Gonepteryx Rhamni Research Articles

Oviposition strategies in Pieridae butterflies and the role of an egg-killing plant trait therein.

Most herbivorous insects are host-plant specialists that evolved detoxification mechanisms to overcome their host plant's toxins. In the evolutionary arms-races between Pieridae butterflies and Brassicaceae plants, some plant species have evolved another defence against the pierids: egg-killing. Underneath the eggs, leaves develop a so-called hypersensitive response (HR)-like cell death. Whether some butterflies have evolved oviposition strategies to counter-adapt against egg-killing remains to be studied. In this study, we assessed the oviposition site location of Pieridae butterflies on their natural host plants. We described the plant tissue on which we located the eggs of the most common Pieridae in the Netherlands: Gonepteryx rhamni, Anthocharis cardamines, Pieris rapae, P. napi, P. brassicae and P. mannii. Additionally, we assessed expression of HR-like cell death in response to the deposited butterfly eggs. We found that both A. cardamines and G. rhamni mainly oviposited on the floral stem and the branch, respectively, and oviposited on host plants from lineages not expected to kill pierid eggs. Accordingly, no HR-like cell death was seen. All Pieris eggs found were located on leaves of their host, the only tissue found to express HR-like cell death. Furthermore, each Pieris species was found to at least occasionally oviposit on Brassica nigra. This was the only plant species in this survey that expressed HR-like cell death in response to the eggs of P. rapae, P. napi and P. brassicae. Our observations demonstrate that HR-like cell death remains an effective defence strategy against these Pieris species and as such did not find evidence for the hypothesized counterstrategies. Surveying certain key species and disentangling the micro-evolution of oviposition strategies within a species would allow us to further investigate potential counter-adaptations that evolved against HR-like cell death. This study provides the basis for further investigation of potential counter-adaptations to egg-killing defences.

Studies on Insect Pollinators of Vinca major (L.) in Shimla Hills, Western Himalaya, India

Pollinators are essential for sustaining the pollination services, because of variation in pollinator community. Therefore, the present investigation were conducted on taxonomy of insect pollinators of medicinally important plant Vinca major (L.) in Shimla Hills. The collections of insect pollinators were made during flowering season i.e. April-May. During this study period a total 11 insect pollinators were recorded, collected and indentified on Vinca major, and their taxonomic studies were made on the basis of their morphological characters and genitalic attributes. Of these 11 species 08 belonged to sub-order Rhoplaocera (Celastrina huegelii, Celastrina lavendularis, Lampides boeticus, Aglais cashmirensis, Pieris brassicae, Pieris canidia, Gonepteryx rhamni neplensis & Hyarotis adrastus), of Order Lepidoptera and 01 each to Hymenoptera (Xylocopa sp.), Coleoptera (Coccinella septumpunctata) and Diptera (Macroglossum pyrrhosticta).

The complete mitochondrial genomes of Aporia crataegi, Gonepteryx rhamni, and Appias remedios (Lepidoptera, Pieridae) and phylogenetic relationship of other Pieridae species.

Pieridae is a butterfly family whose higher classification has not yet been settled. In this research, we used completed mitochondrial genome sequence to identify and analyze the relationship of three subfamilies of Pieridae (Dismorphiinae, Coliadinae and Pierinae). We have sequenced the complete mitogenomes of Aporia crataegi, Gonepteryx rhamni, and Appias remedios, which are 15,148 bp, 15,118 bp and 15,223 bp in length, respectively. These three mitogenomes have a typical set of 37 genes and an A+T-rich region. The protein-coding genes, excluding the cox1 gene, in the three mitogenomes are initiated by the typical ATN codons. The putative initiation codon for the cox1 gene in the three species is CGA. tRNA genes in the three Pieridae mitogenomes show typical clover leaf structures, except tRNASer(AGN) which lacks the dihydrouridine (DHU) stem. The length of A+T-rich region varies in the three species, which leads to the variation in the mitochondrial genome size. The characterizations of the three mitogenomes enrich our knowledge on the Lepidopteran mitogenome and provide us genetic information to reconstruct the phylogenetic tree. Finally, the phylogenetic analyses using mitogenome data showed that the relationship among three subfamilies of Pieridae is (Dismorphiinae + (Coliadinae + Pierinae)).

Environmental effects on the shape variation of male ultraviolet patterns in the Brimstone butterfly (Gonepteryx rhamni, Pieridae, Lepidoptera).

The males of the Brimstone butterfly (Gonepteryx rhamni) have ultraviolet pattern on the dorsal surfaces of their wings. Using geometric morphometrics, we have analysed correlations between environmental variables (climate, productivity) and shape variability of the ultraviolet pattern and the forewing in 110 male specimens of G. rhamni collected in the Palaearctic zone. To start with, we subjected the environmental variables to principal component analysis (PCA). The first PCA axis (precipitation, temperature, latitude) significantly correlated with shape variation of the ultraviolet patterns across the Palaearctic. Additionally, we have performed two-block partial least squares (PLS) analysis to assess co-variation between intraspecific shape variation and the variation of 11 environmental variables. The first PLS axis explained 93% of variability and represented the effect of precipitation, temperature and latitude. Along this axis, we observed a systematic increase in the relative area of ultraviolet colouration with increasing temperature and precipitation and decreasing latitude. We conclude that the shape variation of ultraviolet patterns on the forewings of male Brimstones is correlated with large-scale environmental factors.

The complete mitochondrial genome of Gonepteryx rhamni (Lepidoptera: Pieridae: Coliadinae)

The complete mitochondrial genome (mitogenome) of Linnaeus Gonepteryx rhamni (Lepidoptera: Pieridae: Coliadinae) is a circular molecule of 15,023 bp in length, containing 37 typical coding genes and one non-coding AT-rich region. Its gene order and content are identical to the common type found in most insect mitogenomes. All protein coding genes (PCGs) start with a typical ATN initiation codon, except for the cox1, which use CGA as its start codon. Nine genes use standard complete termination codon (TAA), whereas the cox1, cox2, nad4 and nad5 genes end with single T. Except for trnS1(AGN), all tRNA genes display typical secondary cloverleaf structures as those of other insects. Additionally, the 371 bp long AT-rich region contains a few structures common to the other lepidopterons, such as the motif ATAGA followed by a 17 bp poly-T stretch, a microsatellite-like (AT)10 element preceded by the ATTTA motif, and a 10 bp poly-A presented immediately upstream of trnM gene.

Butterfly species identification by branch length similarity entropy

We previously developed a shape recognition methodology that uses “branch length similarity” (BLS) entropy, which is defined as a simple branching network consisting of a single node and branches. The simple network is referred to as a “unit branching network” (UBN). Our approach involves obtaining BLS entropy profiles from UBNs created by joining each pixel in the outline of a shape with every other pixel in the shape's border. The profiles successfully characterize the shapes by comparing their BLS entropy profiles. Presently, we modified this approach to facilitate its application to butterfly species identification by partitioning and weighting the entropy profile. As a test, we identified the butterfly species Colias erate, Parnassius bremeri, Eurema hecabe, Gonepteryx rhamni, and Papilio maackii. Each species group consisted of 10 specimens. We used wing shape to identify a species. We extracted evenly spaced x–y coordinates of boundary pixels for the wing shapes in a counter counterclockwise direction. The number of the pixels was 749. We then sequentially partitioned 749 x–y pairs into 15 groups, calculated entropy profiles for the groups, and weighted the profiles. The profiles were combined in order, resulting in a single weighted BLS entropy profile for a wing's shape. Subsequently, we statistically compared the correlation coefficient among the weighted BLS profiles. Our experimental results showed that this method was statistically successful for butterfly species identification. The advantage of the partitioning and weighting process in shape recognition is also discussed.

Declines in Common, Widespread Butterflies in a Landscape under Intense Human Use

Analyses of species' population losses typically show a dichotomy between strongly affected, rare, and localized species and apparently unaffected, common, and widespread species. We analyzed 16 years (1992-2007) of butterfly transect count data from The Netherlands in a reevaluation of the trends of common, widespread species. Fifty-five percent (11 of 20 species) of these species suffered severe declines in distribution and abundance. Overall, cumulative butterfly abundance declined by around 30%. Some of the species in decline used to be omnipresent in gardens and parks, and 2 of the species were previously considered agricultural pests. Based on their declines over the last 16 years, 2 of the 20 species (Lasiommata megera and Gonepteryx rhamni) reached endangered status in The Netherlands under the IUCN (International Union for Conservation of Nature) population-decline criterion, and 2 species (Inachis io and Thymelicus lineola) met vulnerable criterion. Butterflies in farmland, urban, and particularly woodland areas showed the largest decline in species abundance. The abundance of species associated with vegetation types found mainly in nature reserves (dunes, heathland, and, to a lesser extent, seminatural grassland) increased or remained stable. The decline of widespread species requires additional conservation strategies in the wider landscape.

Antennal responses to floral scents in the butterflies Inachis io , Aglais urticae (Nymphalidae), and Gonepteryx rhamni (Pieridae)

To better understand the biological role of floral scents for butterflies, electrophysiological responses to floral scents were investigated using combined gas chromatography and electroantennographic detection (GC-EAD). The antennal responses of three butterfly species, Aglais urticae L. (Nymphalidae), Inachis io L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae) to floral scent compounds from both natural and synthetic mixtures were examined. Floral scents were collected from the butterfly nectar plants Cirsium arvense (L.) (Asteraceae), and Buddleja davidii Franchet cv. (Loganicaeae) with dynamic head-space methods on Tenax-GR and eluted with pentane. These eluates, composed of natural floral scent blends, represent an array of compounds in their natural state. In the GC-EAD analyses eleven compounds were identified from C. arvense with the benzenoid compound phenylacetaldehyde in highest abundance. Seventeen compounds were identified from B. davidii with the irregular terpene oxoisophorone in highest abundance. Thirty-nine synthetic floral scent compounds were mixed in pentane, in equal amounts; about 35 ng were allowed to reach the antennae. The butterflies showed antennal responses to most of the floral scent compounds from both natural and synthetic blends except to the highly volatile monoterpene alkenes. Certain benzenoid compounds such as phenylacetaldehyde, monoterpenes such as linalool, and irregular terpenes such as oxoisophorone, were emitted in relatively large amounts from C. arvense and B. davidii, and elicited the strongest antennal responses. These compounds also elicited strong antennal responses when present in the synthetic scent blends. Thus, the butterflies seem to have many and /or sensitive antennal receptors for these compounds, which points to their biological importance. Moreover, these compounds are exclusively of floral scent origin. For B. davidii, which depends highly on butterflies for pollination, the exclusive floral scent compounds emitted in high abundance could be the result of an adaptive pressure to attract butterflies.

Foraging responses in the butterflies Inachis io , Aglais urticae (Nymphalidae), and Gonepteryx rhamni (Pieridae) to floral scents

For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants.

Marginal range expansion in a host‐limited butterfly species Gonepteryx rhamni

Summary1. The British distribution of the butterfly Gonepteryx rhamni (L.) follows closely the range of its natural host plants, Rhamnus catharticus L. and Frangula alnus Miller, suggesting that it is one of the few British butterflies that has a host‐limited distribution. In North Wales, this species has its range margin, and it was recorded only occasionally in a 35‐km2 area prior to the 1980s. Frangula alnus bushes were planted in the area in about 1986, allowing the hypothesis that G. rhamni would expand its range following increased host plant availability to be tested.2. From 1996 to 1998, the distribution of the butterfly and its host plants, R. catharticus (native), Rhamnus alaternus L. (introduced), and F. alnus (introduced to the area but native to Britain), was mapped in the study area. It was found that the butterfly was more widespread than any of its host plants. Frangula alnus was the most widespread of the host plants, and received most eggs, suggesting that the carrying capacity of the habitat would have increased substantially following the planting of this species. Gonepteryx rhamni was able to complete its lifecycle on both introduced species in the study area.3. A mark–release–recapture study showed that adult G. rhamni moved an average of 512 m, and 50% of movements were further than 400 m; these values are underestimates. The relatively high mobility of this species suggests that it probably perceives host plants and nectar sources as resource patches (patchy population) in this fragmented landscape, and this population now represents a satellite population of the butterfly's main distribution in Britain.4. The results presented here confirm empirically the host‐limited distribution of G. rhamni, which expanded following the planting of extra host plants.

Early Male Emergence and Reproductive Phenology of the Adult Overwintering Butterfly Gonepteryx rhamni in Sweden

In insects males usually emerge before females. This is known as protandry and can be understood in terms of sexual selection acting on males to maximise the number of matings, or on females to minimise the prereproductive period. A corollary of protandry theory is that males should not emerge before females when the advantage of protandry is nullified, as is the case when diapause intervenes between adult eclosion and the mating season. The brimstone butterfly, Gonepteryx rhamni, has such a life cycle, and in Sweden it develops from egg to adult during April/May to July, and overwinters in the adult stage before the onset of the mating season in April the following year. The first objective of this paper was to test the predictions that (1) there should be no difference between the sexes in egg to adult development time, and that (2) males should emerge from hibernation earlier than females, i.e. protandry should be expected during the mating season. A second objective was to test what proximate mechanisms were responsible for the eventual sex difference in time of emergence from hibernation. Protandry theory shows that late emerging males are heavily penalized in terms of mating opportunities, and so male G. rhamni may benefit from being able to mate directly after emerging from hibernation, whereas females may not be in such a hurry to start laying eggs because the bud break of the host plant Frangula alnus occurs in late May. Hence male G. rhanmi may have to allocate their fat reserves both to winter survival and sperm production, whereas females can most likely allocate all their fat reserves to winter survival, and so the third objective was to document the reproductive phenology of males and females and to test if male G. rhamni suffered higher winter mortality than females. As expected from theory there was no sex difference in egg to adult development time, whereas three years of census data from the field showed that the first males emerged from hibernation on average 21 d earlier than the first females (with the peak of male abundance occurring on average 8 d before that of female peak abundance). The earlier male emergence from hibernation seemed to be the result of the combined effects of males having a slightly lower temperature threshold for flight, and males being ready to fly as soon as environmental conditions were suitable after 4 months of hibernation, whereas females seemed to require a higher temperature summation before being ready to fly. Dissections of males and females throughout the year showed that male sperm production started soon after eclosion with males entering hibernation with a substantial sperm complement, and experiments showed that both males and females mated within minutes after having spent at least 6 months in hibernation at 4°C. By contrast, female eggs were small and unyolked throughout winter, and egg development started only after females had emerged from hibernation. Females brought from hibernation, and mated, towards the end of March started to lay eggs only after spending between 7 and 14 d at room temperature. Although males divide their fat reserves between survival and reproduction during winter, there was no difference in winter survival between the sexes under laboratory conditions, and the sex ratio assessed in the field did not differ before versus after hibernation giving no support for the idea that males suffer higher winter mortality than females.

Thermal characterization of butterfly wings—1. Absorption in relation to different color, surface structure and basking type

1. 1. Spectral integral reflectance, transmittance and the resulting absorption of intact and descaled butterfly wings of the black-winged Pachliopta aristolochiae (Papilionidae), the white-winged Pieris brassicae (Pieridae), and the yellow-winged Gonepteryx rhamni (Pieridae) were determined between 350 and 800 nm. 2. 2. Whereas in the black forewing of the dorsal basking Pachliopta almost all incident light is absorbed nearly independent of the wavelength and thus converted into heat, the white forewing of the body basking Pieris absorbs less than 20% in the visible range of the spectrum. 3. 3. The yellow hindwing of the lateral basking Gonepteryx absorbs to a higher degree than the Pierid wing, but—due to the sparsely arranged scales—transmittance is clearly increased (40–50% between 525 and 800 nm). 4. 4. The varying thermal characteristics of the different wings with reference to the color and arrangement of the scales and the different basking strategies of the butterflies are discussed.

The use of chemical composition as a population marker in insects: a study of the Brimstone butterfly

ABSTRACT. The elemental composition of 249 adult Brimstone butterflies (Gonepteryx rhamni L.) was studied by wavelength dispersive X‐ray fluorescence spectrometry, to determine whether site/population‐specific differences could be detected. The chemical elements studied were K, Ca, Fe, Cu, Zn, Ti, P, CI and S. Marked differences were found in the chemical composition of the two sexes, and between individuals caught on different sites and in different years and seasons. Multivariate analyses of the data using principal component analysis and canonical variate analysis showed a clear separation of the butterflies by sex, and within each sex, by seasons. Of the elements studied, the main discriminating ones for this butterfly were K, Ca, P, Zn, CI and S. Whilst differences in chemical composition appear to result from variations in the composition of soil and food‐plants in different localities, any site/population‐specific differences are soon masked by the passage of time, as adults age and feed. This makes the technique of limited value in the study of dispersal in this species.

Haemolymph flows in the wings of pierid butterflies visualized by vital staining (insecta, lepidoptera)

The flow of stained haemolymph was photographed in the wings of resting Pieris rapae, Pieris brassicae, and Gonepteryx rhamni under UV-radiation at definite intervals after abdominal application of fluorescent tetracycline. There is no circular route in the wing. All wing veins are supplied with stained haemolymph from their own bases without preference to single veins. In freely resting Pieris with intact wings, most veins are completely stained after 20 min. The staining pattern supports the existence of an oscillating haemolymph supply mechanism in the wing veins and shows that the cross vein and encircling sinus are not essential in the supply of the longitudinal veins. Inflow of stained haemolymph into the wing membrane begins about 1 h after application and is generally completed within 12 h in Pieris. The wing membrane is supplied with fluid by diffusion and — especially under low relative humidity — additionally by haemolymph substitution of evaporated water. This mechanism is associated with the disadvantages of water loss and probably salt withdrawal from the body. The puddling behaviour of butterflies might help in restoring these postulated deficits. It is hypothesized that haemolymph substitution of water evaporated from the wing membrane is a preadaptation for accumulation of defensive toxins and pheromones in the wing membranes, especially in diurnal and basking Lepidoptera. The veinal system of 5-day-old young summer specimens of Gonepteryx stains more intensely than that of 4–5-month-old specimens just before entering hibernation. The transition of stained haemolymph from vein to membrane is reduced in this species, probably as an adaptation for water retention during diapauses.

’Gynandromorphic Effect‘ and the Optical Nature of Hidden Wing-pattern in Gonepteryx rhamni L. (Lepidoptera, Pieridae)

G. A. MAZOKHIN-PORSHNYAKOV1, examining the optical characters of insect wings in connexion with the visual perception of insects, and taking photographs using ultra-violet rays, found that the smooth, yellow surface of the Gonepteryx rhamni wing has a real pattern which is invisible to the human eye. He also found that such a pattern in the full sense is presented in males only. The wing of the female absorbs ultra-violet rays by the entire surface and then looks dark. Mazokhin-Porshnyakov supposed that this hidden pattern may be considered as a character of taxonomic value. This assumption has been partly supported by my work2.

Warning Colours

MY friend Prof. Meldola has drawn my attention to a communication by Mr. F. E. Beddard in NATURE of November 26 (p. 78), in which the view is expressed that the brimstone butterfly (Gonepteryx rhamni) is rendered protected or unpalatable by the yellow pigment of its wings being due to a substance formed as “a urinary pigment,” and that the coloration is “a consequence of the deposition in the integument of bitter pigments.”