Brian A. Gill

Morphological taxonomy, DNA barcoding, and species diversity in southern Rocky Mountain headwater streams

Abstract: Elevation gradients allow scientists to observe changes in fauna over a range of abiotic conditions. A variety of trends in aquatic insect diversity patterns across elevation have been reported. However, many of these studies are confounded because they include streams at lower elevations, which are often larger in size and more polluted than their higher-elevation counterparts. Moreover, such studies always relied solely on morphological delineation of taxa, thereby potentially overlooking cryptic diversity. We reduced these limitations by sampling only minimally impacted wadeable streams across an elevation gradient and by combining morphological taxonomy with deoxyribonucleic acid (DNA) barcoding to identify taxa. We collected numerically abundant Ephemeroptera, Plecoptera, and Trichoptera (EPT) from single streams at ∼200-m elevation intervals across >1000-m transects in 3 watersheds draining the eastern slope of the Colorado Rocky Mountains. Based on morphology alone, we identified 49 numerically abundant EPT morphospecies across 26 sites. Using DNA barcoding, we found 69 distinct lineages that probably represent distinct species. EPT species richness was highest at midelevations, and rates of turnover along elevation transects showed no consistent elevation trend or trend among ecological zones defined by vegetation. &bgr;-diversity across sites at comparable elevations in different watersheds showed a negative trend with increasing elevation that was marginally significant for DNA barcode taxa (p = 0.051) but not for morphospecies. Furthermore, significant (p < 0.05) differences in taxon richness, turnover, and lateral &bgr;-diversity values generated by DNA barcoding underscore the ability of molecular tools to quantify patterns in aquatic insect diversity across elevations.

Climate variability predicts thermal limits of aquatic insects across elevation and latitude

Summary Janzens extension of the climate variability hypothesis posits that increased seasonal variation at high latitudes should result in greater temperature overlap across elevations, and favor wider thermal breadths in temperate organisms compared to their tropical counterparts. We tested these predictions by measuring stream temperatures and thermal breadths (i.e. the difference between the critical thermal maximum and minimum) of 62 aquatic insect species from temperate (Colorado, USA) and tropical (Papallacta, Ecuador) streams spanning an elevation gradient of ca. 2000m. Temperate streams exhibited greater seasonal temperature variation and overlap across elevations than tropical streams, and as predicted, temperate aquatic insects exhibited broader thermal breadths than tropical insects. However, elevation had contrasting effects on patterns of thermal breadth. In temperate species, thermal breadth decreased with increasing elevation because CTMAX declined with elevation while CTMIN was similar across elevations. In tropical insects, by contrast, CTMAX declined less sharply than CTMIN with elevation, causing thermal breadth to increase with elevation. These macrophysiological patterns are consistent with the narrower elevation ranges found in other tropical organisms, and they extend Janzens climate variability hypothesis to freshwater streams. Furthermore, because lowland tropical aquatic insects have the narrowest thermal breadths of any region, they may be particularly vulnerable to short-term extreme changes in stream temperature. This article is protected by copyright. All rights reserved.

Cryptic species diversity reveals biogeographic support for the ‘mountain passes are higher in the tropics’ hypothesis

The ‘mountain passes are higher in the tropics’ (MPHT) hypothesis posits that reduced climate variability at low latitudes should select for narrower thermal tolerances, lower dispersal and smaller elevational ranges compared with higher latitudes. These latitudinal differences could increase species richness at low latitudes, but that increase may be largely cryptic, because physiological and dispersal traits isolating populations might not correspond to morphological differences. Yet previous tests of the MPHT hypothesis have not addressed cryptic diversity. We use integrative taxonomy, combining morphology (6136 specimens) and DNA barcoding (1832 specimens) to compare the species richness, cryptic diversity and elevational ranges of mayflies (Ephemeroptera) in the Rocky Mountains (Colorado; approx. 40°N) and the Andes (Ecuador; approx. 0°). We find higher species richness and smaller elevational ranges in Ecuador than Colorado, but only after quantifying and accounting for cryptic diversity. The opposite pattern is found when comparing diversity based on morphology alone, underscoring the importance of uncovering cryptic species to understand global biodiversity patterns.

Genetic diversity and gene flow decline with elevation in montane mayflies

Montane environments around the globe are biodiversity ‘hotspots’ and important reservoirs of genetic diversity. Montane species are also typically more vulnerable to environmental change than their low-elevation counterparts due to restricted ranges and dispersal limitations. Here we focus on two abundant congeneric mayflies (Baetis bicaudatus and B. tricaudatus) from montane streams over an elevation gradient spanning 1400 m. Using single-nucleotide polymorphism genotypes, we measured population diversity and vulnerability in these two species by: (i) describing genetic diversity and population structure across elevation gradients to identify mechanisms underlying diversification; (ii) performing spatially explicit landscape analyses to identify environmental drivers of differentiation; and (iii) identifying outlier loci hypothesized to underlie adaptive divergence. Differences in the extent of population structure in these species were evident depending upon their position along the elevation gradient. Heterozygosity, effective population sizes and gene flow all declined with increasing elevation, resulting in substantial population structure in the higher elevation species (B. bicaudatus). At lower elevations, populations of both species are more genetically similar, indicating ongoing gene flow. Isolation by distance was detected at lower elevations only, whereas landscape barriers better predicted genetic distance at higher elevations. At higher elevations, dispersal was restricted due to landscape effects, resulting in greater population isolation. Our results demonstrate differentiation over small spatial scales along an elevation gradient, and highlight the importance of preserving genetic diversity in more isolated high-elevation populations.

Freshwater vertebrate and invertebrate diversity patterns in an Andean-Amazon basin: implications for conservation efforts

The Napo Basin in Ecuador is an important drainage of the Amazon Basin, the most biodiverse ecosystem for freshwater species. At the same time, this basin has conspicuous information gaps on its biodiversity patterns and human threats. Here, we estimated the diversity distribution patterns of freshwater vertebrates and invertebrates in the Napo Basin, as a tool for present and future management and conservation efforts. Also, we assessed the spatial congruence of the diversity patterns observed between aquatic vertebrates and invertebrates. For this, we compiled occurrence records for 481 freshwater vertebrate species (amphibians, birds, mammals, reptiles, and fish), and 54 invertebrate families obtained across an altitudinal gradient of the basin (200–4500 m). Using these occurrence records and environmental variables, we modeled the distribution of each vertebrate species and invertebrate family. Then, we stacked these distributions to build species richness maps for vertebrates, and a family richness map for invertebrates. We found that the most diverse areas for vertebrate species are the lowlands (<600 m), whereas richness of invertebrate families peaks at higher elevations (lower montane forests). Congruence among species richness patterns of the five vertebrate groups was high (r = 0.66), with fish being the best predictor for vertebrates (r = 0.78). However, congruence decreased at higher elevations (r = 0.14), suggesting that specific species or habitat-based approaches should be used in the highlands. Also, we found a high correlation between species and family richness of freshwater invertebrates (r = 0.66), suggesting that family richness of invertebrates could be used as a surrogate of species richness in this basin. We highlight this correlation because, at the watershed scale, it allows working with family groups where species-level taxonomy is challenging. Our results provide the first comprehensive representation of freshwater biodiversity patterns at high resolution in an Andean-Amazon basin, and calls attention to the need for incorporating different taxonomic groups when assessing diversity patterns. Given these different diversity patterns, conservation programs for this basin should incorporate both vertebrate and invertebrate groups as biodiversity indicators. Finally, our study provides a practical methodological guidance in the estimation of freshwater diversity in regions of scarce information with high conservation priority, such as the Andean-Amazon basins.

Discovery of new populations and DNA barcoding of the Arapahoe snowfly Arsapnia arapahoe (Plecoptera: Capniidae)

The Arapahoe Snowfly, Arsapnia arapahoe (Nelson & Kondratieff)was recently discovered in six different first-order streams outside of the Cache la Poudre River Basin where it was previously considered endemic. Specimens of A. arapahoe were always collected in much lower relative abundance, 1.09% (±2.3SD), than other sympatric adult capniids. The first mitochondrial deoxyribonucleic acid (DNA) barcodes for A. arapahoe and A. coyote (Nelson & Baumann) are presented and compared with those of A. decepta. DNA barcoding was not able to differentiate between A. arapahoe and A. decepta Banks but it was able to indicate that A. coyote is specifically distinct.