Cephalotes mariadeandrade, Oliveira & Powell & Feitosa, 2021

Oliveira, Aline Machado, Powell, Scott & Feitosa, Rodrigo Machado, 2021, A taxonomic study of the Brazilian turtle ants (Formicidae: Myrmicinae: Cephalotes), Revista Brasileira de Entomologia 65 (3), No. e 20210028, pp. 1-52 : 37-39

Cephalotes mariadeandrade newspecies

urn:lsid:zoobank.org:act:76BDEF90-0721-48AA-A898-1F9A92280852 ( Figs. 42 a-c, 43a-c, 44a-c, 45a-c, 48)

Holotype: BRAZIL,MG, Uberlândia , -19.033630 -48.318400, 860m, 03.viii.2016. S. Powell et al. col. /C16-64, 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].

Diagnosis: Amember of pinelii speciesgroup. Workerswith lateral lamellar extensions of the propodeum on the declivous face but not on the dorsal face ( Fig. 42c ). 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. 42c ). 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.42c ). Gaster with sparse short appressed simple hairs, anterior lamellar expansions glabrous ( Fig. 42c ).

Headslightlywiderthanlong(CI110-123),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.42c ).Antennaewithatwo-segmentedclub. Lateroventralmarginsofheadwithposteriorcarinaeextending beyondthe eyesuntilvertexalcorners.Vertexalcornerswithanarrowlamellarexpansion.

Mesosomaalmostflatinlateralview,withagentlyascendantpronotum ( Fig.42b ). Indorsalview, lateralmarginsofpronotum withconcavelamellar expansions, broader anteriorly; promesonotalgroove absent, instead of it is an elevation on dorsum of pronotum ( Fig.42c ). 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. 42c ). 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.42b ).Postpetiolenarrowerthanpetiole ( Fig. 42b ), with lateral lamellar expansions, dorsum of postpetiole with a transverse elevation ( Fig. 42c ), subpostpetiolar process pronounced and compressed anteroposteriorly ( Fig. 42b ).

Gaster suboval, deeply concave anteromedially, with broad anterior lamellar expansions, not extending posteriorly in a carina ( Fig. 42c ).

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. 43a ); frontal lobes weakly rugose. Pronotum scabrous ( Fig. 43c ). Meso- and metanotum, meso- and metapleura foveate-microalveolate. Petiole and postpetiole with shallow foveae. Legs, and gaster as in the workers ( Fig. 43c ).

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. 43a ). Dorsum of head disc shaped and convex, strongly tuberculate with a posteriorly pair of distinct larger tubercle ( Fig. 43a, 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. 43b, c ).

In lateral view, pronotum ascending, pronotal carina well marked and crenulate, mediallyinterrupted ( Fig. 43c ). Indorsal view, anterior marginof pronotum gentlyrounded, lateral margins pointed anteriorly. Mesonotumandpropodeumdiscontinuousandflat ( Fig. 43b ). Mesonotum, propodeum, legs, petiole, postpetioleand gaster as in workers; except by the narrow lamellar expansions on the declivous face of propodeum ( Fig. 43c ).

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.44b ). Propodeumfoveate.Petioleandpostpetiolewith shallowfoveae ( Fig.44c ).

Pilosity as in the soldier, but hairsmore abundant on propodeum.

Headasinthesoldier (CI106-110).Thetwoposteriorocellipositioned in the posterior tubercular elevations ( Fig.44a ).

Indorsalview,anteriormarginofpronotumslightlyrounded,narrower thaninthesoldiers,lateralmarginswithapairofdenticlespointedforwards, pronotalcarinaweaklydeveloped,crenulate,mediallyinterrupted ( Fig.44c ). 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.44c ).Laterally,mesopleuralgroovedividing anepisternumandkatepisternum;metapleuradividedinuppermetapleura andlowermetapleurabyagroove;metapleuropropodealgrooveimpressed ( Fig.44b ).Indorsalview,posteriormarginofpropodeumconcavewithapair ofshort,bluntdenticles( Fig.44c ).Wingvenationasinthegenusdescription.

Indorsalview,petiolesubrectangular, lateralmarginsstraight,without lamellar expansions and without dorsal projections, subpetiolar process narrow ( Fig.44c ).Postpetiolelongerthan petiole,withapairoflateralbroad opaqueprojections( Fig.44c ),withoutprojectionondorsum,subpostpetiolar process pronounced and compressed anteroposteriorly.

Gaster protruding anteriorly ( Fig. 44c ).

Malemeasurements (N=4):HL0.58-0.60; HW0.78-0.81; EL0.30-0.31; PW0.80-0.84; WL 1.24-1.28;PTL 0.21-0.25;PTW 0.31-0.34; PPL 0.20-0.22; PPW0.33-0.39;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. 45b ).

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. 45a ). Ocelli positioned at a central elevation. Antenna as the genusdescription.

Indorsalview,anteriormarginof pronotumstronglyrounded ( Fig.45c ). Mesoscutumsubtriangular;notaulipresent;parapsidiallinesfeeblyvisibleand parallel;transscutallinereachinglateralmarginsof mesosoma;scutoscutellar groovedeeplyimpressed,arched,andscrobiculate;scutellumwelldelimitedand broaderanteriorly;axillaeroundedposteriorly( Fig.45c ).Laterally,mesopleural groove dividing anepisternum and katepisternum; metapleura divided in uppermetapleuraandlowermetapleurabyagroove;metapleuropropodeal groove inconspicuous ( Fig. 45b ). 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. 45c ).

First tergiteofgasterasbroadasmesosoma,occupyinghalforlessthan the totallength of gaster ( Fig. 45b ), without anterior projection ( Fig. 45c ).

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.

Distribution: Minas Gerais, Brazil.

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.

Figures

Figure 42 Worker of Cephalotes mariadeandrade. A: frontal view. B: lateral view. C: dorsal view. Brazil:MG, Uberlândia.

Figure 43 Soldier of Cephalotes mariadeandrade. A: frontal view. B: lateral view. C: dorsal view. Brazil: MG, Uberlândia.

Figure 44 Gyne ofCephalotes mariadeandrade. A:frontal view.B:lateral view.C:dorsal view.Brazil: MG, Uberlândia.

Figure45 Male ofCephalotes mariadeandrade. A:frontal view.B:lateral view.C:dorsal view.Brazil:MG, Uberlândia.

Figure54 Themaximum cladecredibility treefrom phylogenetic analysesperformed byPriceet al.(2016) basedonmorphologicalandmoleculardata.The tree has beentrimmedtoinclude onlythose species knownfromBrazil.Thelabelsforthespecies groupsfollowour proposals,including thesynonymies,andtheprevious namesforthesynonymized groupsareshown ingrey.