Exploratory comparison of flower visiting behavior and pollination ability of mason bees, bumble bees, and honey bees.

This study explored the flower visiting behaviors and pollination abilities of mason bees (Osmia excavata Alfken (Hymenoptera: Megachilidae)), bumble bees (Bombus terrestris (Linnaeus, 1758) (Hymenoptera: Apidae)), and Italian honey bees (Apis mellifera ligustica Spinola (Hymenoptera: Apidae)) in apple orchards in early spring in Jinan (located in the central region of Shandong) and Yantai (located in the Peninsula of Shandong). We compared the pollen collection patterns, flower visiting behavior, flying speed, and effects on apple pollination of the 3 types of bees. The frequencies of flower visits were significantly higher for mason bees (12.89/min in Jinan and 10.63/min in Yantai) than bumble bees and Italian honey bees in the 2 regions. The single flower residence times were significantly higher for Italian honey bees (8.22 s in Jinan and 9.43 s in Yantai), but Italian honey bees were most affected by the climate. The 3 bees differed significantly in terms of the amount of apple pollen collected and their effects on the fruit setting rate in apples (mason bees > bumble bees > Italian honey bees). The results showed that the mason bee was the most suitable pollinating species for spring apple orchards; Bumble bees were more suitable as alternative pollinators during cloudy and low temperatures; Italian honey bees were able to take advantage of their large number of worker bees in sunny and warm weather. Compared to individual bee species, a combination of 2 or 3 species of bees might be more advantageous in dealing with complex and variable weather conditions.

Foraging Behavior of Two Pollen Wasp Species of the Genus Celonites Latreille, 1802 (Hymenoptera: Vespidae: Masarinae), from the Altai Mountains.

Celonites kozlovi Kostylev, 1935, and C. sibiricus Gusenleitner, 2007, coexist in semi-deserts of the Altai Mountains. The trophic relationships of these pollen wasp species to flowers are largely unknown. We observed the flower visits and behaviors of wasps on flowers; pollen-collecting structures of females were studied using SEM; the taxonomic position of these two species was ascertained with the barcoding sequence of the mitochondrial COI-5P gene. Celonites kozlovi and C. sibiricus form a clade together with C. hellenicus Gusenleitner, 1997, and C. iranus Gusenleitner, 2018, within the subgenus Eucelonites Richards, 1962. Celonites kozlovi is polylectic in the narrow sense, collecting pollen from flowers of plants belonging to five families (with the predomination of Asteraceae and Lamiaceae) using diverse methods for both pollen and nectar uptake. In addition, this species is a secondary nectar robber, which has not been observed in pollen wasps before. The generalistic foraging strategy of C. kozlovi is correlated with an unspecialized pollen-collecting apparatus on the fore-tarsi. In contrast, C. sibiricus is broadly oligolectic, predominantly collecting pollen from flowers of Lamiaceae. Its specialized foraging strategy is associated with apomorphic behavioral and morphological traits, particularly specialized pollen-collecting setae on the frons, which enable indirect pollen uptake using nototribic anthers. These adaptations in C. sibiricus evolved independently of similar specializations in the Celonites abbreviatus-complex. Celonites kozlovi is re-described, and males are described for the first time.

Conserving diversity in Irish plant-pollinator networks.

Beneficial insects provide valuable services upon which we rely, including pollination. Pollinator conservation is a global priority, and a significant concern in Ireland, where over half of extant bee species have declined significantly in recent decades. As flower-visiting insects rely on flowering plants, one way to conserve and promote pollinator populations is to protect high-quality habitat. We analyzed the structure of insect-flower interactions from multiple habitat categories in a large database of interactions from Ireland. Our primary goals were to compare spatial and temporal variation in Irish network structures, compare Irish networks to published networks from other countries, and provide evidence-based recommendations for pollinator conservation in Ireland by identifying well-visited plant species that may promote high pollinator diversity, abundance, and functional complementarity. Habitat types within Ireland differed substantially: seminatural grasslands had the highest pollinator species richness and largest number of unique pollinator species, while intensively managed habitats exhibited negative asymmetry (more plant than pollinator species). This negative asymmetry is notable because most plant-pollinator networks exhibit a positive asymmetry. Within intensively managed habitats, agricultural and urban habitats differed. Urban habitats had the highest number of non-native plant species while agricultural habitats had the lowest pollinator species richness. We also found Irish networks varied across the growing season, where July had the highest plant and insect species richness. When comparing Irish networks to published networks from other countries, we found Irish networks had a higher ratio of plant species to pollinator species, and that this difference was most evident in agricultural habitats. This ratio means the typical network asymmetry (more pollinator than plant species) was flipped (more plant than pollinator species) in the Irish network. We conclude that conserving seminatural grasslands in Ireland will be an essential component of pollinator conservation and identify thirty-five plant species important for restoring seminatural habitats.

Predators or Herbivores: Cockroaches of Manipulatoridae Revisited with a New Genus from Cretaceous Myanmar Amber (Dictyoptera: Blattaria: Corydioidea).

Manipulator modificaputis Vrsanský and Bechly, 2015 (Manipulatoridae, Corydioidea) is a purported predatory cockroach from Cretaceous Myanmar amber, based on a single male. It is distinctive by the nimble head, elongate pronotum and legs, and particularly by the extraordinarily long maxillary palpi. In the present study, we redescribe Manipulator modificaputis based on six new fossils including males and females, and comment on the original description. The closely related Manipulatoides obscura gen. & sp. nov. is proposed on the basis of five fossils, including males and females. It differs from Manipulator in weaker spination of the legs, including the type-C forefemoral spination instead of the type-A of Manipulator. Some undetermined adults and nymphs are also described. We discuss the ethology of Manipulatoridae and speculate that they might feed on flowers. They are unlikely to be specialized predators since they lack necessary weaponry for capturing prey; in contrast, their unique morphotype appears to be suitable for efficient foraging and locomotion amid flowering twigs. The possibility of being kleptoparasites of the spider-web is also discussed. In addition, regenerated four-segmented tarsi are found from the new species.

Biology and ecology of the Oriental flower-breeding Drosophila elegans and related species.

Animals adapt to their environments in the course of evolution. One effective approach to elucidate mechanisms of adaptive evolution is to compare closely related species with model organisms in which knowledge of the molecular and physiological bases of various traits has been accumulated. Drosophila elegans and its close relatives, belonging to the same species group as the model organism D. melanogaster, exhibit various unique characteristics such as flower-breeding habit, courtship display, territoriality, sexual dimorphism, and colour polymorphism. Their ease of culturing and availability of genomic information makes them a useful model for understanding mechanisms of adaptive evolution. Here, we review the morphology, distribution, and phylogenetic relationships of D. elegans and related species, as well as their characteristic flower-dependent biology, food habits, and life-history traits. We also describe their unique mating and territorial behaviours and note their distinctive karyotype and the genetic mechanisms of morphological diversity that have recently been revealed.

CropPol: A dynamic, open and global database on crop pollination.

Seventy five percent of the world's food crops benefit from insect pollination. Hence, there has been increased interest in how global change drivers impact this critical ecosystem service. Because standardized data on crop pollination are rarely available, we are limited in our capacity to understand the variation in pollination benefits to crop yield, as well as to anticipate changes in this service, develop predictions, and inform management actions. Here, we present CropPol, a dynamic, open, and global database on crop pollination. It contains measurements recorded from 202 crop studies, covering 3,394 field observations, 2,552 yield measurements (i.e., berry mass, number of fruits, and fruit density [kg/ha], among others), and 47,752 insect records from 48 commercial crops distributed around the globe. CropPol comprises 32 of the 87 leading global crops and commodities that are pollinator dependent. Malus domestica is the most represented crop (32 studies), followed by Brassica napus (22 studies), Vaccinium corymbosum (13 studies), and Citrullus lanatus (12 studies). The most abundant pollinator guilds recorded are honey bees (34.22% counts), bumblebees (19.19%), flies other than Syrphidae and Bombyliidae (13.18%), other wild bees (13.13%), beetles (10.97%), Syrphidae (4.87%), and Bombyliidae (0.05%). Locations comprise 34 countries distributed among Europe (76 studies), North America (60), Latin America and the Caribbean (29), Asia (20), Oceania (10), and Africa (7). Sampling spans three decades and is concentrated on 2001-2005 (21 studies), 2006-2010 (40), 2011-2015 (88), and 2016-2020 (50). This is the most comprehensive open global data set on measurements of crop flower visitors, crop pollinators and pollination to date, and we encourage researchers to add more datasets to this database in the future. This data set is released for non-commercial use only. Credits should be given to this paper (i.e., proper citation), and the products generated with this database should be shared under the same license terms (CC BY-NC-SA).

Biodiversity Survey of Flower-Visiting Spiders Based on Literature Review and Field Study.

Many arthropods exhibit flower-visiting behavior, including a variety of spider species. However, as spiders are assumed to be strictly predatory, flower-visiting spiders are an often neglected group. We conducted a systematic biodiversity study of flower-visiting spiders based on published papers and field surveys. Most previous studies have focused on the herbivorous behavior of flower-visiting spiders (nectivory or pollinivory) and their effects on host flowers (tritrophic interactions with flower-visiting insects). In our field survey, we utilized standard transect walks (active sampling) and colored pan traps (passive sampling) to investigate species occurrence, diurnal and seasonal variation, and flower color preference of flower-visiting spiders. From the transect walks, crab spider species were found to be the dominant flower-visiting spiders and, based on all spider species, juvenile visitors were significantly more common than adults. Furthermore, in terms of spider number and species richness, tulips were the preferred flower to visit. For the pan traps, wolf spiders were found to be the dominant spider species. No significant differences were observed in the number of spiders caught in different colored pans, suggesting that color may not be an important flower trait in regard to spider preference. To the best of our knowledge, this study is the first to propose the term 'flower-visiting spiders' and conduct a systematic investigation of their diversity. However, this is preliminary research and further studies are required, especially as biodiversity is often closely linked to survey sites and ecotopes.

1. Generally speaking, flower-visiting spiders focuses on the herbivorous behavior (nectivory or pollinivory) and their effects on host flowers (tritrophic interactions with flower-visiting insect). 2. The authenticity of flower-visiting spiders collected by active sampling was higher than passive trapping; juvenile spider visitors were significantly more common than adults. 3. Our study is the first to propose the term 'flower-visiting spiders', and further research is needed.

Inter-Individual Nectar Chemistry Changes of Field Scabious, Knautia arvensis.

Nectar is crucial to maintain plant-pollinator mutualism. Nectar quality (nutritional composition) can vary strongly between individuals of the same plant species. The factors driving such inter-individual variation have however not been investigated closer. We investigated nectar quality of field scabious, Knautia arvensis in different grassland plant communities varying in species composition and richness to assess whether nectar quality can be affected by the surrounding plant community. We analyzed (with high performance liquid chromatography) the content of carbohydrates, overall amino acids, and essential amino acids. Amino acid and carbohydrate concentrations and proportions varied among plant individuals and with the surrounding plant community but were not related to the surrounding plant species richness. Total and individual carbohydrate concentrations were lowest, while proportions of the essential amino acids, valine, isoleucine, leucine (all phagostimulatory), and lysine were highest in plant species communities of the highest diversity. Our results show that K. arvensis nectar chemistry varies with the composition of the surrounding plant community, which may alter the taste and nutritional value and thus affect the plant's visitor spectrum and visitation rate. However, the strong inter-individual variation in nectar quality requires additional studies (e.g., in semi-field studies) to disentangle different biotic and abiotic factors contributing to inter-individual nectar chemistry in a plant-community context.

Aurum, a new monotypic genus of Cetoniinae (Coleoptera: Scarabaeidae) endemic to Western Australia, Australia.

The flower beetle genus Aurum new genus (Coleoptera: Scarabaeidae: Cetoniinae: Schizorhinini: Schizorhinina) is described. Aurum is endemic to Australia, and contains a single species, Aurum leonorensis new species, from Western Australia. The genus is compared to the most closely related Australian flower beetle genera, Diaphonia Newman, 1840, Aphanesthes Kraatz, 1880, and Chondropyga Kraatz, 1880. Ecological observations and the collecting localities are detailed for Aurum leonorensis.

Refuges from fire maintain pollinator-plant interaction networks.

Fire is a major disturbance factor in many terrestrial ecosystems, leading to landscape transformation in fire-prone areas. Species in mutualistic interactions are often highly sensitive to disturbances like fire events, but the degree and complexity of their responses are unclear. We use bipartite insect-flower interaction networks across a recently burned landscape to explore how plant-pollinator interaction networks respond to a recent major fire event at the landscape level, and where fire refuges were present. We also investigate the effectiveness of these refuges at different elevations (valley to hilltop) for the conservation of displaced flower-visiting insects during fire events. Then, we explore how the degree of specialization of flower-visiting insects changes across habitats with different levels of fire impact. We did this in natural areas in the Greater Cape Floristic Region (GCFR) biodiversity hotspot, which is species rich in plants and pollinators. Bees and beetles were the most frequent pollinators in interactions, followed by wasps and flies. Highest interaction activity was in the fire refuges and least in burned areas. Interactions also tracked flower abundance, which was highest in fire refuges in the valley and lowest in burned areas. Interactions consisted mostly of specialized flower visitors, especially in refuge areas. The interaction network and species specialization were lowest in burned areas. However, species common to at least two fire classes showed no significant difference in species specialization. We conclude that flower-rich fire refuges sustain plant-pollinator interactions, especially those involving specialized species, in fire-disturbed landscape. This may be an important shelter for specialized pollinator species at the time that the burned landscape goes through regrowth and succession as part of ecosystem recovery process after a major fire event.

First record of the genus Empidideicus Becker, 1907 (Diptera, Mythicomyiidae) in China and the Oriental Region, with description of a new species.

Empidideicus Becker, 1907 is newly recorded from China and the Oriental Region, with one new species, E.pentagonius sp. n., described and illustrated. Observations are provided on the biology of E.pentagonius sp. n. visiting flowers of Stemonamairei (Levl.) Krause (Liliflorae, Stemonaceae). A key to the genera of Mythicomyiidae known to occur in China is provided.

The Lordiphosa denticeps species group (Diptera: Drosophilidae) in China, with redescriptions of four known species and descriptions of nine new species.

Based on specimens collected from Yunnan, Xizang, and Taiwan in China, nine new species of the Lordiphosa denticeps species group, L. mikioides sp. nov., L. kimurai sp. nov., L. anthophilia sp. nov., L. yangi sp. nov., L. tibetica sp. nov., L. medogensis sp. nov., L. hamatispina sp. nov., L. secula sp. nov., and L. spatulata sp. nov., were described and four known species, L. denticeps (Okada Sasakawa), L. neokurokawai (Singh Gupta), L. ramula Zhang, and L. tripartita (Okada), were redescribed. In addition, we provided a key to all species of this species group. Males of three new species, L. mikioides, L. kimurai, and L. anthophilia, have distinct sex-combs consisting of black, stout teeth on the 1st and 2nd tarsomeres of foreleg; the large, longitudinal sex-comb on the 1st tarsomere is similar to those seen in the L. miki species group and the subgenus Sophophora Sturtevant of the genus Drosophila Fallén. Two of these and another new species were collected at flowering Impatiens L. (Balsaminaceae) stands: all specimens of L. anthophilia and L. medogensis directly from flowers, and some specimens of L. kimurai by net sweeping. The presence of large, longitudinal sex-combs and the flower-visiting habit were reported for the first time in the L. denticeps group.

Learning in two butterfly species when using flowers of the tropical milkweed Asclepias curassavica: No benefits for pollination.

PREMISE OF THE STUDY: The ability of insect visitors to learn to manipulate complex flowers has important consequences for foraging efficiency and plant fitness. We investigated learning by two butterfly species, Danaus erippus and Heliconius erato, as they foraged on the complex flowers of Asclepias curassavica, as well as the consequences for pollination. METHODS: To examine learning with respect to flower manipulation, butterflies were individually tested during four consecutive days under insectary conditions. At the end of each test, we recorded the number of pollinaria attached to the body of each butterfly and scored visited flowers for numbers of removed and inserted pollinia. We also conducted a field study to survey D. erippus and H. erato visiting flowers of A. curassavica, as well as to record numbers of pollinaria attached to the butterflies' bodies, and surveyed A. curassavica plants in the field to inspect flowers for pollinium removal and insertion. KEY RESULTS: Learning improves the ability of both butterfly species to avoid the nonrewarding flower parts and to locate nectar more efficiently. There were no experience effects, for either species, on the numbers of removed and inserted pollinia. Heliconius erato removed and inserted more pollinia than D. erippus. For both butterfly species, pollinium removal was higher than pollinium insertion. CONCLUSION: This study is the first to show that Danaus and Heliconius butterflies can learn to manipulate complex flowers, but this learning ability does not confer benefits to pollination in A. curassavica.

Most ornamental plants on sale in garden centres are unattractive to flower-visiting insects.

BACKGROUND: Gardeners and park managers seeking to support biodiversity in urban areas often plant ornamentals attractive to flower-visiting insects. There is a huge diversity of garden plant varieties, and some recommendations are available as to which are attractive to insects. However, these are largely not based on rigorous empirical data. An important factor in consumer choice is the range of varieties available for purchase. In the UK, garden centres are a key link in the supply chain between growers and private gardens. This study is the first to determine the proportions of flowering ornamentals being sold that are attractive to flower-visiting insects. METHODS: We surveyed six garden centres in Sussex, UK, each over two days in 2015, by making 12 counts of insects visiting patches of each ornamental plant on display for sale that was in bloom. To provide a consistent baseline among different locations, we brought with us and surveyed marjoram (Origanum vulgare) plants in pots, which are known to be attractive to a wide range of flower-visiting insects. The attractiveness of plant varieties to insects was then expressed in two ways: the absolute number and relative to that on marjoram ('marjoram score'), both per unit area of plant cover. In addition, we noted whether each variety was recommended as pollinator-friendly either via a symbol on the label, or by being included in the Royal Horticultural Society's 'Perfect for Pollinators' list. Furthermore, we compared the attractiveness of plants that are typically grown for more than one year versus only one year. RESULTS: We surveyed 59-74 plant varieties in bloom across the six garden centres. In each garden centre, the distributions of variety attractiveness were highly skewed to the right, with most varieties being relatively unattractive, and few varieties highly attractive to flower-visiting insects. The median attractiveness of varieties with a recommendation was 4.2× higher than that of varieties without. But, due to the large variation there was a substantial number of both poor varieties that had a recommendation and good varieties that did not. Median attractiveness of multi-year plants was 1.6× that of single-year plants, with a similar overlap in distributions. DISCUSSION: Our study demonstrates the practicality of carrying out plant surveys in garden centres. Garden centres display large numbers of varieties for sale, most of which are in bloom. Furthermore, data gathered in garden centres appear to correlate well with data gathered in two previous studies in Sussex for plants established in gardens. Although it is unclear whether the varieties being sold in garden centres are a fair representation of varieties that are actually grown by gardeners, our results suggest that there might be considerable scope for making parks and gardens considerably more insect-friendly through judicious variety choices.

Investigating the impacts of field-realistic exposure to a neonicotinoid pesticide on bumblebee foraging, homing ability and colony growth.

The ability to forage and return home is essential to the success of bees as both foragers and pollinators. Pesticide exposure may cause behavioural changes that interfere with these processes, with consequences for colony persistence and delivery of pollination services.We investigated the impact of chronic exposure (5-43 days) to field-realistic levels of a neonicotinoid insecticide (2·4 ppb thiamethoxam) on foraging ability, homing success and colony size using radio frequency identification (RFID) technology in free-flying bumblebee colonies.Individual foragers from pesticide-exposed colonies carried out longer foraging bouts than untreated controls (68 vs. 55 min). Pesticide-exposed bees also brought back pollen less frequently than controls indicating reduced foraging performance.A higher proportion of bees from pesticide-exposed colonies returned when released 1 km from their nests; this is potentially related to increased orientation experience during longer foraging bouts. We measured no impact of pesticide exposure on homing ability for bees released from 2 km, or when data were analysed overall.Despite a trend for control colonies to produce more new workers earlier, we found no overall impacts of pesticide exposure on whole colony size. Synthesis and applications. This study shows that field-realistic neonicotinoid exposure can have impacts on both foraging ability and homing success of bumblebees, with implications for the success of bumblebee colonies in agricultural landscapes and their ability to deliver crucial pollination services. Pesticide risk assessments should include bee species other than honeybees and assess a range of behaviours to elucidate the impact of sublethal effects. This has relevance for reviews of neonicotinoid risk assessment and usage policy world-wide.

Mouthparts and nectar feeding of the flower visiting cricket Glomeremus orchidophilus (Gryllacrididae).

Glomeremus orchidophilus (Gryllacrididae) is a flower visiting cricket on the tropical island La Réunion. This species is the only Orthoptera shown to be a pollinator of a plant. We studied its nectar feeding behavior and mouthpart morphology in detail. Since G. orchidophilus possesses biting-and-chewing mouthparts, our objective was to find behavioral and/or structural specializations for nectar-feeding. The comparative analysis of feeding behavior revealed that fluid is taken up without movements of the mouthparts in Glomeremus. A comparative morphological examination of two Glomeremus species, together with several representatives of other Gryllacrididae and other Ensifera taxa revealed subtle adaptations to fluid feeding in Glomeremus. All representatives of Gryllacrididae were found to possess a distinct patch of microtrichia at the tip of their galeae. However, in Glomeremus a channel is formed between the distal components of the maxillae and the mandibles on each side of the body. Micro-CT and SEM examination revealed a longitudinal groove that extends over the galea beginning at the patch of microtrichia in the studied Glomeremus species. We hypothesize that the microtrichia take up fluid by capillarity and the action of the cibarium and pharyngeal pumps transports fluid along the channels between the maxillae and mandibles into the preoral cavity. These mouthpart features allow nectar uptake from flowers that is unique in Orthoptera.

Comparative morphology of the mouthparts of the megadiverse South African monkey beetles (Scarabaeidae: Hopliini): feeding adaptations and guild structure.

Although anthophilous Coleoptera are regarded to be unspecialised flower-visiting insects, monkey beetles (Scarabaeidae: Hopliini) represent one of the most important groups of pollinating insects in South Africa's floristic hotspot of the Greater Cape Region. South African monkey beetles are known to feed on floral tissue; however, some species seem to specialise on pollen and/or nectar. The present study examined the mouthpart morphology and gut content of various hopliine species to draw conclusions on their feeding preferences. According to the specialisations of their mouthparts, the investigated species were classified into different feeding groups. Adaptations to pollen-feeding included a well-developed, toothed molar and a lobe-like, setose lacinia mobilis on the mandible as well as curled hairs or sclerotized teeth on the galea of the maxillae. Furthermore, elongated mouthparts were interpreted as adaptations for nectar feeding. Floral- and folial-tissue feeding species showed sclerotized teeth on the maxilla, but the lacinia was mostly found to be reduced to a sclerotized ledge. While species could clearly be identified as floral or folial tissue feeding, several species showed intermediate traits suggesting both pollen and nectar feeding adaptations. Mismatches found between mouthpart morphology and previously reported flower visiting behaviours across different genera and species requires alternative explanations, not necessarily associated with feeding preferences. Although detailed examinations of the mouthparts allowed conclusions about the feeding preference and flower-visiting behaviour, additional morphological and behavioural investigations, combined with greater taxon sampling and phylogenetic data, are still necessary to fully understand hopliine host plant relationships, related to monkey beetle diversity.

Structure of the lepidopteran proboscis in relation to feeding guild.

Most butterflies and moths (Lepidoptera) use modified mouthparts, the proboscis, to acquire fluids. We quantified the proboscis architecture of five butterfly species in three families to test the hypothesis that proboscis structure relates to feeding guild. We used scanning electron microscopy to elucidate the fine structure of the proboscis of both sexes and to quantify dimensions, cuticular patterns, and the shapes and sizes of sensilla and dorsal legulae. Sexual dimorphism was not detected in the proboscis structure of any species. A hierarchical clustering analysis of overall proboscis architecture reflected lepidopteran phylogeny, but did not produce a distinct group of flower visitors or of puddle visitors within the flower visitors. Specific characters of the proboscis, nonetheless, can indicate flower and nonflower visitors, such as the configuration of sensilla styloconica, width of the lower branches of dorsal legulae, presence or absence of dorsal legulae at the extreme apex, and degree of proboscis tapering. We suggest that the overall proboscis architecture of Lepidoptera reflects a universal structural organization that promotes fluid uptake from droplets and films. On top of this fundamental structural organization, we suggest that the diversity of floral structure has selected for structural adaptations that facilitate entry of the proboscis into floral tubes.

Adult Behavior of Tirumala limniace (Lepidoptera: Danaidae).

Tirumala limniace Cramer as an ornamental butterfly is utilized in butterfly garden, in this article we study their adult activities include flight, foraging, courtship, mating, and oviposition. We found that males spent 22.1% of its time flying, 14.1% foraging, 63.8% in courtship and mating. And females spent 30.8% of its time flying, 10.1% foraging, 57.1% in courtship and mating, and 2% ovipositing. Adults emerged from pupae when temperatures were above 23.5°C and eclosion took only ∼1 min, typically followed by a small amount of flight on the first day. Flight activity peaked from the ninth to the thirteenth day after eclosion, and there were two daily peak flight times: 10:00-13:00 and 15:00-18:00. The peak of flower-visiting activity was from the eighth to the thirteenth day after eclosion, and there were two daily peak foraging times: 11:00-12:00 and 16:00-17:00. Flight and foraging frequency and time were positively correlated and both were closely related to temperature, with very little flight or foraging below 18°C and an increase at temperatures above 22°C. Courtship and mating took place on the sixth day after eclosion, while oviposition occurred the following day. Oviposition occurred over 8 d, and the shortest time of a single oviposition was 2 s. The average life expectancy of males was 16.5 d, while that of females was 15 d.

Evolution of extreme proboscis lengths in Neotropical Hesperiidae (Lepidoptera).

Exaggerated morphologies have evolved in insects as adaptations to nectar feeding by natural selection. For example, the suctorial mouthparts of butterflies enable these insects to gain access to floral nectar concealed inside deep floral tubes. Proboscis length in Lepidoptera is known to scale with body size, but whether extreme absolute proboscis lengths of nectar feeding butterflies result from a proportional or disproportional increase with body size that differs between phylogenetic lineages remains unknown. We surveyed the range of variation that occurs in scaling relationships between proboscis length and body size against a phylogenetic background among Costa Rican Hesperiidae. We obtained a new record holder for the longest proboscis in butterflies and showed that extremely long proboscides evolved at least three times independently within Neotropical Hesperiidae. We conclude that the evolution of extremely long proboscides results from allometric scaling with body size, as demonstrated in hawk moths. We hypothesize that constraints on the evolution of increasingly long butterfly proboscides may come from (1) the underlying scaling relationships, i.e., relative proboscis length, combined with the butterfly's flight style and flower-visiting behaviour and/or (2) developmental constraints during the pupal phase. Lastly, we discuss why butterflies did not evolve similar scaling relationships as hawk moths.