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Ecology and Biogeography of Dung-Beetle Communities (Coleoptera, Scarabaeoidea) in an Iberian Mountain Range

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Ecology and Biogeography of Dung-Beetle Communities (Coleoptera, Scarabaeoidea) in an Iberian Mountain Range Journal of Biogeography (1992) 19, 677-691 Ecology and biogeography of dung-beetle communities (Coleoptera, Scarabaeoidea) in an Iberian mountain range F. MARTIN-PIERA, C. M. VEIGA* and J.M. LOB0 Departamento de Biodiversidad; Museo Nacional de Ciencias Naturales (C.S.I. C.); c/ Jose' Gutie'rrez Abascal2; 28006-Madrid, Spain and "Ca'tedra de Entomologia, Facultad de Biologia, Universidad Complutense de Madrid, Ciudad Universitaria; 28040-Madrid, Spain Abstract. The role of the Eastern Mountains of the Iberian richness, characteristic species, systematic composition and Central System (EMICS) as both a barrier to the dispersion EP was found to relate mainly with altitude. Principally of, and ecologic refuge for, Scarabaeoidea dung beetles is endemic, European or wider-range species of the Aphodiidae studied. An Ecogeographic Patterns comparison method (EP and Geotrupidae families, prevented from dispersing toward = set of ecologic features common to, and present geographic lower altitudes by a climatic discontinuity, colonize the range shared by, a group of species) to weight the historical higher altitudes, having taken refuge in the EMICS after and ecological causes of geographic distribution is proposed. Pleistocene glaciations. (iii) The explanation of the causes of This method is based on the assumption that species now the present vertical distribution of Scarabaeoidea dung sharing geographic ranges, with common structural and non- beetles in the EMICS must take into account the historical structural features, also have a (to some extent) common ev- events there, and major modifications of the geographic dis- olutionary history. tribution of the species inhabiting it. The data studied indicate that (i) fauna of the N and S The agreement of inferences drawn from EP analysis with slopes are identical. Euroturanian and Mediterranean species, those drawn from paleontologic and phylogenetic evidence and those which belong to the Scarabaeidae family, predomi- (for some species) supports the use of EP, and validates the nated in all samples. Except for very few species, the assumption made in their application. EMICS have not impeded dispersal of Scarabaeoidea dung beetles; minor regional differences should be explained in Key words. Eco-geographical patterns, dung-beetles, Scara- terms of ecological causes. (ii) The variation in abundance, baeoidea, distribution, historical biogeography, Iberia. INTRODUCTION Non-biotic environmental factors (mountains, rivers, deserts, seas) may act either as a barrier to dispersal, or as a This paper deals with the biogeographical topics of (i) bar- refuge for different taxonomic groups. Now, as Udvardy riers, (ii) refuges, and (iii) the role played by historical and (1969) rightly argued, the distinction between geographical ecological causes in species distribution. and ecological- barriers is a semantic one. In fact, a mountain There have been many attempts to arrive at acceptable defi- chain accommodates an environmental mosaic which may be nitions for the terms dispersal and migration. Dispersal an ecologic barrier for some species or a refuge for others. means a scattering of a population leading to an increase in The Pleistocene climates of ice-free regions (and hence the mean distance between individuals (Danthanarayana, their biotic histories) were greatly affected by the great cli- 1986) as a result of (i) short-distance non-migratory move- matic oscillations of the glaciaVinterglacia1 cycles. Several ments (behavioural range); (ii) long-distance migratory colder and warmer intervals within glacial stages (stadia1 and movements (ecological range), and (iii) truly migratory interstadial) have been recognized and named (Turner & movements and/or repeated short- andor long-distance Hannon, 1988). In response to these climatic changes of the movements during more than one generation (geographical Quaternary, large-scale alterations have taken place in the range); see Southwood (1962) and Hanski (1980a). Over a geographical ranges of species of Coleoptera (Coope, 1979). large space-time scale the geographic range of a species may During warm periods (interglaciaVinterstadia1) the mountain be extended by combinations of these three kinds of move- ranges and other areas within ice-free regions became eco- ments, affected by variations in the balance between intrinsic, logical refuges for northern biotas which were widespread in migration-favouring attributes (dispersal-favouring: struc- colder periods (Halffter, 1987; Turner & Hannon, 1988). ture, physiology, ecology, behaviour, etc.) and extrinsic, en- This effect has probably been heightened by the transversal vironmental factors (both biotic, species interaction; and non- orientation of the Eastern Mountains of the Iberian Central biotic, climate, topography, history, etc.). System (EMICS). Aubert (1963), Gangwere & Morales 678 F. arti in-piera, C.M. Veiga and J.M. ~obo Agacino (1970) and Pascual (1978), among others, investi- (2) Are EMICS an ecological refuge for some species of gated the possible effect of Iberian mountain range orienta- this group of Coleoptera ? tion on the relative abundance of endemisms. (3) Have historic factors played an important role in the When attempting to explain the distribution of a species, spatial distribution variations of coprophagous Scarabaeoi- one must consider both historical and ecological factors dea throughout EMICS ? (Blondel, 1979). The former are taken here to be those that greatly modify the geographical range of a species over many generations through dispersal. The latter are factors METHODOLOGY that scarcely modify the geographical range. Thus historical Sampling design factors are measured on a geological scale and take effect over an evolution time period, while ecological factors are EMICS were divided into three sampling sectors: East, measured on a habitat scale, over ecological time. Central and West (Fig. 1). In each sector three sampling sta- Historical and ecological information can be 'decoded' tions were established on a north-south transect; two sta- from its species carriers, but rarely can the historical contri- tions at similar altitudes on both slopes (at 1200 m. in the bution to variations of geographical range be distinguished lower limit of the Supramediterranean phytoclimate approxi- from the ecological. mately) and the third higher, on the upper limit of the Supra- As is widely accepted by biologists, the closer the mediterranean phytoclimate (for botanical terminology see phylogenetic relationships among species, the greater the Rivas-Martinez, 1987 and Rivas-Martinez et al., 1987), likelihood of a common evolutionary history. Usually, which might or might not correspond with the mountain phylogenetic inferences are based on structural features since crests. knowledge of ecology, behaviour and other non-structural The easternmost transect was situated in the Macizo de features is less complete than that of morphology. Neverthe- Ayllbn; Riaza (1240 m: R) and Colmenar de la Sierra less, common non-structural features, and shared geographic (1210 m: CS) were, respectively, the north and south sta- range among species, may also be indicative of a (to some tions. The highest point was in Puerto de la Quesera extent) common evolutionary history. If this is so, we would (1600 m), where three samples of different biomes were then be enabled to draw historical inferences based on these collected: mat-grasses, ~ardusstricta (PQMl), Pyrenean oak common ecologic and geographic features. woodland, Quercus pyrenaica (PQQp), and beech wood- We define the Ecogeographic Pattern (EP) of the species land, Fagus sylvatica (PQFs). as the set of ecologic features common to, and present geo- In the central transect, set up in the Macizo Central de graphic ranges shared by, a group of species (see Methodo- Guadarrama, the north and south stations were El Robledo logy for particular definitions). Therefore, common EP (RO) and Navacerrada (N) respectively, both at 1170 m. among species is taken to mean a common (at least partially) The highest altitude station was in the Puerto de Cotos where evolutionary history. Then species EP become a means of two samples were collected: the first in a closed pine wood- facilitating comparisons of the fauna of different areas, and land, Pinus sylvestris (CT1:1630 m), and the other in mixed may also provide a means for breaking down the contribu- zones of mat-grasses, pines, and broom, Cytisus purgans tion to distribution variations into their historic and ecologic (CT2: 1760 m). components. Thus EP would provide an alternative line of Finally, the western transect was established in the Sierra historical biogeography research in cases where phylogenetic de Malagbn by fixing the stations in Aldeavieja (A) and data are lacking. Navalperal de Pinares (NP), both at 1200 m. The highest A biogeographic sampling of areas might produce the station, in Puerto de la Cruz de Hierro (CH), was on the line following data sets. of the mountain crest (1560 m). (1) The EP and species are common. Obviously no bioge- Besides the above-mentioned three transects, two samples ographic inferences are possible. (TI: 1220 m, grasslands, pastures; T2: 1240 m, closed (2) Species are not identical, but are characterized by Pyrenean oak woodland) were collected in the westernmost similar EP. Although the species involved have adapted dif- station, El Tiemblo, on the southern slope. This station is in ferently to ecological conditions, they probably have a the Valley of the
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