Skipper Butterflies Research Articles

The genome sequence of the Essex Skipper butterfly, Thymelicus lineola (Ochsenheimer, 1808)

We present a genome assembly from an individual female Essex Skipper butterfly, Thymelicus lineola (Arthropoda; Insecta; Lepidoptera; Hesperiidae). The genome sequence spans 511.80 megabases. Most of the assembly is scaffolded into 30 chromosomal pseudomolecules, including the Z and W sex chromosomes. The mitochondrial genome has also been assembled and is 17.24 kilobases in length.

Reduced palatability, fast flight, and tails: decoding the defence arsenal of Eudaminae skipper butterflies in a Neotropical locality.

Prey often rely on multiple defences against predators, such as flight speed, attack deflection from vital body parts, or unpleasant taste, but our understanding on how often and why they are co-exhibited remains limited. Eudaminae skipper butterflies use fast flight and mechanical defences (hindwing tails), but whether they use other defences like unpalatability (consumption deterrence) and how these defences interact have not been assessed. We tested the palatability of 12 abundant Eudaminae species in Peru, using training and feeding experiments with domestic chicks. Further, we approximated the difficulty of capture based on flight speed and quantified it by wing loading. We performed phylogenetic regressions to find any association between multiple defences, body size, and habitat preference. We found a broad range of palatability in Eudaminae, within and among species. Contrary to current understanding, palatability was negatively correlated with wing loading, suggesting that faster butterflies tend to have lower palatability. The relative length of hindwing tails did not explain the level of butterfly palatability, showing that attack deflection and consumption deterrence are not mutually exclusive. Habitat preference (open or forested environments) did not explain the level of palatability either, although butterflies with high wing loading tended to occupy semi-closed or closed habitats. Finally, the level of unpalatability in Eudaminae is size dependent. Larger butterflies are less palatable, perhaps because of higher detectability/preference by predators. Altogether, our findings shed light on the contexts favouring the prevalence of single versus multiple defensive strategies in prey.

The Impact of Wildflower Habitat on Insect Functional Group Abundance in Turfgrass Systems.

Urbanization is rapidly influencing the abundance and diversity of arthropods. Within urban systems, managed turfgrass is a prominent land cover which can support only a limited number of arthropod groups. To allow for more arthropod biodiversity and to support beneficial insects within turfgrass, increasing numbers of land managers are choosing to partially convert turf habitat to wildflower habitat using commercially available seed mixes. However, the population dynamics of arthropod groups in these systems are poorly known, with consequentially little information on best long-term practices for managing wildflower habitats in turfgrass systems. To address this gap, we sampled insects using pan traps in turfgrass systems pre- and post-implementation of wildflower habitats and examined the change in abundance of several insect families and functional guilds. Insect groups had variable responses to wildflower habitat implementation, with some groups such as sweat bees and skipper butterflies showing a decline two years post-implementation. Other groups, such as predatory flies, were relatively more abundant one and two years post-implementation. These variable responses point to the need for more research on the long-term effects of wildflower habitats on beneficial insects in turfgrass habitats.

Can species distribution models using remotely sensed variables inform reintroductions? Trialling methods with Carterocephalus palaemon the Chequered Skipper Butterfly

Despite large numbers of reintroduction projects taking place and the high cost involved, there is a generally low success rate. Insects in particular are understudied within reintroduction ecology, with guidelines focusing on more iconic vertebrate taxa. Species distribution models (SDMs) examine the associations between species observations and environmental variables to find the conditions in which populations could survive. This study utilises two frequently used SDM approaches, a regression model (general linear model (GLM)) and a machine learning method (MaxEnt) to model habitat suitability for Chequered Skipper, Carterocephalus palaemon, butterflies, which are being reintroduced to Northamptonshire following extinction in England. We look at how SDMs using widespread remotely sensed variables could be used to inform the reintroduction process by finding areas of suitable habitat that were previously overlooked. These remotely sensed variables have the potential to inform reintroductions without extensive on the ground research as they cover huge areas and are widely available. We found that both models are successful in discriminating between presences and absences, using only a limited number of explanatory variables. We conclude that these wide-scale SDMs are useful as a first step in the decision-making process in determining appropriate sites for reintroductions, but that they are less accurate when establishing precisely where species should be placed.

Skipper butterflies (Hesperiidae: Lepidoptera) common to Sulawesi (Indonesia) and India, with their host plants

The present communication deals with the Skipper butterflies (Hesperiidae: Lepidoptera) common to Sulawesi (Indonesia) and India with their common names, host plants and updating.

Immature stages, new host plant records and shelter structures of Troyus phyllides (Röber, 1925) and Thoon ponka Evans, 1955 in the Peruvian Amazon (Lepidoptera: Hesperiidae: Hesperiinae: Hesperiini)

We describe here for the first time the complete immature life cycles and shelter structures of two Neotropical skipper butterflies in the subtribe Moncina, Troyus phyllides (Röber, 1925) and Thoon ponka Evans 1955, along with new natural host plant records for these species at Finca Las Piedras, Madre de Dios, Peru. Four eggs and a preantepenultimate larva of T. phyllides, as well as three eggs of Thoon ponka were collected in nature and each passed through five larval instars to adulthood. Troyus phyllides fed on a herbaceous species, Lasiacis ligulata Hitchcock & Chase (Poaceae: Panicoideae: Paniceae), while T. ponka fed on two congeneric herbaceous bamboo species, Pariana lunata Nees and Pariana sp. (Poaceae: Bambusoideae: Olyreae). We present photos of all immature stages and host plants, as well as illustrations of the shelter structures and the head capsules for each of these two species.

New taxa of skipper butterflies (Lepidoptera: Hesperiidae) from tropical Africa

Three new species, Apallaga klaudiae sp. n., Gorgyra ziama sp. n., Andronymus teresae sp. n. and a new subspecies, Eagris tetrastigma lomana ssp. n., of skipper butterflies (Lepidoptera: Hesperiidae) from the tropical forest zone of Africa are described in comparison with their presumed closest relatives.

The Chinese species of skipper butterflies in the tribe Tagiadini Mabille, 1878 (Lepidoptera: Hesperiidae): insights from phylogeny, hostplants, and biogeography

The Chinese species of skipper butterflies in the tribe Tagiadini Mabille, 1878 (Lepidoptera: Hesperiidae): insights from phylogeny, hostplants, and biogeography

Sympiesis notata (Hymenoptera: Chalcidoidea: Eulophidae), a gregarious parasitoid of Hesperiidae (Lepidoptera), and some records of other Chalcidoidea associated with skipper butterflies in the Western Palaearctic

Sympiesis notata is a gregarious ectoparasitoid of skipper butterfly caterpillars. Other Lepidoptera larvae in leaf mines and leaf folds are attacked by the same morphospecies, but on these smaller hosts S. notata is usually a solitary parasitoid. Other Chalcidoidea recorded as reared from hesperiid hosts are mostly species of Brachymeria (Chalcididae) and Elasmus (Eulophidae), which are primary parasitoids, and Catolaccus (Pteromalidae) which are pseudohyperparasitoids attacking cocooned ichneumonoid primary parasitoids. Species of Pteromalus, Baryscapus and Tetrastichus are also reported as secondary parasitoids from hesperiid hosts.

Genomic DNA sequencing reveals two new North American species of Staphylus (Hesperiidae: Pyrginae: Carcharodini).

Two new skipper butterfly (Hesperiidae) species are described from the United States: Staphylus floridus Grishin, sp. n. (type locality in Florida, Volusia Co.) and Staphylus ecos Grishin, sp. n. (type locality in Texas, Brewster Co.). They are cryptic and hence escaped recognition. They differ from their sister species by the relative size and morphology of genitalia and by genotype-including and beyond the COI barcode-thus suggesting genetic isolation that argues for their species-level status. A lectotype is designated for Helias ascalaphus Staudinger, 1876. Staphylus opites (Godman & Salvin, 1896), stat. rest. is a species-level taxon and not a synonym of Staphylus vincula (Plötz, 1886), while Pholisora iguala Williams & Bell, 1940, syn. n. is a junior subjective synonym of S. vincula.

Explosive Cenozoic radiation and diversity-dependent diversification dynamics shaped the evolution of Australian skipper butterflies

AbstractAustralia was predominantly tropical for most of the Early Cenozoic, then transitioned to a cooler and drier climate in the Oligocene. In response to this increasing aridity, some lineages either adapted to xeric ecosystems, contracted to increasingly fragmented mesic refugia, or went extinct. However, the lack of macroevolutionary studies at a continental scale precludes a better understanding of Australian biodiversity patterns and processes during the Cenozoic. Here, we infer a robust, dated phylogenomic tree for a radiation of Australian endemic butterflies, the Trapezitinae skippers, to test the impact of biotic and abiotic drivers on Cenozoic diversification dynamics in Australia. These butterflies originated during the Eocene (c. 42 Mya) in the mesic biome of Australia. Trapezitinae exploded in diversity during a cool, dry period in the Late Oligocene and Early Miocene, then experienced a sharp deceleration in speciation. Xeric ecosystems appear to have been colonized more recently, supporting the hypothesis of arid and semi-arid biomes as evolutionary sinks. Temperature-dependent and phytophagy-dependent diversification models received little support. Instead, we find evidence for diversity-dependent processes with declining diversification in Trapezitinae likely linked to limited ecological opportunities following a rapid initial burst of diversification.

Pollination and Floral Biology of a Rare Morning Glory Species Endemic to Thailand, Argyreia siamensis.

Argyreia siamensis is extremely rare, and very little is known about its reproduction. The species has colorful flowers that seem likely to attract pollinators, but population sizes are typically small (<30 individuals). To determine whether poor reproduction contributes to its rarity, we investigated its mating system and potential pollinators in two populations. We also examined the staminal trichomes and floral nectary to investigate their role in pollinator attraction. The mating system was assessed with a bagging experiment and pollinator visits were recorded with action cameras. Additionally, we tested the staminal trichomes and floral nectary for terpenes and flavonoids and examined floral nectary micromorphology via scanning electron microscope and compound light microscope. Our results reveal that A. siamensis is self-incompatible and dependent on pollinators; the western population was pollinated by bees (Meliponini and Amegilla), while the eastern population was mainly pollinated by skipper butterflies (Hesperiidae). Both staminal trichomes and the floral nectary appear to contribute to pollinator attraction through the presence of terpenes and flavonoids (in both secretory structures) and nectariferous tissue and nectarostomata (in the nectary). Our results indicate that A. siamensis has reliable and effective pollinators and that insufficient pollination is likely not a primary cause of its rarity.

The omission of critical data in the pursuit of ‘revolutionary’ methods to accelerate the description of species

The omission of critical data in the pursuit of ‘revolutionary’ methods to accelerate the description of species

A genomic perspective on the taxonomy of the subtribe Carcharodina (Lepidoptera: Hesperiidae: Carcharodini).

We obtained whole genome shotgun sequences and phylogenetically analyzed protein-coding regions of representative skipper butterflies from the genus Carcharodus Hübner, [1819] and its close relatives. Type species of all available genus-group names were sequenced. We find that species attributed to four exclusively Old World genera (Spialia Swinhoe, 1912, Gomalia Moore, 1879, Carcharodus Hübner, [1819] and Muschampia Tutt, 1906) form a monophyletic group that we call a subtribe Carcharodina Verity, 1940. In the phylogenetic trees built from various genomic regions, these species form 7 (not 4) groups that we treat as genera. We find that Muschampia Tutt, 1906 is not monophyletic, and the 5th group is formed by currently monotypic genus Favria Tutt, 1906 new status (type species Hesperia cribrellum Eversmann, 1841), which is sister to Gomalia. The 6th and 7th groups are composed of mostly African species presently placed in Spialia. These groups do not have names and are described here as Ernsta Grishin, gen. n. (type species Pyrgus colotes Druce, 1875) and Agyllia Grishin, gen. n. (type species Pyrgus agylla Trimen, 1889). Two subgroups are recognized in Ernsta: the nominal subgenus and a new one: Delaga Grishin, subgen. n. (type species Pyrgus delagoae Trimen, 1898). Next, we observe that Carcharodus is not monophyletic, and species formerly placed in subgenera Reverdinus Ragusa, 1919 and Lavatheria Verity, 1940 are here transferred to Muschampia. Furthermore, due to differences in male genitalia or DNA sequences, we reinstate Gomalia albofasciata Moore, 1879 and Gomalia jeanneli (Picard, 1949) as species, not subspecies or synonyms of Gomalia elma (Trimen, 1862), and Spialia bifida (Higgins, 1924) as a species, not subspecies of Spialia zebra (Butler, 1888). Sequencing of the type specimens reveals 2.2–3.2% difference in COI barcodes, the evidence that combined with wing pattern differences suggests a new status of a species for Spialia lugens (Staudinger, 1886) and Spialia carnea (Reverdin, 1927), formerly subspecies of Spialia orbifer (Hübner, [1823]).

Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae).

We obtained and analyzed whole genome data for more than 160 representatives of skipper butterflies (family Hesperiidae) from all known subfamilies, tribes and most distinctive genera. We found that two genera, Katreus Watson, 1893 and Ortholexis Karsch, 1895, which are sisters, are well-separated from all other major phylogenetic lineages and originate near the base of the Hesperiidae tree, prior to the origin of some subfamilies. Due to this ancient origin compared to other subfamilies, this group is described as Katreinae Grishin, subfam. n. DNA sequencing of primary type specimens reveals that Ortholexismelichroptera Karsch, 1895 is not a female of Ortholexisholocausta Mabille, 1891, but instead a female of Ortholexisdimidia Holland, 1896. This finding establishes O.dimidia as a junior subjective synonym of O.melichroptera. Furthermore, we see that Chamunda Evans, 1949 does not originate within Pyrginae Burmeister, 1878, but, unexpectedly, forms an ancient lineage of its own at the subfamily rank: Chamundinae Grishin, subfam. n. Finally, a group of two sister genera, Barca de Nicéville, 1902 and Apostictopterus Leech, [1893], originates around the time Hesperiinae Latreille, 1809 have split from their sister clade. A new subfamily Barcinae Grishin, subfam. n. sets them apart from all other Hesperiidae.

Genomes of skipper butterflies reveal extensive convergence of wing patterns

For centuries, biologists have used phenotypes to infer evolution. For decades, a handful of gene markers have given us a glimpse of the genotype to combine with phenotypic traits. Today, we can sequence entire genomes from hundreds of species and gain yet closer scrutiny. To illustrate the power of genomics, we have chosen skipper butterflies (Hesperiidae). The genomes of 250 representative species of skippers reveal rampant inconsistencies between their current classification and a genome-based phylogeny. We use a dated genomic tree to define tribes (six new) and subtribes (six new), to overhaul genera (nine new) and subgenera (three new), and to display convergence in wing patterns that fooled researchers for decades. We find that many skippers with similar appearance are distantly related, and several skippers with distinct morphology are close relatives. These conclusions are strongly supported by different genomic regions and are consistent with some morphological traits. Our work is a forerunner to genomic biology shaping biodiversity research.

An urban wildlife habitat experiment: conservation implications of altering management regimes on animals and plants along urban and rural rights-of-way

Abstract Biodiversity along rights-of-way (ROWs) can decline due to local-scale management, including frequent urban mowing and spraying, or with increasing amount of urban infrastructure surrounding those grassy spaces. Distinguishing effects of mowing regime from effects of surrounding urban land are necessary to determine what management strategies effectively conserve wildlife in different landscapes, and to justify changes in management that could increase populations of weeds. We used a manipulative Before/After-Control/Impact (BACI) experiment in 17 transmission line ROWs during 2007–2009, along an urbanization gradient, to disentangle effects of mowing frequency and the amount of urban land (buildings, hard surfaces like concrete, asphalt) surrounding ROWs. In the BACI study, we halted mowing and spraying for 1 year in five urban ROWs, introduced two rounds of mowing per year in three rural ROWs, and compared vegetation and arthropods found within these manipulated ROWs and within three urban and six rural control ROWs. European skipper butterflies Thymelicus lineola, lepidopteran biomass in herbaceous vegetation, milkweeds (Asclepias spp.) and Canada thistle Cirsium arvense increased when mowing and spraying were halted for one year. Conversely, monarch butterflies Danaus plexippus, legume cover and dandelion Taraxacum officinale increased when mowing was introduced to rural ROWs. To increase taller butterfly resource plants while still controlling weeds within urban ROWs, we recommend reducing management frequency within the interior of ROWs while maintaining frequent management along ROW borders that are adjacent to infrastructure.

DNA Barcoding and Taxonomic Challenges in Describing New Putative Species: Examples from Sootywing and Cloudywing Butterflies (Lepidoptera: Hesperiidae)

DNA barcoding has resulted in the ‘discovery’ of a vast number of new species and subspecies. Assigning formal scientific names to these taxa remains a major challenge. Names sometimes are newly designated. Alternatively, available valid names can be resurrected from synonymy, based on barcode analyses together with classical taxonomic characters. For the most part, however, new putative species revealed by barcoding studies go undescribed. This situation is most often attributed to insufficient taxonomic expertise with the authors conducting the study, together with a critical lack of formally trained taxonomists. However, even with formal training, and additional supportive data from morphological, ecological or life history characters, other factors can arise that impede new species descriptions. In the present paper, several specific taxonomic challenges that have arisen from barcode analyses in two groups of skipper butterflies (Lepidoptera: Hesperiidae), the Sootywings (Pholisora catullus and P. mejicanus) and the Coyote Cloudywing (Achalarus toxeus) are highlighted and discussed. Both P. catullus and A. toxeus show relatively large intraspecific genetic divergences of barcodes (2–3%) which suggests the possibility of previously unrecognized cryptic speciation within each group. Some of the challenges to providing formal names and clarifying taxonomic status of these cryptic taxa could be largely overcome by (1) barcoding type specimens, (2) clarifying imprecise and often vague or suspect type localities, and (3) by conducting in-depth comparative studies on genitalic morphology.

Hostplant change and paleoclimatic events explain diversification shifts in skipper butterflies (Family: Hesperiidae)

BackgroundSkippers (Family: Hesperiidae) are a large group of butterflies with ca. 4000 species under 567 genera. The lack of a time-calibrated higher-level phylogeny of the group has precluded understanding of its evolutionary past. We here use a 10-gene dataset to reconstruct the most comprehensive time-calibrated phylogeny of the group, and explore factors that affected the diversification of these butterflies.ResultsAncestral state reconstructions show that the early hesperiid lineages utilized dicots as larval hostplants. The ability to feed on monocots evolved once at the K-Pg boundary (ca. 65 million years ago (Mya)), and allowed monocot-feeders to diversify much faster on average than dicot-feeders. The increased diversification rate of the monocot-feeding clade is specifically attributed to rate shifts in two of its descendant lineages. The first rate shift, a four-fold increase compared to background rates, happened ca. 50 Mya, soon after the Paleocene-Eocene thermal maximum, in a lineage of the subfamily Hesperiinae that mostly fed on forest monocots. The second rate shift happened ca. 40 Mya in a grass-feeding lineage of Hesperiinae when open-habitat grasslands appeared in the Neotropics owing to gradual cooling of the atmospheric temperature.ConclusionsThe evolution of monocot feeding strongly influenced diversification of skippers. We hypothesize that although monocot feeding was an intrinsic trait that allowed exploration of novel niches, the lack of extensive availability of monocots comprised an extrinsic limitation for niche exploration. The shifts in diversification rate coincided with paleoclimatic events during which grasses and forest monocots were diversified.

European Skipper Butterfly (&lt;i&gt;Thymelicus lineola&lt;/i&gt;) Associated with Reduced Seed Development of Showy Lady’s-slipper Orchid (&lt;i&gt;Cypripedium reginae&lt;/i&gt;)

It has been suggested that European Skipper butterflies (Thymelicus lineola) trapped in the lips of the Showy Lady’s-slipper orchid (Cypripedium reginae) may interfere with pollination. This could occur through blockage of the pollinator pathway, facilitation of pollinator escape without pollination, and/or disturbance of the normal pollinators. A large population of the orchid at an Ottawa Valley site provided an opportunity to test the interference hypothesis. The number of trapped skippers was compared in 475 post-blooming flowers with regard to capsule development and thus seed development. The presence of any skippers within flowers was associated with reduced capsule development (P = 0.0075), and the probability of capsule development was found to decrease with increasing numbers of skippers (P = 0.0271). The extent of a negative effect will depend on the abundance of the butterflies and the coincidence of flowering time and other factors. Counts of skippers trapped in flowers were found to follow closely a negative binomial distribution (P = 0.8656).