Gunnar Brehm

Global Warming, Elevational Range Shifts, and Lowland Biotic Attrition in the Wet Tropics

Many studies suggest that global warming is driving species ranges poleward and toward higher elevations at temperate latitudes, but evidence for range shifts is scarce for the tropics, where the shallow latitudinal temperature gradient makes upslope shifts more likely than poleward shifts. Based on new data for plants and insects on an elevational transect in Costa Rica, we assess the potential for lowland biotic attrition, range-shift gaps, and mountaintop extinctions under projected warming. We conclude that tropical lowland biotas may face a level of net lowland biotic attrition without parallel at higher latitudes (where range shifts may be compensated for by species from lower latitudes) and that a high proportion of tropical species soon faces gaps between current and projected elevational ranges.

Tropical forests are not flat: how mountains affect herbivore diversity

Ecologists debate whether tropical insect diversity is better explained by higher plant diversity or by host plant species specialization. However, plant-herbivore studies are primarily based in lowland rainforests (RF) thus excluding topographical effects on biodiversity. We examined turnover in Eois (Geometridae) communities across elevation by studying elevational transects in Costa Rica and Ecuador. We found four distinct Eois communities existing across the elevational gradients. Herbivore diversity was highest in montane forests (MF), whereas host plant diversity was highest in lowland RF. This was correlated with higher specialization and species richness of Eois/host plant species we found in MF. Based on these relationships, Neotropical Eois richness was estimated to range from 313 (only lowland RF considered) to 2034 (considering variation with elevation). We conclude that tropical herbivore diversity and diet breadth covary significantly with elevation and urge the inclusion of montane ecosystems in host specialization and arthropod diversity estimates.

Comprehensive Molecular Sampling Yields a Robust Phylogeny for Geometrid Moths (Lepidoptera: Geometridae)

Background The moth family Geometridae (inchworms or loopers), with approximately 23 000 described species, is the second most diverse family of the Lepidoptera. Apart from a few recent attempts based on morphology and molecular studies, the phylogeny of these moths has remained largely uninvestigated. Methodology/Principal Findings We performed a rigorous and extensive molecular analysis of eight genes to examine the geometrid affinities in a global context, including a search for its potential sister-taxa. Our maximum likelihood analyses included 164 taxa distributed worldwide, of which 150 belong to the Geometridae. The selected taxa represent all previously recognized subfamilies and nearly 90% of recognized tribes, and originate from all over world. We found the Geometridae to be monophyletic with the Sematuridae+Epicopeiidae clade potentially being its sister-taxon. We found all previously recognized subfamilies to be monophyletic, with a few taxa misplaced, except the Oenochrominae+Desmobathrinae complex that is a polyphyletic assemblage of taxa and the Orthostixinae, which was positioned within the Ennominae. The Sterrhinae and Larentiinae were found to be sister to the remaining taxa, followed by Archiearinae, the polyphyletic assemblage of Oenochrominae+Desmobathrinae moths, Geometrinae and Ennominae. Conclusions/Significance Our study provides the first comprehensive phylogeny of the Geometridae in a global context. Our results generally agree with the other, more restricted studies, suggesting that the general phylogenetic patterns of the Geometridae are now well-established. Generally the subfamilies, many tribes, and assemblages of tribes were well supported but their interrelationships were often weakly supported by our data. The Eumeleini were particularly difficult to place in the current system, and several tribes were found to be para- or polyphyletic.

Ordinating tropical moth ensembles from an elevational gradient: a comparison of common methods

The analysis of beta diversity (inter-habitat diversity) of very species-rich and incompletely sampled tropical arthropod communities requires the choice of appropriate statistical tools. The performance of the three commonly employed ordination methods, correspondence analysis (CA), detrended correspondence analysis (DCA), and non- metric multidimensional scaling (NMDS), was compared on a large empirical data set of geometrid moths sampled along an altitudinal gradient in an Andean montane rain forest. Despite the high species richness and incompleteness of the ensembles, all methods depicted the same, readily interpretable patterns. Both CA and NMDS showed an arch-like structure, which hints at an underlying coenocline, whereas this arch was computationally eliminated in DCA. For this particular data set, CA and NMDS both provided convincing results while the detrending algorithm of DCA did not improve the interpretability of the data. Of the large number of similarity indices available to be used in combination with NMDS, the binary Sorensen and the abundance-based Normalized Expected Species Shared (NESS) index were tested. Performance of the indices was measured by comparing stress, a measure of poorness-of-fit in NMDS. NMDS ordinations with lowest values of stress were achieved by the NESS index with the parameter m set to its maximum (mmax). In contrast, ordinations based on NESS values with the parameter m set to 1 (identical with Morisitas index), had consistently higher stress values and performed worse than ordinations using Sorensens index. Hence, if high values of m can be achieved in similar data sets, the NESS index with mmax is recommended for ordination purposes and Morisitas index should be avoided.

DNA barcoding-based species delimitation increases species count of Eois (Geometridae) moths in a well-studied tropical mountain forest by up to 50%

Abstract  The genus Eois comprises an important part of megadiverse assemblages of geometrid moths in mountain rainforests of southern Ecuador. In this study we report: (i) on the construction of a DNA barcode library of Eois for identification purposes; and (ii) the exploration of species diversity through species delimitation by pair‐wise distance thresholds. COI barcode sequences were generated from 408 individuals (at least 105 species) collected on a narrow geographic scale (∼40 km2) in the Reserva Biológica San Francisco. Analyses of barcode sequence divergence showed that species delimitations based solely on external morphology result in broad overlap of intra‐ and interspecific distances. Species delimitation at a 2% pair‐wise distance threshold reveals a clear barcoding gap. Fifty‐two previously unrecognized species were identified, 31 of which could only be distinguished by an integrative taxonomy approach. Twelve additional putative species could only be recognized by threshold‐based delimitation. Most splits resulted in two or three newly perceived cryptic taxa. The present study increased the number of Eois species recorded from that small area of Andean mountain forest from 102 to 154 (morphology‐ plus integrative taxonomy‐based) or even 166 (sequence‐based), leaving the species accumulation curve still far from reaching an asymptote. Notably, in no case did two or more previously distinguished morphospecies have to be lumped. This barcode inventory can be used to match larvae to known adult samples without rearing, and will therefore be of vital help to extend the currently limited knowledge about food plant relationships and host specialization.

Caterpillars and Host Plant Records for 59 Species of Geometridae (Lepidoptera) from a Montane Rainforest in Southern Ecuador

Abstract During four months of field surveys at the Reserva Biológica San Francisco in the south Ecuadorian Andes, caterpillars of 59 Geometridae species were collected in a montane rainforest between 1800 and 2800m altitude and reared to adults. The resulting data on host plant affiliations of these species was collated. The preimaginal stages of 58 and adult stages of all 59 species are depicted in colour plates. Observations on morphology and behaviour are briefly described. Five species, documented for the first time in the study area by means of larval collections, had not been previously collected by intensive light-trap surveys. Together with published literature records, life-history data covers 8.6% of the 1271 geometrid species observed so far in the study area. For 50 species these are the first records of their early stages, and for another 7 the data significantly extend known host plant ranges. Most larvae were collected on shrubs or trees, but more unusual host plant affiliations, such as ferns (6 geometrid species) and lichens (3 geometrid species), were also recorded. Thirty-four percent of the caterpillars were infested by wasp or tachinid parasitoids.

Diversity and ensemble composition of geometrid moths along a successional gradient in the Ecuadorian Andes

Little is known about the change of species-rich tropical insect communities along habitat gradients. Diversity and species richness of geometrid moths were investigated at 15 anthropogenically disturbed sites in a montane area in the Andes of southern Ecuador representing a successional gradient. These plots were compared with six closed-forest understorey sites. We collected a total of 23,720 individuals representing 868 morphospecies. Local diversity increased with forest recovery and decreased with increasing distance to the natural forest. 18.6% of all species were found as unique singletons. The mean proportions of local singletons differed significantly between three succession classes. Forest understorey showed a higher proportion of singletons than early and late successional stages. Ordination of the moth samples showed a clear separation of geometrid ensembles at successional sites vs. the forest understorey sites. Patterns of species turnover were influenced by the degree of habitat openness, and to a lesser extent by elevation. However, faunal differences were not related to geographical distances between the sampling sites. In conclusion, geometrid moth ensembles of regenerating Andean montane forest remain diverse, but change significantly in composition relative to adjacent natural forest, whereas the diversity and composition of the geometrid fauna are far more strongly affected in non-forested habitats and abandoned pastures.

A Comparison of Manual and Automatic Moth Sampling Methods (Lepidoptera: Arctiidae, Geometridae) in a Rain Forest in Costa Rica

Abstract Sampling with UV fluorescent light tubes is a commonly used technique both in applied and basic insect studies. Our study compares the performance of two such methods: manual sampling (light towers) and automatic sampling (funnel light traps). The abundance, diversity, and body size of moths representing two species-rich families (Lepidoptera: Arctiidae, Geometridae) were analyzed in a lowland rain forest in Costa Rica (La Selva Biological Station, 10.4° N, 84.0° W) during 2003 and 2004. Light towers were equipped with two 15-W UV fluorescent tubes and were operated for 3 h in 16 nights. Traps equipped with single 8-W fluorescent tubes were run throughout 20 nights in the understory of the forest. In addition, parallel trap sampling was carried out in the canopy. A total of 1,238 arctiid moths representing 162 species and 1,769 geometrid moths representing 196 species were collected. In Geometridae, tower samples were significantly larger than trap samples. Towers also attracted a higher overall number of species. Very small geometrids (particularly of the subfamily Sterrhinae) were under-represented in trap samples, suggesting that this method is biased toward larger species. In arctiid moths, there were no significant differences in either the sample sizes, the number of species or in the size of the individuals sampled. Diversity calculated as Fisher’s α was similar for towers and understory traps in both families. A major component of diversity was added with canopy trap samples for arctiid moths, but not for geometrid moths. In conclusion, ground-based tower sampling proved to be the most suitable method for geometrid moths, and trap sampling including both understory and canopy for arctiid moths. For full moth species inventories, a combination of both approaches is recommended.

Molecular phylogeny of Eois (Lepidoptera, Geometridae): evolution of wing patterns and host plant use in a species-rich group of Neotropical moths.

Strutzenberger, P., Brehm, G., Bodner, F. & Fiedler K. (2010). Molecular phylogeny of Eois (Lepidoptera, Geometridae): evolution of wing patterns and host plant use in a species‐rich group of Neotropical moths. —Zoologica Scripta, 39, 603–620.

Turning Up the Heat on a Hotspot: DNA Barcodes Reveal 80% More Species of Geometrid Moths along an Andean Elevational Gradient

We sampled 14,603 geometrid moths along a forested elevational gradient from 1020–3021 m in the southern Ecuadorian Andes, and then employed DNA barcoding to refine decisions on species boundaries initially made by morphology. We compared the results with those from an earlier study on the same but slightly shorter gradient that relied solely on morphological criteria to discriminate species. The present analysis revealed 1857 putative species, an 80% increase in species richness from the earlier study that detected only 1010 species. Measures of species richness and diversity that are less dependent on sample size were more than twice as high as in the earlier study, even when analysis was restricted to an identical elevational range. The estimated total number of geometrid species (new dataset) in the sampled area is 2350. Species richness at single sites was 32–43% higher, and the beta diversity component rose by 43–51%. These impacts of DNA barcoding on measures of richness reflect its capacity to reveal cryptic species that were overlooked in the first study. The overall results confirmed unique diversity patterns reported in the first investigation. Species diversity was uniformly high along the gradient, declining only slightly above 2800 m. Species turnover also showed little variation along the gradient, reinforcing the lack of evidence for discrete faunal zones. By confirming these major biodiversity patterns, the present study establishes that incomplete species delineation does not necessarily conceal trends of biodiversity along ecological gradients, but it impedes determination of the true magnitude of diversity and species turnover.