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Principal objectives

•  Create an integrative approach to study understand speciation mechanisms in scarab beetles by combining phylogenetics, biogeography, phylogeography, and ecology.

•  Establish species boundaries within the endemic dung beetle tribe Eucraniini to be used for monitoring and habitat recovery measures in the Monte biogeographic province.

•  Provide empirical evidence for biogeographic processes that may have affected the speciation events of insects in the Monte biogeographic province.

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Project Summary

This project will investigate the evolutionary radiation of a group of endemic dung beetles from South America through monographic research and applying an integrative method in systematics and biogeography. This research will provide evidence for evolutionary trends in dung beetles, historical and ecological biogeography, and conservation priorities for southern South America.

Members of the dung beetle tribe Eucraniini (4 genera, 16 species) have traits that make them of interest to evolutionary biologists: they are flightless (except 1 species), have a unique ecological specialization on dry dung pellets, and have a unique food relocating behavior. Species of Eucraniini are endemic to desert-scrub habitats of the Monte biogeographic province in Argentina. These regions have high levels of endemism, are exploited by humans, and are extremely fragile. Populations of eucraniine species are isolated geographically and often exhibit different morphological character states from neighboring populations.

Speciation is one of the most intriguing questions in evolutionary biology. To date, most attempts to infer speciation modes have compared geographic ranges in relation to phylogenies without considering the environmental or temporal space. The proposed research will implement a new approach that integrates phylogenetics, biogeography, phylogeography, and ecology in order to understand speciation in the Eucraniini, a monophyletic clade of South American dung beetles.

Species are considered the fundamental units in studies of ecology, systematics, conservation biology, and macroevolution. However, the questions "What is a species?" and "How does speciation work?" are still two of the most intriguing questions in evolutionary biology. By applying modern methods in systematics and biogeography to define species and study speciation in dung beetles, I will be able to provide empirical evidence to address these crucial questions. The proposed research will establish species boundaries within this unique group of dung beetles that could be used forconservation monitoring and habitat recovery measures in the Monte biogeographic province.

Results of this project will provide a much-needed foundation in our understanding of distribution patterns, endemism, and conservation in the Monte biogeographic province of southern South America.

Eucraniini food relocation behavior

Adults run in zig-zags or randomly from their burrow (apparently searching for food). To carry the food, the beetles grasp it with the foretibiae and run forward using only their middle and hind legs. This behavior is known only for members of the tribe Eucraniini.

When foraging, species of Glyphoderus and Anomiopsoides, run on four legs, keeping their forelegs motionless and in a horizontal position with respect to the surface, a bahvarior that is known for members if these two genera among all beetles.

In a second method of food relocation bahavior, called “butting,” pieces of dung (not formed into a ball) are rolled by the beetles walking forward and pushing it with their head and forelegs. This method was observed in Eucranium arachnoides Brullé.

When the entrance of the burrow is reached, the beetles enter by walking forward rather than backward as most of the dung beetles do. The burrow is always previously dug and it is variable in depth.

Detailed descriptions of the biology, food relocation behavior, and evolution of this unique group of dung beetles are discused in Zunino et al. (1989, 1991), Monteresino and Zunino (2003), Ocampo (2003, 2005), and Ocampo and Philips (2005).


Monteresino, E. M. and M. Zunino. 2003. Sobre el comportamiento de la alimentación de Eucraniini (Coleoptera: Scarabaeidae: Scarabaeinae). (Eds. G. Onore, P. Reyes Castillo, and M. Zunino) pp. 75-80. Sociedad Entomologica Aragonesa. Zaragoza, Spain.

Ocampo, F. C. 2004. Food relocation behavior and synopsis of the southern South American genus Glyphoderus Westwood (Scarabaeidae: Scarabaeinae: Eucraniini). The Coleopterists Bulletin 58: 295-305.

Ocampo, F. C. 2005. Revision of the southern South American endemic genus Anomiopsoides Blackwelder 1944 (Coleoptera: Scarabaeidae: Scarabaeinae: Eucraniini) with description of its food relocation behavior. Journal of Natural History 39: 2537-2557.

Ocampo, F. C and T. K. Philips. 2005. Food relocation behavior of the Argentinian dung beetle genus Eucranium Brullé and comparison with the southwest African Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae: Scarabaeiniae). Revista de la Sociedad Entomológica Argentina 64: 53-59.

Zunino, M., E. Barbero, and M. Luzzatto. 1989.   Food relocation behavior in Eucraniina beetles (Scarabaeidae) and the constraints of xeric environment. Tropical Zoology 2: 235-240.

Zunino, M., E. Barbero, C. Palestrini, and M. Luzzatto. 1991. La taxocenosi a scarabeidi coprofagi xerico: materiali per un'analisi biogeografica storico-causale. Biogeographia 15: 41-47.

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Images

The following images illustrate some important characteristics of the Eucraniini biology and behavior that make them unique among all dung beetles.

1. Specialization for arid habitats.
2. Flightlessness (except one species).
3. Specialization on dry dung, ussually dung pellets.
4. Bizarre behavior of relocating food by grasping it with the forelegs and running forward with it, and sometimes butting it with thier forelegs and head.

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Ecraniini habitat in La Rioja (top) and Mendoza provinces in Argentina. The landscape consists of sandy plains and plateaus and is characterized by the presence of mountain chains that define several, longitudinal valleys.
     
Ennearabdus lobocephalus next
to its burrow.
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Ennearabdus lobocephalus digging its burrow close to a rodent nest (not seen) where its finds its favorite food, dung pellets.
     
Eucranium arachnoides digging its new burrow after several attempts to carry a large piece of dry horse dung to its previously dug burrow. This specimen was observed carrying this large piece
of dung by "butting" it with its head
and forelegs.
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The same specimen as above using its head and pronotum
to bulldoze the sand out
of its burrow.

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Glyphoderus sterquilinus foraging
for dung pellets on the sandy soils
of Mendoza, Argentina. While foraging, species of Anomiopsoides and Glyphoderus tipically run
on four legs.
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Female of Glyphoderus monticola
carrying a rodent pellet between
its forelegs while its walks
on the remaining four legs.

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Videos  
These videos were recorded during a field trip conducted in Mendoza, Argentina in January 2003, by Federico Ocampo (except indicated). Copyright © 2003 by Federico Ocampo.
   
Video 1 Eucranium arachnoides carrying a dung pellet 1
Video 2 Eucranium arachnoides carrying a dung pellet 2
Video 3 Glyphoderus sterquilinus foraging
Video 4 Anomiopsoides heteroclyta foraging
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