To probe its environment, the flying insect controllably flexes, twists, and maneuvers its antennae by coupling mechanical deformations with the sensory output. We question how the materials properties of insect antennae could influence their performance. A comparative study was conducted on four hawkmoth species: Manduca sexta, Ceratomia catalpae, Manduca quinquemaculata, and Xylophanes tersa. The morphology of the antennae of three hawkmoths that hover while feeding and one putatively non-nectar-feeding hawkmoth (Ceratomia catalpa) do not fundamentally differ, and all the antennae are comb-like (i.e., pectinate), markedly in males but weakly in females. Applying different weights to the free end of extracted cantilevered antennae, we discovered anisotropy in flexural rigidity when the antenna is forced to bend dorsally versus ventrally. The flexural rigidity of male antennae was less than that of females. Compared with the hawkmoths that hover while feeding, Ceratomia catalpae has almost two orders of magnitude lower flexural rigidity. Tensile tests showed that the stiffness of male and female antennae is almost the same. Therefore, the differences in flexural rigidity are explained by the distinct shapes of the antennal pectination. Like bristles in a comb, the pectinations provide extra rigidity to the antenna. We discuss the biological implications of these discoveries in relation to the flight habits of hawkmoths. Flexural anisotropy of antennae is expected in other groups of insects, but the targeted outcome may differ. Our work offers promising new applications of shaped fibers as mechanical sensors. STATEMENT OF SIGNIFICANCE: Insect antennae are blood-filled, segmented fibers with muscles in the two basal segments. The long terminal segment is muscle-free but can be flexed. Our comparative analysis of mechanical properties of hawkmoth antennae revealed a new feature: antenna resistance to bending depends on the bending direction. Our discovery replaces the conventional textbook scenario considering hawkmoth antennae as rigid rods. We showed that the pectinate antennae of hawkmoths behave as a comb in which the bristles resist bending when they come together. This anisotropy of flexural resistance offers a new mode of environmental sensing that has never been explored. The principles we found apply to other insects with non-axisymmetric antennae. Our work offers new applications for shaped fibers that could be designed to sense the flows.

Pollinators are needed for the reproduction of Echites umbellatus, and only sphingid moths have mouthparts long enough to reach the nectar at the bottom of the species\' long, twisted floral tube. Though plants produce many flowers over a period of several months, one observes very few fruits in nature. We asked: (1) Are plants self-compatible, or do they need pollen from another individual to set fruit and seed? (2) Are cross-pollinations between unrelated individuals more successful than crosses with relatives? (3) How does the relatedness of pollen and ovule parent plants affect fruit set, seed number, and seed quality? We investigated the breeding system of E. umbellatus by collecting fruits from seven sites, growing plants and performing hand pollinations over a period of several years, collecting and measuring fruits and counting seeds. Echites umbellatus is self-incompatible, though some individuals produce fruit by self-pollination. Cross-pollinations between unrelated individuals set the most fruit (59%), and those that were self-pollinated set the least (9%). Fruit set from cross-pollinations between related individuals was intermediate (32%). Although the number of seeds per fruit did not differ significantly among pollination treatments, fruits from self-pollinations had substantially fewer viable seeds than outcrossed fruits, with fruits from sibling crosses being intermediate. There were higher levels of self-compatibility in the fragment populations compared with plants from intact habitats. Self-incompatibility may explain why fruit set is low in this plant species; future investigation into the breakdown of self-incompatibility in smaller populations is warranted.

Capparis spinosa is a widespread charismatic plant, in which the nocturnal floral habit contrasts with the high visitation by diurnal bees and the pronounced scarcity of hawkmoths. To resolve this discrepancy and elucidate floral evolution of C. spinosa, we analyzed the intrafloral patterns of visual and olfactory cues in relation to the known sensory biases of the different visitor guilds (bees, butterflies, and hawkmoths).
We measured the intrafloral variation of scent, reflectance spectra, and colorimetric properties according to three guilds of known visitors of C. spinosa. Additionally, we sampled visitation rates using a motion-activated camera.
Carpenter bees visited the flowers eight times more frequently than nocturnal hawkmoths, at dusk and in the following morning. Yet, the floral headspace of C. spinosa contained a typical sphingophilous scent with high emission rates of certain monoterpenes and amino-acid derived compounds. Visual cues included a special case of multisensory nectar guide and color patterns conspicuous to the visual systems of both hawkmoths and bees.
The intrafloral patterns of sensory stimuli suggest that hawkmoths have exerted strong historical selection on C. spinosa. Our study revealed two interesting paradoxes: (a) the flowers phenotypically biased towards the more inconsistent pollinator; and (b) floral display demands an abundance of resources that seems maladaptive in the habitats of C. spinosa. The transition to a binary pollination system accommodating large bees has not required phenotypic changes, owing to specific eco-physiological adaptations, unrelated to pollination, which make this plant an unusual case in pollination ecology.

Plants whose flowers open at night but remain open during the day also attract diurnal flower visitors, potentially boosting their pollination rates and providing resources that can support diverse arthropod communities. The rough-leaf velvetseed, Guettarda scabra (Rubiaceae), is an evergreen shrub that thrives only in the imperiled pine rockland habitat in south Florida. Its white, tubular, and fragrant flowers open during late afternoon, exhibiting traits strongly associated with the attraction of nocturnal hawkmoths (Sphingidae). Flowers of G. scabra remain open until the following morning, becoming available to a wider array of visitors, bringing into question the expectation that sphingophilous flowers are visited mainly by hawkmoths. To evaluate whether the flowers of G. scabra are mainly visited by nocturnal hawkmoths and understand the role of this plant in the pine rockland habitat, we characterized the arthropod fauna associated with its flowers during the morning, evening, and at night. We found that most flower visitors were diurnal insects of the orders Hymenoptera and Lepidoptera, although we observed other arthropod groups too. Visitation at night was dominated by two species of hawkmoths. Nectar was the main resource used by the arthropod community during this study. Legitimate visitation and nectar-robbing were the behaviors most frequently observed among the flower visitors. Our results suggest that flowers of the night-blooming G. scabra constitute an important food source for both diurnal and nocturnal arthropod fauna in the fire-dependent pine rocklands of southern Florida. Our study provides novel data to support efforts to conserve and protect pine rocklands and the plants and animals that inhabit them.

To meet energetic and osmotic demands, animals make dynamic foraging decisions about food quality and quantity. In the wild, foraging animals may be forced to consume a less preferred or sub-optimal food source for long periods of time. Few choice feeding assays in laboratory settings approximate such contingencies. In this study the foraging behaviors of the hawkmoth Manduca sexta were measured when adult moths were placed within different relative humidity (RH) environments (20%, 40%, 60% and 80% RH) and provided with only one of the following experimental nectars: 0% (water), 12% or 24 % w/V sucrose solutions. Overall, ambient humidity influenced survivorship and foraging behaviors. Moth survivorship increased at higher ambient humidity regardless of experimental nectar. Moths that had access to experimental nectar imbibed large volumes of fluid regardless of what nectar was offered when placed at the lowest humidity (20% RH). However, when placed at the highest humidity (80% RH), moths imbibed higher volumes of fluid when given access to experimental nectar with sucrose in comparison with water. RH also influenced daily foraging behaviors: peak nectar consumption occurred earlier at lower RH levels. Consistent with previous studies in which moths could choose among nectar solutions, total energy intake was not affected by ambient RH under no-choice conditions. However, the proportion of time spent foraging and total energy consumption were significantly reduced across all RH levels in no-choice assays, when compared with previous studies of choice assays under the same conditions. Our results show that even when M. sexta moths are presented with limited options, they can alter their foraging behavior in response to environmental changes, enabling them to meet osmotic and/or energetic demands.

UNASSIGNED: Hawkmoths consist of species where most adults are nocturnal, but there are some day-flying genera. Hawkmoth species have a wide variety of life-history traits, comprising species with adults (mostly nectarivorous though with some exceptions, honey-feeding), but there are also species that do not feed at all. The nectarivorous species are an important component of tropical ecosystems, with significant roles as major pollinators of both crops and wild flora with the pollination done by the adult stage. Pollinators are in decline world-wide and there is need for baseline data to provide information about their conservation strategies. Species occurrence data from Museum collections have been shown to be of great value as a tool for prioritising conservation actions in Africa. The National Museums of Kenya (NMK) have a large and active entomology collection that is in continuous growth. The NMK\'s collection of hawkmoths had not been digitised prior to 2017. This moth family Sphingidae includes about 1,602 species and 205 genera worldwide (Kitching et al. 2018) with the majority of these species occurring in Africa. These moth species can also be used as indicators in biodiversity assessments as they can be easily sampled and identified. However, hawkmoths have rarely been surveyed over the long term for this purpose. Long-term datasets are of unquestionable significance for understanding and monitoring temporal changes in biodiversity. These hawkmoth data have addressed one of the most significant challenges to insect conservation, the lack of baseline information concerning species diversity and distribution and have provided key historic hawkmoth species diversity and distribution data that can be used to monitor their populations in the face of climate change and other environmental degradation issues that are facing the world today. The publication of the hawkmoth species occurrence data records in GBIF has enhanced data visibility to a wider audience promoting availability for use.
UNASSIGNED: The hawkmoth (Lepidoptera: Sphingidae) collection at the National Museums of Kenya was digitised from 2017 - 2020 and this paper presents details of species occurrence records as in the insect collection at the NMK, Nairobi, Kenya.The collection holds 5,095 voucher specimens consisting of 88 genera and 208 species. The collection covers the period between 1904 and 2020.The geographical distribution of the hawkmoths housed at the NMK covers East Africa at 81.41%, West Africa at 7.20%, Southern Africa at 6.89%, Central Africa at 4.02% and North Africa at 0.2%.

Within-species variation in pollinator behavior is widely observed, but its causes have been minimally investigated. Pollinator sex is associated with large differences in behavior that may lead to predictable differences in flower foraging, but this expectation has not been explicitly tested. We investigate sex-associated differences in nectar-foraging behavior of the hawkmoth Hyles lineata, using pollen on the proboscis as a proxy for flower visitation. We tested two predictions emerging from the literature: (1) the sexes differ in the flower species they visit, (2) females are more specialized in flower choice. We also examined potential drivers underlying these predictions by performing field and laboratory experiments to test whether males (3) switch among flower species more frequently, or (4) fly farther and therefore encounter more species than females. Consistent with prediction (1), pollen load composition differed between the sexes, indicative of visitation differences. Contrary to prediction (2), females consistently carried more species-rich pollen loads than males. (3) Both sexes switched between flower species at similar rates, suggesting that differences in floral fidelity are unlikely to explain why females are less specialized than males. (4) Males flew longer distances than females; coupled with larger between-site differences in pollen composition for females, this result suggests that sex differences in mobility influence foraging, and that females may forage more frequently and in smaller areas than males. Together, our results demonstrate that sex-associated foraging differences can be large and consistent over time, and highlight the importance of sex as a driver of variation in pollinator behavior.

To date, a relatively complete classification of Sphingidae (Lepidoptera) has been generated, but the phylogeny of the family remains need to be fully resolved. Some phylogenetic relationships within Sphingidae still remains uncertain, especially the taxonomic status of the subfamily Langiinae and its sole included genus and species, Langia zenzeroides. To begin to address this problem, we generated nine new complete mitochondrial genomes, including that of Langia, and together with that of Theretra oldenlandiae from our previous study and 25 other Sphingidae mitogenomes downloaded from GenBank, analyzed the phylogenetic relationships of Sphingidae and investigated the mitogenomic differences among members of the Langiinae, Sphinginae, Smerinthinae and Macroglossinae. The mitogenomes of Sphingidae varied from 14995 bp to 15669 bp in length. The gene order of all newly sequenced mitogenomes was identical, containing 13 protein-coding genes, two ribosomal RNA genes, 22 transfer RNA genes and the A + T-rich region. Nucleotide composition was A + T biased, and all the protein-coding genes exhibited a positive AT-skew, which was reflected in the nucleotide composition, codon, and amino acid usage. The A + T-rich region was comprised of nonrepetitive sequences, which contained regulatory elements related to the control of replication and transcription. We analyzed concatenated gene sequences, with third codon positions of protein coding genes and rRNAs excluded, using Maximum Likelihood and Bayesian Inference techniques. All four currently recognized subfamilies were recovered as monophyletic but in contrast to the most recent studies, our preferred tree placed Langiinae as the first subfamily to diverge within Sphingidae rather as sister to Smerinthinae + Sphinginae. Our results also support the removal of the genus Barbourion from the smerinthine tribe Ambulycini to an unresolved position in \"Smerinthinae incertae sedis\".

Orchids attach their pollinaria (cohesive masses of pollen) to specific body parts of flower visitors, but usually not to the hairy and scaly body parts of flower-visiting moths, because hairs and scales are easily detached. We demonstrate that pollinaria of Habenaria sagittifera (Orchidaceae) are transferred among flowers on the hairy thoraxes of moths in Japan. Diurnal and nocturnal insects visited the orchid flowers. However, pollinaria were attached only to the hairy thoraxes of plusiine moths (Lepidoptera: Noctuidae). All pollinaria were directly attached to the ventral thorax surfaces at the bases of hairs. Orchid spur lengths matched plusiine proboscis lengths so that nectar-feeding moths contacted the viscidia (sticky pads of pollinaria) and stigma. Other flower visitors did not contact the viscidia or stigmas while feeding on nectar. Habenaria sagittifera appears to have a floral morphology that is adaptive for the transfer of pollinaria on the thoraxes of plusiine moths.

The color and patterns of animal-pollinated flowers are known to have effects on pollinator attraction. In this study, the relative importance of flower color and color contrast patterns on pollinator attraction was examined in two pollinator groups, swallowtail butterflies and hawkmoths using two Hemerocallis species; butterfly-pollinated H. fulva and hawkmoth-pollinated H. citrina, having reddish and yellowish flowers in human vision, respectively. Flowers of both species have UV bullseye patterns, composed of UV-absorbing centers and UV-reflecting peripheries, known to function as a typical nectar guide, but UV reflectance was significantly more intense in the peripheries of H. citrina flowers than in those of H. fulva flowers. Comparison based on the visual systems of butterflies and hawkmoths showed that the color contrast of the bullseye pattern in H. citrina was more intense than that in H. fulva. To evaluate the relative importance of flower color and the color contrast of bullseye pattern on pollinator attraction, we performed a series of observations using experimental arrays consisting of Hemerocallis species and their hybrids. As a result, swallowtail butterflies and crepuscular/nocturnal hawkmoths showed contrasting preferences for flower color and patterns: butterflies preferred H. fulva-like colored flower whereas the preference of hawkmoths was affected by the color contrast of the bullseye pattern rather than flower color. Both crepuscular and nocturnal hawkmoths consistently preferred flowers with stronger contrast of the UV bullseye pattern, whereas the preference of hawkmoths for flower color was incoherent. Our finding suggests that hawkmoths can use UV-absorbing/reflecting bullseye patterns for foraging under light-limited environments and that the intensified bullseye contrast of H. citrina evolved as an adaptation to hawkmoths. Our results also showed the difference of visual systems between pollinators, which may have promoted floral divergence.