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The Mojave poppy bee, Perdita meconis Griswold (Hymenoptera: Anthophila: Andrenidae), is a species of conservation concern that is restricted to the eastern Mojave Desert of North America. It is a specialist pollinator of two poppy genera, Arctomecon and Argemone (Papaveraceae), and is being considered for listing under the US Endangered Species Act along with one of its pollinator hosts, the Las Vegas bearpoppy (Arctomecon californica). Here, we present a near chromosome-level genome of the Mojave poppy bee to provide a genomic resource that will aid conservation efforts and future research. We isolated DNA from a single, small (<7 mm), male specimen collected using non-ideal preservation methods then performed whole-genome sequencing using PacBio HiFi technology. After quality and contaminant filtering, the final draft genome assembly is 327 Mb, with an N50 length of 17.5 Mb. Annotated repetitive elements compose 37.3% of the genome, although a large proportion (24.87%) of those are unclassified repeats. Additionally, we annotated 18,245 protein-coding genes and 19,433 transcripts. This genome represents one of only a few genomes from the large bee family Andrenidae and one of only a few genomes for pollinator specialists. We highlight both the potential of this genome as a resource for future research, and how high-quality genomes generated from small, non-ideal (in terms of preservation) specimens could facilitate biodiversity genomics.
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Adaptive coloration among animals is one of the most recognizable outcomes of natural selection. Here, we investigate evolutionary drivers of white coloration in velvet ants (Hymenoptera: Mutillidae), which has previously been considered camouflage with the fruit of creosote bush (Larrea tridentata). Our analyses indicate instead that velvet ants evolved white coloration millions of years before creosote bush was widespread in North America's hot deserts. Furthermore, velvet ants and the creosote fruit exhibit different spectral reflectance patterns, which appear distinct to potential insectivorous predators. While the white coloration in velvet ants likely did not evolve as camouflage, we find that white-coloured species remain cooler than their red/orange relatives, and therefore we infer the white coloration likely evolved in response to Neogene desertification. This study shows the importance of cross-disciplinary investigation and of testing multiple hypotheses when investigating evolutionary drivers of adaptive coloration.
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Hormigas , Mimetismo Biológico , Animales , Ecología , América del NorteRESUMEN
Chemically mediated plant-herbivore interactions contribute to the diversity of terrestrial communities and the diversification of plants and insects. While our understanding of the processes affecting community structure and evolutionary diversification has grown, few studies have investigated how trait variation shapes genetic and species diversity simultaneously in a tropical ecosystem. We investigated secondary metabolite variation among subpopulations of a single plant species, Piper kelleyi (Piperaceae), using high-performance liquid chromatography (HPLC), to understand associations between plant phytochemistry and host-specialized caterpillars in the genus Eois (Geometridae: Larentiinae) and associated parasitoid wasps and flies. In addition, we used a genotyping-by-sequencing approach to examine the genetic structure of one abundant caterpillar species, Eois encina, in relation to host phytochemical variation. We found substantive concentration differences among three major secondary metabolites, and these differences in chemistry predicted caterpillar and parasitoid community structure among host plant populations. Furthermore, E. encina populations located at high elevations were genetically different from other populations. They fed on plants containing high concentrations of prenylated benzoic acid. Thus, phytochemistry potentially shapes caterpillar and wasp community composition and geographic variation in species interactions, both of which can contribute to diversification of plants and insects.
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Lepidópteros/fisiología , Fitoquímicos/metabolismo , Piperaceae/parasitología , Animales , Variación Genética , Lepidópteros/genética , Modelos Biológicos , Parásitos/fisiología , Fitoquímicos/química , Hojas de la Planta/química , Análisis de Componente Principal , Especificidad de la EspecieRESUMEN
The genetic and ecological factors that shape the evolution of animal diets remain poorly understood. For herbivorous insects, the expectation has been that trade-offs exist, such that adaptation to one host plant reduces performance on other potential hosts. We investigated the genetic architecture of alternative host use by rearing individual Lycaeides melissa butterflies from two wild populations in a crossed design on two hosts (one native and one introduced) and analysing the genetic basis of differences in performance using genomic approaches. Survival during the experiment was highest when butterfly larvae were reared on their natal host plant, consistent with local adaptation. However, cross-host correlations in performance among families (within populations) were not different from zero. We found that L. melissa populations possess genetic variation for larval performance and variation in performance had a polygenic basis. We documented very few genetic variants with trade-offs that would inherently constrain diet breadth by preventing the optimization of performance across hosts. Instead, most genetic variants that affected performance on one host had little to no effect on the other host. In total, these results suggest that genetic trade-offs are not the primary cause of dietary specialization in L. melissa butterflies.
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Adaptación Fisiológica/genética , Evolución Biológica , Mariposas Diurnas/genética , Variación Genética , Herbivoria , Animales , Planta del Astrágalo , Mariposas Diurnas/fisiología , Femenino , Genoma de los Insectos , Genotipo , Larva/fisiología , MasculinoRESUMEN
The asidine darkling beetles (Coleoptera: Tenebrionidae: Asidini) are a diverse tribe of flightless tenebrionids found in many arid and sub-arid habitats around the world. The 263 currently described North American species are contained in ten genera, all of which are restricted to the western half of the continent. The Asidini, like all members of the subfamily Pimeliinae, lack defensive glands. Instead, several phenotypic traits occur within the tribe that may help limit predation. These include the contrasting defensive strategies of crypsis, through either background matching or pattern disruption, and Batesian mimicry of the chemically defended genus Eleodes. Dorsal elytral morphology was assessed between 53 North American asidine species and 13 common Eleodes model species using multiple methodologies to assess similarities between species in the two groups that might indicate mimetic relationships. A phylogeny of the North American asidines is used to map the occurrence of differing defensive strategies within the tribe. Crypsis is reconstructed as the ancestral state, with two origins for Batesian mimicry and multiple reversals. The combination of strongly to weakly cryptic species and varying levels of mimetic fidelity to Eleodes model species make the asidines a promising lineage upon which to further explore the evolution of defensive phenotypes.
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The genitalia of male insects have been widely used in taxonomic identification and systematics and are potentially involved in maintaining reproductive isolation between species. Although sexual selection has been invoked to explain patterns of morphological variation in genitalia among populations and species, developmental plasticity in genitalia likely contributes to observed variation but has been rarely examined, particularly in wild populations. Bilateral gynandromorphs are individuals that are genetically male on one side of the midline and genetically female on the other, while mosaic gynandromorphs have only a portion of their body developing as the opposite sex. Gynandromorphs might offer unique insights into developmental plasticity because individuals experience abnormal cellular interactions at the genitalic midline. In this study, we compare the genitalia and wing patterns of gynandromorphic Anna and Melissa blue butterflies, Lycaeides anna (Edwards) (formerly L. idas anna) and L. melissa (Edwards) (Lepidoptera: Lycaenidae), to the morphology of normal individuals from the same populations. Gynandromorph wing markings all fell within the range of variation of normal butterflies; however, a number of genitalic measurements were outliers when compared with normal individuals. From these results, we conclude that the gynandromorphs' genitalia, but not wing patterns, can be abnormal when compared with normal individuals and that the gynandromorphic genitalia do not deviate developmentally in a consistent pattern across individuals. Finally, genetic mechanisms are considered for the development of gynandromorphism in Lycaeides butterflies.
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Mariposas Diurnas/anatomía & histología , Alas de Animales/anatomía & histología , Animales , Mariposas Diurnas/crecimiento & desarrollo , Femenino , Genitales/anatomía & histología , Genitales/crecimiento & desarrollo , Masculino , Estados Unidos , Alas de Animales/crecimiento & desarrolloRESUMEN
There is a need for a reliable statistical test which is appropriate for assessing cospeciation of more than two phylogenies. We have developed an algorithm using a permutation method that can be used to test for and infer tri-trophic evolutionary relationships of organisms given both their phylogenies and pairwise interactions. An overall statistic has been developed based on the dominant eigenvalue of a covariance matrix, and compared to values of the statistic computed when tree labels are permuted. The resulting overall p-value is used to test for the presence or absence of cospeciation in a tri-trophic system. If cospeciation is detected, we propose new test statistics based on partial correlations to uncover more details about the relationships between multiple phylogenies. One of the strengths of our method is that it allows more parasites than hosts or more hosts than parasites, with multiple associations and more than one parasite attached to a host (or one parasite attached to multiple hosts). The new method does not require any parametric assumptions of the distribution of the data, and unlike the old methods, which utilize several pairwise steps, the overall statistic used is obtained in one step. We have applied our method to two published datasets where we obtained detailed information about the strength of associations among species with calculated partial p-values and one overall p-value from the dominant eigenvalue test statistic. Our permutation method produces reliable results with a clear procedure and statistics applied in an intuitive manner. Our algorithm is useful in testing evidence for three-way cospeciation in multiple phylogenies with tri-trophic associations and determining which phylogenies are involved in cospeciation.
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Modelos Genéticos , Algoritmos , Animales , Bacterias/genética , Evolución Biológica , Simulación por Computador , Interpretación Estadística de Datos , Especiación Genética , Interacciones Huésped-Parásitos/genética , Isópteros/genética , Isópteros/microbiología , Filogenia , SimbiosisRESUMEN
Arctomecon californica, the Las Vegas bear poppy, is a rare plant found only in the eastern Mojave Desert of North America. Because of recent declines in the populations of this endemic plant, conservationists are currently seeking protection for A. californica under the US Endangered Species Act. In this study, we investigated the natural history of A. californica and documented insect visitors potentially pollinating A. californica in Clark County, Nevada. We find that the populations of A. californica fluctuate from year to year, with many populations declining by over 90% from 2021 to 2022. The pollinator communities of A. californica also vary from year to year. In some years, specialist bees in the genus Perdita make up the majority of pollinators, while in other years, generalist bees like Apis mellifera and Hylaeus dominate. Furthermore, we confirm what previous work has suggested, that A. californica requires pollinators to set seed, yet not all insect visitors are good pollinators. This work provides useful natural history information about the Las Vegas bear poppy, which will be informative to conservationists designing strategies to protect this imperiled species.
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Despite the broad recognition of mimicry among bumble bees, distinct North American mimicry rings have yet to be defined, due in part to the prevalence of intermediate and imperfect mimics in this region. Here we employ a generalization approach using human perception to categorize mimicry rings among North American bumble bees. We then map species distributions on North American ecoregions to visually test for geographic concordance among similarly-colored species. Our analyses suggest that there are five mimicry rings in the North American bumble bee mimicry complex, and one broadly distributed group of mixed and intermediate color forms. We describe the Black Mimicry Ring, Black-cloaked Mimicry Ring, Eastern Yellow Mimicry Ring, Red Mimicry Ring, and Western Yellow Mimicry Ring as well as the mixed group. We then test these hypothesized mimicry rings by examining other insects that participate in these mimicry rings. Describing these mimicry rings is a vital step that will enable future analyses of imperfect mimicry, intermediate mimicry, and additional analyses of other insects that mimic bumble bees.
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Mimetismo Biológico , Humanos , Abejas , Animales , América del Norte , PercepciónRESUMEN
Understanding the ecology of extinction is one of the primary challenges facing ecologists in the 21st century. Much of our current understanding of extinction, particularly for invertebrates, comes from studies with large geographic coverage but less temporal resolution, such as comparisons between historical collection records and contemporary surveys for geographic regions or political entities. We present a complementary approach involving a data set that is geographically restricted but temporally intensive: we focus on three sites in the Central Valley of California, and utilize 35 years of biweekly (every two weeks) surveys at our most long-sampled site. Previous analyses of these data revealed declines in richness over recent decades. Here, we take a more detailed approach to investigate the mode of decline for this fauna. We ask if all species are in decline, or only a subset. We also investigate traits commonly found to be predictors of extinction risk in other studies, such as body size, diet breadth, habitat association, and geographic range. We find that population declines are ubiquitous: the majority of species at our three focal sites (but not at a nearby site at higher elevation) are characterized by reductions in the fraction of days that they are observed per year. These declines are not readily predicted by ecological traits, with the possible exception of ruderal/non-ruderal status. Ruderal species, in slightly less precipitous decline than non-ruderal taxa, are more dispersive and more likely to be associated with disturbed habitats and exotic hosts. We conclude that population declines and extirpation, particularly in regions severely and recently impacted by anthropogenic alteration, might not be as predictable as has been suggested by other studies on the ecology of extinction.
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Mariposas Diurnas , Extinción Biológica , Animales , Biodiversidad , California , Geografía , Dinámica PoblacionalRESUMEN
The influence of historical climatic and geological changes on patterns of species diversification was investigated in a widely distributed group of North American nocturnal mutillids (Hymenoptera: Mutillidae: Sphaeropthalminae), with particular focus on Pleistocene glacial cycles and earlier patterns of Neogene mountain building. We collected molecular data from two nuclear intergenic regions (internal transcribed spacer regions 1 and 2; approximately 2600 bp in total) to produce the first phylogeny of nocturnal Nearctic mutillids. Bayesian inference of the combined data returned a well-resolved tree with posterior probabilities of over 95% for most nodes. This tree suggested the monophyly of the nocturnal, primarily Nearctic, Sphaeropthalminae genera, but the paraphyly of the three largest genera (Odontophotopsis, Photomorphus and Sphaeropthalma). Dates of species divergences were obtained using r8s (PL and NPRS) and BEAST with the date of Dominican amber set at three different dates (15 Ma, 20 Ma, and 45 Ma) to account for uncertainty in the fossil age. The derived dates ranged from the Pleistocene to the Middle Miocene, suggesting that both recent Pleistocene glaciation cycles and older orogenic events, albeit to a somewhat greater extent, were both causes of major diversification in western North America. Examination of other phylogeographical studies using North American desert taxa indicated that diversification patterns are explained by either mountain building or Pleistocene climate change, depending on the taxa in question.
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Hormigas/genética , Evolución Molecular , Filogenia , Animales , Hormigas/clasificación , Teorema de Bayes , Cambio Climático , ADN Espaciador Ribosómico/genética , Geografía , Modelos Genéticos , América del Norte , Análisis de Secuencia de ADNRESUMEN
Grand Staircase-Escalante National Monument is a federally protected area found in central southern Utah. Designated in 1996 by President William J. Clinton, it was recently reduced in size by President Donald J. Trump in a proclamation that turned the one large monument into three smaller ones. A long-term, standardized study of the bees had been conducted from 2000-2003, revealing 660 species. The bee communities of the area are characterized by being spatially heterogeneous; most of the bees occur in isolated areas, with only a few being both abundant and widespread. Here we examine what affect the recent resizing of the monument has on the number, and ecology, of the bees now excluded from monument boundaries. Using the new monument boundaries and the geographic coordinates associated with each bee, we derived new species lists for each of the three monuments, and compared them to each other, and to the excluded lands. All three monuments now protect unique faunas, with Bray-Curtis similarity values not exceeding 0.59%. Each monument now harbors species not found in the other two monuments. We found that 84 bee species are no longer protected by any of the three monuments. These 84 species were not concentrated in one area that is now excluded, but were scattered throughout the newly excluded lands. For some of the excluded bee species, there is no evidence that they are rare or imperiled, being widespread throughout the west. However, there is a concentration of bees in the southern and eastern former monument lands that represent range extensions from nearby hot deserts. In addition to numerous range extensions, the list of excluded bees also contains several undescribed species (newly discovered species that have not yet been named and described by taxonomists) and morphospecies (individuals that are morphologically distinct, but that require additional research before species designations can be made). This indicates that the bee communities housed in these excluded areas would benefit from additional scientific inquiry. The areas now excluded from monument protections house a greater proportion of the original GSENM bee community than any of the three new monument units. We conclude this paper by discussing what the smaller monuments might mean for bee conservation in this hot spot of bee biodiversity and suggest that bee communities here and elsewhere should be taken into account when conservation decisions are being made.
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Our goals were to explore the relationship between biogeography and the evolution of fire-adaptive syndromes in the genus Pinus. We used a previously published time-calibrated phylogeny and conducted ancestral trait reconstruction to estimate the likely timing of diversification in Pinus, and to determine when fire-adaptive syndromes evolved in the lineage. To explore trait conservation among fire syndromes and to investigate historical biogeography, we constructed ancestral state reconstructions using the program RASP and estimated the degree of conservatism for fire-adapted traits in the program BaTS. Our reconstructions suggest that the Bering land bridge, which connected North America and Asia, probably played a major role in early pine evolution. Our estimates indicated that fire-adaptive syndromes seem to have evolved more frequently in New World taxa and probably are related to the uplift of major North American mountain ranges. Our data suggest that certain geographically widespread adaptations to fire evolved repeatedly, possibly due to localized changes in climate and environment, rather than resulting from large dispersal events of pre-adapted individuals.
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The stings of bees, wasps, and ants are something that catches the attention of anyone that experiences them. While many recent studies have focused on the pain inflicted by the stings of various stinging wasps, bees, or ants (Hymenoptera: Aculeata), little is known about how the length of the sting itself varies between species. Here, we investigate the sting length of a variety of aculeate wasps, and compare that to reported pain and toxicity values. We find that velvet ants (Hymenoptera: Mutillidae) have the longest sting compared to their body size out of any bee, wasp, or ant species. We also find that there is no link between relative sting length and pain; however, we did find an inverse relationship between relative sting length and toxicity with taxa having shorter relative stings being more toxic. While we found a significant relationship between host use and relative sting length, we suggest that the long sting length of the velvet ants is also related to their suite of defenses to avoid predation.
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Interest in bees has grown dramatically in recent years in light of several studies that have reported widespread declines in bees and other pollinators. Investigating declines in wild bees can be difficult, however, due to the lack of faunal surveys that provide baseline data of bee richness and diversity. Protected lands such as national monuments and national parks can provide unique opportunities to learn about and monitor bee populations dynamics in a natural setting because the opportunity for large-scale changes to the landscape are reduced compared to unprotected lands. Here we report on a 4-year study of bees in Grand Staircase-Escalante National Monument (GSENM), found in southern Utah, USA. Using opportunistic collecting and a series of standardized plots, we collected bees throughout the six-month flowering season for four consecutive years. In total, 660 bee species are now known from the area, across 55 genera, and including 49 new species. Two genera not previously known to occur in the state of Utah were discovered, as well as 16 new species records for the state. Bees include ground-nesters, cavity- and twig-nesters, cleptoparasites, narrow specialists, generalists, solitary, and social species. The bee fauna reached peak diversity each spring, but also experienced a second peak in diversity in late summer, following monsoonal rains. The majority of GSENM's bees are highly localized, occurring in only a few locations throughout the monument, and often in low abundance, but consistently across the four years. Only a few species are widespread and super-abundant. Certain flowering plants appear to be inordinately attractive to the bees in GSENM, including several invasive species. GSENM protects one of the richest bee faunas in the west; the large elevational gradient, incredible number of flowering plants, and the mosaic of habitats are all likely contributors to this rich assemblage of bees.
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Africa has the most tropical and subtropical land of any continent, yet has relatively low species richness in several taxa. This depauperate nature of the African tropical fauna and flora has led some to call Africa the "odd man out." One exception to this pattern is velvet ants (Hymenoptera: Mutillidae), wingless wasps that are known for Müllerian mimicry. While North American velvet ants form one of the world's largest mimicry complexes, mimicry in African species has not been investigated. Here we ask do African velvet ant Müllerian mimicry rings exist, and how do they compare to the North American complex. We then explore what factors might contribute to the differences in mimetic diversity between continents. To investigate this we compared the color patterns of 304 African velvet ant taxa using nonmetric multidimensional scaling (NMDS). We then investigated distributions of each distinct mimicry ring. Finally, we compared lizard diversity and ecoregion diversity on the two continents. We found that African female velvet ants form four Müllerian rings, which is half the number of North American rings. This lower mimetic diversity could be related to the relatively lower diversity of insectivorous lizard species or to the lower number of distinct ecoregions in Africa compared to North America.
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Hormigas/fisiología , Mimetismo Biológico , África , Animales , América del NorteRESUMEN
Velvet ants are a group of parasitic wasps that are well known for a suite of defensive adaptations including bright coloration and a formidable sting. While these adaptations are presumed to function in antipredator defense, observations between potential predators and this group are lacking. We conducted a series of experiments to determine the risk of velvet ants to a host of potential predators including amphibians, reptiles, birds, and small mammals. Velvet ants from across the United States were tested with predator's representative of the velvet ants native range. All interactions between lizards, free-ranging birds, and a mole resulted in the velvet ants survival, and ultimate avoidance by the predator. Two shrews did injure a velvet ant, but this occurred only after multiple failed attacks. The only predator to successfully consume a velvet ant was a single American toad (Anaxyrus americanus). These results indicate that the suite of defenses possessed by velvet ants, including aposematic coloration, stridulations, a chemical alarm signal, a hard exoskeleton, and powerful sting are effective defenses against potential predators. Female velvet ants appear to be nearly impervious to predation by many species whose diet is heavily derived of invertebrate prey.
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The stinging hymenopteran velvet ants (Mutillidae) and bumble bees (Apidae: Bombus spp.) have both undergone extensive diversification in aposematic color patterns, including yellow-red hues and contrasting dark-light body coloration, as a result of Müllerian mimicry. Understanding the genetic and developmental mechanisms underlying shifts in these mimetic colors requires characterization of their pigmentation. In this study, a combination of solubility, spectrophotometry, and melanin degradation analysis are applied to several color forms and species of these lineages to determine that orange-red colors in both lineages are comprised of primarily dopamine-derived pheomelanins. Until a few recent studies, pheomelanins were thought not to occur in insects. These results support their potential to occur across insects and particularly among the Hymenoptera. Shifts between black and orange-red colors, such as between mimetic color forms of bumble bee Bombus melanopygus, are inferred to involve modification of the ratios of dark eumelanins to red pheomelanins, thus implicating the melanin pathway in mimetic diversification. This discovery highlights the need to focus on how pheomelanins are synthesized in the insect melanin pathway and the potential for new pigments to be found even in some of our most well-known insect systems.