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Discussion

Understanding global patterns of biodiversity distribution is at the core of macroecology (e.g., Gaston 2000; Gaston and Blackburn 2000; Crisp et al. 2009), and represents a basis for identifying regions that should be the focus for conservation (e.g., Myers et al. 2000). In our study we used six different measures to assess large-scale patterns of diversity for three butterfly genera in the Neotropics. These measures capture different attributes of biodiversity and their simultaneous use contributes to a better picture of how they are related. We found that the patterns of distribution of species richness, phylogenetic diversity and mimicry diversity remain relatively consistent across different ithomiine genera. However, sensitivity to extinction related to mutualistic interactions strongly varies across regions and shows incongruence across the groups studied here.

Hotspots of Species Richness and Phylogenetic Diversity in the Neotropics

For the three genera studied here, the eastern part of the Andes is one of the regions with highest species richness and phylogenetic diversity (PD and ES) while the poorest regions are the lower-Amazon, the Cerrado and the Atlantic-forest. Napeogenes and Oleria show a relatively similar secondary peak of diversity in the upper-Amazon. By contrast, Ithomia exhibits low diversity in the upper-Amazon but maximum species richness in Central-America. The latter pattern is due mainly to the diversification of a single clade in Central-America, which explains the relatively intermediate values of PD and ES (i.e., not maximum values). Interestingly, the Central-America diversity peak also corresponds to a mountainous region.

Within each genus, species richness, PD and ES show a very strong pattern of covariation. This is likely due to the fact that these indices are summed across all species in a grid cell and are therefore strongly influenced by species richness (see Rodrigues et al. 2005; Davies and Cadotte 2011). This may be particularly important for the three genera studied here because their phylogenetic trees are rather balanced, resulting in no major differences in phylogenetic diversity among species (see Rodrigues et al. 2005 for an analysis of PD in this respect). But, as shown in previous works, the congruence among different indices is not perfect throughout the spectrum of species richness distribution. Here, differences among measures are more obvious in areas with intermediate or intermediate to low species richness. Differences between species and phylogenetic diversity are likely to be common for relatively low species richness areas, because such areas could harbour distantly related species and/or phylogenetically distinctive species, resulting in high PD and ES values. For example, Arponen and Zupan (chapter “Representing Hotspots of Evolutionary History in Systematic Conservation Planning for European Mammals”) found major differences between phylogenetic diversity and species richness for mammals in areas of low diversity in the north of Europe.

MPD captures the average relatedness of the pairs of species in each grid cell, and high values indicate the presence of pairs of distant relatives in species assemblages. As a mean value it is independent of species richness, but its variance increases with low species richness. However, it provides useful information related to the diversification history of a clade. For example, the increase of MPD for Oleria, from northwest toward southeast, is explained by phylogenetically independent colonisation of these regions.

One of the first studies investigating the usefulness of ithomiines as biogeographic indicators suggested that they could be good surrogates of total butterfly diversity in lowland Neotropical forests (Beccaloni and Gaston 1995). Our results are mostly consistent with that suggestion, since peaks in richness in the eastern Andes and upper Amazon, as identified here, have also been reported in Heliconius butterflies (Rosser et al. 2012) and the genus Adelpha (Willmott 2003).

Studies on other taxa have also found a pattern of high diversity in the upper Amazon, based on various different measures. For example, using a dataset of 50 clades (López-Osorio and Miranda-Esquivel 2010), found that species richness and evolutionary distinctiveness of several groups of vertebrates and some groups of insects and plants are high in the southern upper Amazon. But, unlike the genera studied here, they also found very high diversity in the Guianas (see also MirandaEsquivel chapter “Support in Area Prioritization Using Phylogenetic Information”). Similarly, Amori et al. 2013 noted that rodent diversity peaks in the upper Amazon, but also found diversity hotspots in the Guianas and Atlantic forest. Primates similarly show increasing diversity from east to west (Da Silva et al. 2005b), as well as birds (Haffer 1990), non-volant mammals (Costa et al. 2000) and plants (Ter Steege et al. 2000). Many factors are likely to contribute to the general high species richness in the western Amazon and Andean foothills. Along the eastern Andes, high turnover in abiotic conditions, habitat types, vegetation and host-plants for phytophagous insects, in addition to topological complexity, may explain a high species turnover within a grid cell, therefore increasing diversity. All these factors are also potential drivers of speciation, which also contributes to increase diversity. The diversification histories across geographical areas also account for patterns of diversity of different organisms. In the case of ithomiine butterflies, previous studies found that Napeogenes, Ithomia and Oleria likely originated in the northern Andes and subsequently diversified throughout both the Andes and the rest of the Neotropics (Elias et al. 2009; de-Silva et al. 2015; de-Silva et al. 2010). Shifts of altitudinal range and colour pattern are also correlated (Chazot et al. 2014) and are involved in speciation events (Jiggins et al. 2006; Elias et al. 2009), and may likely have increased speciation rate in montane regions. In addition, hostplant diversity has been proposed to drive diversification in phytophagous insects (Janz et al. 2006) and particularly in ithomiines (Willmott and Freitas 2006), whose Solanaceae hostplants are most diverse in the Andes and the upper Amazon, and to a lesser extent in the Atlantic Forest (Knapp 2002; PBI Solanum Project; nhm.ac.uk/ research-curation/research/projects/solanaceaesource/). Understanding the ecology and the diversification history of different groups of organisms may therefore lead to a better explanation of diversity patterns in the Neotropics and improve conservation strategies. For this reason, no single group of organism can be a good indicator of general patterns of diversity. Approaches that rely on a wide range of taxa (e.g., López-Osorio and Miranda-Esquivel 2010) are more powerful in this respect.

 
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