Marianne Espeland

Diversity dynamics in New Caledonia: towards the end of the museum model?

BackgroundThe high diversity of New Caledonia has traditionally been seen as a result of its Gondwanan origin, old age and long isolation under stable climatic conditions (the museum model). Under this scenario, we would expect species diversification to follow a constant rate model. Alternatively, if New Caledonia was completely submerged after its breakup from Gondwana, as geological evidence indicates, we would expect species diversification to show a characteristic slowdown over time according to a diversity-dependent model where species accumulation decreases as space is filled.ResultsWe reanalyze available datasets for New Caledonia and reconstruct the phylogenies using standardized methodologies; we use two ultrametrization alternatives; and we take into account phylogenetic uncertainty as well as incomplete taxon sampling when conducting diversification rate constancy tests. Our results indicate that for 8 of the 9 available phylogenies, there is significant evidence for a diversification slowdown. For the youngest group under investigation, the apparent lack of evidence of a significant slowdown could be because we are still observing the early phase of a logistic growth (i.e. the clade may be too young to exhibit a change in diversification rates).ConclusionsOur results are consistent with a diversity-dependent model of diversification in New Caledonia. In opposition to the museum model, our results provide additional evidence that original New Caledonian biodiversity was wiped out during the episode of submersion, providing an open and empty space facilitating evolutionary radiations.

Ancient Neotropical origin and recent recolonisation: Phylogeny, biogeography and diversification of the Riodinidae (Lepidoptera: Papilionoidea)

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.

Pleistocene climate change promoted rapid diversification of aquatic invertebrates in Southeast Australia

BackgroundThe Pleistocene Ice Ages were the most recent geohistorical event of major global impact, but their consequences for most parts of the Southern hemisphere remain poorly known. We investigate a radiation of ten species of Sternopriscus, the most species-rich genus of epigean Australian diving beetles. These species are distinct based on genital morphology but cannot be distinguished readily by mtDNA and nDNA because of genotype sharing caused by incomplete lineage sorting. Their genetic similarity suggests a Pleistocene origin.ResultsWe use a dataset of 3858 bp of mitochondrial and nuclear DNA to reconstruct a phylogeny of Sternopriscus using gene and species trees. Diversification analyses support the finding of a recent rapid speciation event with estimated speciation rates of up to 2.40 species per MY, which is considerably higher than the proposed average rate of 0.16 species per MY for insects. Additionally, we use ecological niche modeling and analyze data on habitat preferences to test for niche divergence between species of the recent Sternopriscus radiation. These analyses show that the species can be characterized by a set of ecological variables referring to habitat, climate and altitude.ConclusionsOur results suggest that the repeated isolation of populations in glacial refugia might have led to divergent ecological adaptations and the fixation of morphological traits supporting reproductive isolation and therefore may have promoted speciation. The recent Sternopriscus radiation fulfills many characteristics of a species flock and would be the first described example of an aquatic insect species flock. We argue that the species of this group may represent a stage in speciation past the species flock condition because of their mostly broad and often non-overlapping ranges and preferences for different habitat types.

Phylogeny of the Polycentropodidae (Insecta: Trichoptera) based on protein-coding genes reveal non-monophyletic genera.

We tested the previous hypotheses of the phylogenetic position and monophyly of the caddisfly family Polycentropodidae. We also tested previous hypotheses about the internal generic relationship within the family by including 15 ingroup genera, many of them also represented by the genotype. All families that were previously taxonomically associated with the polycentropodids were included in the analysis. The total data set of 2225 bp representing sequences of combined nuclear and mitochondrial genes and 171 taxa, was analyzed using Bayesian inference. We found strong support for a monophyletic Polycentropodidae with Ecnomidae as the closest sister group. The recently erected families Kambaitipsychidae and Pseudoneureclipsidae were monophyletic and distantly related to the Polycentropodidae. Within Polycentropodidae, monophyly and validity of the genera Neucentropus, Neureclipsis, Cyrnus, Holocentropus, Tasmanoplegas, Pahamunaya, Cernotina and Cyrnellus was strongly supported, while the genera Polycentropus, Polyplectropus, Plectrocnemia, Placocentropus and Nyctiophylax were all polyphyletic. The New Caledonian species were polyphyletic and represented three distinct clades. The sister group to the New Caledonian clades are from Australia, New Zealand and Chile, respectively. The Vanuatu species evolved after dispersal from the Fiji Islands. New internal primers for cytochrome oxidase I sequences of Trichoptera are introduced.

Diversity Dynamics in Nymphalidae Butterflies: Effect of Phylogenetic Uncertainty on Diversification Rate Shift Estimates

The species rich butterfly family Nymphalidae has been used to study evolutionary interactions between plants and insects. Theories of insect-hostplant dynamics predict accelerated diversification due to key innovations. In evolutionary biology, analysis of maximum credibility trees in the software MEDUSA (modelling evolutionary diversity using stepwise AIC) is a popular method for estimation of shifts in diversification rates. We investigated whether phylogenetic uncertainty can produce different results by extending the method across a random sample of trees from the posterior distribution of a Bayesian run. Using the MultiMEDUSA approach, we found that phylogenetic uncertainty greatly affects diversification rate estimates. Different trees produced diversification rates ranging from high values to almost zero for the same clade, and both significant rate increase and decrease in some clades. Only four out of 18 significant shifts found on the maximum clade credibility tree were consistent across most of the sampled trees. Among these, we found accelerated diversification for Ithomiini butterflies. We used the binary speciation and extinction model (BiSSE) and found that a hostplant shift to Solanaceae is correlated with increased net diversification rates in Ithomiini, congruent with the diffuse cospeciation hypothesis. Our results show that taking phylogenetic uncertainty into account when estimating net diversification rate shifts is of great importance, as very different results can be obtained when using the maximum clade credibility tree and other trees from the posterior distribution.

Description of a new genus for Euptychia hilara (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae)

Based on external morphology, food plant records for caterpillars, and molecular analysis, Euptychia hilara (C. Felder & R. Felder, 1867) is removed from Euptychia Hübner 1818. A new genus, Inbio Nakahara & Espeland gen. nov., is proposed for this taxon. Inbio hilara comb. nov. is a member of a monophyletic clade containing Cyllopsis Felder, 1869, Paramacera Butler, 1868, and Atlanteuptychia Freitas, Barbosa & Mielke, 2013, although it can be morphologically distinguished from these genera. Lectotypes for Neonympha hilara C. Felder & R. Felder, 1867 and Euptychia anacleta Butler, 1877 (a synonym of E. hilara) are designated herein.

Phylogeny of the Aphnaeinae: myrmecophilous African butterflies with carnivorous and herbivorous life histories

The Aphnaeinae (Lepidoptera: Lycaenidae) are a largely African subfamily of 278 described species that exhibit extraordinary life‐history variation. The larvae of these butterflies typically form mutualistic associations with ants, and feed on a wide variety of plants, including 23 families in 19 orders. However, at least one species in each of 9 of the 17 genera is aphytophagous, parasitically feeding on the eggs, brood or regurgitations of ants. This diversity in diet and type of symbiotic association makes the phylogenetic relations of the Aphnaeinae of particular interest. A phylogenetic hypothesis for the Aphnaeinae was inferred from 4.4 kb covering the mitochondrial marker COI and five nuclear markers (wg, H3, CAD, GAPDH and EF1α) for each of 79 ingroup taxa representing 15 of the 17 currently recognized genera, as well as three outgroup taxa. Maximum Parsimony, Maximum Likelihood and Bayesian Inference analyses all support Heaths systematic revision of the clade based on morphological characters. Ancestral range inference suggests an African origin for the subfamily with a single dispersal into Asia. The common ancestor of the aphnaeines likely associated with myrmicine ants in the genus Crematogaster and plants of the order Fabales.

Anchored phylogenomics illuminates the skipper butterfly tree of life

BackgroundButterflies (Papilionoidea) are perhaps the most charismatic insect lineage, yet phylogenetic relationships among them remain incompletely studied and controversial. This is especially true for skippers (Hesperiidae), one of the most species-rich and poorly studied butterfly families.MethodsTo infer a robust phylogenomic hypothesis for Hesperiidae, we sequenced nearly 400 loci using Anchored Hybrid Enrichment and sampled all tribes and more than 120 genera of skippers. Molecular datasets were analyzed using maximum-likelihood, parsimony and coalescent multi-species phylogenetic methods.ResultsAll analyses converged on a novel, robust phylogenetic hypothesis for skippers. Different optimality criteria and methodologies recovered almost identical phylogenetic trees with strong nodal support at nearly all nodes and all taxonomic levels. Our results support Coeliadinae as the sister group to the remaining skippers, the monotypic Euschemoninae as the sister group to all other subfamilies but Coeliadinae, and the monophyly of Eudaminae plus Pyrginae. Within Pyrginae, Celaenorrhinini and Tagiadini are sister groups, the Neotropical firetips, Pyrrhopygini, are sister to all other tribes but Celaenorrhinini and Tagiadini. Achlyodini is recovered as the sister group to Carcharodini, and Erynnini as sister group to Pyrgini. Within the grass skippers (Hesperiinae), there is strong support for the monophyly of Aeromachini plus remaining Hesperiinae. The giant skippers (Agathymus and Megathymus) once classified as a subfamily, are recovered as monophyletic with strong support, but are deeply nested within Hesperiinae.ConclusionsAnchored Hybrid Enrichment sequencing resulted in a large amount of data that built the foundation for a new, robust evolutionary tree of skippers. The newly inferred phylogenetic tree resolves long-standing systematic issues and changes our understanding of the skipper tree of life. These resultsenhance understanding of the evolution of one of the most species-rich butterfly families.

Molecular phylogeny of Sericostomatoidea (Trichoptera) with the establishment of three new families

We inferred the phylogenetic relationships among 58 genera of Sericostomatoidea, representing all previously accepted families as well as genera that were not placed in established families. The analyses were based on five fragments of the protein coding genes carbamoylphosphate synthetase (CPSase of CAD), isocitrate dehydrogenase (IDH), Elongation factor 1a (EF‐1a), RNA polymerase II (POL II) and cytochrome oxidase I (COI). The data set was analysed using Bayesian methods with a mixed model, raxml, and parsimony. The various methods generated slightly different results regarding relationships among families, but the shared results comprise support for: (i) a monophyletic Sericostomatoidea; (ii) a paraphyletic Parasericostoma due to inclusion of Myotrichia murina, leading to synonymization of Myotrichia with Parasericostoma; (iii) a polyphyletic Sericostomatidae, which is divided into two families, Sericostomatidae sensu stricto and Parasericostomatidae fam.n.; (iv) a polyphyletic Helicophidae which is divided into Helicophidae sensu stricto and Heloccabucidae fam.n.; (v) hypothesized phylogenetic placement of the former incerta sedis genera Ngoya, Seselpsyche and Karomana; (vi) a paraphyletic Costora (Conoesucidae) that should be divided into several genera after more careful examination of morphological data; (vii) reinstatement of Gyrocarisa as a valid genus within Petrothrincidae. A third family, Ceylanopsychidae fam.n., is established based on morphological characters alone. A hypothesis of the relationship among 14 of the 15 families in the superfamily is presented. A key to the families is presented based on adults (males). Taxonomic history, diagnosis, habitat preference and distribution data for all sericostomatoid families are presented.

Seven new species of Chimarra (Trichoptera Philopotamidae) from Malawi

For the first time species of caddisflies in the genus Chimarra Stephens 1829 are reported from Malawi. The following new species are described: Chimarra zombaensis, C. flaviseta, C. chichewa, C. circumverta, C. mulanjae, C. psittacus and C. calidopectoris. The descriptions add to the knowledge of Afrotropical diversity in the order Trichoptera.