J. HYM. RES. Vol. 17(1), 2008, pp. 64–82

Revisionary Studies on the Enigmatic Neotropical Stegomyrmex Emery, 1912 (: Formicidae: ), With the Description of Two New Species

RODRIGO M. FEITOSA,CARLOS R. F. BRANDA˜ O AND JORGE L. M. DINIZ (RMF, CRFB) Museu de Zoologia da Universidade de Sa˜o Paulo, Av. Nazare´ 481, Sa˜o Paulo, SP, 04263-000, Brazil; RMF email: [email protected]; CRFB email: [email protected] (JLMD) Campus Jataı´, Unidade Jatoba´, Universidade Federal de Goia´s, BR 364, km 192, Jataı´, GO, 75800-000, Brazil; email: [email protected] ______

Abstract.—The recent increase in leaf litter sampling effort in Neotropical wet forests has revealed new and interesting records of the highly specialized myrmicine ant genus Stegomyrmex Emery, previously considered as extremely rare. We present a modified diagnosis for the genus and describe Stegomyrmex bensoni n. sp. and S. olindae n. sp., based on, respectively, workers, males, and gyne (central-north Brazil) and on a single worker (northern Brazil). Stegomyrmex vizottoi Diniz (southeastern Brazil) is redescribed and compared with S. olindae n. sp.; these species present significant differences in size, sculpturation, and sting apparatus morphology. The males of S. vizottoi are described for the first time. A key for workers and queens and a distribution map for the five know Stegomyrmex species are provided. ______

Stegomyrmex is the sole representative of species, S. manni, from Barro Colorado the peculiar and exclusively Neotropical Island, Panama, and agreed with Wheeler’s myrmicine tribe Stegomyrmecini (Bolton placement of the genus in an individual 2003). These ants have been considered tribe. extremely rare by many authors, perhaps Bernard (1951) and Lenko (1965) com- due to their cryptobiotic habits enhanced mented on the morphological resemblance by peculiar soil-binding pilosity (Ho¨lldo- of Stegomyrmex to some Attini. However, bler and Wilson 1986), and by the foraging Brown (1949), Brown and Kempf (1960), technique they employ (Diniz and Branda˜o and Ho¨lldobler and Wilson (1986) consid- 1993). However, recent collections employ- ered stegomyrmecine ants more closely ing large-scale sampling (e.g. Agosti et al. related to Basicerotini than to Attini or 2000) have revealed that they are relatively Dacetini, mainly by the presence of deep common inhabitants of the dense leaf litter antennal scrobes and the soil-binding of Neotropical forests. pilosity. Dlussky and Fedoseeva (1988) Emery (1912) described Stegomyrmex considered Stegomyrmex as incertae sedis with a single species, S. connectens, based in Myrmicinae, without further discus- on a gyne and a male from Peru and sion. However, in the last proposals of Bolivia respectively. Emery included Ste- Bolton (1994, 2003, 2006 et al.), Stegomyr- gomyrmex in the Dacetini based on gyne mex is placed in its own tribe within the characters. Wheeler (1922) established a Myrmicinae. Bolton (2003) commented new tribe, Stegomyrmicini (sic), with Ste- that the structure of the promesonotum gomyrmex as its only member. He separat- may suggest a relationship between Ste- ed it from the Dacetini mainly by the shape gomyrmecini and Pheidolini, but that of mandibles and wing venation. Smith there is no undisputed evidence for this (1946) described the second Stegomyrmex yet. VOLUME 17, NUMBER 1, 2008 65

Lenko (1965) found a worker of S. METHODS vizottoi (identified by him as S. manni)in This study was based on the available the gizzard of a Conopophaga lineata Wied specimens in the collection of the Museu (Aves, Conopophagidae). Ho¨lldobler & de Zoologia da Universidade de Sa˜o Paulo, Wilson (1986) commented on the pre- Sa˜o Paulo, Brazil, which is believed to hold sumed role of the soil-binding hairs of most of the known Stegomyrmex specimens. basicerotine and stegomyrmecine ants in Depository collections are referred to by enhancing their camouflage to predators. the following acronyms: Diniz (1990) was the first to revise the BMNH – The Natural History Museum, of Stegomyrmecini, describing London, UK. the third species of the genus, Stegomyrmex CASC – California Academy of Sciences, vizottoi, based on workers and a gyne from San Francisco, California, USA. Brazil and Paraguay. In the same work, CPDC – Centro de Pesquisas do Cacau, Diniz commented on the relatively slow Itabuna, Bahia, Brazil. movements of stegomyrmecine ants. JLMD – Laborato´rio de Zoologia, Cam- Diniz and Branda˜o (1993) were the first to pus Jataı´, Universidade Federal de Goia´s, describe the nesting habits of Stegomyrmex, Brazil. based on observations on colonies of S. vizottoi LACM from Mirassol, state of Sa˜o Paulo, Brazil, – Los Angeles County Museum describing nest architecture, population dis- of Natural History, Los Angeles, Califor- tribution among nest chambers, different nia, USA. workerbehaviorsateachpartofthenest, MCSN – Museo Civico di Storia Natur- and the foraging habits of the workers, which ale ‘‘Giacomo Doria’’, Genoa, Italy. exploit the environment surrounding their MPEG – Museu Paraense Emı´lio Goeldi, nests singly, searching for myriapod eggs. Bele´m, Para´, Brazil. Recent surveys of leaf litter ants in the MZSP – Museu de Zoologia da Uni- Brazilian Atlantic forest and in sparse versidade de Sa˜o Paulo, Sa˜o Paulo, Brazil. localities of central and northern Brazil USNM – National Museum of Natural revealed several Stegomyrmex specimens, History – Smithsonian Institution, Wash- including a remarkable new species de- ington, DC, USA. scribed here, and extending considerably The terms for external morphology and the known distribution range of S. vizottoi. surface sculpturing follow, respectively, Our analysis of S. vizottoi along its distri- Bolton (1994, 2000) and Harris (1979). The bution shows, however, that, as presently terms for wing venation follow Brown and accepted, it includes two distinct species, Nutting (1950). The reproductive females recognizable by the surface sculpture, by are here called ‘‘gynes’’, as suggested by morphometry, and by differences in the De Andrade and Baroni Urbani (1999). sting apparatus, as we fully describe and Measurements were obtained with a comment on below. micrometric reticule and using the scale In this paper we offer taxonomic notes of a scanning electron microscope (SEM). on the peculiar ant genus Stegomyrmex, All measurements are given in mm, and based on the study of the specimens the abbreviations used are: deposited in the Museu de Zoologia da HW: head width; the maximum width of Universidade de Sa˜o Paulo ant collection, the head capsule, measured in full face literature information, and enriched by view, at a median transverse line that unpublished observations. We also de- touches the superior margins of the com- scribe two new species and comment on pound eyes. new records and information regarding HL: head length; the maximum measur- these seldom collected ants. able length of head capsule excluding the 66 JOURNAL OF HYMENOPTERA RESEARCH mandible, measured in full face view, in a in xylene, and then mounted in Canada straight line from the midpoint of the balsam for observation and illustration anterior clypeal margin to the midpoint of under optical microscope. The terms for the vertexal margin. sting apparatus morphology follow Kugler SL: antennal scape length; the chord (1978). length of the antennal scape, excluding Coordinates of localities were obtained the basal condyle and its peduncle. from the information on the specimens WL: mesosoma length (Weber’s length); labels and after consulting the ENCARTA the diagonal length of mesosoma in profile, World AtlasH (Microsoft); they were plot- from the midpoint of the anterior pronotal ted on the distribution map generated by declivity to the posterior basal angle of the the software ArcView 3.2 GISH. metapleuron. When citing label data, we present PL: petiole length; the longitudinal axis additional information between brackets, of petiole in lateral view. explanation of codes on the labels, eventual PPL: postpetiole length; the longitudinal corrections to the misprints, and reference axis of postpetiole in lateral view. to the notebooks from which we took GL: gaster length; the maximum length information regarding the localities and/ of gaster in lateral view, excluding sting. or the biology of the species. TL: total length; the summed length of HL (plus the closed mandibles), WL, PL, RESULTS PPL, and GL. CI: cephalic index. HW x 100/HL. Stegomyrmex Emery, 1912 SI: scape index. SL x 100/HW. Stegomyrmex Emery, 1912: 99. Gyne. Type The SEM images of Stegomyrmex speci- species: Stegomyrmex connectens, by mono- mens were obtained from a single speci- typy. Emery, 1912: 101 (placement in Dace- men of each species. The specimens were tini); Emery, 1914: 42 (placement in Dace- previously cleaned in acetone, critical- tini); Forel, 1917: 246 (placement in Dacetini); point dried in a Balzer (Bal-TecH CPD Emery, 1924: 314 (placement in Dacetini, 030), and sputtered over with gold (Bal- diagnosis, catalogue); Wheeler, 1922: 668 TecH SCD 050). After that, the specimens (establishment of Stegomyrmecini [as Stego- were mounted on the tip of metallic myrmicini]); Donisthorpe, 1943: 727 (place- triangles using silver glue and then affixed ment in Dacetini, list of type specimens); Smith, 1946: 286 (revision); Brown and to stubs for the electron microscopy. The Kempf, 1960: 162 (systematic notes); Lenko, images were obtained under several mag- 1965: 201 (distribution and biology); Kempf, nifications (40 to 300x), according to the 1972: 242 (catalogue); Wheeler and Wheeler, size of the specimen and/or structure 1985: 258 (tribal classification); Ho¨lldobler observed. Finally, the images were edited and Wilson, 1986: 16 (pilosity); Dlussky & (Adobe PhotoShop 7.0H) to enhance some Fedoseeva, 1988: 81 (incertae sedis in Myrmi- brightness and contrast details. cinae); Diniz, 1990: 277 (revision, species We studied also the sting apparatus of key); Ho¨lldobler and Wilson, 1990: 15 (tribal the species from which we had enough classification); Branda˜o, 1991: 379 (cata- individuals. The sting was obtained by logue); Diniz and Branda˜o, 1993: 301 (biol- rehydrating ants in 70% ethanol, extracting ogy); Bolton, 1994: 106 (catalogue); Bolton, 1995a: 1052 (census); Bolton, 1995b: 392 the terminal segments from the gaster, (catalogue); Serna, 2002: 217 (first record clearing them in 55–60uC lactophenol for for Colombia); Bolton, 2003: 255 (diagnosis, five minutes (or longer if necessary), synoptic classification); Ferna´ndez and Os- rinsing twice in 70% ethanol, and twice in pina, 2003: 49 (census); Ferna´ndez, 2003: 325 95% ethanol. After the clearing process, the (genera list for Neotropics); Bolton et al., sting apparatus was dismembered, soaked 2006 (catalogue). VOLUME 17, NUMBER 1, 2008 67

Worker. Monomorphic. 5 to 6.5 mm in from the integument sculpture; placed on length. Reddish brown to black. Integu- the sides of head immediately beneath the ment thick, shining and in general densely antennal scrobe, but visible when head is areolate, except for S. bensoni. Pilosity in full face view. conspicuous and bizarre; hairs varying Mesosoma, in dorsal view, slender, from short, subdecumbent and filiform to widest at the level of the anterior area of long, suberect, and variably branched; pronotum. Promesonotum evenly rounded mandibles sparsely covered by long fili- in profile, dome-like; anterosuperior corner form hairs; anterior margin of clypeus of anepisternum set much lower than the bearing one or two pairs of very long adjacent surface, forming a deep fovea; setae, reaching half the length of mandi- promesonotal suture almost obsolete in bles, but without an isolated median seta; some individuals. Mesonotum elongate appendages covered by short decumbent with posterior portion sloping down; me- hairs and by a fine and dense appressed tapropodeal impression relatively broad pubescence; inferior corners of pronotum and usually shallow, except for S. bensoni. with a dense row of plumose hairs. Propodeum, in side view, variably convex Head subtrapezoidal with vertexal mar- dorsally, and with the declivity sinuous; gin slightly depressed to slightly convex; propodeal spiracles low on side and raised occipital corners angulate; broader pos- in prominent, subcylindrical protuberanc- terad. Palpal formula 2:2. Labrum bilobed. es; propodeal spines short and more or less Mandible triangular, long, strongly curved acute; propodeal lobes large and usually down apically and with the blades crossing projected over the petiolar peduncle. Legs apically when mandibles are closed; mas- relatively long; femora and tibiae moder- ticatory border multidenticulate (total den- ately incrassated; tarsal claws simple; tal count 12–15), with the apical tooth metatibial spurs absent. longer than the preceding ones. Median Petiole long and pedunculate, with about portion of clypeus narrow, flat and vertical, twice the length of the postpetiole; petiolar not bicarinate, quite narrowly inserted node variably convex in profile; ventral between the frontal lobes. Frontal lobes carina present and bearing 0–2 blunt enormously expanded anterolaterally and anterior projections. Postpetiole approxi- projected far out over the lateral portions mately as long as broad, globose, without of clypeus and mandibles. Each frontal ventral process. Gaster oval, without basal lobe covering dorsally a very deep anten- shoulder; tergite of abdominal segment IV nal scrobe; in full face view, space between (first gastral) not broadly overlapping frontal lobes narrowest near the middle of sternite on gaster ventral surface. head, revealing the compound eyes; clyp- Sting apparatus. Spiracular plate with eus and basal portion of mandible entirely spiracle placed ventrally; anal plate with concealed by the frontal lobes. Frontal area several sensillae; lancet with a pair of impressed, glabrous and smooth, the ante- functional valves; furcula with indistinct rior suture obsolete. A shallow groove, dorsal arms. almost devoid of any sculpture, present on Gyne. Like conspecific worker, with the each side of the head dorsum, extending modifications expected for myrmicine from the frontal area to the occipital corner gynes. Anterior ocellus slightly larger than of head, the two grooves meeting anterior- posterior ones. Notauli and parapsidial ly, forming a noticeable V. Antenna with 12 lines usually indistinct from surrounding segments, with a three-segmented club; sculpture; prescutellum with central area antenal scape slender, curved basally and indistinct, scutoscutellar sulcus shallowly broader at apex. Compound eye exceed- impressed, with transversal rugulae vary- ingly small, oval in shape, almost indistinct ing in number and forming distinct cells; 68 JOURNAL OF HYMENOPTERA RESEARCH lateral wing of prescutellum not projecting Mesosoma robust; prescutellum separat- laterally; scutellum semicircular, with its ed from scutellum by an impression with posterior half always sloping down and short longitudinal rugae. Scutellum narrow with posterior border concave; propodeal posterad. Metanotum narrow, with blunt spines shorter than in conspecific workers. median tumosity. Propodeum dorsal face Forewing with distinct and strongly flat, steeply sloping posterad, unarmed. colored stigma; longitudinal veins Sc+R, Legs slender, middle and hind tibiae SR, M+Cu, and A present. Cells R, Cu and without apical spurs; tarsal claws 1M closed. Hind wing with Sc+R extending slender and simple. Wing venation as in shortly beyond point where they connect to the gynes. M, which extends as a tubular vein up to Petiole clavate, pedunculate, and with a the wing distal border; Cu cell closed and long, low, rounded node. Postpetiole as very short; six to eight submedian hamuli. broad as long, attached to the gaster by Male. Dark brown to black, with ap- almost its full width. Gaster elongate, with pendages and gaster usually lighter. Integ- first segment occupying most of its length; ument densely sculptured, opaque or visible apical segments subequal in length. nearly so, except for the postpetiole and Comments.—We revise the Stegomyrmex gaster which are smooth and shining; diagnosis presented by Diniz (1990) in appendages very finely punctate. Pilosity order to include information on the shape composed of fine hairs, whitish to golden, of the head and on the structure of the mostly curved or suberect on body, sparser alate’s mesosoma, besides features present on metasoma. Apressed pubescence on in S. bensoni n. sp and S. olindae n. sp. antennae and legs. Apomorphies for Stegomyrmecini defined Head broadest across compound eyes, by Bolton (2003) hold true for the new narrowed anteriorly; median portion of species. vertexal margin usually weakly convex; Despite the recent information regarding occipital corners rounded; ocelli promi- these seldom collected ants, the phyloge- nent. Mandible relatively developed and netic position of Stegomyrmex remains truly subtriangular; masticatory border multi- enigmatic. The affinities with Dacetini and denticulate, with the apical tooth much Attini, proposed in the past, seem improb- more developed than the others. Clypeus able by the significant differences in habits broad. Frontal lobes not so developed as in and morphology. Despite the body sculp- the conspecific gynes and workers, but turation patterns and the presence of concealing the antennal insertions, forming specialized pilosity approximating Stego- a short and shallow antennal scrobe. myrmex and Basicerotini (Ho¨lldobler and Antennae long and slender with 13 seg- Wilson 1986), the possibility of homoplasy ments; scapes relatively short. can not be presently discarded.

REVISED KEY TO THE STEGOMYRMEX SPECIES (WORKERS AND GYNES)

1. Integument of mesosoma predominantly smooth and shining; body covered mainly by sparse aggregations of somewhat curved and multibranched hairs; metapropo- deal groove deeply impressed; petiole without anteroventral spines (state of Para´, Brazil) ...... S. bensoni sp. n. – Integument of mesosoma predominantly sculptured, areolate; body covered mainly by sparse and erect clavate setae; metapropodeal groove only moderately impressed; petiole with at least one anteroventral spine ...... 2 VOLUME 17, NUMBER 1, 2008 69

2. Petiole with two anteroventral spines; inferior margin of pronotum with a row of filiform hairs; known only from the gyne (Peru and Bolivia) . . . S. connectens Emery – Petiole with a single anteroventral spine; inferior margin of pronotum with a row of plumose hairs ...... 3 3. Promesonotum much higher than propodeum, in lateral view; propodeal spines blunt and directed posteriorly; in dorsal view propodeal spiracles strongly projected laterally (Costa Rica, Panama and Colombia) ...... S. manni Smith – Promesonotum slightly higher than propodeum, in lateral view; propodeal spines subtriangular, acute and directed upwards; in dorsal view propodeal spiracles not strongly projected laterally ...... 4 4. Mesosoma length $ 1.59 mm; mesosoma partially sculptured, with foveae sparsely set on the polished integument; metapropodeal impression without a projecting tubercle; nucal area predominantly smooth; in dorsal view basal face of propodeum relatively narrow (northern Argentina, Paraguay and southeastern Brazil)...... S. vizottoi Diniz – Mesosoma length , 1.59 mm); mesosoma strongly sculptured, with the integument completely areolate; metapropodeal impression with a projecting tubercle; nucal area predominantly sculptured; in dorsal view basal face of propodeum relatively broad (central-north Brazil) ...... S. olindae sp. n.

Stegomyrmex bensoni n. sp. and shining, except for a few scattered (Figs 1, 7) punctures at the inferior portion of meso and metapleuron; legs smooth and rather Holotype worker.—BRAZIL: Para´, Canaa˜ opaque; petiole and postpetiole smooth and dos Caraja´s(06u44949’’S, 50u219050W) shining with some sparse piligerous punc- (Gruta NV06) 22–28.ii.2005 (Andrade & tures; dorsum of gaster feebly shining and Arnoni) [MZSP]. with sparse and fine punctuation. Worker description.—HW 1.26; HL 1.09; Pilosity golden and extremely diverse; ML 0.61; SL 0.84; WL 1.77; PL 0.78; PPL 0.45; sparse filiform hairs covering the dorsum GL 1.70; TL 6.40; CI 115.56; SI 66.35. Color of mandible, external borders of frontal reddish brown. Basal portion of mandible carinae, antenal scapes, legs, dorsum of finely and densely striate, with large and mesosoma and metasoma; long, slightly sparse piligerous punctures, apical portion curved, moderately clavate hairs present and masticatory border mostly smooth and on dorsum of head and promesonotum; shining; inner surface of antennal scrobes short, curved, branched hairs present on with fine, dense, transversal and concentric head occipital corners, dorsal surface of striation; dorsal surface of head predomi- legs and gaster; posteroventral corners of nantly smooth and shining, with scattered head, anterior and lateral portions of punctures near the vertexal border; margin promesonotum, dorsum of metanotum of frontal lobes finely areolate-rugose; central and propodeum, ventral and lateral faces portion of each frontal lobe virtually trans- of waist and anterior portion of gaster (in lucent, so that is possible to observe the inner special the sternite) with aggregations of surface of the antennal scrobes near the long, multibranched (plumose), curved insertions of antennae; antennae opaque and hairs, so that the integument is hardly finely punctate; lateral and ventral surface of visible in these areas. head deeply areolate; occipital face of head Head vertexal margin convex in the smooth and shining except for the nucal middle. Compound eyes exceedingly collar which is regularly and deeply scrobi- small, with circa three almost indistinct culate; mesosoma almost entirely smooth facets at the maximum diameter. 70 JOURNAL OF HYMENOPTERA RESEARCH

Fig. 1. Stegomyrmex bensoni n. sp., worker. A, head in full face view; B, SEM close-up view of promesonotal multibranched hairs; C, habitus.

Promesonotum strongly convex dorsal- petiolar peduncle without anterior projec- ly; metapropodeal groove deeply im- tions. Postpetiole strongly convex dorsally pressed; propodeal spines short and sub- and without ventral processes. Gaster oval triangular, entirely covered by the propo- and robust. deum pilosity; propodeal spiracle wide Gyne.—Unknown. open, and moderately projected laterad; Male.—Unknown. propodeal lobes subquadrate and weakly Etymology.—Species named after the projected over the petiolar peduncle. prominent Brazilian-American ecologist, Petiole elongate, slightly arched, with a Woodruff Whitman Benson, well known prominent rounded node; ventral carina of for the study of ecological interactions in VOLUME 17, NUMBER 1, 2008 71

Brazilian ecosystems. He organized for and 1 gyne) [LACM]; Mato Grosso: Sto. several years a field course on Ecology Antoˆnio de Leverger, A´ guas Quentes (High (graduate program of the Universidade Cerrado) (n. 0184) 26.x.1984 (J.C. Trager) (1 Estadual de Campinas, Sa˜o Paulo), given worker) [MZSP]; Minas Gerais: Timo´teo, P.E. for several years in the Serra dos Caraja´s, do Rio Doce (TM3–8) 07.v.2005 (Esteves, F.A.) (1 worker) [MZSP]; Tocantins: Palmeirante (Mata Para´, where generations of Brazilian grad- Ciliar/Cerrada˜o) (07u529250S, 47u319480W) 10– uate students had their first field experi- 15.xii.2001 (Albuquerque and Silva) (1 worker) ence, and where the unique known spec- [MZSP]; same data (1 worker) [CASC]; Aragua- imen of S. bensoni was found. cema (08u599200S, 49u409410W) 16–30.xi.2005 Comments.—The peculiar multibranched (Silva,R.R.andFeitosa,R.M.)(1worker) pilosity, allied to the deep metapropodeal [MZSP]; same data (1 worker) [MPEG]; same groove, and the absence of anteroventral data (1 worker) [USNM]. petiolar projections, easily separate this Worker description.—Holotype (workers species from all others in the genus. N5 8); HW 1.09 (1.04–1.22); HL 0.97 (0.92– The single worker known thus far was 1.07); ML 0.49 (0.46–0.49); SL 0.74 (0.69– captured by our colleague arachnologist, 0.80); WL 1.43 (1.33–1.53); PL 0.68 (0.61– Renata Andrade, while searching for cav- 0.70); PPL 0.39 (0.33–0.44); GL 1.36 (1.24– ernicolous pseudoscorpions in Serra dos 1.50); TL 5.32 (5.01–5.70); CI 113.75 (104.88– Caraja´s, southeastern state of Para´ (Ama- 114.63); SI 66.59 (63.83–67.44). Dark brown zon region), Brazil. The finding represents to ferruginous, with appendages some- the first record of a stegomyrmecine ant in what lighter. Mandible finely and densely the Amazon Region. striate, with large and sparse piligerous Despite the fact that the only know punctures, except for the masticatory bor- specimen was collected inside a cave, near der and dorsum of apical portion which its mouth, there is no undisputed evidence are smooth and shining; inner surface of that Stegomyrmex bensoni is restricted to this antennal scrobes punctate and with fine habitat. transversal striation; central disc of head and external margin of frontal lobes dense- Stegomyrmex olindae sp. n. ly areolate-rugose; oblique lateral grooves (Figs 2, 3, 6, 7) of head, frontal area and posterior portion Holotype worker.—BRAZIL: Tocantins, of frontal lobes with smooth areas and Palmeiras do Tocantins (06u409120S, sparse punctuation; anterior portion of 47u319480W) (Winkler n.3) 14–19.i.2005 (Sil- frontal lobes shallowly areolate and with va, R.R. and Silvestre, R.) [MZSP]. irregular longitudinal rugulae; antennae Stegomyrmex vizottoi Diniz, 1990: 290 (in opaque and finely punctate; lateral, ventral part). and occipital surfaces of head deeply areolate; mesosoma (including the anterior Paratypes.—BRAZIL: Bahia: Ilhe´us, CEPEC- coxae), petiole, and postpetiole entirely a´rea Zoolog. (Km22 Ilhe´us-Itabuna) x.1986 (J. and deeply areolate; legs opaque and Delabie) (1 worker) [MZSP]; Porto Seguro, E.E. weakly sculptured; surface of gaster deep- Pau Brasil (16u239330S, 39u109990W) (Winkler ly and densely foveolate. n.1) 16.vi.2000 (Santos, J.R.M and Soares, J.C) (1 Pilosity cream-colored. Body covered by worker) [MZSP]; Maranha˜o: Ac¸ailaˆndia, Horto abundant, long, slightly stiffened, moder- Faz. Pompe´ia (04u529300S, 47u179400W) 13– ately clavate hairs, somewhat shorter in the 22.ii.2006 (Silva, R.R. and Feitosa, R.M.) (1 worker) [MZSP]; Estreito, Fazenda Itaueiras external borders of frontal lobes, antenal (06u31954’’S, 47u729160W) 07–13.i.2005 (Silva, scapes, and legs; mandible with long, R.R. and Silvestre, R.) (2 workers and 1 gyne) sparse filiform setae; short, curved, plu- [MZSP]; same data (1 worker) [BMNH]; same mose hairs present on the posteroventral data (1 worker) [CDPC]; same data (1 worker corners of head, inferior and lateral por- 72 JOURNAL OF HYMENOPTERA RESEARCH tions of pronotum, and more rarely on the sensilla sparsely distributed over the plate lateral surfaces of waist; occipital face of dorsum. Oblong plate with long posterior head and lateral surface of mesonotum, apodema; subterminal tubercle with metanotum, and propodeum virtually gla- rounded apex; postincision well devel- brous. oped. Gonostylus one-segmented and Vertexal border gently convex and with with six chaetae, five subequal in length a discrete concavity medially. Compound and one extremely long; terminal sector eyes with circa six facets at maximum short and membranous, with dorsoterm- diameter. inal and companion chaetae present. Promesonotum strongly convex dorsal- Triangular plate as long as broad, without ly, in lateral view; promesonotal suture tubercles or projections. Lancets with distinct only in the lateral faces of prome- functional valves; sensorial barbles absent; sonotum; anepisternum set lower than the dorsal and ventral margins converging adjacent surface; metapropodeal groove towards the apex. Sting shaft weakly relatively large, moderately impressed sclerotized, probably not perforating; dor- and with a median triangular projection; sum of valve chamber indistinct in pro- propodeal spines subtriangular, directed file; internal apophysis absent; basal con- upwards and with the posterior faces nection gently concave; anterolateral pro- enlarged medially; propodeal spiracles cesses well developed, as broad as the relatively wide, and considerably projected furcula lateral arms; campaniform sensilla posterad; propodeal lobes rounded and absent. Dorsal arm of furcula relatively moderately projected over the petiolar reduced, indistinct; lateral arms well peduncle. In dorsal view, the propodeum developed; fulcral articulation connected is relatively broad, slightly narrower than to the sting basis only by its lateral the promesonotum. corners. Petiole elongate, gently arched, with a Gyne.—(N5 2); HW 1.24–1.26; HL 1.04; relatively long rounded node; ventral ML 0.52–0.53; SL 0.78; WL 1.82; PL 0.80– carina of peduncle with a well-developed 0.83; PPL 0.46–0.49; GL 1.82–1.84; TL 6.50– anterior projection. Postpetiole with a long 6.51; CI 118.60–120.93; SI 62.12–62.75. Like and moderately convex dorsal face, with- conspecific worker, with the modifications out ventral projections. Gaster oval and expected for myrmicine gynes. Plumose robust. hairs restricted to the posteroventral corner Sting apparatus (Fig. 3): Spiracular plate of head and inferior corner of pronotum. subquadrate, not extending towards the Compound eyes with circa 11 facets at medial connection; margin of medial maximum diameter; propodeal spines connection sclerotized; dorsal notch ab- drastically reduced; posterior face of pro- sent; spiracle relatively wide and set close podeum vertical in side view, reaching the to the posterior margin of plate; anterior propodeal lobes in a rounded angle. Wings apodema narrow with the medial region unknown. with a distinct angle; ventral edge vesti- Male.—Unknown. gial, marked only by a weak projection. Etymology.—This species is named after Quadrate plate with the dorsal region as Florinda Gonzaga Teixeira, a long-term broad as the ventral region, except for the and always large-hearted steward of the apodema; apodema area smaller than the MZSP ant lab, at the occasion of her plate body; dorsal margin convex; apex of retirement. She prefers to be called ‘‘Dona anterodorsal corner rounded; posterior Olinda’’, hence the specific name. margin complete. Anal plate with the arc Comments.—While examining specimens rounded and strongly sclerotized; apical of Stegomyrmex vizottoi from the MZSP margin rounded and weakly definite; anal collection, one of us (RMF) noticed that VOLUME 17, NUMBER 1, 2008 73

Fig. 2. Stegomyrmex olindae n. sp., worker (SEM). A, head in full face view; B, close-up view of nucal area; C, habitus. there was a morphologically distinct sub- of Bahia (northeastern Brazil) presented the group of individuals, all collected in the lateral faces of the mesosoma more densely northern range of S. vizottoi distribution. sculptured and the basal face of propodeum Diniz (1990) already mentioned, while relatively enlarged in dorsal view in relation commenting on the original description of to other specimens, but he considered these S. vizottoi, that a specimen from Ilhe´us, state characteristics as geographical variations 74 JOURNAL OF HYMENOPTERA RESEARCH

Fig. 3. Stegomyrmex olindae n. sp., worker sting apparatus. A, sting and furcula in profile; B, sting and furcula in dorsal view; C, oblong plate; D, gonostylus; E, anal plate; F, spiracular plate; G, triangular plate; H, quadrate plate. and considered this specimen to belong to S. tumosity, and by the differences in the vizottoi. The study of the recently collected sting apparatus morphology. stegomyrmecine specimens deposited in Stegomyrmex olindae has been recorded in the MZSP collection, led us to recognize sparse localities in the Brazilian states of this specimen and several others as a Bahia, Maranha˜o, Minas Gerais, Mato different and undescribed species. Grosso, and Tocantins (central-north Bra- This species can be separated from the zil). The specimens are usually collected in related S. vizottoi by: the smaller size the leaf litter of mature wet forests. (Fig. 6), the nucal area and the whole Nothing is known about its natural history. mesosoma densely areolate-rugose, the The MZSP collection received recently metapropodeal groove bearing a median two stegomyrmecine males from Itabirito, VOLUME 17, NUMBER 1, 2008 75 state of Minas Gerais, Brazil. Despite the sculptured and the lateral surfaces mostly fact that these ants have been collected smooth and shining with a few sparse within the range of S. olindae, they are very foveae; anterior coxae, petiole, and post- similar to the males of S. vizottoi, differing petiole entirely and deeply areolate-rugose; only in discrete details of wing venation. legs opaque and weakly sculptured; sur- So, we decided not to include these males face of gaster finely and deeply foveolate. in the present study until we have addi- Pilosity cream-colored. Body covered by tional material from Minas Gerais. slightly stiffened, moderately clavate hairs, somewhat shorter in the external borders of Stegomyrmex vizottoi Diniz 1990 frontal lobes, antennal scapes, and legs; (Figs 4, 5, 6, 7) mandibles with long, sparse filiform setae; Stegomyrmex vizottoi Diniz, 1990: 290. Holotype posterior portion of ventral surface of head worker. BRAZIL: Sa˜o Paulo, Mirassol x.1971 and anteroinferior portion of pronotum (J.L.M. Diniz) (JLMD code 361) (MZSP code with short, curved, plumose hairs; occipital 11.029) [MZSP] (examined). Diniz and Bran- face of head and lateral surface of mesono- da˜o, 1993: 301 (biology). tum, metanotum, and propodeum glabrous. Stegomyrmex manni Smith, 1946: 288 (in part). Head relatively broad in frontal view. Lenko, 1965: 201 (distribution and biology); Vertexal margin slightly convex and with a Kempf, 1972: 242 (catalogue). discrete concavity medially; eyes with circa six facets at maximum diameter. Worker description.—Holotype (workers Promesonotum relatively elongate in 5 N 7); HW 1.26 (1.17–1.33); HL 1.07 (1.00– dorsal view, strongly convex dorsally in 1.14); ML 0.58 (0.51–0.61); SL 0.80 (0.76– lateral view; promesonotal suture distinct 0.85); WL 1.65 (1.59–1.77); PL 0.78 (0.70– only in the lateral faces of promesonotum; 0.80); PPL 0.46 (0.41–0.49); GL 1.60 (1.48– anepisternum set lower than the adjacent 1.70); TL 6.14 (5.72–6.48); CI 118.18 (116.67– surface; metapropodeal groove large, shal- 118.60); SI 63.46 (61.82–67.96). Dark brown lowly impressed and without median to black, with ferruginous appendages. projections; propodeal spines relatively Mandible finely and densely striate, with short, subtriangular, with the apexes di- large and sparse piligerous punctures, rected upwards and with the posterior except for the masticatory border and faces straight; propodeal spiracles weakly dorsum of apical portion which are smooth projected posterad; propodeal lobes round- and shining; inner surface of antennal ed and projected over the petiolar pedun- scrobes feebly striate and with punctation cle. In dorsal view, propodeum as narrow restricted to the region of antennal inser- as the promesonotum. tions; central disc of head and external Petiole elongate, slightly arched, with a margin of frontal lobes moderately areo- long rounded node; ventral carina of late; oblique lateral grooves of head, frontal peduncle with a well-developed anterior area and posterior portion of frontal lobes projection. Postpetiole globose, with the predominantly smooth and shining, with a dorsal face gently convex, without ventral few sparse punctures; anterior portion of projections. Gaster oval and robust. frontal lobes weakly areolate and with Sting apparatus (Fig. 5): Spiracular plate irregular longitudinal rugulae; antennae subquadrate, not extending towards the opaque and finely punctate; lateral and medial connection; margin of medial con- ventral surfaces of head deeply areolate; nection sclerotized; dorsal notch present; occipital face of head mostly smooth and spiracle moderately wide and placed close shining, except for the nucal collar which is to the posteroventral region of plate; uniformly scrobiculate; mesosoma partial- anterior apodema enlarged medially and ly foveolate, with the dorsum densely with an apical tubercle; ventral edge well 76 JOURNAL OF HYMENOPTERA RESEARCH developed. Quadrate plate with the dorsal SR extending distally beyond the stigma as a region broader than the ventral region, tubular vein for most of its length; M and Cu excluding the apodema; apodema area also extending distally, initially as tubular smaller than the plate body; dorsal margin veins and then as spectral veins almost flattened and sloped; anterodorsal corner reaching the distal wing border; anal vein of apex acute; posterior margin divided. not extending beyond CU cell. Hind wing Anal plate with the arc strongly sclero- with Sc+R extending shortly beyond the tized; apical margin triangular and well point where they connect to M, which definite; anal sensillae equally developed extends as a spectral vein to the wing distal and restricted to the posterior border of border; basally M+Cu does not continue as a plate. Oblong plate with short posterior tubular vein beyond the junction with 1M; apodema; subterminal tubercle acute api- anal vein drastically reduced; seven sub- cally; postincision well developed. Gonos- median hamuli. tylus one-segmented and with five chaetae Male.—(N5 4); HW 0.92–0.98; HL 0.80– subequal in length; terminal sector short 0.85; ML 0.21–0.24; SL 0.29–0.30; WL 1.89– and membranous, with dorsoterminal and 2.04; PL 0.90–0.95; PPL 0.41–0.49; GL 1.82– companion chaetae present. Triangular 1.89; TL 6.07–6.41; CI 114.29–115.15; SI plate as long as broad; only the median 30.00–31.58. Slightly smaller and slenderer tubercle is present. Lancets with functional than conspecific gynes. Color black with valves; sensorial barbles absent; distal appendages and gaster somewhat lighter. portion weakly sclerotized, probably not Integument opaque and densely areolate- perforating; dorsal and ventral margins rugose, except for the postpetiole and converging towards the apex; outer dorsal gaster, which are smooth and shining; wall absent. Sting shaft weakly sclerotized, appendages very finely punctate. Dorsum not perforating; dorsum of valve chamber of head and mesosoma densely covered by indistinct in profile; internal apophysis whitish, fine, suberect hairs, sparser over long and well sclerotized, extending along the dorsum of the metasoma; appendages the dorsum of valve chamber; basal con- with short, subdecumbent hairs. nection strongly concave; anterolateral Head broadest across large bulging processes well developed, narrowed medi- compound eyes (situated at the head ally, and as broad as the furcula lateral midlength) rather suddenly narrowed in arms; campaniform sensilla absent. Dorsal front of eyes and tapering moderately arm of furcula indistinct; lateral arms well anterad; median portion of vertexal margin developed; fulcral articulation connected weakly convex; occipital corners rounded; to the sting basis only by its lateral corners. ocelli prominent. Mandible relatively de- Gyne.—(N5 1); HW 1.46; HL 1.19; ML veloped, subtriangular, with slightly 0.65; SL 0.95; WL 2.00; PL 0.92; PPL 0.56; GL curved outer borders, rapidly converging 1.99; TL 7.31; CI 122.45; SI 65.00. Like in apical half; gently down curved; masti- conspecific worker, with the modifications catory border bearing circa seven serial expected for myrmicine gynes. Plumose teeth, with the apical tooth much more hairs restricted to the posteroventral corner developed than the others. Clypeus broad of head and inferior corner of pronotum. and truncate in front. Frontal lobes not so Compound eyes with circa 13 facets at developed as in the conspecific gynes and maximum diameter; propodeal spines re- workers, but concealing the antennal in- duced; posterior face of propodeum slightly sertions. Frontal carinae short and not inclined in side view, reaching the propo- expanded laterally, forming a short and deal lobes in rounded angles. Forewing with shallow antennal scrobe. Antennae long strongly colored stigma; longitudinal vein and slender with 13 segments; scape very Sc+R nebulous when reaching the stigma; short, only about twice as long as broad. VOLUME 17, NUMBER 1, 2008 77

Fig. 4. Stegomyrmex vizottoi Diniz, worker (SEM). A, head in full face view; B, close-up view of nucal area; C, habitus.

Prescutum with more or less distinct longitudinal rugae. Scutellum narrow pos- anteromedian carina; notauli shallow and terad. Metanotum narrow, with blunt me- complete, with transversal costulae. Para- dian tumosity. Propodeum with dorsal face psidial furrows as fine shining lines; parap- flat, steeply sloping posterad, and unarmed. sides more or less impressed behind, but Legs slender, middle and hind tibiae with- each with a sharp, raised posterolateral out apical spurs; tarsal claws slender and margin. Prescutellum separated from scu- simple. Wings brownish, with opalescent tellum by an impression bearing short bluish reflections; venation as in the gynes. 78 JOURNAL OF HYMENOPTERA RESEARCH

Fig. 5. Stegomyrmex vizottoi Diniz, worker sting apparatus. A, sting and furcula in profile; B, sting and furcula in dorsal view; C, oblong plate; D, gonostylus; E, anal plate; F, spiracular plate; G,triangular plate; H, quadrate plate.

Petiole clavate, with anterior peduncle smooth and shining, and propodeum distinct and long, low, rounded node; relatively narrow in dorsal view. Stegomyr- anteroventral projection vestigial. Postpe- mex vizottoi is the only species in the genus tiole as broad as long in dorsal view and for which both sexes and castes are known. slightly broader posteriorly than anteriorly, It has been registered from localities in attached to gaster by almost its full width. northern Argentina, Paraguay, and states Gaster somewhat elongate. of Santa Catarina, Parana´, and Sa˜o Paulo, Comments.—Thisspeciesisuniquely southeastern Brazil (Fig. 7). characterized by the combination of rela- Diniz and Branda˜o (1993) published tively large size (TL $ 6.00 mm), meso- observations on the foraging and nesting soma partially sculptured but for the habits of Stegomyrmex vizottoi from Miras- lateral faces which are almost entirely sol, Sa˜o Paulo, Brazil. They observed that VOLUME 17, NUMBER 1, 2008 79 workers are specialized predators of spir- the main chamber. It is interesting to note obolid millipede eggs and forage solitarily, that in the relatively mild winters in moving slowly in shaded areas, probing in southeastern Brazil, populations of colo- small cracks and cavities in the soil. The nies decrease sharply even in the laborato- ants seem to use their faces like a shovel, ry, and the ants close the nest’s entrance, tucking their antennae in the antennal relaying entirely on the millipede eggs scrobes and pushing soil away with the collected in the previous season for nutri- face. When a millipede egg is found, it is tion during the winter time, the eggs being grasped by the undersurface of the man- taken from the pile one at a time, to be dibles and pressed against the gular face of consumed by all individuals arranged in a the worker’s head, which then returns to circle. This helps to make these ants the nest. These ants are virtually unnoticed unnoticed for several months a year, to the naked eye, not only because they contributing to their rareness status. forage individually, but because of their slow movements, and because as they age Examined material [All deposited in MZSP].— their integument becomes covered by a BRAZIL: Parana´: Rio Azul (1.000 m) x.1959 (F. thin but hard layer of mud. Moreover, Plaumman) (1 paratype worker) (MZSP collec- workers can feign death for several min- tion n. 3147); Santa Catarina: Blumenau, P.E. utes when disturbed. Nascentes (27u069150S, 49u099140W) (Winkler Diniz and Branda˜o (1993) were the first samples) 30.iii.2001 (Silva, R.R. and Eberhardt, F.) (2 workers); Sa˜o Bento do Sul, APA Rio to examine a Stegomyrmex nest in detail. Vermelho (26u219510S, 49u169160W) (Winkler The S. vizottoi small and perfectly rounded samples) (Silva, R.R. and Eberhardt, F.) (8 nest entrance (0.4 cm in diameter) was in a workers and 1 gyne); Seara (24u079S, 52u189W) vertical soil bank, leading to a single (Winkler sample) v-xii.1998 (Roge´rio R. Silva) (1 sinuous tunnel extending about 40 cm to worker); Sa˜o Paulo: Anhembi, Faz. Barreiro a secondary chamber, in the roof of which Rico (in gizzard of Conopophaga lineata Wied, they found a funnel leading to the main 1831) ii.1964 (E. Dante) (MZSP collection n. chamber. Along the tunnel, which had 3470) (1 paratype worker); Mirassol (collected several small dead ends, they found three manually in soil) 10.x.1971 (Diniz, J.L.M.) enlargements, where the returning workers (MZSP collection n. 11029) (JLMD collection n. apparently stop to clean the eggs before 361) (1 paratype worker); xii.1976 (Diniz, J.L.M.) reaching the secondary chamber. The (JLMD collection n. 1226) (2 paratype gynes); cleaning process continues at the second- 13.ii.1987 (Diniz, J.L.M.) (MZSP collection n. 10924) (1 worker); same data (JLMD collection ary chamber, and the eggs are piled in the no. 544), (1 worker); (20u 509S, 49u 309W) (nest in main chamber only when completely soil) 11.xi.1991 (J.L.M. Diniz) (2 workers and 13 cleaned. The main chamber contained the males); Ribeira˜o Preto, Mata Santa Tereza, colony dealate gyne, its brood and a pile of 10.xii.1985 (C.G. Froelich) (1 worker). fully cleaned millipedes’ eggs. The second- ary chamber contained the millipede egg Accounts of the other Stegomyrmex shells, never found in the main chamber. species The total worker population in this partic- ular nest was 76. A second nest contained Stegomyrmex connectens Emery, 1912 some 300 workers, 22 alate gynes, seven Stegomyrmex connectens Emery, 1912: 51. Holo- dealate gynes, and brood. Colonies trans- type gyne. PERU: Vilcanota [MCSN] (not ferred to gypsum laboratory nests with examined). conditions and architecture similar to that of natural nests, adapted easily to the new Comments.—Stegomyrmex connectens is conditions and even constructed the fun- the type species of the genus and remains nel-like structure linking the secondary to known only by a single gyne collected in 80 JOURNAL OF HYMENOPTERA RESEARCH

Fig. 6. The relationship between head width and mesosoma length for the worker caste in S. vizottoi Diniz and S. olindae n. sp. The best-fitting regression line is plotted for each species. There is no significant difference for regression slope or elevation of the two lines. However, there is a significant difference between head width (F1,28 5 81.554, P 5 0.000) or mesosoma length (F1,28 5 163.75, P 50.000) between the species.

Vilcanota, Peru, and a male tentatively Stegomyrmex manni Smith, 1946 assigned to this species, from Mapiri, Stegomyrmex manni Smith, 1946: 288. Holotype Bolivia. In the original description, Emery worker. PANAMA: Canal Zone, Barro Colo- (1912) mentioned that the male might rado Island, ix.1941, (James Zetek) (Zetek belong to a different Stegomyrmex species, code 4879) (USNM code 57305) [USNM] (not although he decided to describe it as S. examined). Serna, 2002: 217 (first record for connectens. Diniz (1990) examined this Colombia). specimen and noticed that it presents some Stegomyrmex connectens Emery, 1912: 101 (in important morphological differences in part). Ho¨lldobler and Wilson, 1986: 16 (pilos- comparison to the conspecific gyne, mainly ity features). in wing venation, pilosity, and by the absence of a second anteroventral spine in Comments.—Stegomyrmex manni has been the petiolar peduncle. These differences, registered in the forest floor of mature and the disjunct distribution of the gyne rainforests in Costa Rica, Panama, and and male specimens, may indicate that Colombia. The combination of the relatively they indeed do not belong to the same high promesonotum in side view, propodeal species. spines directed posteriorly, and propodeal The gyne of S. connectens can be imme- spiracles strongly projected laterally sepa- diately recognized and separated from the rate this species from the other in the genus. other species in the genus by the presence Ho¨lldobler and Wilson (1986) illustrated of two anteroventral projections in the and discussed the special soil-binding petiolar peduncle. setae of S. manni (cited as S. connectens). VOLUME 17, NUMBER 1, 2008 81

Fig. 7. Distribution map for the known Stegomyrmex species.

Longino (2007) published very good the statistical tests and prepared Fig. 6; Rosa da Silva pictures of S. manni workers. He found and one of us (RMF) collected stegomyrmecine speci- mens in different expeditions supported by OIKOS specimens of S. manni in Winkler samples Pesquisa Aplicada Ltda; we would like to make a of sifted litter from the forest floor collected special reference to Dr. Fa´bio Olmos and Dr. Jose´ at Manuel Antonio National Park, Carara Fernando Pacheco, who arranged these trips. We would Biological Reserve, Pen˜ as Blancas east of also like to thank several individuals, institutions and Monteverde, and Hitoy Cerere Biological agencies that helped or supported other collecting trips that yielded material we studied. The present work was Reserve, all in Costa Rica. According to in part supported by the Conselho Nacional de him, Ronald Vargas collected an alate gyne Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq). at La Selva Biological Station. LITERATURE CITED

ACKNOWLEDGEMENTS Agosti, D., J. D. Majer, L. E. Alonso, and T. R. Schultz. 2000. Ants: Standard methods for measuring and We thank Renata Andrade for depositing in the monitoring . Smithsonian Institution MZSP ant collection the single known specimen of S. Press, Washington, DC. bensoni.LaraM.Guimara˜es took the SEM images and Bernard, F. 1951. Super-Famille des Formicoidea Gla´ucia Marconato made the Stegomyrmex bensoni Pp. 997–1104 in: Grasse´,P.P.Traite´ de Zoologie. drawing. Dr. Roge´rio Rosa da Silva kindly conducted Tome 10. Fasc. 2. Paris. 82 JOURNAL OF HYMENOPTERA RESEARCH

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