Temperature is thought to be a key factor influencing global species richness patterns. We investigate the link between temperature and diversification in the butterfly family Pieridae by combining next generation DNA sequences and published molecular data with fine-grained distribution data. We sampled nearly 600 pierid butterfly species to infer the most comprehensive molecular phylogeny of the family and curated a distribution dataset of more than 800,000 occurrences. We found strong evidence that species in environments with more stable daily temperatures or cooler maximum temperatures in the warm seasons have higher speciation rates. Furthermore, speciation and extinction rates decreased in tandem with global temperatures through geological time, resulting in a constant net diversification.
We propose a new genus of plexippine jumping spiders from the Western Ghats of India based on the new species Ghatippuspaschimagen. et sp. nov. While it bears a superficial resemblance to Pancorius in body form and Hyllus in membrane-bearing embolus, our UCE phylogenomic data-the first to resolve broad relationships within the Plexippina-as well as morphological features justify its status as a new genus. In addition to the molecular data and morphological descriptions, we provide photographs of living specimens of Ghatippuspaschimagen. et sp. nov. and information on their natural history.
A new species, Cigaritis conjuncta sp. nov. (Lepidoptera: Lycaenidae), is described based on five male and four female specimens from Honey Valley, Kodagu District, Karnataka, which is part of the Western Ghats biodiversity hotspot, India. Cigaritis conjuncta sp. nov. is distinguished from all other congeners in India and Sri Lanka based on the following combination of diagnostic characters in the male: (a) dorsal forewing outer half, costal margin and nearly upper half of the cell black without orange markings, the remaining wing shining deep blue, (b) dorsal hindwing tornus pale orange-red with two black spots, (c) ventral forewing bands at end of discal cell conjoined, and (e) ventral hindwing subbasal and discal bands composed of spots that are not separated but conjoined to form broad bands that have irregular outlines, leaving only a narrow background colour in between. Female is similar on the ventral side, but entirely dark brown above. Male and female genitalia are dissected and figured for two paratypes of each sex, and natural history notes on the species are provided.
Butterflies , Lepidoptera , Male , Female , Animals , India , Biodiversity
A new species, Piarosoma arunachalensis sp. nov., is described and illustrated based on external characters and the genitalia of two male specimens from Tale Wildlife Sanctuary, Lower Subansiri District, Arunachal Pradesh, India. The new species is distinguished from other Piarosoma by a combination of: a) presence of large hyaline patches on the wings, b) the absence of any white or yellow markings on the patagia and the abdominal segments, and c) differences in male genitalia.
Lepidoptera , Moths , Male , Animals , Genitalia , Genitalia, Male , India , Animals, Wild
The range of the butterfly genus Pseudocoladenia includes several biodiversity hotspots, such as the Himalayas, mountains of southwestern China, and Sundaland. However, the taxonomic status of some of its species/subspecies remain controversial, and no previous phylogenetic or biogeographic analyses have been conducted. Herein, we determined the systematic relationships and biogeographic history of this genus by reconstructing its phylogeny based on six genes and 69 specimens as representatives of all known species/subspecies. Two species delimitation methods (Bayes Poisson Tree Processes and Bayesian Phylogenetics and Phylogeography) were also employed to assess the status of each taxon. Based on these results and morphological evidence, we identified 12 species and three subspecies in the genus and subsequently classified these into three species groups: P. fatih, P. dea, and P. dan. Five taxa, P. sadakoe (Sonan and Mitono, 1936) stat. nov., P. celebica (Fruhstorfer, 1909) stat. nov., P. fulvescens (Elwes and Edwarda, 1897) stat. nov., P. eacus (Latreille, 1823) stat. nov., and P. fabia (Evans, 1949) stat. nov. were all recognized as independent species. Additionally, two taxa, P. eacus sumatrana (Fruhstorfer, 1909) comb. nov. and P. eacus dhyana (Fruhstorfer, 1909) comb. nov., were placed under P. eacus (Latreille, 1823) stat. nov. as subspecies. Another new species distributed in N. Yunnan, Pseudocoladenia yunnana Fan, Cao & Hou sp. nov., was discovered and described. Divergence time and ancestral range estimation indicated that the most recent common ancestor of Pseudocoladenia was distributed in the Himalayas-Hengduan Mountain region and Indochina and diverged approximately 14.00 Ma. Continuous and episodic dispersal, vicariance, and extinction were used to determine the current geographic distribution of the genus. The P. fatih group had a prominently disjunct distribution between the Himalaya-Hengduan Mountain and Taiwan. Meanwhile, the P. dan group was first derived in Indochina and subsequently dispersed into the southeastern Asian archipelagoes. This study provides a reference for the evolutionary route of transoceanic distributed species in Asia and elaborates on the causes of biodiversity.
Butterflies , Animals , Phylogeny , Butterflies/genetics , Bayes Theorem , China , Asia
Prey seldom rely on a single type of antipredator defence, often using multiple defences to avoid predation. In many cases, selection in different contexts may favour the evolution of multiple defences in a prey. However, a prey may use multiple defences to protect itself during a single predator encounter. Such "defence portfolios" that defend prey against a single instance of predation are distributed across and within successive stages of the predation sequence (encounter, detection, identification, approach (attack), subjugation and consumption). We contend that at present, our understanding of defence portfolio evolution is incomplete, and seen from the fragmentary perspective of specific sensory systems (e.g., visual) or specific types of defences (especially aposematism). In this review, we aim to build a comprehensive framework for conceptualizing the evolution of multiple prey defences, beginning with hypotheses for the evolution of multiple defences in general, and defence portfolios in particular. We then examine idealized models of resource trade-offs and functional interactions between traits, along with evidence supporting them. We find that defence portfolios are constrained by resource allocation to other aspects of life history, as well as functional incompatibilities between different defences. We also find that selection is likely to favour combinations of defences that have synergistic effects on predator behaviour and prey survival. Next, we examine specific aspects of prey ecology, genetics and development, and predator cognition that modify the predictions of current hypotheses or introduce competing hypotheses. We outline schema for gathering data on the distribution of prey defences across species and geography, determining how multiple defences are produced, and testing the proximate mechanisms by which multiple prey defences impact predator behaviour. Adopting these approaches will strengthen our understanding of multiple defensive strategies.
Ecology , Predatory Behavior , Animals , Phenotype
Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants.
Butterflies , Phylogeny , Animals , Biological Evolution , Butterflies/genetics
The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.
Butterflies , Animals , Phylogeny , Australia , Butterflies/genetics , Biological Evolution , Asia, Southeastern
Multitrait adaptive evolution is shaped by factors such as phylogenetic and functional constraints as well as the intensity and direction of selection. The tempo and mode of such multitrait evolution can differentially impact the assembly of biological communities. Batesian mimicry, in which undefended prey gain a fitness advantage by evolving a resemblance to aposematic models, involves adaptive evolution of multiple traits such as color patterns and flight morphology. To elucidate the evolutionary mechanisms of such multitrait adaptations, we evaluated the tempo and mode of adaptive convergence in flight morphology and color patterns in mimetic butterfly communities. We found that compared with Batesian mimics or nonmimetic sister species, models showed significantly faster rates of aposematic trait evolution, creating adaptive peaks for mimicry. At the community level, the degree of mimetic resemblance between mimics and models was positively correlated with the rate of character evolution, but independent of phylogenetic relatedness. Monomorphic mimics and female-limited mimics converged on the color patterns of models to a similar degree, showing that there were no constraints on mimetic trait evolution with respect to sex-specific selections. Convergence was driven by the greater lability of color patterns, which evolved at significantly faster rates than the phylogenetically conserved flight morphological traits, indicating that the two traits evolve under differential selection pressures and/or functional and genetic constraints. These community-wide patterns show that during the assembly of a community, the tempo of adaptive evolution is nonlinear, and specific to the underlying functional relationships and key traits that define the community.
Biological Mimicry , Butterflies , Animals , Female , Male , Biological Evolution , Biota , Butterflies/genetics , Phylogeny , Predatory Behavior
Long-term socioeconomic progress requires a healthy environment/ecosystem, but anthropogenic activities cause environmental degradation and biodiversity loss. Constant ecological monitoring is, therefore, necessary to assess the state of biodiversity and ecological health. However, baseline data are lacking even for ecologically sensitive regions such as the Western Ghats. We looked at the seasonality and polyphenism of butterflies of the central Western Ghats to obtain baseline population patterns on these charismatic taxa. We recorded 43118 individuals (175 species) using fortnightly time-constrained counts for two consecutive years and found the peak abundance (49% of the total individuals) in the post-monsoon period (October to January). Seasonal abundance was correlated with the overall increase in species richness. Habitat differences were stronger than seasonality as samples clustered based on sites. Several species also displayed polyphenism with distinct distributions of wet and dry season forms. Seasonal equitability and indicator species analysis showed distinct inter-species differences in seasonality patterns. This work provides key baseline data on the seasonal dynamics of butterflies of the Western Ghats in the context of climate change and conservation. It will help monitor this ecologically sensitive region using butterflies.
Butterflies , Ecosystem , Humans , Animals , Seasons , Biodiversity , India
We describe a new species of dragonfly, Burmagomphus chaukulensis sp. nov., based on four males and two females from Chaukul, Sindhudurg, Maharashtra located in the Western Ghats biodiversity hotspot of India. The newly described species is diagnosed from its congeners by a combination of following characters: the shape of male caudal appendages (prominent lateral spines on cerci) and the lateral thoracic markings (stripe on mesepisternum reduced and pointed anteriorly). Additionally, B. chaukulensis is unique in terms of the shape of hamuli, strongly angulated with spines at both sides. An updated identification key to Indian Burmagomphus spp. is also provided.
Dipterocarpaceae , Odonata , Animals , Body Size , Female , India , Male
Contamination of a genetic sample with DNA from one or more nontarget species is a continuing concern of molecular phylogenetic studies, both Sanger sequencing studies and next-generation sequencing studies. We developed an automated pipeline for identifying and excluding likely cross-contaminated loci based on the detection of bimodal distributions of patristic distances across gene trees. When contamination occurs between samples within a data set, a comparison between a contaminated sample and its contaminant taxon will yield bimodal distributions with one peak close to zero patristic distance. This new method does not rely on a priori knowledge of taxon relatedness nor does it determine the causes(s) of the contamination. Exclusion of putatively contaminated loci from a data set generated for the insect family Cicadidae showed that these sequences were affecting some topological patterns and branch supports, although the effects were sometimes subtle, with some contamination-influenced relationships exhibiting strong bootstrap support. Long tip branches and outlier values for one anchored phylogenomic pipeline statistic (AvgNHomologs) were correlated with the presence of contamination. While the anchored hybrid enrichment markers used here, which target hemipteroid taxa, proved effective in resolving deep and shallow level Cicadidae relationships in aggregate, individual markers contained inadequate phylogenetic signal, in part probably due to short length. The cleaned data set, consisting of 429 loci, from 90 genera representing 44 of 56 current Cicadidae tribes, supported three of the four sampled Cicadidae subfamilies in concatenated-matrix maximum likelihood (ML) and multispecies coalescent-based species tree analyses, with the fourth subfamily weakly supported in the ML trees. No well-supported patterns from previous family-level Sanger sequencing studies of Cicadidae phylogeny were contradicted. One taxon (Aragualna plenalinea) did not fall with its current subfamily in the genetic tree, and this genus and its tribe Aragualnini is reclassified to Tibicininae following morphological re-examination. Only subtle differences were observed in trees after the removal of loci for which divergent base frequencies were detected. Greater success may be achieved by increased taxon sampling and developing a probe set targeting a more recent common ancestor and longer loci. Searches for contamination are an essential step in phylogenomic analyses of all kinds and our pipeline is an effective solution. [Auchenorrhyncha; base-composition bias; Cicadidae; Cicadoidea; Hemiptera; phylogenetic conflict.].
Hemiptera , Animals , Hemiptera/genetics , High-Throughput Nucleotide Sequencing , Insecta/genetics , Phylogeny
The collection of caterpillar fungus accounts for 50-70% of the household income of thousands of Himalayan communities and has an estimated market value of $5-11 billion across Asia. However, Himalayan collectors are at multiple economic disadvantages compared with collectors on the Tibetan Plateau because their product is not legally recognized. Using a customized hybrid-enrichment probe set and market-grade caterpillar fungus (with samples up to 30 years old) from 94 production zones across Asia, we uncovered clear geography-based signatures of historical dispersal and significant isolation-by-distance among caterpillar fungus hosts. This high-throughput approach can readily distinguish samples from major production zones with definitive geographical resolution, especially for samples from the Himalayan region that form monophyletic clades in our analysis. Based on these results, we propose a two-step procedure to help local communities authenticate their produce and improve this multi-national trade-route without creating opportunities for illegal exports and other forms of economic exploitation. We argue that policymakers and conservation practitioners must encourage the fair trade of caterpillar fungus in addition to sustainable harvesting to support a trans-boundary conservation effort that is much needed for this natural commodity in the Himalayan region.
Fungi , Asia , Geography
Three new species of the Asian genus Mata Distant, 1906 (Hemiptera: Cicadidae) viz. Mata lenonia sp.nov.; Mata ruffordii sp.nov. and Mata meghalayana sp.nov. are described from Indian state of Meghalaya. Keys and taxonomic descriptions of these species are provided with detailed accounts of their natural history and acoustics.
Hemiptera , Acoustics , Animals , India , Reptiles
Molecular phylogenies have yielded strong support for many parts of the amphibian Tree of Life, but poor support for the resolution of deeper nodes, including relationships among families and orders. To clarify these relationships, we provide a phylogenomic perspective on amphibian relationships by developing a taxon-specific Anchored Hybrid Enrichment protocol targeting hundreds of conserved exons which are effective across the class. After obtaining data from 220 loci for 286 species (representing 94% of the families and 44% of the genera), we estimate a phylogeny for extant amphibians and identify gene tree-species tree conflict across the deepest branches of the amphibian phylogeny. We perform locus-by-locus genealogical interrogation of alternative topological hypotheses for amphibian monophyly, focusing on interordinal relationships. We find that phylogenetic signal deep in the amphibian phylogeny varies greatly across loci in a manner that is consistent with incomplete lineage sorting in the ancestral lineage of extant amphibians. Our results overwhelmingly support amphibian monophyly and a sister relationship between frogs and salamanders, consistent with the Batrachia hypothesis. Species tree analyses converge on a small set of topological hypotheses for the relationships among extant amphibian families. These results clarify several contentious portions of the amphibian Tree of Life, which in conjunction with a set of vetted fossil calibrations, support a surprisingly younger timescale for crown and ordinal amphibian diversification than previously reported. More broadly, our study provides insight into the sources, magnitudes, and heterogeneity of support across loci in phylogenomic data sets.[AIC; Amphibia; Batrachia; Phylogeny; gene tree-species tree discordance; genomics; information theory.].
Fossils , Genomics , Animals , Anura , Humans , Phylogeny
A new species, Zographetus dzonguensis sp. nov., is described based on three male specimens from Upper Dzongu, North Sikkim District, Sikkim, India. The new species is closely similar to Z. pangi from Guangdong and Z. hainanensis from Hainan, China, from which it is distinguished based on the following combination of external characters: (a) forewing with the white spot at the base of space M3 being sharply pointed at the inner edge and conspicuously more elongated, and (b) on underside of hindwing, all chocolate-brown spots being smaller, in Z. dzonguensis sp. nov. compared with both Z. pangi and Z. hainanensis. In Z. dzonguensis sp. nov., the male genitalia may be distinguished as follows: (c) tegumen and uncus are almost equal in length, (d) the dorsal outline of tegumen is relatively flat, and (e) saccus is distinctly bent upward as angle between vinculum and saccus is more acute than in Z. pangi and Z. hainanensis.
Lepidoptera , Moths , Animals , Genitalia, Male , India , Male , Sikkim
Adaptive phenotypes often arise by rewiring existing developmental networks. Co-option of transcription factors in novel contexts has facilitated the evolution of ecologically important adaptations. doublesex (dsx) governs fundamental sex differentiation during embryonic stages and has been co-opted to regulate diverse secondary sexual dimorphisms during pupal development of holometabolous insects. In Papilio polytes, dsx regulates female-limited mimetic polymorphism, resulting in mimetic and non-mimetic forms. To understand how a critical gene such as dsx regulates novel wing patterns while maintaining its basic function in sex differentiation, we traced its expression through metamorphosis in P. polytes using developmental transcriptome data. We found three key dsx expression peaks: (i) eggs in pre- and post-ovisposition stages; (ii) developing wing discs and body in final larval instar; and (iii) 3-day pupae. We identified potential dsx targets using co-expression and differential expression analysis, and found distinct, non-overlapping sets of genes-containing putative dsx-binding sites-in developing wings versus abdominal tissue and in mimetic versus non-mimetic individuals. This suggests that dsx regulates distinct downstream targets in different tissues and wing colour morphs and has perhaps acquired new, previously unknown targets, for regulating mimetic polymorphism. Additionally, we observed that the three female isoforms of dsx were differentially expressed across stages (from eggs to adults) and tissues and differed in their protein structure. This may promote differential protein-protein interactions for each isoform and facilitate sub-functionalization of dsx activity across its isoforms. Our findings suggest that dsx employs tissue-specific downstream effectors and partitions its functions across multiple isoforms to regulate primary and secondary sexual dimorphism through insect development.
Three new species of Protosticta Selys, 1885 (Odonata: Zygoptera: Platystictidae) from the Western Ghats biodiversity hotspot in India: P. cyanofemora sp. nov. (wet evergreen forests in Shendurney Wildlife Sanctuary, Kollam, Kerala and Kalakkad Mundanthurai Tiger Reserve, Tirunelveli, Tamil Nadu), P. myristicaensis sp. nov. (Myristica swamp at Kathalekan, Shivamogga, Karnataka) and P. sholai sp. nov. (montane sholas of Upper Manalar, Meghamalai Wildlife Sanctuary, Theni, Tamil Nadu) are described and illustrated. We compare these three new species with other Protosticta spp. from the Western Ghats based on new material and provide comprehensive differential diagnoses with determination key for males of all species occurring in the Western Ghats. The taxonomic validity of P. mortoni Fraser, 1922 is established with fresh specimens from Hassan, Karnataka, and rediscovery of P. rufostigma Kimmins, 1958 is reported from Kanyakumari Wildlife Sanctuary, Tamil Nadu.
Odonata , Animals , Animals, Wild , Biodiversity , Forests , India , Male
A new species, Metallolophia taleensis, sp. nov., is described and illustrated from Tale Wildlife Sanctuary, Lower Subansiri District, Arunachal Pradesh, India, based on male specimens. A second species, Metallolophia opalina (Warren, 1893), is recorded from India after more than a century, extending its range eastwards into Arunachal Pradesh, India.
Moths , Animals , India , Male
Movement may fundamentally alter morphology and reproductive states in insects. In long-distance migrants, reproductive diapause is associated with trade-offs between diverse life-history traits such as flight morphology and lifespan. However, many non-diapausing insects engage in shorter resource-driven dispersals. How diapause and other reproductive states alter flight morphology in migrating versus dispersing insects is poorly understood. To find out, we compared flight morphology in different reproductive states of multiple butterfly species. We found that dispersers consisted of ovulating females with higher egg loads compared with non-dispersing females. This trend was in stark contrast with that of migrating female butterflies in reproductive diapause, which made substantially higher investment in flight tissue compared with reproductively active, non-migrating females. Thus, long-distance migration and shorter resource-driven dispersals had contrasting effects on flight morphology and egg loads. By contrast, male flight morphology was not affected by dispersal, migration or associated reproductive states. Thus, dispersal and migration affected resource allocation in flight and reproductive tissue in a sex-specific manner across relatively mobile versus non-dispersing individuals of different species. These findings suggest that dispersals between fragmented habitats may put extra stress on egg-carrying females by increasing their flight burdens.
Butterflies , Animals , Ecosystem , Female , Flight, Animal , Humans , Insecta , Longevity , Male , Reproduction
