Progress 10/01/07 to 09/30/12
Outputs
OUTPUTS: During this period additional specimens of Hypherpes beetles were collected for future DNA analysis. Three target species were the focus data collection for morphometric analysis to begin a study of sexual size dimorphisms and character displacement among Californian species. In conjunction with an ongoing NSF funded project more than 2000 Californian Pterostichus lama specimens were images and their data entered into the Essig Museum database. Of these Pterostichus several hundred have been used for capture of morphometric data. Three undergraduate students were trained in morphometric and phylogenetic methods using specimens and data from the project. Preliminary surveys were done in local parks and on public lands. PARTICIPANTS: PI Kipling Will TARGET AUDIENCES: Biologists, in particular systematists and taxonomists. The project provides taxonomic knowledge and identify reference specimens PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.
Impacts
Numerous specimen identifications were done for individuals and museums and so provided comparative material for use by other researchers. Preliminary surveys were done in local parks and on public lands. Data from the morphometric analyses are too incomplete to report results.
Publications
- A new species of Lesticus Dejean, 1828 (Coleoptera, Carabidae) from the Finisterre Range, Papua New Guinea and a key to the genera of pterostichine-like Harpalinae of New Guinea K Will, DH Kavanaugh. 2012. ZooKeys, 27
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Progress 01/01/11 to 12/31/11
Outputs
OUTPUTS: During this period approximately 10,000 specimens of carabid beetles were prepared and sorted. In conjunction with an ongoing NSF funded project approximately 1000 Californian Pterostichus specimens were images and their data entered into the Essig Museum database. Of these Pterostichus several hundred have been used for capture of morphometric data. Three undergraduate students were trained in morphometric and phylogenetic methods using specimens and data from the project. PARTICIPANTS: PI Kipling Will TARGET AUDIENCES: Biologists, in particular systematists and taxonomists. The project provides taxonomic knowledge and identify reference specimens. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.
Impacts
Numerous specimen identifications were done for individuals and museums and so provided comparative material for use by other researchers.
Publications
- Will, K.W. 2011. Taxonomic review of the Pterostichini and Loxandrini fauna of New Caledonia (Coleoptera, Carabidae). ZooKeys 147: 337-397.
- Will, K.W. 2011. Chaetauchenium Tschitcherine, 1900 (Carabidae: Pterostichini) a new species and new status for a lineage from the Valdivian temperate rainforest. Gayana (Concepc.) 75(2): 192-197.
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Progress 01/01/10 to 12/31/10
Outputs
OUTPUTS: Pterostichine taxa were collected for DNA extraction in Australia. Six months were spent focusing on field work and resulted in approximately 150 sampling sites visited during the reporting period and about 5,000 specimens being collected. Preliminary sorting and identification was done and high-priority specimens were targeted for DNA sequencing. Related beetles that were collected during field work but are not the subject the pterostichine project have been sent to students and experts in the United States and Australia to be used in a variety of studies. Digital images of exemplar carabid species were taken for use in electronic identification keys. PARTICIPANTS: The Australian National Insect Collection, CSIRO, Canberra and UCB Essig Museum of Entomology, contribute and receive specimens from this study. Collaborators and contacts include Dr. Stephen Cameron, Dr. Adam Slipinslki (CSIRO), Dr. Geoff Monteith (QM) and David Maddison (Oregon State U.). Training opportunities as part of this project were given to Traci Grzymala and Matt VanDam (graduate students). Students were trained in lab techniques related to preparation and DNA extraction/PCR. TARGET AUDIENCES: Not relevant to this project. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.
Impacts
Sampling Pterostichine carabid beetles for North America and worldwide was the focus during this period. The current matrix of DNA data that targets the phylogeny worldwide was expanded for numerous rare taxa. A significant amount of student training was done in the lab, e.g. genomic extraction, PCR and in the field, e.g. collecting, specimen preservation and preparation.
Publications
- No publications reported this period
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Progress 01/01/09 to 12/31/09
Outputs
OUTPUTS: OUTPUTS: Based on data collected during this period the subgenus Hypherpes is found to form a monophyletic group with exemplars of other Northern Hemisphere pterostichine subgenera based on an analysis of morphology and 28S rDNA, COI and COII mtDNA sequence data. A significant amount of student training was done including activities in the lab, e.g. whole genomic extractions, PCR, morphological dissections, larval rearing and in the field, e.g. collecting, specimen preservation and preparation. Presentations of results from this research were given at local, national and international entomological meetings. Related beetles that were collected during field work but are not the subject the pterostichine project have been sent to students and experts in the United States to be used in a variety of studies. Digital images of exemplar carabid species were taken for use in electronic identification keys. Students were trained in the development of interactive electronic identification keys and some of these have been made available via the internet. PARTICIPANTS: Partner organizations include The California Academy of Science and UCB Essig Museum of Entomology. Both contribute and receive specimens from this study. Collaborators and contacts include David Kavanagh (CAS), James Labonte (Oregon Dept. of Agr.) and David Maddison (Oregon State U.). Training opportunities as part of this project were given to Matt VanDam (graduate student), Mae Hou (undergraduate), Elizabeth Morrill (undergraduate) and Andrea Gadon (undergraduate) Undergraduate and graduate students have gained laboratory and field experience. Students were instructed in extraction and PCR methods, sequence data analysis and collection, identification and rearing of beetles. Using beetles, other insects and spiders, graduate students were instructed of the development of Lucid interactive electronic keys. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.
Impacts
Activities during this period were primarily data accumulation and specimen collecting and curation. Major findings are expected during a subsequent. analysis phase, but preliminary results suggest that the worldwide fauna of pterostichines is monophyletic including the Hypherpes clade. A surprising result that forces reevaluation of the polarity of character evolution in the group.
Publications
- No publications reported this period
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Progress 01/01/08 to 12/31/08
Outputs
OUTPUTS: DNA sequencing efforts increased during this period and we continue to build the database of DNA characters for Pterostichine carabid beetles for North America and worldwide. The current matrix of DNA data that targets species in the subgenus Hypherpes and exemplars of other North American pterostichine subgenera was completed and analyzed. For this exemplar sample of pterostichine species 28S rDNA and COI and COII mtDNA sequence data were used to reconstruct a phylogenetic hypothesis for generic and subgeneric taxa putatively in or related to the western North American subgenus Hypherpes. The evolution of eye reduction and its relationship to genetic divergence and life history was studied. Using these data biogeographical patterns of Leptoferonia species in the western United States and between transcontinental populations of Hypherpes were studied. A significant amount of student training was done in the lab, e.g. genomic extraction, PCR dissections and larval rearing and in the field, e.g. collecting, specimen preservation and preparation. Presentations of results from this research were given at local, national and international entomological meetings. Related beetles that were collected during field work but are not the subject the pterostichine project have been sent to students and experts in the United States to be used in a variety of studies. Digital images of exemplar carabid species were taken for use in electronic identification keys. Students were trained in the development of interactive electronic identification keys and some of these have been made available via the internet. PARTICIPANTS: Partner organizations include The California Academy of Science and UCB Essig Museum of Entomology. Both contribute and receive specimens from this study. Collaborators and contacts include Aman Gill (SUNY), David Kavanagh (CAS), James Labonte (Oregon Dept. of Agr.) and David Maddison (U of AZ). Training opportunities as part of this project were given to Ainsley Seago (graduate student), Steven Lew (graduate student), Mae Hou (undergraduate), Myfanwy Rowlens (undergraduate) and Stephanie Skophammer (undergraduate) Undergraduate and graduate students have gained laboratory and field experience. Students were instructed in extraction and PCR methods, sequence data analysis and collection, identification and rearing of beetles. Using beetles, other insects and spiders, graduate students were instructed of the development of Lucid interactive electronic keys. TARGET AUDIENCES: Keys and information developed under this project are part of student thesis projects or side projects that have been released to the public via our web pages. The general public, campus community, local businesses and state government agencies have all submitted to the PI specimens of Californian ground beetles for identification. Because of the efforts of this project the PI is able to provide free diagnostic services to these units and people. In most cases concerns that these beetles may pose a risk to human health and/or be an economic threat (pest) were eliminated and suggested changes to ameliorate the nuisance were adopted. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.
Impacts
The monophyly of Pterostichus is equivocal as the position of the subgenus Bothriopterus, depending on methods of sequence alignment and gap region treatment, moves outside of the genus. Pterostichus is found to be monophyletic in the combined data analysis if Cyclotrachelus and Tapinopterus are included in a larger concept of the genus. No taxon previously suggested as a close relative of Hypherpes was found to be closely related to the subgenus. The sister-group of Hypherpes remains unclear, but there is some support for a clade of Pseudoferonina + Cryobius as the adelphotaxon. Taxa included in current classifications of Hypherpes compose a group that is in fact a complex of Hypherpes sensu stricto and two other subgenera, Leptoferonia and Anilloferonia. Our analyses show that these three taxa are well supported as subgenera and reciprocally monophyletic, with the only change to previous taxonomic concepts of included species being the transfer of Pterostichus rothi from Anilloferonia to Leptoferonia. In Leptoferonia the DNA data support all species groups established by Hacker using morphological characters, with the exception of the inopinus-group. We found that significant reduction of the compound eyes has occurred independently at least five and possibly seven times in the Hypherpes complex. As many as five separate instances of eye reduction may have occurred in Leptoferonia alone. A concentrated changes test was used to show that there is a significant correlation between microphthalmy and autapomorphic sequence data as represented by longer than average terminal branch lengths based on Bayesian estimates of change per site. However, taxon pair contrasts show no consistent pattern of absolute difference of evolutionary rate or directionality of differences between small-eyed taxa and their sister species or sister clade. Repeated patterns of allopatric distributions are found for species-pairs of Leptoferonia, which consist of divisions along a north/south axis near the Pacific Coast and in the Sierra Nevada Range, or east/west divisions between coastal species and inland or Sierran species. In addition to allopatric biogeographic patterns, instances of sympatry in closely related species are interpreted to have been the result of two reduced-eye species moving into the deep litter and soil layer, thereby ecologically differentiating from near-surface leaf-litter and log dwelling species. Pterostichus morionides, which is restricted to the Sierra Nevada Mountains in western North America, is found to be sister to P. adoxus and P. tristis, the only species of Hypherpes in eastern North America. This grouping was further tested by using a subset of taxa for 18S rDNA, CAD and wg sequence data and was found in some or all most-parsimonious trees for these data. In cases where they did not form a clade, they usually formed a convex group. Although counterintuitive due to the unusual disjunct biogeographic connection of these two areas and the generally dissimilar form of the adults, the mta-clade is very well supported by the DNA sequence data.
Publications
- Will, K.W. and Gill, A.S. 2008. Phylogeny and classification of Hypherpes auctorum (Coleoptera: Carabidae: Pterostichini: Pterostichus). Annals of Carnegie Museum. 77(1)93-127.
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Progress 01/01/07 to 12/31/07
Outputs
OUTPUTS: I continue to build the database of DNA characters for Pterostichine carabid beetles. The existing DNA sequence data collection that targets species in the subgenus Hypherpes and other North American pterostichine subgenera has been expanded and is contributing to the development of worldwide phylogenetic hypothesis for the tribe. Over 400 sequences have been submitted to Genbank (28s, COI, COII, CAD, WG, 18S). Sampling is underway to broaden the species-level and population-level samples from across California. Select taxa have had all available specimens data-based and georeferenced so that plotting of distributions based on these records can be access via the internet. Four highly endemic, nearly blind, soil-dwelling species were newly described from the Californian fauna. An interactive Lucid key to species of Leptoferonia is under development. I have finished most of the character scoring and assembled images of all male genitalia and secondary sexual characters needed
for identification. One of the most significant features shared by all ground beetles is the chemical defense system they employ. All beetles have a pair of glands in their abdomen that emit foul smelling acidic compounds when they are threatened. In order to investigate how much fluid is discharged and in what relative amounts of these compounds are release in response to attack I undertook a study in which beetles were weighed before and after spraying in order to quantify the amount sprayed, the amount of various compounds in the spray were determined and this related to quantities that elicit alarm reactions in ants. I have begun investigating products of the male accessory glands. These glandular fluids are probably essential for reproductive success, however, the constituents, seasonal variation and what role they play in females is unknown. Californian pterostichines provide an excellent model system for this study.
PARTICIPANTS: principal investigator Kipling Will Students participating in and trained in project Aman Gill - Undergraduate UCB Ainsley Seago - Graduate UCB Steven Lew - Graduate UCB Collaborators and contacts Dr. Athula Attygalle - Stevens Technical Institute, Hoboken, NJ
Impacts
Based on the phylogenetic analyses the Holarctic genus Pterostichus was found to not be monophyletic, however, the western North American subgenera Hypherpes, Leptoferonia and Anilloferonia where found to be reciprocally monophyletic. This has important implications for studies of the group world wide. Investigation of the biosynthetic pathway of tiglic and related acidic compounds in pterostichine beetles has led to a greater understanding of a pathway of synthesis and our methods and results have been picked up and used by other researchers investigating the same or similar compounds in other organisms.
Publications
- Will, Kipling W. 2007. Four new species of the subgenus Leptoferonia Casey (Coleoptera, Carabidae, Pterostichus Bonelli) from California. Proceedings of the California Academy of Sciences. 58(4):49-57.
- Attygalle, A.B., Wu, X. and & Kipling W. Will. 2007. Biosynthesis of tiglic, ethacrylic, and 2-methylbutyric acids in a carabid Beetle, Pterostichus (Hypherpes) californicus. Journal of Chemical Ecology. 33:963-970.
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Progress 01/01/06 to 12/31/06
Outputs
The primary accomplishment this year was the completion of DNA sequence data collection for target species in the ground beetle pterostichine subgenus Hypherpes and from exemplars of nearly all North American pterostichine subgenera. Additionally, putative related species from Europe and Asia were also added to the molecular data matrix. A total of 157 terminals for six sequence regions (28s, COI, COII) have been sequenced. A peculiar relationship between Sierran and Appalachian species was detected in preliminary analyses and so an additional three loci have been sequenced for a subset of taxa (CAD, WG, 18s). As a group previously recognized, but currently synonymized with Hypherpes by other workers, Leptoferonia is found to be monophyletic in all analyses. This is significant, as within the Leptoferonia species clade, eye-reduction and subterranean life history has evolved at least four separate times. A paper has been submitted that describes four new species of
Californian Leptoferonia. Because 28s is a variable length sequence primary alignment is a significant issue. Using this dataset I have completed a series of analyses to better understand the general phenomenon and impact of different alignments.
Impacts
This work shows significant promise in using morphological and multiple DNA sequence data sources to develop phylogenetic hypotheses for Hypherpes in California. This analysis will allow for revision of existing species, development of identification tools and description of new species. These data are vital to understanding the biodiversity of California.
Publications
- Will, K.W. 2006. A new species of the New Caledonian genus Sphodrosomus Perroud (Coleoptera: Carabidae: Pterostichini). Memoirs of the Queensland Museum.
- Cameron, S., Rubinoff, D. and Will, K. 2006. Who Will Actually Use DNA Barcoding and What Will it Cost? Systematic Biology. 55(5):844-847.
- Park, J. K., Dam, H.T and Will, K. 2006. Carabidae from Vietnam (Coleoptera). Journal of Asia-Pacific Entomology. 9(2):1-20.
- Will, K.W. 2006. Book review: Evolution of the Insects. Grimaldi, D. and M. Engel. 2005. Bioscience 56(6):538-539.
- Rubinoff, D., Cameron, S. and Will, K. 2006. Are plant DNA barcodes a search for the Holy Grail? Trends in Ecology and Evolution 21(1):1-2.
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Progress 01/01/05 to 12/31/05
Outputs
The primary accomplishment this year was the near completion of DNA sequence data collection for target species in the ground beetle pterostichine subgenus Hypherpes and from exemplars of nearly all North American pterostichine subgenera. Additionally, putative related species from Europe and Asia were also added to the molecular data matrix. A total of 136 terminals for three sequence regions (28s, COI, COII) have been sequenced. A peculiar relationship between Sierran and Appalachian species was detected in preliminary analyses and so an additional three loci have been sequenced for a subset of taxa (H3, WG, 18s). As a group previously recognized, but currently synonymized with Hypherpes by other workers, Leptoferonia is found to be monophyletic in all analyses. This is significant, as within the Leptoferonia species clade, eye-reduction and subterranean life history has evolved at least four separate times. I have been able to recognize four undescribed species of
Leptoferonia and two new species of Hypherpes (in the restricted sense) using morphological and molecular data. An electronic, interactive key (Lucid) to the species of Leptoferonia is under construction and is largely functional. Several field trips have been conducted this year to various sites in California and Idaho for extensive sampling and now nearly every species of Leptoferonia is now represented in the analysis.
Impacts
This work shows significant promise in using morphological and multiple DNA sequence data sources to develop phylogenetic hypotheses for Hypherpes in California. This analysis will allow for revision of existing species, development of identification tools and description of new species. These data are vital to understanding the biodiversity of California.
Publications
- Will, K.W. and J.K. Liebherr. 2005. Removal of Hannaphota distincta Landin (Coleoptera: Carabidae) from Pterostichini to Platynini. The Coleopterists Bulletin. 59(4): 448-449.
- Will, K.W, Mishler, B.D. and Q.D.Wheeler. 2005. The perils of DNA barcoding and the need for integrative taxonomy. Systematic Biology. 54(5):844-851.
- Will, K.W. 2005. Book review: The Genus Carabus in Europe. A Synthesis. by Turin, H., L. Penev & A. Casale (eds.). Coleopterists Bulletin. 59(2):403-404.
- Will, K.W. 2005. New tribal and generic placement for taxa of Pterostichini (auct.) (Coleoptera: Carabidae) from the Juan Fernandez Archipelago, Chile with taxonomic notes on Trirammatus Chaudoir, 1838. The Pan-Pacific Entomologist. 81(1/2):68-75.
- Arndt, E., Beutel, R. G.Beutel & Will, K. 2005. 7.8. Carabidae Latreille, 1802. in Handbook of Zoology, Vol. IV Arthropoda: Insecta (series ed. By N.P. Kristensen and R.G. Beutel). Part 38. Coleoptera, Vol. 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga (partim) (vol. ed. by R.G. Beutel and R.A.B. Leschen). Walter De Gruyter, Berlin, New York.
- Will. K.W. 2005. The Neotropical genera Oxycrepis Reiche and Stolonis Motschulsky: a taxonomic review, key to the described species and description of new Stolonis species from Ecuador (Coleoptera: Carabidae: Loxandrini). Zootaxa. 1049:1-17.
- Will, K.W. 2005. Book review: The Types of Carabidae (Coleoptera) in the Zoologisch Museum Amsterdam, Predominantly the Collection ]"Oskar Vogt". A. de Boer. Annals of the Entomological Society of America. 98(3):415.
- Will, K.W., J.K.Liebherr, D.R.Maddison and J.Galian. 2005. Absence asymmetry: the evolution of monorchid beetles (Insecta: Coleoptera: Carabidae). Journal of Morphology. 264:75-93.
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Progress 01/01/04 to 12/31/04
Outputs
Work focused on sampling DNA sequence data and rearing larvae of beetles in the Californian subgenus Hypherpes and in other putatively related subgenera. Continued study of morphological systems and sampling led to the discovery of three undescribed, nearly blind soil-dwelling species unique to California. Over 20 different sites throughout the state were sampled for specimens and hundreds of individuals suitable for rearing larvae and/or DNA extraction were collected. In the course of this sampling and study of museum material, three rare deep-soil species were found that have not yet been described or named. Unlike their typical surface and near-surface dwelling relatives, these beetles have extremely reduced eyes and are quite pale in color. These unusual species have very restricted ranges and peculiar life histories, and they are very important for building an overall understanding of California biodiversity and will be useful in developing any conservation plan
that includes information on invertebrates. A phylogenetic analysis of DNA sequence data from three gene regions (COI, COII and 28s) was done for 77 terminals that represent a nearly complete matrix of exemplars for major groups of pterostichine ground beetles in North America with an emphasis on species in the Californian subgenus Hypherpes. This analysis shows that although there is little morphological evidence, the DNA data suggest that Hypherpes is a natural group of species more closely related to each other than any of them are to other groups of similar beetles. This is an important first step in being able to completely revise all species in the group and develop a sound understanding of the evolutionary history of the species in California. Larvae in this group are little known and virtually un-identifiable. We successfully established a rearing protocol and have begun associating adults and larvae so that ultimately species can be identified for all life stages. Larvae
potentially have characteristics that will inform us about the evolution of the group. Larvae are also very likely significant predators so the ability to identify them in agro-ecosystems will be important.
Impacts
This work shows significant promise in using morphological and multiple DNA sequence data sources to develop phylogenetic hypotheses for Hypherpes in California. This analysis will allow for revision of existing species, development of identification tools and description of new species. These data are vital to understanding the biodiversity of California.
Publications
- Attygalle B.A., Wu, X., Ruzicka, J., Rao, S., Garcia, S., Herath, K., Meinwald, J., Maddison, D.R. and Will, K.W. 2004. Defensive chemicals of two species of Trachypachus Motschulski. Journal of Chemical Ecology, 30(3):577-588.
- Will, K.W. 2004. New species of the South American loxandrine genus Metoncidus Bates (Coleoptera: Carabidae). Revista Brasileira de Entomologia. 48(4):441-446.
- Will, K.W. 2004. A remarkable new species of Trirammatus Chaudoir (Coleoptera: Carabidae: Pterostichini) from the Valdivian forest of Chile. Zootaxa. 758:1-9. http://www.mapress.com/zootaxa/2004f/zt00758.pdf
- Will, K.W. 2004. Review and cladistic analysis of the generic-level taxa of Morionini Brulle (Coleoptera: Carabidae). The Pan-Pacific Entomologist. 79(3/4):212-229. ***interactive Lucid and Lucid Phoenix keys for Morionini genera at *** http://nature.berkeley.edu/~kiplingw/Morionini
- Will, K.W. and Rubinoff, D. 2004. Myth of the molecule: DNA barcodes for species cannot replace morphology for identification and classification. Cladistics. 20(1):47-55.
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Progress 01/01/03 to 12/31/03
Outputs
During this period I acquired an additional 6000 specimens of Pterostichus (Hypherpes) species from museums in North America and collected fresh material. Four pilot studies were conducted using this material, 1. biosynthesis of naturally produced defensive chemicals, 2. survey of variation in P. lama, 3. preliminary analysis of Hypherpes taxa using DNA sequences data and 4. investigation of chromosome variation. All ground beetles produce irritating chemicals from pygidial glands. Compounds are exuded as a defense against predators. This ability is thought to be essential to their evolutionary success. To better understand how they produce these compounds, individuals of P. californicus were injected with labeled and unlabeled amino acids. Incorporation of labeled amino acids into the resulting defensive mixture was tracked. This allowed us to deduce the chemical pathway used to make the defensive secretions. Additionally, we induced production of compounds not part
of their normal synthesis. Though very preliminary, these results suggest a potentially important new line of research. In order to assess variation in morphology and DNA sequence data, a pilot study was done on P. lama, a large-sized and widespread species in California. This provides an estimate of variation expected for the subgenus overall and is an important first step toward revising the group. Ultimately, revision will make identification of specimens possible for forest and agro-ecologists state-wide. From this initial study we now know, for this exemplar species, that morphological variation is lower than DNA sequence variation for our selected marker. Based on this we have begun implementing a refined sampling strategy. Preliminary results suggest population-level structure is important in understanding the group. Based on the initial study of P. lama and targeting a series of taxa thought to represent the breadth of genus in North America, an initial phylogenetic hypothesis
was developed based on the mitochondrial gene COI. Results indicate many of the putatively included species are in fact members of the Hypherpes while several geographically separated, but morphologically similar, taxa are possibly more related to other groups. We progressed toward integrating karyotype data into the broader Hypherpes analysis. Karyotype data such as chromosome number, size, sex chromosome morphology and sex determination mechanism are used to inform phylogenetic hypotheses within Carabids. Karyotypes must be obtained from fresh gonadal tissue. Thus far, we have been accruing California Hypherpes species and dissecting males to obtain testes. We have established a protocol to view chromosomes, digitally photograph and then rearranged the images to present a karyotype. Once a suitable sampling of Hypherpes specimens has been obtained, dissected and karyotyped, a comparative analysis will be possible. So far we have successfully examined two species, P. lama and P.
morionides. No published data exists for these taxa and these are the first Hypherpes species to have their karyotype examined.
Impacts
This year's studies show significant promise in using morphological, chromosome and mtDNA data to develop phylogenetic and population-level hypotheses for Hypherpes in California, which is the first step toward species-level revision of the group. Studies on defensive chemicals show for the first time the biosynthetic pathway for secretions of the beetles and suggest a potent new area of research.
Publications
- Will, K.W. 2003. Insects as research tools. In Encyclopedia of Insects. Resh, V.H. and R. T. Carde eds. Academic Press.
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Progress 01/01/02 to 12/31/02
Outputs
I have acquired beetle specimens (Pterostichus (Hypherpes) species) from 10 museums in North America, accumulating over 9,000 individuals for this study and designated an additional 15,000 specimens in the California Academy of Sciences for later study. The subgenus clearly has numerous undescribed species, most from California. Many Hypherpes species occur in and near agricultural systems and working with this material will result in tools for identification of these generalist predators and provide a database of their distribution. I have collected specimens from numerous sites in California and Oregon for use in molecular sequencing, internal organ morphology and chromosome studies. Preliminary data for all these systems has been collected with an emphasis on P.(Hypherpes) lama, the largest species of pterostichine ground beetle in North America. This species' wide latitudinal range, from British Columbia to southern California, and elevation range, sea-level to
3,000m, makes it ideal for initial study to determine the approximate rate of character change to expect in the subgenus. Before selecting particular gene sequences or character systems to focus on for the revision of the group, it is essential to use a few exemplars to gage the utility of the characters. Specifically, studies of the male genitalia have been done for specimens from across the entire range of P. (Hypherpes) lama. The structure of various characteristic of this organ are usually useful in delimiting species and often provide clues regarding population structure and breeding biology. Despite the broad range of P.(Hypherpes) lama, and apparent separation of the various populations, the male structures appear highly conserved within the species. Mitochondrial DNA sequencing has been done for a many beetles that are presumed near relatives of Hypherpes, a dozen exemplar taxa in the subgenus and six populations of P.(hypherpes) lama. These data will be combined with
morphological data to develop a phylogenetic hypotheses for the group and address species and population boundaries. Based on this type of analysis the group's phylogeny, its history in California and how they have come to be distributed where they are can be deduced. Parallel to this study I have collected distantly related but ecologically similar pterostichine beetles in Chile. During field work I visited seven national parks in Chile to sample the ground beetle fauna. Specimens were collected for morphological, DNA and defensive chemical analyses. These southern taxa share many similarities to Hypherpes and by working on both systems simultaneously I will better be able to separate the effects of inheritance of characters and ecological adaptations in pterostichine ground beetles.
Impacts
Pterostichine ground beetles are relatively large and abundant generalist predators found commonly in wide variety of habitats in California and much of western North America. Some species are important predators of pest insects in agro-ecosystems, while others are restricted to threatened habitats. Describing and understanding ptersotichine beetles has consequences for both land usage and pest management.
Publications
- Will, K. W. 2002. Revision of the New World abariform genera Neotalus n. gen. and Abaris Dejean (Coleoptera: Carabidae: Pterostichini (Auctorum)). Annals of Carnegie Museum 71(3) 143-213.
- Will, K.W. and Liebherr, J.K. 2002. A new tribal placement for Glyptolenus mirabilis (Straneo) (Coleoptera: Carabidae), comb. n., and description of G. straneoi, sp. n. from Ecuador. Studies on Neotropical Fauna and Environment 37(1): 59-64.
- Will, K.W. 2001. A new species of Harpalus Latreille (Coleoptera: Carabidae) from southeastern North America. Coleopterists Bulletin 55(4): 447-452.
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