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Butterfly nectaring flowers: Butterfly morphology and flower form
Abstract
The profitability of butterfly foraging depends in part on the corolla depth and clustering of flowers, and the tongue length, body mass and wing loading of butterflies. Interactions among these attributes of flowers and butterflies were investigated, using data from a field study in Cornwall and from Porter et al. (1992). The maximum corolla depth from which a butterfly can feed depends on tongue length, which correlates with the more easily measured attributes of body mass and wing loading. Small, short-tongued butterflies did not visit deep flowers. The quantity of nectar sugar per flower necessary for profitable foraging depends on foraging costs, which are expected to correlate with wing loading. Butterfly species with a high wing loading generally confined their visits to flowers that were clustered or very nectar-rich. Butterfly species with a low wing loading included solitary and less nectar-rich flowers in their diet. Body mass and wing loading affect a butterfly's load-carrying capacity (limiting the distance between fuelling stops) and cooling rate (limiting the distance between stops for basking or endothermic warming), and will therefore influence the capacity for floral selectivity and for migration and dispersal. Body mass, wing loading and tongue length characterised families or subfamilies of butterflies. For example vanessine nymphalids, with their long tongues and high wing loading, visited the deep, massed flowers of Buddleja davidii, but lycaenids, with their short tongues and low wing loading, did not. These often visited members of the Asteraceae. Eupatorium cannabinum, with massed flowers offering abundant and accessible nectar, was visited by butterflies of all tongue lengths and both high and low wing loading. These findings may help to inform habitat management for butterfly nectaring flowers.
... Nectar concentration and colour pattern of the flowers are important factors for the pollinators' flower preferences (Watt et al., 1974;Pivnick and McNeil, 1985;May, 1985;Tiple et al., 2006). Although the preferences depend on matching between the flowers' morphological characters such as corolla length (Kingsolver and Daniel, 1979) and butterfly morphology such as proboscis length and wing loading (Porter et al., 1992;Corbet, 2000). Scaling relationships of various body structures compared to body size, as well as adaptive deviations from typical allometric patterns, are prevalent in the animal world (Thompson, 1917;Kunte, 2007). ...
... The multiplicity and benefits of this type of deviation from the allometric growth condition have been well established (Schmidt-Nielson, 1984;Nilsson, 1988;Emlen, 2001). In the case of butterflies and moths, body size function is a strong predictor of proboscis length (Corbet, 2000). Some nectar-sucking species confer incongruently longer probosces in comparison to body size (Kunte, 2007). ...
... In the case of nectar thieves, those with a longer proboscis have a great advantage (Kunte, 2007), because a longer proboscis can suck nectar from deeper and more tubular flowers (May, 1992), which respond with higher-standing nectar crops (Brown et al., 1978;Grant and Grant, 1983;Kodric-Brown et al., 1984;May, 1988;Haber and Frankie, 1989). Many temperate butterfly species with high wing loading restrict their visits to plants by visiting those with clustered flowers, while those with low wing loading prefer solitary flowers (Corbet, 2000). Butterfly species with larger bodies are heavier and require larger wings to support them in flight (Dennis, 1993). ...
Butterfly species are regarded as one of the most important members of the plant-pollinator guild. They mainly feed on nectar, and occasionally they rely on pollen. It was reported that butterfly species collect nectar from a greater array of plants. Emperical studies demonstrate that morphological variables of the butterfly species play an important role in foraging. Four morphological variables and two indices, viz., proboscis length, wing span, body length, and weight, and proboscis index and the wing load index of the butterfly species, were used to check their effect on foraging behavior on two plants, viz., Lantana camara (LCA) and Tridax procumbens (TPR), for the current study. Wing load index emerged as the most sensitive factor for foraging on these two plants. Both plants have the highest rate of visits by the butterfly species, with Lantana camara being more frequently visited than Tridax procumbens (TPR). We can conclude that the information gained from this study may help to conserve and sustain the butterfly community in the wild, and this, in turn, may also help to facilitate conservation strategies for the naturally growing nectaring plant species.
... However, their choice of flowers is not random and do not feed indiscriminately from any flower they find (Tiple et al. 2006). They often exhibit distinct flower preferences that can differ between species (Tudor et al. 2004) and also are known to differ in the flowering species they visit (Corbet 2000, Tudor et al. 2004. Even when several species of flowers are simultaneously in blossoms, most flowervisiting insects utilize not all flowers but specific ones for energy resources (Dobson 1994). ...
... Pollinators use plant cues (floral traits) to distinguish between morphologically identical plants to facilitate exploitation of nectar sources (Chittka and Raine 2006). Among the floral cues patterns of flowers and time of anthesis (Dosa 1999, Sourakov 2012; shape (Gunathilagaraj et al. 1998), size (Waser and Price 1983) and clustering (Corbet 2000) are very important. Pollinators perceive diverse floral signals and cues and modify their foraging behaviour in response, thereby affecting both their own foraging success and the plants' reproductive prospects. ...
... Competition for nectar resources among butterflies clearly occurs (Porter et al. 1992). The assemblage of attributes that influences flower choice also influences other aspects of butterfly biology such as flight costs and rate of energy intake (Corbet 2000). This means that pollinators have to make economic choices about what type of flowers they visit to increase benefits (energy intake) compared to energy consumption, Amaya-Marquez (2009). ...
Foraging behaviour of nine papilionid butterflies have been studied from two moist deciduous sal forests viz. Bhawal and Madhupur from central part of Bangladesh. Papilionid butterflies were found to visit 156 flowering plant species belong to 39 plant families. Dependency of papilionid butterflies on flowering plants varied from family to family with high dependency on family Asteraceae (15%) followed by Rubiaceae (8%), Rutaceae (6%), Verbenaceae (6%), Apocynaceae (6%) and Fabaceae (6%). Although most of the plant species (72 species, 47.68%) were visited by single butterfly species but some were also visited by multiple species of butterflies (e.g., Chromolaena odorata by 9 species, followed by Mikania cordata and Lantana camara by 8 and 7 species, respectively). Ninety eight percent (153 out of 156 species) of the nectar resources were dicotyledonous in origin but two percent plant species (three out of 156 species under three families) were monocotyledonous. Six floral attributes such as plants life forms, nature, perennation, origin, corolla shape and flower massing contributed butterflies significantly to visit plants for nectaring. J. Biodivers. Conserv. Bioresour. Manag. 2020, 6(2): 55-68
... Such butterfly-pollination syndrome (psychophilous) traits can be identified in many plants with wing pollination systems. Psychophilous traits are variable but the traditional description includes red, pink and mauve colours and flat-topped inflorescences or wide, flat petal margins to act as a relatively large landing platform (Faegri and Van der Pijl, 1979;Corbet, 2000;Willmer, 2011). Nectar is the sole reward for butterflypollinated flowers and is often hidden at the base of narrow, long tubes or spurs (Corbet, 2000). ...
... Psychophilous traits are variable but the traditional description includes red, pink and mauve colours and flat-topped inflorescences or wide, flat petal margins to act as a relatively large landing platform (Faegri and Van der Pijl, 1979;Corbet, 2000;Willmer, 2011). Nectar is the sole reward for butterflypollinated flowers and is often hidden at the base of narrow, long tubes or spurs (Corbet, 2000). Flowers frequently have nectar guides and nectar has low volume and low solute concentration (Faegri andVan der Pijl, 1979, Willmer, 2011). ...
... Insect foraging efficiency depends on handling time, time spent travelling between flowers and the cost of moving between sources on a single flower. Inter-flower travel costs are relatively low for massed flowers such as those in compact inflorescences, as these allow foragers to move between flowers by walking, rather than flying (Corbet, 2000). Gloriosa superba produces solitary flowers with a long pedicle and often only a few flowers per plant are produced, with sequential opening of flowers to prevent overlap in receptivity (Gupta and Raina, 2001). ...
Background and aims:
Complex modifications of angiosperm flowers often function for precise pollen placement on pollinators and to promote cross-pollination. We explore the functional significance of the unusually elaborate morphology of Gloriosa superba flowers which are divided into five male and one hermaphrodite meranthia (functional pollination units of a single flower).
Methods:
We used controlled pollination experiments, floral measurements, pollen load analyses and visitor observations in four populations of G. superba in South Africa to determine the breeding system, mechanism of pollination and role of flower orientation in promotion of cross-pollination.
Key results:
We established that G. superba is self-compatible, but reliant on pollinators for seed production. Butterflies, in particular the pierid Eronia cleodora, were the primary pollinators (>90% of visitors). Butterflies brush against the anthers and stigma during nectar feeding and pollen is carried on their ventral wing surfaces. Butterfly scales were positively correlated with the number of pollen grains on stigmas. We demonstrate that the styles were orientated toward clearings in the vegetation and we confirm that the highest proportion of initial visits was to hermaphrodite meranthia pointing towards clearings.
Conclusion:
The flower morphology of G. superba results in effective pollen transfer on the wings of butterfly visitors. The style-bearing hermaphrodite meranthium of the flowers orientates towards open spaces in the vegetation thus increasing the probability that butterflies land first on the hermaphrodite meranthium. This novel aspect of flower orientation is interpreted as a mechanism that promotes cross-pollination.
... Compared with the flower-visiting species, the proboscis of the non-flower-visiting species do not taper as much, and the galeal apex is somewhat rounded. There is a positive correlation between body length and proboscis length both in nectar-feeding and non-nectar feeding butterflies, and nectar-feeders have a tendency to show an allometric relationship [33][34][35][36]. The proboscis length of these ten investigated species is between 3.26-22.84 ...
... mm, accounting for 16%-90% of body length and 10%-90% of forewing length. The profitability of butterfly foraging depends in part on the corolla depth and clustering of flowers and the tongue length, body mass and wing loading of butterflies [35,37,38]. Short-tongued butterflies do not visit flowers with deep corollas, while butterflies with a light wing load generally prefer clustered or nectar-rich flowers [35,39]. ...
... The profitability of butterfly foraging depends in part on the corolla depth and clustering of flowers and the tongue length, body mass and wing loading of butterflies [35,37,38]. Short-tongued butterflies do not visit flowers with deep corollas, while butterflies with a light wing load generally prefer clustered or nectar-rich flowers [35,39]. The morphology of sensilla reflects taxonomic relationships and has been used to infer phylogenetic relationships [36,40,41]. ...
The ultrastructure of the sensilla on the proboscis of ten species of butterflies, Iphiclides podalirius, Parara guttata, Colias fieldii, Celastrina oreas, Sasakia charonda, Tirumala limniace, Acraea issoria, Stichophthalma neumogeni, Callerebia suroia, and Libythea celtis, among five families were investigated using scanning electron microscopy. They were compared to reveal the morphological differences in the proboscis sensilla among these butterflies. Four distinct types of sensilla were found on the proboscis among these species. The types of proboscis sensilla of I. podalirius and T. limniace were sensilla chaetica, sensilla coeloconica, and sensilla basiconica. The types in the other eight species were sensilla chaetica, sensilla styloconica, and sensilla basiconica. The number of sensilla styloconica on the proboscis of non-flower-visiting species was greater than that of flower-visiting species.
... Size matching between floral nectar tube depth and pollinator proboscis length has been shown to be an important determinant of plant-pollinator interactions. In general, flowers with deep nectar tubes restrict access of nectar to visitors with long mouthparts (Haber & Frankie 1989;Corbet 2000;Stang et al. 2007). For specialised and co-dependent species pairs, evolutionary theory predicts that (1) there should be strong natural selection for pollinator proboscides to exceed floral nectar tubes in order to access all the nectar within the flowers; and (2) that selection on flowers should favour nectar tubes that are longer than their pollinator's proboscis because this ensures contact with the reproductive parts of the flowers and thus maximises pollen transfer (e.g. ...
... This finding supports the idea that, in contrast to Hymenoptera, Lepidoptera are mainly opportunistic nectar feeders, using nectar resources as they become available (Shreeve 1992). Other floral traits, such as display or blossom size might also be more important for butterfly flower choice (Corbet 2000). Table 1 Results of the generalised additive mixed model (GAMM) predicting the effect of degree of mismatch, flower head density and flower head display size on interaction probability, fitting a binomial (link=logit) distribution. ...
... Being able to predict plant-pollinator interactions and understand the underlying mechanisms is of great importance for understanding (the structure of) interaction networks. Generally, short-tongued insects are not able to visit deep-tubed flowers (Haber & Frankie 1989;Corbet 2000;Stang et al. 2007) and the absence of these 'forbidden' interactions are often assumed to results in a nested network structure (Stang et al. 2006;Olesen et al. 2011;Ibanez 2012). However, our results indicate that interactions between long-tongued pollinators and short-tubed flowers do not occur often. ...
A long‐standing question in ecology is how species interactions are structured within communities. Although evolutionary theory predicts close size matching between floral nectar tube depth and pollinator proboscis length of interacting species, such size matching has seldom been shown and explained in multispecies assemblages. Here, we investigated the degree of size matching among Asteraceae and their pollinators and its relationship with foraging efficiency. The majority of pollinators, especially Hymenoptera, choose plant species on which they had high foraging efficiencies. When proboscides were shorter than nectar tubes, foraging efficiency rapidly decreased because of increased handling time. When proboscides were longer than nectar tubes, a decreased nectar reward rather than an increased handling time made shallow flowers more inefficient to visit. Altogether, this led to close size matching. Overall, our results show the importance of nectar reward and handling time as drivers of plant–pollinator network structure.
... Vanessa butterflies visiting Buddleja davidii), whereas species with low wing loading and short proboscis more often visit flower species with shallow corollas (e.g. lycaenid species visiting Asteraceae; Corbet, 2000). Wing loading refers to the ratio of wing area to body mass and relates to flight power (Shreeve et al., 2009). ...
... Such a preference is thought to be particularly important as adult compensatory feeding mechanism after poor larval growth conditions (Rusterholz & Erhardt, 2000;Mevi-Sch€ utz & Erhardt, 2003, Erhardt & Mevi-Sch€ utz, 2009). Butterfly preference for nectar with high relative sucrose concentrations has also been demonstrated (Erhardt, 1992;Rusterholz & Erhardt, 1997, 2000. We observed C. jacea to have slightly more nectar per floret than T. pratense. ...
... First, T. pratense may be more difficult to handle. Maniola jurtina has a proboscis of c. 10.5 mm long (Corbet, 2000). The corolla tube of T. pratense is c. 8.6 mm long (Comba et al., 1999), whereas the tube of C. jacea varies considerably between 6 mm (J. ...
Butterflies are often considered as opportunistic nectar consumers that visit a range of flower species. The degree of specialisation in foraging behaviour and flower choice may, however, vary considerably at the inter‐specific level, from highly specialised to generalist species. In generalist nectar use, there can be intra‐specific variation in the availability and use of floral resources (diversity and abundance) among different populations. Knowing the preferences of nectar‐feeding butterflies can increase the understanding of ecological relationships and resource use and help in developing better strategies for butterfly conservation.
By means of behavioural tracking, we identified foraging patterns of Maniola jurtina butterflies in nectar‐rich and nectar‐poor grasslands. For both males and females, we showed significant preference for the brown knapweed Centaurea jacea and thistles ( Cirsium sp.) in nectar‐rich, extensively managed grasslands. In the absence of these preferred species, as is typically the case in nectar‐poor intensively managed sites, M. jurtina nectared on Trifolium pratense and Leucanthemum vulgare . Flower visits were on average shorter on T. pratense compared to C. jacea . These observational results were confirmed by a field experiment in which a wild flower nectar array with C. jacea and T. pratense was introduced.
In extensively managed grasslands, foraging bouts were generally confined to patches of flowering plants and did not involve long flights between flower visits. In intensively managed meadows, butterflies performed longer flights, and ignored more flowers between two consecutive visits. Despite the variety of flower species visited, M. jurtina showed a significant preference for a few nectar sources in a context‐dependent way.
... Butterflies of both species would go to most flowers for nectars. Corbet (2000) and Tiple et al. (2009) pointed out that butterflies can use almost any flowers that can be reached by the proboscis (Corbet, 2000;Tiple et al., 2009 Title section. This includes the title of the paper (all capitalized), author's full name, author's institution and address (all with first letters capitalized), and e-mail address of the corresponding author. ...
... Butterflies of both species would go to most flowers for nectars. Corbet (2000) and Tiple et al. (2009) pointed out that butterflies can use almost any flowers that can be reached by the proboscis (Corbet, 2000;Tiple et al., 2009 Title section. This includes the title of the paper (all capitalized), author's full name, author's institution and address (all with first letters capitalized), and e-mail address of the corresponding author. ...
Troides helena and Pachliopta adamas utilize the same food plant species: Aristolochia acuminata. For the purpose of captive breeding and conservation, it is desirable to find out whether they can co-exist in captivity. Captive breeding research was conducted on the butterfly species within the period of October 2016 to September 2019. In total, 1,361 individuals were observed. Data on adult emergence of the species is presented to show population trends. Both species co-existed poorly at the facility when food plants were limited. It took 45.9 days for T. helena helena and 32.6 days for Pc. adamas adamas from egg to imago stage. Habitat enrichment can encourage the species to come and establish the population.
... For instance, corolla length may alter nectar accessibility and selection by insect species (Wäckers, 2004;Campbell et al., 2012;Balzan et al., 2014). The presence of inflorescences implies clustered resources, which can present advantages in foraging (Corbet, 2000). Reciprocally, insect traits also modify their interaction with plants. ...
... This was in spite that lepidopteran species have long proboscises, which allow them to obtain flower resources from deep corollas. But they also have high wing loads, so inflorescence represents the advantage of clustered foraging resources, and might explain the pattern observed in this study (Corbet, 2000). ...
1 Flower strips can promote and conserve beneficial insects in agroecosystems. Knowingwhich
groups are favoured and which plant traits affect visitation rates by beneficial
insects is important for the design of plant strips.
2 We established 21 Native Flower Strips (NFS) in avocado orchards in Central Chile.
NFS contained 7–11 plant species, with variable corolla length and flowering period,
to promote beneficial insects. We assessed flight activity of ladybirds (Coccinellidae)
and bees (Apiformes) in sites adjacent to and far from NFS within avocado orchards.
Additionally, we evaluated flower visitation for the main flower visitor groups
(Apiformes, Lepidoptera, Diptera, and Coleoptera) to the plant species in NFS
according to season and corolla length.
3 We found almost six times greater flight activity of coccinellids and bees in NFS
than sites far from NFS within avocado orchards. Visitation rates of pollinator groups
varied according to corolla length and season. Diptera, Coleoptera, and Lepidoptera
consistently visited short corolla flowers. NFS were highly visited in summer and
autumn, when avocados were not in bloom.
4 NFS in orchards should be encouraged by policymakers because they support
beneficial insects that could deliver ecosystem services and contribute to local
biological conservation.
... So, corolla depth is a key factor that limits exploitation by nectar feeding butterflies. In case of temperate butterfly species with high wing loading restricts their visits to plants that contain clustered flowers for rich nectar supplies and in case of butterflies with low wing loading chose solitary and less rich nectar plants in their diet [25]. Larger bodied butterflies with higher wing loading require richer, plenty of food and so, possess longer tongues for accessing a wide range of flowering plants. ...
... Compatibility is the fundamental driver in case of plant-butterfly species association. From plant side nectar source attractiveness determined largely by nectar quality, corolla depth, nectar supply and flower clustering [25] and in case of butterfly species access of the host plant for nectar source depend mainly on proboscis length but wing loading is main factor for suitability of exploitation [30]. The foraging efficiency of the butterfly species depends mainly on their handling time, so greater the handling time lower the foraging efficiency [26]. ...
The study of foraging behaviour of butterflies has been an important focus among the lepidopterists for years. Although established as invasive weed plant, the role Lantana camara as an important host plant as well as a food plant for the butterfly species is now well-known. The present study aims to interpret the role of morphological variables of the visitor butterflies in relation to their foraging behaviour on Lantana camara. Butterflies with longer proboscis were more efficient in foraging on flowers of Lantana camara than the butterflies with shorter ones, when the species containing proboscis same as or longer than their body length were excluded. Longer relative proboscis length (RPL) have selective advantage for foraging over Lantana camara up to a certain point (RPL = 0.881) and after that larger relative proboscis length often obstructs foraging behaviour as the handling time increases and these butterflies face more resistance against incoming nectar than the others.The residuals (body weight-wing span) also act as important factor.The handling time of the butterflies on Lantana camara shows a negative relationship with the residuals. The findings suggests that in case of Lantana camara as a nectaring plant butterfly species with longer proboscis have selective advantage for exploitation of the resource than the species with shorter ones and plantation of Lantana camara may be useful for conserving the long tongued butterfly species.
... For this reason, studies of flight aerodynamics focus on the evaluation of parameters such as the capacity of the individual to carry its own body mass and produce sufficient force to enable flight (i.e. wing loading) [2,[8][9][10], the proportion of flight muscle as an estimator of the musculature necessary to support the mass of the individual as well as its capacity for acceleration [8,11] and descriptors of the wing shape such as the aspect ratio [2,11,12]. ...
... In addition, total dry mass was determined in each individual, as well as the mass of the thorax and abdomen (both separately) with no legs or wings, using an analytical balance (Sartorius CP124S) of precision 0.01 g. The total dry body mass and wing area values were used to calculate the wing loading of each individual as an estimator of the carrying capacity of the wing membranes [2,[8][9][10]: ...
Flight morphological variations and its consequences on animal performance are common in winged insects. In the butterfly Heliconius charithonia, sex-related differences in the wing morphological design have been described resulting in differences in foraging behavior, daily flight distances and flight aerodynamics. It has been suggested that these differences should be reflected in the metabolic capacities and energetic budgets associated with flight in both sexes. In this study, we analyzed the relationship between wing morphological variation and metabolic performance, flight aerodynamics and energetic reserves in females and males of Heliconius charithonia over two years. The results confirm the presence of wing shape sexual dimorphism, but also show an unexpected sex-related annual variation in wing shape, mirrored in the metabolic condition (resting metabolic rate) of individuals. However, contrary to expectation, intersexual variations in wing shape are not related to differences between the sexes in terms of flight aerodynamics, flight metabolic rates, or energetic reserves (carbohydrates, lipids and proteins). Our results indicate a considerable plasticity in H. charithonia wing shape, which we suggest is determined by a trade-off between environmental pressures and reproductive restriction of each sex, maintaining an optimum flight design. Finally, similarities in metabolic rates between young and older males and females in both years may be a consequence of the ability of Heliconius species to feed on pollen.
... During bloom, plants have many pink and white flowers which produce large amounts of both pollen and nectar (Gyan & Woodell, 1987b;Fowler et al., 2016); a recent study found that bramble flowers have the fifth highest nectar sugar content per flower per day out of 175 species for which nectar data were available (Baude et al., 2016). The bowl-shaped flower is typical of Rosaceae, with open petals and no corolla tube (Corbet, 2000), so that nectar and pollen are easily accessible to insects with either long or short tongues. Indeed, bramble flowers are visited by many insect groups (Balfour et al., 2015;Baldock et al., 2019) including bees (Goulson et al., 2005;Falk & Lewington, 2015), butterflies (Sparks & Parish, 1995;Corbet, 2000;Tudor et al., 2004), hoverflies (Drabble & Drabble, 1927;Lucas et al., 2018) and non-syrphid Diptera (Drabble & Drabble, 1927). ...
... The bowl-shaped flower is typical of Rosaceae, with open petals and no corolla tube (Corbet, 2000), so that nectar and pollen are easily accessible to insects with either long or short tongues. Indeed, bramble flowers are visited by many insect groups (Balfour et al., 2015;Baldock et al., 2019) including bees (Goulson et al., 2005;Falk & Lewington, 2015), butterflies (Sparks & Parish, 1995;Corbet, 2000;Tudor et al., 2004), hoverflies (Drabble & Drabble, 1927;Lucas et al., 2018) and non-syrphid Diptera (Drabble & Drabble, 1927). Rubus fruticosus has a long flowering period, which typically extends from May to September in the United Kingdom (Streeter et al., 2009;Baude et al., 2016) although flowers can be found into November (V.W. and F.R., personal observations). ...
• Bramble (Rubus fruticosus L. agg.) is a common summer‐flowering plant native to the United Kingdom. Multiple complementary approaches were used to evaluate its ecological value to the honeybee (Apis mellifera ), bumblebees (Bombus spp.) and other flower‐visiting insects in Sussex, England.
• Regional surveys of insect groups at seven sites across 2 years showed that foraging activity on bramble was dominated by honeybees (60.2%; n = 28 surveys) and bumblebees (17.4%), compared to non‐Apis/Bombus bees (2.8%), hoverflies (Syrphidae, 7.9%), non‐syrphid flies (0.6%), butterflies (6.4%), wasps (0.4%) and beetles (4.4%). Foraging insect community structure was highly similar spatially, but varied significantly between survey months (June and July).
• In detailed local surveys at one rural and one urban location, there was a diverse range of insect taxa foraging on the bramble flowers, including species of conservation concern (Bombus humilis , Coenonympha pamphilus and Limenitis camilla ).
• Pollen trapping at 12 honeybee hives in four locations showed that an average of 31% of pollen pellets collected by honeybees from late May to early August were bramble, with a peak of 66–86% per location.
• Bramble was present in 54 out of 60 200 × 200 m randomly selected grid squares surveyed over a large area across Sussex. Plants were recorded in multiple habitat types in both urban and rural areas.
• Bramble is sometimes considered an undesirable plant or a “thug” that outcompetes other wild flowers; however, these findings confirm that it is highly valuable for flower‐visiting insects. Wherever conflicts of interest and management strategies allow, bramble should be maintained and promoted for wildlife and insect conservation.
... nih.gov/ij) on thresholded images. Where possible the right fore and hind wings were measured, summed and doubled for total wing area without considering overlap during flight [48,49] -if a wing was damaged we measured the least damaged alternative wing. Wing loading was calculated as N/m 2 following Corbet [49], where wing load = (body mass(kg) * gravitational field (g = 9.80665 m/s 2 ))/wing area (m 2 ). ...
... Where possible the right fore and hind wings were measured, summed and doubled for total wing area without considering overlap during flight [48,49] -if a wing was damaged we measured the least damaged alternative wing. Wing loading was calculated as N/m 2 following Corbet [49], where wing load = (body mass(kg) * gravitational field (g = 9.80665 m/s 2 ))/wing area (m 2 ). We used both dry weights of intact butterflies and the dry weights of lipidextracted butterflies to calculate intact and lean wing loads. ...
Running title: Monarch alternative migration: We collected 434 adult monarchs and surveyed milkweeds for immature monarchs in southwest Michigan, USA in order to test the hypothesis that monarchs are temporally variable, sequential partial migrants rather than partial migrants that may be spatially separated. Adult size, wing wear, female egg counts, fat content and sequestered chemical defenses were measured in monarchs across an entire season from spring migrant arrival, through breeding, until autumn migrant departure. We predicted that a population characterized by starting from all migrants and no residents, through breeding residents, to all migrants and no residents should show life history measures consistent with changes in these proportions. Results show that female monarch spring migrants arrive with chorionated eggs and high wing loads in both intact and fat-extracted adults. Wing loads of both males and females decrease during the summer and increase again immediately before autumn departure, when the fat content of all adults increases markedly. The high fat content of spring arrivals is also characteristic of migrants. Cardenolide content of adults showed a similar pattern of high content in spring arrivals, a decrease in the summer and then an accumulation of cardenolide defenses in adults in late summer just before migratory departure. We conclude that these results are consistent with temporally variable, sequential partial migration in a short-lived insect that contrasts with spatially variable partial migration in longer-lived vertebrates.
... In addition to being important ecologically, butterflies are known as a symbol of peace, happiness, and love and are of high ornamental and economic value [1]. Globally, a great deal of work has been done on butterfly classification [2][3][4][5][6][7][8][9][10], habitat utilization of butterflies as a resource [11][12][13][14][15][16][17][18][19], and butterfly conservation [19][20][21][22][23][24]. These studies are of great significance for the conservation of biodiversity, responsible utilization of butterfly resources, understanding the feeding habits of butterflies, conservation of butterfly habitat, and artificial breeding of rare butterflies. ...
... In butterfly foraging, vision plays an important role [5][6] and is vital in finding food and choosing spawning grounds [5]. Previous studies have found that butterflies' colour preference depends on sex [7], concentration [8][9][10][11], and pattern [12][13], and different butterflies forage on different types of flowers [14][15][16][17][18]. Behavioral studies in laboratory settings have demonstrated that yellow is the preferred flower colour of pierids [19]. ...
In addition to their ornamental value, butterflies provide value to flowering plants through pollination, which can improve their heterosis. Studying the relationship between butterflies and flowers is useful for understanding the adaptive relationship between them. By observing the pollinating and feeding behaviour of Aporia bieti in the Xiama forest, Gansu, we explored the adaptive relationship between A. bieti proboscis size and the corolla tube length of the flowers. To eliminate interference from the butterfly's sense of smell, we used foam flowers instead of real flowers. The results indicated that (1) of the red, yellow, pink, purple, and white flowers tested, yellow flowers were preferred; (2) there was no distinct difference in the preferences of A. bieti for large- (45 mm) and small- (25 mm) sized flowers; and (3) A. bieti proboscis length was significantly related to the corolla tube length of the flowers fed on by the butterfly, which suggests co-evolution between butterfly proboscis length and the length of the pollen tube.
... Foraging depends in part on the depth of the corolla (linked to the effort to reach nectar) and the clustering of flowers. Moreover, the length of proboscis limits the range of flowers from which nectar can be extracted (Corbet, 2000). There is a very strong correlation between the length of corolla tube and the sucrose concentration of floral nectar (Tiedge & Lohaus, 2017): indeed, flowers with short corollas produce nectars rich in glucose and fructose with little sucrose (which make them less prone to evaporation) (Baker & Baker, 1983). ...
Nutritional ecology examines the environmental effects on nutritional needs, food intake and foraging behaviors, and the use of nutrients ingested by animals. Adults of many insects’ species feed on nectars rich in sugars allowing them to match the nutritional needs necessary for reproduction. Among insects, Lepidoptera are often considered opportunistic foragers that visit a wide variety of available flowers, although with some preferences. While nutritional ecology of diurnal Lepidoptera is beginning to be explored, very little work focuses on nocturnal species because they are complicated to study in the wild. To address this, we used new laboratory approaches to study feeding behaviors (number and duration of visits to artificial flowers, food preferences associated with the texture and odors of the flowers) as well as gustatory detection by antennae (proboscis extension reflex) in the male crop pest moth Agrotis ipsilon. We showed that (i) food responsiveness is age-dependent and increases mainly with sugar quantity and marginally with sugar quality, (ii) diet quality impacts feeding behaviors in the first days of adulthood, and (iii) male moths choose their food through floral cues. Taken together, these data allow to define this species as a generalist forager with a preference for flowers with sugary nectars rich in sucrose, fructose, and glucose. Our results thus provide considerable information on the close links between food sources and nutritional ecology in this species, which is important for guiding future studies on their behavioral ecology, population dynamics, as well as for population monitoring and for regional pest management.
... Pieridae is a large butterfly family closely associated with flowering plants, adept at extracting nectar from flowers with various morphologies [19,20]. Although the types of sensilla on their proboscides are similar to those found in most nectarivorous lepidopterans [21][22][23], the proboscis structure of Pieridae shows significant divergence [2,9,24]. ...
Morphological studies of the proboscis can provide valuable insights into the evolutionary adaptations of Lepidoptera. However, such research is relatively limited for Pieridae, a family that is significant both ecologically and economically. In this study, scanning electron microscopy (SEM) was employed to investigate the proboscis and associated sensilla of adult C. erate, with an emphasis on ultrastructural details and potential sex-specific differences. The proboscis in both sexes exhibits a similar structure, consisting of two elongated maxillary galeae connected by dorsal and ventral legulae. It is divided into three distinct zones based on morphological changes in the dorsal legulae. The outer surface of the proboscis is characterized by spike-like microbumps on transverse ridges, while the inner surface (food canal) is smooth throughout all three zones, with regularly spaced transverse grooves. Three types of sensilla comprising five subtypes were identified: sensilla chaetica (sc), sensilla basiconica (sb1, sb2, and sb3), and sensilla styloconica (ss). The functional implications of these morphological features are also discussed briefly.
... However, we also found specialization to be common in nectar plant interactions, with 26% of butterfly species visiting three or fewer nectar plants. Our result supports previous findings that butterflies often have specialized nectar plant relationships due to specific chemical requirements or the coevolution of specialized pollination systems (Nilsson 1988;Lipani 1990;Corbet 2000;Fenster et al. 2004;Hardy et al. 2007;Krenn 2010;Jain et al. 2016;Levin et al. 2017). Importantly, we found that the composition of host and nectar plants differed significantly in all ecoregions, a pattern that was stronger when only the top 10% most important plants were considered. ...
Anthropogenic global climate change can disrupt plant-pollinator interactions by altering the traits, phenologies, and distributions of interacting species, exacerbating insect declines and compromising ecosystem function. However, most research has focused on diurnal pollinators, and little is known about the prevalence, importance, and vulnerability of nocturnal moth pollination. This knowledge gap limits our ability to predict and mitigate the effects of climate change and other stressors on moths and their pollination services. In this dissertation, I investigate the ecology of moth pollination interactions, how moths and their host and nectar plants will be impacted by climate change, and how to apply this knowledge in conservation strategies. I focus on native plants and moths in California, a biodiversity hotspot that is particularly impacted by climate change. I employ techniques ranging from greenhouse experiments to DNA metabarcoding to explore impacts spanning the levels of functional traits to ecological networks. In Chapter 1, I document hundreds of previously undescribed moth pollen-transport interactions along an elevational gradient spanning desert to conifer forest. I also find that moths are smaller, less diverse, and more sensitive to the simulated loss of their nectar plants in hotter and drier conditions. In Chapter 2, I reveal that experimental warming and drought alter diel patterns of floral nectar quantity and quality in a generalist plant. This may differentially affect interactions with diurnal and nocturnal pollinators, scaling up to alter the structure and stability of plant-pollinator interaction networks. In Chapter 3, I analyze and compare Lepidoptera-host and -nectar plant interaction networks across California, revealing structural differences and spatial patterns that inform management priorities. I also analyze species roles in networks to identify spatially-explicit keystone plant species to be used in butterfly and moth conservation efforts. Together, my results reveal that moth pollination interactions are diverse, complex, and vulnerable to climate change, and that data-driven conservation strategies can help protect them. Ultimately, this dissertation highlights the importance of considering the nocturnal components of plant-pollinator networks in research and management.
... (Barbosa 2013). The choice of plants by butterflies is influenced by a variety of visual stimuli, including flower shape and size, petal colour, bloom structure or cluster and floral density (McCall and Primack 1992;Waser et al. 1996;Corbet 2000;Tudor et al. 2004;Tiple et al. 2005;Rani and Raju 2016). In addition to visual stimuli, floral scent, the volume of petals and the components and concentrations of floral nectar all have a collective role in flower attraction (May 1985;Kunte 2000;Tiple et al. 2005;Jürgens et al. 2014). ...
Butterflies have a wide spectrum of colour vision, and changes in flower colour influence both the visiting and nectaring (the act of feeding on flower nectar) events of them. However, the spontaneous behavioural response of butterflies while foraging on real flowers is less characterised in wild conditions. Hence, this study intends to investigate flower colour affinity in wild butterflies in relation to nectaring frequency (NF) and nectaring duration (ND). Six distinct flower colours were used to study spontaneous nectaring behaviour in 20 species of subtropical butterflies. Both NF and ND greatly varied in the flower colours they offered. Yellow flowers were frequently imbibed by butterflies for longer durations, followed by orange, while red, pink, white and violet flowers were occasionally nectared in shorter bouts. Though butterflies have a general tendency to nectar on multiple flower colours, the Nymphalids were more biased towards nectaring on yellow flowers, but Papilionids preferred both yellow and orange, while the Pierids were likely to display an equal affinity for yellow, orange and violet flowers as their first order of preference. Even if the blooms are associated with higher nectar concentrations or a significant grade reward, the butterfly may prefer to visit different-coloured flowers instead. Flower colour choice appears to be a generalist phenomenon for butterflies, but their specialist visiting nature was also significant. Nymphalid representatives responded to a wider variety of floral colour affinities than Pierid and Papilionid species. The colour preference of butterflies aids in the identification of flowers during foraging and influences subsequent foraging decisions, which ultimately benefits pollination success. The current information will support the preservation and conservation of butterflies in their natural habitats.
... The observation showed that adults P. demoleus visited Zinnia flowers the most, whilst P. polytes visited A. leptosus the most. Preference on flowers visited by both butterfly species was not observed in much detail as adult butterflies in general would take almost any flowers (Courtney, 1986;Shreeve, 1992) with corolla depth that can be reached by the proboscis (Corbet, 2000;Tiple et al., 2009). At a particular time when there were so many butterflies of different species in the dome, additional 10% sugar solution was placed on Hibiscus flowers and both butterfly species would also take it. ...
Papilio demoleus L. and P. polytes L. are common butterflies and distribute almost throughout Indonesia. Both species are attractive in butterfly gardens, but may be considered as pests in Citrus plantations. This research aimed to obtain data on their biology, on how the species thrive in captivity, and to assess the alternative host plants. Captive breeding research on these two species was conducted at the butterfly research facility within the period of September 2016 to February 2019, with 482 individuals of P. demoleus and 2,334 individuals of P. polytes reared, of which 292 individuals of P. demoleus and 560 individuals of P. polytes have complete informative data. The average duration of eggs was 3.7 days for P. demoleus on Citrus spp., 3.68 days for P. polytes on Citrus spp., and 3.48 days for P. polytes on Micromelum minutum. The duration of larvae varied between 13–19 days for both species. Prepupal stage lasted for 1 day for all observed individuals. Incidental observation at home during the pandemic COVID-19 added some insights that the pupation happened between 18:00-19:00. The duration of pupae varied between 9–14 days for both species. The total duration of pre-adult stages for both species was between 26–38 days. Adults at the butterfly dome could live up to 19 days for P. demoleus and 39 days for P. polytes. Recognition of alternative host plants is very useful for the improvement of species management in butterfly gardens and in Citrus plantations.
... Plant species selection for NFS considered the following criteria: i) perennial and endemic to the Mediterranean-climate vegetation of central Chile; ii) species which in combination exhibit a long flowering period, including avocado flowering period, offering flower resources also when avocado is not in bloom (Russo et al., 2013); iii) a combination of species with short and long corollas which could benefit different type of insects (Balzan et al., 2014;Campbell et al., 2012;W€ ackers, 2004); and iv) the inclusion of species that offer flower resources clustered in dense inflorescences (Asteraceae and Apiaceae), which can also favor some insects (Corbet, 2000). A few species failed to establish (exhibiting 2% of final coverage), and thus NFS were composed of seven to eight species (Table 1) that grew in mixed culture within the strip. ...
Pollination is an essential ecosystem service for pollinator-dependent crops and plant communities. Apis mellifera L. is by far the most commonly used species to obtain this service in agriculture. However, there is growing evidence of the importance for crop yields of the service provided by wild bees and non-bee insect pollinators. Establishing flower resources in agricultural landscapes is a management practice that can increase insect pollinator populations and improve crop yields. We established perennial native flower strips (NFS) in four avocado orchards in central Chile during autumn 2017. We monitored flower visitors and counted newly formed fruits in avocados near and far from NFS in spring 2019, to assess flower visitor groups, flower visitation rates and fruit numbers. Only A. mellifera visited avocado flowers within bees, whereas both the managed A. mellifera and wild bees were the main visitors to the NFS. NFS increased visitation rates to adjacent avocado of flies, wasps and the sum of all non-managed flower visitors (i.e. excluding A. mellifera). However, there were no differences in the rates of A. mellifera visitation to avocados near and far from NFS. Avocado fruit numbers were higher among avocados near NFS than among those farther away. This difference could be due to better pollination by the increased visits to avocado flowers by flies or other wild insects. Therefore, NFS could contribute to crop fruit number, as well as conservation of native flora, wild bees and non-bee pollinators on fruit farms in the “Central Chile” biodiversity hotspot.
... Since variation in butterfly visual sensitivity is coarsely associated with variation in butterfly wing pattern (Stavenga et al. 2001;Briscoe et al. 2010;Everett et al. 2012), comparing the colors of flowers visited by different colored butterflies is a first step toward assessing whether butterflies are partitioning food sources in a color-dependent way at a community scale. While other floral traits, such as shape, odor, and amount of nectar reward, may also be important components of flower selection (Corbet 2000;Tang et al. 2013), butter-flies use approximately two-thirds of their brains to process visual information (Snell-Rood et al. 2009;Heinze and Reppert 2012). Thus, visual information is likely to be particularly important for flower selection by butterflies. ...
Community science, which engages students and the public in data collection and scientific inquiry, is often integrated into conservation and long-term monitoring efforts. However, it has the potential to also introduce the public to, and be useful for, sensory ecology and other fields of study. Here we describe a community science project that exposes participants to animal behavior and sensory ecology using the rich butterfly community of Northwest Arkansas, USA. Butterflies use visual signals to communicate and to attract mates. Brighter colors can produce stronger signals for mate attraction but can also unintentionally attract negative attention from predators. Environmental conditions such as weather can affect visual signaling as well, by influencing the wavelengths of light available and subsequent signal detection. However, we do not know whether the signals butterflies present correlate broadly with how they behave. In this study, we collaborated with hundreds of students and community members at the University of Arkansas (UARK) and the Botanical Gardens of the Ozarks (BGO) for over 3.5 years to examine relationships among wing pattern, weather, time of day, behavior, and flower choice. We found that both weather and wing color influenced general butterfly behavior. Butterflies were seen feeding more on cloudy days than on sunny or partly cloudy days. Brown butterflies fed or sat more often, while white butterflies flew more often relative to other butterfly colors. We also found that there was an interaction between the effects of weather and wing color on butterfly behavior. Furthermore, butterfly color predicted the choice of flower colors that butterflies visited, though this effect was influenced by observer group (UARK student or BGO participant). These results suggest that flower choice may be associated with butterfly wing pattern, and that different environmental conditions may influence butterfly behavior in wing-pattern-specific ways. They also illustrate one way that public involvement in behavioral studies can facilitate the identification of coarse-scale, community-wide behavioral patterns, and lay the groundwork for future studies of sensory niches.
... Species with shorter tongues do not visit flowers with deeper corolla tubes. Hence, short tongued Lycaenids like Silver lines are inclined towards Asteraceae members with short corolla tubes like Eupatorium odoratum and Solidago canadensis (Corbet, 2000). Monkey puzzle butterfly (Rathinda amor) and Ape fly (Spalgis epeus) were strongly attracted to Noni flower nectar. ...
Abstract
Aim: The present study was taken up to assess the food resources and butterfly diversity in Botanical Garden, Tamil Nadu Agricultural University.
Methodology: The survey was conducted in targeted locality from September 2015 – October 2019 and butterflies sightings were recorded using a digital camera (Nikon D7200 with kit lens 18-105 mm, Tamron 200-300 apo dg macro lens).
Results: Totally 95 species of butterflies were recorded in the garden. The count of Nymphalidae was 30 which is highest among all families, followed by Lycaenidae (29 species), Pieridae (15 species), Hesperiidae (10 species) and Papilionidae (11 species) were recorded. The relative diversity of Nymphalidae (31.57 %) and Lycaenidae (30.52 %) were higher. For Pieridae, RD value accounted for 15.78 % and 11.57 % for Papilionidae. RD value was least for the Hesperiidae family (10.52 %). Out of 95 butterflies, the host plants of 88 butterflies were present in the garden itself. Fifteen nectar plants belonging to eleven families were most preferred nectar sources.
Interpretation: The present study gives the information of butterfly and its food plant diversity in the garden. The obtained information will help in planting of additional host and nectar plant resources to attract rare species. It also sheds light on the importance of survival of prominent nectar plants throughout the year for maintaining the butterfly abundance in garden.
Keywords: Butterfly, Conservation, Diversity, Host plant
... No proboscis-tube relationship has been found in other studies (e.g. 9,12,52 ), although some studies found larger butterflies visiting larger flowers as well 17,18 . Nevertheless, although preferring flowers with longer tubes, the long-proboscid lepidopterans were not more specialised with respect to their food-niche breadth (relative number of visited plant species) in our study. ...
Butterflies and moths are conspicuous flower visitors but their role in plant-pollinator interactions has rarely been quantified, especially in tropical rainforests. Moreover, we have virtually no knowledge of environmental factors affecting the role of lepidopterans in pollination networks. We videorecorded flower-visiting butterflies and hawkmoths on 212 plant species (> 26,000 recorded hrs) along the complete elevational gradient of rainforests on Mount Cameroon in dry and wet seasons. Altogether, we recorded 734 flower visits by 80 butterfly and 27 hawkmoth species, representing only ~ 4% of all flower visits. Although lepidopterans visited flowers of only a third of the plant species, they appeared to be key visitors for several plants. Lepidopterans visited flowers most frequently at mid-elevations and dry season, mirroring their local elevational patterns of diversity. Characteristics of interaction networks showed no apparent elevational or seasonal patterns, probably because of the high specialisation of all networks. Significant non-linear changes of proboscis and forewing lengths were found along elevation. A positive relationship between the lengths of proboscis of hesperiid butterflies and tube of visited flowers was detected. Differences in floral preferences were found between sphingids and butterflies, revealing the importance of nectar production, floral size and shape for sphingids, and floral colour for butterflies. The revealed trait-matching and floral preferences confirmed their potential to drive floral evolution in tropical ecosystems.
... At the same time, this finding supports the use of proboscis length as a morphological indicator of resource utilization in butterflies. Similar findings were recorded in the study by Corbet, (2000) which showed that the maximum corolla depth of potential nectar plants limits the species feeding on them to those with sufficiently long proboscises; short-tongued butterfly species are therefore unable to feed on deep flowers. Moreover, Sultana et al. (2017) found that the proboscis had significant role in the coevolution between butterflies and their nectaring plants. ...
Floral attributes often influence the foraging choices of nectar-feeding butterflies, given the close association between plants and these butterfly pollinators. The diversity of butterflies is known to a large extent in Nepal, but little information is available on the feeding habits of butterflies. This study was conducted along the periphery of Rupa Wetland from January to December 2019 to assess butterfly species diversity and to identify the factors influencing their foraging choices. In total, we recorded 1535 individuals of 138 species representing all six families. For our examination of butterfly-nectar plant interactions, we recorded a total of 298 individuals belonging to 31 species of butterfly visiting a total of 28 nectar plant species. Overall, total butterfly visitation was found to be significantly influenced by plant category (her-baceous preferred over woody), floral color (yellow white and purple preferred over pink), and corolla type (tubular preferred over nontubular). Moreover, there was a significant positive correlation between the proboscis length of butterflies and the corolla tube length of flowers. Examining each butterfly family separately revealed that, for four of the families (Lycaenidae, Nymphalidae, Papilionidae, and Pieridae), none of the tested factors (flower color, plant category, and corolla type) were shown to significantly influence butterfly abundance at flowers. However, Hesperidae abundance was found to be significantly influenced by both flower color (with more butterflies observed at yellow flowers than purple) and flower type (with more butterflies observed at tubular flowers than nontubular flowers). Our results reveal that Rupa Lake is a suitable habitat for butterflies, providing valuable floral resources. Hence, further detailed studies encompassing all seasons, a greater variety of plants, and other influential factors in different ecological regions are fundamental for creating favorable environments to sustain important butterfly pollinators and help create balanced wetland ecosystems. K E Y W O R D S corolla depth, Lepidoptera, nectar plants, proboscis length, species diversity
... Adult butterflies have antennae, compound eyes, a proboscis, six legs, a hard exoskeleton, a pair of forewings and hind wings, and it poses a three section in their body (Schappert ,2000). The characteristic of a flower may also play a vital role in the feeding behaviour.The nectar of a flower is the main source of nutrition for adult butterfly nutrition (Omura and Honda ,2005).In searching food, Butterflies are the ecological diverse group of insects showing the complex foraging behaviour (Sourakov et al, 2012) .It choose its nectaring plant according to the colour ,odour and ( De Vries et al, 1999) the foraging and feeding behaviour of butterfly expends on the proboscis length which limits the range of flowers from the nectar extracting (Corbet, 2000). There were more number of research were carried out by most of the researches about the morphology and taxonomy of different butterfly but works on the feeding and Nectaring behaviour of butterfly with the time was not great in the field. ...
The butterflies are diurnal and very attractive organisms with the aesthetical and economical values. Ecologist use the butterflies as the model organism to study the impact of the habitat loss and climatic change. Preliminary study of the butterflies and its feeding behaviour not as great in Batticaloa district.Therefore,The present study was carried out to assess the feeding behaviour in Monarch butterfly during the morning and evening hours of a day. The objective of this study was to assess the feeding behaviour of butterfly in morning and evening hours and the detailed ethogram steps of feeding behaviour of butterfly.The present study was carried out in a selected home garden in Batticaloa during the period of February to March 2016.The Monarch butterfly was observed with 10 minutes interval per hour.The time taken for each feeding was recorded in seconds for both morning and evening hours. Totally 15 trials were taken on each attempt during the period of study. The time taken for the feeding in each time was recorded by digital stop watch and Canon EOS digital camera.Data were analysed statistically using Microsoft Excel 2013 for Chi-square test.Results (X 2 ,p< 0.05) of the present study revealed that the feeding behaviour of Monarch butterflies were significantly high in morning hours in comparing with the evening hours
... At the same time, this finding supports the use of proboscis length as a morphological indicator of resource utilization in butterflies. Similar findings were recorded in the study by (Corbet, 2000) which showed that the maximum corolla depth of potential nectar plants limits the species feeding on them to those with sufficiently long proboscises; short-tongued butterfly species are therefore unable to feed on deep flowers. Moreover, (Sultana et al. , 2017) found that the proboscis had significant role in the co-evolution between butterflies and their nectaring plants. ...
... However, Papilio polytes and several other butterfly species have been recorded to take nectar from this plant (Raju et al. 2001, table 1) as observed also with other butterfly species in the enclosure. Flower preference by P. peranthus was not investigated in detail as the adults in general would visit almost any flowers with corolla depth that could be reached by the proboscis (Corbet, 2000;Tiple et al., 2009). Based on daily observations inside the enclosure, newly emerged butterflies of this species do not visit flowers for nectar on the day of emergence. ...
Papilio peranthus is endemic to Indonesia, where it occurs on several islands and island groups. This beautiful butterfly is extensively traded, thus efforts to breed this species are very desirable. Captive breeding research was conducted on P. peranthus during September 2016 to December 2018. In total, 221 individuals were available for observation. Data on the life cycle of the species, together with observations on females being approached for mating, and female oviposition after mating, are presented. The result demonstrate that P. peranthus is not monogamous. Observations on other biological aspects are also reported.
... Liquid-feeding behavior plays an important role in bringing water, sugar, amino acids, organic acids, proteins, fats, vitamins, minerals and other minor components to organisms (Corbet, 2000;Bertazzini et al., 2010). Some studies have investigated the drinking patterns of animals that use tongues or tongue-like devices to load fluid (Kingsolver and Daniel, 1983;Winter and von Helversen, 2003;Krenn, 2010;Karolyi et al., 2014). ...
The morphological characters of honeybees were measured using a Scan Photo technique (SPT) that involves the use of a stereo binocular microscope and Photoshop program combined together. The body parts of a honeybee worker were scanned as images after they have been dissected. Photoshop program was used to view the images and their characters were measured using the ruler of the software program. While measuring the chosen morphometric characters, a significant difference was observed when the SPT and using a binocular microscope were compared. The measurements of 7 morphological characters of honeybee workers from nectar-feeding colonies were compared with those of Pepsi cola feeding colonies. There are more than 24 different subspecies of honeybees (Apis mellifera) classified based on their morphological characteristics. As a result, morphological characteristics are usually considered when classifying honeybees. Many authors for various reasons have used body morphological characteristics as well as sets of wings to classify subspecies of honeybees. Such characteristics have been defined overtime and sourced from diverse studies. Of all body morphological characteristics studied so far, wing venation characteristics have been given more attention. Up to now in Saudi Arabia, there are no specific review articles that focus mainly on Apis mellifera jemenitica body morphological characteristics. Consequently, information gathered from sampling and measurement methods, and factors affecting these characteristics especially mouthparts is to recognize the possibility of finding any changes in configuration as a result of feeding on soft drink. The morphology and functional anatomy of the mouthparts are examined by dissection, light microscopy, and supplemented by feeding type observations. This manuscript focuses on the differentiation of the proboscis which is formed by the glossa, in context with feeding from natural nectar and soft drink. In the literature, morphological innovations have been described for insects, particularly forbees.These innovations are critical for the production of differentiation in proboscis such as proboscis hair formation and modification. The present study showed that the proboscis of honeybees feeding on Pepsi is wider, thicker and very hairy compared to the natural feeding honeybee's proboscis. The morphological structures of the head, proboscis, hind leg, and wings were measured for the two groups of bees, the first naturally nourished with nectar, and the second feeding on the drink Pepsi Cola. The results showed significant differences in nectar and Pepsi feeding in different length measurements of head (18.
... Floral nectaries are either easily accessible or deeply embedded within corolla tubes or other floral structures such as spurs (Bernardello, 2007). In species with hidden nectar, the relative shape and size of the visitor's mouthparts compared to the shape and size of the nectarholding structure (e.g., corolla tube) are essential in determining the visitor's access to the nectar and its efficiency as a pollinator (Corbet, 2000;Inouye, 1980;Stang, Klinkhamer, & van der Meijden, 2007). ...
The foraging behavior of bees is a complex phenomenon that depends on numerous physical features of flowers. Of particular importance are accessibility of floral rewards, floral proportions, symmetry and orientation. The flowers of Roepera are characterized by the presence of staminal scales (SS), which play an important role in nectar protection. We studied two species of Roepera with different symmetry and flower orientation, which are mainly visited by honeybees (Apis mellifera). We aimed to show how the foraging behavior of honey bees is affected by the function of SS, floral symmetry and orientation. The foraging behavior was documented by video photography. Handling time, access to nectar, percentage of pollen/nectar foraging, percentage of pollen contact and pollen deposition site on the honey bee's body were assessed. The morphometric features of the honey bees and flowers were analyzed. We found that the SS restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The relative proportions of SS and proboscis length appear to be crucial for the success of pollinators. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency. We found that the staminal scales restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency.
... Species preferring sodium-rich substrates (like rotting fish or moist surfaces) had a significantly higher WL. A similar effect was observed among European nectar-feeding butterflies, where species with high WL forage mainly on aggregated or unusually nectar-rich flowers, while low-WL species were more likely to include solitary or less nectar-rich flowers in their diet (Corbet, 2000). It is unclear whether the higher WL observed in species relying on nutrient-rich resources reflects increased selection on flight performance in species with intensive foraging activities or conversely selection on resource acquisition in species with a high WL. ...
Butterflies display extreme variation in wing shape associated with tremendous ecological diversity. Disentangling the role of neutral versus adaptive processes in wing shape diversification remains a challenge for evolutionary biologists. Ascertaining how natural selection influences wing shape evolution requires both functional studies linking morphology to flight performance, and ecological investigations linking performance in the wild with fitness. However, direct links between morphological variation and fitness have rarely been established. The functional morphology of butterfly flight has been investigated but selective forces acting on flight behaviour and associated wing shape have received less attention. Here, we attempt to estimate the ecological relevance of morpho-functional links established through biomechanical studies in order to understand the evolution of butterfly wing morphology. We survey the evidence for natural and sexual selection driving wing shape evolution in butterflies, and discuss how our functional knowledge may allow identification of the selective forces involved, at both the macro-and micro-evolutionary scales. Our review shows that although correlations between wing shape variation and ecological factors have been established at the macro-evolutionary level, the underlying selective pressures often remain unclear. We identify the need to investigate flight behaviour in relevant ecological contexts to detect variation in fitness-related traits. Identifying the selective regime then should guide experimental studies towards the relevant estimates of flight performance. Habitat, predators and sex-specific behaviours are likely to be major selective forces acting on wing shape evolution in butterflies. Some striking cases of morphological divergence driven by contrasting ecology involve both wing and body morphology, indicating that their interactions should be included in future studies investigating co-evolution between morphology and flight behaviour.
... This creates a nested hierarchy that is ordered by gape width and fruit size (Wheelwright 1985;Rey et al. 1997;Burns 2006Burns , 2013. Size coupling has also been documented in a variety of plant-pollinator groups, including solitary bees (Armbruster & Guinn 1989), bumblebees (Harder 1985), long-proboscid flies (Goldblatt & Manning 2000), butterflies (Corbet 2000) and moths (Haber & Frankie 1989;Agosta & Janzen 2005). Little is known about the role species traits play in plant-bird pollinator network topology. ...
... Analyses of the four focal butterfly species showed that more monarch and gulf fritillary adults were observed in plots with exotic plant species. The presence of highly attractive plant species with larger and brighter floral displays, such as butterfly bush (Buddleia davidii) present in the exotic plots, may have influenced monarch and gulf fritillary abundance (Corbet, 2000;Matteson & Langellotto, 2011). Also, exotic ornamentals tend to bloom over a longer duration (Garbuzov & Ratnieks, 2014b), a trend we observed in our study plots. ...
• Pollinator declines have motivated efforts to plant nectar and host plants for butterflies and other pollinators, but whether gardens promote pollinator conservation requires further investigation.
• We established garden plots to determine whether plant type (native vs. exotic) and weed maintenance (low or high) influence adult butterfly abundance and species richness, and the abundance and survival of immature stages of four species (monarch, queen, black swallowtail, and gulf fritillary). To assess how predator activity and mortality of immature stages might differ inside gardens compared to other suitable habitat patches, we compared caterpillar survival on sentinel host plants placed within and outside of plots.
• The abundance and species richness of adult butterflies (all taxa) increased with the number of flowering plants per plot, but did not depend on plant type or weeding treatment. Exotic plots had greater adult monarch and gulf fritillary abundances, greater monarch and queen egg counts, and lower black swallowtail caterpillar counts relative to native plots.
• The survival of immature stages (egg to instar 5) ranged from 2 to 13% and was similar across plots, except for gulf fritillary larvae, which had higher survival in exotic plots. Monarch caterpillar mortality risk was higher on sentinel plants placed inside plots, relative to those outside of plots.
• This study suggests that garden plots attracted a diversity of adult butterflies and supported the reproduction of focal species. Given lower immature monarch survival within versus outside of plots, further work is needed to examine natural enemy pressure within pollinator gardens.
... Long proboscis observed in Hesperiidae is attributed to the coevolution with flowers having corolla tube (Kunte 2000). Maximum corolla depth of potential nectar plants limits species feeding on them to those with sufficiently long tongues; short-tongued butterfly species are therefore unable to feed on deep flowers and corolla depth has been shown to place a limit to exploitation by nectar feeding butterflies (Corbet 2000). The species with longest proboscis utilize the highest range of corolla tube depths. ...
p>An attempt was made to examine butterfly proboscis length and their significance in carrying out activities of the butterflies in relation to their nectar plants. Observations were made in seven selected areas (viz. Satchari, Modhupur, Rema-kalenga, Shaltila, Bhawal National Park, and Botanical and Zoological gardens of the Curzon hall area) from July 2014 to June 2015. Thirty four butterfly species of seven families, viz. Hesperiidae, Nymphalidae, Danaidae, Papilionidae, Pieridae, Lycaenidae and Satyridae, were observed. The strategic activities of proboscis in different butterflies were examined when they were used in foraging activity. The proboscis length of butterflies was measured during the study period. Nectar plants were identified in the laboratory. The corolla length of the nectar plants was measured in the field and found a good relation with the proboscis length of the butterflies. Among the observed species, hesperiid butterflies comparatively contain long proboscis, and can visit flowers up to 28 mm long corolla tube. The results obtained, indicate that the butterfly proboscis had significant role in co-evolution between the butterfly species and the flowers of the nectar plants.
J. Biodivers. Conserv. Bioresour. Manag. 2017, 3(1): 93-102 </p
... Thus, butterflies and many species of flowering plants are interdependent [32] and thus, butterfly diversity clearly indicated the floral diversity of an ecosystem. Further, butterflies select nectar plants to get energy requirements [35] . They generally prefer clustered flowers compared to solitary flowers to obtain good amount of nectar with in short time followed by little expenditure of energy. ...
Butterflies are opportunistic foragers feed on various plants. To record nectar plants of few butterfly species, systematic field survey was conducted by employing visual count method (VCM) and an all out search method (AOSM) at agriculture ecosystems of Chamarajanagar District during 2013 and 2014. Total 86 flowering plant species which belong to 27 plant families were visited by four butterfly species namely: P. polytes, G. agamemnon, A. merione and J. hierta for nectar collection. Interestingly, weeds have contributed more (49%) nectar to these butterfly species and it was followed by shrubs (14%), herbs (13%), trees (8%) and climbers (2%). Compositae (10 plant species) and Acanthaceae (8 plant species) families were visited more often for nectar collection compared to other families by these butterflies. Number of plant species visited by these butterfly species for nectar collection is recorded in detail. However there was a significant difference (F=11.048; P>0.05) existed between nectar plants visited by butterfly species. Further, color of the flower matters a lot while collecting nectar. Yellow, white, pink and blue coloured flowers were visited more often and there exited a significant difference (F= 30.117; P> 0.05) between colored flowers visited by butterfly species. Furthermore, floral calendar was prepared by using 86 flowering plants blooming period. It could help understand the locally available flora with different flower colours as source of food for few butterfly species and emphasized the need of herbaceous flora conservation to restore native butterfly species amidst agriculture ecosystems.
... Odor sometimes acts as a synergist with color as the important cue for foraging (Ômura & Honda 2005). The usefulness of butterfly foraging depends on corolla depth and proboscis length, which limits the range of flowers from which nectar can be extracted (Porter et al. 1992;Corbet 2000). ...
A study was conducted during the flowering season of Ziziphus mauritiana from September 2015 to October 2015 in Belgachi Railgate Para, Chuadanga, Bangladesh. The study recorded 265 individuals of 39 butterfly species belonging to five families and 32 genera nectar feeding on Z. mauritiana flowers. Amongst the families, Lycaenidae was dominant with 33.33% (n=13). Amongst the species Parnara bada (Moore, 1878) (Hesperiidae) was the most dominant species followed by Ypthima baldus (Fabricius, 1775) (Nymphalidae). Virachola isocrates (Fabricius, 1793) (Lycaenidae) spent the maximum time (60–120 sec) nectar feeding on Z. mauritiana amongst the 39 species sampled.
... In this situation, the scenarios are not foreseeable. Zheng et al. [46] found AR = 3.18 for Vanessa cardui (Nymphalidae), Dudley [21] [47] found WL values from 0.56 to 0.83 N/m 2 in four European species of Pieridae and from 1.20 to 1.80 N/m 2 in five species of Nymphalidae, three being perchers. Fric et al. [48] (2006) discussed AR and WL but did not show raw data. ...
... The use of specific nectar sources is one indicator for the ecological specialization in insects (Waddington, 1980;Murphy et al., 1984;Baguette & Nève, 1994;Biesmeijer & Tóth, 1998;Corbet, 2000;Tudor et al., 2004). Most butterfly species are considered to be opportunistic in the general use of these resources (Sharp et al., 1974;Loertscher et al., 1995;Dosa, 1999;Bakowski & Boron, 2005). ...
High mountain ecosystems are a challenge for the survival of animal and plant species, which have to evolve specific adaptations to cope with the prevailing extreme conditions. The strategies to survive may reach from opportunistic to highly adapted traits. One species successfully surviving under these conditions is the here studied butterfly Erebia nivalis. In a mark-release-recapture study performed in the Hohe Tauern National Park (Austria) from 22 July to 26 August 2013, we marked 1386 individuals and recaptured 342 of these. For each capture event, we recorded the exact point of capture and various other traits (wing conditions, behaviour, nectar sources). The population showed a partial protandrous demography with the minority of males emerging prior to the females, but the majority being synchronised with them. Males and females differed significantly in their behaviour with males being more flight active and females nectaring and resting more. Both sexes showed preferences for the same plant species as nectar sources, but this specialisation apparently is the result of a rapid individual adaptation to the locally available flowers. Estimates of the realised dispersal distances predicted a comparatively high amount of long-distance flights, especially for females. Therefore, the adaptation of Erebia nivalis to the unpredictable high mountain conditions might be a mixture of opportunism and specialised traits. This article is protected by copyright. All rights reserved.
This is a critical moment for land use policy globally, with many countries (e.g. the UK and the European Union) currently undertaking significant green reforms of their agricultural policies. Despite their importance for maintaining agricultural outputs and plant diversity, the effects of artificial soil enrichment on pollinators remain poorly understood. Our two-year study at the world’s longest-running ecological experiment, Park Grass, Rothamsted, examines the relationship between soil fertilisation, grassland yield and biodiversity. Our data show a large and significant negative effect of the major plant nutrients (NPK) on the abundance, species richness and functional diversity of both pollinators and flowering plants. The results also indicate a large and significant trade-off between productivity and biodiversity. Our findings are a salutary reminder of the challenge in reconciling conflicting aims in farmland management and strongly suggest that financial incentives are necessary to offset yield reductions to improve biodiversity outcomes in agricultural grasslands.
Aims: The present work aims to analyze the environmental drivers of diversity in the butterfly community in the western Qinling Mountains.
Methods: In the autumn of 2020 and spring and summer of 2021, we investigated butterfly diversity in the western region of Qinling Mountains using line transects across multiple habitat types in 15 sampling areas. We used trend and extrapolation analyses for estimating α diversity, and non-metric multidimensional scaling (NMDS) and cluster analyses for β diversity. For determining drivers of butterfly diversity, we fit environmental factors to diversity indices using a generalized additive model (GAM).
Results: We observed a total of 8,898 individuals representing 169 species, 84 genera, and 5 families. Of these families, the highest number of individuals were from Pieridae (N = 3,671), and the most number of species were from Nymphalidae (N = 80). We found that α diversity was highest during the summer and in coniferous and broad-leaved forests. For β diversity, we found the highest degree of similarity between coniferous and broad-leaved forest and deciduous broad-leaved forest, the low similarity between seasons, and that species are concentrated in spring and summer but relatively dispersed in autumn. The GAM fitted curves demonstrated several key relationships between environmental factors and butterfly diversity, including: (1) plant heterogeneity was correlated with butterfly community diversity; (2) an ambient temperature between 24℃ and 30℃ underlined a higher Pielou evenness index and a more stable butterfly community structure; and (3) humidity between 70% and 85% was associated with a higher Simpson index.
Conclusion: Butterfly community composition and diversity in the western region of Qinling Mountains were closely related to habitat type and have a distinct chronological relationship with seasons. Plant cover, abundance, humidity, and temperature are important factors in maintaining the diversity of butterfly species on a regional scale.
Abundance of pierid butterflies were estimated from June to September and their foraging pattern on Ver-nonia cinerea was recorded. Among the five butterfly species Appias albina was observed to be most frequent visitor of Vernonia cinerea comparatively than Leptosia nina (Psyche), Eurema hecabe (Common grass yellow), Catopsilia pomona (Common emigrant) and Catopsilia pyranthe (Mottled emigrant). The foraging activity pattern of these butterflies showed a definite pattern of foraging schedule by showing a resource partitioning. One sample t-test and month wise ANOVA indicated significant difference (p<0.05) for each butterfly species in different timeslots and how is co-related with blooming period of Vernonia cinerea. Our study indicates the importance of weeds like Vernonia cinerea in butterfly conservation project. The present study provides preliminary information on the interaction between native plants and butterflies.
Avantages et désavantages du buddleia
Most of our understanding of the effects of climate warming on insect body size comes from laboratory experiments. Whether these studies predict patterns in nature is largely unknown.
Here we examine the relevance of laboratory warming experiments for wild populations of the butterfly Pieris rapae . We tested two predictions: (i) butterflies reared at warmer temperatures in the laboratory should attain smaller adult sizes and have reduced flight ability, and (ii) in nature, this trait combination should lead to smaller butterflies visiting fewer flowers and accumulating less pollen.
Overall, we found that warm‐reared butterflies were indeed smaller and flew more slowly compared to colder‐reared conspecifics. Additionally, wild‐caught small butterflies carried fewer, and a lower diversity of pollen grains compared to larger butterflies.
Our warming experiments thus largely predicted pollen collection patterns in wild P. rapae .
This study demonstrates that increased temperatures will likely have important consequences for butterfly‐plant interactions in nature.
The Arabian Peninsula is a hotspot for bee diversity but studies looking at plant-insect interactions there remain rare. A network analysis of insects visiting wildflowers in the Hajar Mountains, Oman was made from the results of eight survey periods between 2016 and 2020. Centrality scores were used to assess the role different groups of potential pollinators play in network topology. A list of 113 insect species visiting 26 plant species has been compiled with Lepidopteran species acting as important connectors within the network. A nested, asymmetric and compartmentalised network was recorded. The order Hymenoptera was the most species rich group, with 46 species recorded followed by Diptera (43 spp.), Lepidoptera (13 spp.), and Coleoptera (11 spp.). Amegilla pyramidalis (Kirby, 1900) (Hymenoptera: Apidae: Antho phorini) is recorded for the first time outside of the Socotra Archipelago. The study is the first specific effort to record the flower visitation behaviour of insects in Oman and gives an overview of the resulting visitation network.
Traditionally managed agricultural lands in Transylvania repeatedly presented a high biodiversity and valuable insect communities. On the other hand, intensified agricultural landscapes have been somewhat less analysed from the perspective of habitat cover and habitat heterogeneity in the same region. The aim of our study was to document the fauna of diurnal lepidoptera in two types of habitats: grasslands and rape field margins, located in landscapes dominated by intensive agricultural practices. In addition we aimed to evaluate the diversity and ecological profile of diurnal lepidoptera in connection with the types and areas of the surrounding habitats. We found that lepidopteran communities in the two habitat types were dominated by species with different ecological profiles: grasslands by mesophilous species and rape field margins by ubiquitous species. The most important habitat types that confered our study plots their local character were the amount of surrounding cropfields, grasslands and tractor tracks. The species richness, abundance and diversity of diurnal lepidoptera were negatively correlated with the homogeneity of the semi-natural habitats in the surrounding landscape. Cuvinte cheie: peisaj agricol intensiv, diversitatea lepidopterelor diurne, omogenitatea habitatelor semi naturale
Herkogamy, the spatial separation of sex organs in hermaphroditic plants, has been proposed as a mechanism to reduce self-pollination and the associated processes of inbreeding and gamete wastage. Longitudinal herkogamy is the most frequent type, with two subtypes: approach herkogamy (anthers below the stigma), which is associated with diverse pollinator arrays, and reverse herkogamy (anthers above the stigma), associated with specialized, long-tongued pollinators. By using a herkogamy index that varied continuously from negative (reverse herkogamy) to positive (approach herkogamy) values, we studied the effect of continuous variation in herkogamy on pollinator attraction, selfing capability and plant fitness across three populations of Lonicera implexa differing in the relative abundance of long-tongued vs. short-tongued pollinators. Reverse herkogamy was significantly more frequent in the population where long-tongued pollinators were dominant than in the other two populations. Agreeing with this, the main floral visitors of L. implexa individuals with small and large herkogamy index were, respectively, long-tongued and short-tongued pollinators. Spontaneous selfing was low and increased with increasing herkogamy index (i.e. with approach herkogamy), although most of it occurred when there was close distance between anthers and stigma. Fruit production was unrelated to the herkogamy index in the population with long-tongued pollinators, but it increased with approach herkogamy (higher herkogamy index) in the other two populations. In contrast, seeds of individuals with reverse herkogamy (smaller herkogamy indices) germinated better. In this species, continuous variation in herkogamy might function as a reproductive strategy, as different morphotypes might be favoured by different pollinator assemblages.
Herkogamy, the spatial separation of sex organs in hermaphroditic plants, has been proposed as a mechanism to reduce self-pollination and the associated processes of inbreeding and gamete wastage. Longitudinal herkogamy is the most frequent type, with two subtypes:approach herkogamy (anthers below the stigma), which is associated with diverse pollinator arrays, and reverse herkogamy (anthers above the stigma), associated with specialized, long-tongued pollinators. By using a herkogamy index that varied continuouslyfrom negative (reverse herkogamy) to positive (approach herkogamy) values, we studied the effect of continuous variation in herkogamy on pollinator attraction, selfing capability, and plant fitness across three populations of Lonicera implexa differing in the relative abundance of long-tongued vs short-tongued pollinators. Reverse herkogamy was significantly more frequent in the population were long-tongued pollinators were dominant than in the other two populations. Agreeing with this, the main floral visitors of L. implexa individuals with small and large herkogamy index were, respectively, long-tongued and short-tonged pollinators. Spontaneous selfing was low and increased with increasing herkogamy index (i.e. with approach herkogamy), although most of it occurred when there was close distance between anthers and stigma. Fruit production was unrelated to the herkogamy index in the population with long-tongued pollinators, but it increased with approach herkogamy (higher herkogamy index) in the other two populations. In contrast, seeds of individuals with reverse herkogamy (smaller herkogamy indices) germinated better. In this species, continuous variation in herkogamy might function as a reproductive strategy, as different morphotypes might be favoured by different pollinator assemblages
Pachliopta aristolochiae plays important role in the ecosystems. Their role as pollinators can be observed from their daily behaviours. This study aimed to observe the behaviours of butterfly P. aristolochiae as well as their food preferences on daily basis. This research was conducted at the Museum of insect and butterfly of Taman Mini Indonesia Indah. Observations were done in two female individuals based on scan sampling method. Behaviour was observed according to perching, flying, nectaring and interaction activities. Our results indicate that P. aristolochiae mostly showed flying (35.9%) and nectaring (30.6%) activities. On the other hand, interaction (9.3%) was among the least behaviours observed during the study. Nectaring activity was mostly shown within 08.00-09.40 am, while perching was observed throughout the day within 11.00-16.00. During observation, P. aristolochiae was recorded in visiting various plants, including Ixora sp., Hibiscus rosa-sinensis, Clerodendrum japonicum, Ochna serrulata, Jatropha curcas and J. Integrifolia.
Several aspects of plant-pollinator interactions are presented in the thesis. It contains a review on the open questions of plant-pollinator interactions from single species to complex networks. The following sections document novel results. Firstly, the conservation of complex pollination networks is addressed through the hierarchy of species’ importance. In addition, the habitat requirements and interactions of a threatened rare pollinator species are explored. In the following chapters, the results from manipulative approaches applied in the field to plant-pollinator interactions are presented. The effect of pollinator’s population decline on pollinators’ foraging for pollen is investigated. Moreover, the way plant species loss impact several aspects of pollinator visitation is presented. Lastly, the impact of species removal on plant-pollinator network topology and on species ability of establishing new interactions is investigated.
A detailed study on the butterfly species diversity was carried out at Omkareshwar at Narmada river bank, in district Khandwa, Madhya Pradesh, India during 2011-13. The pattern of butterflies abundance and species richness are studied in wild (forest, scrub and grassland). A total of 70 butterfly species belonging to 8 families of order Lepidoptera were recorded during the study period. The family Nymphalidae, represented by 16 species was the most dominant followed by Papilionidae 07 Pieridae (11 species), Danaidae 05, Satyridae 08, Riodinidae 01, Lycaenidae 14 and Hespriidae 08 species. From the conservation point of view, the study area is undisturbed and rich in flora and fauna species.
Mass-specific oxygen consumption of euglossine bees during free hovering flight is inversely related to body mass, varying from 66mlO2g−1h−1 in a 1−0·10 bee to 154mlO2g−1h−1 in a 0·10-g bee. Individuals of Eulaema and Eufreisea spp. have smaller wings and higher wing stroke frequency and energy metabolism at any given mass than bees of Euglossa spp. or Exaerete frontalis. Calculated aerodynamic power requirements represent only a small fraction of the energy metabolism, and apparent flight efficiency [aerodynamic power (= induced + profile power)/power input] decreases as size declines. If efficiency of flight muscle = 0·2, the mechanical power output of hovering bees varies inversely with body mass from about 480 to 1130 W kg−1 of muscle. These values are 1·9 to 4·5 times greater than previous predictions of maximum mechanical power output (Weis-Fogh & Alexander, 1977; see also Ellington, 1984c). Mass-specific energy expenditure per wing stroke is independent of body mass and essentially the same for all euglossines. Differences in energy metabolism among bees having similar body mass is primarily related to differences in wing stroke frequency. Scaling of energy metabolism in relation to mass is generally similar to the relationship for sphingid moths despite the fact that bees have asynchronous flight muscle whereas moths have synchronous muscle.
Maximum lift production during takeoff in still air was determined for a wide variety of insects and a small sample of birds and bats, and was compared with variation in morphology, taxonomy and wingbeat type. Maximum lift per unit flight muscle mass was remarkably similar between taxonomic groups (54–63 N kg−1), except for animals using clap-and-fling wingbeats, where muscle mass-specific lift increased by about 25 % (72–86 N kg−1). Muscle mass-specific lift was independent of body mass, wing loading, disk loading and aspect ratio. Birds and bats yielded results indistinguishable from insects using conventional wingbeats. Interspecific differences in short-duration powered flight and takeoff ability are shown to be caused primarily by differences in flight muscle ratio, which ranges from 0.115 to 0.560 among species studied to date. These results contradict theoretical predictions that maximum mass-specific power output and lift production should decrease with increasing body mass and wing disk loading.
Summary 1. We assess the importance of body mass and the minimum ambient temperature at which foraging occurs in determining the warm-up rates and thoracic temperatures in flight at an air temperature of 22 °C of 55 species of bee (Hymenoptera: Apoidea) from six families adapted to a variety of thermal environments. 2. To control for the effects of taxonomic differences in the relationships between these variables, we use multiple regression incorporated in the phylogen- etic regression method developed by Grafen (1989). 3. The prediction made by May (1976) that for very small heterotherms warm- up rate will correlate positively with body mass is confirmed when the effects of phylogeny and the thermal environment to which the bee is adapted have been controlled for. The relationship between warm-up rate and body mass within the Apoidea is thus not an extension to lower body masses of the relationship found for heterothermic vertebrates. 4. Having controlled for the effects of body mass in our analyses, we demonstrate that bees able to fly at lower ambient temperatures have higher thoracic temperatures and warm-up rates than bees adapted to wanner environ- ments. 5. There is some suggestion that kleptoparasitic bees, being freed from the need to forage in order to provision cells, have lower warm-up rates than provisioning species. 6. The significance of these relationships in the ecology of bees is discussed in relation to studies of body temperatures and warm-up rates in bees and other insects.
This book was conceived to mark the Silver Jubilee of the British Butterfly Conservation Society. Interest in the conservation of butterflies has increased so rapidly that it is difficult to relate to the situation 25 years ago. Butterflies were on the decline in Britain, Europe and elsewhere but we lacked data on the extent of the decline and the underlying reasons, leaving us unable to implement effective conservation measures. An early recognition of the plight of British butterflies and moths led to the foundation of the society by a small group of conservationists in 1968. Today the society has over 10000 members, owns a number of reserves and sponsors research, conservation and monitoring activities at the local and national level. As part of the Silver Jubilee celebrations an international symposium was held at Keele University in September 1993 entitled 'Ecology and Conservation of Butterflies'. This symposium clearly showed how much important work has been done in recent years and also gave me the impression that the subject had reached a watershed. This was not because the decline of butterflies has stopped or even slowed down, far from it, the threat to our butterflies continues to increase from habitat destruction and intensification of land use. The watershed is in our understanding of the relationship between butterflies and their habitat.
This identification guide to the vascular flora of Britain and Ireland is drawn up from actual plant material and covers all natives, naturalized plants, crop plants and recurrent casuals: 2990 species and 197 extra subspecies are treated fully, with 559 hybrids and marginal species mentioned more briefly. The information, for each family or similar taxon, is presented in the form of an introductory summary of characteristics generally followed by a dichotomous key to genera; for each genus or similar taxon, a brief summary is followed by a dichotomous key to species and then by individual descriptions of the keyed species. These descriptions include other species not mentioned in the keys, as well as hybrids and subspecies. They also give information on status, habitat, distribution and frequency of occurrence or rarity, and indicate endemic or extinct plants. Within the book are interspersed 150 pages of illustrations and photographs of difficult groups. There is a glossary of terms used and an index combining common and Linnean names. -J.W.Cooper
Although there are indications that Lepidoptera had started to decrease early in the twentieth century (Entomologischer Verein Alpstein, 1989), they have declined throughout Europe most markedly during the past few decades. This decrease is so strong that it has been noticed not only by lepidopterists but even by lay-persons. According to red data lists, about 40–50% of all Lepidoptera species occurring in Germany and Austria are endangered, and 2–5% of the species occurring in these countries have already become extinct (Ebert and Falkner, 1978; Wagener et al., 1979; Pretscher, 1984; Gepp, 1981; Huemer, 1994; Huemer, Reichl and Wieser, 1994). In Switzerland, 39% of all butterfly species are endangered (Gonseth, 1987), although no recent extinctions have been reported so far. However, in some areas of Switzerland the percentage of endangered butterfly species is distinctly higher. According to Bryner (1987), 53% of the butterfly species originally occurring in the Seeland (area between the lakes of Biel and Murten) have already become extinct, 27% are endangered, leaving only 20% of mainly trivial species unthreatened. The figure for moths, although much less well known, is probably similar to that of butterflies.
Four woody species from the family Rosaceae share the same habitat and pollinators in Wytham Wood, Oxford. They are similar in floral morphology but Rosa canina L. differs in that the only reward it offers is pollen whereas the others offer both nectar and pollen. The three species which offer nectar are separated temporally in their flowering and the possible evolution of this separation is discussed.
1. The effects of biotic and abiotic factors on the distribution of three species of satyrid butterfly (Aphantopus hyperantus, Pyronia tithonus and Maniola jurtina) in arable field margins were studied in a block of arable farmland on a North Hampshire farm. 2. Non-floral factors affecting distribution included the degree of shelter, insolation, width of hedgebank or grass verge, and uncultivated habitat. Farm tracks adjacent to the field boundaries exerted a negative effect on abundance. 3. Floral variables affecting distribution included the abundance of flowers of bramble (Rubus fruticosus), thistle-like Compositae, marjoram (Origanum vulgare) and mayweeds (Matricaria spp.) in conservation headlands. Negative variables included abundance of cow parsley (Anthriscus sylvestris) and old man's beard (Clematis vitalba); although the latter was negative for A. hyperantus and M. jurtina, it was a positive variable for P. tithonus. 4. The distribution of butterflies in arable field margins is discussed in relation to landscape structure, microclimate and resources.
This paper examines whether the use of 14 plant species as nectar sources by eight species of bumble bees related systematically to differences in bee morphology. I predicted that a particular bee should have fed from a given plant species if the bee was physically more similar to the other bees visiting that plant species than to bees on any other species. Glossa (=tongue) length, body mass, and wing length all influence a bumble bee's foraging ability and its choice of flowers and were therefore included in the analysis. Morphological differences between bees were associated with use of different plant species; however, the role of bee morphology in flower choice was most evident when preferred plant species bloomed abundantly. The interaction between morphology and flower choice was also influenced by plant species richness, season, the plant species visited, and the species of bee; but was not affected by the time of day that the bee was foraging, overall bee density, or the bee's caste. Bee species with long glossae had access to nectar in a greater variety of flowers than those with short glossae, and they tended to feed from a larger number of plant species. Also, their use of a particular species was less predictable. Discrimination between bees using different plant species depended on joint consideration of several morphological characters: no character alone accurately separated the bees.
1. Flower visits by bumble-bees were recorded in a national survey throughout Britain in 1987 and 1988. 2. To avoid misidentifications, recorders were asked to name bumble-bees in terms of five colour groups. Each group included one or two common and widespread species, which are probably responsible for major patterns of flower usage, and several rarer forms. 3. A simple selectivity score ranks plant taxa in terms of the visits they receive from bumble-bees of each colour group. High scores may be due to high local flower abundance or to preference by the bees, and indicate plant taxa that may be worth encouraging to enhance bumble-bee populations locally. 4. A group-specific selectivity score highlights differences between bumble-bee groups in flower choice. It could be used to indicate which bumble-bee groups visit particular crops, and to plan vegetation management to favour particular bumble-bee species selectively. 5. Bumble-bees allocate a disproportionately high percentage of their visits to perennial plants of later successional stages. Annual plants of newly disturbed land receive relatively few visits. The implications for management are discussed.
This study examines flower choice by two syntopic butterfly species, Agraulis vanillae (Nymphalidae) and Phoebis sennae (Pieridae), and the energy consequences of foraging selectivity. Foraging selectivity is influenced by proboscis of P. sennae is @?~1 cm longer than that of A. vanillae, it is able to forage at long-corolla flowers (three Ipomoea spp.) inaccessible to A. vanillae. Relative to other nectar sources exploited by the two species, these Impomoea species are highly rewarding energetically. Phoebis sennae increases its energy intake relative to A. vanillae by exercising greater discrimination between high- and low-reward nectar sources, avoiding several species with small nectar volumes that are avidly visited by A. vanillae. Consequently, the average foraging profit of P. sennae is nearly 4 times that by A. vanillae. These differences in foraging selectivity energy intake are closely related to energy storage and depletion rates, measured as age-related changes in gross lipid contents of adult butterflies. Adult activities in the less selective species, A. vanillae, are fueled to a greater degree with lipid reserves set aside at metamorphosis; A. vanillae emerge from pupae with 13-16% of dry mass allocated to lipids, and these lipid stores decline at a variable rate throughout adult life. By contrast, in P. sennae lipid accumulated adult feeding provides the largest proportion of energy stores. P. sennae emerge from the pupa with only 6% of the dry mass allocated to lipids, and accumulate lipid stores to levels as high as 40% of dry mass through most of adult life, depleting these lipid reserves only in late adulthood. Further, A. vanillae show significant sexual dimorphism in lipid content throughout life, with females containing greater quantities, while P. sennae show no such dimorphism. These results are discussed with respect to Boggs' model concerning the allocation of larval-derived resources to reproduction.
1. Three major determinants of bees' nectar-foraging patterns are the maximum depth at which nectar is accessible (dependent on bee tongue length); the minimum profitable energy content per flower (dependent on foraging costs and hence on bee body mass); and the minimum threshold temperature for flight. These form the axes of a three-dimensional template, the competition box. 2. Thresholds of depth, cost and temperature can be identified for each species of bee, delimiting domains within the box compatible with foraging by each bee species. 3. Changes through a day in temperature and in the depth and sugar content of nectar define the nectar trajectory through the box for a flower species in relation to microclimate and bee activity. 4. This approach allows systematic appraisal of the roles of accessibility, energy content and temperature in interspecific competition among bees for nectar, and draws attention to situations in which additional factors may be important. 5. By superimposing a bee-species-specific template on a representative nectar trajectory for a given flower species in a given situation, it is possible to make reasoned guesses about the expected interactions of foraging bee species with a novel flower species and with each other in a novel situation. Such guesses are required for evaluating the expected impact of honey-bees and other introduced pollinators on the pollination system of crops and natural vegetation. 6. Use of the competition box is illustrated in relation to dawn-to-dusk studies involving measurements of nectar concentration and volume, microclimate, flower form, and the tongue lengths and foraging activity of bees of different species.
Twenty-four plant species native or naturalized in Britain were grown in the Cambridge University Botanic Garden, UK and evaluated as potential resources for nectar-foraging bees, butterflies and hoverflies. In ten plant species a series of measurements were made, at regular intervals from dawn to dusk, of nectar secretion rate and standing crop, and in all species insect visits were monitored throughout daylight hours. The study revealed differences between plant species in the composition of the assemblage of insect visitors, and in the magnitude and temporal distribution of the nectar reward. In some cases we found interesting correlations between temperature and secretion rates or patterns of insect visits. Species that received numerous insect visits in our study are potentially valuable forage plants that might be planted by gardeners to support local pollinator populations. Deep flowers whose nectar is accessible to long-tongued bumblebees (Bombus hortorum, B. pascuorum) but not to honeybees may provide long-tongued pollinators with a resource refuge relatively free from honeybee competition. Features that make some of those plant species particularly interesting to observe in the garden include robbing by short-tongued bumblebees inSaponaria,baseworking by honeybees on closed flowers inMalva sylvestris, and apparent displacement of bumblebees by territorial behaviour of the solitary beeAnthidium manicatumon species ofStachys.
Two types of selectively were observed in 4 of 13 groups of 10-15 bees: a preference for larger than average flowers when all flowers of a particular species were shallower than the lengths of the bees' glossae; or a positive correlation between glossa length and the depth of flowers visited when glossa length approximately equalled flower depth. An energetic model was developed based jointly on the dependence of a bee's probing time on glossa length and flower depth, and a positive correlation between flower depth and nectar production. Selective behaviour was reflected in very different patterns of size-related flower visitation, which depended on the particular characteristics of both bees and flowers. -from Author
A study was made of the activities of pollinating insects in relation to the flowering phenology of plants near Banff, Alberta in 1968. From snow-melting time (early May) to the end of May, pollinating insects were abundant and competed actively for relatively scarce pollen and nectar resources. Then, in early June some "cornucopian species" (Salix spp. and Taraxacum officinale L.) began flowering abundantly offering virtually unlimited supplies of nectar and pollen. Pollinating insects abandoned the spring flowers and were attracted to these cornucopian species. It is postulated that in the presence of cornucopian species natural selection would favor the evolution of earlier- or later-flowering populations of all those species that compete with the cornucopians for pollinators. It was found that in the spring, insects competed for flowers but that after the cornucopian species had completed their flowering (throughout the summer), flowers competed for pollinating insects. /// Исследовали активность насекомых-опылителей в эависимости от фенологии цветения растений в районе Банфф (Альберта) в 1968 г. С момента таяния снега (в начале мая) до конца мая насекомые встречались в большом количестве, и среди опылителей воэникала конкуренция иэ-эа пыльцы и нектара, т.к. цветущих растений было немного. Затем в начале июня начинают цвести некоторые "иэобильные" виды с большим количеством цветов (Salix spp. и Taraxacum officinale L.). соэдавая неограниченные эапасы пыльцы и нектара. Насекомые-опылители начинали лэбегать весенние цветы, привлекаясь больше к зтим "иэобильным" видам. Предполагается, что при цветении зтих "иэобильных' растений естественный отбор стимулирует зволюцию видов, цветущих раньше или поэднее "иэобильных" растений и конкурирующих с последними иэ-эа насекомых-опылителей. Установлено, что весной насекомые конкурируют иэ-эа цветов, но в период цветения иэобильных цветами растений (в течение всего лета) цветы конкурируют иэ-эа насекомых-опылителей.
Analysis of a 16-mm cinefilm of feeding bumblebees (Bombus) indicated that the glossa is the functional unit of proboscis length. This was corroborated by observations of the depth to which wild bees probed Impatiens biflora and by data presented in the literature. Accurate measurement of the glossa length of bees that died with their proboscides folded is facilitated by straightening the labium by dissection. Regression equations describing the relation between glossa and wing length, derived from bees from a single colony, provide sufficient accuracy to allow prediction of glossa length for that species in a field study.
This paper contrasts proximate physiological explanations for the relation between nectar production per flower, the number of flowers in an inflorescence and flower size with an evolutionary explanation based on competition for pollinators. Nine species in the Fabaceae with papilionaceous flowers and eight species in the Ericaceae with urceolate flowers were analysed separately. Intraspecific patterns of nectar production (production uncorrelated or positively correlated with inflorescence size and flower length) indicate that source-sink dynamics and variation in plant size and/or vigour primarily affect nectar secretion rates within species. Interspecific patterns (production per flower declines with increasing inflorescence size) are also most consistent with a physiological explanation, namely the packaging of a limited allocation of photosynthate. Selection on the competing species to offer shared pollinators equivalent rewards seems to have had little effect on interspecific patterns of nectar production. The frequency-dependent nature of pollinator attraction is probably the major constraint on evolutionary responses to competition for pollinators.
Butterfly conservation may require active management of reserves. The role of collectors in butterfly declines is probably overrated.
1. Daily patterns of activity, thoracic temperature (Tth) and thermoregulatory behaviour in relation to environmental conditions were studied in the European skipper Thymelicus lineola (Ochsenheimer) adults.
2. Daily activity was limited mainly by Tth, which in turn was dependent on air temperature (Ta) and sunlight. However, when light intensity fell to < 100 W m−2, skippers also became inactive.
3. Tth was sometimes as much as 12°C above Ta, and this was most pronounced under cool conditions when basking activity predominated.
4. Black globe temperature (Tbg) which, in the absence of wind, is influenced by both Ta, and radiant heat load in a manner similar to a basking butterfly, was used to approximate the maximum Tth attained by T.lineola through basking.
5. Both males and females basked at Tbg >20°C, and if this temperature was not attained, skippers remained inactive. As Tbg increased, basking activity declined more rapidly in males than in females. Basking bouts were also shorter in males.
6. Males also flew, fed and courted females at Tbg >20°C, while females only fed and flew at Tbg >22°C. The percentage of both sexes feeding and flying, and courting in the case of males, were positively correlated with Tbg.
7. In warm weather, males divided their time equally between flying and feeding, while females spent the majority of their time feeding. Feeding bouts were shorter and flying bouts were longer in males than in females. Flight duration in males was positively correlated with Tbg.
8. Skippers avoided, and, consequently laid fewer eggs in, shady areas.
9. Sexual differences in activity patterns in relation to weather reflect differences in the reproductive requirements of the two sexes.
I studied flower selection and foraging energetics of Agraulis vanillae L. (Nymphalidae) and Phoebis sennae (Pieridae), two butterfly species common to north central Florida. I identified the major nectar resources exploited by several populations of these butterflies and, for each plant species, measured available nectar volumes and concentrations, corolla lengths, and density. I quantified foraging behaviour of each butterfly species at each nectar source (flower visitation rate and percentage of foraging time in flight), and used these data to estimate the net rate of energy intake of each butterfly species at each nectar source.
Estimated mean energy contents of individual flowers of the eleven exploited plant species spanned three orders of magnitude, ranging between 0.015 and 9.27 joules. Mean energy content of individual flowers was strongly correlated with mean foraging profit of both butterfly species.
Mean nectar volume strongly influenced energy content and varied widely within and among species, ranging from 0.0076 to 1.853 μ1. Nectar concentration varied between 17.1% and 40.4% sucrose‐equivalents. Nectar volume was the best single predictor of foraging profitability (correlation coefficients of 0.994 and 0.984 for Phoebis and Agraulis respectively). Corolla length also strongly affected foraging profitability for both butterfly species; flower species with longer corollas were generally more profitable.
Flower density and nectar concentration showed weak or nonsignificant associations with foraging profitability.
The usefulness and limitations of these floral characteristics as bases for foraging selectivity, and the selective pressures foraging butterflies might place on the visited plants are discussed.
The time required for a bumble bee to visit a flower is affected by the length of the bee's glossa and its body weight, and by the depth of the flower and the volume of nectar it contains. Probing time is comprised of two components: access time and ingestion time. Access time increases linearly with flower depth, but ingestion time varies with flower depth only in flowers deeper than the length of the bee's glossa, due to a decline in the rate of ingestion of nectar. Probing time therefore increases gradually with increasing depth for flowers shallower than the bee's glossa, but beyond that depth it increases much more rapidly. The relation of probing time to flower depth influences the foraging efficiency and choice of flowers by bumble bees.
Thesis (Ph. D.) - University of Cambridge, 1982. 269pp.
This thesis is comprised of studies of life history variability and foraging ecology of bumblebees (Bombus spp.). Life history is investigated largely by means of regular censuses of the species inhabiting Wicken Fen National Nature Reserve and the Cambridge University Botanic Garden. Emergence from hibernation is related to temperature variables, and seasonal patterns of flight activity and pollen collection are described. Certain species pairs are shown of differ markedly in relative timing, others in the length of the colony cycle. Generally greater similarity is found between the phenology of a species in different years, than between species in the same year. Length of colony cycle is related to latitude, and two groups of species are distinguished: those in which the cycle increases with the length of the season, and those in which there is little change. Various lines of evidence are presented suggesting that B. hortorum and B. pratorum are sometimes bivoltine.
The other main area of study comprises aspects of foraging activity. Firstly, foraging is considered in relation to morphological factors, including flower arrangement, flower orientation and both inter- and infra-specific variation in tongue length. The existence of a relationship between worker tongue length and foraging activity is supported, within and between species. However, in the latter case the relationship depends upon the inclusion of a single very long-tongued species (B. hortorum). Species with similar tongue lengths are found to differ markedly in their use of different flower arrangements: evidence of a relationship between foraging behaviour and activity of the flight muscle enzymes is proposed as a possible explanation. Males generally visit shorter corollae, and make a larger proportion of visits to massed flowers, than their workers.
Secondly, foraging is related to physical characteristics of sugar solutions and nectar. There is no detectable tongue-length related variation in the concentration maximising rate of uptake of sugar solutions, but rate is dependent on body size. Nectar characteristics are considered in relation to flower morphology: a survey of 91 bumblebee visited plant species shows that nectar availability is positively correlated with corolla depth. This is considered as an explanation for correlations between tongue length and corolla depth. Bumblebees of all species are most often found collecting nectar between 30-50% w/w. Under conditions of fluctuating microclimate, corolla length is shown to influence the length of time that nectar remains within a given concentration range.
Thirdly, temporal variation in the distribution of foraging effort is considered as a function of nectar availability. When nectar per flower is relatively abundant in one or other of two alternative plant species, the observations correspond quite closely with a simple prediction that individuals select between flower types on the basis of maximising rate of energy gain. However, the correspondence between observed and predicted disappears when nectar is relatively scarce in both sources.
Lastly the foraging studies are drawn together by a model, designed to help clarify the relative importance of different components of a foraging trip in determining the overall rate of gain of energy by the colony (as opposed to rate of gain to the individual while sucking).
Consequences of central place foraging are shown to generate testable hypotheses concerning the assumption that energy maximisation is the criterion used when selecting between flower types. The thesis concludes with a short discussion of the findings in the context of the comparative ecology of the species.
A new statistical method called the phylogenetic regression is proposed that applies multiple regression techniques to cross-species data. It allows continuous and categorical variables to be tested for and controlled for. The new method is valid despite the problem that phylogenetically close species tend to be similar, and is designed to be used when information about the phylogeny is incomplete. Information about the phylogeny of the species is assumed to be available in the form of a working phylogeny, which contains multiple nodes representing ignorance about the order of splitting of taxa. The non-independence between species is divided into that due to recognized phylogeny, that is, to phylogenetic associations represented in the working phylogeny; and that due to unrecognized phylogeny. The new method uses one linear contrast for each higher node in the working phylogeny, thus applying the ‘radiation principle’. For binary phylogenies the method is similar to an existing method. A criterion is suggested in the form of a simulation test for deciding on the acceptability of proposed statistical methods for analysing cross-species data with a continuous y-variable. This criterion is applied to the phylogenetic regression and to some other methods. The phylogenetic regression passes this test; the other methods tested fail it. Arbitrary choices have to be made about the covariance structure of the error in order to implement the method. It is argued that error results from omitted but relevant variables, and the implications for those arbitrary choices are discussed. One conclusion is that the dates of splits between taxa, even supplemented by rates of neutral gene evolution, do not provide the ‘ true ’ covariance structure. A pragmatic approach is adopted. Several analytical results about the phylogenetic regression are given, without proof, in a mathematical appendix. A computer program has been written in GLIM to implement the phylogenetic regression, and readers are informed how to obtain a copy.
- R L H Dennis
Dennis, R. L. H., 1977. The British Butterflies. Their Origin and
Establishment. E. W. Classey Ltd, Faringdon.
Butterfly conservation on arable farmland
- R E H Feber
- Smith
Feber, R. E. & H. Smith, 1995. Butterfly conservation on arable
farmland. In: A. S. Pullin (ed), Ecology and Conservation of
Butterflies. Chapman & Hall, London, pp. 84-97.
Butterflies East of the Great Plains
- P A G O Opler
- Krizek
Opler, P. A. & G. O. Krizek, 1984. Butterflies East of the Great
Plains. The Johns Hopkins University Press, Baltimore.
Flowering phenology, flower colour and mode of reproduction of Prunus spinosa L
- K S R J Yeboah Gyan
- Woodell
Yeboah Gyan, K. & S. R. J. Woodell, 1987a. Flowering phenology, flower colour and mode of reproduction of Prunus spinosa
L. (blackthorn);
dog rose); and Rubus fruticosus L. (bramble) in Oxfordshire
- L Rosa Canina
Rosa
canina L. (dog rose); and Rubus fruticosus L. (bramble) in
Oxfordshire, England. Functional Ecology 1: 261-268.
Nectar production, sugar content
- K S R J Yeboah Gyan
- Woodell
Yeboah Gyan, K. & S. R. J. Woodell, 1987b. Nectar production,
sugar content, amino acids and potassium in Prunus spinosa L.,
Crataegus monogyna Jacq. and Rubus fruticosus L. at Wytham,
Oxfordshire. Functional Ecology 1: 251-259.