ࡱ>  []RSTUVWXYZ5@ =Pbjbj22 ~xXXGE42 6!6!6!6!T!D2 D!6####&:&& 3D5D5D5D5D5D5D$XFRHYD5%&55YD##nDAAA5H&##CA53DALAC(C#! TMeJ6!:C?CDD0DCDI>DIC2 2 DIC8&+tA+/1&&&YDYD2 2 6!A2 2 6!The ants of North America: the genus Mycocepurus (Hymenoptera: Formicidae) William P. Mackay1 *, Jean-Michel Maes2, Patricia Rojas Fernndez3 and Gladys Luna2 1Centennial Museum, Department of Biological Sciences, The University of Texas, El Paso, TX 79968 * Correspondent:  HYPERLINK "mailto:wmackay@utep.edu" wmackay@utep.edu 2Museo Entomolgico, Apartado Postal 527, Len, Nicaragua,  HYPERLINK "mailto:jmmaes@ibw.com.ni" jmmaes@ibw.com.ni 3Departamento Biologa de Suelos, Instituto de Ecologa A.C. CP. 91000, Apartado Postal 63 Xalapa, Veracruz,. Mxico Submitted xx October 2003, Accepted XXX, Published XXX _______________________________________________________________________ Abstract We provide a review of the North American ants (north of Colombia) of the ant genus Mycocepurus, including keys to the workers and females, illustrations and distribution maps. The distribution of M. tardus is extended to Nicaragua and Costa Rica. The female of M. curvispinosus is described. Resumen Se revisan las especies del gnero Mycocepurus de Norte Amrica (al norte de Colombia). Se incluyen claves para la identificacin de las obreras y las hembras, ilustraciones y mapas de distribucin. Se amplia hacia el norte la distribucin de M. tardus, incluyendo ahora Nicaragua y Costa Rica y se describe la hembra de M. curvispinosus. Keywords: fungus-growing ants, tribe Attini, tropical rain forest, Neotropics, Atta sexdens ________________________________________________________________________ Introduction Ants of the genus Mycocepurus are among the most easily recognized among the fungus-growing ants of the tribe Attini. They are monomorphic ants, characterized by the numerous spines on most bodily surfaces, including a pair of occipital spines (Fig. 1), and usually 2 - 3 pairs on the pronotum, 5 - 6 pairs on the mesonotum and 2 pairs on the propodeum (Fig. 3), with the last pair being the most developed. The dorsum of the petiole has 2 pairs of spines (Fig. 6). There is a definite circlet or crown of spines on the promesonotum (Fig. 2). No other genus in the New World has this crown of spines on the promesonotum. This genus is a member of the tribe Attini, a monophyletic group of fungus-growing, predominantly Neotropical ants (Chapela et al. 1994). It apparently occupies a divergent phylogenic position, based on its unusual mandibular gland secretion (Blum et al. 1981). The genus Mycocepurus uses a variety of organic matter as a substrate for their fungal gardens, ranging from dry leaves and caterpillar dung (Kempf 1963) to fruit matter on the seeds of Hymenaea courbaril (Caesalpiniaceae), which facilitates germination of the plant (Oliveira et al. 1995).  Figure.  SEQ Fig. \* ARABIC 1. Head (frontal view) of a worker of M. curvispinosus (modified from Mackay, 1998). The genus consists of a small group (5 species) of cryptic ants, which are found from Mxico south to Brazil and Argentina (Kempf, 1963, Mackay, 1998). Two of the species are relatively common and widely distributed from Mxico to Brasil (M. goeldii Forel and M. smithii Forel), one is known from Mxico to Panam (M. curvispinosus Mackay), another from Nicaragua south to Panam (M. tardus Weber), and the remaining species is presently known only from the type locality (M. obsoletus Emery from Santarm, Par, Brazil).  Figure  SEQ Fig. \* ARABIC 2. Pronotum and mesonotum (top view) of a worker of M. smithi, showing the crown of spines on the promesonotum (modified from Mackay, 1998).  Figure  SEQ Fig. \* ARABIC 3. Mesosoma (side view) of a worker of M. goeldi, showing the variety of spines. Results The following key will separate the workers of the species of Mycocepurus: 1. Pronotum with pair of anterior pronotal spines (Fig. 3), which defines anterior border of crown of spines on promesonotum (Fig. 3); propodeal spines usually straight (Fig. 8) .. 2  Figure  SEQ Fig. \* ARABIC 4. Promesonotum (top view) of a worker of M. goeldii (modified from Mackay, 1998).  Figure  SEQ Fig. \* ARABIC 5. Promesonotum (top view) of a worker of M. curvispinosus (modified from Mackay, 1998). - Pronotum lacking pair of anterior pronotal spines (Figs. 5 and 6); propodeal spines strongly curved upwards (Fig. 6) . .... curvispinosus Mackay Figure  SEQ Fig. \* ARABIC 6. Mesosoma and petiole (side view) of the holotype worker of M. curvispinosus (modified from Mackay, 1998). 2(1). Pair of promesonotal spines present in center of crown (Fig. 7) ... 3 Figure  SEQ Fig. \* ARABIC 7. Mesosoma of a worker of M. smithii as seen obliquely from the side. - Promesonotal spines lacking in center of crown, although raised area or low crest may be present (Fig. 2) . 4 3(2). Inferior lateral pronotal spine poorly developed or absent (Fig. 9); Central America ... tardus Weber - Inferior lateral pronotal spine well developed (Fig. 3); Brazil and Argentina ... goeldii Forel 4(2). Promesonotal spines low and blunt (Fig. 8); known only from the type locality in Brazil (Par: Santarm) . obsoletus Emery  Figure  SEQ Fig. \* ARABIC 8. Mesosoma (side view) of the holotype worker of M. obsoletus (modified from Emery, 1913). - Promesonotal spines high and sharp (Fig. 7); widely distributed (Mxico: Jalisco, Nayarit, Tabasco south throughout Central America to Brazil) smithii Forel Figure  SEQ Fig. \* ARABIC 9. Mesosoma (side view) of a worker of M. tardus. The following key may separate the females of the species of Mycocepurus (the females of M. tardus and M. obsoletus are unknown, and the characters are extrapolated from the workers): 1. Mxico, Central America, and northern South America . 2 - Brasil and Argentina 4 2(1). Propodeal spines slightly upturned (Fig. 10); lateral pronotal spine large and thick (Fig. 10); inferior lateral tooth tiny .. curvispinosus Mackay  Figure  SEQ Fig. \* ARABIC 10. Mesosoma, petiole and postpetiole (side view) of a female of M. curvispinosus (Ro Frijolito, Panam, CWEM). The inset shows the scutellum and propodeum as seen from above. - Propodeal spines straight or slightly bent downward (Figs. 11, 12) . 3 3(2). Common and widely distributed; propodeal spines slightly bend downward (Fig. 11); lateral pronotal spine poorly developed (Fig. 11); inferior lateral pronotal spine tiny smithii Forel  Figure  SEQ Fig. \* ARABIC 11. Mesosoma (side view) of a female of M. smithi (Parque Nacional Soberana, Panam, CWEM). - Rarely collected, but widely distributed; propodeal spines probably straight (female unknown) . tardus Weber 4(1). Inferior lateral tooth well developed, about same size as lateral pronotal tooth (Fig. 12) ... goeldii Forel Figure  SEQ Fig. \* ARABIC 12. Mesosoma (side view) of a female of M. goeldii (Rio Claro, Brasil, CWEM). - Inferior lateral pronotal tooth poorly developed, obsolete or at least much smaller than lateral pronotal tooth (Fig. 11) .. 5 5(4). Common and widely distributed (Mxico south to Brasil; Caribbean); lateral pronotal spine moderately well developed (Fig. 11) ...... smithii (Forel) - Known only from the state of Par, Brasil; lateral pronotal spine probably poorly developed or even absent .... obsoletus Emery Mycocepurus curvispinosus Mackay Figs. 1, 5, 6, 10; Map 1 Mycocepurus curvispinosus Mackay, 1998:423-424 + Figs. 5, 6 & 9, Costa Rica, Loma Barbudal, worker Discussion The worker of this species is easily recognized, as it lacks the anterior pronotal spines (Fig. 6). Additionally, the propodeal spines are strongly curved upwards (Fig. 6). No other species has this combination of characters. The female can be recognized by the large lateral pronotal spines, and the slightly upturned propodeal teeth (Fig. 10). It is relatively small (total length slightly more than 2.5 mm), smaller than the females of M. smithii. The first two characteristics easily separate this species from M. smithii and would probably separate it from the unknown females of M. tardus. Description of female (mm): Head length (HL, anterior edge of median lobe of clypeus to midpoint of posterior margin) 0.71, head width (HW, maximum, at posterior edge of eye) 0.67, scape length (SL, excluding basal condyle) 0.58, eye length (maximum) 0.17, Webers length (anterior border of pronotum to posterior angle of metapleuron) 1.07, cephalic index (HL/HW X 100) 95, scape index (SL/HL X 100) 81. Mandible apparently with 5 teeth; frontal lobes expanded, covering insertions of antennae; sides of head nearly parallel, slightly narrowed in region of eyes, which extend past sides of head; posterior margin of head concave, occipital spines poorly developed; scape barely reaches posterior lateral corner; lateral pronotal spine well-developed, with wide base, inferior lateral pronotal spine poorly developed; scutellum angulate posteriorly, overhanging metanotum; propodeal spines well-developed, slightly upturned, with broad base; subpeduncular process poorly developed, peduncle elongated, apex of petiole with two sets of spines. Anterior margin of clypeus with several, erect hairs, dorsum of head with several, erect hairs, hairs on scape decumbent, dorsum of mesosoma with erect hairs, petiole nearly lacking erect hairs, postpetiole and gaster with abundant, erect hairs, hairs on tibiae mostly suberect. Entire ant very roughly sculptured, except for mandibles, which are striate and moderately shining, much of rough sculpture, especially on dorsum of gaster, arranged in poorly defined striate. Ferrugineous red. The single female specimen is deposited in the collection of William and Emma Mackay (CWEM). Distribution Mxico (Veracruz: Reserva Ecolgica La Mancha, Chiapas: 24 k SW Cintalapa), Costa Rica (Heredia: Loma Barbudal); Panam (Coln, Ro Frijolito in El Parque Nacional Soberana, female). Map  SEQ Map \* ARABIC 1. Mycocepurus curvispinosus. Habitat Found in a variety of communities, ranging from slashed and burned areas, sub-deciduous forests to tropical rain forests. Biology Most specimens have been collected in pitfall traps, or litter samples. This species lives in nests of M. smithii in Mxico (La Mancha, Veracruz). The workers are slow and timid and forage together with those of M. smithi. A single, dealate female was collected in Panam on 18-v-1995. Mycocepurus smithii (Forel) Figs. 2, 7, 11; Map 2 Atta (Mycocepurus) smithii Forel, 1893:370, Antilles, worker Trachymyrmex attaxenus Eidmann, 1936:85, worker, female (Kempf, 1963:425) Mycocepurus bolivianus Weber, 1938:155 (Kempf, 1963:425) Atta (Mycocepurus) smithii var. borinquenensis Wheeler, 1907:718 (Kempf, 1963:425) Mycocepurus smithii var. eucarnitae Forel, 1913:235 (Kempf, 1963:425) Mycocepurus manni Weber, 1938:156 (Kempf, 1963:425) Mycocepurus reconditus Borgmeier, 1937:246 (Kempf, 1963:425) Atta (Mycocepurus) smithii var. tolteca Wheeler, 1907:718 (Kempf, 1963:425) Mycocepurus smithii var. trinidadensis Weber, 1937:378 (Kempf, 1963:425) Discussion This is one of the two species, in which the workers lack the well-developed promesonotal spines in the center of the crown (Fig. 2). Generally there is a very low, sharp crest in this region. In most cases, it can be separated from the other species that lacks these spines, the Brazilian M. obsoletus, on the basis of the distributions. Otherwise, it can be separated by the lateral pronotal spines, which are long and sharp (Fig. 7), not short and blunt is in M. obsoletus (Fig. 8). Specimens that have bumps in the center of the crown of spines, could be confused with M. tardus, but can be usually separated as M. tardus has definite promesonotal spines (Fig. 9). The females have small lateral pronotal spines, and tiny inferior lateral pronotal spines (Fig. 11). The females are slightly larger than those of M. curvispinosus (~ 3 mm total length). Additionally the propodeal spines are bent slightly downward (Fig. 11). These characters would easily separate it from females of M. curvispinosus, but may not work to separate it from the unknown females of M. tardus, which occur in the same areas. Distribution Mxico (Jalisco: 39 k N Colima; Nayarit: Tepic, 54 k S Rosamorada; Tabasco: 10 k N Crdenas; Veracruz: Reserva Ecolgica La Mancha, 26 k S Puente Tampico, Arroyo Seco Acayucn), Honduras, Costa Rica, Panam (Parque Nacional Soberana near Gamboa), Colombia (Valle: Buga, Sevilla), Cuba, Puerto Rico (Ro Grande at El Verde Field Station), Per, Trinidad, Guianas, Bolivia (Ro Ben), Brasil (Par, Bahia, Mato Grosso, Goias, Minas Gerais, Rio de Janeiro, So Paulo: Ilha dos Buzios, Esprito Santo), Argentina (Formosa). Map  SEQ Map \* ARABIC 2. Mycocepurus smithi. Habitat Ranging from open, disturbed areas, burned rain forest, sub-deciduous tropical forests, to tropical rain forests, and moist gullies. Biology Kempf (1963) reviewed the biology of this species. It nests only in the soil (Torres 1989), often with the entrance surrounded by a turret. Specimens in Mxico are often found nesting under stones. The nest cavity is very small, only a few cm in diameter, and is connected with several fine, threadlike, tunnels. The nest may be as deep as a meter. There are often several nests in a single area, which suggests a single large nest with multiple entrances. The colonies are polygynous. The small, sluggish workers carry dry leaves, caterpillar dung (Kempf, 1963) and bat dung (Levins et al. 1973) back to the nest, where they culture an unidentified fungus, which is almost identical to that used by the fungus-growing ant Cyphomyrmex minutus (Torres 1984). Populations vary greatly over time, and they may be abundant in some areas (Torres 1989, Majer, 1997). This species may be associated with the leaf-cutting ant Atta sexdens, and is often found in the same habitat as M. goeldii, with which it has mixed mating swarms, although hybridization apparently does not occur. This species tolerates a number of other species in its nest, including M. curvispinosus, Solenopsis sp. (thief ants) and Acropyga sp. It forages together with M. curvispinosus. Mycocepurus tardus Weber Fig. 8; Map 3 Mycocepurus tardus Weber, 1940:416, Panama, worker Discussion The worker can be recognized by the pair of promesonotal spines located in the middle of the circlet of teeth on the promesonotum (Fig. 9). It can be separated from the other two species (M. curvispinosus and M. goeldii) which also have these teeth, based on several different characters. It differs from the Mexican and Central American M. curvispinosus, as the anterior pronotal spines (Fig. 3) are well developed (lacking in M. curvispinosus - Fig. 6), and propodeal spines are straight (Fig. 9), and directed somewhat vertically. It can be easily separated from the Brazilian and Argentinean M. goeldii on the basis of the distributions, and in that the inferior lateral pronotal tooth is poorly developed (well-developed in M. goeldii). Distribution Nicaragua (Zelaya: Bluefields), Costa Rica (Puntarenas: Cerro Helado), south to Panam (Panam: Barro Colorado Island).  Map  SEQ Map \* ARABIC 3. Mycocepurus tardus. Habitat Lowland rain forest. Biology Unknown. Acknowledgments We thank Robert Anderson for the gift of several of the specimens. Emma Mackay critically reviewed the manuscript. Literature cited Blum, M, Brand J and Amante E. 1981. o-aminoacetophenone: identification in a primitive fungus-growing ant (Mycocepurus goeldii). Experientia 37:816-817. Borgmeier T. 1937. Formigas novas ou pouco conhecidas da Amrica do sul e Central, principalmente do Brasil (Hym. Formicidae). Archivos do Instituto de Biologia Vegetal, Rio de Janeiro. 3:217-255 + 5 plates. Chapela I, Rehner S, Schulz T and Mueller U. 1994. Evolutionary history of the symbiosis between fungus-growing ants and their fungi. Science 266:1691-1694. Eidmann H. 1936. kologisch-faunistische Studien an sdbrasilianischen Ameisen. Arbeiten ber Physiologische und angewandte Entomologie aus Berlin-Dahlem 3:81-114. Emery C. 1913. tudes sur les Myrmicinae. V. Les genres des Attini; descriptions de nouvelles formes de Mycocepurus et de Myrmicocrypta. Annales de la Socit Entomologique de Belgique 57:250-262. Forel A. 1893. Formicides de l'Antille St. Vincent. Rcoltes par Mons. H. H. Smith. Transactions of the Entomological Society of London 1893:333-418. Forel A. 1913. Fourmis d'Argentine du Brsil, du Guatemala & de Cuba reues de M. M. Bruch, Prof. v. Ihering, Mlle Bez, M. Peper et M. Rovereto. Bulletin de la Socit Vaudoise des Sciences Naturelles 49:203-250. Kempf W 1963. A review of the ant genus Mycocepurus Forel, 1893 (Hymenoptera: Formicidae). Studia Entomologica 6:417-432. Levins R, Pressick M and Heatwole H. 1973. Coexistence patterns in insular ants. American Scientist 61:463-472. Mackay W. 1998. Dos especies nuevas de hormigas de la tribu Attini de Costa Rica y Mxico: Mycetosoritis vinsoni y Mycocepurus curvispinosus (Hymenoptera: Formicidae). Revista de Biologa. Tropical 46:421-426. Major J, Delabie J and McKenzie N. 1997. Ant litter fauna of forest, forest edges and adjacent grassland in the Atlantic rain forest region of Bahia, Brazil. Insectes Sociaux 44:255-266. Oliveira P, Galetti M, Pedroni F and Morellato L. 1995. Seed cleaning by Mycocepurus goeldii ants (Attini) facilitates germination in Hymenaea courbaril (Caesalpiniaceae). Biotropica 27:518-522. Torres J. 1984. Niches and coexistence of ant communities in Puerto Rico: repeated patterns. Biotropica 16:284-295. Torres J. 1989. The status of the fungus-grower ants (Hymenoptera: Formicidae) in Puerto Rico and adjacent islands. Journal of Agriculture of the University of Puerto Rico 73:401-403. Weber N. 1937. The biology of the fungus growing ants. Part. l. New forms. Revista de Entomologia 7:378-409 + 11 Figs. Weber N. 1938. The biology of the fungus-growing ants. Part IV. Additional new forms; Part V. The Attini of Bolivia. Revista de Entomologia 9:154-206 + 21 Figs. Weber N. 1940. The biology of the fungus-growing ants. Part VI. Key to Cyphomyrmex, new Attini and a new guest ant. Revista de Entomologia 11:406-427 + 16 Figs. Wheeler W. 1907. The fungus-growing ants of North America. Bulletin of the American Museum of Natural History 23:669-807 + 5 plates and 31 Figs.   3. M. goeldii 9. M. obsoletus 12. M. goeldii %0L]^_`rs    ; < = M N O P ! " ; = G ηypjpp hd=@aJhd=@6]aJ hd=@H*aJhd=@0J6>*OJQJ]^J!hd=@6B* OJQJ]^Jph*jhd=@6B* OJQJU]^Jphhd=@0J6]jhd=@6U]jhd=@6U]hd=@6H*] hd=@H*hd=@5H*\ hd=@5\ hd=@6]hd=@)KL O < = t H I ``$a$$`a$`$a$Ohd=@U hd=@] hd=@6 hI6hd=@jhd=@U<    IZ[zSftuy 1234er$+134;<PQRSjyhd=@Uh9ejV+h9eUhIOhd=@56\] hd=@o(hIj'vhh<U hd=@5\ hd=@] hd=@6h9emHnHujhd=@Uhd=@Bz563=>=? 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