The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient


  • Gunnar Brehm,

    Corresponding author
    1. Department of Animal Ecology I, University of Bayreuth, Universitätsstraße 30, D-95440 Bayreuth, Germany,
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    • Present address: Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, Erbertstraße 1, 07743 Jena, Germany.

  • Robert K. Colwell,

    1. Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269-3043, USA, and
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  • Jürgen Kluge

    1. Department of Systematic Botany, University of Göttingen, Untere Karspüle 2, 37073 Göttingen, Germany
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*Correspondence: Gunnar Brehm, Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, Erbertstraße 1, 07743 Jena, Germany. E-mail:


Aim  The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored.

Location  The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas.

Methods  Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance-based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity.

Results   A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump-shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction-based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r2 in Geometridae. The potential effects of geometric constraints (mid-domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small-ranged species showed stronger deviations from model predictions than large-ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small-ranged species. We hypothesize that small-ranged species (at least of the Ennominae) may tend to be host specialists, whereas large-ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern.

Conclusions  The results clearly show that moth diversity shows a hump-shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.