Systematic revision of the North American velvet ant genus Invreiella Suárez (Hymenoptera: Mutillidae) with description of eleven new species.

The velvet ant genus Invreiella Suárez, 1966 is redescribed and revised. Females are grouped into five species-groups, with eleven new species described based on females: I. acuminata Waldren, sp. nov., I. australis Waldren, sp. nov., I. bimaculata Waldren, sp. nov., I. breviclypeata Waldren, sp. nov., I. chihuahuensis Waldren, sp. nov., I. cuernavaca Waldren, sp. nov., I. erythrocephala Waldren, sp. nov., I. manleyi Waldren, sp. nov., I. mesomexicana Waldren, sp. nov., I. suarezi Waldren, sp. nov., and I. tequila Waldren, sp. nov. One new combination is included, I. cephalargia (Mickel, 1924), comb. nov., formerly placed in Pseudomethoca Ashmead, 1896. Additionally, I. curoei Quintero Cambra, 2011 is synonymized with I. cardinalis (Gerstaecker, 1874), syn. nov., and I. megacantha (Cockerell Casad, 1894), stat. resurr., is treated as a nomen dubium. The lectotype of Mutilla satrapa Gerstaecker, 1874 originally designated by C. Mickel is here validated. An illustrated key to species is included.

New and unusual host records for North American and South American spider wasps (Hymenoptera: Pompilidae).

New and unusual host records for 133 species and subspecies of Pompilidae predominantly from the southwestern United States, Mexico, Central America, and South America are presented in modified taxonomic order. First-time species host records are given for Calopompilus Ashmead, Pepsis Fabricius, Hemipepsis Dahlbom, Priocnessus Banks, Entypus Dahlbom, Pompilocalus Roig-Alsina, Sphictostethus Kohl, Auplopus Spinola, Ageniella Banks, Eragenia Banks, Aporus Spinola, Poecilopompilus Ashmead, Tachypompilus Ashmead, Anoplius Dufour, Priochilus (Fabricius) and Notocyphus Smith. New host spider families are introduced for Calopompilus, Pepsis, Hemipepsis, Priocnessus, Entypus, Cryptocheilus Panzer, Priocnemis Schiødte, Auplopus, Ageniella, Eragenia, Aporus, Tachypompilus, Anoplius, Priochilus and Notocyphus. Eight host spider families are reported from the Western Hemisphere for the first time: Halonoproctidae (Notocyphus dorsalis dorsalis Cresson); Dipluridae (Pepsis pretiosa Dahlbom, P. montezuma Smith, P. infuscate Spinola, P. atripennis Fabricius, P. martini Vardy, Priocnessus vancei Waichert and Pitts); Nemesiidae (Pepsis pallidolimbata Lucas, P. viridis Lepeletier, P. spp., Pompilocalus hirticeps (Guérin), Sphictostethus gravesii (Haliday), S. striatulus Roig-Alsina, Priocnemis oregona Banks); Barychelidae (Eragenia sp.); Paratropididae (Pepsis stella Montet); Trechaleidae (Hemipepsis toussainti (Banks), Entypus unifasciatus cressoni (Banks), Tachypompilus ferrugineus (Say), Tachypompilus unicolor cerinus Evans, Priochilus gloriosum (Cresson); Desidae (Ageniella accepta (Cresson), Sphictostethus isodontus Roig-Alsina) and Selenopidae (Priochilus scrupulum (Fox), Tachypompilus erubescens (Taschenberg) or xanthopterus (Rohwer)). The first known host records for the rare South American pompilid genera Chirodamus (Lycosidae: Lycosa sp.) and Herbstellus (Nemesiidae: Diplothelopsis cf bonariensis Mello-Leitão) are presented.

Re-examination of Australian and Oriental species of Minagenia Banks, 1934 (Hymenoptera: Pompilidae), with a new record for the genus in Australia and a new species description.

The genus Minagenia Banks, 1934 is reported for the first time from Australia. A new species, M. lasallei Decker, Rodriguez Pitts, sp. nov. (Australia: from Queensland to Tasmania), is described and illustrated. A key to the four species from the Australasian and Oriental regions is given. Three species are removed from Asian Minagenia: granulosa Tsuneki, 1989, comb. nov., to the genus Priocnemis Schiødte, 1837; pempuchiensis Tsuneki, 1989, comb. nov., to the genus Nipponodipogon Ishikawa, 1965; and assimilis Smith, 1861, comb. nov., to the genus Dichragenia Haupt, 1950. Minagenia alticola Tsuneki, 1989 is a member of an undescribed genus. Placement of this species is maintained in Minagenia until the new genus can be described in a future work. Minagenia brevicornis (Banks, 1934) is redescribed.

Addition to the taxonomic and distribution records of Abernessia Arlé (Hymenoptera: Pompilidae).

Species of spider wasps (Hymenoptera: Pompilidae) are diverse in the tropical regions. The Neotropical genus Abernessia Arlé, 1947 was described from São Paulo and later found in Espírito Santo, Bahia and Minas Gerais, Brazil. Herein, we establish a neotype for the type-species A. irmgardae Arlé, discuss the previous distribution records of Abernessia in South America, add new records for Brazil, and expand the recorded distribution to San Pedro, Paraguay. The male of Abernessia giga is described and illustrated. Finally, an updated key to species known from males is provided.

Thistledown velvet ants in the Desert Mimicry Ring and the evolution of white coloration: Müllerian mimicry, camouflage and thermal ecology.

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.

Redescription of Neanoplius Banks (Hymenoptera: Pompilidae).

Neanoplius Banks, 1947 is a monotypic genus of Neotropical spider wasp. It has been ignored in taxonomic works since its description. We redescribe the genus and type species, provide illustrations, provide additional characters to help separate it from other closely related genera, and provide a discussion of probable evolutionary relationships of this interesting genus.

Review of the genus Lomachaeta Mickel, 1936 (Hymenoptera: Mutillidae) with new species and sex associations.

New species and sex associations are proposed in the genus Lomachaeta Mickel, 1936. Six new species are described: Lomachaeta calamondin Williams, L. eironeia Williams, L. juanita Williams, L. meloi Williams, L. osita Williams, and L. warneri Williams. The previously unknown females of five species are described: L. beadugrimi (Pitts Manley, 2004); L. chionothrix Pitts Manley, 2004; L. hedera Williams Pitts, 2009; L. hyphantria Pitts Manley, 2004; and L. vacamuerta Williams Pitts, 2009. The previously unknown male of L. vianai Casal, 1969 is described and L. megomicron Williams Pitts, 2009 is placed as its synonym. Various new country and state records are presented. New keys and habitus pictures for all the 24 known species are provided.

Revision of the bee group Anthophora (Micranthophora) (Hymenoptera: Apidae), with notes on potential conservation concerns and a molecular phylogeny of the genus.

Anthophora (Micranthophora) comprises the largest subgenus of Anthophora in the Western Hemisphere, with 26 species. Though previously synonymized with the Anthophora (Heliophila), A. (Micranthophora) is here confirmed as a subgenus, morphological and multi-gene molecular phylogenetic evidence refute this and the synonymy of A. (Micranthophora) to A. (Heliophila) is formally rejected. Characters for (Micranthophora) are given and a key to species is provided. Seven new species are described: Anthophora (Micranthophora) caudata Orr, sp. nov.; Anthophora (Micranthophora) chihuahua Orr, sp. nov.; Anthophora (Micranthophora) escalante Orr, sp. nov.; Anthophora (Micranthophora) parkeri Orr, sp. nov.; Anthophora (Micranthophora) rara Orr, sp. nov.; Anthophora (Micranthophora) striata Orr, sp. nov.; and Anthophora (Micranthophora) timberlakei Orr, sp. nov. Further, four new synonyms are enacted: A. arthuri and A. flexipes to A. albata, A. nigritula to A. exigua, and A. xanthochlora to A. pachyodonta. Distributional and phenological data, as well as known details of nesting biology and floral specialization, are provided. Future research directions and species of potential conservation interest are also discussed.

Phylogeny and population genetic analyses reveals cryptic speciation in the Bombus fervidus species complex (Hymenoptera: Apidae).

Bumble bees (Bombus Latrielle) are significant pollinators of flowering plants due to their large body size, abundant setae, and generalist foraging strategies. However, shared setal coloration patterns among closely and distantly related bumble bee species makes identification notoriously difficult. The advent of molecular genetic techniques has increased our understanding of bumble bee evolution and taxonomy, and enables effective conservation policy and management. Individuals belonging to the North American Bombus fervidus species-complex (SC) are homogenous in body structure but exhibit significant body color phenotype variation across their geographic distribution. Given the uncertainty of the genealogical boundaries within the SC, some authors have synonymized all members of the B. fervidus SC within a single taxon, while others propose an alternative two taxa hypothesis. Operating under the phylogenetic species concept, our analysis supports the hypothesis that there are two independent lineages of bumble bees within the B. fervidus SC. With the current evidence, however, it is not possible to assign valid names to either of them, because both lineages include the color phenotypes found in the original species descriptions of B. fervidus and B. californicus. Cryptic speciation does not seem to be the product of Müllerian mimicry between the clades, because diverging coloration patterns are observed when the distribution of the clades overlaps. Furthermore, within each lineage there is evidence for strong population differentiation that is correlated with geographic distribution rather than color phenotype. In our study, we demonstrate the importance of obtaining a broad sample of multiple populations when conducting lower-level phylogenetic analyses. In addition to improving our knowledge of bumble bee diversification patterns, characterizing the evolutionary history of these pollinators provides the foundation needed to guide contemporary conservation assessments and management strategies.

Stinging wasps (Hymenoptera: Aculeata), which species have the longest sting?

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.

Taxonomic contributions to Ageniella Banks, 1912 (Hymenoptera: Pompilidae) from Brazil.

Ageniella Banks is the second most diverse genus in the tribe Ageniellini (Pompilidae: Pepsinae) with about 200 valid names. Ageniella is known to be paraphyletic; yet, no revision has been made. Lack of a taxonomic catalogue and of identification keys has further delayed taxonomic studies. Additionally, many of the currently valid names should be treated as synonyms. This present study is the first attempt to organize the species of Ageniella, focusing on the species occurring in Brazil. A total of six subgenera and 40 species of Ageniella occur in Brazil. Of these, 25 species are endemic to Brazil. The new synonyms are proposed for (valid names are first): Alasagenia Banks, 1944 =Lissagenia Banks, 1946, syn. nov.; Ageniella (Alasagenia) erichsoni Banks, 1944 =Priophanes major Banks, 1945, syn. nov.; Ageniella (Ameragenia) zeteki (Banks, 1925) =Priophanes marcida Banks, 1946, syn. nov.; Ageniella (Ameragenia) agitata (Smith, 1873), comb. nov. =Salius (Priocnemis) setaceicornis Fox, 1897, syn. nov.; Ageniella (Ameragenia) sanguinolenta (Smith, 1864) =Agenia ruficeps Smith, 1864, syn. nov.; =Ageniella alternata Banks, 1946, syn. nov.; Ageniella (Priophanes) dolorosa (Banks, 1946), comb. nov. =Priophanes plagosa Banks, 1946, syn. nov.; Ageniella (Ameragenia) fabricii (Banks, 1944) =Ameragenia notabilis Banks, 1946, syn. nov. A new combination is proposed for: Ageniella (Alasagenia) cursor (Smith, 1873), comb. nov. (from Agenia Schiødte); Ageniella (Ameragenia) clypeata (Fox, 1897), comb. nov. (from Salius Fabricius), Ageniella (Ameragenia) rutila (Fox, 1897), comb. nov. (from Salius), Ageniella (Priophanes) tegularis (Fox, 1897), comb. nov. (from Salius), Ageniella (Priophanes) rufitarsis (Fox, 1897), comb. nov. (from Salius), Ageniella (Ameragenia) citricornis (Fox, 1897), comb. nov. (from Salius), Ageniella (Ameragenia) serrula (Fox, 1897), comb. nov. (from Salius), Ageniella (Priophanes) cingulata (Fox, 1897), comb. nov. (from Agenia). A new name Ageniella (Ameragenia) banksii Waichert, nom. nov. is proposed for Ameragenia festina Banks, 1946, secondary homonym of Ageniella festina Banks, 1917, and Ageniella (Priophanes) otiosa is reinstate, stat. resurr.

The geological record and phylogeny of spider wasps (Hymenoptera: Pompilidae): A revision of fossil species and their phylogenetic placement.

Accurate fossil identification has become increasingly relevant with the widespread use of phylogenetic divergence time estimation methods, which rely on fossil data to determine clade hard-minimum ages. Here we revise, diagnose and illustrate known spider wasp (Hymenoptera: Pompilidae) fossil species and place them within the latest Pompilidae phylogenetic hypothesis. Ceropalites infelix Cockerell, from the Florissant Fossil Beds (Priabonian), is no longer recognized as Pompilidae, but as Aulacidae. Agenioideus saxigenus (Cockerell) comb. nov., Deuteragenia wettweri (Statz) comb. nov., Caputelus scudderi (Cockerell, 1906) comb. nov., Pepsinites avitula (Cockerell, 1941) comb. nov., Pepsinites contentus (Theobald, 1937) comb. nov., Pepsinites florissantensis (Cockerell, 1906) comb. nov., Pepsinites laminarum (Rohwer, 1909) comb. nov., Pepsinites scelerosus (Meunier, 1919) comb. nov., Pepsinites cockerellae (Rohwer, 1909) comb. nov., Pompilinites coquandi (Theobald, 1937) comb. nov., Pompilinites depressus (Statz, 1936) comb. nov., Pompilites incertus (Theobald, 1937) comb. nov., Pompilites induratus (Heer, 1849) comb. nov., Pompilites fasciatus (Theobald, 1937) comb. nov., and Pompilites senex comb. nov. are new combinations. Twenty-three fossil species of spider wasps are now recognized in 13 genera. Four new genera are proposed: Caputelus Waichert & Pitts gen. nov., Pompilites Rodriguez gen. nov., Pompilinites Rodriguez & Waichert gen. nov., and Pepsinites Rodriguez & Waichert gen. nov., of which the three latter are collective-group names for fossils with taxonomic uncertainty. One species of fossil spider wasp is described: Deuteragenia catalunyia Rodriguez, Waichert & Pitts sp. nov., from the Bellver deposits in Catalonia, Spain. Five of the 23 known species can be used to determine hard-minimum age for calibrations of genera stem-groups (Agenioideus, Anoplius, Cryptocheilus, Deuteragenia, Priocnemis). The fossil belonging to the stem-group of the tribe Ageniellini (Chubutholites) is not recommended for calibration because of the high uncertainty in its age and taxonomy. The remaining taxa can be assigned to the lineage comprising Pompilinae + Pepsinae (12 species) or crown-group Pompilidae (four species).

More new records of spider wasps from Colombia (Hymenoptera, Pompilidae).

Aporinellus Banks, Austrochares Banks and Dicranoplius Haupt are new generic records for Colombia, as well as the species Dipogon ariel Banks, Evagetes peruana Banks, and Euplaniceps notabilis (Smith). Five new combinations are formally endorsed: Aimatocare argentinica (Banks), comb. n.; Aimatocare longula (Banks), comb. n.; Aimatocare imitator (Evans), comb. n.; Aimatocare impensa (Evans), comb. n.; Aimatocare vitrea (Fox), comb. n. Although these names have been used in Pompilidae, no formal nomenclatural act had been proposed. The presence of Chirodamus paramicola Roig-Alsina, previously reported with uncertainty, is confirmed. Finally, a new combination for Euplaniceps notabilis (Smith), comb. n. is proposed based on molecular phylogenetics and morphological data. The Colombian fauna of Pompilidae sums up to 38 genera and approximately 150 species.

Phylogenomic Insights into the Evolution of Stinging Wasps and the Origins of Ants and Bees.

The stinging wasps (Hymenoptera: Aculeata) are an extremely diverse lineage of hymenopteran insects, encompassing over 70,000 described species and a diversity of life history traits, including ectoparasitism, cleptoparasitism, predation, pollen feeding (bees [Anthophila] and Masarinae), and eusociality (social vespid wasps, ants, and some bees) [1]. The most well-studied lineages of Aculeata are the ants, which are ecologically dominant in most terrestrial ecosystems [2], and the bees, the most important lineage of angiosperm-pollinating insects [3]. Establishing the phylogenetic affinities of ants and bees helps us understand and reconstruct patterns of social evolution as well as fully appreciate the biological implications of the switch from carnivory to pollen feeding (pollenivory). Despite recent advancements in aculeate phylogeny [4-11], considerable uncertainty remains regarding higher-level relationships within Aculeata, including the phylogenetic affinities of ants and bees [5-7]. We used ultraconserved element (UCE) phylogenomics [7, 12] to resolve relationships among stinging-wasp families, gathering sequence data from >800 UCE loci and 187 samples, including 30 out of 31 aculeate families. We analyzed the 187-taxon dataset using multiple analytical approaches, and we evaluated several alternative taxon sets. We also tested alternative hypotheses for the phylogenetic positions of ants and bees. Our results present a highly supported phylogeny of the stinging wasps. Most importantly, we find unequivocal evidence that ants are the sister group to bees+apoid wasps (Apoidea) and that bees are nested within a paraphyletic Crabronidae. We also demonstrate that taxon choice can fundamentally impact tree topology and clade support in phylogenomic inference.

A new bee species that excavates sandstone nests.

Humanity has long been fascinated by animals with apparently unfavorable lifestyles [1]. Nesting habits are especially important because they can limit where organisms live, thereby driving population, community, and even ecosystem dynamics [2]. The question arises, then, why bees nest in active termite mounds [3] or on the rim of degassing volcanoes, seemingly preferring such hardship [4]. Here, we present a new bee species that excavates sandstone nests, Anthophora (Anthophoroides) pueblo Orr (described in Supplemental Information, published with this article online), despite the challenges already inherent to desert life. Ultimately, the benefits of nesting in sandstone appear to outweigh the associated costs in this system.

Association of the female of Perdita (Xeromacrotera) cephalotes (Cresson), and a replacement name for Perdita bohartorum Parker (Hymenoptera: Andrenidae).

The monotypic Perdita subgenus Xeromacrotera Timberlake is currently known only from the male sex. Here, the female of Perdita (Xeromacrotera) cephalotes (Cresson, 1878) is associated, resulting in two new junior synonyms of P. (X.) cephalotes: Perdita (Procockerellia) excellens Timberlake, 1958, new synonym and Perdita (Cockerellia) autumnalis Timberlake, 1977, new synonym. The true number of maxillary palpi are reported, and updated subgeneric and species diagnoses are provided. In addition, Perdita boharti Portman & Griswold, new name, is designated as a replacement name for the primary junior homonym P. bohartorum Parker, 1983 (not P. bohartorum Timberlake, 1956) and is assigned to the related subgenus Cockerellia Ashmead.

Molecular phylogeny of Pompilinae (Hymenoptera: Pompilidae): Evidence for rapid diversification and host shifts in spider wasps.

Pompilinae is one of the largest subfamilies of spider wasps (Pompilidae). Most pompilines are generalist spider predators at the family level, but some taxa exhibit ecological specificity (i.e., to spider-host guild). Here we present the first molecular phylogenetic analysis of Pompilinae, toward the aim of evaluating the monophyly of tribes and genera. We further test whether changes in the rate of diversification are associated with host-guild shifts. Molecular data were collected from five nuclear loci (28S, EF1-F2, LWRh, Wg, Pol2) for 76 taxa in 39 genera. Data were analyzed using maximum likelihood (ML) and Bayesian inference (BI). The phylogenetic results were compared with previous hypotheses of subfamilial and tribal classification, as well as generic relationships in the subfamily. The classification of Pompilus and Agenioideus is also discussed. A Bayesian relaxed molecular clock analysis was used to examine divergence times. Diversification rate-shift tests accounted for taxon-sampling bias using ML and BI approaches. Ancestral host family and host guild were reconstructed using MP and ML methods. Ancestral host guild for all Pompilinae, for the ancestor at the node where a diversification rate-shift was detected, and two more nodes back in time was inferred using BI. In the resulting phylogenies, Aporini was the only previously proposed monophyletic tribe. Several genera (e.g., Pompilus, Microphadnus and Schistonyx) are also not monophyletic. Dating analyses produced a well-supported chronogram consistent with topologies from BI and ML results. The BI ancestral host-use reconstruction inferred the use of spiders belonging to the guild "other hunters" (frequenting the ground and vegetation) as the ancestral state for Pompilinae. This guild had the highest probability for the ML reconstruction and was equivocal for the MP reconstruction; various switching events to other guilds occurred throughout the evolution of the group. The diversification of Pompilinae shows one main rate-shift coinciding with a shift to ground-hunter spiders, as reconstructed by the BI ancestral character-state analysis.

North American velvet ants form one of the world's largest known Müllerian mimicry complexes.

Color mimicry is often celebrated as one of the most straightforward examples of evolution by natural selection, as striking morphological similarity between species evolves in response to a shared predation pressure. Recently, a large North American mimetic complex was described that included 65 species of Dasymutilla velvet ants (Hymenoptera: Mutillidae). Beyond those 65 species, little is known about how many species participate in this unique Müllerian complex, though several other arthropods are thought to be involved as Müllerian mimics (spider wasps) and Batesian mimics (beetles, antlions, and spiders; see references in). Müllerian mimicry is similarity in appearance or phenotype among harmful species, while Batesian mimicry is similarity in which not all species are harmful. Here, we investigate the extent of the velvet ant mimicry complex beyond Dasymutilla by examining distributional and color pattern similarities in all of the 21 North American diurnal velvet ant genera, including 302 of the 361 named species (nearly 84%), as well as 16 polymorphic color forms and an additional 33 undescribed species. Of the 351 species and color forms that were analyzed (including undescribed species), 336 exhibit some morphological similarities and we hypothesize that they form eight distinct mimicry rings (Figure 1A; Supplemental Information). Two of these eight mimicry rings, red-headed Timulla and black-headed Timulla, were not documented in earlier assessments of mimicry in velvet ants, and are newly described here. These findings identify one of the largest known Müllerian mimicry systems worldwide and provide a novel system to test hypotheses about aposematism and mimicry, especially those regarding the evolution of imperfect mimicry.

Description of a new species and species-group of Sphaeropthalma Blake Hymenoptera: Mutillidae) with an updated classification of the genus.

A study of Sphaeropthalma Blake from the southwestern USA has revealed a highly autapomorphic undescribed species. This species is described here as S. tapio Pitts & Sadler, sp. nov., and is placed in a new species-group, the S. tapio species-group. This species seems to be found only at higher altitudes in southern Arizona. It is compared to other similar Sphaeropthalma species-groups and is likely most closely related to members of the S. imperialis species-group. An updated species-group classification of Sphaeropthalma is provided.

New additions to the Honduran fauna of spider wasps (Hymenoptera: Pompilidae) with the description of two species.

Two new species of pepsine spider wasps, Calopompilus setaerotundus Waichert & Pitts, sp. nov. and Priocnemis (Umbripennis) wasbaueri Waichert & Pitts, sp. nov., are described and illustrated. Calopompilus Ashmead, Priocnemis Schiødte and sixteen additional species are newly reported from Honduras. A new combination is proposed for Ageniella fragilis (Fox), comb. nov., which is transferred from Agenia Schiødte.