A new Heraclides swallowtail (Lepidoptera,Papilionidae) from North America is recognized by the pattern on its neck

Heraclides rumiko Shiraiwa & Grishin, sp. n. is described from southwestern United States, Mexico, and Central America (type locality: USA, Texas, Duval County). It is closely allied to Heraclides cresphontes (Cramer, 1777) and the two species are sympatric in central Texas. The new species is diagnosed by male genitalia and exhibits a nearly 3% difference from Heraclides cresphontes in the COI DNA barcode sequence of mitochondrial DNA. The two Heraclides species can usually be told apart by the shape and size of yellow spots on the neck, by the wing shape, and the details of wing patterns. “Western Giant Swallowtail” is proposed as the English name for Heraclides rumiko. To stabilize nomenclature, neotype for Papilio cresphontes Cramer, 1777, an eastern United States species, is designated from Brooklyn, New York, USA; and lectotype for Papilio thoas Linnaeus, 1771 is designated from Suriname. We sequenced DNA barcodes and ID tags of nearly 400 Papilionini specimens completing coverage of all Heraclides species. Comparative analyses of DNA barcodes, genitalia, and facies suggest that Heraclides oviedo (Gundlach, 1866), reinstated status, is a species-level taxon rather than a subspecies of Heraclides thoas (Linnaeus, 1771); and Heraclides pallas (G. Gray, [1853]), reinstated status, with its subspecies Heraclides Papilio bajaensis (J. Brown & Faulkner, 1992), comb. n., and Heraclides anchicayaensis Constantino, Le Crom & Salazar, 2002, stat. n., are not conspecific with Heraclides astyalus (Godart, 1819).     

Phylogeny and biogeography of Sassafras (Lauraceae) disjunct between eastern Asia and eastern North America

Sassafras (Lauraceae) consists of three species disjunct between eastern Asia (S. tzumu and S. randaiense) and eastern North America (S. albidum). Phylogenetic analysis based on sequences of nuclear ribosomal ITS and three chloroplast non-coding regions (rpl16, trnL-F, and psbA-trnH) showed that Sassafras is monophyletic and that the eastern North American S. albidum is sister to the clade of its two eastern Asian counterparts. Their intercontinental divergence was estimated to be 13.80 ± 2.29−16.69 ± 2.52 million years ago (mya) using the penalized likelihood method with the ITS and three chloroplast markers. Biogeographic analyses combined with fossil evidence suggest that Sassafras has a relict distribution in the Northern Hemisphere without a Gondwanan link. The divergence time of the two eastern Asian species (the continental Chinese Sassafras tzumu and S. randaiense endemic to Taiwan) is estimated to be 0.61 ± 0.75−2.23 ± 0.76 mya. Sassafras randaiense from Taiwan was most likely derived from an ancestor from continental China.     

Intercontinental genetic divergence of Castanea species in eastern Asia and eastern North America

Castanea is one of the many plant genera with a disjunct distribution pattern between eastern Asia and eastern North America. Five species from three sections of the genus were investigated to examine genetic divergence between eastern Asian and eastern North American species. A total of 62 native populations were sampled for allelic variation at isozyme loci. The Chinese chestnut C. mollissima had the highest genetic variability, while the American C. dentata had the lowest genetic variability. The highest intracontinental genetic identities were observed between the Allegheny and Ozark chinkapins (0.931) and between C. mollissima and C. seguinii (0.870), while lower identities were detected between the American C. pumila and C. dentata (0.720-0.729). In intercontinental comparisons, genetic identities of 0.505, 0.495 and 0.507 were observed between the American chestnut and the Chinese C. mollissima, C. seguinii and C. henryi, respectively, whereas the Ozark chinkapin C. pumila var. ozarkensis had lower identities of 0.469, and 0.435 with C. mollissima and C. seguinii, respectively, but a slightly higher identity of 0.520 with C. henryi, the Chinese chinkapin. Divergence times were estimated at 10-13 million years before present between C. dentata and C. mollissima, and C. pumila var. ozarkensis and C. henryi.     

Intercontinental biogeography of subfamily Orontioideae (Symplocarpus, Lysichiton, and Orontium) of Araceae in Eastern Asia and North America

Symplocarpus, Lysichiton, and Orontium (Orontioideae) are three of the few north temperate genera of the primarily tropical Araceae. Symplocarpus is disjunctly distributed in eastern Asia (3 spp.) and eastern North America (1 sp.); Lysichiton has an intercontinental discontinuous distribution in eastern Asia (1 sp.) and northwestern North America (1 sp.); and the monotypic Orontium is restricted to eastern North America. Phylogenetic analysis of the trnL-F and ndhF sequences supports (1) the monophyly of both Symplocarpus and Lysichiton, (2) the sister-group relationship of Symplocarpus and Lysichiton, and (3) the clade of Orontium, Symplocarpus, and Lysichiton. Although Symplocarpus shows a much wider disjunction than Lysichiton, the estimated divergence time of the former [4.49+/-1.69 or 6.88+/-4.18 million years ago (mya)] was similar to that of the latter (4.02+/-1.60 or 7.18+/-4.33 mya) based on the penalized likelihood and the Bayesian dating methods, respectively. Eastern Asia was suggested to be the ancestral area of the Symplocarpus-Lysichiton clade based on the dispersal-vicariance analysis. Our biogeographic results support independent migrations of Symplocarpus and Lysichiton across the Bering land bridge in the late Tertiary (Pliocene/late Miocene). Fossil evidence suggests Orontioideae dated back to the late Cretaceous in the temperate Northern Hemisphere (72 mya). The relative rate test shows similar substitution rates of the trnL-F sequences between the proto and the true aroids, although the latter has substantially higher species diversity. The proto Araceae perhaps suffered from a higher rate of extinction in the temperate zone associated with periods of climatic cooling in the Tertiary.     

A partitioned likelihood analysis of swallowtail butterfly phylogeny (Lepidoptera:Papilionidae)

Although it is widely agreed that data from multiple sources are necessary to confidently resolve phylogenetic relationships, procedures for accommodating and incorporating heterogeneity in such data remain underdeveloped. We explored the use of partitioned, model-based analyses of heterogeneous molecular data in the context of a phylogenetic study of swallowtail butterflies (Lepidoptera: Papilionidae). Despite substantial basic and applied study, phylogenetic relationships among the major lineages of this prominent group remain contentious. We sequenced 3.3 kb of mitochondrial and nuclear DNA (2.3 kb of cytochrome oxidase I and II and 1.0 kb of elongation factor-1 alpha, respectively) from 22 swallowtails, including representatives of Baroniinae, Parnassiinae, and Papilioninae, and from several moth and butterfly outgroups. Using parsimony, we encountered considerable difficulty in resolving the deepest splits among these taxa. We therefore chose two outgroups with undisputed relationships to each other and to Papilionidae and undertook detailed likelihood analyses of alternative topologies. Following from previous studies that have demonstrated substantial heterogeneity in the evolutionary dynamics among process partitions of these genes, we estimated evolutionary parameters separately for gene-based and codon-based partitions. These values were then used as the basis for examining the likelihoods of possible resolutions and rootings under several partitioned and unpartitioned likelihood models. Partitioned models gave markedly better fits to the data than did unpartitioned models and supported different topologies. However, the most likely topology varied from model to model. The most likely ingroup topology under the best-fitting, six-partition GTR + gamma model favors a paraphyletic Parnassiinae. However, when examining the likelihoods of alternative rootings of this tree relative to rootings of the classical hypothesis, two rootings of the latter emerge as most likely. Of these two, the most likely rooting is within the Papilioninae, although a rooting between Baronia and the remaining Papilionidae is only nonsignificantly less likely.     

A new species of Herpetogramma (Lepidoptera, Crambidae, Spilomelinae) from eastern North America

Herpetogramma sphingealissp. n., a new species of Crambidae (Lepidoptera), is described from Québec, Canada. The species is included in the genus Herpetogramma Led., 1863, a genus in the subfamily Spilomelinae. Adults and genitalia of this species are described and illustrated, as well as those of Herpetogramma aeglealis (Walker, 1859) and Herpetogramma thestealis (Walker, 1859), and adults of the semi-melanic form of Herpetogramma aeglealis are illustrated.     

Molecular Phylogeny and Historical Biogeography of the Neotropical Swarm-Founding Social Wasp Genus Synoeca (Hymenoptera: Vespidae)

The Neotropical Region harbors high biodiversity and many studies on mammals, reptiles, amphibians and avifauna have investigated the causes for this pattern. However, there is a paucity of such studies that focus on Neotropical insect groups. Synoeca de Saussure, 1852 is a Neotropical swarm-founding social wasp genus with five described species that is broadly and conspicuously distributed throughout the Neotropics. Here, we infer the phylogenetic relationships, diversification times, and historical biogeography of Synoeca species. We also investigate samples of the disjoint populations of S. septentrionalis that occur in both northwestern parts of South America through Central American and the Brazilian Atlantic rainforests. Our results showed that the interspecific relationships for Synoeca could be described as follows: (S. chalibea + S. virginea) + (S. cyanea + (S. septentrionalis/S. surinama)). Notably, samples of S. septentrionalis and S. surinama collected in the Atlantic Forest were interrelated and may be the result of incomplete lineage sorting and/or mitochondrial introgression among them. Our Bayesian divergence dating analysis revealed recent Plio-Pleistocene diversification in Synoeca. Moreover, our biogeographical analysis suggested an Amazonian origin of Synoeca, with three main dispersal events subsequently occurring during the Plio-Pleistocene.     

Intercontinental union of Argentine ants: behavioral relationships among introduced populations in Europe, North America, and Asia

Many invasive ants, including the Argentine ant Linepithema humile, form expansive supercolonies, within which intraspecific aggression is absent. The behavioral relationships among introduced Argentine ant populations at within-country or within-continent scales have been studied previously, but the behavioral relationships among intercontinental populations have not been examined. The present study investigated the levels of aggression among intercontinental Argentine ant populations by transporting live ants from Europe and California to Japan and conducting aggression tests against Japanese populations. Workers from the dominant supercolonies of Europe and California did not show aggressive behavior toward workers from the dominant supercolony of Japan, whereas they fought vigorously against workers from minor supercolonies. The three massive supercolonies, together with Argentine ants from Macaronesia, may be the largest non-aggressive unit formed by a social insect species in which intraspecific aggression exists. Absence or low levels of aggression at transcontinental scale, which may have derived from low genetic variation, may help introduced Argentine ants maintain expansive supercolonies. The lack of aggression implies possible frequent exchanges of individuals among the intercontinental populations mediated by human activities.     

A comparison of the taxonomic richness of temperate plants in East Asia and North America

The taxonomic richness of seed plants at different taxonomic levels was compared between temperate East Asia and North America at both continental and semi-continental scales. In each comparison, land area and latitude range were adjusted to a comparable level between the two continental regions. East Asia is significantly more diverse than North America. In general, differences in taxonomic diversity arise at and below the genus level. At the continental scale, East Asia has 1.3 and 1.5 times as many genera and species, respectively, as North America. The northern part of East Asia has 1.1 times as many species as the northern part of North America. At the genus level, the northern part of East Asia is less diverse than the northern part of North America by a factor of 0.94. This pattern indicates that the diversity bias between the two continental regions results from the flora of southern East Asia. The diversity differences between East Asia and North America are not homogenously distributed across different plant groups. At the species level, East Asia had significantly more species than expected in magnoliids, alismatids, Liliidae, ranunculids, and rosids and had significantly less species in the Commelinidae, Caryophyllidae, and euasterids than North America.     

Free Tropospheric Transport of Microorganisms from Asia to North America

Microorganisms are abundant in the troposphere and can be transported vast distances on prevailing winds. This study measures the abundance and diversity of airborne bacteria and fungi sampled at the Mt. Bachelor Observatory (located 2.7?km above sea level in North America) where incoming free tropospheric air routinely arrives from distant sources across the Pacific Ocean, including Asia. Overall deoxyribonucleic acid (DNA) concentrations for microorganisms in the free troposphere, derived from quantitative polymerase chain reaction assays, averaged 4.94?×?10^?5?ng DNA m^?3 for bacteria and 4.77?×?10^?3?ng DNA m^?3 for fungi. Aerosols occasionally corresponded with microbial abundance, most often in the springtime. Viable cells were recovered from 27.4?% of bacterial and 47.6?% of fungal samples (N?=?124), with 49 different species identified by ribosomal DNA gene sequencing. The number of microbial isolates rose significantly above baseline values on 22–23 April 2011 and 13–15 May 2011. Both events were analyzed in detail, revealing distinct free tropospheric chemistries (e.g., low water vapor, high aerosols, carbon monoxide, and ozone) useful for ruling out boundary layer contamination. Kinematic back trajectory modeling suggested air from these events probably originated near China or Japan. Even after traveling for 10?days across the Pacific Ocean in the free troposphere, diverse and viable microbial populations, including presumptive plant pathogens Alternaria infectoria and Chaetomium globosum, were detected in Asian air samples. Establishing a connection between the intercontinental transport of microorganisms and specific diseases in North America will require follow-up investigations on both sides of the Pacific Ocean.     

A chloroplast phylogeny of Arisaema (Araceae) illustrates Tertiary floristic links between Asia, North America, and East Africa

The evolution of Arisaema is reconstructed, based on combined sequences (2048 aligned bases) from the chloroplast trnL intron, trnL-trnF spacer, and rpl20-rps12 spacer obtained for species from all 11 sections, including sectional type species and geographically disjunct East African and North American/Mexican species. Analyses were rooted with a representative sample of the closest outgroups, Pinellia and Typhonium, to rigorously test the monophyly of Arisaema. Sections in Arisaema are mostly based on leaf, stem, and inflorescence characters and, with one exception, are not rejected by the molecular data; however, statistical support for sectional relationships in the genus remains poor. Section Tortuosa, which includes eastern North American A. dracontium and Mexican A. macrospathum, is demonstrably polyphyletic. The third New World species, A. triphyllum, also occurs in eastern North America and groups with a different Asian clade than do A. dracontium/A. macrospathum. The genus thus appears to have entered North America twice. Fossil infructescences similar to those of A. triphyllum are known from approximately 18 million-year-old deposits in Washington State and can serve to calibrate a molecular clock. Constraining the age of A. triphyllum to 18 million years (my) and applying either a semiparametric or an ultrametric clock model to the combined data yields an age of approximately 31-49 my for the divergence of A. dracontium/A. macrospathum from their Asian relatives and of 19-32 my for the divergence between African A. schimperianum and a Tibetan/Nepalese relative. The genus thus provides an example of the Oligocene/Miocene floristic links between East Africa, Arabia, the Himalayan region, China, and North America. The phylogeny also suggests secondary loss of the environmental sex determination strategy that characterizes all arisaemas except for two subspecies of A. flavum, which have consistently bisexual spathes. These subspecies are tetraploid and capable of selfing, while a third subspecies of A. flavum is diploid and retains the sex-changing strategy. In the molecular trees, the sex-changing subspecies is sister to the two non-sex-changing ones, and the entire species is not basal in the genus.     

Phylogeny and Biogeography of Gibbons: A Dispersal-Vicariance Analysis

Phylogenetic relationships within Hylobatidae are controversial. Numerous studies based on molecular, morphological and behavioral characteristics have provided conflicting results. I reanalyzed published cytochrome b gene sequence data to provide a new estimate of gibbon phylogeny. My results indicate that Nomascus, Symphalangus and Hoolock are successively more closely related to Hylobates. Molecular clock analyses provide estimates of divergence times within Hylobatidae, indicating that the radiation dates to ca. 10.5 million years ago. Scientists have little understanding of the biogeographic history of gibbons, largely because of a sparse fossil record. I combined the estimate of gibbon phylogeny with distribution data in a dispersal-vicariance analysis and present a new scenario for the pattern and timing of gibbon radiation.     

Butterflies and skippers (Lepidoptera,Diurna) from the Republic of Mordovia

According to the data obtained, the Mordovian fauna of diurnal lepidopterans comprises 75 species, including Hesperiidae (7 species), Papilionidae (5), Pieridae (10), Satyridae (10), Nymphalidae (27), and Lycaenidae (16). Based on the quantitative assessments, the species were characterized as rare (1–5 ind. in the collection), scarce (6–30), common (31–170), and abundant (171–940). Species with temperate (52 species or 69.3%) and subboreal (12 species or 16.0%) ranges predominated in the latitudinal distribution pattern. The total and background species diversity of the Diurna was the highest in the forests (63 species) and meadows (52), and the lowest on the steppe slopes (6). Among the species revealed, 14 were rare: Iphiclides podalirius, Zerynthia polyxena, Parnassius apollo, Driopa mnemosyne, Colias erate, Coenonympha hero, Neptis sappho, Apatura iris, Limenitis populi, Nymphalis antiopa, Fixsenia pruni, Lycaena helle, Cupido minimus, and Maculinea arion.     

Morphological Characters Are Compatible with Mitogenomic Data in Resolving the Phylogeny of Nymphalid Butterflies (Lepidoptera: Papilionoidea: Nymphalidae)

Nymphalidae is the largest family of butterflies with their phylogenetic relationships not adequately approached to date. The mitochondrial genomes (mitogenomes) of 11 new nymphalid species were reported and a comparative mitogenomic analysis was conducted together with other 22 available nymphalid mitogenomes. A phylogenetic analysis of the 33 species from all 13 currently recognized nymphalid subfamilies was done based on the mitogenomic data set with three Lycaenidae species as the outgroups. The mitogenome comparison showed that the eleven new mitogenomes were similar with those of other butterflies in gene content and order. The reconstructed phylogenetic trees reveal that the nymphalids are made up of five major clades (the nymphaline, heliconiine, satyrine, danaine and libytheine clades), with sister relationship between subfamilies Cyrestinae and Biblidinae, and most likely between subfamilies Morphinae and Satyrinae. This whole mitogenome-based phylogeny is generally congruent with those of former studies based on nuclear-gene and mitogenomic analyses, but differs considerably from the result of morphological cladistic analysis, such as the basal position of Libytheinae in morpho-phylogeny is not confirmed in molecular studies. However, we found that the mitogenomic phylogeny established herein is compatible with selected morphological characters (including developmental and adult morpho-characters).     

Corn borer (Lepidoptera: Noctuidae and Crambidae) resistance of main races of maize from North America

Resistance to corn borers, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae) and Sesamia nonagrioides (Lefebvre) (Lepidoptera: Noctuidae) in maize, Zea mays L., populations is partial, and more resistant populations are needed. The objective of this research was to compare resistance to corn borers of the main maize races from North America. Twenty open-pollinated maize populations belonging to the races Southern Dent, Corn Belt Dent, and Northern Flint, and three check populations, were evaluated under artificial infestation with S. nonagrioides and O. nubilalis. None of the populations had complete resistance. Northern Flint had the lowest yield under corn borer infestation, whereas Southern Dent had the highest yield but also the largest damage. Corn Belt Dent had a shorter growing cycle and similar yield of infected plants than Southern Dent. The checks had intermediate yield and resistance and were not significantly different from Corn Belt Dent for any trait. The Southern Dent populations Tennessee Red Cob and White Dent (PI221885 and PI311232) could be used as sources of tolerance to corn borers, although they are not expected to provide great gains compared with the levels of tolerance already present in some Corn Belt Dent and European Flint populations and would require adaptation to short growing cycle. The Corn Belt Dent synthetic BS17 had the highest yield and general agronomic performance under corn borer infestation, along with Rustler and Silver King, and the European Flint composite EPS13.     

Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera, Papilionidae) based on morphology and seven genes

We tested the taxonomic utility of morphology and seven mitochondrial or nuclear genes in a phylogenetic reconstruction of swallowtail butterflies in the subfamily Parnassiinae. Our data included 236 morphological characters and DNA sequences for seven genes that are commonly used to infer lepidopteran relationships (COI+COII, ND5, ND1, 16S, EF-1alpha, and wg; total 5775 bp). Nuclear genes performed best for inferring phylogenies, particularly at higher taxonomic levels, while there was substantial variation in performance among mitochondrial genes. Multiple analyses of molecular data (MP, ML and Bayesian) consistently produced a tree topology different from that obtained by morphology alone. Based on molecular evidence, sister-group relationships were confirmed between the genera Hypermnestra and Parnassius, as well as between Archon and Luehdorfia, while the monophyly of the subfamily was weakly supported. We recognize three tribes within Parnassiinae, with Archon and Luehdorfia forming the tribe Luehdorfiini Tutt, 1896 [stat. rev.]. Three fossil taxa were incorporated into a molecular clock analysis with biogeographic time constraints. Based on dispersal-vicariance (DIVA) analysis, the most recent common ancestor of Parnassiinae occurred in the Iranian Plateau and Central Asia to China. Early diversification of Parnassiinae took place at the same time that India collided into Eurasia, 65-42 million years ago.     

Butterflies (Lepidoptera,Diurna) in temperate forests of the southern Far East of Russia

The species groups of butterflies in temperate forests and adjoining forest-steppes of the Russian Far East are analyzed. The contribution of species with different distribution types into the total species composition is considered. A classification analysis of the species lists of sixteen local faunas from Amur and Sakhalin Provinces, Khabarovsk and Primorskii Territories, and the Jewish Autonomous Region was performed. A uniform faunal core is shown to be present in the forests (including deciduous ones) of the Russian Far East temperate zone.     

Field bioassays show heterospecific mating preference asymmetry between hybridizing North American Papilio butterfly species (Lepidoptera: Papilionidae)

Few studies of interspecific mating preferences of naturally hybridizing species have been done in the field. Yet this is the only way potentially critical habitat-specific factors can be included in mating behavior evaluations. We conducted mate selection (male preference) studies using tethered pairs of heterospecific, size-matched virgin yellow females of Papilio glaucus and P. canadensis with free-flying male P. glaucus populations in south-central Florida, and with free-flying male P. canadensis populations in northern Michigan. Florida males clearly preferred the conspecific P. glaucus females of the pairs, as indicated by the touches/attempted matings and by the actual copulations observed. In 1997, 66.7% of the total attempts (n=168) and 94.2% of all copulations (n=69) were with the conspecific. In 1998 the same pattern of conspecific male preference was observed, with 67.3% of the touches/attempts (n=49) and 100% of the copulations (n=30) being with the P. glaucus females. In contrast, the natural populations of P. canadensis males did not show the expected conspecific preference. In fact, with wild P. canadensis males in 1997 a very strong preference was observed for the heterospecific P. glaucus females of the pairs, accounting for 75.8% of all touches/attempted matings (n=483) and 81.7% of all copulations (n=476). The strong asymmetry in interspecific mating preferences among these hybridizing Papilio species may reflect an ancestral trait in P. glaucus females that elicits strong preference in P. canadensis males, due perhaps to previously possessed male sensory bias or "good genes" indicators in the females. However, reinforcement of reproductive isolation in or near the hybrid zone of species overlap (sympatry) cannot be ruled out. Electronic Publication     

Male-like females, mimicry and transvestism in butterflies (Lepidoptera: Papilionidae)

Abstract. When a butterfly species has a polymorphic female, with one of the forms closely resembling the male, it is customary to suppose that this form is ancestral, and that the ‘odd’ forms have arisen later. R. I. Vane-Wright, on the other hand, has suggested that in some species the male-like form may be a ‘transvestite’ female, the ancestral form of the female having been strikingly unlike the male. As later-derived forms are usually, but not always, genetically dominant to ancestral forms, we can make some choice between these hypotheses by discovering the dominance relations of the male-like and the ‘odd’ forms of the female. In the mimetic Papilio aegeus the male-like form is shown to be recessive to the ‘odd’ (mimetic) form, as has essentially been the case in all other butterflies so far investigated. Papilio phorcas is now shown to be the exception: the ‘odd’ (non-mimetic) form is recessive to the male-like form. We conclude that usually the male-like form is ancestral, but that P.phorcas may be an authentic example of ‘transvestism’, or the ‘transfer’ of male epigamic colour to the female of the species. The yellow, male-like pattern of the mimetic Papilio dardanus may be dominant or recessive to the mimetic forms according to the genetic background: largely recessive in Madagascar, and southern and western Africa, dominant to most forms in Ethiopia, and probably dominant to one mimetic form but recessive to the others in Kenya. All female dardanus patterns, both mimetic and yellow, are strongly dominant to both female phorcas patterns in P. dardanus × P. phorcas hybrids (P. ‘nandina’). The simplest explanation of this situation is that the male-like pattern of dardanus is ancestral, and that dominance has become locally reversed in Ethiopia. The dominance relations, and the sex- or autosomal-linkage of two forms can be determined without pedigree-breeding, simply by observing a few offspring each from a large number of wild-caught females.