Source: UNIVERSITY OF CALIFORNIA, DAVIS submitted to NRP
SYSTEMATIC AND EVOLUTIONARY STUDIES ON ANTS (HYMENOPTERA: FORMICIDAE)
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
0085801
Grant No.
(N/A)
Cumulative Award Amt.
(N/A)
Proposal No.
(N/A)
Multistate No.
(N/A)
Project Start Date
Oct 1, 2011
Project End Date
Sep 30, 2016
Grant Year
(N/A)
Program Code
[(N/A)]- (N/A)
Recipient Organization
UNIVERSITY OF CALIFORNIA, DAVIS
410 MRAK HALL
DAVIS,CA 95616-8671
Performing Department
Entomology and Nematology
Non Technical Summary
Ants are the world's most successful social insects. They impose a strong ecological footprint in many communities in their varied roles as scavengers, predators, granivores, and herbivores. There is a word for "ant" in most languages, reflecting their ubiquity and distinctiveness to humans. The Chinese have employed ants for biological control for almost 1700 years, and there is continued interest in the use of ants as predators in agroecosystems. On the other hand, some ants especially introduced species have become serious economic pests, causing millions of dollars worth of damage per year in the United States. In California the exotic Argentine ant (Linepithema humile) is a serious urban, environmental and agricultural pest, and the red imported fire ant (Solenopsis invicta) poses similar threats. Improved knowledge of the taxonomy, phylogeny and distribution of ants will help us to deal more effectively with the challenges posed by pest species, and will contribute to a better understanding of the roles that ants play in natural ecosystems. Worldwide there are more than 13,000 described species of ants, but there are numerous additional species awaiting discovery and description. Moreover the evolutionary relationships among species are still being clarified. Application of molecular methods (analysis of DNA sequences) is revolutionizing our understanding of the "tree of life" (phylogeny) of ants. This project aims to improve the species-level taxonomy and phylogeny of ants, using a combination of morphological and molecular characters. New information about the identification and distribution of ants is published in conventional journals and online (www.antweb.org). A global approach to problems in ant systematics is especially important in California where the most significant pest ants are non-native species originating in distant parts of the world.
Animal Health Component
(N/A)
Research Effort Categories
Basic
(N/A)
Applied
(N/A)
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
21131101130100%
Goals / Objectives
Specific objectives of this project are (1) publication of taxonomic monographs and identification keys that permit reliable identification of ant species; (2) analysis of phylogenetic relationships among ants and evaluation of the factors that have been important in their evolutionary history; and (3) provision of accurate and detailed online information about ant diversity, especially with regard to the California ant fauna. Additional goals include (4) the rapid identification of ant specimens submitted for determination, and (5) the training of students in ant systematics.
Project Methods
Species-Level Taxonomy Improvement of taxonomic knowledge of ants requires access to large, geographically extensive samples of specimens from throughout the world. This material is either collected in the field by the P.I. or borrowed from museum collections. The ant specimens are then carefully scrutinized for morphological variation. Measurements are taken, qualitative characters are assessed, and a search is made for phenotypic discontinuities that would indicate the existence of more than one species. Particular attention is paid to male genitalia because differences in such features usually reflect reproductive isolation. Molecular (DNA sequence) data can be helpful in the delimitation of species boundaries and in the testing of existing hypotheses about species limits. Several nuclear genes are used in the P.I.'s lab for this purpose. We also collaborate with the Biodiversity Institute of Ontario (University of Guelph) in a project to barcode North American ants using a 658 bp fragment of the mitochondrial gene COI. Phylogenetic Relationships The field of phylogenetics is in a state of renaissance, as new analytical methods are developed which improve the power of phylogenetic inference. In studies on ants two principal classes of evidence are used: morphology and DNA sequences. Terminal taxa are chosen usually species or exemplar taxa and data is gathered on morphological and molecular variation. This is codified in a taxon-by-character matrix, and then subject to phylogenetic analysis using model based approaches such as maximum likelihood (ML) and Bayesian inference. The strength of support for different components of the tree is evaluated with the bootstrap and with Bayesian posterior probabilities. The resulting ant phylogenies can be applied to other problems such as delimitation of species boundaries, comparative analysis of trait evolution, estimation of dates of origin of higher taxa, and testing of biogeographic hypotheses. At a practical level such phylogenies offer predictive power about the characteristics of ants. For example, we could ask: Where in the ant "family tree" do invasive species emerge Are they confined to particular clades What are the traits shared by them Ant Identification Each year the P.I. receives thousands of ant specimens for identification from personnel at various state, federal, non-governmental, and international agencies. These identification requests are processed as promptly as possible. This is made possible by having access to the relevant literature and to a comprehensive collection of ants. An essential component of this research project is the ongoing maintenance of this large reference collection. The collection is global in scope but particular emphasis is placed on coverage of the ants of California and adjacent regions. In order to disseminate information to the public about ant identification and ant diversity a web site has been established which provides Automontage images of all of the ant species of California (http://www.antweb.org/california.jsp). This web site is frequently updated and modified, and has the potential for future integration of additional information.

Progress 10/01/11 to 09/30/16

Outputs
Target Audience:Target audiences include (1) other scientists working on the systematics, ecology and evolution of Hymenoptera; (2) personnel in state, federal and other institutions requiring assistance with ant identification, including pest species; and (3) members of the general public interested in, or requiring information about, ants and ant control. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?UC Davis Entomology graduate students Bonnie Blaimer, Michael Branstetter, Marek Borowiec, Matthew Prebus and Brendon Boudinot received research experience in ant taxonomy, molecular phylogenetics and bioinformatics. Every year the PI participated in Ant Course (http://research.calacademy.org/ent/courses/ant), providing training in ant systematics and biology to hundreds of students from around the world. How have the results been disseminated to communities of interest?Publication of peer-reviewed articles. Submission of collection data to AntWeb (www.antweb.org). What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? Over the duration of this five-year project we completed comprehensive studies on the phylogeny and taxonomy of ants. Underpinned by morphological and molecular analyses (now based on next-generation sequencing technology) and by field work carried out on all major continents, we published contributions on a diverse array of taxa, including army ants, dracula ants, myrmicine ants, and formicine ants. We also published an analysis of the co-evolution of ants and plants; a review paper on ant phylogeny and evolution; and a seminal phylogenomic paper on the relationship of ants to other stinging Hymenoptera. Many ant specimens were databased and the collection information was uploaded to AntWeb (www.antweb.org). Contributions were made to a new community-based online catalogue of the world's ant species, AntCat (www.antcat.org).

Publications

  • Type: Journal Articles Status: Published Year Published: 2016 Citation: Boudinot, B. E.; Probst, R. S.; Brand�o, C. R. F.; Feitosa, R. M.; Ward, P. S. 2016. Out of the Neotropics: newly discovered relictual species sheds light on the biogeographic history of spider ants (Leptomyrmex, Dolichoderinae, Formicidae). Systematic Entomology 41:658-671. doi:10.1111/syen.12181
  • Type: Journal Articles Status: Published Year Published: 2016 Citation: Ward, P. S.; Blaimer, B. B.; Fisher, B. L. 2016. A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinomyrmex. Zootaxa 4072:343-357. doi:10.11646/zootaxa.4072.3.4
  • Type: Journal Articles Status: Published Year Published: 2016 Citation: Ward, P. S.; Fisher, B. L. 2016. Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae). Systematic Entomology 41:683-693. doi:10.1111/syen.12186
  • Type: Journal Articles Status: Published Year Published: 2016 Citation: Ward, P. S.; Brady, S. G.; Fisher, B. L.; Schultz, T. R. 2016. Phylogenetic classifications are informative, stable and pragmatic: the case for monophyletic taxa. Insectes Sociaux 63:489-492 doi: 10.1007/s00040-016-0516-9


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

Outputs
Target Audience: Entomologists and members of the general public. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? UC Davis Entomology graduate students Marek Borowiec, Michael Branstetter, Matthew Prebus and Brendon Boudinot received research experience in ant taxonomy, molecular phylogenetics and bioinformatics. In August 2014 the PI participated as a lecturer in Ant Course (http://research.calacademy.org/ent/courses/ant), interacting with a diverse array of students and researchers. Research was supported by NSF grant DEB-1354996 (Ant Diversity of the MesoAmerican Corridor (ADMAC)), which includes collaborators at the University of Utah. How have the results been disseminated to communities of interest? Publication of peer-reviewed articles. Submission of collection data to AntWeb (www.antweb.org). What do you plan to do during the next reporting period to accomplish the goals? 1. Continue systematic and phylogenetic studies of ants, including the generation of new information on the distribution and identity of California ants (http://www.antweb.org/california.jsp). 2. Publish research findings in peer-reviewed journals. 3. Contribute additional data to web sites such as AntWeb and AntCat. 4. Continue NSF-supported collaborative work on ant diversity of the MesoAmerican corridor (ADMAC), including incorporation of new phylogenomic data.

Impacts
What was accomplished under these goals? Research continued on the systematics and evolution of ants (Hymenoptera: Formicidae). This involved a combination of field work in North America and southeast Asia, specimen curation and identification, and molecular phylogenetics. Papers were published on the relationship of ants to other stinging (aculeate) Hymenoptera; the evolution of army ants (dorylines); and the phylogeny and evolution of ants as a whole. Many ant specimens were databased, and the collection information was uploaded to AntWeb (www.antweb.org). Contributions were made to a new community-based online catalogue of the world's ant species, AntCat (www.antcat.org).

Publications

  • Type: Journal Articles Status: Published Year Published: 2014 Citation: Brady, S. G.; Fisher, B. L.; Schultz, T. R.; Ward, P. S. 2014. The rise of army ants and their relatives: diversification of specialized predatory doryline ants. BMC Evolutionary Biology 14:93 doi:10.1186/1471-2148-14-93.
  • Type: Journal Articles Status: Published Year Published: 2014 Citation: Ward, P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology Evolution and Systematics 45:23-43.
  • Type: Journal Articles Status: Published Year Published: 2013 Citation: Johnson, B. R.; Borowiec, M. L.; Chiu, J. C.; Lee, E. K.; Atallah, J.; Ward, P. S. 2013. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. Current Biology 23:2058-2062.


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

Outputs
Target Audience: Entomologists and members of the general public. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? UC Davis Entomology graduate students Marek Borowiec, Michael Branstetter, and Matthew Prebus received research experience in ant taxonomy, molecular phylogenetics and bioinformatics. In August 2013 the PI participated as a lecturer in Ant Course (http://research.calacademy.org/ent/courses/ant), interacting with a diverse array of students and researchers. Research was supported by NSF grant DEB-0842204 (Ants of the Southwest Indian Ocean and East Africa), which involves collaborators at the California Academy of Sciences, the University of Mississippi and the University of Guelph. How have the results been disseminated to communities of interest? Nothing Reported What do you plan to do during the next reporting period to accomplish the goals? 1. Continue systematic and phylogenetic studies of ants, including the generation of new information on the distribution and identity of California ants (http://www.antweb.org/california.jsp). 2. Publish research findings in peer-reviewed journals. 3. Contribute additional data to web sites such as AntWeb and AntCat. 4. In collaboration with colleagues at the University of Utah, initiate a new project (ADMAC) on ant diversity in the Meso-American corridor.

Impacts
What was accomplished under these goals? Research continued on the systematics and evolution of ants (Hymenoptera: Formicidae). This involved a combination of field work in North and South America, specimen curation and identification, and molecular phylogenetics. The relationships of ants to other stinging (aculeate) Hymenopteras was investigated using newly generated genomic data. Many ant specimens were databased, and the collection information was uploaded to AntWeb (www.antweb.org). Contributions were made to a new community-based online catalogue of the world's ant species, AntCat (www.antcat.org).

Publications


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

    Outputs
    OUTPUTS: Research continued on the taxonomy, phylogeny and biogeography of ants (Hymenoptera: Formicidae). This involved a combination of field work, specimen curation and identification, molecular phylogenetic analysis, and publication of findings. This year (2012) field studies were carried out in California, Nevada, Utah, Canada, and Uganda, resulting in the acquisition of 229 accessions and approximately 3500 ant specimens. These collections are being processed and databased, with information uploaded to AntWeb (www.antweb.org). In addition, identification services were provided to individuals at various institutions including UC Extension, the Essig Museum, University of Utah, and the University of Reading. Results of research were published in two scientific articles. PARTICIPANTS: UC Davis Entomology graduate students Marek Borowiec, Michael Branstetter, Bonnie Blaimer and Matthew Prebus assisted with the molecular phylogenetics research. In August 2012 the PI participated as a lecturer in Ant Course (http://research.calacademy.org/ent/courses/ant), interacting with a diverse array of students and researchers. Research was supported by NSF grant DEB-0842204 (Ants of the Southwest Indian Ocean and East Africa), which involves collaborators at the California Academy of Sciences, the University of Mississippi and the University of Guelph. The PI was also involved in a collaborative project on the leaf litter arthropods of Mesoamerica (LLAMA), working with colleagues at the University of Utah and the Canadian Museum of Nature (Ottawa). TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

    Impacts
    The content of AntWeb (www.antweb.org) is being continually updated, including new information on the distribution and identity of California ants (http://www.antweb.org/california.jsp). As the designated "California Ant Curator" on this web site the PI receives and fields numerous inquiries from the general public about ants.

    Publications

    • Ward, P. S.; Sumnicht, T. P. 2012. Molecular and morphological evidence for three sympatric species of Leptanilla (Hymenoptera: Formicidae) on the Greek island of Rhodes. Myrmecological News 17:5-11.
    • Seltmann, K. C.; Yoder, M. J.; Miko, I.; Forshage, M.; Bertone, M. A.; Agosti, D.; Austin, A. D.; Balhoff, J. P.; Borowiec, M. L.; Brady, S. G.; Broad, G. R.; Brothers, D. J.; Burks, R. A.; Buffington, M. L.; Campbell, H. M.; Dew, K. J.; Ernst, A. F.; Fernandez-Triana, J. L.; Gates, M. W.; Gibson, G. A. P.; Jennings, J. T.; Johnson, N. F.; Karlsson, D.; Kawada, R.; Krogmann, L.; Kula, R. R.; Mullins, P. L.; Ohl, M.; Rasmussen, C.; Ronquist, F.; Schulmeister, S.; Sharkey, M. J.; Talamas, E.; Tucker, E.; Vilhelmsen, L.; Ward, P. S.; Wharton, R. A.; Deans, A. R. 2012. A hymenopterists' guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research 27:67-88.


    Progress 01/01/11 to 12/31/11

    Outputs
    OUTPUTS: Research continued on various aspects of the taxonomy, phylogeny and biogeography of ants (Hymenoptera: Formicidae). This involved a combination of field work, specimen curation and identification, molecular phylogenetic analysis, and publication of findings. This year (2011) field studies were carried out in California, Arizona, Washington, Canada, and Hong Kong, resulting in the acquisition of 182 accessions and approximately 2100 ant specimens. A subset of these specimens was DNA barcoded using a 658 bp fragment of the mitochondrial gene COI. Data from these collections is in the process of being uploaded to AntWeb (www.antweb.org). In addition identification services were provided to individuals at various institutions including University of Utah, Evergreen College (Olympia, Washington), Smithsonian Institution, and Universidade Federal do Rio Grande do Norte (Brazil). Results of research were published in several scientific articles. PARTICIPANTS: UC Davis Entomology graduate students Marek Borowiec, Michael Branstetter, Bonnie Blaimer and Matthew Prebus assisted with the molecular phylogenetics research. Undergraduate students also worked in the lab, receiving training in sorting and identification of ants and other insects from bulk field samples. Research was supported by NSF grant EF-0431330 (Ant Tree of Life Project), carried out in collaboration with colleagues at the California Academy of Sciences (San Francisco) and the Smithsonian Institution (Washington, DC); and by a second NSF-funded project, ASWEA (Ants of the Southwest Indian Ocean and East Africa), which involves collaborators at the California Academy of Sciences, the University of Mississippi and the University of Guelph. The PI was also involved in a collaborative project on the leaf litter arthropods of Mesoamerica (LLAMA), working with colleagues at Evergreen College (Olympia, Washington) and the Canadian Museum of Nature (Ottawa). TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

    Impacts
    The content of AntWeb (www.antweb.org) is being continually updated, including new information on the distribution and identity of California ants (http://www.antweb.org/california.jsp). As the designated "California Ant Curator" on this web site the PI receives and answers numerous inquiries from the general public about ants.

    Publications

    • Roura-Pascual, N.; Hui, C.; Ikeda, T.; Leday, G.; Richardson, D. M.; Carpintero, S.; Espadaler, X.; Gomez, C.; Guenard, B.; Hartley, S.; Krushelnycky, P.; Lester, P. J.; McGeoch, M. A.; Menke, S. B.; Pedersen, J. S.; Pitt, J. P. W.; Reyes, J.; Sanders, N. J.; Suarez, A. V.; Touyama, Y.; Ward, D.; Ward, P. S.; Worner, S. P. 2011. The relative roles of climatic suitability and anthropogenic influence in determining the pattern of spread in a global invader. Proceedings of the National Academy of Sciences U. S. A. 108: 220-225.
    • Ward, P. S. 2011. Integrating molecular phylogenetic results into ant taxonomy. Myrmecological News 15: 21-29.


    Progress 01/01/10 to 12/31/10

    Outputs
    OUTPUTS: This is an ongoing project on the taxonomy and phylogeny of ants (Hymenoptera: Formicidae), which entails a combination of field work, specimen curation and identification, molecular phylogenetic analysis, and publication of findings. This year (2010) field studies were carried out in California, Nevada, North Carolina, Honduras and Borneo, resulting in the acquisition of 265 accessions and approximately 3000 ant specimens. A subset of these specimens was DNA barcoded using a 658 bp fragment of the mitochondrial gene COI. Data from these collections is in the process of being uploaded to AntWeb (www.antweb.org). In addition identification services were provided to individuals at various institutions including University of California Cooperative Extension, Florida Atlantic University, Field Museum (Chicago), Evergreen College (Olympia, Washington), Smithsonian Institution, Wake Forest University (North Carolina), CSIRO Tropical Ecosystem Centre (Darwin) and CNRS, Montpellier, France. Publications appearing this year covered research on ant taxonomy, phylogeny and biogeography, including a review paper on this topic published in the book "Ant Ecology" (Oxford University Press). PARTICIPANTS: UC Davis Entomology graduate students Marek Borowiec, Michael Branstetter and Bonnie Blaimer assisted with the molecular phylogenetics research. Undergraduate students also worked in the lab, receiving training in sorting and identification of ants and other insects from bulk field samples. NSF-supported research on the Ant Tree of Life is carried out in collaboration with colleagues at the California Academy of Sciences (San Francisco) and the Smithsonian Institution (Washington, DC). The PI is also involved in a collaborative project on the leaf litter arthropods of Mesoamerica (LLAMA), working with colleagues at Evergreen College (Olympia, Washington) and the Canadian Museum of Nature (Ottawa). Another NSF-funded project, ASWEA (Ants of the Southwest Indian Ocean and East Africa) involves collaborators at the California Academy of Sciences, the University of Mississippi and the University of Guelph. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

    Impacts
    The content of AntWeb (www.antweb.org) is being continually updated, including new information on the distribution and identity of California ants (http://www.antweb.org/california.jsp). As the designated "California Ant Curator" on this web site the PI receives and answers numerous inquiries from the general public about ants.

    Publications

    • Ward, P. S. 2009. Taxonomy, phylogenetics and evolution. Pp. 3-17 in: Lach, L., Parr, C. L. & Abbott, K. (eds). Ant ecology. Oxford: Oxford University Press, xvii + 410 pp.
    • Ward, P. S.; Brady, S. G.; Fisher, B. L.; Schultz, T. R. 2010. Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology 59: 342-362.
    • Lucky, A.; Ward, P. S. 2010. Taxonomic revision of the ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Zootaxa 2688: 1-67.


    Progress 01/01/09 to 12/31/09

    Outputs
    OUTPUTS: Studies continued on the taxonomy and phylogeny of ants (Hymenoptera: Formicidae). Field work was carried out in California, Arizona, Florida, Maine and eastern Canada, resulting in the collection of 75 accessions and approximately 1000 ant specimens. A subset of these specimens was DNA barcoded using a 658 bp fragment of the mitochondrial gene COI. Molecular phylogenetic research on ants continued at a global scale, under the auspices of the NSF-supported Ant AToL (Assembling the Tree of Life) Project, of which Ward is the lead PI. A study on the phylogeny and biogeography of dolichoderine ants was completed and accepted for publication. Publications appearing this year covered research on ant/plant mutualisms, the relationship between carpenter ants (Camponotus spp.) and their microbial symbionts, and the taxonomy of selected ant groups. Species identifications of ants were provided to personnel at various institutions, including the University of California Cooperative Extension, UC White Mountain Research Station, California Academy of Sciences, University of Nevada at Reno, Arizona State University, Florida Atlantic University, Smithsonian Institution, Wake Forest University (North Carolina), Universidad Autonoma Agraria Antonio Narro (Mexico), and CNRS, Montpellier, France. PARTICIPANTS: UC Davis Entomology graduate students Andrea Lucky, Michael Branstetter and Bonnie Blaimer assisted with the molecular phylogenetics research. Undergraduate students also worked in the lab, receiving training in sorting and identification of ants and other insects from bulk field samples. NSF-supported research on the Ant Tree of Life is carried out in collaboration with colleagues at the California Academy of Sciences (San Francisco) and the Smithsonian Institution (Washington, DC). The PI is also involved in a collaborative project on the leaf litter arthropods of Mesoamerica (LLAMA), working with colleagues at Evergreen College (Olympia, Washington) and the Canadian Museum of Nature (Ottawa). Another NSF-funded project, ASWEA (Ants of the Southwest Indian Ocean and East Africa) involves collaborators at the California Academy of Sciences and the University of Guelph. TARGET AUDIENCES: The target audience for the Ants of California web site (http://www.antweb.org/california.jsp) is the public of California. This site is designed to assist members of the public in identifying ants found in California. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

    Impacts
    The content of AntWeb (www.antweb.org) is being continually updated, including new information on the distribution and identify of California ants (http://www.antweb.org/california.jsp). As the designated "California Ant Curator" on this web site the PI receives and answers numerous inquiries from the general public about ants.

    Publications

    • Kautz, S.; Lumbsch, H. T.; Ward, P. S.; Heil, M. 2009. How to prevent cheating: a digestive specialization ties mutualistic ants to their ant-plant partners. Evolution 63:839-853.
    • Ward, P. S.; Brady, S. G. 2009. Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa 2063:46-54.
    • Ward, P. S. 2009. The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae). Journal of Hymenoptera Research 18:285-304.
    • Wernegreen, J. J.; Kauppinen, S. N.; Brady, S. G.; Ward, P. S. 2009. One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology 9:292 doi:10.1186/1471-2148-9-292.


    Progress 01/01/08 to 12/31/08

    Outputs
    OUTPUTS: Ongoing research continued to generate new knowledge about the species-level taxonomy and phylogeny of ants (Hymenoptera: Formicidae). Field samples of ants (215 accessions, approximately 1500 specimens) were collected by the PI in California, Nevada, southeastern United States, Canada, Honduras, and Venezuela. These collections are being processed and incorporated into the UC Davis collection (Bohart Museum of Entomology). The ant specimens are being used for both alpha-taxonomy and for phylogenetic studies under the rubric of the NSF-supported Ant AToL (Assembling the Tree of Life) project, of which Ward is the lead PI. Many new ant taxa were sequenced for a series of protein-encoding nuclear genes (including abdominal-A, long wavelengths rhodopsin, elongation factor 1-alpha F2, arginine kinase, enolase, and CAD), and these data promise to enhance our understanding of the evolutionary relationships of these organisms. Ants collected at Algodones Dunes in Imperial County were identified, as part of a larger survey of the insect fauna of this site by the Bohart Museum. Species identifications of ants were provided to personnel at various institutions, including the University of California Cooperative Extension, California Department of Food & Agriculture, USGS San Diego Field Station, University of California at San Diego, California Academy of Sciences, and Florida State Collection of Arthropods. PARTICIPANTS: UC Davis Entomology graduate students Andrea Lucky, Michael Branstetter and Bonnie Blaimer assisted with the molecular phylogenetics research. NSF-supported research on the Ant Tree of Life is carried out in collaboration with colleagues at the California Academy of Sciences (San Francisco) and the Smithsonian Institution (Washington, DC). The PI is also involved in a collaborative project on the leaf litter arthropods of Mesoamerica (LLAMA), working with colleagues at Evergreen College (Olympia, Washington) and the Canadian Museum of Nature (Ottawa). TARGET AUDIENCES: The target audience for the Ants of California web site (http://www.antweb.org/california.jsp) is the public of California. This site is designed to assist members of the public in identifying ants found in California. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

    Impacts
    New information about the identity and distribution of ants was added to AntWeb (www.antweb.org), including pages devoted specifically to the ants of California (http://www.antweb.org/california.jsp). The AntWeb site is a popular one, and the PI received and responded to many inquiries from the California public about ants or ant-like insects that they have encountered. Feedback from the public indicates that most of these inquires are sent to the PI because he is listed as a "California Ant Curator" (i.e., expert) on the web page.

    Publications

    • Ward, P. S. 2008. [Review of: Fisher, B. L. & Cover, S. P. 2007. Ants of North America: a guide to genera. Berkeley: University of California Press, xiv + 194 pp.] Myrmecological News 11:135-136.


    Progress 01/01/07 to 12/31/07

    Outputs
    Taxonomic and phylogenetic studies were carried out on selected groups of ants (Hymenoptera: Formicidae), with particular emphasis on the subfamilies Dolichoderinae and Pseudomyrmecinae. This involved a combination of field work in California, Arizona and southeastern United States, and laboratory-based analyses of morphology and DNA sequence variation. The Ward lab continued to be the lead group for the NSF-sponsored Ant AToL (Assembling the Tree of Life) Project. Species identifications of ants were provided to personnel at various institutions, including the University of California Cooperative Extension, California Department of Food & Agriculture, California Department of Health Services, Santa Barbara Museum of Natural History, California Academy of Sciences, and Stanford University. Publications included a general review of progress in ant systematics (Ward, 2007c), an appraisal of the contribution of Edward O. Wilson to the field (Ward, 2007a), and a specialized study of phylogenetic relationships among ants of the genus Leptanilloides and other taxa (Ward, 2007b). A web site providing color images of all the ant species in California (http://www.antweb.org/california.jsp) was updated and improved.

    Impacts
    Research on ant systematics provides the foundational knowledge that makes possible a better understanding of ant biology. The "Ants of California" web site has generated strong interest from the public, in part because of the high quality color images which facilitate identification of ants. Accurate identification of ants is crucial for effective monitoring of potential invasive species. A better knowledge of ant phylogeny will allow more effective prediction about the traits of related species, and how these might be exploited for control purposes.

    Publications

    • Ward, P. S. 2007a. Edward O. Wilson and his contributions to ant systematics. Memoirs of the American Entomological Institute 80: 3-7.
    • Ward, P. S. 2007b. The ant genus Leptanilloides: discovery of the male and evaluation of phylogenetic relationships based on DNA sequence data. Memoirs of the American Entomological Institute 80: 637-649.
    • Ward, P. S. 2007c. Phylogeny, classification and species-level; taxonomy of ants. Zootaxa 1668: 549-563.


    Progress 01/01/06 to 12/31/06

    Outputs
    Long-term systematic studies on ants were continued, with emphases on (1) the ant fauna of southwestern United States, (2) the ant subfamily Pseudomyrmecinae, and (3) the phylogeny of ants. Field work was carried out in California, Nevada, Arizona, Fiji and Australia. This allowed further modification and improvement of a web site covering the ants of California (http://www.antweb.org/california.jsp) and other regions of the world. Species identifications of ants were provided to colleagues at the California Academy of Sciences; Chaffey College, California; Stanford University, California; USDA Forest Service (Region 5); US National Park Service (Yosemite National Park); Smithsonian Institution; Florida Atlantic University; Boston University; McGill University, Canada; Royal Zoological Society of Antwerp, Belgium; Universitat Duisburg-Essen, Germany; and Universidad Nacional de Colombia, Bogota. Publications included a general review of ant biology for the layperson (Ward, 2006a) and a taxonomic revision of the ant genus Tetraponera in the Afrotropical region (Ward, 2006b). In addition, a landmark paper on ant phylogeny was published (Brady et al., 2006). Based on the most comprehensive molecular phylogenetic data set ever assembled for ants, this study provides a well-calibrated timeline for the evolution of ants and evaluates areas of ant phylogeny that remain uncertain.

    Impacts
    Research on ant systematics provides the foundational knowledge that makes possible a better understanding of ant biology. The 'Ants of California' web site has generated strong interest from the public, in part because of the high quality color images which facilitate identification of ants. Accurate identification is crucial for effective monitoring of potential invasive species. A better knowledge of ant phylogeny will allow more effective prediction about the traits of related species, and how these might be exploited for control purposes.

    Publications

    • Brady, S. G.; Fisher, B. L.; Schultz, T. R.; Ward, P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences U. S. A. 103: 18172-18177.
    • Ward, P. S. 2006a. Ants. Current Biology 16: 152-155.
    • Ward, P. S. 2006b. The ant genus Tetraponera in the Afrotropical region: synopsis of species groups and revision of the T. ambigua-group (Hymenoptera: Formicidae). Myrmecologische Nachrichten 8: 119-130.


    Progress 01/01/05 to 12/31/05

    Outputs
    A comprehensive review of the ants of California was completed and published. This publication includes a key to the ant genera of California and a standardized list of the species occurring in the state; and it complements a web site ((http://www.antweb.org/california.jsp) devoted to the provision of high quality images of California ants. Other research carried out during the year 2005 included intensive molecular phylogenetic analyses of selected ant taxa, aimed at resolving the relationships among the major lineages of ants (supported by an NSF 'Assembling the Tree of Life' grant); morphological studies of army ants; analyses of ant species intercepted in quarantine and the properties that make them good invaders; and specimen-based taxonomic research on the ant subfamily Pseudomyrmecinae. Species identifications of ants were provided to colleagues at UC Cooperative Extension, California Department of Food and Agriculture, San Jose State University, University of Illinois Urbana-Champaign, Harvard University, Stanford University, Florida Atlantic University and Universidad Nacional de Colombia.

    Impacts
    As a result of this research the ants of California can be more readily identified and we now have improved understanding of what makes certain ant species successful invaders. The 'Ants of California' web site continues to attract strong interest from members of the public and it has facilitated K-12 school programs in ant monitoring and identification. Basic research on the taxonomy and phylogeny of ants underpins these more applied aspects of ant knowledge.

    Publications

    • Ward, P. S. 2005. A synoptic review of the ants of California (Hymenoptera: Formicidae). Zootaxa 936:1-68.
    • Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R. 2005. Assembling the ant "Tree of Life" (Hymenoptera: Formicidae). Myrmecologische Nachrichten 7:87-90.
    • Brady, S. B., Ward, P. S. 2005. Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae). Systematic Entomology 30:593-618.
    • Suarez, A. V., Holway, D. A.; Ward, P. S. 2005. The role of opportunity in the unintentional introduction of nonnative ants. Proceedings of the National Academy of Sciences U. S. A. 102:17032-17035.


    Progress 01/01/04 to 12/31/04

    Outputs
    Taxonomic studies were carried out on selected groups of ants, including the subfamily Pseudomyrmecinae and California species in the genera Camponotus, Forelius, Leptothorax, Monomorium and Temnothorax. This work involved a combination of field studies in California, Nevada, Arizona, and Australia; molecular phylogenetic analyses; and specimen-based museum research. Species identifications of ants were provided to colleagues at UC Cooperative Extension, California Department of Food and Agriculture, USGS Western Ecological Research Center, Orma Smith Museum of Natural History (Caldwell, ID) and various universities including Cornell University, Harvard University, Arizona State University, Utah State University, Humboldt State University and University of Nevada. In collaboration with Brian Fisher at the California Academy of Sciences, improvements were made to the AntWeb site, which provides (among other services) information and images of the ants of California (http://www.antweb.org/california.jsp). A large ($1.5 million) Tree of Life grant was awarded to the PI from the National Science Foundation for support of ant systematics.

    Impacts
    As a result of this research our knowledge of the ant fauna of California continues to improve. The AntWeb site has attracted strong interest from members of the California public who want to learn more about ant diversity and biology. Basic research on the species-level taxonomy and phylogeny of ants enhances our ability to identify species, both native and introduced, and to develop appropriate strategies for controlling pest ants.

    Publications

    • Ward, P. S., Downie, D. A. 2004. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology [published online December 2004; DOI: 10.1111/j.1365-3113.2004.00281.x]


    Progress 01/01/03 to 12/31/03

    Outputs
    Long-term research continued on the systematics and evolutionary biology of ants. Field work was carried out in California, Arizona, Nevada, Utah, Colorado and Central America, to collect ant specimens for taxonomic and biogeographic studies. Species identifications were provided to investigators at the National Park Service, the US Geological Survey, and at various universities including UC Davis, UC Berkeley, California State University at Chico, College of Marin, Chaffey College, Arizona State University, and University of Nevada. In collaboration with a colleague at the California Academy of Sciences, a web site was set up depicting high quality color images of all the species of ants in California (http://www.antweb.org/california.jsp). Systematic papers were published on ants of the subfamilies Pseudomyrmecinae and Myrmeciinae.

    Impacts
    Field work has led to improved understanding of the distribution and identity of ants from California and adjacent regions. The antweb site has been widely praised as a useful online tool for conservation biologists who wish to use ants as indicator taxa. Basic research on ant systematics enhances our ability to identify species, and to apply intelligent control measures against pest ants.

    Publications

    • Ward, P. S., Brady, S. G. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17:361-386.
    • Ward, P. S. 2003. Subfamilia Pseudomyrmecinae. Pp. 331-333 in Fernandez, F. (ed.) Introduccion a las hormigas de la region Neotropical. Bogota, Colombia: Instituto de Investigacion de Recursos Biologicos Alexander von Humboldt, xxvi + 398 pp.


    Progress 01/01/02 to 12/31/02

    Outputs
    Research was carried out on various aspects of ant systematics. Field work was conducted in California (17 counties), Nevada and Arizona, resulting in substantial additions to the ant collection of the Bohart Museum of Entomology. The P.I. visited the Museum of Comparative Zoology, Harvard University to examine type specimens in that collection. This helped to clarify the identity of several California ant species. In addition, substantial numbers of ant specimens were identified for personnel at various state, federal and international institutions, including UC Cooperative Extension, the National Park Service, the US Geological Survey, and the Florida State Collection of Arthropods. Systematic research focused on the ant genera Pseudomyrmex, Leptothorax and Formica. Papers were published on the taxonomy and biogeography of the ants of Baja California (Johnson & Ward, 2002; Boulton & Ward, 2002; Ward & Boulton, 2002).

    Impacts
    Basic research on the taxonomy, biology and distribution of ants enhances our ability to identify species, and to apply intelligent control measures against pest ants. Moreover, ants are seen as increasingly useful in bio-monitoring studies and taxonomic knowledge is vital in this regard.

    Publications

    • Gronlund, C. J., Deangelis, M. D., Pruett-Jones, S., Ward, P. S., Coyne, J. A. 2002. Mate grasping in Drosophila pegasa. Behaviour 139:545-572.
    • Johnson, R. A., Ward, P. S. 2002. Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview. Journal of Biogeography 29:1009-1026.
    • Boulton, A. M., Ward, P. S. 2002. Ants. Pp. 112-128 in Case, T. J., Cody, M. L. & Ezcurra, E. (eds.) A new island biogeography of the Sea of Cortes. Oxford: Oxford University Press, xvii + 669 pp.
    • Ward, P. S., Boulton, A. M. 2002. Checklist of the ants of the Gulf of California islands. Pp. 545-553 in Case, T. J., Cody, M. L. & Ezcurra, E. (eds.) A new island biogeography of the Sea of Cortes. Oxford: Oxford University Press, xvii + 669 pp.


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

    Outputs
    Research continued on the species-level taxonomy, phylogeny and distribution of ants (Hymenoptera: Formicidae). This included field work to collect ants for systematic analyses in Florida, Arizona and in the following California counties: Lake, Napa, Placer, San Diego, Shasta, Siskiyou, Solano, Sonoma, Trinity, Yolo and Yuba. Particular emphasis was placed on collecting representatives of the genera FORMICA and PSEUDOMYRMEX. A museum visit to the Smithsonian Institution permitted examination and identification of ant specimens in that collection. Substantial amounts of material were also studied from other institutions in North America, Latin America, Asia, Australia and Europe. The P.I. identified numerous ant specimens that were sent for identification by personnel in state, federal and other agencies. This included a large volume of ant specimens that were collected as part of a biodiversity survey in southern California by the Biological Resources Division of the US Geological Survey. The P.I. also served as a taxonomic consultant for a similar project by the National Park Service in the Santa Monica Mountains National Recreation Area. A study of the composition of ant communities living in leaf litter was published (Ward, 2000) in addition to a monographic revision of the ant genus TETRAPONERA in the Oriental and Australian biogeographic regions (Ward, 2001).

    Impacts
    The taxonomic studies carried out by the P.I. improve our ability to identify ants and to understand their geographical distribution and ecology. This is important not only in the context of recognizing pestiferous species of ants, but also because native ants are being increasingly used as "bioindicators" of environmental quality (as in the USGS and NPS projects mentioned above).

    Publications

    • Ward, P. S. 2000. Broad-scale patterns of diversity in leaf litter ant communities. Pp. 99-121 in: Agosti, D., Majer, J. D., Alonso, L. E., Schultz, T. R. (eds.) Ants. Standard methods for measuring and monitoring biodiversity. Washington: Smithsonian Institution Press, xix + 280 pp.
    • Ward, P. S. 2001. Taxonomy, phylogeny and biogeography of the ant genus TETRAPONERA (Hymenoptera: Formicidae) in the Oriental and Australian regions. Invertebrate Taxonomy 15:589-665.


    Progress 01/01/00 to 12/31/00

    Outputs
    Systematic and biological investigations were carried out on several groups of ants. Scrutiny of a complex of species in the ant genus PHEIDOLe helped to resolve some longstanding taxonomic problems among populations occurring in California and Baja California (Ward, 2000). A survey of the ants of Santa Cruz Island was published (Wetterer et al., 2000). Additional progress was made on the ant fauna of California through field collections by the P.I. in the following counties: Alpine, Colusa, Lake, Lassen, Los Angeles, Mariposa, Napa, Nevada, Plumas, San Diego, Santa Cruz, Sonoma, Stanislaus, Tehama, and Yolo. Work continued on a global revision of the ants in the subfamily Pseudomyrmecinae, through examination of museum material in various collections. Identification keys for various groups of pseudomyrmecine ants were placed on the web (http://entomology.ucdavis.edu/faculty/ward/psintro.html). A preliminary molecular phylogenetic analysis of CAMPONOTUs suggested that this genus may not be monophyletic (Brady, Gadau and Ward, 2000). Research was initiated on the nesting habits of the Argentine ant (LINEPITHEMA HUMILE) and on the feasibility of using trap nests to reduce population densities of this pest. The P.I. received numerous requests for assistance with ant identification and several thousand ant specimens were examined and identified in the course of responding to these requests.

    Impacts
    Taxonomic knowledge of ants is far from complete, compromising our ability to identify species (both pestiferous and beneficial). Research by the P.I. is designed to remove this impediment, and permit reliable species identification. In addition, fundamental data on the taxonomy, biology and distribution of ants is critically important as we assess changes to community composition brought about by changes in the environment.

    Publications

    • Wetterer, J. K., Ward, P. S., Wetterer, A. L., Longino, J. T., Trager, J. C. and Miller, S. E. 2000. Ants (Hymenoptera: Formicidae) of Santa Cruz Island, California. Bulletin. Southern California Academy of Sciences 99:25-31.
    • Ward, P. S. 2000. On the identity of PHEIDOLE VASLITII Pergande (Hymenoptera: Formicidae), a neglected ant from Baja California. Journal of Hymenoptera Research 9:85-98.
    • Brady, S. G., Gadau, J. and Ward, P. S. 2000. Systematics of the ant genus CAMPONOTUS (Hymenoptera: Formicidae): a preliminary analysis using data from the mitochondrial gene cytochrome oxidase I. Pp. 131-139 in: Austin, A. D., Dowton, M. (eds.) Hymenoptera. Evolution, biodiversity and biological control. Collingwood, Victoria: CSIRO Publishing, xi + 468 pp.


    Progress 01/01/99 to 12/31/99

    Outputs
    Research was conducted on several aspects of ant systematics, evolution and biogeography. Work continued on revisionary studies of the ant subfamily Pseudomyrmecinae, including an assessment of type specimens and other material in European museums. A revision of the PSEUDOMYRMEX VIDUUS group was published (Ward, 1999b). A taxonomic study of a species complex in the ant genus NEIVAMYRMEX was also published (Ward, 1999a). Investigations of phylogenetic relationships among species in the ant genus CAMPONOTUS were carried out (Brady, Gadau & Ward, 1999; Gadau, Brady & Ward, 1999). Taxonomic analyses of ants from California and Baja California involved the examination of a large quantity of specimens in various collections including the Bohart Museum of Entomology, California Academy of Sciences, California Department of Food and Agriculture, Natural History Museum of Los Angeles County, and Museum of Comparative Zoology (Harvard University). Field surveys of ants were carried out in several California parks and reserves including McNee Ranch State Park, San Pedro Valley County Park, Black Diamond Mines Regional Preserve, McKenzie Table Mountain Preserve, and Mary Elizabeth Miller Black Mountain Preserve. Voucher specimens from these surveys were identified and deposited in the Bohart Museum, and reports on the ant faunae were sent to the respective management authorities. Several thousand ant specimens were submitted for identification by individuals from other institutions, and these were duly processed.

    Impacts
    Basic taxonomic research of the kind described above permits reliable identification of ant species. Detailed studies on the ant fauna of California being carried out by the Investigator also provide baseline information on the composition of the indigenous ant fauna, and a yardstick against which to measure changes as a result of habitat destruction and invasion by exotic species.

    Publications

    • Brady, S. G., Gadau, J., Ward, P. S. 1999. Is the ant genus CAMPONOTUS paraphyletic? Pp. 27-28 in: 4th International Hymenopterists Conference, 6-11th January 1999, Canberra, Australia. Program and abstracts. Glen Osmond, South Australia: 4th International Hymenopterists Conference, 96 pp. [Abstract].
    • Gadau, J., Brady, S. G., Ward, P. S. 1999. Systematics, distribution, and ecology of an endemic California CAMPONOTUS QUERCICOLA (Hymenoptera: Formicidae). Annals of the Entomological Society of America 92:514-522.
    • Ward, P. S. 1999a. Deceptive similarity in army ants of the genus NEIVAMYRMEX (Hymenoptera: Formicidae): taxonomy, distribution and biology of N. CALIFORNICUS (Mayr) and N. NIGRESCENS (Cresson). Journal of Hymenoptera Research 8:74-97.
    • Ward, P. S. 1999b. Systematics, biogeography and host plant associations of the PSEUDOMYRMEX VIDUUS group (Hymenoptera: Formicidae), TRIPLARIS- and TACHIGALI-inhabiting ants. Zoological Journal of the Linnean Society 126:451-540.


    Progress 01/01/98 to 12/31/98

    Outputs
    Research was carried out in several areas related to ant systematics and evolution. Taxonomic work continued on the genus PSEUDOMYRMEX, with emphasis on those species that live in cavities (domatia) of specialized "ant-plants" and protect their host plant from herbivory. The PSEUDOMYRMEX VIDUUS group was revised, with the consequent recognition of 13 species, of which three are new. A phylogenetic analysis of this group, combined with data on worker behavior and hostplant associations, suggests that aggressive behavior evolved before the development of mutualistic ant/plant relationships. Another taxonomic study was concerned with resolving the identities of several closely related species of army ants in the genus NEIVAMYRMEX. A detailed morphometric analysis demonstrated that N. CALIFORNICUS is a distinct species, not conspecific with N. NIGRESCENS. Confusion between these two species has arisen because in northern California N. NIGRESCENS has evolved features of morphology (especially integument sculpture) that make it superficially similar to N. CALIFORNICUS. A third systematic study involved a preliminary assessment of relationships in the large genus CAMPONOTUS, using molecular (DNA sequence) data. Field collections of ants were made at multiple locations in California, with the aim of improving our knowledge of ant distribution and ecology in the state. The ants from these collections have been processed, identified and deposited in the Bohart Museum of Entomology.

    Impacts
    (N/A)

    Publications

    • WARD, P.S. 1998. Phylogeny and biogeography of host plant associations in Neotropical pseudomyrmecine ants (Hymenoptera: Formicidae). P. 499 IN Social insects at the Turn of the Millennium, Schwarz & Hogendoorn (eds.), Proc. XIII Internat. Cong. IUSSI,


    Progress 01/01/97 to 12/01/97

    Outputs
    Research was continued on the systematics and distribution of ants (Hymenoptera: Formicidae). Surveys were carried out in a number of California parks and reserves including the Pine Hill Ecological Reserve, Carrizo Plains Natural Area, Frank Raines Park (Del Puerto Canyon), San Bruno Mountain Park, and Elliott Reserve (San Diego County). These field collections yielded new details on the distribution and ecology of California ants. A taxonomic study of several species of army ants in the genus NEIVAMYRMEX was initiated. Morphometric analysis helped to clarify the taxonomic status of NEIVAMYRMEX CALIFORNICUS, which was shown to be a distinct species, endemic to California and heretofore confused with the more widespread NEIVAMYRMEX NIGRESCENS. Research on the systematics and biogeography of the Neotropical ant genus PSEUDOMYRMEX continued. A revision of the PSEUDOMYRMEX VIDUUS group was completed, along with an analysis of its coevolution with specialized ant-plants. A new hypothesis on the origin of ant soldiers--that they originated from ant queens rather than workers--was evaluated and found to be poorly supported. A review of the evidence supports the view that ant soldiers represent a modified form of the worker caste (results published in the journal NATURE). Curation of the ant collection in the Bohart Museum of Entomology continued apace. Large numbers of ant specimens (approximately 3570) were submitted for identification by individuals from other institutions.

    Impacts
    (N/A)

    Publications

    • WARD, P.S. 1997. Ant soldiers are not modified queens. Nature 385:


    Progress 01/01/96 to 12/30/96

    Outputs
    Research in ant systematics was conducted on several fronts. Field work was carried out in California and Brazil, and important new collections were amassed. The ant fauna of a CDFG (California Department of Fish & Game) gabbro site near Shingle Springs, El Dorado County, was surveyed and several outstanding ant finds resulted. Highlights include the occurrence of MYRMICA RUGIVENTRIS (a very rare species, previously unrecorded from northern California) and the co-occurrence of three species of army ants (NEIVAMYRMEX species). Systematic analysis of the Neotropical ant genus PSEUDOMYRMEX continued, and one paper was published on this topic. A comprehensive bibliography of the literature on ant systematics was published and I began making available electronic versions of the bibliography to interested researchers. In Brazil I attended a meeting on the use of ants as indicator taxa in environmental monitoring, and I presented a summary paper on the global distribution and diversity patterns of leaf litter ants. Curation of the ant collection in the Bohart Museum of Entomology continued apace. Large numbers of ant specimens (approximately 3110) were also submitted to me for identification by individuals from other institutions.

    Impacts
    (N/A)

    Publications

    • Olson, D.M. and Ward, P.S. 1996. The ant fauna (Hymenoptera: Formicidae) of Kirindy Forest (tropical dry forest) in western Madagascar. Pp. 161-164 in Ganzhorn, J. U., Sorg, J.-P. (eds.). Ecology and economy of a tropical dry forest in Mada
    • Ward, P.S., Bolton, B.B., Shattuck, S.O., and Brown, W.L., Jr. 1996. A bibliography of ant systematics. Univ. Calif. Publ. Entomol. 116:1-417.
    • Ward, P.S. 1996. A new workerless social parasite in the ant genus PSEUDOMYRMEX (Hymenoptera: Formicidae), with a discussion of the origin of social parasitism in ants. Syst. Entomol. 21:253-263.


    Progress 01/01/95 to 12/30/95

    Outputs
    Research continued in ant systematics with a focus on the New World genus PSEUDOMYRMEX. Field work in northern Argentina produced important material for taxonomic revision, including a new species of workerless social parasite. A phylogenetic analysis of the parasite and its host species (another species of PSEUDOMYRMEX) shows that they are not closely related, a result which contradicts a long-standing generalization ("Emery's Rule") that social insect parasites evolve from their hosts. Compilation of a large computerized database of the literature on ant systematics was completed. The file contains approximately 8000 records. A manuscript ("A bibliography of ant systematics") was submitted for publication and is currently in review. The bibliography will also be made available electronically.

    Impacts
    (N/A)

    Publications


      Progress 01/01/94 to 12/30/94

      Outputs
      Several areas of research in ant systematics were pursued. A large collection ofants from Bolivia (from field work carried out in December 1993) was curated and analyzed. More than 50 species of PSEUDOMYRMEX were recognized, including several new species and a valuable series of specimens in the taxonomically difficult P. PALLIDUS group. Revisionary work was continued on these and other pseudomyrmecine ants. Another area of emphasis was the ongoing development of a bibliographic database (ANTBIB) on the literature of ant systematics and evolution. The database is about 95 percent complete and should be available for publication within a year.

      Impacts
      (N/A)

      Publications


        Progress 01/01/93 to 12/30/93

        Outputs
        Systematic studies were continued on the ant subfamily Pseudomyrmecinae. Field studies were carried out in Madagascar (focusing on the genus TETRAPONERA) and in the Neotropics (with emphasis on the Bolivian species of PSEUDOMYRMEX). Collections from these areas will contribute to future taxonomic revision of TETRAPONERA and PSEUDOMYRMEX by helping to clarify the limits of intra- and interspecific variation. The field work in Madagascar also uncovered a new genus of ant--a bizarre, pale, eyeless subterranean form which is unlike any other known ant. A morphological analysis suggests that the new ant (christened ADETOMYRMA) is cladistically a member of the primitive ponerine tribe, Amblyoponini. The strange morphology of this ant has caused a reevaluation of the basal phylogenetic relationships within Formicidae. ADETOMYRMA was collected in Zombitse Forest, a tract of tropical dry forest which is under severe threat from human activities (slash-and-burn agriculture). Discovery of ADETOMYRMA has been reported in several conservation newsletters, and has helped to focus attention on Madagascar's threatened dry forests.

        Impacts
        (N/A)

        Publications


          Progress 01/01/92 to 12/30/92

          Outputs
          Research was focused on two areas: 1) A systematic study of PSEUDOMYRMEX ants inDominican amber. This led to the description of 11 new fossil species. An analysis of their relationships revealed an affinity with relict species in upland regions of Central America, a finding consistent with the assertion that early in the Tertiary Period Hispaniola was adjacent to, or part of, the Central American mainland. The fossil study also highlighted a marked loss in species diversity of arboreal ants on Hispaniola in Recent times. 2) A taxonomic revision of the obligate acacia-ants (PSEUDOMYRMEX FERRUGINEUS group) which have a mutualistic relationship with swollen-thorn acacias. Several new species were described and their phylogenetic relationships elucidated. Cladistic analysis demonstrated that the ants and their host-plants have experienced "diffuse" coevolution, not one-on-one cooperation.

          Impacts
          (N/A)

          Publications


            Progress 01/01/91 to 12/30/91

            Outputs
            Systematic and biological research on ants (Hymenoptera: Formicidae) has encompassed the following areas: (i) continued species-level studies of the genera PSEUDOMYRMEX and TETRAPONERA, (ii) an investigation of the higher phylogeny of ants at the subfamily and tribal levels, in collaboration with B. Bolton (BMNH) and C. Baroni Urbani (Basel), (iii) continued surveys of the distribution and taxonomy of California ants, (iv) collaborative studies on ant/plant/coccid relationships in Old and New World rainforests, and (v) further development of a bibliographic database on ant systematics. A new subfamily classification of the Formicidae, based on the results of (ii), is probably the most significant achievement of this period. Field work was carried out in Ecuador in August 1991 for the purpose of gathering ant specimens for taxonomic analysis. This trip yielded valuable material, including new species of PSEUDOMYRMEX.

            Impacts
            (N/A)

            Publications


              Progress 01/01/90 to 12/30/90

              Outputs
              Studies were conducted on the biology, taxonomy, and evolution of ants, with emphasis on the subfamily Pseudomyrmecinae. A generic revision of the subfamily resulted in the description of a new genus (MYRCIDRIS) whose only known species is confined to the Amazon rain forests. Phylogenetic analysis suggests that this is part of an ancient lineage of ants which may predate the rift between Africa and South America. Distinguishing features of the two principal pseudomyrmecine genera (PSEUDOMYRMEX, TETRAPONERA) were clarified for the first time. Phylogenetic relationships among the major groups (subfamilies) of ants were explored in a pilot study which involved comparative morphological study of workers, queens, and males. The results suggest the need for a rearrangement of the existing classification of ants, and point to new hypotheses about their evolution. These systematic studies are necessarily global in scope. Field work to collect material for taxonomic analysis was carried out in the United States, Costa Rica, Madagascar, and east Africa.

              Impacts
              (N/A)

              Publications


                Progress 01/01/89 to 12/30/89

                Outputs
                Research continued on the biology and systematics of ants, with emphasis on the subfamily Pseudomyrmecinae. The major species groups of Pseudomyrmex were reevaluated and diagnosed, and two such groups were subjected to detailed taxonomic revision. Taxonomic analysis of other Pseudomyrmex species led to clarification of species limits, particularly among those taxa associated with specialized ant-plants (myrmecophytes). A phylogenetic analysis of pseudomyrmecine ants associated with domatia-bearing plants permitted inferences to be made about the evolution of ant-plant mutualisms. The results indicated the repeated origin of the domatia-inhabiting trait, and considerable "host-plant" switching. Field studies to gather material for taxonomic analysis were carried out in Australia, southeast Asia, and Madagascar, during a sabbatical leave (7/88 to 7/89).

                Impacts
                (N/A)

                Publications


                  Progress 01/01/88 to 12/30/88

                  Outputs
                  This project involves a series of long-term studies on ant taxonomy and biology.Research included a species-level study of the genus Pseudomyrmex. Five previously unrecognized species were described, 39 species names were synonymized, and 7 species had there status reevaluated. Detailed morphological analysis needed for a higher-level phylogenetic analysis of the Pseudomyrmecinae were continued. Additionally, a study was begun on the Australasian genus Leptomyrmex. The unusual biology of members of this genus may reveal valuable biogeographical information.

                  Impacts
                  (N/A)

                  Publications


                    Progress 01/01/87 to 12/30/87

                    Outputs
                    Research continued on the systematics of the ant subfamily Pseudomyrmecinae, including both higher-level phylogenetic analysis and species-level revisionary studies on selected species groups in the genus Pseudomyrmex. Museum visits to collections in Europe helped to clarify nomenclatural problems; field studies were carried out in Mexico and South America. A study of the introduced Argentine ant in California was completed. This ant was shown to have a strongly deleterious effect on the indigenous ant fauna, in areas where it has become established. I also conducted a taxonomic study on mesic elements in the western Nearctic ant fauna, with emphasis on the genera Amblyopone, Proceratium, and Smithistruma. New species were discovered and described in the last two genera. These species evidently represent the remnants of a fauna which was formerly more widespread in the western United States during wetter climatic periods.

                    Impacts
                    (N/A)

                    Publications


                      Progress 01/01/86 to 12/30/86

                      Outputs
                      This project involves a series of long-term studies of ant taxonomy and biology. Studies focused on the systematics of the ant subfamily Pseudomyrmecinae, with particular emphasis on a generic revision, and alpha-taxonomy of the Pseudomyrmex elongatus and P. subtilissimus species groups. Work continued on the ecology and social behavior of ants, including investigations on the primitive ponerine genus, Rhytidoponera, and on the introduced Argentine ant, Iridomyrmex humilis. The distribution of I. humilis in natural habitats of the lower Sacramento Valley and its impact on the native ant fauna were examined. As a side project, I assisted in the analysis of phylogenetic relationships among clones of Trypanosoma cruzi, the causative agent of Chagas' disease. This study revealed a complex, multiclonal population structure, which has significant medical implications.

                      Impacts
                      (N/A)

                      Publications


                        Progress 01/01/85 to 12/30/85

                        Outputs
                        This project involves a series of continuing studies on the taxonomy and biologyof ants (Hymenoptera:Formicidae). During 1985, research was focused on the systematics of ants in the subfamily Pseudomyrmecinae. A revision of the Nearctic Pseudomyrmex revealed ten species, two of which were new to science. One of the new species is a workerless social parasite, the first recorded for the subfamily. An analysis of taxonomic congruence in New Caledonian ants of the genus Rhytidoponera was published. Studies continued on the distribution of the Argentine ant in the lower Sacramento Valley, and its effect on the native ant fauna.

                        Impacts
                        (N/A)

                        Publications


                          Progress 01/01/84 to 12/30/84

                          Outputs
                          This project involves several studies on systematic relationships and populationbiology of ants (Hymenoptera: Formicidae). During 1984 research was focussed on (i) an analysis of taxonomic congruence in New Caledonian ants of the genus Rhytidoponera, (ii) a taxonomic revision of the North American species of Pseudomyrmex (Quaest. Entomol., in press), (iii) an investigation of the distribution and abundance of the Argentine ant in natural habitats of northern California.

                          Impacts
                          (N/A)

                          Publications


                            Progress 01/01/83 to 12/30/83

                            Outputs
                            This project is concerned with the systemistics and biology of social insects, with particular emphasis on ants (Hymenoptera:Formicidae). Current (1983) research has been focussed on: (i) genetic relatedness and social organization in primitive ponerine ants; (ii) phylogenetic relationships in the New World genus, Pseudomyrmex. An investigation into Argentine ant (Iridomyrmex humilis) distribution in natural habitats of northern California was also initiated.

                            Impacts
                            (N/A)

                            Publications


                              Progress 01/01/82 to 12/30/82

                              Outputs
                              Work has continued on the systematics and evolutionary biology of various socialHymenoptera, with emphasis on the systematics and population structure of ants (Formicidae). A taxonomic revision of the genus Rhytidoponera in New Caledonia is nearing completion. Field studies of the New World genus Pseudomyrmex were carried out in Arizona, Texas, and Florida resulting in the acquisition of valuable material for electrophoresis and phylogenetic analysis. A taxonomic and genetic study of isozyme variation in Tetranychus spider-mites (which have a haplodiploid genetic system like Hymenoptera) has revealed genetic markers which should prove useful in studying the biology of these pestiferous species.

                              Impacts
                              (N/A)

                              Publications


                                Progress 01/01/81 to 12/30/81

                                Outputs
                                Investigations have been carrie out on the systematics and population biology ofseveral groups of arthropods: An analysis of morphological and allozyme variation in 15-20 New Caledonian species of the ant genus Rhytidoponera revealed considerable concordance among classifications based on disparate data sets. Numerical taxonomic analysis and SEM studies of morphology continue. As part of a long-term project on ants of the Pseudomyrmex pallidus group, field studies were carried out in Texas and Arizona, resulting in the accrual of new data on colony composition and nest-site ecology. Preparatory to a study of genetic relatedness in honey bee (Apis mellifera) colonies, electrophoretic assays for two polymorphic enzyme systems (malate dehydrogenase and esterase) were developed. An examination of electrophoretically detectable variation in malate dehydrogenase enzymes in spider mites (Tetranychus spp.) revealed the first reported allozyme polymorphism in these organisms, and provided genetic data on population structure and systematic relationships.

                                Impacts
                                (N/A)

                                Publications