Tropical lowland areas have often been seen as the centres of terrestrial species proliferation, but recent evidence suggests that young species may be more frequent in montane areas. Several montane ...speciation modes have been proposed, but their relative frequencies and predominant evolutionary sequence remain unclear because so few biogeographic and phylogenetic studies have tested such questions. I use morphological data to generate a phylogenetic hypothesis for all 11 species of the riodinid butterfly genus Ithomiola (Riodininae: Mesosemiini: Napaeina). These species are shown here to be all strictly geographically and elevationally allo- or parapatrically distributed with respect to their closest relatives in lowland and montane regions throughout the Neotropics. The overwhelming pattern in Ithomiola is of repeated upward parapatric speciation across an elevational gradient, and the genus appears to provide the clearest example to date of vertical montane speciation. All of the young derived species are montane and all of the old basal species are confined to the lowlands, supporting the hypothesis of montane regions largely as 'species pumps' and lowland regions as 'museums'. Possible reasons for the post-speciation maintenance of parapatric ranges in Ithomiola are discussed.
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•The first higher-level molecular phylogeny of riodinid butterflies is presented.•Ancient Neotropical origin with dispersal into the Old World and back to the Neotropics.•Initial ...diversification by far predates the start of the climatic fluctuations in the Pleistocene.•The Neotropical Styx and Corrachia are part of the Old World subfamily Nemeobiinae.•Diversification has increased in New World taxa and decreased in Old World taxa.
We present the first dated higher-level phylogenetic and biogeographic analysis of the butterfly family Riodinidae. This family is distributed worldwide, but more than 90% of the c. 1500 species are found in the Neotropics, while the c. 120 Old World species are concentrated in the Southeast Asian tropics, with minor Afrotropical and Australasian tropical radiations, and few temperate species. Morphologically based higher classification is partly unresolved, with genera incompletely assigned to tribes. Using 3666bp from one mitochondrial and four nuclear markers for each of 23 outgroups and 178 riodinid taxa representing all subfamilies, tribes and subtribes, and 98 out of 145 described genera of riodinids, we estimate that Riodinidae split from Lycaenidae about 96Mya in the mid-Cretaceous and started to diversify about 81Mya. The Riodinidae are monophyletic and originated in the Neotropics, most likely in lowland proto-Amazonia. Neither the subfamily Euselasiinae nor the Nemeobiinae are monophyletic as currently constituted. The enigmatic, monotypic Neotropical genera Styx and Corrachia (most recently treated in Euselasiinae: Corrachiini) are highly supported as derived taxa in the Old World Nemeobiinae, with dispersal most likely occurring across the Beringia land bridge during the Oligocene. Styx and Corrachia, together with all other nemeobiines, are the only exclusively Primulaceae-feeding riodinids. The steadily increasing proliferation of the Neotropical Riodininae subfamily contrasts with the decrease in diversification in the Old World, and may provide insights into factors influencing the diversification rate of this relatively ancient clade of Neotropical insects.
Aim
The biodiversity crisis has highlighted the need to assess and map biodiversity in order to prioritize conservation efforts. Clearwing butterflies (tribe Ithomiini) have been proposed as ...biological indicators for habitat quality in Neotropical forests, which contain the world's richest biological communities. Here, we provide maps of different facets of Ithomiini diversity across the Neotropics to identify areas of evolutionary and ecological importance for conservation and evaluate their overlap with current anthropogenic threats.
Location
Neotropics.
Methods
We ran species distribution models on a data set based on 28,986 georeferenced occurrences representing 388 ithomiine species to generate maps of geographic rarity, taxonomic, phylogenetic and Müllerian mimetic wing pattern diversity. We quantified and mapped the overlap of diversity hotspots with areas threatened by or providing refuge from current anthropogenic pressures.
Results
The eastern slopes of the Andes formed the primary hotspot of taxonomic, phylogenetic and mimetic diversity, with secondary hotspots in Central America and the Atlantic Forest. Most diversity indices were strongly spatially correlated. Nevertheless, species‐poor communities on the Pacific slopes of the Andes also sheltered some of the geographically rarest species. Overall, tropical montane forests that host high species and mimetic diversity as well as rare species and mimicry rings appeared particularly under threat.
Main conclusions
Remote parts of the Upper Amazon may act as refuges against current anthropogenic pressures for a limited portion of Ithomiini diversity. Furthermore, it is likely that the current threat status may worsen with ongoing climate change and deforestation. In this context, the tropical Andes occupy a crucial position as the primary hotspot for multiple facets of biodiversity for ithomiine butterflies, as they do for angiosperms, tetrapods and other insect taxa. Our results support the role of ithomiine butterflies as a suitable flagship indicator group for Neotropical butterfly diversity and reinforce the position of the tropical Andes as a flagship region for biodiversity conservation in general, and insect and butterfly conservation in particular.
A fully resolved cladogram for 19 species in the Charis cleonus group of riodinid butterflies, which have closely parapatric ranges throughout the Amazon basin, is used to derive an area cladogram ...for the region. This represents the first comprehensive species-level analysis using insects and results in a hypothesis of Amazonian area relationships that is the most detailed to date. The Charis area cladogram is interpreted as supporting an historical vicariant split between the Guianas and the remainder of the Amazon and then between the upper and lower Amazon. The latter two clades can be further divided into the six most widely recognized areas of endemism and even smaller endemic centers within these, some of which, especially along the Madeira and lower Amazon Rivers, have never been previously hypothesized for butterflies. The overall pattern of historical interrelationships indicated is Guiana + ((Rondônia + (Pará + Belém)) + (Imeri + (Napo + Inambari))). The area relationships for riodinid butterflies show substantial congruence with those presented from the literature for amphibians, reptiles, birds, primates, rodents, and marsupials, suggesting a common vicariant history for these organisms. A summary area cladogram generated by combining area cladograms for all the aforementioned groups of organisms indicated the pattern of historical interrelationships to be (Guiana + (Rondônia + (Pará + Belém))) + (Imeri + (Napo + Inambari)). Charis cleonus group species distributions are noticeably larger around the upland periphery of Amazonia and smaller in the central and lower regions. A significant positive correlation between the proportion of range area above 100 m and total range size for each species is used to suggest that past sea-level rises may explain smaller range sizes in low-lying regions and that riverine barriers have been important in shaping the current distribution of C. cleonus group species. Corresponding Editor: T. Smith
The 'aegrota species group' of the Neotropical nymphalid genus Caeruleuptychia
Forster, 1964, in addition to three other superficially similar, enigmatic species in the genus, are revised. A ...lectotype is designated for Euptychia aegrota Butler, 1867, E. aetherialis
Butler, 1877
stat. rev., E. helios
Weymer, 1911 and E. pilata Butler, 1867, and C. aetherialis is resurrected from its synonymy with C. aegrota. Caeruleuptychia helios caelestissima
Brévignon, 2010, syn. nov., and Magneuptychia keltoumae
Brévignon & Benmesbah, 2012, syn. nov. are both regarded as junior subjective synonyms of C. helios (Weymer, 1911), as a result of the discovery and first illustration of the female of this taxon. The female of C. aegrota is also described and illustrated for the first time, and three new species, C. trembathi Willmott, Nakahara, Hall & Neild, sp. nov., C. scripta Nakahara, Zacca & Huertas, sp. nov., and C. maryzenderae Lamas & Nakahara, sp. nov. are described and named. We analyze morphological and molecular data separately, in addition to combining morphological data with molecular data, to provide the first phylogenetic hypothesis for the taxa treated in this revision.
A new species, Adelpha margarita Willmott & Hall, new species, is described from Andean cloud forest habitats from southern Ecuador to Bolivia. Adelpha margarita garleppi Willmott, new subspecies, is ...described for southern Peruvian and Bolivian individuals, which differ from the nominate subspecies in having complete orange postdiscal bands on the dorsal surface. The new species differs from related species in the Adelpha serpa group in wing pattern, DNA sequence data and habitat. A lectotype is designated for Adelpha seriphia therasia Fruhstorfer, because the type series of this name contains individuals of both A. seriphia and A. margarita. Neighbor-joining and maximum parsimony analyses of 579 bp of the mitochondrial cytochrome oxidase I (COI) ‘barcode’ region, for 27 Adelpha specimens representing 9 species and 13 taxa, suggest that the closest relative to A. margarita is the Central American to west Andean taxon A. seriphia godmani Fruhstorfer. The DNA sequence data, coupled with a re-analysis of museum specimens, suggest that Adelpha godmani should be treated as a distinct species (revised status). Finally, a new subspecies, Adelpha justina pichincha Willmott & Hall, new subspecies, is described from Pichincha province in western Ecuador.
We describe a new species of extinct riodinid butterfly, Voltinia dramba, from Oligo-Miocene Dominican amber (15-25 Myr ago). This appears to be the first butterfly to be taxonomically described from ...amber, and the first adult riodinid fossil. The series of five specimens represents probably the best-preserved fossil record for any lepidopteran. The phenomenon of extant Voltinia females ovipositing on arboreal epiphytes probably explains the discovery of multiple female V. dramba specimens in amber. Voltinia dramba appears to be one of many extinct butterfly species on Hispaniola. The northwestern Mexican distribution of the explicitly hypothesized sister species, the extant V. danforthi, supports the hypothesis that V. dramba reached Hispaniola by the 'proto-Greater Antillean arc', dating the divergence of V. dramba and V. danforthi to 40-50 Myr ago. This date is contemporaneous with the oldest known butterfly fossils, and implies a more ancient date of origin for many of the higher-level butterfly taxa than is often conceded.
The small Neotropical riodinid genus Minstrellus (Lepidoptera: Riodinidae) is described and revised. It seems to be most closely related to Pseudonymphidia Callaghan, in the incertae sedis section of ...the Riodininae. A phylogenetic analysis of its four species, using 11 characters of adult morphology, generated a single most parsimonious cladogram with the following structure: grandis + (nivosa + (leucotopus + emphatica)). The species watkinsi d’Abrera, like all the taxa of Minstrellus, was described in Ematurgina Röber (a synonym of Synargis Hübner) and briefly subsequently treated in Zelotaea Bates, but it is here transferred to Machaya Hall & Willmott (n. comb.). The taxon emphatica Stichel is raised from a subspecies of leucotopus Stichel to the rank of species (n. stat.), and the name candida Le Cerf is synonymized with emphatica (n. syn.).
Inventory of the caterpillars, their food plants and parasitoids began in 1978 for today's Area de Conservacion Guanacaste (ACG), in northwestern Costa Rica. This complex mosaic of 120 000 ha of ...conserved and regenerating dry, cloud and rain forest over 0-2000 m elevation contains at least 10 000 species of non-leaf-mining caterpillars used by more than 5000 species of parasitoids. Several hundred thousand specimens of ACG-reared adult Lepidoptera and parasitoids have been intensively and extensively studied morphologically by many taxonomists, including most of the co-authors. DNA barcoding -- the use of a standardized short mitochondrial DNA sequence to identify specimens and flush out undisclosed species -- was added to the taxonomic identification process in 2003. Barcoding has been found to be extremely accurate during the identification of about 100 000 specimens of about 3500 morphologically defined species of adult moths, butterflies, tachinid flies, and parasitoid wasps. Less than 1% of the species have such similar barcodes that a molecularly based taxonomic identification is impossible. No specimen with a full barcode was misidentified when its barcode was compared with the barcode library. Also as expected from early trials, barcoding a series from all morphologically defined species, and correlating the morphological, ecological and barcode traits, has revealed many hundreds of overlooked presumptive species. Many but not all of these cryptic species can now be distinguished by subtle morphological and/or ecological traits previously ascribed to 'variation' or thought to be insignificant for species-level recognition. Adding DNA barcoding to the inventory has substantially improved the quality and depth of the inventory, and greatly multiplied the number of situations requiring further taxonomic work for resolution.
A new riodinid species in the tribe Symmachiini, Lucillella arcoirisa Hall & Willmott n. sp., is described from the eastern Andes of Ecuador. All five true members of the genus Lucillella Strand, ...1932, are figured, and their known geographic ranges are mapped. Lucillella is hypothesized to be monophyletic with the exclusion of suberra Hewitson, 1877, and the sister genus is hypothesized to be Esthemopsis C. & R. Felder, 1865. The new Lucillella species was discovered in the forest canopy using the single rope technique. The equipment and procedures used in this canopy access method are described and illustrated.
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