Heterospecific courtship,minority effects and niche separation between cryptic butterfly species

Species interacting in varied ecological conditions often evolve in different directions in different local populations. The butterflies of the cryptic Leptidea complex are sympatrically distributed in different combinations across their Eurasian range. Interestingly, the same species is a habitat generalist in some regions and a habitat specialist in others, where a sibling species has the habitat generalist role. Previous studies suggest that this geographically variable niche divergence is generated by local processes in different contact zones. By varying the absolute and relative densities of Leptidea sinapis and Leptidea juvernica in large outdoor cages, we show that female mating success is unaffected by conspecific density, but strongly negatively affected by the density of the other species. Whereas 80% of the females mated when a conspecific couple was alone in a cage, less than 10% mated when the single couple shared the cage with five pairs of the other species. The heterospecific courtships can thus affect the population fitness, and for the species in the local minority, the suitability of a habitat is likely to depend on the presence or absence of the locally interacting species. If the local relative abundance of the different species depends on the colonization order, priority effects might determine the ecological roles of interacting species in this system.     

Defense against Pieris rapae in cabbage plants induced by Bemisia tabaci biotype B

Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) and Pieris rapae L. (Lepidoptera: Pieridae) are serious pests of vegetables, that can occur simultaneously on a single cabbage plant [Brassica oleracea var. capitata L. (Brassicaceae)]. We determined whether pre‐feeding or infestation by B. tabaci on cabbage could induce physiological and biochemical responses of the plant against P. rapae. Developmental time, length, and weight of immature P. rapae, and defense‐related plant compounds (SOD, superoxide dismutase; POD, peroxidase; CAT, catalase; APX, ascorbate peroxidase) were measured. Development of P. rapae larvae was 21% slower on B. tabaci‐pre‐infested plants than on plants without B. tabaci infestation. When feeding on B. tabaci‐pre‐infested plants, 22% of P. rapae larvae pupated as compared with 83% on B. tabaci‐free plants. Weights of P. rapae from first to fourth instars that fed on B. tabaci‐pre‐infested plants were also reduced, whereas those of fifth instars and pupae were not. Similarly, body length of P. rapae from first to fourth instars was affected by B. tabaci pre‐infestation, whereas that of the fifth instars was not. Peroxidase and APX activities of the B. tabaci‐pre‐infested plants increased more than SOD and CAT. Peroxidase and SOD activities of B. tabaci and P. rapae co‐infested plants increased as compared with those of P. rapae‐infested plants; however, CAT and APX activities were not different between B. tabaci‐ and P. rapae‐infested plants. These results showed that B. tabaci infestation had a negative effect on P. rapae when they occurred simultaneously on the same host plant. The implications of the induced plant changes on the herbivore are discussed.     

Phylogeography and population genetics of pine butterflies: Sky islands increase genetic divergence

The sky islands of southeastern Arizona (AZ) mark a major transition zone between tropical and temperate biota and are considered a neglected biodiversity hotspot. Dispersal ability and host plant specificity are thought to impact the history and diversity of insect populations across the sky islands. We aimed to investigate the population structure and phylogeography of two pine‐feeding pierid butterflies, the pine white (Neophasia menapia) and the Mexican pine white (Neophasia terlooii), restricted to these “islands” at this transition zone. Given their dependence on pines as the larval hosts, we hypothesized that habitat connectivity affects population structure and is at least in part responsible for their allopatry. We sampled DNA from freshly collected butterflies from 17 sites in the sky islands and adjacent high‐elevation habitats and sequenced these samples using ddRADSeq. Up to 15,399 SNPs were discovered and analyzed in population genetic and phylogenetic contexts with Stacks and pyRAD pipelines. Low genetic differentiation in N. menapia suggests that it is panmictic. Conversely, there is strong evidence for population structure within N. terlooii. Each sky island likely contains a population of N. terlooii, and clustering is hierarchical, with populations on proximal mountains being more related to each other. The N. menapia habitat, which is largely contiguous, facilitates panmixia, while the N. terlooii habitat, restricted to the higher elevations on each sky island, creates distinct population structure. Phylogenetic results corroborate those from population genetic analyses. The historical climate‐driven fluxes in forest habitat connectivity have implications for understanding the biodiversity of fragmented habitats.     

Effects of Pieris host species on life history parameters in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

Host specificity and host selection by insect parasitoids are hypothesized to be correlated with suitability of the hosts for parasitoid development. The present study investigates the correlation between host suitability and earlier studied host-finding behaviour of two closely related braconid larval parasitoid species, the generalist Cotesia glomerata (L.) and the specialist C. rubecula (Marshall) (Hymenoptera: Braconidae). We compared the capability of both parasitoid species to parasitize and develop in three Pieris host species, i.e. P. brassicae (L.), P. rapae (L.) and P. napi (L.) (Lepidoptera: Pieridae). In laboratory experiments, we measured the effect of host species on fitness parameters such as survival, development, sex ratio and size of parasitoid progeny. The results show that C. glomerata is capable of developing in the three host species, with significant differences in parasitoid survival, clutch size and adult weight among Pieris species. The host range for development was more restricted for C. rubecula. Although C. rubecula is physiologically able to develop in P. brassicae larvae, parasitoid fitness is negatively affected by this host species, compared to its most regular host, P. rapae. A comparison of the present data on host suitability with earlier studies on host-searching behaviour suggests that the host-foraging behaviour of both parasitoid species not only leads to selection of the most suitable host species for parasitoid development, but also plays a significant role in shaping parasitoid host range.     

Phylogenetic systematics of Colotis and associated genera (Lepidoptera: Pieridae): evolutionary and taxonomic implications

We investigated the genetic diversity and phylogenetic placement of the butterflies in the genus Colotis and eight related pierid genera using sequence information from two mitochondrial and two nuclear genes. To establish the status of species, we initially barcoded 632 specimens representative of all genera and most species and subspecies in those genera. A subset was then selected for phylogenetic analysis where additional gene regions were sequenced: 16S rRNA (523 bp), EF‐1α (1126 bp) and wg (404 bp). DNA barcode results were largely congruent with the traditional classification of species in the Colotis group, but deep splits or lack of genetic divergence in some cases supported either species‐level differentiation or synonymy. Despite using information from four genes, the deeper nodes in our phylogeny were not strongly supported, and monophyly of the ‘Colotis group’ and the genera Colotis and Eronia could not be established. To preserve the monophyly of Colotis, we revive the genus Teracolus for three outlying species previously in Colotis (i.e. Colotis eris, Colotis subfasciatus and Colotis agoye), as well as the genus Afrodryas for Eronia leda. The position of Calopieris is unresolved although it appears to be well outside the molecular variation in Colotis (s.l.). A dispersal/vicariance analysis suggested that major diversification in Colotis (s.str.) occurred in Africa with subsequent dispersal to India and Madagascar.     

Butterfly community ecology: the influences of habitat type,weather patterns,and dominant species in a temperate ecosystem

We compared variation in butterfly communities across 3 years at six different habitats in a temperate ecosystem near Boulder, Colorado, USA. These habitats were classified by the local Open Space consortium as Grasslands, Tallgrass, Foothills Grasslands, Foothills Riparian, Plains Riparian, and Montane Woodland. Rainfall and temperature varied considerably during these years. We surveyed butterflies using the Pollard‐Yates method of invertebrate sampling and compared abundance, species richness, and diversity across habitats and years. Communities were most influenced by habitat, with all three quantitative measures varying significantly across habitats but only two measures showing variation across years. Among habitats, butterfly abundance was higher in Plains Riparian sites than in Montane Woodland or Grassland sites, though diversity was lowest in Plains Riparian areas. Butterfly species richness was higher in Foothills Riparian sites than it was in all but one other habitat (Tallgrass). Among years, butterfly abundance and species richness were lower during the year of least rainfall and highest temperatures, suggesting a substantial impact of the hot, dry conditions. Across habitats and years, butterfly abundance was consistently high at Plains Riparian and Foothills Riparian sites, and richness and diversity were consistently high in Foothills Riparian areas. These two habitats may be highly suitable for butterflies in this ecosystem, regardless of weather conditions. Generally low abundance and species richness in Montane Woodlands sites, particularly in 2002, suggested low suitability of the habitat to butterflies in this ecosystem, and this may be especially important during drought‐like conditions. Finally, to examine the effect that the presence of the very abundant non‐native species Pieris rapae L. (Lepidoptera: Pieridae) has on these communities, we re‐analyzed the data in the absence of this species. Excluding P. rapae dramatically reduced variation of both butterfly abundance and diversity across habitats, highlighting the importance of considering community membership in analyses like ours.     

Effects of Neem EC on gas exchange, tracheal ventilation, and water loss in diapausing pupae of Pieris brassicae

Effects of Neem EC (The Indian Neem Tree Company^TM, 1% azadirachtin) on gas exchange cycles, tracheal ventilation, and water loss in diapausing pupae of the large white butterfly, Pieris brassicae L. (Lepidoptera: Pieridae), were studied using a constant volume respirometer combined with an infrared probe actograph. The non‐treated pupae displayed discontinuous gas exchange cycles (DGC) with a trend coinciding with the bursts of carbon dioxide (CO₂) release, active tracheal ventilation, and the heartbeat periods. Two independent forms of tracheal ventilation were observed, relatively vigorous abdominal shaking movements and weak abdominal pulsations. The ability to respond to mechanical excitation with abdominal movements was entirely lost on the 2nd day after treatments with Neem EC, and also a reduced tendency to use a DGC was observed. During 2–3 days after treatments, the DGCs and gas exchange microcycles were entirely lost, as was active ventilation. Before treatments, body mass loss, that is, water loss, was 0.6–0.9 mg g^?1 day^?1. After the treatments, water loss increased to 3–5 mg g^?1 day^?1. The pupae remained alive for 10–15 days after the treatments and died after having lost about 50% of their initial body mass. The absence of heartbeats measured during at least 4–5 h was the main criterion for ascertaining death of pupae. The results suggested that respiratory failures, that is, the loss of cyclic gas exchange, evoked by Neem EC were the primary cause of lethal desiccation. Thus, the hypothesis that the cyclic gas exchange is an adaptation for restricting water losses in insects was supported.     

中国蝴蝶新种, 新亚种及新记录种（Ⅲ）（鳞翅目）

记述中国粉蝶科1新种,4新亚种,1新型,1中国新记录种及6个《中国蝶类志》未记述的已知种,所有模式标本保存在西北农林科技大学昆虫博物馆。     

Receptor based models for spontaneous colour choices in flies and butterflies

Colour is one of several stimuli used by herbivorous insects in host choice. Insects have between 2 and 5 different types of photoreceptors to catch quanta of different wavelengths of the spectrum. Many insects have been shown to possess opponent neural interactions between the receptors that enable them to see colour. I present simple models to describe colour choices as functions of the receptor quantum catches and linear interactions of the receptor types. Models are applied to data sets obtained from own experiments and from the literature, on Pieris brassicae and P. rapae (Lepidoptera, Pieridae), Papilio aegeus (Papilionidae), Dacus oleae (Diptera, Tephritidae) and Eristalis tenax (Syrphidae). In fruit flies, detection of green fruit is based on an inhibitory interaction between a green-sensitive receptor type and a blue-sensitive receptor type. This might explain the preference many herbivorous insects have for yellow over green stimuli. Pollen feeding in hoverflies might have evolved from yellow pollen being a super-normal stimulus for herbivorous insects. In butterflies, an additional red-receptor is involved in the colour choice for an oviposition substratum and leads to them choosing green and not yellow. The models introduced in this study open new perspective for a physiological understanding of the design of visual stimuli for monitoring and trapping pest insects.     

Influence of visual stimuli on host location in the butterfly, Eurema hecabe

The influence of visual stimuli on female host location was investigated in Eurema hecabe (Lepidoptera: Pieridae). Mated females with no oviposition experience were presented with artificial plant models treated with the methanol extract of the host plant, Lespedeza cuneata (Fabaceae). When models of different colours were presented to females, they landed and deposited eggs predominantly on the yellow-green model. However, the size of the model was not important. When females were presented with simple square models and a model on which small squares were assembled, they predominantly chose the assembly model. This suggests that female discern the pattern which resembles the leaf of their main host plants, which consists of many small leaflets. However, the whole image of a leaf of the host plant was not an essential cue for host location, as females also chose the square with the central space more frequently than that without the space. We suggest that the long contour of a complicated form plays a role in female host location.