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TERRITORIAL BEHAVIOR of the SUBSOCIAL a Transversal Que Es Medida No Está BEETLE HELISCUS TROPICUS UNDER LABORATORY De Prominencia (Figs

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TERRITORIAL BEHAVIOR of the SUBSOCIAL a Transversal Que Es Medida No Está BEETLE HELISCUS TROPICUS UNDER LABORATORY De Prominencia (Figs MEx. 70 ( 1986) nte delimitada por dos abruptos pro­ TERRITORIAL BEHAVIOR OF THE SUBSOCIAL a transversal que es medida no está BEETLE HELISCUS TROPICUS UNDER LABORATORY de prominencia (figs. 9 y 14). En S. CONDITIONS (COLEOPTERA, PASSALIDAE) :mcavidad central del borde postero­ ' menos profunda y lateralmente está nares sin embargo la placa transver­ ¡ueada por dos abruptas prominencias ,, neto, en relación con la ausencia de JORGE V ALENZUELA-GONZÁLEZ lENTOS Centro de Investigaciones Ecológicas del Sureste nstituciones por el' préstamo de mate­ Apartado Postal 36 Dr Randall T. Schuh (AMNH); Dr. Tapachula, Chiapas 30700 ra. Ursula Gollner-Scheiding (ZMB); MEXICO 1mann (MABR); Dr. Abraham Wil­ 'A); Al Dr. W. R. Dolling (BMNH) >rindados en el curso de este estudio. ional de Ciencia y Tecnología (CO permitió visitar diversos museos sud- al M. en C. Pedro Reyes-Castillo y >ermitirme trabajar durante mi año ' de Ecología, A. C. CITADA eteropterorum Bruxellensis 11. Coreidae, 'em. S oc. Entamo/. Belgique. 22: 154-155. ae in the Rijksmuseum van Natuurlijke Meded. 18:187. 'ntius St<il, con descripción de un nuevo era-Heteroptera-Coreidae-Nema topodini) . r. Zool. (En prensa). e general des Hemipteres Tome /l. He- 1 1' Academie Royal e de Belgique: 85 ~venska Vet. Ak. Handl 12:181-192. Ofv. Vet-Ak. Fort. 16(10) :471-472. Folia Entomológica Mexicana No. 70: 53-63 (1986) ,K. Svensksa Vet-Ak. H andl. 9 (1) : 207. of Hemiptera-Heteroptera in the collec­ l: 103 y 142-143. Recibido para publicación: 20 enero 1986. Aceptado para publicación: 26 mayo 1986. 53 54 FoLIA ENTOMOL. MEx. 70 ( 1986) VALENZ zation, which occurs in dec< together and cooperate in th .After their eclosion, the you help their progenitors with later they disperse and form 1 Castillo, 1975; Schuster 197 RESUMEN González & Castillo, 1984; 1985) Generally these insect: El comportamiento territorial de H. tropicus, su variación estacional y la part1c1pa­ Gray, 1946; Schusted 1975:: ción de cada uno de los sexos en dicho comportamiento fueron estudiados por medio Sorne observations carried de la introducción de pasalidos "extraños" en nidos huéspedes instalados en el labora­ torio. Se encontró que únicamente los nidos colectados en la época de reproducción suggest the existence of a se: presentan este comportamiento. Este se manifiesta por medio de la producción de reac­ junctus (Illiger). Other stU< ciones agonísticas ostensibles (embestidas, mordidas y huidas) al contacto entre los Schuster ( 1975a) with the sa huéspedes y los intrusos. En todos los casos observados los huéspedes iniciaron la agre­ sión y los intrusos presentaron un comportamiento de huida. La mayor parte de estas affinis ( Percheron), also su¡ interacciones fueron unidireccionales (de los huéspedes a los intrusos) ; sin embargo, species. N evertheless, in all 1 ocasionalmente, se observaron también algunas interacciones agonísticas recíprocas que sional. In previous publicati tuvieron el mismo resultado que las anteriores. Ambos sexos participan en la defensa del nido, pero la agresión es más frecuentemente dirigida hacia los individuos del mismo productive behaviour (Valen: sexo. También se observó que algunos de los intrusos pueden ser tolerados por las pa­ lenzuela-González, 1985) of 1 rejas territoriales; con los datos disponibles no es posible explicar el origen de estas to the tribe Proculini and variaciones en el comportamiento. present study we report th PALABRAS CLAVE: Comportamiento Territorial, Passalidae, Insectos subsociales, carried out under laboratory México. torial behaviour of these im sex in this same behaviour. ARSTRACT Territorial behavior, its seasonal variation and the partJc1pation of each sex were studied in H. tropicus by introducing "foreign" passalids into host nests imtalled in the laboratory. Only nests collected in the reproductive season exhibited this behaviour, The insects were collecte' which comprises obvious agonistic reaction ( attacks, bites and retreats on contact be­ of Berriozabal, Chiapas, Me tween the host and intruders. In all observed cases the hosts initiated the aggression and type of nests utilized for thc intruders presented the escape response. The majority of these interactions were uni­ directional (from host to intruder), but occasionally sorne reciproca! agonistic interac­ described ( Valenzuela-Gonz: ctions were observed that nevertheless had the same end result. Both sexes participated With the object of studying in nest defense but aggression was more frequently directed at individuals of the same­ sects, a series of experimenl sex. It was also observed that sorne intruders were tolerated by the host pair; with the data available it is not possible to explain the origin of this variation in behaviour. were introduced into host r nests were occupied by pass KEY WORDS: Territorial behaviour, Passalidae, Subsocial Insects, Mexico. so that there was no modifi was also utilised for the ex¡: lNTRODUCTION After their installation in in complete darkness to allc Passalids are saproxylophagous subsocial beetles (Wilson, 1971). The repro­ roundings. Following this p ductive unit begins with the formation of a monogamous pair; aftcr fertili- were started using individm ( 1986) VALENZUELA. BEHAVIOR OF Heliscus 55 zation, which occurs in decaying tree trunks, both members of the pair stay together and cooperate in the construction of galleries and care of the young. After their eclosion, the young adults remain for sorne time in the nest and help their progenitors with the care of the younger members of the brood; la ter they disperse and form new nests ( Ohaus, 1900, 1909; Gray, 1946; Reyes­ Castillo, 1975; Schuster 1975a; Reyes-Castillo & Halffter, 1983; Valenzuela­ González & Castillo, 1984; Valenzuela-González 1985; Schuster & Schuster 1985) Generally these insects have a seasonal life-cycle ( Pearse et al.) 1936; .."''""'u" estacional y la participa­ Gray, 1946; Schusted 1975a; Valenzuela-González, 1985). estudiados por medio instalados en el labora­ Sorne observations carried out by Gray ( 1946) and Mullen & Hunter ( 1973), en la época de reproducción suggest the existence of a seasonal territorial behaviour in Odontotaenius dis­ medio de la producción de reac­ junctus (Illiger). Other studies varried out by Alexander et al. ( 1963) and y huidas) al contacto entre los los huéspedes iniciaron la agre­ Schuster (1975a) with the same species and by the latter author with Passalus huida. La mayor parte de estas affinis ( Percheron), also support the existen ce of this behaviour in the two a los intrusos) ; sin embargo, species. Nevertheless, in al! these cases the observations were scant and occa­ agonísticas recíprocas que sexos participan en la defensa sional. In previous publications we have reported various aspects of the re­ hacia los individuos del mismo productive behaviour (Valenzuela-Rodríguez & Castillo, 1984) and biology (Va­ lenzuela-González, 1985) of Heliscus tropicus (Percheron), a passalid belonging to the tribe Proculini and having a wide distribution in Mexico. In the present study we report the results obtained from a series of experiments Passalidae, Insectos subsociales, carried out under laboratory conditions with the object of studying the terri­ torial behaviour of these insects, its seasonal variability and the role of each sex in this same behaviour. METHODS part1c1pation of each sex were into host nests installed in the The insects were collected in "El Suspiro", a locality in the municaplity of Berriozabal, Chiapas, Mexico. The location, techniques of collection and type of nests utilized for the observations of the insects have been previously reciproca! agonistic interac­ described (Valenzuela-González & Castillo, 1984; Valenzuela-González 1985). Both sexes participa ted With the object of studying the existence of territorial behaviour in these in­ at individuals of the same­ sects, a series of experiments was carried out where "instrusive" individuals by the host pair; with the of this variation in behaviour. were introduced into host nests previously installed in the laboratory. These nests were occupied by passalids al! collected from the same nest in the field Insects, Mexico. so that there was no modification of the host group. The original nest wood was also utilised for the experiments. After their installation in the laboratory, the insects remained for five days in complete darkness to allow them to become accustomed to their new sur­ roundings. Following this period, the homospecific introduction experiments pair; after fertili- were started using individuals collected at the same time and place as those 56 FoLIA ENTOMOL. MEx. 70 (1986) VALENZl from the host nest. Thus insects of unknown age were used, although to reduce the influence of this factor only individuals that had aquired adult colour were utilized. Typically, with this species the adults attain their characteristic 1) Description of behavio black colour at approximately four months. ( 21.6%) aggressive interactio During the agonistic interactions between passalids, obvious acts physical initiated the aggression and 1 aggression such as attacks and hites were seen, although usually accompanied escape response to the aggre: by other acts including exploratory movements, antennal tapping and stri­ these encounters were marke dulations (Gray, 1946; Alexander et at., 1963; Mullen & Hunter, 1973; Schus­ sistence of the aggression, it ü ter, 1975a; Valenzuela-González & Castillo, 1984). However, these latter acts finity in the host nest the fin; al so manifest
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