BIOTROPICA 34(1): 144±154 2002 Response of Dung Beetle Diversity to Human-induced Changes in a Tropical Landscape1 Gonzalo Halffter 2 and Lucrecia Arellano 3 Instituto de EcologõÂa, A.C., Departamento de EcologõÂa y Comportamiento Animal, Apartado Postal 63, 91000, Xalapa, Veracruz, MeÂxico ABSTRACT This paper examines dung beetle communities in remnant patches of tropical deciduous forest at Veracruz, Mexico, as a case study of the effects of tropical deforestation on biodiversity. The two study areas have a common biogeographic history and similar macroclimatic conditions and have been modi®ed by human activities to different extents. The main difference between them is that cattle, which provide the dung beetle's food supply, are present in only one of the areas. Comparison of the dung beetle faunas sheds light on the relative importance of forest cover versus food supply as the principal factor determining the structure and diversity of the fauna. This comparison, which we broaden through an examination of data from other localities in central Veracruz, permits us to speculate about what happens to biodiversity when a tropical deciduous forest undergoes modi®cation of varying type and intensity. Where tree cover has been most modi®ed, native forest species undergo local extinction and are replaced by open area species. On the whole, there has not been a net reduction in species richness (gamma diversity) in the fragmented landscape of central Veracruz, although local species richness (alpha diversity) has diminished. RESUMEN En este trabajo se presenta una aproximacioÂn de lo que ocurre con la biodiversidad estudiada a traveÂs de un grupo indicador (los escarabajos del estieÂrcol) en dos remanentes de bosque tropical caducifolio del estado de Veracruz, MeÂxico. Los dos sitios comparten la misma historia biogeogra®ca, condiciones macroclimaÂticas semejantes, y ser bosques parcialmente modi®cados por la actividad humana. La diferencia principal esta en la oferta de alimento para los Scarabaeinae, porque solamente en uno de los lugares hay ganado vacuno. La comparacioÂn de la fauna de Scara- baeinae de los dos lugares nos permite senÄalar que la cubierta forestal, y no la oferta de alimento, es el principal elemento conformador de la estructura y diversidad del gremio. Esta comparacioÂn, ampliada con datos de otros puntos de Veracruz centro nos permite especular lo que ocurre con la biodiversidad (representada por los Scarabaeinae) al modi®carse el bosque tropical caducifolio en distintas formas e intensidades. Es relevante la sobrevivencia de las especies propias del bosque a nivel paisaje (diversidad gama), aunque puedan desaparecer en parte a escala puntual. Asõ como su reemplazo por especies helio®las en los puntos en que la vegetacioÂn arboÂrea ha sido maÂs modi®cada. En conjunto el paisaje fragmentado y diverso de Veracruz centro no senÄala una peÂrdida de especies, aunque puntualmente (diversidad alfa) sõ ocurra. Key words: biodiversity indicator groups; human disturbance; Mexico; Scarabaeinae; tropical deciduous forest. FEW PUBLICATIONS MEASURE BIODIVERSITY LOSS IN DIF- portant loss of biodiversity. There is no doubt that FERENT TAXA, especially in the tropics, despite the this view (which has led to much speculation about fact that much has been published about loss of the extinction of species) is realistic at the small species due to human activities (Murphy 1989, spatial and temporal scales at which the changes Brown & Brown 1992, Santos Filho 1995). Almost take place. These conclusions, however, may not be all studies have aimed at determining the local ef- directly extrapolated to the landscape level, which fects of habitat destruction (e.g., for arthropods, see is the scale at which extinction and survival gen- Klein 1989, Shure & Phillips 1991, Holloway et erally occur (except in the case of restricted-range al. 1992, Majer & Beeston 1996). The conclusions endemics). of these studies are alarming: few species from the There are two main reasons why the conclu- original communities survive and there is an im- sions of small-scale studies cannot be extrapolated directly to the much larger scale of landscapes. 1 Received 8 Mar 2000; revision accepted 10 March First, the physical matrix may be heterogeneous, 2001. especially in landscapes with complex topography. 2 E-mail: [email protected] This heterogeneity (orographic, edaphologic) does 3 E-mail: [email protected] not permit human activities to occur uniformly 144 Dung Beetles and Tropical Deforestation 145 throughout the landscape. Second, not all human of large species and burrowers, greater total bio- activities cause radical changes to natural commu- mass, and lower equitability. nities. In the tropics, a forest may be destroyed over a large area to provide land for plantations or for METHODS livestock, but remnant patches are often left rela- tively unmodi®ed. In such cases, human activities STUDY AREAS. The two study areas Jalcomulco produce very heterogeneous landscapes in which and Rancho Real Minero (RRM) are located natural communities are fragmented but do not (198199390±198269390N, 968449080±968359180W) disappear completely. These landscapes provide a 20 km apart in the lower foothills of central Ve- challenge to biodiversity studies, because at inter- racruz (Fig. 1). The original vegetation of both ar- mediate levels of disturbance, diversity may actually eas was tropical deciduous forest (Castillo-Campos increase (Connell 1975, 1978). 1995). Jalcomulco has a remnant of original veg- In this paper, we examine how biodiversity re- etation with patches of different successional stages. sponds to habitat fragmentation at the local and The patch we studied (which covers ca 500 ha) was landscape levels using dung beetles (Coleoptera: on a southward-facing slope and had an average 2 Scarabaeidae: Scarabaeinae) as an indicator group density of 12 trees and 8 shrubs per 100 m . There (Forsyth et al. 1998 and references therein; Halffter were no cattle on this slope, although cattle were 1998). We studied dung beetle communities in two present in small numbers in the ¯at valley bottom. areas of central Veracruz that were once covered RRM measures 300 ha and contains patches of for- continuously by tropical deciduous forest and sub- est with an average density of 5 trees and 15 shrubs 2 jected to different types of human disturbance. We per 100 m interspersed with pastureland contain- compared them with each other and to other dung ing species introduced 28 years ago (e.g., Sorghum beetle communities in central Veracruz. halapanse and Cynodon plectostachyus). On average, Veracruz is one of three states in Mexico that 40 head of cattle fed on a rotational basis in the pasture as well as in the forest patches. have extremely high biodiversity (a megadiverse area); however, it is also a region with one of the SAMPLING METHODS. Four different trap types highest amounts of habitat destruction. Contrary were used to sample beetles. (1) The surface excre- to what might be expected, and partly due to the ment trap consisted of a 1-liter plastic container complex orography of central Veracruz (with a gra- buried in the soil so that its rim was at ground dient of 0 to 4250 m elev. over 80 km), the land- level. Each trap contained 150 g of fresh cow ma- scape has not been affected in a uniform way by nure placed in a nylon stocking in the upper part human activities as would have occurred if the land of the container, and the bottom of the container were ¯at. The main consequence has been that was ®lled with 250 ml of water containing deter- habitats persist as patches in the least accessible ar- gent. We placed 40 traps in each study area for a eas and the landscape is extremely heterogeneous. 24-hour interval and 168 for periods of 12 hours These patches, some of them quite extensive, have in order to distinguish between diurnal (0600± been in¯uenced by human activities to varying de- 1800 h) and nocturnal (1800±0600 h) activities. grees, making Veracruz an appropriate location to (2) The surface carrion trap was constructed from study the effects of fragmentation on biological di- a container similar to the preceding trap but was versity. baited with 150 g of squid meat. The number of Our working hypothesis was that the amount traps and the sampling periods were the same as and quality of the food supply is the main factor with the previous traps. (3) The bottom excrement, determining species richness and trophic structure or bottom carrion trap, was baited with 150 g of in dung beetle communities. To test this hypoth- human excrement or 150 g of squid but was placed esis, we compared one area with and one without on a layer of soil in the bottom of the plastic con- cattle (large human-induced change in food sup- tainer. In this case, 20 traps of each type were left ply), other environmental conditions being similar. for a 24-hour period. (4) The permanent necrotrap We expected to ®nd a large difference in the species (NTP 80; MoroÂn & TerroÂn 1984), which was de- composition and trophic structure of the two areas. signed to remain functional for long periods, had We did not expect to ®nd signi®cant differences in bait consisting of 150 g of squid meat and was left necrophagous beetles; among coprophagous bee- it in the ®eld for 30 days. There were four traps tles, we expected the area with cattle to have a per study area. We protected the ®rst three types much larger number of species, higher abundances from rain by covering them with a plastic plate at 146 Halffter and Arellano FIGURE 1. Location of the study areas. Beetles represent sampling sites; arabic numbers: points of comparison used in Table 2. 10 cm height from the rim of the trap. Finally, and uals was signi®cantly different in relation to light as a complementary method, we placed on two intensity (t 5 132, P 5 0.044).