Progress 07/01/12 to 06/30/17

Outputs
Target Audience:The target audiences for this research include evolutionary biologists, entomologists, taxonomists, biogeographers, geologists, and academics. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?I have mentored 6 graduate students and 8 undergraduate students in the aspects of this project in the past year. How have the results been disseminated to communities of interest?The results have been disseminated via 32 publications and 20 presentations at various scientific meetings held in the USA. What do you plan to do during the next reporting period to accomplish the goals?This is the final report for this project. Specific objectives for my new project include: 1. Continue to extract DNA from further nocturnal and diurnal species, begin gathering phylogenomic data (UCEs), which is new for this project, from all nocturnal species that have not been sequenced yet. 2. Perform a phylogenetic analysis using new phylogenomic data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions extending into Central America determining the timing of speciation events and establishing corresponding geological events.

Impacts
What was accomplished under these goals? 1. We extracted DNA from samples, amplified and sequenced the mitochondrial gene (cytochrome c oxidase I [COI], nuclear gene (opsin [op]) and two nuclear intergenic regions (internal transcribed spacer regions 1&2 [ITS1&2]) from 70 nocturnal species and 5 outgroups. 2. We completed the Mark-Recapture and cage studies in southern Utah this summer after expanding it to include Chyphotidae and Brachycistidinae. 3. We were unable to determine hosts for nocturnal wasp species by locating would-be hosts' nests, collecting hosts and rearing host cocoons and pupae that may harbor nocturnal wasps this year. 4. We curated (mounted, labeled and identified) over 350,000 specimens of nocturnal wasps and related Hymenoptera from North and Central America collected this summer or left over from last summer. 5. We described two new species of pepsine spider wasps, Calopompilus setaerotundus Waichert & Pitts, sp. nov. and Priocnemis (Umbripennis) wasbaueri Waichert & Pitts, sp. nov. 6. Eragenia Banks was revalidated (stat. rev.) and applied to 16 species, eight of which were newly described: Eragenia bella Waichert & Pitts sp.n., E. carinata Waichert & Pitts sp.n., E. dentata Waichert & Pitts sp.n., E. oliva Waichert & Pitts sp.n., E. pseudomicans Waichert & Pitts sp.n., E. setosa Waichert & Pitts sp.n., E. rotunda Waichert & Pitts sp.n.,and E. villosa Waichert & Pitts sp.n. 7. Taxonomic changes were made to Drepanaporus based on the LWRh phylogenetic results and a new species of Drepanaporus, Drepanaporus bachata sp. nov, was newly described based on both molecular and morphological characters for both male and female specimens. 8. Psorthaspis was determined to mimic diurnal mutillids and the species of this spiderwasp was added to currently defined Dasymutilla mimicry rings. 9. We incorporated specimens brought by the merger of the American Entomological Institute in the summer of 2016. 10. Accomplishments from publications include a study of Sphaeropthalma Blake from the southwestern USA that revealed a highly autapomorphic undescribed species that I described as S. tapio Pitts & Sadler, sp. nov., and placed in a new species-group, the S. tapio species-group. 11. We reconstructed a molecular phylogeny of Pompilidae from Bayesian and maximum-likelihood analyses of four nuclear molecular markers (elongation factor-1 α F2 copy, longwavelength rhodopsin, RNA polymerase II, and 28S ribosomal RNA), performed Bayesian divergence-time estimation analysis using four calibration points and ancestral-area reconstruction with a Bayesian binary Markov chain Monte Carlo method. New relationships for spider wasps were discovered, and new subfamilial delimitations are proposed and discussed based on the phylogeny. The origin of Pompilidae was c. 43.3 Mya, probably in the Nearctic region. Most of the extant subfamilies originated from the late Eocene to the Oligocene, and their current distributions are the product of various dispersal events that occurred over the course of ∼40 Mya. This is the first phylogenetic reconstruction of Pompilidae from molecular characters, with broad geographical and taxonomic sampling. 12. We discovered that North American velvet ants form one of the world's largest known Müllerian mimicry Complexes. 13. We identified over 50,000 specimens of spider wasps from Central and South America, which were mounted and curated from the previous three summers. 14. We investigated the role of taxon sampling in causing incongruent results between two recent phylogenomic studies of stinging wasps (Hymenoptera: Aculeata), a diverse lineage that includes ants, bees and the majority of eusocial insects. Using target enrichment of ultraconserved element (UCE) loci, the largest aculeate phylogenomic data set to date has been assembled, sampling 854 loci from 187 taxa, including 30 out of 31 aculeate families, and a diversity of parasitoid outgroups. We analyzed the complete matrix using multiple analytical approaches, and also performed a series of taxon inclusion/exclusion experiments, in which we analyzed taxon sets identical to and slightly modified from the previous phylogenomic studies. The results provide a highly supported phylogeny for virtually all aculeate lineages sampled, supporting ants as sister to Apoidea (bees+apoid wasps), bees as sister to Philanthinae+Pemphredoninae (lineages within a paraphyletic Crabronidae), Melittidae as sister to remaining bees, and paraphyly of cuckoo wasps (Chrysidoidea). Divergence dating analyses estimate ages for aculeate lineages in close concordance with the fossil record. Our analyses also demonstrate that outgroup choice and taxon evenness can fundamentally impact topology and clade support in phylogenomic inference. 15. We found that spider wasps, which had long been proposed to originate in the Late Cretaceous based on the Burmese amber fossil Bryopompilus interfector, did not belong to Pompilidae or any other described hymenopteran family. Instead, we placed it into a 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).

Publications

• Type: Conference Papers and Presentations Status: Other Year Published: 2012 Citation: Pitts, J., Systematics and Biodiversity of Hymenoptera in Boreal and Hemiboreal Ecosystems Symposium, "Molecular Phylogenetics of the Vespoidea (Hymenoptera)," Smithsonian, Tovetorp forskningsstation, Sweden, Aug. 7, 2012. (August 11, 2012)
• Type: Conference Papers and Presentations Status: Other Year Published: 2012 Citation: Pitts, J., HymCourse, "Systematics of Chrysidoidea and Vespoidea (Hymenoptera)," Smithsonian, Tovetorp forskningsstation, Sweden, Aug. 7, 2012. (August 7, 2012)

Progress 10/01/15 to 09/30/16

Outputs
Target Audience:Target Audience The target audiences for this research include evolutionary biologists, entomologists, taxonomists, biogeographers, geologists, and academics. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?Opportunities I have mentored 2 graduate students and 2 undergraduate students in the aspects of this project in the past year. How have the results been disseminated to communities of interest?Dissemination The results have been disseminated via five publications and seven presentations at various scientific meetings held in the USA. What do you plan to do during the next reporting period to accomplish the goals?Plan of Work Specific objectives include: 1. Continue to extract DNA from further nocturnal species, begin gathering phylogenomic data (UCEs), which is new for this project, from all nocturnal species that have not been sequenced yet. 2. Perform a phylogenetic analysis using new phylogenomic data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions determining the timing of speciation events and establishing corresponding geological events.

Impacts
What was accomplished under these goals? Accomplishments Major activities completed this year were to identify over 50,000 specimens of spider wasps from Central and South America, which were mounted and curated from the previous three summers. Additionally, we have continued to incorporate specimens sent to us from the American Entomological Institute and are preparing for the merger of this collection with our, which will occur in the summer of 2016. Additionally, I have investigated the role of taxon sampling in causing incongruent results between two recent phylogenomic studies of stinging wasps (Hymenoptera: Aculeata), a diverse lineage that includes ants, bees and the majority of eusocial insects. Using target enrichment of ultraconserved element (UCE) loci, the largest aculeate phylogenomic data set to date has been assembled, sampling 854 loci from 187 taxa, including 30 out of 31 aculeate families, and a diversity of parasitoid outgroups. We analyzed the complete matrix using multiple analytical approaches, and also performed a series of taxon inclusion/exclusion experiments, in which we analyzed taxon sets identical to and slightly modified from the previous phylogenomic studies. The results provide a highly supported phylogeny for virtually all aculeate lineages sampled, supporting ants as sister to Apoidea (bees+apoid wasps), bees as sister to Philanthinae+Pemphredoninae (lineages within a paraphyletic Crabronidae), Melittidae as sister to remaining bees, and paraphyly of cuckoo wasps (Chrysidoidea). Divergence dating analyses estimate ages for aculeate lineages in close concordance with the fossil record. Our analyses also demonstrate that outgroup choice and taxon evenness can fundamentally impact topology and clade support in phylogenomic inference. Concerning fossils, spider wasps had long been proposed to originate in the Late Cretaceous based on the Burmese amber fossil Bryopompilus interfector. A morphological examination of this fossil was performed and determined that it does not belong to Pompilidae or any other described hymenopteran family. Instead, it is placed into a 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).

Publications

• Type: Journal Articles Status: Published Year Published: 2016 Citation: Britton, J. M., Robert, D. J., Karen, M. E., Long, J. N. (2016). Herbivory and advance reproduction influence quaking aspen regeneration response to management in southern Utah, USA. Canadian Journal of Forest Research, 46, 674-682.
• Type: Journal Articles Status: Published Year Published: 2016 Citation: Windmuller-Campione, M. A., Long, J. N. (2016). Limber Pine (Pinus flexilis James), a Flexible Generalist of Forest Communities in the Intermountain West. PLoS ONE, 11(8), 1-16.
• Type: Conference Papers and Presentations Status: Other Year Published: 2016 Citation: Presentations Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "Systematics of the "well-known" family Chyphotidae (Hymenoptera).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2016 Citation: Presentations Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "The black-headed conundrum: Species boundaries in Chyphotes (Hymenoptera: Chyphotidae)," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)

Progress 10/01/14 to 09/30/15

Outputs
Target Audience:The target audiences for this research include evolutionary biologists, entomologists, taxonomists, biogeographers, geologists, and academics. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?Opportunities I have mentored 6 graduate students and 2 undergraduate students in the aspects of this project in the past year. How have the results been disseminated to communities of interest?Dissemination The results have been disseminated via five publications and four presentations at various scientific meetings held in the USA. What do you plan to do during the next reporting period to accomplish the goals?Plan of Work Specific objectives include: 1. Continue to extract DNA from further nocturnal species, begin gathering phylogenomic data (UCEs), which is new for this project, from all nocturnal species that have not been sequenced yet. 2. Perform a phylogenetic analysis using new phylogenomic data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions determining the timing of speciation events and establishing corresponding geological events.

Impacts
What was accomplished under these goals? Accomplishments Major activities completed this year were to identify over 100,000 specimens of nocturnal wasps from North and Central America mounted and curated from the previous two summers. Additionally, we incorporated specimens sent to us from the American Entomological Institute and are preparing for the merger of this collection with our, which will occur in the summer of 2016. Accomplishments from publications include a study of Sphaeropthalma Blake from the southwestern USA that revealed a highly autapomorphic undescribed species that I described as S. tapio Pitts & Sadler, sp. nov., and placed in a new species-group, the S. tapio species-group. I reconstruct a molecular phylogeny of Pompilidae from Bayesian and maximum-likelihood analyses of four nuclear molecular markers (elongation factor-1 α F2 copy, longwavelength rhodopsin, RNA polymerase II, and 28S ribosomal RNA), performed Bayesian divergence-time estimation analysis using four calibration points and ancestral-area reconstruction with a Bayesian binary Markov chain Monte Carlo method. New relationships for spider wasps were discovered, and new subfamilial delimitations are proposed and discussed based on the phylogeny. The origin of Pompilidae was c. 43.3 Mya, probably in the Nearctic region. Most of the extant subfamilies originated from the late Eocene to the Oligocene, and their current distributions are the product of various dispersal events that occurred over the course of ∼40 Mya. This is the first phylogenetic reconstruction of Pompilidae from molecular characters, with broad geographical and taxonomic sampling. I discovered that North American velvet ants form one of the world's largest known Müllerian mimicry Complexes. Lastly, I revised Priocnemella Banks, which is a small Neotropical genus within Ageniellini with remarkable morphological variation between species. Based on molecular phylogenetic analyses, Eragenia Banks was revalidated (stat. rev.) and applied to 16 species, eight of which are newly described: Eragenia bella Waichert & Pitts sp.n., E. carinata Waichert & Pitts sp.n., E. dentata Waichert & Pitts sp.n., E. oliva Waichert & Pitts sp.n., E. pseudomicans Waichert & Pitts sp.n., E. setosa Waichert & Pitts sp.n., E. rotunda Waichert & Pitts sp.n., E. villosa Waichert & Pitts sp.n., E. abdominalis (Smith), E. amabilis (Taschenberg), E. aureicornis (Smith), E. coerulipes (Smith), E. congrua (Fox), E. isolata (Banks), E. micans (Fabricius), and E. tabascoensis (Cameron).

Publications

• Type: Other Status: Published Year Published: 2015 Citation: Refereed Journal Articles J. R., Pitts, J., von Dohlen, C. D. (2015). Historical biogeography of the widespread spider wasp tribe Aporini (Hymenoptera: Pompilidae). Journal of Biogeography, 42(3), 495-506.
• Type: Other Status: Published Year Published: 2015 Citation: Refereed Journal Articles Waichert, C., Pitts, J., von Dohlen, C. D. (2015). Resurrection, revision and molecular phylogenetics of Eragenia Banks with implications for Ageniellini systematics (Hymenoptera: Pompilidae). Systematic Entomology, 40(2), 291321.
• Type: Conference Papers and Presentations Status: Other Year Published: 2015 Citation: Presentations Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "Systematics of the "well-known" family Chyphotidae (Hymenoptera).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2015 Citation: Presentations Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "The black-headed conundrum: Species boundaries in Chyphotes (Hymenoptera: Chyphotidae)," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)

Progress 10/01/13 to 09/30/14

Outputs
Target Audience: Target Audience Academics Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? Opportunities I have mentored 5 graduate students and 3 undergraduate students in the aspects of this project in the past year. How have the results been disseminated to communities of interest? Dissemination The results have been disseminated via eight publications and five presentations at various scientific meetings held in the USA. What do you plan to do during the next reporting period to accomplish the goals? Plan of Work Specific objectives include: 1. Continue to extract DNA from further nocturnal species, amplifying the mitochondrial gene (cytochrome c oxidase I [COI], nuclear genes (wingless [Wg]) and two nuclear intergenic regions (internal transcribed spacer regions 1&2 [ITS1&2]) from them and the two nuclear intergenic regions (internal transcribed spacer regions 1&2 [ITS1&2]) all the remaining species that have not been sequenced yet. 2. Extract and amplify COI for female specimens to associate the sexes using molecular data describing any previously unknown sexes or new species. We have an additional 18 females of Brachycistidinae for which this can be done. 3. Perform a phylogenetic analysis using molecular data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions determining the timing of speciation events and establishing corresponding geological events.

Impacts
What was accomplished under these goals? Accomplishments Major activities completed this year were to curate (mount, label and identify) over 50,000 specimens of nocturnal wasps from North and Central America collected this summer or left over from previous summers. Additionally, we incorporated the remaining collection of Hymenoptera from Los Angeles County Museum into the main USU insect collection. These new specimens will be integral to this study. Accomplishments from publications include Two new species of pepsine spider wasps, Calopompilus setaerotundus Waichert & Pitts, sp. nov. and Priocnemis (Um-bripennis) wasbaueri Waichert & Pitts, sp. nov., are described and illustrated. Eragenia Banks is revalidated (stat. rev.) and applied to 16 species, eight of which are newly described: Eragenia bella Waichert & Pitts sp.n., E. carinata Waichert & Pitts sp.n., E. dentata Waichert & Pitts sp.n., E. oliva Waichert & Pitts sp.n., E. pseudomicans Waichert & Pitts sp.n., E. setosa Waichert & Pitts sp.n., E. rotunda Waichert & Pitts sp.n.,and E. villosa Waichert & Pitts sp.n. Taxonomic changes were made to Drepanaporus based on the LWRh phylogenetic results and a new species of Drepanaporus, Drepanaporus bachata sp. nov, was newly described based on both molecular and morphological characters for both male and female specimens. Psorthaspis was determined to mimic diurnal mutillids and the species of this spiderwasp was added to currently defined Dasymutilla mimicry rings. Environmental niche mapping (ENM) and traditional morphological analyses was used to determine the validity of the subjective synonymy of Anthophora (Heliophila) curta Provancher with the senior synonym A. squammulosa Dours. New records of genera and species of spider wasps from Colombia were documented.

Publications

• Type: Journal Articles Status: Published Year Published: 2014 Citation: C. W., J. R., Pitts, J. (2014). *Waichert, C., *J. Rodriguez, and J. Pitts. 2014. New additions to the Honduran fauna of spider wasps (Hymenoptera: Pompilidae) with the description of two species. Zootaxa 3873: 590-600. Zootaxa, 3873, 590-600.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: C. W., Pitts, J., von Dohlen, C. D. (2014). *Waichert, C., C. von Dohlen, and J. Pitts. 2014. Resurrection, revision and molecular phylogenetics of Eragenia Banks with implications for Ageniellini systematics (Hymenoptera: Pompilidae). Systematic Entomology (2014) DOI: 10.1111/syen.12101
• Type: Journal Articles Status: Published Year Published: 2014 Citation: F. K., Pitts, J., N. E. (2014). Kurczewski, F.E., J.P. Pitts, N.B. Elliott. 2014. Annotated list of spider wasps from The Bahamas, with description of a new species of Tachypompilus (Hymenoptera: Pompilidae). Caribbean Naturalist 5: 1-28. Caribbean Naturalist, 5, 1-28.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: J. R., Pitts, J., von Dohlen, C. D. (2014). *Rodriguez, J., Pitts, J. P. and von Dohlen, C. D. 2014. Historical biogeography of the recently diverged widespread spider wasp tribe Aporini (Hymenoptera: Pompilidae). Journal of Biogeography. Journal of Biogeography.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: J. R., von Dohlen, C. D., Pitts, J. (2014). *Rodriguez, J., von Dohlen, C. D. & Pitts, J. P. 2014. 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. Annals of the Entomological Society of America 107: 709-720. UAES 8650. Annals of the Entomological Society of America, 107, 709-720.
• Type: Journal Articles Status: Published Year Published: 2014 Citation: J. R., Pitts, J., von Dohlen, C. D., Wilson, J. S. (2014). Rodriguez, J., von Dohlen, C. D., Pitts, J. P. and Wilson J. S. 2014. M�llerian mimicry as a result of codivergence between velvet ants and spider wasps. PLoS One. PLoS One
• Type: Journal Articles Status: Published Year Published: 2014 Citation: Orr, M., J. K., T. G., Pitts, J. (2014). *Orr, M.C., *J. Koch, T. Griswold and J. Pitts. 2014. Taxonomic utility of niche models in validationg species concepts: A case study on Anthophora (Heliophila) (Hymneoptera: Apidae). Zootaxa 3846: 411-429. Zootaxa, 3846, 411-429
• Type: Conference Papers and Presentations Status: Other Year Published: 2014 Citation: Pitts, J. (Author Only), von Dohlen, C. D. (Author Only), Rodriguez, J. (Presenter & Author), Entomological Society of America National Meeting,, "Is nesting behavior evolving in a stepwise evolution of increasing behavioral complexity in spider wasps? The case of the subfamily Pompilinae (Hymenoptera: Pompilidae)," Entomological Society of America, Entomological Society of America National Meeting, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2014 Citation: Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "Systematics of the "well-known" family Chyphotidae (Hymenoptera).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2014 Citation: Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting,, "The black-headed conundrum: Species boundaries in Chyphotes (Hymenoptera: Chyphotidae)," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Journal Articles Status: Published Year Published: 2014 Citation: V. C.-H., Pitts, J., J. R., c. w., F. F. (2014). *Castro-Huertas, V., J.P. Pitts, *J. Rodriguez, *C. Waichert, F. Fern�ndez. 2014. New Records of spider wasps (Hymenoptera, Pompilidae) from Colombia. ZooKeys 443: 35-44. ZooKeys, 449, 35-44.
• Type: Conference Papers and Presentations Status: Other Year Published: 2014 Citation: Rodriguez, J. (Presenter & Author), Pitts, J. (Author Only), Bond, J., von Dohlen, C. D. (Author Only), Entomological Society of America National Meeting,, "Molecular phylogeny of Pompilinae (Hymenoptera: Pompilidae): Evidence for rapid diversification and host shifts in spider wasps," Entomological Society of America, Portland, OR. (November 2014 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2014 Citation: Pitts, J. (Author Only), von Dohlen, C. D. (Author Only), Rodriguez, J. (Presenter & Author), Entomological Society of America National Meeting,, "The origin and diversification of spider wasps (Hymenoptera: Pompilidae).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)

Progress 01/01/13 to 09/30/13

Outputs
Target Audience: Academics Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? I have mentored 5 graduate students and 5 undergraduate students in the aspects of this project in the past year. How have the results been disseminated to communities of interest? The results have been disseminated via seven publications and four presentations at various scientific meetings held in the USA. What do you plan to do during the next reporting period to accomplish the goals? Specific objectives include: 1. Collect fresh specimens of nocturnal wasps to give baseline data on diversity, and abundance, and to obtain specimens for generating molecular, taxonomic and natural history data. 2. Extract DNA from samples, amplify and sequence the mitochondrial gene (cytochrome c oxidase I [COI], nuclear genes (wingless [Wg]) and two nuclear intergenic regions (internal transcribed spacer regions 1&2 [ITS1&2]) from all nocturnal species and outgroups. 3. Associate the sexes using molecular data describing any previously unknown sexes or new species. 4. Perform a phylogenetic analysis using molecular data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions determining the timing of speciation events and establishing corresponding geological events. 5. Continue Mark-Recapture and cage studies in southern Utah expanding it to include Chyphotidae and Brachycistidinae. 6. Determine hosts for nocturnal wasp species by locating would-be hosts' nests, collecting hosts and rearing host cocoons and pupae that may harbor nocturnal wasps. Now that a fresh batch of specimens have been identified, I will gather further molecular data, both from species not extracted thus far and from specimens from unexpected geographical areas.

Impacts
What was accomplished under these goals? Specific objectives include: 1. Collect fresh specimens of nocturnal wasps to give baseline data on diversity, and abundance, and to obtain specimens for generating molecular, taxonomic and natural history data. 2. Extract DNA from samples, amplify and sequence the mitochondrial gene (cytochrome c oxidase I [COI], nuclear genes (wingless [Wg]) and two nuclear intergenic regions (internal transcribed spacer regions 1&2 [ITS1&2]) from all nocturnal species and outgroups. 3. Associate the sexes using molecular data describing any previously unknown sexes or new species. 4. Perform a phylogenetic analysis using molecular data and use the recovered evolutionary relationships for subsequent biogeographical analysis of the desert regions determining the timing of speciation events and establishing corresponding geological events. 5. Continue Mark-Recapture and cage studies in southern Utah expanding it to include Chyphotidae and Brachycistidinae. 6. Determine hosts for nocturnal wasp species by locating would-be hosts' nests, collecting hosts and rearing host cocoons and pupae that may harbor nocturnal wasps. Major activities completed this year were to curate (mount, label and identify) over 100,000 specimens of nocturnal wasps from North and Central America collected this summer or left over from last summer. Additionally, the I acquired a large collection of Hymenoptera from Los Angeles County Museum in trade for our extra Diptera. This collection amounted to 740 drawers and a large proportion of nocturnal wasps. One third of this collection has been incorporated into the main USU insect collection. These new specimens will be integral to this study. Lastly, data from a third gene (wingless) has been sequenced from approximately 300 species of nocturnal wasps and a significant paper is in progress.

Publications

• Type: Conference Papers and Presentations Status: Other Year Published: 2013 Citation: Pitts, J. (Author Only), von Dohlen, C. D. (Author Only), Rodriguez, J. (Presenter & Author), Entomological Society of America National Meeting, "Is nesting behavior evolving in a stepwise evolution of increasing behavioral complexity in spider wasps? The case of the subfamily Pompilinae (Hymenoptera: Pompilidae)," Entomological Society of America, Entomological Society of America National Meeting, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2013 Citation: Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting, "Systematics of the "well-known" family Chyphotidae (Hymenoptera).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2013 Citation: Pitts, J. (Author Only), Sadler, E. (Presenter & Author), Entomological Society of America National Meeting, "The black-headed conundrum: Species boundaries in Chyphotes (Hymenoptera: Chyphotidae)," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)
• Type: Conference Papers and Presentations Status: Other Year Published: 2013 Citation: Pitts, J. (Author Only), von Dohlen, C. D. (Author Only), Rodriguez, J. (Presenter & Author), Entomological Society of America National Meeting, "The origin and diversification of spider wasps (Hymenoptera: Pompilidae).," Entomological Society of America, Knoxville, TN, 2012. (2012 - Present)

Progress 01/01/12 to 12/31/12

Outputs
OUTPUTS: This year's results (the first 6 months) were disseminated via one publication and four oral presentations. I have continued to mentor two undergraduates and three graduate students in desert biogeography and insect taxonomy, specifically instructing them in identification of nocturnal wasps. These individuals will be the only ones in the United States to have these skills. PARTICIPANTS: "3 graduate students: Cecilia Waichert, Juantia Rodriguez, and Emily Sadler; 2 undergraduates: Alex Kelley and Lauren Peless worked on the project. I am collaborating with the following individuals: Dr. Andy Austin (The University of Adelaide, South Australia); Dr. Neil Evenhuis (Bishop Museum of Natural History, Hawaii); Dr. Anthony Harris (Otago Museum, Dunedin, New Zealand); Dr. Jorge Horta (Tam. Mexico); Dr. Arkady Lelej (Vladivostok Russia); Dr. Donald G. Manley (Clemson, SC); Dr. Frank D. Parker (Retired); Dr. Eduardo F. dos Santos (Sao Paulo Brasil); Dr. Christian Schmid-Egger (Herrsching - Germany); Dr. Michael Sharkey (University of Kentucky, Lexington KY); Dr. Akira Shimizu (Tokyo, Japan); Dr. Carol D. von Dohlen (Dept. Biology, USU); Dr. Raymond Wahis (Gembloux, Belgium); and Dr. Marius Wasbauer (Retired)." TARGET AUDIENCES: The target audiences for this research include evolutionary biologists, entomologists, taxonomists, biogeographers, and geologists. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
This project will expand our knowledge of speciation in the deserts by using unrelated groups of wasps that differ from velvet ants, on which most of the previous evolutionary work has been completed. Experiments performed this summer indicate that these wasps are more abundant than previously thought, with one group having over 250,000 individuals per hectare during peak season. As all of these wasps are parasitic, some on bees, they greatly impact the ecology of the deserts they live in.

Publications

• Pitts, J., Rodriguez, J., & Restrepo-Giraldo, C., (2012). Temporal activity patterns of the spider wasp Pepsis montezuma Smith (Hymenoptera: Pompilidae) in a disturbed lower montane rainforest (Manizales, Colombia): Psyche, 2012: 1-4. (Published).