Showing papers by "Niklas Wahlberg published in 2012"

What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity

Abstract: The latitudinal diversity gradient (LDG) is one of the most striking ecological patterns on our planet. Determining the evolutionary causes of this pattern remains a challenging task. To address this issue, previous LDG studies have usually relied on correlations between environmental variables and species richness, only considering evolutionary processes indirectly. Instead, we use a phylogenetically integrated approach to investigate the ecological and evolutionary processes responsible for the global LDG observed in swallowtail butterflies (Papilionidae). We find evidence for the 'diversification rate hypothesis' with different diversification rates between two similarly aged tropical and temperate clades. We conclude that the LDG is caused by (1) climatically driven changes in both clades based on evidence of responses to cooling and warming events, and (2) distinct biogeographical histories constrained by tropical niche conservatism and niche evolution. This multidisciplinary approach provides new findings that allow better understanding of the factors that shape LDGs.

Molecular phylogenetics of Erebidae (Lepidoptera, Noctuoidea)

Abstract: As a step towards understanding the higher-level phylogeny and evolu- tionary affinities of quadrifid noctuoid moths, we have undertaken the first large-scale molecular phylogenetic analysis of the moth family Erebidae, including almost all subfamilies, as well as most tribes and subtribes. DNA sequence data for one mitochondrial gene (COI ) and seven nuclear genes (EF-1α, wingless, RpS5, IDH, MDH, GAPDH and CAD) were analysed for a total of 237 taxa, principally type genera of higher taxa. Data matrices (6407 bp in total) were analysed by parsimony with equal weighting and model-based evolutionary methods (maximum likelihood), which revealed a well-resolved skeleton phylogenetic hypothesis with 18 major lineages, which we treat here as subfamilies of Erebidae. We thus present a new phylogeny for Erebidae consisting of 18 moderate to strongly supported subfami- lies: Scoliopteryginae, Rivulinae, Anobinae, Hypeninae, Lymantriinae, Pangraptinae, Herminiinae, Aganainae, Arctiinae, Calpinae, Hypocalinae, Eulepidotinae, Toxocamp- inae, Tinoliinae, Scolecocampinae, Hypenodinae, Boletobiinae and Erebinae. Where possible, each monophyletic lineage is diagnosed by autapomorphic morphological character states, and within each subfamily, monophyletic tribes and subtribes can be circumscribed, most of which can also be diagnosed by morphological apomorphies. All additional taxa sampled fell within one of the four previously recognized quadrifid families (mostly into Erebidae), which are now found to include two unusual monobasic taxa from New Guinea: Cocytiinae (now in Erebidae: Erebinae) and Eucocytiinae (now in Noctuidae: Pantheinae).

Cretaceous origin and repeated tertiary diversification of the redefined butterflies

Abstract: Although the taxonomy of the ca 18 000 species of butterflies and skippers is well known, the family-level relationships are still debated. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of the superfamilies Papilionoidea, Hesperioidea and Hedyloidea to date based on morphological and molecular data. We reconstructed their phylogenetic relationships using parsimony and Bayesian approaches. We estimated times and rates of diversification along lineages in order to reconstruct their evolutionary history. Our results suggest that the butterflies, as traditionally understood, are paraphyletic, with Papilionidae being the sister-group to Hesperioidea, Hedyloidea and all other butterflies. Hence, the families in the current three superfamilies should be placed in a single superfamily Papilionoidea. In addition, we find that Hedylidae is sister to Hesperiidae, and this novel relationship is supported by two morphological characters. The families diverged in the Early Cretaceous but diversified after the Cretaceous‐Palaeogene event. The diversification of butterflies is characterized by a slow speciation rate in the lineage leading to Baronia brevicornis, a period of stasis by the skippers after divergence and a burst of diversification in the lineages leading to Nymphalidae, Riodinidae and Lycaenidae.

Unprecedented ichneumonid parasitoid wasp diversity in tropical forests

Abstract: The megadiverse parasitoid wasp family Ichneumonidae (Hymenoptera) is classically considered an exception to the extensively studied latitudinal diversity gradient: the majority of ichneumonid species are described from temperate regions. The gradient has been hypothesized to be dependent on the biology of the wasps, but recently questions of sampling and description biases have been raised. Here, we show with primary data that the species richness of Ichneumonidae is markedly underestimated in tropical areas and that latitudinal diversity patterns in the family remain uncharacterized. We discovered a startling 177 likely undescribed orthocentrine species with relatively low sampling effort in the forests of Central America and Amazonian Ecuador, over three times the previously known orthocentrine diversity in the world's tropics. Species accumulation curves reveal that we are just beginning to unveil the true extent of tropical orthocentrine diversity. We also found evidence for cryptic species; our DNA analysis revealed additional species not easily distinguishable using morphological characteristics. The difficulty in establishing species richness patterns of Ichneumonidae probably follows from the relative lack of taxonomic expertise and the low density of ichneumonid species throughout the landscape.

Climate‐driven diversity dynamics in plants and plant‐feeding insects

Abstract: The origin of species-rich insect–plant food webs has traditionally been explained by diversifying antagonistic coevolution between plant defences and herbivore counter-defences. However, recent studies combining paleoclimatic reconstructions with time-calibrated phylogenies suggest that variation in global climate determines the distribution, abundance and diversity of plant clades and, hence, indirectly influences the balance between speciation and extinction in associated herbivore groups. Extant insect communities tend to be richest on common plant species that have many close relatives. This could be explained either by climatedriven diffuse cospeciation between plants and insects, or by elevated speciation and reduced extinction in herbivore lineages associated with expanding host taxa (resources). Progress in paleovegetation reconstructions in combination with the rapidly increasing availability of fossil-calibrated phylogenies provide means to discern between these alternative hypotheses. In particular, the Diffuse cospeciation scenario predicts closely matching main diversification periods in plants and in the insects that feed upon them, while the Resource abundance-dependent diversification hypothesis predicts that both positive and negative responses of insect diversity are lagged in relation to host-plant availability. The dramatic Cenozoic changes in global climate provide multiple possibilities for studying the mechanisms by which climatic shifts may drive diversity dynamics in plants and insect herbivores.

Evolutionary history of the recruitment of conserved developmental genes in association to the formation and diversification of a novel trait

Abstract: The origin and modification of novel traits are important aspects of biological diversification. Studies combining concepts and approaches of developmental genetics and evolutionary biology have uncovered many examples of the recruitment, or co-option, of genes conserved across lineages for the formation of novel, lineage-restricted traits. However, little is known about the evolutionary history of the recruitment of those genes, and of the relationship between them -for example, whether the co-option involves whole or parts of existing networks, or whether it occurs by redeployment of individual genes with de novo rewiring. We use a model novel trait, color pattern elements on butterfly wings called eyespots, to explore these questions. Eyespots have greatly diversified under natural and sexual selection, and their formation involves genetic circuitries shared across insects. We investigated the evolutionary history of the recruitment and co-recruitment of four conserved transcription regulators to the larval wing disc region where circular pattern elements develop. The co-localization of Antennapedia, Notch, Distal-less, and Spalt with presumptive (eye)spot organizers was examined in 13 butterfly species, providing the largest comparative dataset available for the system. We found variation between families, between subfamilies, and between tribes. Phylogenetic reconstructions by parsimony and maximum likelihood methods revealed an unambiguous evolutionary history only for Antennapedia, with a resolved single origin of eyespot-associated expression, and many homoplastic events for Notch, Distal-less, and Spalt. The flexibility in the (co-)recruitment of the targeted genes includes cases where different gene combinations are associated with morphologically similar eyespots, as well as cases where identical protein combinations are associated with very different phenotypes. The evolutionary history of gene (co-)recruitment is consistent with both divergence from a recruited putative ancestral network, and with independent co-option of individual genes. The diversity in the combinations of genes expressed in association with eyespot formation does not parallel diversity in characteristics of the adult phenotype. We discuss these results in the context of inferring homology. Our study underscores the importance of widening the representation of phylogenetic, morphological, and genetic diversity in order to establish general principles about the mechanisms behind the evolution of novel traits.

Variation of Basal EROD Activities in Ten Passerine Bird Species – Relationships with Diet and Migration Status

Abstract: Inter-specific differences in animal defence mechanisms against toxic substances are currently poorly understood. The ethoxyresorufin-O-deethylase (EROD) enzyme plays an important role in defence against toxic chemicals in a wide variety of animals, and it is an important biomarker for environmental contamination. We compared basal hepatic EROD activity levels among ten passerine species to see if there is inter-specific variation in enzyme activity, especially in relation to their diet and migration status. Migratory insectivores showed higher EROD activity compared to granivores. We hypothesize that the variable invertebrate diet of migratory insectivores contains a wider range of natural toxins than the narrower diet of granivores. This may have affected the evolution of mixed function oxidases (MFO) system and enzyme activities. We further tested whether metabolic rates or relative liver size were associated with the variation in detoxification capacity. We found no association between EROD activity and relative (per mass unit) basal metabolic rate (BMR). Instead, EROD activity and relative liver mass (% of body mass) correlated positively, suggesting that a proportionally large liver also functions efficiently. Our results suggest that granivores and non-migratory birds may be more vulnerable to environmental contaminants than insectivores and migratory birds. The diet and migration status, however, are phylogenetically strongly connected to each other, and their roles cannot be fully separated in our analysis with only ten passerine species.

Exploration of data partitioning in an eight-gene data set: phylogeny of metalmark moths (Lepidoptera, Choreutidae)

Abstract: Molecular data sets for phylogenetic inference continue to increase in size, especially with respect to the number of genes sampled. As more and more genes are included in analyses, the importance of partitioning the data to avoid problems that can arise from underparameterization becomes more apparent. With an eight-gene data set from 38 metalmark moth species (12 genera represented) and three outgroups, we explored different data partitioning strategies and their influence on convergence and mixing of Markov Chains Monte Carlo in a Bayesian setting. We found that in larger data sets, with an increase in the number of partitions that are made a priori (e.g. by gene and codon position), convergence and mixing become poor. This problem can be overcome by using a recently published algorithm in which homologous sites are grouped into blocks with similar evolutionary rates that can then be modelled as separate data subsets. Using this novel approach to data partitioning, our analyses resolve with strong support relationships among the genera of metalmark moths. Support for the monophyly of the family, the two subfamilies and all genera except Hemerophila is strong. Hemerophila is broken into two separate clades, Hemerophila sensu stricto and another well-supported clade. To render Hemerophila monophyletic, we describe a new genus, Ornarantia Rota, gen. nov., and transfer 18 species from Hemerophila to it. The type species of Ornarantia is Hemerophila laciniosella Busck, 1914. (Less)

Diversification of Morpho butterflies (Lepidoptera, Nymphalidae): a re‐evaluation of morphological characters and new insight from DNA sequence data

Abstract: This study compiles previously published morphological, colour and behavioural characters and includes new DNA sequence data for eight markers (one mitochondrial and seven nuclear) to re-evaluate phylogenetic relationships and estimate times of divergence for Morpho butterflies using parsimony and Bayesian methods. We note an effect of missing data on phylogenetic inference and calculations of Partitioned Bremer Support. Morphology and DNA trees were moderately congruent, and the combined analyses of all data included elements of both sources. Both morphology and DNA support the monophyly of Morpho and the early separation of the sister pair M. marcus plus M. eugenia, but trees from different data sources are congruent mostly at derived nodes, and differ at several internal nodes. The analyses of combined data indicate that Morpho is composed of four clades each of which include one or more previously proposed subgenera. The subgenera Pessonia and Morpho were not monophyletic, and to address this issue we propose that Pessonia, syn.nov. be subsumed within Morpho. The ancestor of Morpho probably arose during the Oligocene, and most diversification seems to have occurred during the late Miocene. S-DIVA analysis suggests eastern Andean region as the ancestral area for Morpho, and that the South American Atlantic Forest was colonized multiple times.

Phylogeography of the threatened butterfly, the woodland brown Lopinga achine (Nymphalidae: Satyrinae): implications for conservation

Abstract: We have studied the phylogeography of the red-listed Palearctic butterfly Lopinga achine (Nymphalidae: Satyrinae) based on 1,450 base pairs of mitochondrial DNA sequences from 86 individuals representing 12 populations. Our results indicate a strong structuring of genetic variation, with among-population differences accounting for ca. 67% of the variation and almost all populations being significantly differentiated from each other. We surmise that the insular nature of populations as well as the low dispersal ability of the species has given rise to such a pattern. The genetic diversity within populations is low compared to that in other butterflies. Our results point to a scenario where the species originated in the Eastern Palearctic and expanded into Europe. Based on the analyses, we suggest that the Czech population merits the highest conservation priority. The two Swedish populations represent a distinct evolutionary lineage, and hence merit high conservation attention. The Estonian and Asian populations had the highest genetic diversity, and although we do not consider them to be under immediate threat, their genetic diversity should be conserved in the long term.

Euptychia boulleti (Le Cerf) n. comb. (Lepidoptera: Nymphalidae: Satyrinae), a rare and endangered butterfly from Southeastern Brazil.

Abstract: This paper discusses the systematic position of the rare and endangered satyrine butterfly Caenoptychia boulleti Le Cerf, the only included species in Caenoptychia (type species), based on adult morphology and molecular data. The results showed that Caenoptychia Le Cerf belongs to the Euptychia Hubner clade, and the genus is synonymized with Euptychia, new synonymy. Euptychia boulleti (Le Cerf) is a new combination. The male genitalia of E. boulleti showed at least one important synapomorphy with the other species of Euptychia, which is the presence of a posterior projection of the tegumen above the uncus. Molecular data reinforces the position of Caenoptychia within the genus Euptychia.

Cuban Calisto (Lepidoptera, Nymphalidae, Satyrinae), a review based on morphological and DNA data.

Abstract: The Cuban species of Calisto are reviewed based on the morphology of adult and immature stages, as well as DNA sequences of six genes (COI, EF1α, wingless, GAPDH, RpS5, CAD). A new species, Calisto occulta sp. n., is described from the northeastern Cuban mountains. Calisto smintheus Bates, 1935 and Calisto bruneri, Michener 1949 are revised and revalidated. A new status, the species level, is proposed for Calisto brochei, Torre 1973, Calisto muripetens, Bates 1939 and Calisto bradleyi, Munroe 1950. The immature stages of Calisto smintheus, Calisto brochei,and Calisto occulta are described for the first time, and those of Calisto herophile, Hubner 1823 are redescribed. Useful morphological characters for adults are the shape and conspicuousness of androconial patch, the number and relative size of white dots on underside of hindwing, the shape of aedeagus, the shape of digitiform projection of genitalia valve, the shape and relative size of tegumen and uncus, the relative size of female genitalia, the height of sterigmal ring dorsal crown of the latter, and the relative size of corpus bursae and ductus bursae. For the immature stages, the most important characters are the color pattern of head capsule, the number and width of longitudinal lines of body, in the larvae; and the color pattern and the absence or presence of dorsal ridges on the abdomen of pupae. The phylogenetic relationships between the Cuban Calisto species are quite robust and well-supported; however, conflict between mitochondrial and nuclear datasets was detected in Calisto brochei, Calisto muripetens and to a lesser degree in Calisto bradleyi.

A taxonomic study of the caddisfly Oxyethira falcata Morton, 1893 (Trichoptera: Hydroptilidae) using genital morphology and DNA barcoding

Abstract: Taxonomists have had problems with the hydroptilid caddisfly, Oxyethira falcate Morton, in the past. Four described taxa have been synonymized with O. falcate due to considerable intra-specific morphological variation of the male genitalia, and due to misinterpretation of some of the structures of these. In the present study specimens resembling morphologically O. boreella Svensson & Tjeder, one of the synonymized taxa, were compared with the true O. falcata using DNA barcoding and studying the male genitalia. Further, a molecular examination of all the Fennoscandian Oxyethira species were carried out, including the rare O. klingstedti Nybom, O. tamperensis Malicky, and O. ecornuta Morton. The results support keeping O. boreella as a synonym of O. falcata . In addition, the DNA analyses showed the presence of monophyletic groups for all of the studied Oxyethira species.