Diversity of Carabidae (Insecta, Coleoptera) in Epiphytic Bromeliaceae in Central Veracruz, Mexico

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Diversity of Carabidae (Insecta, Coleoptera) in Epiphytic Bromeliaceae in Central Veracruz, Mexico COMMUNITY AND ECOSYSTEM ECOLOGY Diversity of Carabidae (Insecta, Coleoptera) in Epiphytic Bromeliaceae in Central Veracruz, Mexico 1,2 2 3 E. MONTES DE OCA, G. E. BALL, AND J. R. SPENCE Environ. Entomol. 36(3): 560Ð568 (2007) ABSTRACT This paper documents the existence of carabid assemblages associated with bromeliads on the Cofre de Perote, Veracruz, Mexico. Based on bromeliads sampled over three altitudinal ranges, the assemblages included at least 26 species with an arboreal lifestyle and another 11 species that are not strictly arboreal. Seven species are new to science, urging us to pay attention to the arboreal fauna in forest conservation studies. Composition of carabid assemblages associated with bromeliads changes with alti- tude. In lowlands, it is comprised almost entirely of species of Lebiini, with the Platynini dominating assemblages found in bromeliads Ͼ1,000 m above sea level. Our data suggest that carabids use bromeliads to reduce stresses associated with drought periods, the exact timing of which depends on altitude. The unexpected low diversity of the carabid fauna associated with bromeliads at middle altitude is explained in terms of anthropogenic conversion of the original forest to pastureland. Given the importance of arboreal elements, further fragmentation of subtropical and tropical mountain forest signiÞcantly threatens overall carabid diversity. Este trabajo presenta la existencia de ensambles de cara´bidos asociados con bromelias en el Cofre de Perote, Veracruz, Me´xico. Con base en un muestreo de bromelias efectuado en tres rangos altitudinales los ensambles incluyeron al menos 26 especies con aÞnidad arbo´rea y otras once sin esa aÞnidad. Siete especies de cara´bidos son nuevas especies con lo cual se llama la atencio´n sobre la importancia de considerar la fauna arbo´rea en los estudios de conservacio´n de bosques. La composicio´ndelos ensambles de cara´bidos asociados a bromelias cambia con la altitud. En tierras bajas se compone casi completamente por especies de la tribu Lebiini en tanto que especies de la tribu Platynini dominan en los ensambles que se encuentran en bromelias por arriba de los 1,000 metros de altitud. Nuestros datos sugieren que los cara´bidos usan las bromelias para reducir el estre´s asociado con perõ´odos de sequõ´a, cuya ocurrencia depende de la altitud. La inesperada baja diversidad de cara´bidos asociados a bromelias que se encontro´ en altitudes medias se explica en te´rminos de la conversio´n de los bosques originales a tierras de pastoreo. Dada la importancia de los elementos arbo´reos, la continua frag- mentacio´n del bosque montano tropical y subtropical amenaza signiÞcativamente la diversidad de cara´bidos. KEY WORDS altitude, bromeliads, carabids, diversity, Mexico The role of epiphytic plants for establishment and than 40 beetle families including ground beetles maintenance of the extraordinary arthropod diver- (Carabidae) have been recorded, even in tree sity in forest canopies has been well documented crowns of old-growth temperate deciduous forest (Richardson 1999, Stuntz et al. 1999, Greeney 2001, (Gering and Crist 2000). For tropical rain forests, Stuntz et al. 2002). For example, Beutelspacher the high number of carabid beetle species sampled (1999) recognized 47 insect families belonging to 14 from the canopy suggests that the diversity of this orders in just one bromeliad species. In particular, family is far higher than previously thought (Lucky beetles are well represented in such forest canopy et al. 2002). Beutelspacher (1972) reported four communities (Floren and Linsenmair 1998). More carabid species with “certain frequency” in epi- phytes, and Murillo et al. (1983) concluded this family was one of the seven most abundant among 1 Corresponding author: Department Biodiversidad y Ecologõ´a An- imal, Instituto de Ecologõ´a, A.C., Km 2.5 Carretera Antigua, a Coate- the bromeliad-associated insect fauna. In an explan- pec no. 351 Congregacio´n El Haya, C.P. 91070 Xalapa, Veracruz, atory summary about the carabids of Mexico, Ball Mexico (e-mail: [email protected]). and Shpeley (2000) highlighted the importance of 2 Department of Biological Sciences, University of Alberta, Ed- the arboreal lifestyle with respect to the diversity of monton, Alberta T6G 2E9, Canada. 3 Department of Renewable Resources, University of Alberta, Ed- the carabid fauna in tropical areas. They showed monton, Alberta T6G 2H1, Canada. that 24% of the 172 genera of carabids extant in 0046-225X/07/0560Ð0568$04.00/0 ᭧ 2007 Entomological Society of America June 2007 MONTES DE OCA ET AL.: CARABIDS AND BROMELIADS IN VERACRUZ,MEXICO 561 Mexico include arboreal species, and many such in which trees harbored many epiphytic bromeliads species have been collected from epiphytes, espe- on trunks and branches. The spatial distribution of cially bromeliads. bromeliads in Veracruz is highly aggregated, but there Bromeliads and their associated fauna are well-de- is no evidence of further spatial segregation of bro- Þned biological systems (Picado 1913, Delamare-De- meliad species within these aggregations (Hietz and bouteville 1948, Beutelspacher 1972, Benzing 1986, Hietz-Seifert 1994, 1995). Therefore, in this study of Willimas and Feltmate 1992, Beutelspacher 1999, Ri- carabid beetles, we considered all bromeliads, regard- chardson 1999). These epiphytic plants have rosetted, less of species, as equivalent potential habitats. overlapping wide leaves that allow water and debris According to Hietz and Hietz-Seifert (1994), one accumulation, which favors establishment of a variety can Þnd 12 bromeliad species (Tillandsia) at the low of organisms. Leaves form a cup or small tank that altitude sites of range A, whereas the species number becomes occupied by water and organic matter increases to 28 (3 Catopsis and 25 Tillandsia)atthe (sprigs, leaves, fruits, seeds, spores, pollen grains, ßow- middle altitude sites on range B. The number of ex- ers, etc.). Water from rain or mist condensation is pected bromeliad species decrease to ϳ10 (1 Catopsis deposited in the tank during the rainy season and and 9 Tillandsia) over range C. Those authors re- slowly evaporates during the dry season. This water is ported no bromeliads at Ͼ3,000 m of elevation. partially absorbed by the plant itself, and it is also used Sampling Protocol. Bromeliads were sampled for by multiple organisms, which in return supply nutri- carabids by at least two people once about every 45 d ents derived from their activities to the plant. during a full year interval (July 2000 to June 2001) in It is reasonable to suppose that tight relationships, each one of the altitudinal ranges. Because some ep- perhaps even mutualisms, could develop between bro- iphytic bromeliads are threatened through illegal meliads and some animals that depend on them for trade of ornamental plants, we sought to minimize particular aspects of their life cycles. For example, the destruction of the plants as follows: (1) we sampled metabolic wastes and remains of dead ants result in a only from one of several possible sites in each altitude large contribution of nutrients for the plants and for during each sampling period, (2) if Þve consecutive the bromeliad inhabitants as well (Rickson 1979, Da- bromeliads were sampled from at least two trees at vidson et al. 1989, Rico-Gray et al. 1989, Benzing 1990, least 10 m apart with failure to Þnd any carabids, Olmsted et al. 1995, Treseder et al. 1995, Benzing sampling was terminated at that site for that sampling 2000). The beneÞts offered by epiphytes to their as- period, (3) a maximum of 25 bromeliads was sampled sociated fauna are hypothesized to include cover and at a site during a particular sampling period, and (4) refuge against predators and/or extreme climatic con- only bromeliads no more than 5 m above the ground ditions, a supply of water, resting sites, or aggregation were sampled (this was also partly a constraint of the sites useful for mating (Benzing 1970, 1984, 1990). cutting pole used). Samples came from isolated trees In this paper, we describe and analyze the carabid or groups of trees at least 10 m apart as they were assemblage associated with epiphytic bromeliads in encountered haphazardly in walking through the sites. central Veracruz, Mexico. We show that there is a Neither tree nor bromeliad species were identiÞed nor regular carabid assemblage associated with bromeli- speciÞcally targeted in this study. ads that includes some species commonly living at Bromeliads were cut and pulled down using exten- ground level. Our data suggest that the association sible branch-cutters and a curved blade and immedi- between carabids and bromeliads has a distinct sea- ately disassembled leaf by leaf and shaken over a sonal component, underscoring the potential role of plastic sheet placed on the ground. Individual carabids these epiphytes in relieving water stress during dry routed by these procedures were captured and pre- seasons. Moreover, this latter effect interacts with served in tubes with vapor of ethyl acetate. To in- altitude so that patterns must be understood in the crease efÞciency of collection, specimens belonging to context of altitudinal zones. We conclude with a gen- other taxa were ignored. Although we conducted no eral discussion about the contribution of the brome- tests of sampling efÞcacy, we felt that we collected liads to the diversity of the carabid fauna of the study virtually all the beetles in the bromeliads. IdentiÞca- area. tion of subsequently pinned specimens was made to the speciÞc level using dichotomous keys (G.E.B. and E.M.O., unpublished data) and by comparison with Materials and Methods previously identiÞed material held at the University of Study Area. This study was conducted on the south- Alberta Strickland Museum (UASM). The material eastern slope of the Cofre de Perote (CP), a mountain from this study is housed in the carabid collection of located in central Veracruz, Mexico, between 19Њ40Ј, the Instituto de Ecologõ´a A.C.
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