Carlos E. Giraldo

Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina

Understanding why species richness peaks along the Andes is a fundamental question in the study of Neotropical biodiversity. Several biogeographic and diversification scenarios have been proposed in the literature, but there is confusion about the processes underlying each scenario, and assessing their relative contribution is not straightforward. Here, we propose to refine these scenarios into a framework which evaluates four evolutionary mechanisms: higher speciation rate in the Andes, lower extinction rates in the Andes, older colonization times and higher colonization rates of the Andes from adjacent areas. We apply this framework to a species‐rich subtribe of Neotropical butterflies whose diversity peaks in the Andes, the Godyridina (Nymphalidae: Ithomiini). We generated a time‐calibrated phylogeny of the Godyridina and fitted time‐dependent diversification models. Using trait‐dependent diversification models and ancestral state reconstruction methods we then compared different biogeographic scenarios. We found strong evidence that the rates of colonization into the Andes were higher than the other way round. Those colonizations and the subsequent local diversification at equal rates in the Andes and in non‐Andean regions mechanically increased the species richness of Andean regions compared to that of non‐Andean regions (‘species‐attractor’ hypothesis). We also found support for increasing speciation rates associated with Andean lineages. Our work highlights the importance of the Andean slopes in repeatedly attracting non‐Andean lineages, most likely as a result of the diversity of habitats and/or host plants. Applying this analytical framework to other clades will bring important insights into the evolutionary mechanisms underlying the most species‐rich biodiversity hotspot on the planet.

North Andean origin and diversification of the largest ithomiine butterfly genus

The Neotropics harbour the most diverse flora and fauna on Earth. The Andes are a major centre of diversification and source of diversity for adjacent areas in plants and vertebrates, but studies on insects remain scarce, even though they constitute the largest fraction of terrestrial biodiversity. Here, we combine molecular and morphological characters to generate a dated phylogeny of the butterfly genus Pteronymia (Nymphalidae: Danainae), which we use to infer spatial, elevational and temporal diversification patterns. We first propose six taxonomic changes that raise the generic species total to 53, making Pteronymia the most diverse genus of the tribe Ithomiini. Our biogeographic reconstruction shows that Pteronymia originated in the Northern Andes, where it diversified extensively. Some lineages colonized lowlands and adjacent montane areas, but diversification in those areas remained scarce. The recent colonization of lowland areas was reflected by an increase in the rate of evolution of species’ elevational ranges towards present. By contrast, speciation rate decelerated with time, with no extinction. The geological history of the Andes and adjacent regions have likely contributed to Pteronymia diversification by providing compartmentalized habitats and an array of biotic and abiotic conditions, and by limiting dispersal between some areas while promoting interchange across others.

Diferenciación morfológica y molecular de Oleria makrena (Hewitson) y Oleria fumata (Haensch) (Lepidoptera: Ithomiinae)

Oleria (Olerinii tribe) is a genus of great ecological and evolutionary importance among the Ithomiinae (Nymphalidae). Species identification is very complex since most of them are involved in mimicry rings and share large morphological similarities. Ithomiinae specimens collected in the Antioquia Southwest - Colombia, showed confuse wing color pattern generating diffi culties to define the presence of one or two species at the sampled region. Collected specimens were studied in detail using male and female genitalia and nucleotide sequences besides traditional character of wing color and venation. Oleria makrena (Hewitson) and Oleria fumata (Haensch) were registered and the relative utility of each character for species identification was defi ned. For males, wing color pattern was enough to differentiate both species, and the separation of specimens by this character was congruent with the separation obtained by using genitalia and DNA sequences. However, for females, wing patterns were not enough for the adequate separation or species and genitalia-based characters were necessary. DNA sequences of a short fragment of the mitochondrial gene ND4 allowed for the rapid and precise differentiation of both species, coinciding with the separation using genitalia for both males and females. The analysis of 579 base pairs (bp) indicated 46 nucleotide differences between both species.

Differences in butterfly (Nymphalidae) diversity between hillsides and hilltop forest patches in the northern Andes

Andean montane butterflies have unique diversity patterns in each mountain system, and the knowledge on how their community diversity changes at local scale has potential important implications on designing regional conservation strategies. The multiplicative partition of the diversity and the effective number of species provides a useful tool in studying the patterns of diversity in heterogeneous Andean cloud forest habitats. Here, we evaluated diversity in three nearby sites in the Andean Central Cordillera of Colombia, two hillsides – on western (Cauca canyon) and eastern (Aburrá valley) slopes – and one hilltop. We collected a total of 1039 individuals belonging to 55 species of butterflies. Alpha diversity differs between hillsides and the hilltop. Beta diversity in the three areas show differences in richness (q0) and in diversity of the order of magnitude of q1 and q2, with the major differences between hillsides and hilltop. This study shows important differences in butterfly (Nymphalidae) assemblages on a local scale, and suggests that any efforts directed towards comprehensive protection of a given area in a heterogeneous mountain landscape must be focused on the complete forested area, not being limited to a hilltop or to a single hillside.

Description of two new species of the Neotropical genus Yphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from the ‘renata clade’

This paper describes two new species of Neotropical butterflies: Yphthimoides blanquita Barbosa, Marín and Freitas sp. nov., from the dry forest of northwestern Colombia, and Yphthimoides nareta Barbosa & Freitas sp. nov. from northeastern Brazil, based on morphological and molecular data. Adult morphology, including wing shape and pattern as well as male genitalia, is described in detail. Furthermore, analysis of the mitochondrial CoxI ‘barcode’ showed that both new species are quite distinct from all similar Yphthimoides species and additionally, the ‘renata clade’ is defined based on the presence of cornuti in the aedeagus. http://zoobank.org/urn:lsid:zoobank.org:pub:22CA85FF-9682-4016-A300-AB11C71CEDA2

Contrasting patterns of Andean diversification among three diverse clades of Neotropical clearwing butterflies

Abstract The Neotropical region is the most biodiverse on Earth, in a large part due to the highly diverse tropical Andean biota. The Andes are a potentially important driver of diversification within the mountains and for neighboring regions. We compared the role of the Andes in diversification among three subtribes of Ithomiini butterflies endemic to the Neotropics, Dircennina, Oleriina, and Godyridina. The diversification patterns of Godyridina have been studied previously. Here, we generate the first time‐calibrated phylogeny for the largest ithomiine subtribe, Dircennina, and we reanalyze a published phylogeny of Oleriina to test different biogeographic scenarios involving the Andes within an identical framework. We found common diversification patterns across the three subtribes, as well as major differences. In Dircennina and Oleriina, our results reveal a congruent pattern of diversification related to the Andes with an Andean origin, which contrasts with the Amazonian origin and multiple Andean colonizations of Godyridina. In each of the three subtribes, a clade diversified in the Northern Andes at a faster rate. Diversification within Amazonia occurred in Oleriina and Godyridina, while virtually no speciation occurred in Dircennina in this region. Dircennina was therefore characterized by higher diversification rates within the Andes compared to non‐Andean regions, while in Oleriina and Godyridina, we found no difference between these regions. Our results and discussion highlight the importance of comparative approaches in biogeographic studies.

Immature stages of Pagyris cymothoe cymothoe (Hewitson, 1855) (Lepidoptera, Danainae, Ithomiini)

This paper describes the immature stages of Pagyris cymothoe cymothoe (Hewitson, 1855) based on samples collected in Venezuela and Colombia in 1991 and 2013, respectively. Eggs were collected on Brugmansiacandida Pers. and Brugmansiasuaveolens (Willd.), and reared on B.suaveolens under laboratory conditions at 25 ± 2°C. Larvae are gregarious, passing through five instars; the duration of life cycle (from egg to adult) lasted about 28 days. Detailed morphology of the egg and first instar chaetotaxy is presented. This new life history is compared with the life histories of Pagyris ulla (Hewitson, 1857) and of the sister genus Placidina dAlmeida, 1928.

Ithomiini Butterflies (Lepidoptera: Nymphalidae) of Antioquia, Colombia

Colombia is one of the most biodiverse countries on the planet. However, economic and scientific investment in completing inventories of its biodiversity has been relatively poor in comparison with other Neotropical countries. Butterflies are the best studied group of invertebrates, with the highest proportion of known to expected species. More than 3,200 species of butterflies have been recorded in Colombia, although the study of the still many unexplored areas will presumably increase this number. This work provides a list of Ithomiini butterflies collected in the department of Antioquia and estimates the total number of species present, based on revision of entomological collections, records in the literature and field work performed between 2003 and 2011. The list includes 99 species and 32 genera, representing 27% of all Ithomiini species. We report 50 species of Ithomiini not formerly listed from Antioquia, and found the highest diversity of ithomiine species to be at middle elevations (900–1,800xa0m). The mean value of the Chao2 estimator for number of species in Antioquia is 115 species, which is close to a predicted total of 109 based on known distributions of other Ithomiini not yet recorded from the department. Nine species are potentially of particular conservation importance because of their restricted distributions, and we present range maps for each species. We also highlight areas in Antioquia with a lack of biodiversity knowledge to be targeted in future studies. This paper contributes to mapping the distribution of the Lepidoptera of Antioquia department in particular and of Colombia in general.