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.     

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.     

The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae)

The generic names Euptychia, Cissia and Argyreuplychia are reviewed in relation to the Cissia confusa species-group in Costa Rica and Trinidad, for which the generic name Cissia Doubleday is formally resurrected. Three new species of Cissia are described, C. confusa itself is resurrected from synonymy, three neotypes and three lectotypes are designated, and six new combinations established. On the basis of early instar morphology three subgroups are recognized and the general biology of the confusa species-group is summarized. A key to species in Costa Rica is provided and the ecology of the adults is reviewed.     

Life history of Manataria maculata (Lepidoptera: Satyrinae) from Costa Rica

The life history and early stages of the satyrine butterfly Manataria maculata are described and illustrated from Costa Rica. Eggs are laid on Lasiacis sp. (Panicoideae), a new non-bamboo host plant for the genus Manataria. The larval stage varied from 23 to 28 days, and the pupal duration was approximately 12 days when reared on Bambusa vulgaris and Guadua angustifolia in captivity at 23-24 degrees C.     

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.     

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.     

Seasonal plasticity in growth and development of the speckled wood butterfly, Pararge aegeria (Satyrinae)

Regulation of growth and development by photoperiod was studied in a population of the speckled wood butterfly, Purarge aegeria L. (Lepidoptera: Satyrinae), from southern Sweden. Individuals were reared in a range of photoperiodic regimes (9L. to 22L) and temperatures (13°C to 21° C). Plasticity was found for important life-history traits- generation time, growth rate and final weight and seasonal regulation of development in response to photoperiod was found to occur at two levels. Purarge aegeria hibernates as a third instar larva or in the pupal stage, cantering one of four major developmental pathways in response to photoperiod: (1) direct development in both the larval and pupal stages, (2) pupal winter diapause with or (3) without a preceding larval summer diapause, or (4) larval winter diapause. In addition to this high-level regulation of individual development, larval growth rate and pupal development rate also appear to be finally regulated by photoperiod within each major pathway. As photoperiods decreased from 22 h to 17 h at 17° C, growth rate among directly developing larvae increased progressively, as was the case for larva? developing according to a univoltine life cycle from 17 h to 14 h. At two photoperiods, 13 h and 16 h (corresponding to shifts between major pathways), both larval and pupal development were extremely variable with the fastest individuals developing directly and the slowest developing with a diapause. This indicates a gradual nature of diapause itself, suggesting that the two level may not he fundamentally different.     

Limited hybridization along a large contact zone between two genetic lineages of the butterfly Erebia medusa (Satyrinae, Lepidoptera) in Central Europe

Genetic lineages evolving during glacial isolation frequently come into contact as the result of postglacial range expansions. Hybridization often occurs along these contact zones. In Europe, the high mountain systems of the Alps and Pyrenees are well known for their hybrid belts. This article studies the contact zone of the Woodland Ringlet Erebia medusa in a Hercynian mountain area in the Czech-German border region not exceeding 1500 m a.s.l. Hybrid populations between an eastern and a western genetic lineage were detected by amova -based tests, principal component analysis and neighbour joining analysis. Over most of the range of the Czech-German Border Mts, the ridges separate the western and the eastern genetic lineage from each other. However, two important hybrid areas were detected: (1) the watershed of the Ohře river in the north-west of that area, a major valley system passing through these mountains and (2) the high plateaux of the Šumava Mts in the south-east, an extended area of high elevation. The location of hybrid populations in geographical vicinity to non-hybrid populations and the generally low F _IS (2.1%) make reduced fitness of hybrid individuals little likely. The hybrid populations have intermediate genetic diversity between the genetically poor western and the genetically rich eastern lineage populations.     

A new species of Cyllopsis (Lepidoptera: Nymphalidae: Satyrinae) from Costa Rica

Cyllopsis emilia Chacón and Nishida, a new satyrine species, is described from a single male specimen from Cerro de la Muerte, San José, Costa Rica. This new species can be distinguished from other species of Cyllopsis by its white coloration.     

A new genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from southern Brazil

This paper describes a new genus and a new species of Euptychiina from open grassland habitats (campos de cima da serra) in southern Brazil. The systematic position of this new taxon is discussed based on morphological and molecular data, and it is considered sister to Taydebis Freitas. Since the campos vegetation is considered endangered due to anthropogenic activities, this butterfly species deserves attention and should be included in future conservation plans for this biome.     

Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data

We have inferred the first empirically supported hypothesis of relationships for the cosmopolitan butterfly subfamily Satyrinae. We used 3090 base pairs of DNA from the mitochondrial gene COI and the nuclear genes EF-1alpha and wingless for 165 Satyrinae taxa representing 4 tribes and 15 subtribes, and 26 outgroups, in order to test the monophyly of the subfamily and elucidate phylogenetic relationships of its major lineages. In a combined analysis, the three gene regions supported an almost fully resolved topology, which recovered Satyrinae as polyphyletic, and revealed that the current classification of suprageneric taxa within the subfamily is comprised almost completely of unnatural assemblages. The most noteworthy findings are that Manataria is closely related to Melanitini; Palaeonympha belongs to Euptychiina; Oressinoma, Orsotriaena and Coenonympha group with the Hypocystina; Miller's (1968). Parargina is polyphyletic and its components group with multiple distantly related lineages; and the subtribes Elymniina and Zetherina fall outside the Satyrinae. The three gene regions used in a combined analysis prove to be very effective in resolving relationships of Satyrinae at the subtribal and tribal levels. Further sampling of the taxa closely related to Satyrinae, as well as more extensive sampling of genera within the tribes and subtribes for this group will be critical to test the monophyly of the subfamily and establish a stronger basis for future biogeographical and evolutionary studies.     

Effect of male body size on sperm precedence in the polyandrous butterfly Pieris napi L. (Lepidoptera: Pieridae)

Male Lepidoptera produce an ejaculate during copulation thatcontains both sperm and accessory gland nutrients and may functionas paternal investment and/or male mating effort Several studieshave examined how ejaculates function as paternal investment,but few have determined the influence of sperm competition onmale investment This study examines the effect of male bodysize on sperm precedence in the polyandrous butterfly Pierisnapi L. We used male body mass as an indicator of the size ofejaculate transferred and found that relative male size hada significant effect on paternity. The offspring of twice-matedfemales showed a low incidence of mixed paternity. Larger malesobtained the majority of fertilizations, and the degree of second-malesperm precedence was influenced by relative body size of matingmales. In general, second mates obtained fewer fertilizationsthe larger the size of the first mate. The interval betweenthe first and second mating was influenced by the size of thefirst male mate Females first mated to small males remated soonerthan females first mated to larger males Our results suggestthat large males may have a selective advantage over small maleswhen both a male's fertilization success and a female's refractoryperiod are influenced by the size of ejaculate transferred.Furthermore, the effect of male body size on the proportionof offspring sired lends support to the hypothesis that spermcompetition has played a major role in the evolution of ejaculatesize.     

Tropical dry and wet season polyphenism in the butterfly Melanitis leda (Satyrinae): Phenotypic plasticity and climatic correlates

Changes in wing pattern, colour, shape and size associated with seasonal polyphenism in Melanitis leda were quantified using a series of 155 butterflies collected by N. Manders on Mauritius in 1905. Butterflies of the wetter period were predominantly of the wet season form with large, well differentiated eyespots, short tails, smaller wings and a characteristic background colour. The dry season form occurred only in the drier period and has much smaller eyespots, longer tails, larger wings and a variable background. Many intermediates occurred, mainly in the drier period. These are associated with an absence of extreme seasonal change in Mauritius. The first principal component (PCI) describing the morphometric and colour data is closely related to the wing form (r = 0.80). Regression analyses using temperature and rainfall data for the 8 weeks before each capture showed that about 40% of variation in PCI could be accounted for by temperature in weeks 2–3 before capture. Many of the characters measured are redundant; a subset of seven morphometric characters yields a closely similar PCI. Analysis of is subset in an additional sample of 70 M. leda from Kenya showed that the seasonal polyphenism overrides a small degree of sexual dimorphism. The results are discussed with regard to seasonal changes in adult activity, resting backgrounds and visual predation. Wing phenotype characters are part of an array of coordinated morphological and life history traits which include ovarian dormancy and fat body development in dry season adults. A partial independence occurs in the proximal control of these traits as indicated, for example, by the larger wing and tail size, and smaller eyespots of the small number of the wet season form captured in the drier period in comparison to those of the wetter months.     

Interspecific variation in mitochondrial serine transfer RNA (UCN) in Euptychiina butterflies (Lepidoptera: Satyrinae): structure and alignment

The nucleotide variation and structural patterns of mitochondrial RNA molecule have been proposed as useful tools in molecular systematics; however, their usefulness is always subject to a proper assessment of homology in the sequence alignment. The present study describes the secondary structure of mitochondrial tRNA for the amino acid serine (UCN) on 13 Euptychiina species and the evaluation of its potential use for evolutionary studies in this group of butterflies. The secondary structure of tRNAs showed variation among the included species except between Hermeuptychia sp1 and sp2. Variation was concentrated in the ribotimidina-pseudouridine-cystosine (TψC), dihydrouridine (DHU) and variable loops and in the DHU and TψC arms. These results suggest this region as a potential marker useful for taxonomic differentiation of species in this group and also confirm the importance of including information from the secondary structure of tRNA to optimize the alignments.     

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

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

Microhabitat use and hindwing phenotype in Hipparchia semele (Lepidoptera, Satyrinae): thermoregulation and background matching

Abstract. 1. Microhabitat use and background matching by adult Hipparchia semele (L.) in a coastal sand dune site in south Wales were studied during the flight period of 1986.
2. Microhabitat use by both sexes is related to temperature and activity. When settling substrates are cool, typically during the early morning, most individuals are located on the warmest basking substrate, lichen patches. After a period of warming, males perch to locate mates on exposed areas of sand where there is a large field of view and where they are visually apparent. Egg-laying females tend to settle on lichen patches throughout the day.
3. Overall, females are maximally visually cryptic when settled on lichen patches with unexposed forewings. Male crypsis on lichen is less effective, but greater on most other substrates, than that of females. It is argued that individuals match lichen patches because this substrate is used when they are most liable to detection and capture by vertebrate predators. Male crypsis is probably a compromise between maximizing protection on lichen patches and requirements for resemblance to other substrates.
4. The underside hindwing phenotype does not match sand. Background matching to this substrate is probably not important because it tends to be used when individuals are active and can effectively use secondary defence mechanisms.
5. It is suggested that the apical eyespot and orange patch on the forewing underside have a dual role in secondary defence, acting as a startling and deflective device, being exposed by disturbed individuals and by those engaged in activities which may disrupt the visual crypsis afforded by the hindwing underside.     

中国粉眼蝶属分类研究（鳞翅目：蛱蝶科：眼蝶亚科）

系统整理分布在中国的粉眼蝶属Callarge Leech,[1892]种类,已知1种4亚种,其中含1中国新纪录亚种Callarge occidentalis fansipana Monastyrskii,2005;通过外部形态和雄性生殖器特征研究,提出将其作为Callargesagitta fansipana Monastyrskii,stat.nov.阐述各亚种的主要特征及地理分布,并附有各亚种雄性外生殖器特征图和成虫照片.