Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences

Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index summarizing incongruence among data partitions, the Partition Congruence Proportion (PCP), is introduced. The significance of the inferred pattern of phylogenetic relationships for comparative chemical ecology of milkweed butterflies is briefly discussed.     

青藏高原高山植物的形态和解剖结构及其对环境的适应性研究进展

高山植物是一类生长于树线以上至雪线的山地植物。揭示高山植物适应环境的形态和结构特征及其内在机制,对研究全球气候变化下,植物对环境的响应和适应具有重要的理论意义。然而,国内在高山植物功能生态学的研究上相对薄弱,已有研究主要集中在对青藏高原高山植物的报道上。结合国外高山植物的相关研究报道,从形态和解剖结构两个方面对青藏高原高山植物的研究进展进行了综述,重点阐述了高山植物的形态、解剖结构及其与环境的适应性关系。植株矮小（有的呈垫状）、叶片小而厚、具有通气组织、栅栏组织多层、机械组织发达、虫媒花性状、线粒体数量多和叶绿体基粒片层少等是这一地区高山植物普遍具有的形态和结构特征。高山植物形成上述结构的特异性是高山特殊综合生态环境长期作用的结果。同时,也是高山植物对高山环境的高度适应。最后,对这一领域存在的问题以及未来研究的重点和方向进行了探讨。目的是引起国内研究者的关注,促进我国高山植物功能生态学的研究与发展。     

Biometrical evidence for adaptations of the salivary glands to pollen feeding in Heliconius butterflies (Lepidoptera: Nymphalidae)

The Neotropical genus Heliconius (Nymphalidae) is unique among butterflies for its pollen-feeding behaviour. With the application of saliva, they extract amino acids from pollen grains on the outside of the proboscis. We predicted that the salivary glands of pollen-feeding Heliconiinae would show adaptations to this derived feeding behaviour. A biometrical analysis of the salivary glands revealed that pollen-feeding butterflies of the genus Heliconius have disproportionately longer and more voluminous salivary glands than nonpollen-feeding Nymphalidae. The first two components in the principal component analysis explained approximately 95% of the total variance. The size-dependent factor score coefficients of body length and salivary gland parameters were predominately represented on axis 1. They significantly discriminated pollen-feeding from nonpollen-feeding heliconiines on that axis. Factor score coefficients for the volume of the secretory region of the salivary glands separated heliconiines from the outgroup species. The detailed biometrical analysis of salivary glands features thus provides strong evidence that the secretory regions of the salivary glands are larger in pollen-feeding butterflies. We concluded that pollen feeding is associated with a high production of salivary fluid.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 604–612.     

Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total-evidence analysis

We investigated the phylogeny of butterflies in the tribe Nymphalini sensu Harvey 1991, comprising the genera Vanessa, Cynthia, Bassaris, Aglais, Inachis, Nymphalis, Polygonia, Kaniska, Antanartia, Hypanartia, Symbrenthia, Mynes and Araschnia . Evidence from the mitochondrial gene ndl, the nuclear gene 'wingless' and from morphology/ ecology/behaviour were used separately and combined to analyse relationships. Phylogenies based on the different types of data agreed in many aspects of basic topology. We show that an analysis of only wing pattern characters (based on Nijhout's homology system) results in a topology broadly similar to the one resulting from analysis of the complete matrix. We found support for a monophyletic Nymphalini, where Hypanartia may be the sister clade to all other genera. Mynes, Symbrenthia and Araschnia together seem to form another basal clade. Evidence presented gives only moderate support for a monophyletic Vanessa in the wide sense, including also Cynthia and Bassaris , but strong support for the monophyly of the largely holarctic clade Aglais + Inachis + Nymphalis + Polygonia + Kaniska + Roddia . Within the latter group there is strong support for a clade consisting of Aglais + Inachis and for a second clade which includes Nymphalis, Polygonia (and its sister clade, the monotypic Kaniska) as well as Roddia l-album (= Nymphalis vaualbum ). As a consequence of this topology, Aglais is recognized as a taxon separate from Nymphalis . We present a hypothesis of species relationships within the focal group of genera. We also analyse and discuss the implications of excluding or including ecological data in phylogenetic tree construction, when the tree is to be used for studies in phylogenetic ecology.     

Biogeography of milkweed butterflies (Nymphalidae: Danainae) and mimetic patterns on tropical Paciflc archipelagos

Distributions of danaine butterfly species and associated mimetic patterns were compared among fifteen archipelagos of the tropical Pacific Ocean, and within five major archipelagos (the Bismarcks, Fiji, East and West Solomon Islands, and Vanuatu). Using both simple and stepwise linear regression analysis, variation in the total number of danaine species and number of mimetic patterns was assessed with respect to island size, isolation and elevation. Relative to interarchipelago distributions, the distribution of danaine species and number of mimetic patterns on islands within archipelagos exhibited less dependence upon interisland distance and island area. Geographical features influencing the number of mimetic patterns were similar to those of danaines as a whole. Analysis of residuals from stepwise linear regression suggested that factors influencing danaine distributions were different from those for non-danaine butterflies. This result is consistent with the hypothesis of enhancement of danaine species establishment through Müllerian mimicry, although other factors such as host plant availability and similar habitat use may also be important.     

Biology and morphology of the immature stages of Hermeuptychia atalanta (Lepidoptera: Nymphalidae: Satyrinae)

This paper describes the morphology of the immature stages (egg, larva and pupa) and larval behavior of the butterfly Hermeuptychia atalanta, a common and widespread species of satyrinae in South America. Eggs are white and spherical, laid singly on leaves of grasses. The first instar has a light-green body and dark-brown head; all remaining instars present body and head capsule light green. Larvae have no spines, and general morphology is similar to other species of the subtribe Euptychiina.     

Carotenoids of butterfly models and their mimics (Lep: Papilionidae and Nymphalidae)

Carotenoid concentration in model swallowtail butterflies is exceptionally high; it is suggested that they may exert a photo-protective function in species storing nitrophenanthrenes. Carotenoid concentration in the viceroy is also high. Again, it is suggested that this represents a protective mechanism for Lepidoptera feeding on Salix which contains salicin. Such a mechanism would not be required by the monarch storing cardenolides.     

The phylogenetics and biochemistry of host-plant specialization in Melitaeine butterflies (Lepidoptera: Nymphalidae)

Butterflies in the tribe Melitaeini (Lepidoptera: Nymphalidae) are known to utilize host plants belonging to 16 families, although most host-plant records are from four families. Of the 16 host-plant families, 12 produce secondary plant metabolites called iridoids. Earlier studies have shown that larvae of several melitaeine species use iridoids as feeding stimulants and sequester these compounds for larval defense. I investigate the evolutionary history of host-plant use in the tribe Melitaeini by testing a recent phylogenetic hypothesis of 65 species representing the four major species groups of the tribe. By simple character optimization of host-plant families and presence/absence of iridoids in the host plants, I find that plant chemistry is a more conservative trait than plant taxonomy. The ancestral host plant(s) of the entire tribe most likely contained iridoids and were likely to be in the plant family Plantaginaceae. A major host shift from plants containing iridoids to plants not containing iridoids has happened three times independently. The results show that the evolution of host-plant use in melitaeines has been (and still is) a dynamic process when considering plant taxonomy, but is relatively stable when considering host-plant chemistry.     

Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae: Limenitis)

The evolution of wing pattern diversity in butterflies has emerged as a model system for understanding the origins and maintenance of adaptive phenotypic novelty. Admiral butterflies (genus Limenitis) are an attractive system for studying wing pattern diversity because mimicry is common among the North American species and hybrid zones occur wherever mimetic and non-mimetic wing pattern races meet. However, the utility of this system has been limited because the evolutionary relationships among these butterflies remain unclear. Here I present a robust species-level phylogeny of Limenitis based on 1911 bp of two mitochondrial genes (COI and COII) and 904 bp of EF1-alpha for all five of the Nearctic species/wing pattern races, the majority of the Palearctic species, and three outgroup genera; Athyma, Moduza (Limenitidini), and Neptis (Limenitidinae: Neptini). Maximum-likelihood and Bayesian analyses indicate that the North American species are a well-supported, monophyletic lineage that is most closely related to the widespread, Palearctic, Poplar admiral (L. populi). Within North America, the Viceroy (L. archippus) is the basal lineage while the relationships among the remaining species are not well resolved. A combined maximum-likelihood analysis, however, indicates that the two western North America species (L. lorquini and L. weidemeyerii) are sister taxa and closely related to the wing pattern subspecies of the polytypic Limenitis arthemis species complex. These results are consistent with (1) an ancestral host-shift to Salicaceae by the common ancestor of the Poplar admiral and the Nearctic admiral lineage, (2) a single colonization of the Nearctic, and (3) a subsequent radiation of the North American forms leading to at least three independent origins of mimicry.     

Simulation of movement and egglaying inDanaus plexippus (Lepidoptera: Nymphalidae)

A model simulating the movement and oviposition of monarch butterflies over a female's life time is presented. The model's predictions compare favourably with observed data and suggest that females who lay eggs in an optimal fashion should have low directionalities in areas with high host plant density (patches and single plants) and high directionalities in areas with low host plant density. The model also provides one means of combining individual animal processes and spatial heterogeneity into population dynamics.     

A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA

Classification of the cosmopolitan butterfly genus Danaus (Nymphalidae: Danainae) is revised at subgeneric, specific and subspecific levels, combining for the first time mitochondrial and nuclear DNA sequence information with morphological data. Tree topologies based on the nuclear genome (allozymes, pheromone components, the morphology of all life history stages and nuclear DNA sequences), on the one hand, and mitochondrial DNA, on the other, are incongruent and challenge the current taxonomy of the genus. Although earlier classifications, based on adult morphology alone, are, in general, well supported by an analysis of total evidence, the mitochondrial phylogeny shows that the species D. chrysippus and its subgenus Anosia are deeply paraphyletic. Subspecies dorippus of D. chrysippus is the basal clade of the genus and is reinstated as the species D. dorippus. The former species D. plexaure is demoted to a subspecies of D. eresimus. The specific status of D. erippus, as distinct from D. plexippus, is tentatively supported. On the strength of the new data, division of the monophyletic genus Danaus s.l. into three subgenera Danaus s.s., Salatura and Anosia is unsustainable and is abandoned. Of the 15 terminal clades (taxa) of Danaus s.l. included in the study, 11 are species that broadly conform to the biological species concept. (The West Indian species D. cleophile, missing from our analysis, is the twelfth species). The remaining terminal clades are subspecies of D. chrysippus comb. nov. and D. dorippus stat. rev. Two sympatric Neotropical species, D. eresimus and D. gilippus, are morphologically distinct and sexually isolated but have nearly identical mitochondrial genomes. In contrast, two partially sympatric Palaeotropical species, D. chrysippus and D. dorippus, are cryptic species that share structural morphology and hybridize but have highly differentiated mitochondrial genomes. D. dorippus is polymorphic for two anciently diverged haplotypes and its history has possibly involved recombinational speciation and/or hybridism. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 191?212.     

Population structure and conservation of the coastal palm Coccothrinax borhidiana (Arecaceae) in the northwest of Cuba

Coccothrinax borhidiana is among the top 50 most threatened plants of Cuba. This palm is endemic to Punta Guanos-Punta Seboruco, Matanzas. Given the increase of gas-oil exploitation in the area where this palm population grows, the current situation of the species could be much more serious than what is known. We characterized the population structure based on a systematic sampling of 100 plots and evaluated the existing threats for re-evaluating its conservation status and proposing a recovery strategy. We sampled 685 individuals, 44.2 % juveniles and 55.8 % adults, whose population structure in 10 size classes did not conform to a log-normal distribution. This indicates that reproduction and/or recruitment processes are reduced, due to deforestation for the gas-oil industry, road construction and the establishment of small-holder, as well as the effect of cattle ranching and frequent fires. However, the higher representation of individuals in the three smaller size classes shows regeneration, which is observed in areas abandoned by the gas-oil industry. Despite this, existing threats have significantly reduced the area of occupancy of the species, and it is estimated that it will continue to decline by up to 80 %, making the species Critically Endangered. For these reasons, immediate management actions are needed to minimize the effect of threats on C. borhidiana, as well as to secure the management and implementation of ex situ conservation strategies that will allow the establishment of a new subpopulation in a nearby and ecologically similar area, which has the necessary legal protection status.     

Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae)

The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1alpha and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies.     

Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s . l ., Argynnina comprising Prokuekenthaliella , Issoria , Brenthis , and all the 'large fritillary' species joined in the genus Argynnis s . l . and a new subtribe Euptoietina comprising only the genus Euptoieta . The classical genus Issoria s . l . is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella . Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas , Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 627–673.     

Structures related to pheromone storage in alar androconia and the female abdominal scent gland of Heliconius erato phyllis,Heliconius ethilla narcaea,and Heliconius besckei (Lepidoptera: Nymphalidae: Heliconiinae)

We describe the morphology of alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei. Androconial scales of Heliconius, which are arranged in overlapping wing bands, release pheromones during courtship, probably through vibratory movements of male wings over the female to induce her to mate. An antiaphrodisiac is produced by glands located in the valves of the male and is transferred during copulation to the yellow dorsal abdominal sac present in the virgin female, causing this sac to emit a scent that reduces the attractiveness of the female for courtship with other males. Stereomicroscopy, SEM, and TEM analyses were conducted to describe the morphology of the internal and external scales and the external abdominal scent sac. The findings revealed different sizes of external androconial scales and an internal group of porous structural vesicles that are probably related to the preservation of internal space, reception and storage of secretions, and elimination of volatiles when the male is actively involved in courtship. Translucent projections on the female abdominal scent sac create open reservoirs for the reception, storage, and emission of antiaphrodisiac volatiles along with stink clubs. Male valve denticles vary in form and probably attach securely to the female sac during mating, thus ensuring secretion transfer. These features are discussed in the context of a comparative analysis.     

渤海湾津唐沿海野生大豆(Glycine soja)种群高盐碱胁迫反应

以渤海湾津唐沿海地带895份野生大豆种群在高盐碱(3%)环境下的全生育期鉴定试验,分析不同种群(株系)的生长发育动态以及对盐碱土壤生态适应性的分化。结果表明,出苗阶段存在3种耐盐碱能力类型:耐高盐碱胁迫发芽出苗;耐低盐碱胁迫发芽出苗;盐胁迫不能发芽出苗。在高盐胁迫下营养生长早期阶段种群死亡率最高,中期以后相对降低,成熟期死亡率较低。种群植株形态建成和生殖生长受到高盐胁迫的强烈抑制,表现早期严重而后期较轻。植株和产量性状的受害程度由大到小依次为:单株的地上干物质重、单株产量、种子数、百粒重和收获指数。株系间变异系数从大到小依次为:单株产量、种子数、荚数、收获指数、地上干物质重、百粒重。研究揭示该地域种群内对盐碱土壤有很高的适应性分化,存在高耐盐碱生态型。     

两种共存网蛱蝶幼期的生命表研究

在河北省赤城县大海坨国家自然保护区内,从2002年到2004年应用生命表方法研究了在同一网络斑块中共存的金堇蛱蝶Euphydryas aurinia和大网蛱蝶Melitaea phoebe两个世代的幼期各阶段的死亡情况,目的是了解影响种群动态的重要因子,为它们的长期保育提供信息。结果表明,金堇蛱蝶幼期总累积死亡率都较小,两个世代分别为59%和72%; 而大网蛱蝶总累积死亡率较高,两个世代分别为89%和80%。影响大网蛱蝶死亡的最重要因子是放牧,两个世代与放牧相关的k值分别为0.559和0.167;尤其是在越冬后大网蛱蝶幼虫组聚集取食阶段,导致两个世代分别有50%和25%的幼虫组死亡。寄生蜂在大网蛱蝶小种群下也分别使两个世代4%和9%的5～6龄期幼虫以及13%和24%的蛹死亡。金堇蛱蝶死亡主要发生在越冬期,与越冬死亡相关的k值两个世代分别为0.073和0.199, 而寄主植物的质量影响越冬期幼虫组死亡; 寄生蜂则对金堇蛱蝶种群的调控作用极小,只有在2003～2004世代有4.0% 的越冬后幼虫被寄生和7%的蛹被寄生。影响两种蛱蝶种群动态的关键因子不同,采取的保护措施应有所不同。在春季减少源斑块内的放牧,对于以源-汇集合种群形式存在的大网蛱蝶种群恢复和增长十分必要; 而对以经典集合种群形式存在的金堇蛱蝶, 通过适当植被管理提高繁殖区域内寄主植物质量,可以提高越冬期幼虫组存活率,有利于其长期持续生存。     

Rates of metabolism in diapausing and reproductively active tropical butterflies, Euploea core and Euploea sylvester (Lepidoptera: Nymphalidae)

Abstract.  Although the ecology of diapause has been widely studied in the field, the underlying physiological responses occurring in tropical diapausing insects remain virtually unexplored. This is especially the case with rates of respiration in diapausing tropical insect species. The present study compares rates of metabolism, as assessed by measurement of carbon dioxide production, between two species of diapausing and reproductively active tropical butterflies, Euploea core (W.S Macleay) and Euploea sylvester (Fabricius) , independent of temperature. Measurement of metabolism over a day-time/night-time regime confirms that these tropical butterflies display a diurnal rhythm in accordance with many other tropical and temperate insect species, regardless of developmental state. In addition, diapausing Euploea butterflies display rates of carbon dioxide production only 28% lower than those of reproductively active butterflies, and can terminate diapause within days of receiving the correct cues. Maintaining a similar metabolic rate throughout diapause, as well as a rapid termination of diapause, would enable these tropical butterflies to respond immediately to larval host plant resources, without the disadvantage of missing optimum conditions, allowing the species to maximize their reproductive potential.