Host plant relationships of an endangered butterfly, Lopinga achine (Lepidoptera: Nymphalidae) in northern Europe

The aim of the present study was to evaluate—in a geographic perspective—the role of host plant as a determinant of habitat quality for Lopinga achine, a satyrine butterfly endangered over much of its European range. Laboratory trials were performed to record host choices made by the ovipositing females as well as by neonate larvae. In rearing experiments, growth performance and mortality on different host plants was determined. Oviposition was found to be indiscriminate but larvae were shown to be able to choose between host plants, with the choices made broadly consistent with growth performance of the larvae on particular hosts. Nevertheless, most grasses and sedges offered were found to support larval development reasonably well. No clear superiority of the previously suggested primary host plant Carex montana could be shown. Importantly, no differences in host plant relationships were found between the populations of Sweden, western Estonia and eastern Estonia. In particular, the larvae originating from eastern Estonian populations developed on C. montana equally well even if the plant is absent from their native habitat. In the context of species conservation, one should conclude that L. achine is polyphagous enough on various grasses and sedges so that the presence of any particular host species cannot be a critical component of habitat quality. Nevertheless, some preference to broad- and soft-leaved hosts, as well as sensitivity to host wilting, may partly explain the butterfly’s preference to moist forest habitats, further emphasizing the central role of habitat management in the conservation practice of this species. In turn, the absence of ecological differences between geographic populations should enable conservationists to successful transfer their experience across national boundaries.     

Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera,Nymphalidae, Satyrinae)

Peña, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. (2010). Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae).—Zoologica Scripta, 39, 243–258. The diverse butterfly subtribe Euptychiina was thought to be restricted to the Americas. However, there is mounting evidence for the Oriental Palaeonympha opalina being part of Euptychiina and thus a disjunct distribution between it (in eastern Asia) and its sister taxon (in eastern North America). Such a disjunct distribution in both eastern Asia and eastern North America has never been reported for any butterfly taxon. We used 4447 bp of DNA sequences from one mitochondrial gene and four nuclear genes for 102 Euptychiina taxa to obtain a phylogenetic hypothesis of the subtribe, estimate dates of origin and diversification for major clades and perform a biogeographic analysis. Euptychiina originated 31 Ma in South America. Early Euptychiina dispersed from North to South America via the temporary connection known as GAARlandia during Eocene–Oligocene times. The current disjunct distribution of the Oriental Palaeonympha opalina is the result of a northbound dispersal of a lineage from South America into eastern Asia via North America. The common ancestor of Palaeonympha and its sister taxon Megisto inhabited the continuous forest belt across North Asia and North America, which was connected by Beringia. The closure of this connection caused the split between Palaeonympha and Megisto around 13 Ma and the severe extinctions in western North America because of the climatic changes of the Late Miocene (from 13.5 Ma onwards) resulted in the classic ‘eastern Asia and eastern North America’ disjunct distribution.     

Immature stages of the butterfly Magneuptychia libye (L.) (Lepidoptera: Nymphalidae, Satyrinae)

The immature stages of the butterfly Magneuptychia libye (L.) are described, and their morphology is compared with other Neotropical Satyrinae.     

Population viability analysis of the butterfly Lopinga achine in a changing landscape in Sweden

Metapopulation theory has generally focused only on the stochastic turn-over rate among populations and assumed that the number and location of suitable habitat patches will remain constant through time. This study combines in a PVA both the deterministic landscape dynamics and the stochastic colonisations and extinctions of populations for the butterfly Lopinga achine in Sweden. With data on occupancy pattern and the rate of habitat change, we built a simulation model and examined five different scenarios with different assumptions of landscape changes for L. achine . If no landscape changes would be expected, around 80 populations are predicted to persist during the next 100 yr. Adding the knowledge that many of the sites are unmanaged and that the host plant will slowly deteriorate as canopies close over, and adding environmental variation and synchrony, showed that the number of populations will decrease to around of 4.3 and 2.8 respectively, with an extinction risk of 34% – quite different from the first scenario based only on the metapopulation model. This study has shown the importance of incorporating both deterministic and stochastic events when making a reliable population viability analysis. Even though one can not expect that the long-term predictions of either occupied patches or extinction risks will be accurate quantitatively, the qualitative implications are correct. The extinction risk will be high if grazing is not applied to more patches than is the case today. The simulations indicate that an absolute minimum of 10–30 top-ranked patches needs to be managed for the persistence of the metapopulation of L. achine in the long term. The same problem of abandoned and overgrowing habitats affects many other threatened species in the European landscape and a similar approach could also be applied to them.     

Phylogeography of the threatened crayfish (genus Austropotamobius) in Italy: implications for its taxonomy and conservation

A nucleotide sequence analysis of a portion of the mitochondrial large ribosomal subunit was performed to define the phylogeography of the threatened crayfish Austropotamobius (Decapoda; Astacidae) in Italy. We collected 61 specimens from 31 localities across the Italian peninsula. For the phylogenetic inference, we combined the 61 Austropotamobius spp sequences obtained from this study with 18 sequences deposited in GenBank and corresponding to Italian, French, Irish, Swiss, and Slovenian locations. Among the analysed sequences, 34 distinct haplotypes were detected. Our results confirmed the presence of both A. pallipes and A. italicus in the Italian peninsula and the existence within the latter species of a strong intraspecific genetic variation, due to the occurrence of four subspecies with a well-defined geographic distribution. From a conservation viewpoint, Italy, with its high haplotype variability, may be considered a 'hot spot' for the genetic diversity of the European native crayfish Austropotamobius. We suggest that re-introduction programs should be conducted with extreme caution in Italy, since not only the two Austropotamobius species but also the four A. italicus subspecies are genetically and taxonomically separate units and require independent conservation plans.     

Patterns and timing of diversification in a tropical montane butterfly genus,Lymanopoda (Nymphalidae,Satyrinae)

Species distributions are a product of contemporary and historical forces. Using phylogenetic and geographic data, we explore the timing of and barriers to the diversification of the Andean butterfly genus Lymanopoda (Nymphalidae, Satyrinae). Clade and species level diversification is coincident with Andean orogeny and Pleistocene glaciation cycles. Lymanopoda has primarily diversified within elevational bands, radiating horizontally throughout the Andes with occasional speciation across elevational boundaries, often associated with ecotones. Narrow elevational ranges and infrequent speciation into adjacent elevational strata suggest that expansion across elevational gradients is relatively difficult. These results are similar to those found in studies of other Andean taxa.     

The species of the genus Zethera Felder (Lepidoptera: Nymphalidae,Satyrinae)

The butterfly genus Zethera Felder, endemic to the Philippines and northern Sulawesi, is shown to include six species, four of which comprise the Z. pimplea (Erichson) superspecies. Type data is given for all nominal taxa, and a key to the species recognised is presented, together with data on their variation and distribution. Attention is drawn to the zoogeographic subregions of the Philippine islands, as originally described by Semper, and illustrated by the Zethera species. Their mimicry is also noted, with particular reference to its sex-limitation in the Z. pimplea superspecies, and non-limitation in Z. incerta (Hewitson) and Z. hestioides C. & R. Felder. The need to demonstrate whether or not Zethera represents a true monophyletic unit is briefly discussed.     

Key traits in a threatened butterfly and its common sibling: implications for conservation

We ask here which traits predispose one species to extreme rarity and possible extinction while a sympatric sibling is geographically widespread. With background knowledge on the level of habitat specialization of the two species, the population structure and movement of the localized and threatened Orachrysops ariadne were compared to those of the common and highly sympatric O. subravus, using mark-release-recapture. Of a total of 290 marked O. ariadne individuals 42.8% were recaptured, while of 631 O. subravus individuals 49.3% were recaptured. The Jolly-Seber model was used to estimate daily population numbers (N _i ), survival rates (ϕ _i ), recruitment rates (B _i ), proportion of marked animals in the total population (∝_i), and the number of marked animals at risk (M _i ). O. ariadne is a remarkably rare animal, averaging only 10 individuals ha⁻¹ within its small, remaining colonies. Average residence times of male adults were generally similar in both species, being just over 5 days. O. ariadne is a strong and rapid back and forth flier, covering mean recapture distances of 157 m, almost twice that of O. subravus, principally in search of scarce nectar sources. In short, the rarity of O. ariadne is not so much to do with behaviour, survivorship or longevity, but rather with limited availability of the specialized habitat patches for both larvae and adults, and, in particular, the extreme scarcity of the host plant. Evidence suggests that there has been very high selection pressure on the key trait of strong flight as a compensation for going down the apparently highly risky path of extreme microhabitat specialization. Of concern for conservation of this rare species is that these rare habitat patches have become increasingly isolated through transformation of the surrounding landscape. Reduction of the barrier effects of agroforestry through creation of linkages between colonies is recommended, especially as O. ariadne is such a strong flier. Such corridors are indeed now being implemented.     

Visual mate-searching behaviour in the evening brown butterfly, Melanitis leda (L.) (Lepidoptera: Nymphalidae)

Abstract As in most animals, male butterflies are generally the more proactive sex with respect to seeking out mating opportunities. In most cases, males conduct their search sometime between mid-morning and mid-afternoon, but a few species are active only before dawn or after sunset. These crepuscular species offer a good opportunity to study how males deal with markedly different visual and thermal conditions. Here, I present data from a 5-month behavioural study of male Melanitis leda (Nymphalidae) at a mate-encounter site in tropical Australia. Males of this species defended perching locations along a forest edge in a similar manner to other diurnally active territorial nymphalids. They generally arrived at these sites after sunset and arrived earlier on evenings that darkened earlier. Actively mate-locating males were only seen at the site during a 25−35 min evening period, during which ambient light levels ranged between 50 and 2600 lux. Only 27% of marked territory residents were recorded again at their location of capture, but fidelity in this 'resighted' group ranged up to 23 days. A sample of males, captured under ambient temperatures of 24.0−27.2°C, maintained a mean thoracic excess of 8.25 ± 0.73°C, but did not appear to 'shiver' in the manner of other crepuscular species. Males courted conspecific females and one mating was observed. This species is an excellent candidate for further research into the evolution of mating tactics in crepuscular butterflies.     

Egg size variation in European Satyrine butterflies (Nymphalidae, Satyrinae)

The interspecific relationships between egg size and female size, sexual size dimorphism, latitude, preoviposition and egg-laying behaviour, was investigated in European Satyrine butterflies by controlling for taxonomic relatedness and by simple cross-species comparisons. Egg size scales positively to female size, although the relationship is not significant (but nearly so) when taxonomic effects are controlled for. Egg size is correlated with latitude. When the effects of latitude are removed, the relationship between egg and female size becomes more evident. Sexual size dimorphism is negatively correlated with egg size, but the effect disappears when adult size is controlled for. Small egg size is related to long preoviposition. No relationship between egg laying mode and egg size appears. The results are interpreted as evidence for positive scaling of egg to female size, with superimposed effects of egg size radiation. The correlation of egg size with latitude suggests that temperature, or season length, is likely to have made an important contribution to egg size selection within this geographic area.     

Phylogeography and population expansion of Western Himalayan highly-erratically migratory painted lady butterfly (Lepidoptera: Nymphalidae: Vanessa cardui)

Understanding population genetics, genetic diversity and the migration pattern of ecologically important species is crucial for effective conservation plans. We studied mitochondrial cytochrome c oxidase subunit-I (COI) gene variation in painted lady butterfly (Vanessa cardui) from different altitudinal gradients of Uttarakhand (Western Himalayas), India. We sequenced COI) gene of 91 samples and 22 COI sequences retrieved from GenBank as representatives of different localities for comparative analysis. We observed a total of 16 haplotypes, of which seven haplotypes were observed in Uttarakhand with intra-species sequence divergence ranges from 0.001 to 0.009. Haplotype (Hd) and nucleotide diversity (π) were comparatively low and averaging 0.635 and 0.002, respectively. The neutrality test, mismatch distribution graph and observed pattern of diversity indices (Hd and π) revealed population expansion. It was further confirmed by the Median Joining Network, which exhibited a star-like topology from the core haplotype, while the Bayesian phylogenetic tree displayed a mixing of haplotypes from different localities to an identical clade. Haplotype organization in the Bayesian tree and Median Joining Network was not clustered as per their topographical affinity, indicating incomplete lineage sorting and late species population development or colonization in the novel area.     

Phylogeography of the bushmaster (Lachesis tnuta: Viperidae): implications for neotropical biogeography, systematics, and conservation

We used mitochondrial gene sequences to reconstruct phylogenetic relationships among subspecies of the bushmaster, Lachesis muta. These large vipers are widely distributed in lowland tropical forests in Central and South America, where three of four allopatric subspecies are separated by montane barriers. Our phylogeny indicates that the four subspecies belong to two clades, the Central American and South American lineages. We use published molecular studies of other taxa to estimate a 'reptilian mtDNA rate' and thus temporal boundaries for major lineage divergences in Lachesis. We estimate that the Central and South American forms diverged 18-6 Mya, perhaps due to the uplifting of the Andes, whereas the two Central American subspecies may have diverged 11-4 Mya with the uprising of the Cordillera de Talamanca that separates them today. South American bushmasters from the Amazon Basin and the Atlantic Forest are not strongly differentiated, perhaps due to episodic gene flow during the Pleistocene, when suitable habitat for this species was at times more continuous. Our results agree with previous evidence that genetic divergence among some neotropical vertebrates pre-dated Pleistocene forest fragmentation cycles and the appearance of the Panamanian Isthmus. Based on morphological, behavioral, and molecular evidence, we recognize three species of Lachesis. In addition to L. muta, the widespread South American form, the Central American forms are treated as distinct species (L. meknocephak and L. stenophrys), each deserving of special conservation status due to restricted distribution and habitat destruction.     

Phylogeography and counter-intuitive inferences in island biogeography: evidence from morphometric markers in the mobile butterfly Maniola jurtina (Linnaeus) (Lepidoptera, Nymphalidae)

Distribution of mobile organisms on near-continent islands is mainly shaped by factors operating over ecological rather geological time. However, the phylogeography of single species has the potential to expose historical factors at work. In the present study, West Mediterranean populations of the butterfly Maniola jurtina are studied using geometric morphometrics. The distribution of the two well established lineages ( Maniola jurtina jurtina in the Atlanto–Mediterranean area and Maniola jurtina janira in the Central–Eastern-Mediterranean area) on 12 islands and the adjoining continents are compared. The south-western lineage unexpectedly occurs on islands close to shores occupied by the eastern lineage. We have modelled the distribution of the lineages using three different hypotheses: (1) a contemporary isolation model, which predicts lineage occupancy of islands is linked to relative distances from neighbouring continental areas; (2) a refugial hypothesis, which predicts one lineage to be the ancestral one for the whole region studied, and then successively replaced over part of it; (3) a changing geography hypothesis, which predicts the two lineages to have evolved in their currently occupied areas, continuously sourcing islands subsequent to the Würm maximum glaciation. Of the three models, the refugial hypothesis is most highly correlated with the observed pattern, suggesting that Mediterranean islands may function as refugia during cold periods, much as the three mainland peninsulas of Iberia, Italy and Greece are known to have done. Thereafter, hybridization on the nearest and smallest islands has occurred, with the entire process supporting the notion of the joint influence of factors in ecological and geological time.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 677–692.     

The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods

We have inferred the most comprehensive phylogenetic hypothesis to date of butterflies in the tribe Satyrini. In order to obtain a hypothesis of relationships, we used maximum parsimony and model‐based methods with 4435 bp of DNA sequences from mitochondrial and nuclear genes for 179 taxa (130 genera and eight out‐groups). We estimated dates of origin and diversification for major clades, and performed a biogeographic analysis using a dispersal–vicariance framework, in order to infer a scenario of the biogeographical history of the group. We found long‐branch taxa that affected the accuracy of all three methods. Moreover, different methods produced incongruent phylogenies. We found that Satyrini appeared around 42 Mya in either the Neotropical or the Eastern Palaearctic, Oriental, and/or Indo‐Australian regions, and underwent a quick radiation between 32 and 24 Mya, during which time most of its component subtribes originated. Several factors might have been important for the diversification of Satyrini: the ability to feed on grasses; early habitat shift into open, non‐forest habitats; and geographic bridges, which permitted dispersal over marine barriers, enabling the geographic expansions of ancestors to new environments that provided opportunities for geographic differentiation, and diversification.

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Further observations on the occurrence and mimicry of Mycalesis drusillodes (Lepidoptera: Nymphalidae,Satyrinae)

A previously unknown population of the Satyrid Mycalesis drusillodes (Oberthür) is described from the Torricelli Mountains, which considerably extends the known range of this New Guinea species. New information is given concerning the two mimetic complexes to which this dual mimic butterfly belongs.     

Phylogenetics of Coenonymphina (Nymphalidae: Satyrinae) and the problem of rooting rapid radiations

We report a rapid radiation of a group of butterflies within the family Nymphalidae and examine some aspects of popular analytical methods in dealing with rapid radiations. We attempted to infer the phylogeny of butterflies belonging to the subtribe Coenonymphina sensu lato using five genes (4398 bp) with Maximum Parsimony, Maximum Likelihood and Bayesian analyses. Initial analyses suggested that the group has undergone rapid speciation within Australasia. We further analyzed the dataset with different outgroup combinations the choice of which had a profound effect on relationships within the ingroup. Modelling methods recovered Coenonymphina as a monophyletic group to the exclusion of Zipaetis and Orsotriaena, irrespective of outgroup combination. Maximum Parsimony occasionally returned a polyphyletic Coenonymphina, with Argyronympha grouping with outgroups, but this was strongly dependent on the outgroups used. We analyzed the ingroup without any outgroups and found that the relationships inferred among taxa were different from those inferred when either of the outgroup combinations was used, and this was true for all methods. We also tested whether a hard polytomy is a better hypothesis to explain our dataset, but could not find conclusive evidence. We therefore conclude that the major lineages within Coenonymphina form a near-hard polytomy with regard to each other. The study highlights the importance of testing different outgroups rather than using results from a single outgroup combination of a few taxa, particularly in difficult cases where basal nodes appear to receive low support. We provide a revised classification of Coenonymphina; Zipaetis and Orsotriaena are transferred to the tribe Eritina.