Juanita Rodriguez

Spider phylogenomics: untangling the Spider Tree of Life

Spiders (Order Araneae) are massively abundant generalist arthropod predators that are found in nearly every ecosystem on the planet and have persisted for over 380 million years. Spiders have long served as evolutionary models for studying complex mating and web spinning behaviors, key innovation and adaptive radiation hypotheses, and have been inspiration for important theories like sexual selection by female choice. Unfortunately, past major attempts to reconstruct spider phylogeny typically employing the “usual suspect” genes have been unable to produce a well-supported phylogenetic framework for the entire order. To further resolve spider evolutionary relationships we have assembled a transcriptome-based data set comprising 70 ingroup spider taxa. Using maximum likelihood and shortcut coalescence-based approaches, we analyze eight data sets, the largest of which contains 3,398 gene regions and 696,652 amino acid sites forming the largest phylogenomic analysis of spider relationships produced to date. Contrary to long held beliefs that the orb web is the crowning achievement of spider evolution, ancestral state reconstructions of web type support a phylogenetically ancient origin of the orb web, and diversification analyses show that the mostly ground-dwelling, web-less RTA clade diversified faster than orb weavers. Consistent with molecular dating estimates we report herein, this may reflect a major increase in biomass of non-flying insects during the Cretaceous Terrestrial Revolution 125–90 million years ago favoring diversification of spiders that feed on cursorial rather than flying prey. Our results also have major implications for our understanding of spider systematics. Phylogenomic analyses corroborate several well-accepted high level groupings: Opisthothele, Mygalomorphae, Atypoidina, Avicularoidea, Theraphosoidina, Araneomorphae, Entelegynae, Araneoidea, the RTA clade, Dionycha and the Lycosoidea. Alternatively, our results challenge the monophyly of Eresoidea, Orbiculariae, and Deinopoidea. The composition of the major paleocribellate and neocribellate clades, the basal divisions of Araneomorphae, appear to be falsified. Traditional Haplogynae is in need of revision, as our findings appear to support the newly conceived concept of Synspermiata. The sister pairing of filistatids with hypochilids implies that some peculiar features of each family may in fact be synapomorphic for the pair. Leptonetids now are seen as a possible sister group to the Entelegynae, illustrating possible intermediates in the evolution of the more complex entelegyne genitalic condition, spinning organs and respiratory organs.

Müllerian Mimicry as a Result of Codivergence between Velvet Ants and Spider Wasps

Recent studies have delineated a large Nearctic Müllerian mimicry complex in Dasymutilla velvet ants. Psorthaspis spider wasps live in areas where this mimicry complex is found and are phenotypically similar to Dasymutilla. We tested the idea that Psorthaspis spider wasps are participating in the Dasymutilla mimicry complex and that they codiverged with Dasymutilla. We performed morphometric analyses and human perception tests, and tabulated distributional records to determine the fit of Psorthaspis to the Dasymutilla mimicry complex. We inferred a dated phylogeny using nuclear molecular markers (28S, elongation factor 1-alpha, long-wavelength rhodopsin and wingless) for Psorthaspis species and compared it to a dated phylogeny of Dasymutilla. We tested for codivergence between the two groups using two statistical analyses. Our results show that Psorthaspis spider wasps are morphologically similar to the Dasymutilla mimicry rings. In addition, our tests indicate that Psorthaspis and Dasymutilla codiverged to produce similar color patterns. This study expands the breadth of the Dasymutilla Müllerian mimicry complex and provides insights about how codivergence influenced the evolution of mimicry in these groups.

Eocene and not Cretaceous Origin of Spider Wasps: Fossil Evidence from Amber

Spider wasps had long been proposed to originate in the Late Cretaceous based on the Burmese amber fossil Bryopompilus interfector. We performed a morphological examination of this fossil and determined that it does not belong to Pompilidae or any other described hymenopteran family. Instead, we place it in the new family Bryopompilidae. The oldest verifiable member of the Pompilidae is from Baltic amber, which suggests that the crown group of the family probably originated in the Eocene, not in the Late Cretaceous as previously proposed. The origin of spider wasps appears to be correlated with an increase in spider familial diversity in the Cenozoic. We also add two genera to the extinct pompilid fauna: Tainopompilus Rodriguez and Pitts gen. nov. and Paleogenia Waichert and Pitts gen. nov., and describe three new species of fossil spider wasps: Anoplius planeta Rodriguez and Pitts sp. nov., from Dominican amber (Burdigalian to Langhian); Paleogenia wahisi Waichert and Pitts sp. nov., from Baltic amber (Lutetian to Priabonian); and Tainopompilus argentum Rodriguez and Pitts sp. nov., from Dominican amber (Chattian to Langhian).

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).

Phylogeography and evolution of the Red Salamander (Pseudotriton ruber)

Phylogeographic studies frequently result in the elevation of subspecific taxa to species given monophyly, or the synonymy of subspecies that are not monophyletic. However, given limited or incongruent datasets, retention of subspecies can be useful to describe hypothesized incipient species or to illustrate interesting biological phenomena driving morphological diversity. Four subspecific taxa have been used to describe largely allopatric geographic variation within the species Pseudotriton ruber, a plethodontid salamander occupying stream and spring habitats across eastern North America: P. r. vioscai occurs in lowland Coastal Plain habitats, while P. r. ruber, P. r. nitidus, and P. r. schencki occupy upland regions in and around the Appalachian Mountains. Pseudotriton ruber co-occurs through its distribution with the aposematic newt Notophthalmus viridescens, and both species are hypothesized to be part of a Müllerian mimicry complex. In this study, we sequenced regions of two mitochondrial (cytochrome b, NADH dehydrogenase subunit 2) and one single copy nuclear protein-coding gene (pro-opiomelanocortin) from individuals sampled across much of the distribution of P. ruber and then used maximum-likelihood and Bayesian phylogenetic inference to test the monophyly of subspecies, reconstruct biogeographic history, and make inferences about morphological evolution. Phylogeographic hypotheses from mitochondrial and nuclear datasets described structure among populations of P. ruber which separated Coastal Plain and upland Appalachian populations, but subspecies were not monophyletic. Biogeographic reconstruction estimated the ancestor of all populations to have occupied and initially diverged in the Coastal Plain during the Pliocene (∼3.6mya), before one lineage subsequently invaded upland areas of Appalachia. Bold bright coloration of high elevation subspecies P. r. nitidus and P. r. schencki appears to have evolved twice. We hypothesize that the Müllerian mimicry complex with N. viridescens and P. ruber may provide a selective mechanism driving the co-evolution of striking bright and dull morphological variation among populations of both species. While P. ruber subspecies were not consistent with our criteria for diagnosing species (monophyly) and therefore could not be elevated to species, we advocate for the retention of subspecies because they describe hypotheses about an incipient species (P. r. vioscai) and how Müllerian mimicry may shape morphological diversity of species.

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

Abstract New records of genera and species of spider wasps (Hymenoptera: Pompilidae) from Colombia are provided. Agenioideus, Cryptocheilus, Evagetes, Mystacagenia, and Xerochares are newly recorded genera from Colombia. Nineteen species are first recorded from Colombia: Aimatocare vitrea (Fox); Ageniella azteca (Cameron); Ageniella curtipinus (Cameron); Ageniella fallax (Arlé); Ageniella hirsuta Banks; Ageniella pilifrons (Cameron); Ageniella pretiosa Banks; Ageniella sanguinolenta (Smith); Ageniella zeteki (Banks); Agenioideus birkmanni (Banks); Aporus (Aporus) cuzco Evans; Aporus (Cosmiaporus) diverticulus (Fox); Aporus (Notoplaniceps) canescens Smith; Euplaniceps exilis (Banks); Euplaniceps herbertii (Fox); Irenangelus clarus Evans; Mystacagenia bellula Evans; Phanochilus nobilitatus (Smith) and Xerochares expulsus Schulz. The following species and genera have their occurence ranges expanded for South America: Ageniella azteca (Cameron); Ageniella zeteki (Banks); Agenioideus birkmanni (Banks); and Xerochares expulsus Schulz; Cryptocheilus Panzer; and Xerochares Evans.

Temporal Activity Patterns of the Spider Wasp Pepsis montezuma Smith (Hymenoptera: Pompilidae) in a Disturbed Lower Montane Rainforest (Manizales, Colombia)

We studied the temporal activity pattern of the spider wasp Pepsis montezuma Smith (Hymenoptera: Pompilidae) in a disturbed lower montane rainforest, which is located in the city of Manizales, Colombia, at an altitude of 2,150 m. Females of this species are diurnal with two peaks of activity: one in the morning and the other in the afternoon. During the morning, nectar foraging occurred at Baccharis latifolia. During the afternoon, females hunted for tarantulas of the genus Pamphobeteus (Araneae: Theraphosidae), which were dragged backwards to the nest by the wasp. The nest was excavated before hunting. This is the first description of the behavior of Pepsis montezuma.

Assessing Species Boundaries and Sex-Associations in the Genus Drepanaporus (Hymenoptera: Pompilidae), with Comparison of the Utility of Cytochrome c Oxidase I and a Nuclear Molecular Marker, and the Description of a New Species of Drepanaporus

ABSTRACT The taxonomy of the Antillean genus Drepanaporus Bradley (Pompilidae) is problematic because of sexual dimorphism and nearly uniform morphology of males across species. Species limits are not well understood, and sexes are not properly associated in all species. In this study, we reassessed morphology, and collected novel molecular data for the purpose of determining species boundaries and establishing sex-associations for all species. Two genes, cytochrome c oxidase (COI) and long-wavelength rhodopsin (LWRh) were amplified for 20 specimens of Drepanaporus, from both females and males. Using LWRh (including both introns and exons) and COI sequences, the relationships of Drepanaporus samples were reconstructed. Sequences obtained for COI showed unusually high divergences and putative introgression, and thus were not used for taxonomic decisions. Taxonomic changes were made based on the LWRh phylogenetic results. A new species of Drepanaporus— Drepanaporus bachata sp. nov—is described herein based on both molecular and morphological characters for both male and female specimens, and a key is provided for the genus for the three species now recognized. We also discuss the usefulness of mitochondrial and nuclear markers in Pompilidae for species delimitations and sex-associations.

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

Abstract 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.