Is male puddling behaviour of tropical butterflies targeted at sodium for nuptial gifts or activity?

An apparent sexual difference in adult feeding behaviour in many species of Lepidoptera relates to puddling on mud, dung and carrion. In most butterfly species, puddling is exclusively a male behaviour. A possible explanation for this division in feeding behaviour is that nutrients derived from puddling are transferred to the female in the spermatophore during mating as a nuptial gift. Sodium derived from puddling has been shown to act as a nuptial gift in a few Lepidoptera species. It can also be used for neuromuscular activity in both males and females and may therefore correlate with flight morphology. In this study, we examine the generality of these two hypotheses in comparative work on a community of African fruit-feeding butterflies. We investigated puddling behaviour of males and females on carrion and dung together with sodium preferences, polyandry, relative wing-size, sexual size dimorphism and sodium concentrations in the bodies and spermatophores of several species. The results show that sodium as a nuptial gift can explain the sexual division in puddling in some species, but not in all. Species in which both sexes puddle transfer little sodium in the nuptial gift, which is consistent with the nuptial gift theory. Wing loading and puddling are not significantly correlated, but the trend followed the direction predicted by the activity hypothesis. However, the sodium concentration in the species with the smallest wing area to thoracic volume (WA/TV) ratio (the largest Charaxes spp.), was relatively low. Moreover, in all investigated species, the sodium concentration was higher in the abdomen than in the thorax. The results are discussed in the light of differences between the sexes in foraging behaviour in both larvae and adults, and with respect to alternative explanations for puddling.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 345–361.     

Mouthparts and feeding of certain ground beetles (Goleoptera: Garabidae)

The feeding apparatus of 17 species of ground beetles (Carabidae), representing 17 genera and 12 tribes has been studied, and the observations correlated with gut content analyses and the ability of certain species to regurgitate pre-oral digestive fluids. The diverse feeding habits of species investigated include zoophagous fluid feeders, zoophagous fragmentary feeders and mixed feeders. In the last-named group, the food intake may be both fluid and fragmentary, and either of animal or of both animal and plant origin.     

Temporal changes in floral resource availability and flower visitation in a butterfly

Foraging affects survival and reproductive success in animals, including flower-visiting insects. Plant-derived floral food resources (i.e. nectar and pollen) may be rapidly changing in space and time and pollinators may need to quickly switch to new resources. Butterflies are suitable model organisms to investigate foraging behaviour of insect pollinators, because they can be easily monitored under natural conditions. We studied flower visitation patterns in the Clouded Apollo butterfly Parnassius mnemosyne in relation to the abundance of available floral resources. We recorded flower visitation daily in individually marked butterflies, listed flowering species and estimated flower abundance categories every 3 days in a single meadow, during five consecutive flight periods. Butterflies visited 35 nectar plants from the 71 species available. Few nectar plants were frequently visited (visit ratios for the annually most visited species: 37–60%), many were scarcely visited and no visits were observed on several abundant species. Flower abundance and visit ratio varied among years and within flight periods. The number of visits increased with flower abundance in the seven most frequently visited plant species, but not in the occasionally visited ones. Beside their choosiness, Parnassius mnemosyne butterflies were able to adjust foraging behaviour to rapidly changing resource distributions. Diet selectivity in adults might increase the vulnerability of this species. However, visitation plasticity may mitigate the effect of the lack of some nectar plants, as complementary resources can be used as alternatives.     

Coevolution between flight morphology,vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight‐related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit‐feeding butterflies sampled in seven sites across 3341 ha. We used wing–thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing–thorax ratio in both sexes. Stouter fast‐flying species preferred the canopy, whereas more slender slow‐flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait–habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms.     

Chromosomal evolution in the South American Riodinidae (Lepidoptera: Papilionoidea)

We give the haploid chromosome numbers of 173 species or subspecies of Riodinidae as well as of 17 species or subspecies of neotropical Lycaenidae for comparison. The chromosome numbers of riodinids have thus far been very poorly known. We find that their range of variation extends from n =?9 to n =?110 but numbers above n =?31 are rare. While lepidopterans in general have stable chromosome numbers, or variation is limited at most a subfamily or genus, the entire family Riodinidae shows variation within genera, tribes and subfamilies with no single modal number. In particular, a stepwise pattern with chromosome numbers that are about even multiples is seen in several unrelated genera. We propose that this variation is attributable to the small population sizes, fragmented populations with little migration, and the behavior of these butterflies. Small and isolated riodinid populations would allow for inbreeding to take place. Newly arisen chromosomal variants could become fixed and contribute to reproductive isolation and speciation. In contrast to the riodinids, the neotropical Lycaenidae (Theclinae and Polyommatinae) conform to the modal n =?24 that characterizes the family.     

甘肃太统一崆峒山国家自然保护区的蝶类群落特征

2006--2010年期间,对太统一崆峒山国家级自然保护区的蝶类进行了研究。采集到蝶类标本共计4800余号,经过整理鉴定出蝶类230种及亚种,隶属10科24亚科126属。其中36属71种为优势类群,占总种数的30．87％;眼蝶科有46种,占20．0％;灰蝶科有37种,占16．1％,为次优势类群;喙蝶科、蚬蝶科为单属单种,属于罕见类群;其他各科为常见类群。在230种蝶类中,属于古北界的种类占总种数的26．5％,属于东洋界的种类占总种数的5．7％,两界兼有种类和广布种类占总种数的67．8％,即广布种类成分最多,东洋界成分最少。230种蝶类隶属126属,其中单种属有75个,占总属的59．5％,包含的种类占总种数的32．6％;多种属有51个,占总属的40．5％,包含的种类数有155种,占总种数的67．4％;属种比值系数为0．548。结果表明,该地区蝶类区系以单种属为主。     

横断山区的蝶类资源与珍稀蝴蝶的保护

横断山区的蝶类561种,隶属于12科,208属.在12科中,种类最多的是蛱蝶科148种,其次是眼蝶科117种,灰蝶科65种,凤蝶科63种,粉蝶科60种,弄蝶科60种,蚬蝶科18种,绢蝶科12种,斑蝶科9种,环蝶科6种,喙蝶科2种,珍蝶科1种.分布于横断山区的珍稀蝴蝶有44种,其中国家Ⅰ级保护的1种,Ⅱ级保护的3种.     

Feeding responses of adult butterflies, Nymphalis xanthomelas, Kaniska canace and Vanessa indica, to components in tree sap and rotting fruits: synergistic effects of ethanol and acetic acid on sugar responsiveness

Exuded tree sap and rotting fruits are important feeding sources for adults of many butterfly species. However, the chemical composition of such rotting foods and the effects of these constituents on butterfly feeding behaviour have rarely been investigated. Chemical analyses revealed that these food sources contain several major components, including fructose, glucose, ethanol and acetic acid. Moreover, 15 amino acids were identified from the exuded tree sap. The gustatory responsiveness to these components was examined in the proboscis of nymphalid adults of three species (Nymphalis xanthomelas, Kaniska canace and Vanessa indica). The first two species primarily feed on sap and rotting fruits, while V. indica frequently visits flowers. Fructose elicited larger feeding responses from these saprophagous butterflies than glucose, which had a feeding-stimulatory effect only on N. xanthomelas at a concentration equivalent to that of the food. Ethanol, acetic acid and the five major amino acids identified in tree sap did not elicit feeding responses by themselves. However, ethanol and/or acetic acid, when mixed with these sugars at equivalent concentrations, evoked feeding and enhanced probing responses in K. canace and V. indica. These results suggest that ethanol and acetic acid, together with sugars at low concentrations, synergistically stimulate butterfly feeding behaviour.     

Functional constraints on the evolution of long butterfly proboscides: lessons from Neotropical skippers (Lepidoptera: Hesperiidae)

Extremely long proboscides are rare among butterflies outside of the Hesperiidae, yet representatives of several genera of skipper butterflies possess proboscides longer than 50 mm. Although extremely elongated mouthparts can be regarded as advantageous adaptations to gain access to nectar in deep‐tubed flowers, the scarcity of long‐proboscid butterflies is a phenomenon that has not been adequately accounted for. So far, the scarceness was explained by functional costs arising from increased flower handling times caused by decelerated nectar intake rates. However, insects can compensate for the negative influence of a long proboscis through changes in the morphological configuration of the feeding apparatus. Here, we measured nectar intake rates in 34 species representing 21 Hesperiidae genera from a Costa Rican lowland rainforest area to explore the impact of proboscis length, cross‐sectional area of the food canal and body size on intake rate. Long‐proboscid skippers did not suffer from reduced intake rates due to their large body size and enlarged food canals. In addition, video analyses of the flower‐visiting behaviour revealed that suction times increased with proboscis length, suggesting that long‐proboscid skippers drink a larger amount of nectar from deep‐tubed flowers. Despite these advantages, we showed that functional costs of exaggerated mouthparts exist in terms of longer manipulation times per flower. Finally, we discuss the significance of scaling relationships on the foraging efficiency of butterflies and why some skipper taxa, in particular, have evolved extremely long proboscides.     

福州国家森林公园蝶类名录(Ⅱ)

本文鉴定出福州国家森林公园蝶类共计 1 0科、1 2 0属、2 0 4种 ,分载名录 ( )、( )篇。名录 ( )作为前者的续篇 ,包含 4科、6 6属、86种。名录 ( )已见本刊 1 0 ( 2 )。红蛱蝶属 Vanessa Fabricius1 1 9小红蛱蝶 V. cardui ( Linnaeus)成虫 6、9月可见。寄主 :荨麻属、水麻属、飞廉属等。1 2 0大红蛱蝶 V.indica( Herbst)成虫 2、5- 6、9- 1 2月可见。寄主 :苎麻、密花苎麻、荨麻属植物等。琉璃蛱蝶属 Kaniska Moore1 2 1琉璃蛱蝶 K.canace( Linnaeus)成虫 1 - 5、9- 1 2月可见。寄主 ;尖叶菝葜、圆锥菝葜、菝葜、马甲菝葜、穿鞘菝…     

Flight morphology of Neotropical butterflies: palatability and distribution of mass to the thorax and abdomen

Summary We test whether palatability of Neotropical butterflies is associated with distribution of mass to the thorax and abdomen. Thoracic mass is predominantly muscle mass, whereas abdominal mass includes organs of digestion, food storage, and reproduction. To escape from predation, butterflies palatable to the rufous-tailed jacamar (Galbula ruficauda) use fast, erratic flight, whereas unpalatable butterflies have defensive chemicals and slow, regular flight patterns. We adjusted for effects of phylogeny and report partial correlations for two levels of analysis: 1) comparisons among-lineage means, which test for correlations between traits of distantly related lineages, and 2) comparisons among deviations from lineage means (or within lineages), which test for correlations between traits of more closely related species.Among lineages for both males (n=10 lineages) and females (n=9), palatability and thoracic mass were positively correlated, whereas palatability and abdominal mass were negatively correlated. An inverse correlation between thoracic and abdominal mass is a consequence of the two segments composing 75% of the total body mass. Predation, indexed by palatability, may select for increased flight speed and thoracic mass at the expense of the abdomen, but relative flight speed and thoracic mass were not significantly correlated.Within lineages (n=45 species for each sex), thoracic mass was uncorrelated with palatability in both sexes. Relative flight speed correlated positively with thoracic mass and negatively with body mass. Palatability and abdominal mass were negatively correlated for males but not females. Hence differences between the sexes in mass distribution suggest differences in reproductive constraints and predation stress.     

Butterfly species diversity and their floral preferences in the Rupa Wetland of Nepal

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 (herbaceous 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.     

Flight behaviour of seabirds in relation to wind direction and wing morphology

We studied flight direction relative to wind direction (hereafter referred to as "flight direction"), the relation between wing morphology and flight behaviour and interspecies relationships in flight behaviour among all major seabird taxa. We calculated wing loading and aspect ratios for 98 species from 1029 specimens. Species were sorted into 13 groups on the basis of similarity in patterns of flight direction. The primary flight direction of Pelecaniformes and Charadriiformes was into and across headwinds. The most common flight direction of Procellariiformes was across wind. Seabirds avoided flying with tailwinds. Wing loading and aspect ratios were positively correlated in Procellariiformes, Pelecaniformes and alcids but negatively correlated in larids. In Procellariiformes, incidence of headwind flight and that of tailwind flight were significantly correlated with wing loading and aspect ratio; species with higher wing loading and aspect ratios flew more often into headwinds and less often with tailwinds. In contrast, the proportion of Pelecaniformes and Charadriiformes flying with tailwinds increased significantly with increased wing loading. Our results demonstrate a close link in seabirds between flight behaviour, wing morphology and natural history patterns in terms of distribution, colony location, dispersal and foraging behaviour.     

Pollen histochemistry and pollen : ovule ratios in Zingiberaceae

BACKGROUND AND AIMS: Pollen grains of 37 species from 11 genera in the family Zingiberaceae were examined to assess qualitatively starch or lipid contents; the pollen grain and ovule numbers per flower and pollen : ovule ratios were also counted and calculated. Pollen : ovule ratios were studied to reveal patterns of variation in the Zingiberaceae. METHODS: Freshly open flowers with dehiscing anthers were collected at random from plants growing in natural habitats or in botanical gardens. Presence of lipids or starch in pollen grains was tested by Sudan solution and IKI solution, respectively, and examined under a microscope. To estimate the pollen and ovule numbers per flower, one anther from each bud was carefully dissected and all pollen grains were counted; ovaries were carefully dissected out of each flower and counted. Whenever possible, at least 10-15 buds were used in the determination. KEY RESULTS: Thirty-three of all the 37 species examined had starchy pollen. Starch was not found in only four species and lipid was not found in only one species; among the four tribes in subfamily Zingiberoideae, all species of Zingibereae and Globbeae had pollen with no starch, Alpineae and Hedychieae had pollen with and without starch, whereas, all species of subfamily Costoideae had starchy pollen with abundant lipids. The mean pollen : ovule ratios in the members of the Zingiberaceae investigated range from 3.25 +/- 1.56 to 616.52 +/- 117.83. CONCLUSIONS: The pollen nutrition types seemed not related to mating systems. The pollen : ovule ratios in members of the Zingiberaceae with the same breeding system are noticeably lower than those recorded by previous authors. The lower pollen : ovule ratios in this family are presumed to be related to the highly efficient pollination systems, mediated by pollen which can be quite glutinous and the relatively large stigma area. In most of the Alpinia species the anaflexistylous flowers have much larger numbers of pollen grains and higher pollen : ovule ratios than the cataflexistylous flowers. There are significant differences in mean pollen grain numbers and pollen : ovule ratios between different life forms but ovule numbers are approximately the same.     

SEXUAL SELECTION AND BUTTERFLY DESIGN—A COMPARATIVE STUDY

Temperate butterflies of 44 species were examined to determine if their mating system (perching and patrolling) affected flight design. To control for spurious effects due to ancestry, 25 of these species were assigned to eight contrasts within which a change in mating system had occurred. In perching species sexual selection was predicted to favor traits associated with high acceleration ability and speed, while in patrolling species traits associated with flight endurance were predicted. In conformance with these expectations males of perching species had larger thorax/body mass ratios, higher wing loadings, and higher aspect ratios than patrolling species. The male mating system affected females in the same direction in the same variables as males. This could be explained by a genetic correlation with males. When removing the covariance between the sexes, only male design was explained by the mating system. The mating system was also associated with different degrees of sexual dimorphism in wing size. This supported the hypothesis that male design was affected by the mating system.     

Relationships between morphological parameters in birds with different flying habits

1. Data on morphological and physiological parameters from 346 species of birds were collected from diverse sources. 2. The observed relationships among some pairs of variables showed differences between the five flight styles into which the studied birds were classified. 3. The characteristic kind of flight for each species is related to four easily obtainable non-invasive parameters: body weight (BW), wing area (WA), wing span (WS) and boyd length (BL). 4. Wing-loading index (BW2/3/WA) gives the most effective variable to describe the flying habits of any species.     

新疆东部天山蝶类多样性及其垂直分布

2006-2008年研究了新疆东部天山蝶类多样性和垂直分布.结果表明:研究区域内共记录蝴蝶7科43属63种,占新疆已记录蝶类种数的24.80％,区系组成主要是古北种,占73％;其次是广布种,占27％,没有发现东洋种.其中蛱蝶科的物种数最多,为11属19种,蚬蝶科的物种数最少,只有1属1种.按海拔将生境分为5个垂直自然带,包括低山灌木草原带、山地森林草原带、亚高山草甸带、高山草甸带、垫状植被带.蝶类物种数和个体数排序为亚高山草甸带＞山地森林草原带＞低山灌木草原带＞高山草甸带＞垫状植被带.采用Shannon-Wiener指数和G-F指数对蝶类物种和科、属的多样性进行了分析评价,结果显示亚高山草甸带的蝶类多样性最为丰富,其次是山地森林草原带和低山灌木草原带,而高山草甸带和垫状植被带的蝶类多样性相对较低,物种和科、属多样性分析结果均一致.蝶类垂直分布明显,物种数和个体数随海拔变化的趋势类似,均为先增加后下降.蝶类区系成分随着海拔升高发生改变,广布种的比例逐渐降低,高山草甸带和垫状植被带只有古北种分布.研究结果显示,生境改变对蝴蝶群落影响明显,保护生境是保护蝴蝶生存的最主要措施.     

The role of size and dominance in the feeding behaviour of coexisting hummingbirds

Interspecific competition can strongly influence community structure and limit the distribution and abundance of species. One of the main factors that determine hummingbird community structure is competition for food. The temporal and spatial distribution of nectar has a strong impact on hummingbird assemblages, shaping foraging niches according to hummingbird dominance and foraging strategy. We investigated whether body size and the degree of aggressive dominance influence feeding behaviour of hummingbirds in a temperate forest in northwestern Mexico (El Palmito, Mexico) when winter migrant hummingbirds are present in the community. First, we determined the dominance status of hummingbirds and evaluated the relationship between dominance and body mass, wing disc loading and migratory status. Secondly, we determined how hummingbird species used plant species differently. Thirdly, we examined whether the most dominant hummingbird species defended floral patches with more energy and/or with a larger number of flowers. At each flower patch, hummingbird species, number of hummingbird interactions, feeding time and number of flowers present were recorded. The total number of calories available within each floral patch was also determined. Our results demonstrate that the dominance hierarchy of 13 hummingbird species (migratory and resident) was correlated with body size but not wing disc loading, and that members of the hummingbird community showed a clear separation in resource use (by plant species). Hummingbirds at the top of the dominance hierarchy defended and fed on the best flower patches, defined by the quantity of calories available. Hence, the feeding behaviour of hummingbirds at El Palmito depends on the abundance of plant species used by hummingbirds and on the amount of energy available from each flower patch. Thus, hummingbird body size, aggressive dominance and defence of quality flower patches determines niche partitioning among species.     

Butterfly visual characteristics and ontogeny of responses to butterflies by a specialized tropical bird

The responses of two adult and three hand-reared, naive young rufous-tailed jacamars (Galbula ruficauda) to local butterflies were studied in feeding experiments. Four behavioural characteristics distinguish jacamars from other less specialized avian predators: (1) Exposed to butterflies for the first time, naive young jacamars would attack butterflies without showing signs of inhibition. Unacceptable butterflies, once captured, were taste-rejected quickly, and most survived the sampling. The few presumably unacceptable butterflies consumed by the birds were not observed to cause vomiting. (2) After gaining some familiarity with butterflies, young birds, like the adults, developed a reluctance to attack. They visually rejected certain classes of butterflies, often failing to attack them during an entire four-hour feeding trial. However, occasional attacks were made on butterflies in these ‘rejected’ classes. When this did occur, the insects proved to be actually easier to catch than those that were more often attacked. Once captured, however, the majority of these butterflies were taste-rejected. (3) For a given butterfly species, most individuals were either consumed or rejected. Thus, each species could be clearly classified as either acceptable or unacceptable to the jacamars. This consistency in jacamar responses resulted in a bimodal acceptability distribution of sympatric butterflies. (4) Young jacamars were capable of rapid associative learning and their responses were closely associated with butterfly visual characteristics in which colour pattern, flight behaviour, and morphology were also closely correlated. Thus, a single butterfly morphological parameter termed body shape (body length/thoracic diameter ratio) can adequately predict the feeding responses of jacamars. Visually detectable traits associated with butterflies possessing chemical defences may represent a balance between the need to signal unambiguously to specialized and/or experienced predators and the need to escape attacks by generalized and/or opportunistic predators. Since the proportion of specialized predators is higher in the tropical rainforest than in other habitat types, we expect greater divergence of morphological and behavioural characteristics between palatable and unpalatable butterflies in rainforest habitats.     

Does Skipping a Meal Matter to a Butterfly's Appearance? Effects of Larval Food Stress on Wing Morphology and Color in Monarch Butterflies

In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success.