Kalathomyrmex emeryi, Klingenberg, C. & Brandão, C. R. F., 2009

Klingenberg, C. & Brandão, C. R. F., 2009, Revision of the fungus-growing ant genera Mycetophylax Emery and Paramycetophylax Kusnezov rev. stat., and description of Kalathomyrmex n. gen. (Formicidae: Myrmicinae: Attini)., Zootaxa 2052, pp. 1-31 : 22-25

Kalathomyrmex emeryi (Forel, 1907) new combination

(Figs. 4, 6 h, i, 7 b)

Myrmicocrypta emeryi Forel , 1907:144 (worker) Syntypes, Colombia, Dibulla, Santo Antonio, Forel col. ( MCSN, examined); Forel 1912: 189 (queen, male) Allotypes; Emery 1913: 251 combination in Cyphomyrmex (Mycetophylax) ; Santschi 1916: 383 combination in Myrmicocrypta (Mycetophylax) ; Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue).

Mycetophylax hummelincki Weber , 1948: 84 (worker) Syntypes, Venezuela, Stat. 121, Cabo Blanco, 19.viii.1936 (P. Wagenaar Hummelinck col.) (not examined); Weber 1958: 263 synonym of Mycetophylax emeryi ; Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue).

Myrmicocrypta emeryi var. arenicola Forel , 1912: 189 (worker, queen) Synytpes, Argentina, Huasan, 1300 m (Bruch col.) (not examined); Emery 1922: 343 combination in Cyphomyrmex { Mycetophylax ); Santschi 1922: 355 combination in Myrmicocrypta (Mycetophylax) ; Kempf 1962: 34 combination in Mycetophylax ; Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue) Bestelmeyer & Wiens 1996: 1231 (ecology); new synonym.

Myrmicocrypta (Mycetophylax) emeryi var. argentina Santschi , 1916: 383 (worker) Syntypes, Argentina, Santiago del Estero, Rio Salado, (Wagner col.), Buenos Aires, (Bruch col.) ( NHMB, examined); Emery 1922: 343 combination in Cyphomyrmex (Mycetophylax) ; Kempf 1972: 146 combination in Mycetophylax ; Bolton 1995: 268 (catalogue); new synonym.

Myrmicocrypta emeryi var. fortis Forel , 1912: 189 (worker) Syntype, Argentina, Huasan, 1300 m (Bruch col.) (MNHB, examined); Santschi 1922: 355 combination in Myrmicocrypta { Mycetophylax ); Bucher, 1974: 63; Kempf 1972: 146 in Mycetophylax ^catalogue); Bolton 1995: 268 (catalogue); new synonym.

Mycetophylax bolivari Weber , 1948: 84 (worker) Syntypes, Venezuela, Anzoategui: Llanos 17 km N of Soledad, 27.i.1935, across the Orinoco River from Ciudad Bolivar (Weber col.), (not examined); Weber 1958: 263 ( Mycetophylax emeryi ssp. bolivari ), Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue); new synonym.

Myrmicocrypta (Mycetophylax) emeryi st. gallardoi Santschi , 1922: 354 (worker) Syntypes, Argentina, Province de Buenos-Ayres, Sierra de la Ventana (Bruch, leg.) ( NHMB, examined); Kempf 1972: 146 combination in Mycetophylax (catalogue); Brandao, 1991: 358 (catalogue); Bolton 1995: 268 (catalogue); new synonym.

Mycetophylax emeryi st. hubrichi Santschi , 1925: 163 (worker) Syntypes, Argentina, Santa Fe, Rosario, (Hubrich, col.) ( NHMB, examined); Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue); new synonym. Mycetophylax emeryi st. weiseri Santschi , 1929: 303 (worker) Syntypes, Argentina, Catamarca, Corral Quemado, (Weiser, col.) ( NHMB, examined); Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue); new synonym.

Mycetophylax glaber Weber , 1948: 85 (worker) Holotype, Bolivia, Tumupasa, (W.M. Mann col.) (not examined); Kempf 1972: 145 (catalogue); Bolton 1995: 268 (catalogue); new synonym. Worker (Figs. 4 a, b, g, 6 h and 7 b)

Range of measurements (in mm) and indices of examined specimens (N = 76): IOD 0.49-0.83; HL 0.54-0.83; CI 85-106; SL 0.48-0.8; SI 84-112; ML 0.23-0.48; MI 34-77; WL 0.70-1.26; PrW 0.26-0.60; PL 0.16-0.30; PPL 0.19-0.35; GL 0.57-0.92; FL 0.52-0.98; TL 2.42-4.03.

Color in general yellowish to reddish-brown, some specimens may be darker. Apex of funiculus, clypeus, and masticatory border of mandibles, head vertex, postpetiole, gaster and femora brownish. Mandibles and tarsi yellowish; rest of the body light reddish brown. Under optical scope, body sculpture densely punctuated with exception of lateral parts of pronotum, where the sculpture is more superficial. Mandible disc shiny with piliferous punctuations. Mesosoma covered by a fine layer of "dirt", visible only in SEM images in a way that the sculpture of the integument is "reprinted" in the dirt-layer. Whole body covered by shiny whitish to golden appressed hairs. Long, flexuous hairs covering the mandibles and gular face.

Head shape quadrate (see CI). Mandibles with five teeth, apical tooth larger, followed by smaller second and third teeth. After the diastema a small fourth tooth followed by a small denticule. Anterior margin of clypeus gently concave. In lateral view, clypeus triangular. In frontal view latero-posterior margin of clypeus strongly produced forwards over the lateral wings of the clypeus, as rounded trenchant ridges, resulting in two large flat circular areas where the antennal scapes articulate. Median portion of clypeus attains posteriorly the posterior level of the antennal insertions in a generally straight sometimes rounded suture, followed by a small impressed triangular frontal area. A shallowly impressed median line running from the frontal area to the vertex. Frontal lobes reduced, barely covering the antennal insertions, their maximum expansion less than a fourth of a longitudinal median head axis and the lateral margins of head, ending posteriorly at the level of the posterior margin of compound eyes. Lateral carinae bordering the internal margins of the compound eyes, fading out a little after their posterior margins. Compound eyes set slightly before the middle of the head, at maximum width with ten ommatidia and at maximum length with 14 ommatidia. Vertexal margin straight, but occasionally with a median impression. Antennae with flattened scapes, surpassing the posterolateral corners of the head when laid back over the head capsule. First funicular segment as long as the second and the third together. Apical end of funiculus with a three segmented club, only a little wider than the other funicular segments. Ventral face of head flat.

Mesosoma. Pronotum without tubercles or distinct protuberances, lateral pronotal margins rounded, without spines or angles. Occasionally the pronotum bearing a blunt obliquely directed spine (specimens from Paraguay and Argentina). Dorsal face of mesonotum evenly rounded, in side view, followed by a small depression and a conical low protuberance. Anepisternum clearly divided from the mesonotum by a carina. In lateral view, basal face of propodeum slightly convex anteriorly, meeting the concave declivous face and about one half shorter than basal face. Propodeal spiracle opening in an angle of 45° in relation to the main body axis. Propodeum with a pair of divergent, short, blunt obtuse angles, directed obliquely upwards. In dorsal view, petiole straight, wider at three fourths of its length, with a vestigially developed ventral process. Postpetiole, in dorsal view, subtriangular, with a large impression at posterior margin, forming two distinct lobes, heart-shaped and dorsoventrally flattened.

Gaster smooth, without protuberances or carinae.

Gyne (Figs. 4 c, d, h and 7 b)

Range of measurements (in mm) and indices of examined specimens (N = 10): IOD 0.54-0.64; HL 0.56-0.63; CI 90-107; SL 0.5-0.56; SI 83-100; ML 0.26-0.34; MI 43-61; WL 0.94-1.02; PL 0.16-0.24; PPL 0.2-0.24; GL 0.84-0.98; TL 3.01-3.36.

Color, pilosity and main morphological traits of head, propodeal spiracle, petiole, postpetiole and gaster similar as in the conspecific workers. Mandible with five teeth, apical tooth bigger than all the others, followed by smaller second and third triangular teeth; fourth tooth triangular and smaller than the second and third, followed by a small denticle. Compound eyes with 15 ommatidia at maximum width and 17 ommatidia at maximum length. Posterior portion of head with three ocelli, the median superficially impressed at frons. The apical funicular segment as long as the three anterior together.

Mesosoma in lateral view with dorsally flattened scutum, in lateral view, dorsum of scutum starts at the anterior third of the pronotum. In dorsal view anterior margin and posterior margin round, the last ending in a carina. Parapsidial lines shallowly impressed. Prescutum reduced, represented only by triangular axillae; scutum-scutellar sulcus deeply impressed. Scutellum subquadrate, anterior third wider than posterior portion, posteriorly rounded. Metanotum reduced, appearing only as small, flattened disc in dorsal view. Katepisternum rectangular; anepisternum only half the size of the katepisternum, subquadrate, and both separated by a distinct groove, ending posteriorly in a carina. Propodeum with a small blunt obliquely upwards directed spine.

Male (Figs. 4 e, f, i, 6 i and 7 b)

Range of measurements (in mm) and indices of examined specimens (N = 10): IOD 0.21-0.34; HL 0.24-0.34; CI 88-100; SL 0.32-0.42; SI 110-152; ML 0.13-0.18; MI 38-53; WL 0.76-0.92; PL 0.10-0.18; PPL 0.12-0.20; GL 0.68-0.82; TL 2.31-2.58.

Color brownish to dark brown. Anterior half of mandibles, funiculus, labrum, legs and apex of gaster light yellow. Under optical scope, body densely punctuated, with exception of shiny gaster, where sculpture is more superficial. Whole body covered by golden, shiny, appressed hairs. Head shape quadrate (see CI). Mandibles with three multidenticulate teeth, apical tooth longer than the others, followed by a smaller second triangular tooth and a denticle, better visible with SEM images (Fig. 6 i). Anterior margin of clypeus straight. Clypeus with a psammophore composed by four setae arising from the meeting of the clypeus and the anteclypeus. Psammophore hairs fine and stiff as long as the length of the two apical segments of funiculus, reaching the apex of the mandibles. In frontal view, clypeus bulging, its posterior margin not forming a trenchant ridge. Frontal lobes reduced, covering only half of the antennal insertions, ending posteriorly at the level of the anterior margin of compound eyes. Lateral carinae marginate the compound eyes anteriorly. Compound eyes set at anterior part of head, at maximum width with 19 ommatidia and at maximum length with 23 ommatidia. Vertexal margin with a median longitudinal impression, resulting from the bulging of the lateral ocelli, posterolateral corners of head rounded. Antennae 13-segmented, scapes rounded, surpassing the posterolateral corners of the head by the length of the three apical segments of the funiculus. First funicular segment with the same length of the second. Funiculus with a three segmented club. The apical funicular segment as long as the two anterior together. Ventral face of head flat.

Mesosoma in dorsal view with rounded scutum, almost covering the whole pronotum. Major width of scutum at tegula. Parapsidial lines distinct and in parallel to the main body axis. Axillae subtriangular and scutum-scutellar sulcus deeply impressed. Scutellum subquadrate to subtriangular, wider anteriorly than posteriorly, posterior margin rounded. Katepisternum subquadrate with inferior margin rounded; anepisternum subtriangular. Apex of procoxa fail to attain the anterior margin of katepisternum level. Propodeum basal face occasionally slightly concave, meeting the declivous face in rounded angles. In lateral view, petiole triangular, node rounded dorsally, ventrally straight. In dorsal view, petiole with a shallowly and wide median impression. Postpetiole much wider posteriorly, heart-shaped; ventrally with a vestigial antero-median denticle. Gaster elongated.

Legs long and filamentous.

Examined material: ARGENTINA: Buenos Aires, Sierra de la Ventana, no coll. date (C. Bruch), 3 w ( NHMB); Catamarca, no coll. date. (Coronel Weiser), 2 w ( NHMB) (Types); Chaco, Chaco National Park, 26°48,522' S 59°36.395' W, 4-6.iv.2003, Scott Solomon col., 2 w, 2 q, 2 m ( SMNK); Cordoba, no coll. data (C. Bruch), 2 w, 1 q ( MZSP); Cruz Alta, 30.v.1965 (E. Bucher), 1 w ( MZSP), Mendoza, Chileiro, no coll. date (Duzione), 3 w ( NHMB)(Cotype), Santiago del Estero, no. coll. date (Merkle), 1 w ( MZSP); Chaco de Santiago del Estero, Rio Salado, no coll data, 1 w ( NHMB) (Holotype of M. emeryi argentina ); Rosario, no coll. date (Hubrich), 1 w ( MZSP), 3 w ( NHMB); La Soledad (Canete), Tucuman: no coll. date. (Weiser), 1 w; 5.xii.1956 (NK 10018) [Nicolas Kusnezov], 3 q ( MZSP); Siete de Abil, Dpto. Burruyacu, 8.vi.1965 (E. Bucher) 2 w ( MZSP); BRAZIL: Amazonas: Manaus, Praia de Tupe (Rio Negro), 3.iii.2002 (C. Rabeling), 32 w ( MZSP), 20.iv.2002 (C. Klingenberg), 14 w ( MZSP); Praia Paricatuba (Rio Negro), 20.iv.2002 (C. Klingenberg) 4 w ( MZSP); Bahia: Juazeiro, x-8.xii.1948 (C. R. Goncalves), 5 w, 1 q ( MZSP); Mato Grosso: Porto Murtinho, vii.1960 (B. Kelber), [4680, 3803], 3 w ( MZSP); Rondonopolis, 6.iii. 1971 (W. Kempf), [6263], 7 w ( MZSP); Tocantins: Goiatins, 9.xi.1999 (C. R. F. Brandao & C. I. Yamamoto), 1 w ( MZSP); Para: Alter do Chao, 2°30'S 54° 57W, 15.vii.1998 (M. F. Leite), 2 w (Heraldo Vasconcelos); Paraiba: Juazeirinho, Soledade, 28.i.1956 (C. R. Goncalves), 4 w ( MZSP); Pernambuco: Araripina, 2-4.i.1973 (R. Montenegro), [8424, 8426, 8434, 8445], 8 w, 2 q ( MZSP); Piaui: Canto do Buriti, 18-22.xi.1991 (C. R. F. Brandao) 23 q, 17 m; 20 km S Floriano, Buriti Sol, 5-12.xi.1991 (C. R. F. Brandao & P. Moutinho), 10 km N Corrente, Faz. Maracuja, 23-27.xi.1991 (C. R. F. Brandao) 1q, 1m ( MZSP); Rio Urucui-Preto, 20.ii.1976 (R. Negrett), [12857], 3 w ( MZSP); Sao Paulo: Sao Manoel, 10.ii, 6.xi, 8.xii.1988, (E. S. Zanetti), 2 q, 1 w ( MZSP); Agudos, ix, 10.x.1958 (R. Mueller), [2752, 2661], 4 w ( MZSP), Agudos, 26.xi.1955 (W. Kempf), 2 w, 1 q ( MZSP); Itirapina, 18.iii.1966 (W. Kempf), [4422], 1 w ( MZSP); GUYANA: Pirara, nest in soil, 4.iv.1996 (Ted Schultz & U. G. Mueller), 4 w [#00303800] ( USNM). PARAGUAY: Boqueron: Parque Nacional Tte. Enciso, Zona Administrativa, 21°13'S 61°40'W, 6-7.viii.1994 (B. Garcete), 1 w ( MZSP); PERU: Huanuco ("Panguana"), Rio Yuyapidris, 16.xii.1984 (M. Verhaagh), 3 w ( SMNK), VENEZUELA: Lara, Barquisimeto to Carora km 19, 29.vi.1971 (W. L. Brown), 15 w ( MZSP).

Comments. Authors often based their descriptions of subspecies and varieties of Kalathomyrmex emeryi on morphological character differences regarded today as minor local variations. It is well known that characters like color and pilosity in general vary within the same ant species and colony. Examples of this usage can be seen in Forel (1912) in his descriptions of Myrmicocrypta emeryi var. arenicola and M. emeryi var. fortis . The two variations differ only by their color, but were collected at the same locality. This may suggest that these taxa are composed of sibling species, but the present criteria do not afford recognition. Another example is given by Santschi (1922), when he states that M. emeryi var. fortis seems to be a transitional form between arenicola and argentina . These examples indicate that the validity of the majority of the described subspecies and variations of K. emeryi is doubtful. The examination of all listed specimens, including the available types, shows that there are variations in color and pilosity, but mostly gradual and sometimes aleatory, and thus we found no ground for a reliable recognition of species or subspecies among then. Although we were not able to locate and examine the type of M. glaber , the description of the species given by Weber (1948) led us to conclude that M. glaber is also identical to K. emeryi .

Information on the biology of this species is scant. As mentioned above, Bucher (1974) described the nest architecture of Paramycetophylax bruchi and K. emeryi ; both species inhabit sandy soil and prefer places devoid of vegetation. We confirmed this observation for K. emeryi , as we found this species nesting at the beaches of the Rio Negro (AM), Brazil. The nests are quite common at the sandy beaches of the river and can be easily located. In the Amazonian rainy season, the nests become covered by water for months. This observation has been confirmed by Dr. M. Verhaagh in Peru (pers. comm.), who found a K. emeryi nest nearby a river bank covered by high water for several days; when the river returned to its normal water level, the ants reopened the entrance and came out of the nest. According to Bucher (1974), the fungus chambers are located from 60 to 100 cm deep, but can be found deeper due to temperature changes during the year. For the fungus substrate, the species forages for feces of other insects, mainly of Lepidoptera. The ants showed their highest activity during the night. However, we observed K. emeryi in frantic activity during the day in several instances, even with relatively high temperatures and under full sun exposure.

Kalathomyrmex emeryi is distributed all over cis-Andean South American. A map of the known records is presented in Fig. 7 b.