fig. 29 a). Head slightly wider thanlong (CI 104 - 109), dorsum weakly convex (Fig. 29 b). Mandibles with a weakly developed lateral angle. Anterior margin of clypeus slightly concave, with a pair of lateral denticles. Frontal carinae notched anteriorly to the eyes, forming a lateral angle (Fig. 29 a). Antennalclubill-defined. Lateroventralmarginsofhead without carinae. Vertexal corners with irregular lamellar expansions (Fig. 29 a). Mesosoma stronglyconvex in lateral view (Fig. 29 b). In dorsal view, lateral margins of pronotum with three pairs of denticles, the anterior one acute and the two posterior obtuse; promesonotal groove absent (Fig. 29 c). Mesonotum with a pair of short blunt denticles. Propodeal grooveabsent. Dorsal and declivousfacesof propodeum continuous, not meeting in a distinct propodeal angle; lateral margins of propodeum with an anterior pair of short blunt denticles, followed by a pair of large and acuteand a rowof minor acutedenticlesnear petiolarinsertion, the number and degree of development of the denticlesvary even between sides of the samespecimen (Fig. 29 c). Femoranotangulated dorsally, mid andhindbasitarsi notflattened, with subparalleldorsalandventral faces. Indorsalview, petiolecompressedanteroposteriorly, indorsal view, anterior margin with a discrete median concavity, lateral spines curved backwards, dorsum with a pair of denticles (Fig. 29 b), subpetiolar process acute anteriorly (Fig. 29 b). Postpetiole slightly longer than petiole, without dorsal projections, lateral spines broad and curved backwards (Fig. 29 c), subpostpetiolar process pronounced and compressed anteroposteriorly (Fig. 29 b). Gaster oval, with a pair of well-developed thick opaque anterior expansions, not extending posteriorly formingalateral lamella (Fig. 29 c). Soldiermeasurements (N = 4): HL 1.96 - 2.09; HW 1.82 - 1.96; EL 0.40 - 0.43; PW 1.75 - 1.94; WL 1.66 - 1.88; PTL 0.32 - 0.38; PTW 0.85 - 0.89; PPL 0.33 - 0.39; PPW 0.88 - 0.90; GL 2.04 - 2.40; HBL 0.44 - 0.48; HBW 0.14 - 0.15; TL 6.31 - 7.13; CI 89.6 - 94.9; OI 21.3 - 21.2; PI 37.6 - 42.2; HBI 29.9 - 33.3. Soldier description: Body black; frontal lobes and apices of each segment of legs brownish to ferruginous (Fig. 30). Mandible, ventralfaceofhead (Fig. 30 a), promesonotumandmesonotum foveate, spacebetween foveaemicroalveolatetosmooth (Fig. 30 c). Dorsum of head with small foveae anteriorly, increasing in diameter posteriorly. Dorsal face of propodeum, petiole andpostpetiole foveate, without space betweenfoveae; declivousface of propodeum microalveolate, withsome striae on upper surface; lateral of mesosoma areolate-rugose (Fig. 30 b). Legs microalveolate, except tibiae which external face is areolate-rugose. Gastermicroalveolate, withafewshortweaklymarkedlongitudinalstriae, originating near the postpetiole insertion (Fig. 30 c). Body with abundant flexuous hairs (Fig. 30 b), except on dorsum of head, whichhastinysuberectsimplehairs (Fig. 30 a). Mandibles, lateraland ventralfaceofhead, meso- andmetapleurawithfewappressedcanaliculate hairs. Anterior margin of clypeus with suberect canaliculate hairs. Gaster with sparse appressed simple hairs (Fig. 30 b, c). Head longer than wide (CI 89.6 - 94.9). Mandibles with a strong longitudinallateralangle. Clypeuswithapairofdenticles (Fig. 30 a). Dorsum of head disc shaped, convex medially (Fig. 30 a). Frontal carinae crenulate convergingposteriorly. Antennalclubill-defined. Roofof antennal scrobes withlateralcarinaeand aposteriordenticle. Lateroventralmarginsofhead without carinae. Vertexal corners forming pointed projections separated of the dorsum cephalic disc (Fig. 30 a, b). In lateral view, pronotum ascending, pronotal carina weakly marked and crenulate (Fig. 30 b, c). In dorsal view, anterior margin of pronotum rounded, lateralmarginswithtwopairsof denticles, theanteriorlyoneacute, theposterior oneobtuse. Mesonotumandpropodeumcontinuousand flat (Fig. 30 b); mesonotumwithapairofbluntroundedprojections; propodeal groove impressed; dorsal and declivous faces of propodeum meeting in a distinct propodeal angle, in dorsal view, lateral margins of propodeum withapairofmedianobtusedenticlesand apairof well-developedspines, directed upwards (Fig. 30 c). Legs as in the worker. Petiole and postpetiole as in the worker. Gasterelongate, withtheanterolateralexpansionsprotrudinganteriorly (Fig. 30 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE8FFDDFFFDFF0BD035FBC6.taxon	discussion	Comments: Thisspeciesissimilar toC. pilosus butcanbe distinguished by the frontal carinae notched anterior to the eyes, forming a lateral angle and by the less abundant and shorter pilosity; in C. pilosus the frontal carinae is evenly straight to slightly depressed anterior to the eyes, but neverforming a lateral angle andthe pilosity is long anddense. Natural history: Cephalotes liviaprado was collected in the Chaco formation. This ecosystem is part of the diagonal zone of seasonallydry open areas in South America. The so called “ Gran Chaco ” occurs in northern Argentina, western Paraguay, southeastern Bolivia, and the extreme western of Mato Grosso do Sul state in Brazil, exclusively in the city Porto Murtinho (Prado, 1993). The vegetation is represented by shrub, deciduous, and spinous vegetation, usually associated with saline soils (Silvaet al., 2000). The Chaco extends from tropicallatitudes (18 ° S) tosubtropical zones (31 ° S), and theclimate is marked by strong seasonality, with more severe summers, and winter frosts (Werneck, 2011). The Brazilian Chaco (Chacosensu stricto) occupy a subregion of the Pantanalbiome, and extends over about 7 % of itsterritory (Silva et al., 2000). Despite its relatively small area in Brazil, this formation is considered of highpriority forconservation (Tálamo and Caziani, 2003). However, the anthropic use in this area for agricultureand logging has been a serious threat for the maintenance of this ecosystem and the species that live in it (Pott and Pott, 2003). Cephalotes liviaprado was collected in the thorns of the tree species Prosopis ruscifolia Griseb. (Fabacea: Mimosoideae), which is endemic to the Chaco (Fuster, 2012). This tree species is adapted to edaphic conditions, and marshy and salty environments, and is known as a pioneer and colonizer plant. It has large stem thorns (10 to 30 length and 2 cmdiameter) and extrafloral nectaries (Vilela and Palacios, 1997) which favors nesting by arboreal ants, like Cephalotes (Fuster, 2012). So far, C. liviaprado has been collected only in this location exclusively associated with P. ruscifolia. If these species areintimately relatedand endemic of Chaco, they must be extremely threatened.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE8FFDDFFFDFF0BD035FBC6.taxon	distribution	Distribution: Mato Grosso do Sul, Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFE8FFDDFFFDFF0BD035FBC6.taxon	etymology	Etymology: The specific epithet, in apposition, is in honor of Livia Pires do Prado, a Brazilian myrmecologist and passionate historian of science, for hercontributions to thetaxonomy of Brazilian ants and the rescue of extremely relevant names and facts involving the history of Brazilian entomologists of all generations. Her work and dedication to science stands as aninspiration to female myrmecologists and students of the history of science alike.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	description	(Figs. 33, 34, 35) The grandinosus groupwas first proposed by De Andrade and Baroni Urbani (1999) including five species: C. persimplex, described by De AndradeinDe Andradeand Baroni Urbani (1999), C. klugi, knownonly for the gyne, C. persimilis, C. grandinosus, and C. foliaceus. These last three composed the formerpinelii group by Kempf (1952), along with C. incertus, C. maculatus, C. pinelii, andC. scutulatus.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	discussion	Grandinosus and pinelii are sister groups in the morphological phylogeny (De Andrade and Baroni Urbani, 1999, see their Fig. 24), sharing many characters, as the strongly dorsoventrally flattened body, dorsum of mesosoma continuous, and lamellarexpansions on mesosoma, petiole, postpetioleand gaster. Both groups differ only by the presence of a lamella on hind femora and the lighter color in grandinosus. In the molecularphylogeny (Price et al., 2016, see their Fig. S 3), the grandinosus andpinelii groups arerecovered asparaphyletic. Cephalotes foliaceus (grandinosus group) is sister to C. sp. 2 (here described as C. mariadeandrade new species, of thepinelii group). The clade formed by the otherspecies of the grandinosus group (C. grandinosuş C. klugi, C. persimplex and C. persimilis) is the sister group of a clade formed by some of the previously designated species of the pinelii group (C. maculatus, C. liepini, C. nilpiei, C. pinelii and C. pileini) (Fig. 54). The morphologicalsimilarities and phylogeneticassociation between speciesof the pinelii andgrandinosus groupssuggest theylikely represent asingle evolutionarylineage (Fig. 54). Evenbeing paraphyletic, these groups are morphologically diagnosable, thus we kept it separate here, since we have not examined the species occurring outside Brazil in this study, what would help to better understand the relationships and limits between these species and redefine these groups.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	diagnosis	Diagnosis: In workers, body strongly flattened dorsoventrally. In dorsal view, dorsum of mesosoma continuous, with lamellar lateral expansions. In workers and soldiers, hindfemora with a ventral and / or dorsal lamellar crest, usually crenulate and narrow (Fig. 2 k, 3 k). Petiole and postpetiole with lamellar lateral expansions. Brazilian species of grandinosus group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	discussion	Cephalotes grandinosus (Smith, 1860) Obsolete combination: Cryptocerus grandinosus = Cephalotes grandinosus nevadensis (Forel, 1912) Obsolete combination: Cryptocerus grandinosus nevadensis Cephalotes klugi (Emery, 1894) Obsolete combination: Cryptocerus klugi Cephalotes persimilis De Andrade, 1999 Cephalotes persimplex De Andrade, 1999	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	description	(Figs. 34 a-f)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFEAFFDFFFFDFBB4D1B4FB2D.taxon	discussion	Note: Cephalotes klugi is known only for the gyne.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE0FFFDFBBCD0C7FBCB.taxon	description	(Fig. 36) The monotypicmanni species groupis here created to accommodate the species C. manni, formerly a member of the basalis species group. This species was described by Kempf (1951), as an “ isolated species ” without any defined group. However, De Andrade and Baroni Urbani (1999) included this species in the basalis group.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE0FFFDFBBCD0C7FBCB.taxon	discussion	Cephalotes manni differs from the species in the basalis group by many characters. All species in the basalis group have lamellar expansions on the margins of the pronotum, in dorsal view, while C. manni has three pairs of dentiform projections. The declivous face of propodeum and anterior face of petiole are flat in the species of the basalis group, and concave in C. manni. The species in thebasalis group have amedian dorsal projection on thehind femora, which is absentin C. manni. In dorsal view, the anterior lamellar expansionsof the gaster of species of basalis group have their apices distant to the postpetiole insertion, while in C. manni the apices of the lamellar expansions are near the postpetiolar insertion. In the morphological phylogeny, C. manni is the sister species of the rest of the basalis group (De Andrade and Baroni Urbani, 1999, see their Fig. 24), but this species was not included in the molecular phylogeny (Price et al., 2016), which could corroborate its phylogenetic position. In Brazil, C. manni occurs only in the North Region, in the Amazon Forest, like some species of basalis group, but not in sympatry with other members of this group.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE0FFFDFBBCD0C7FBCB.taxon	diagnosis	Diagnosis: In workers, in lateral view, eyes occupying morethan 1 / 3 of the length of the head. In dorsal view, margin of pronotum with three pairs of lamellar dentiform projections. Fore femora very increased in size relative to the mid and hind femora. Lateral margins of the declivous face with broad lamellar expansion extending to the posterior dorsal spines of propodeum towards to petiolar insertion (Fig. 2 m). Insoldiers, mesonotal spines curveddorsally. Posterior femora without projections, metatibiae never marginated. Brazilian species of manni group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE0FFFDFBBCD0C7FBCB.taxon	synonymic_list	Cephalotes manni (Kempf, 1951)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE2FCB3F8F0D196FE39.taxon	description	(Figs. 37, 38, 39) The pallens group was proposed by De Andrade and Baroni Urbani (1999) with 10 species. It includesC. jamaicensis, C. pallens, C. patellaris, C. porrasi, C. variansand thenew species describedby De AndradeinDe Andradeand Baroni Urbani (1999) (C. decolor, C. decoloratus, C. pallidoides, C. pallidus, and C. pellans). Of these species, only five occur in Brazil – C. pallens, C. pallidoides, C. pallidus, C. patellaris, andC. pellans.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE2FCB3F8F0D196FE39.taxon	description	In addition to the characters employed by De Andrade and Baroni Urbani (1999), here we provide an illustrated identification key for workers based on characters as the shape of frontal carinae in relation to eyes, and sculpture patterns on pleura. Regarding the identification key for soldiers, the most informative characters are the sculpture and pilosity patterns.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE2FCB3F8F0D196FE39.taxon	discussion	The pallens group is sister to the clade formed by the grandinosus and pinelii groups (Fig. 54), with all of these groups sharing a worker body shape that is strongly flattened dorsoventrally.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE2FCB3F8F0D196FE39.taxon	diagnosis	Diagnosis: In worker, body strongly flatteneddorsoventrally. Body colorreddish brown. In frontal view, vertexalcorner extending laterally overhangingtheeye (Fig. 2 d). In soldier, infrontalview, cephalicdorsum completelycovering the mandibles to form a “ dish ” head type (Fig. 38). Brazilian species of pallens group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD4FFE2FCB3F8F0D196FE39.taxon	discussion	Cephalotes pallens (Klug, 1824) Obsolete combinations: Cryptocerus pallens, Zacryptocerus pallens = Cephalotes araneolus (Smith, 1853) Obsolete combination: Cryptocerus araneolus = Cephalotes discocephalus Smith, 1853 Obsolete combination: Cryptocerus discocephalus Cephalotes pallidoides De Andrade, 1999 Cephalotes pallidus De Andrade, 1999 Cephalotes patellaris (Mayr, 1866) Obsolete combination: Cryptocerus patellaris Cephalotes pellans De Andrade, 1999	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD7FFE4FFDFFF0BD764FC64.taxon	description	(Figs. 40, 41, 42, 43, 44, 45, 46, 47, 48) Thepinelii group was first proposed byKempf (1952), including six species (C. foliaceus, C. grandinosus, C. incertus, C. maculatus, C. pinelii, and C. scutulatus). Later, De Andradeand Baroni Urbani (1999) separated it in two groups, grandinosus and pinelii. The species C. foliaceus, and C. grandinosus were included in the grandinosus group, and the other four species in thepinelii group, along with C. kukulcan andthree new species described by De Andrade in De Andrade and Baroni Urbani (C. liepini, C. pileini and C. nilpiei). In the morphological phylogeny (De Andrade and Baroni Urbani, 1999, see their Fig. 24), the grouppinelii isrecovered as monophyletic, with the species C. kukulcan, C. scutulatus and C. incertus forming a deeply nested grouping within the clade. In the molecular phylogeny (Priceet al., 2016, see their Fig. S 3), the relationshipsbetween C. kukulcan, C. scutulatus and C. incertus were supported, but they were recovered as sister group of the texanus and bimaculatus groups, not related with the other species of the pinelii group. The remaining species of the pinelii group were recovered as related to the grandinosus group (Fig. 54), rendering the pinelii group proposed by De Andrade and Baroni Urbani (1999) polyphyletic. Morphology and molecular data suggest that C. kukulcan, C. scutulatus and C. incertus might form an additional group within Cephalotes, but the first two species were not included in this study, since they not occur in Brazil. On the other hand, the species of pinelii that occur in Brazil and are closely related to the grandinosus group likely form a monophyletic group with it, as discussed under the grandinosus group section.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD7FFE4FFDFFF0BD764FC64.taxon	diagnosis	Diagnosis: In workers, body strongly flattened dorsoventrally. In dorsal view, dorsum of mesosoma continuous, with lamellar lateral expansions. In workers and soldiers, hind femora without crest, lamellae, or angles (Fig. 2 l, 3 l). Petiole and postpetiole with lamellar lateral expansions. Brazilian species of pinelii group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD7FFE4FFDFFF0BD764FC64.taxon	discussion	Cephalotes incertus (Emery, 1906) Obsolete combination: Cryptocerus incertus Cephalotes liepini De Andrade, 1999 Cephalotes maculatus (Smith, 1876) Obsolete combination: Cryptocerus maculatus = Cephalotes grandinosusmagdalenensis (Forel, 1899) Obsoletecombination: Cryptocerus grandinosusmagdalenensis = Cephalotes maculatus nanus (Forel, 1912) Obsolete combination: Cryptocerus maculatus nanus Cephalotes mariadeandrade new species Cephalotes nilpiei De Andrade, 1999 Cephalotes pinelii (Guérin-Méneville, 1844) Obsolete combination: Cryptocerus pinelii	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD7FFE4FFDFFF0BD764FC64.taxon	description	(Figs. 40 a-i)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD0FFE7FCB3FF0BD687FF63.taxon	materials_examined	Holotype: BRAZIL, MG, Uberlândia, - 19.033630 - 48.318400, 860 m, 03. viii. 2016. S. Powell et al. col. / C 16 - 64, DZUP 550137 (worker) [DZUP]. Paratypes: samedataas holotype: DZUP 550138 (1 worker, 1 soldier, 1 male), DZUP 550139 (1 worker, 1 soldier, 1 gyne), DZUP 550140 (1 worker, 1 male), DZUP 550141 (1 worker, 1 soldier, 1 gyne), DZUP 550142 (1 worker, 2 soldiers) [DZUP], DZUP 550145 (1 worker, 1 soldier, 1 gyne), DZUP 550146 (1 worker, 1 soldier, 1 male) [MZSP], DZUP 550147 (1 worker, 1 soldier) [MPEG], DZUP 550143 (1 worker, 1 male), DZUP 550144 (1 worker, 1 soldier, 1 gyne) [USNM].	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD0FFE7FCB3FF0BD687FF63.taxon	diagnosis	Diagnosis: Amember of pinelii speciesgroup. Workerswith lateral lamellar extensions of the propodeum on the declivous face but not on the dorsal face (Fig. 42 c). Soldier with cephalic disc covered with many tubercular elevations (Fig. 43 a-c). Worker measurements (N = 11): HL 0.78 - 0.84; HW 0.89 - 0.99; EL 0.25 - 0.28; PW 0.70 - 0.78; WL 0.84 - 0.89; PTL 0.13 - 0.19; PTW 0.48 - 0.53; PPL 0.14 - 0.19; PPW 0.51 - 0.62; GL 0.94 - 1.08; HBL 0.35 - 0.38; HBW 0.09; TL 2.89 - 3.16; CI 110 - 123; OI 26.3 - 29.9; PI 25.0 - 37.5; HBI 23.3 - 25.0. Worker description: Body dark brown; frontal lobes and lamellar expansions of pronotum, propodeum, petiole, postpetioleand gaster yellowish to translucent. Gaster darker than the rest of body (Fig. 42). Mandibles rugose-microalveolate. Dorsum of head and mesosoma foveate, space between foveae smooth; ventral face of head areolatemicroalveolate. Legs andgaster microalveolate. Body with appressed canaliculate hairs, more abundant on dorsum of pronotum (Fig. 42 c). Mandibles and anterior margin of clypeus with suberect clavate hairs. Declivous face of propodeum glabrous. Petiole and postpetiolewith hairs forming a transverse dorsal strip; anterior and posterior portions glabrous (Fig. 42 c). Gaster with sparse short appressed simple hairs, anterior lamellar expansions glabrous (Fig. 42 c). Headslightlywiderthanlong (CI 110 - 123), dorsumveryconvexmedially, with two longitudinal elevations posteriorly, and depressions in front of eyes. Mandibles witha weaklydeveloped lateral angle. Anterior margin of clypeusconcavewithoutdenticles. Frontal carinaestraitanteriorlyof eyes; pointedupwardsovereyes (Fig. 42 c). Antennaewithatwo-segmentedclub. Lateroventralmarginsofheadwithposteriorcarinaeextending beyondthe eyesuntilvertexalcorners. Vertexalcornerswithanarrowlamellarexpansion. Mesosomaalmostflatinlateralview, withagentlyascendantpronotum (Fig. 42 b). Indorsalview, lateralmarginsofpronotum withconcavelamellar expansions, broader anteriorly; promesonotalgroove absent, instead of it is an elevation on dorsum of pronotum (Fig. 42 c). Mesonotum with a pair of short denticles pointed upwards. Propodeal groove deeply impressed. Dorsal and declivous faces of propodeum well differentiated, meeting in a distinct propodeal angle; lateral margins of the dorsum of propodeum withoutlamellarexpansions, declivousfacewithlaterallamellarexpansions pointed upwards (Fig. 42 c). Femora not angulated dorsally, mid and hind basitarsi notflattened, with subparallel dorsal and ventral faces. Indorsalview, petiolecompressedanteroposteriorly, anteriormargin withadiscretemedianconcavity, withlaterallamellarexpansionsbroader posteriorly, dorsally with pair of obtuse denticles, subpetiolar process broaderandroundedanteriorly (Fig. 42 b). Postpetiolenarrowerthanpetiole (Fig. 42 b), with lateral lamellar expansions, dorsum of postpetiole with a transverse elevation (Fig. 42 c), subpostpetiolar process pronounced and compressed anteroposteriorly (Fig. 42 b). Gaster suboval, deeply concave anteromedially, with broad anterior lamellar expansions, not extending posteriorly in a carina (Fig. 42 c). Soldiermeasurements (N = 9): HL 0.91 - 1.00; HW 1.09 - 1.19; EL 0.25 - 0.28; PW 0.94 - 1.13; WL 0.97 - 1.13; PTL 0.15 - 0.19; PTW 0.50 - 0.56; PPL 0.18 - 0.21; PPW 0.54 - 0.64; GL 1.04 - 1.19; HBL 0.32 - 0.38; HBW 0.10 - 0.11; TL 3.27 - 3.65; CI 118 - 122; OI 21.2 - 24.4; PI 29.5 - 36.4; HBI 26.7 - 33.7. Soldier description: Body dark brown; frontal lobes and lamellar expansions of pronotum, propodeum, petiole, postpetiole and gaster yellowish to translucent. Gaster dark brown with an anterior and a posterior pair of small yellowish spots (Fig. 43). Mandiblesalveolate. Ventralfaceofhead areolate-microalveolate; dorsum of head with many tubercular elevations, space between tubercles scabrous (Fig. 43 a); frontal lobes weakly rugose. Pronotum scabrous (Fig. 43 c). Meso- and metanotum, meso- and metapleura foveate-microalveolate. Petiole and postpetiole with shallow foveae. Legs, and gaster as in the workers (Fig. 43 c). Pilosity as in the workers, except by anterior portion of head with erect clavate hairs, dorsum of head with canaliculate hairson tubercles, and dorsum of pronotum with sparse canaliculate hairs. Head slightly wider than longer (CI 118 - 122). Mandibles with a strong longitudinal lateral angle. Clypeus without denticles (Fig. 43 a). Dorsum of head disc shaped and convex, strongly tuberculate with a posteriorly pair of distinct larger tubercle (Fig. 43 a, b). Frontal carinae crenulate laterally. Antennaewithatwo-segmented club. Roof of antennal scrobes with alateral crenulate carinae. Lateroventral margins of headwith a posteriorcarinae extending beyondthe eyes until vertexal corners. Vertexal corners forming a pair of broad projections pointed upwards (Fig. 43 b, c). In lateral view, pronotum ascending, pronotal carina well marked and crenulate, mediallyinterrupted (Fig. 43 c). Indorsal view, anterior marginof pronotum gentlyrounded, lateral margins pointed anteriorly. Mesonotumandpropodeumdiscontinuousandflat (Fig. 43 b). Mesonotum, propodeum, legs, petiole, postpetioleand gaster as in workers; except by the narrow lamellar expansions on the declivous face of propodeum (Fig. 43 c). Gyne measurements (N = 4): HL 1.13 - 1.18; HW 1.22 - 1.27; EL 0.30 - 0.31; PW 1.27 - 1.34; WL 1.63 - 1.88; PTL 0.25 - 0.32; PTW 0.50 - 0.51; PPL 0.30 - 0.32; PPW 0.70 - 0.72; GL 1.80 - 1.88; HBL 0.44 - 0.50; HBW 0.13 - 0.15; TL 5.18 - 5.49; CI 106 - 110; OI 23.4 - 25.2; PI 50 - 63.4; HBI 27.5 - 33.3. Gyne description: Bodyblack; frontal lobes anteriorly translucent; dorsum of head, pronotum and legs brownish toyellowish. Firstgastral tergite with an anterior and a posterior pair of whitish spots, each one occupying lessthan one third of the firsttergite length; posteriormargins of eachtergite and sternite yellowish (Fig. 44). Sculpturing of mandibles, dorsal and ventral face of head, pronotum, legs, and gaster as in the soldier. Dorsum of mesonotum, mesopleuron, upper metapleura foveate-microalveolate, lower metapleura rugosemicroalveolate (Fig. 44 b). Propodeumfoveate. Petioleandpostpetiolewith shallowfoveae (Fig. 44 c). Pilosity as in the soldier, but hairsmore abundant on propodeum. Headasinthesoldier (CI 106 - 110). Thetwoposteriorocellipositioned in the posterior tubercular elevations (Fig. 44 a). Indorsalview, anteriormarginofpronotumslightlyrounded, narrower thaninthesoldiers, lateralmarginswithapairofdenticlespointedforwards, pronotalcarinaweaklydeveloped, crenulate, mediallyinterrupted (Fig. 44 c). Dorsally, mesoscutumsubtriangular, anteriormarginrounded; notauli absent; parapsidiallinesfeeblyvisibleandparallel; transscutallineimpressed, reaching the lateral margins of mesosoma; scutoscutellargroove deeply impressed, arched, and scrobiculate; scutellumwell delimited and broader anteriorly; axillaeroundedposteriorly (Fig. 44 c). Laterally, mesopleuralgroovedividing anepisternumandkatepisternum; metapleuradividedinuppermetapleura andlowermetapleurabyagroove; metapleuropropodealgrooveimpressed (Fig. 44 b). Indorsalview, posteriormarginofpropodeumconcavewithapair ofshort, bluntdenticles (Fig. 44 c). Wingvenationasinthegenusdescription. Indorsalview, petiolesubrectangular, lateralmarginsstraight, without lamellar expansions and without dorsal projections, subpetiolar process narrow (Fig. 44 c). Postpetiolelongerthan petiole, withapairoflateralbroad opaqueprojections (Fig. 44 c), withoutprojectionondorsum, subpostpetiolar process pronounced and compressed anteroposteriorly. Gaster protruding anteriorly (Fig. 44 c). Malemeasurements (N = 4): HL 0.58 - 0.60; HW 0.78 - 0.81; EL 0.30 - 0.31; PW 0.80 - 0.84; WL 1.24 - 1.28; PTL 0.21 - 0.25; PTW 0.31 - 0.34; PPL 0.20 - 0.22; PPW 0.33 - 0.39; GL 0.75 - 0.81; HBL 0.38 - 0.44; HBW 0.06 - 0.08; TL 3.03 - 3.13; CI 130 - 138; OI 36.9 - 40.1; PI 66.7 - 74.8; HBI 15.6 - 18.2. Male description: Body brownish to black; femora brownish to yellowish, tibiae and tarsi yellowish (Fig. 45). Mandibles, headanddorsumof mesosomamicroalveolatewithsparse shallowfoveae. Propleuraandmesopleuronweaklystriate; metapleuraand propodeum rugose. Legs, petiole, postpetioleandgaster microalveolate. Body with long and flexuous hairs, more abundant on mandibles, dorsum of head and mesosoma. Legs with long hairs only in the internal face of coxae andfemur. Hairs shorter on gaster (Fig. 45 b). Head wider than longer (CI 130 - 138), broader posteriorly. Mandibles with one apical tooth, lateral angle weakly developed to absent. Clypeus with a median elevation, posterior margin rounded. Frontal carinae not extendingposteriorly. The central portion of head strongly elevated. Eyes occupying more than half of head length (Fig. 45 a). Ocelli positioned at a central elevation. Antenna as the genusdescription. Indorsalview, anteriormarginof pronotumstronglyrounded (Fig. 45 c). Mesoscutumsubtriangular; notaulipresent; parapsidiallinesfeeblyvisibleand parallel; transscutallinereachinglateralmarginsof mesosoma; scutoscutellar groovedeeplyimpressed, arched, andscrobiculate; scutellumwelldelimitedand broaderanteriorly; axillaeroundedposteriorly (Fig. 45 c). Laterally, mesopleural groove dividing anepisternum and katepisternum; metapleura divided in uppermetapleuraandlowermetapleurabyagroove; metapleuropropodeal groove inconspicuous (Fig. 45 b). Dorsal face of propodeum concave, with rounded lateralangles. Wings asin genusdescription. Indorsalview, petioleandpostpetioleconcaveanteriorly, withlateral acute projections near to the anterior margin, without dorsal projections, subpetiolar andsubpostpetiolar processweakly developed (Fig. 45 c). First tergiteofgasterasbroadasmesosoma, occupyinghalforlessthan the totallength of gaster (Fig. 45 b), without anterior projection (Fig. 45 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD0FFE7FCB3FF0BD687FF63.taxon	discussion	Comments: This species differs from others in pineliigroup by the presence of two longitudinal elevations on the posterior portion of head dorsum, lackoflaterallamellarexpansionsondorsumofpropodeumdeclivous face, and body robust, not strongly flattened dorsoventrally. Cephalotes mariadeadrade is the species C. sp. 2 in the molecular phylogenies by Priceetal. (2014, 2016), recoveredatthebaseoftheparaphyleticgrouping formed by the grandinosus and pinelii groups (Fig. 54). Naturalhistory: AcompletecolonyofC. mariadeandrade specieswas collected in Uberlândia (Minas Gerais) in a region of Neotropical savanna (biome Cerrado). The colony occupied four individual nests on the same treeoccupiedbyanaggressivespeciesofCrematogaster of similarsizeand dark brown color. Workers of C. mariadeandrade were observed running inthewell-establishedforagingtrails of theCrematogaster species, which couldindicateaparasiticrelationbetweenthesespecies, likethatdescribed to CephalotesspecularisandCrematogasterampla Forel, 1912. Inthatcase, C. specularis mimics the behavior and posture of their host C. ampla, so they can use the foraging networks and exploit food resources of their hyperaggressivehost withoutbeingnotice (Powell etal., 2014). Cephalotes mariadeandrade workers also moved like the Crematogaster workers whose trails they integrated with and they were similarly undetected by the Crematogaster foragers.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD0FFE7FCB3FF0BD687FF63.taxon	distribution	Distribution: Minas Gerais, Brazil.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD0FFE7FCB3FF0BD687FF63.taxon	etymology	Etymology: Thespecific epithet, inapposition, isinhonorof Maria de Andrade, a Braziliantaxonomist, who dedicatedyearsof workconducting themostcomprehensivestudyof thegenusCephalotes todate. Thisseminal contribution provided the basisfor the development of the present study and the foundation upon which all current Cephalotes research is based.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD3FFE9FFDFFC66D65BFCD2.taxon	description	Figs. 46 a-c, 47 a-c, 48	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD3FFE9FFDFFC66D65BFCD2.taxon	materials_examined	Holotype: BRAZIL, Rio de Janeiro, Parque Nacional de ltatiaia, 15. xii. 1966, 950 - 1000 m, H. Reichardt [MZSP] [examined]. Additional material examined: Brazil, MG, Santana do Riacho, Capão, 1253 m, - 19.206740 - 43.495630, S. Powell, 31. vii. 2018 / C 18 - 208 (3 gynes, 3 males) [DZUP]. Gyne measurements (N = 3): HL 1.73 - 1.75; HW 1.55; EL 0.35; PW 1.43 - 1.48; WL 2.15 - 2.20; PTL 0.63 - 0.68; PTW 0.28 - 0.35; PPL 0.78 - 0.80; PPW 0.30 - 0.38; GL 2.60 - 2.70; HBL 0.54 - 0.58; HBW 0.18 - 0.20; TL 7.48 - 8.02; CI 88.6 - 89.8; OI 22.6; PI 84.4 - 84.6; HBI 33.3 - 37.0. Worker and soldier descriptions can be found in De Andrade and Baroni Urbani, 1999, page 381. Gyne (first description): Body black to brownish; frontal lobes, lateral margins of pronotum and propodeum, and legs brownish to ferruginous. First gastral tergite with an anterior and a posterior pair of yellowish spots, each one occupying more than one third of the first tergite length (Fig. 46). Mandibles and dorsum of head foveate; anterolateral portion of cephalic disc rugose with sparse foveae (Fig. 46 a); ventral face of head longitudinally rugose-microalveolate. Declivous face of propodeum, legs and gaster microalveolate. Dorsum of mesosoma, petiole and postpetiole microalveolate with shallow foveae (Fig. 46 c); lateral of mesosoma microalveolate. Propleura and lower surface of metapleura weakly rugose. Anepisternum and katepisternum weakly foveate (Fig. 46 b). Dorsum of gaster anteriorly finely rugose (Fig. 46 c). Body with appressed canaliculate hairs. Mandibles, mesosoma, petiole and postpetiole with short erect simple hairs; mandibles and anterior margin of clypeus with short erect clavate hairs (Fig. 46 a). Gaster with appressed simplehairs and someshort erecthairs (Fig. 46 b). Head longer thanwide (CI 88.6 - 89.8). Mandibleswith a longitudinal lateral angle. Clypeus without pair of denticles (Fig. 46 a). Dorsum of head discshaped, slightly concave anteriorly. Frontal carinaecrenulate anteriorly. Antennae with a three-segmented club. Roof of antennal scrobes with lateral carinae without projections. Lateroventral margins of head without carinae. Vertexal corners forming a pair of broad projections (Fig. 46 c). Indorsalview, anteriormarginofpronotumslightlyrounded, anterolateral margins with blunt projection, pronotal carina weakly developed, crenulate, mediallyinterrupted (Fig. 46 c). Dorsally, mesoscutumsubtriangular, anteriormargin convex; notauli absent; parapsidial lines visible and parallel; transscutal line reaching lateral margins of mesosoma; scutoscutellar groove impressed, arched, and scrobiculate; scutellum well delimited and broader anteriorly; axillae acute posteriorly (Fig. 46 c). Laterally, mesopleural groove dividing anepisternum and katepisternum; metapleura divided in upper metapleura and lower metapleura by groove; metapleuropropodeal groove not impressed (Fig. 46 b). In dorsal view, posterior margin of propodeum concave with a pair of short, blunt denticles (Fig. 46 c). Wings unknown. In dorsal view, petiole subrectangular, anterior margin concave and lateral margins with a tiny denticle (Fig. 46 c), subpetiolar process narrowwithan acuteprojection anteriorly (Fig. 46 b). Postpetiole longer and wider than petiole, without dorsal elevations, with lateral blunt projections (Fig. 46 c), subpostpetiolar process short and compressed anteroposteriorly (Fig. 46 b). Gaster elongatewith anarrow anteriorlamellar expansion (Fig. 46 c). Malemeasurements (N = 3): HL 0.58 - 0.64; HW 0.93 - 1.00; EL 0.38; PW 1.03 - 1.10; WL 1.48 - 1.60; PTL 0.40 - 0.45; PTW 0.30 - 0.32; PPL 0.40 - 0.45; PPW 0.28 - 0.30; GL 0.93 - 0.98; HBL 0.54 - 0.56; HBW 0.8; TL 3.82 - 4.07; CI 153 - 166; OI 37.5 - 40.5; PI 125 - 140; HBI 14.3 - 14.8. Male (first description): Body predominantly black; apices of femora, tibiae andtarsi, and edges of each sternite and tergite brownish to yellowish (Fig. 47 b). Mandibles, head, pronotum and metanotum microalveolate with sparseshallow foveae (Fig. 47). Anepisternum, katepisternum and lower mesopleuron rugose; upper mesopleuron and propodeum scabrous (Fig. 47 b). Legs, petiole, postpetiole and gaster weakly microalveolate (Fig. 47 c). Body with long and flexuous hairs, more abundant on mandibles, dorsum of head anddorsum of mesosoma. Legswith long hairs only in the internal face of coxae and femur. Hairs shorter on gaster (Fig. 47 b). Head widerthan longer (CI 153 - 166), broader posteriorly. Mandibles with one to three apical teeth, lateral angle absent (Fig. 47 a). Clypeus with a median elevation, posteriormargin rounded. Frontal carinae not extending posteriorly. The central portion ofhead slightly elevated. Eyes occupying more than half of head length (Fig. 47 a). Ocelli positioned at a central elevation. Antenna as the genus description. Indorsalview, anteriormarginofpronotumslightlyrounded, laterally angular (Fig. 47 c). Mesoscutum subtriangular; notauli deeply impressed; parapsidial linesvisible andparallel; transscutal linereaching the lateral margins of mesosoma; scutoscutellar groove deeply impressed, arched, and scrobiculate; scutellum well delimited and broader anteriorly; axillae with posterior denticles (Fig. 47 c). Laterally, mesopleuralgroove dividinganepisternumand katepisternum; metapleuradivided inupper metapleura and lower metapleura by a groove; metapleuropropodeal groove inconspicuous (Fig. 47 b). Dorsal face of propodeum straight and with a rounded angle (Fig. 47 b). Wings as in the genus description. In dorsal view, petiole and postpetiole wider than longer, laterally rounded, without lateral or dorsal projections, subpetiolar and subpostpetiolar process weakly developed (Fig. 47 c). Gaster as long as mesosoma. The first tergite narrower than mesosoma and occupying half or less than the total length of gaster (Fig. 47 b), without anterior projection. (Fig. 47 c).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFDDFFEAFFDFFC29D197FA87.taxon	description	(Figs. 49, 50, 51) The spinosus group by Kempf (1951) comprised the De Andrade and Baroni Urbani’s laminatus and pusillus groups, which share the followingcharacters: anterolateralpronotum angulate, separatefrom the pronotal expansions; propodeum with twopairs of spinesor denticles, with the posterior one longer thananterior; dorsal anddeclivous faces of propodeum differentiated, meeting in a distinct propodeal angle; gaster with distinct anterolateral lamellate borders.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFDDFFEAFFDFFC29D197FA87.taxon	discussion	Despite the shared characters, De Andradeand Baroni Urbani (1999) separated the former Kempf’sspinosus groupinlaminatus andpusillus groups. Thesynapomorphy forthepusillus group is the absence of fine reticulation on thecephalic ventralface and theabsenceof angulate hind femora. However, both characters are the same in the laminatus group, which includes the remaining species of Kempf’s spinosus group and C. duckei, that hadbeen consideredan isolated speciesby Kempf (1951). The synapomorphy for laminatus group, according to De Andrade and Baroni Urbani (1999), is the vertexal corners with a truncate lamella, butthis character is alsopresent in other groups, including thepusillus group. The characters used by Kempf (1951) tojoin the members of the pusillus and laminatus groups within the spinosus group seem more robust. Concordantly, the molecular phylogeny by Priceet al. (2016, see their Fig. S 3) recovered C. pusillus and C. columbicus within the laminatus group, rendering the laminatus group paraphyletic. Based on this molecular and morphological evidence, we here recognize the members of pusillus and laminatus groups as a unique group, under the name “ pusillus ”, since C. pusillus is the oldest species in the group (Fig. 54). Most species inthisgroup occurin Brazil, except for C. christopherseni, known so far only from Colombia, Venezuela, and Panama, and for C. columbicus, known from Colombia and Venezuela.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFDDFFEAFFDFFC29D197FA87.taxon	diagnosis	Diagnosis: In workers and soldiers, dorsal and declivous faces of propodeum differentiated, meeting in a distinct propodeal angle; propodeum with two pairs of spines, the posterior longer than the anterior (Fig. 2 f). Mid and hind basitarsi not flattened. Brazilian species of pusillus group	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFDDFFEAFFDFFC29D197FA87.taxon	discussion	Cephalotes duckei (Forel, 1906) Obsolete combination: Cryptocerus duckei Cephalotes inaequalis (Mann, 1916) Obsolete combination: Cryptocerus inaequalis Cephalotes laminatus (Smith, 1860) Obsolete combination: Cryptocerus laminatus Cephalotes minutus (Fabricius, 1804) Obsolete combination: Cryptocerus minutus = Cephalotes cognatus (Smith, 1862) Obsolete combination: Cryptocerus cognatus = Cephalotes exiguus (Smith, 1867) Obsolete combination: Cryptocerus exiguus = Cephalotes quadrimaculatus (Klug, 1824) Obsolete combination: Cryptocerus quadrimaculatus = Cephalotes volxemi (Emery, 1878) Obsolete combination: Cryptocerus volxemi Cephalotes pusillus (Klug, 1824) Obsolete combination: Cryptocerus pusillus = Cephalotes caustica (Pohl & Kollar, 1832) Obsolete combination: Formica caustica = Cephalotes elongatus (Klug, 1824) Obsolete combination: Cryptocerus elongatus = Cephalotes obtusus (Smith, 1858) Obsolete combination: Cryptocerus obtusus = Cephalotes pusillus brevispinosa (Santschi, 1921) Obsolete combination: Cryptocerus pusillus brevispinosa Cephalotes simillimus (Kempf, 1951) Obsolete combination: Paracryptocerus simillimus Cephalotes spinosus (Mayr, 1862) Obsolete combination: Cryptocerus spinosus = Cephalotes laminatus peruvianus (Forel, 1911) Obsolete combination: Cryptocerus laminatus peruvianus = Cephalotes punctatus (Mayr, 1862) Obsolete combination: Cryptocerus punctatus	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFDDFFEAFFDFFC29D197FA87.taxon	description	(Figs. 50 a-i)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDFF0BD6FCFBA0.taxon	description	(Fig. 52)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDFF0BD6FCFBA0.taxon	description	The solidus group is monotypic. Its unique species was described by Kempf (1974) as Zacryptocerus solidus, within the angustus group. The angustus and solidus groups share the dorsal and declivous faces of propodeum continuous, not meeting in a distinct propodeal angle, but differ by the mesonotum and propodeum unarmed in solidus, while in angustus there is a pair of denticles on the mesonotum and a series of denticles on propodeum.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDFF0BD6FCFBA0.taxon	discussion	In the morphological phylogeny by De Andrade and Baroni Urbani (1999, see their Fig. 24), C. solidus isnot related with theangustus group species, being more closelyrelated to the pusillus group. Nevertheless, most of the characters used in their morphological analysis are based on soldiers. However, C. solidus is a rarely collected species, with only five workers known so far from northern Brazil (Fig. 52). This species was not included in the molecular phylogeny (Price et al., 2016).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDFF0BD6FCFBA0.taxon	diagnosis	Diagnosis: Dorsal and declivous faces of propodeum continuous, not meeting in a distinct propodeal angle. Metanotum and propodeum unarmed (Fig. 2 e).	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDFF0BD6FCFBA0.taxon	diagnosis	Brazilian species of solidus group Cephalotes solidus (Kempf, 1974) Obsolete combination: Zacryptocerus solidus	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	description	(Fig. 53)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	discussion	The monotypicumbraculatus group was created by Kempf (1958 a). He discussed the position of the species, which shares characters with the angustus and pinelii groups, but is different enough to be separated from them. Kempf’s proposal was corroborated by the morphological and molecular phylogenies (De Andrade and Baroni-Urbani, 1999, see their Fig. 24; Priceet al., 2014, 2016, see their Fig. S 3), both phylogenies recover this species as a monophyletic lineage. The recent molecular phylogenies byPrice et al. (2014, 2016) recovered umbraculatus as a sister group of the clade formed by the angustus, fiebrigi and prodigiosus groups (Fig. 54). Cephalotes umbraculatus is broadly distributed in Central America and north of South America.	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	diagnosis	Diagnosis: In workers, body with appressed hairs. Dorsal and declivous face of propodeum continuous, not meeting in a distinct propodeal angle, with a variable number of spines. Dorsum of petiole without denticles. Gaster yellowish with a cross-shaped dark macula, anteriorly with lamellarexpansions (Fig. 2 p). In soldiers, in dorsal view, pronotum crest with pointed edges (Fig. 3 f). Brazilian species of umbraculatusgroup	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	discussion	Cephalotes umbraculatus (Fabricius, 1804) Obsolete combination: Cryptocerus umbraculatus = Cephalotes elegans (Smith, 1853)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	description	Obsolete combination: Cryptocerus elegans	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
038587E5FFD8FFECFFFDF8B2D1DCFB92.taxon	synonymic_list	= Cephalotes flavomaculatus (Mayr, 1862)	en	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: 1-52, DOI: 10.1590/1806-9665-RBENT-2021-0028, URL: http://dx.doi.org/10.1590/1806-9665-rbent-2021-0028
