Source: UNIVERSITY OF CALIFORNIA, DAVIS submitted to NRP
SYSTEMATICS AND EVOLUTION OF SCALE INSECTS (HEMIPTERA: COCCOIDEA)
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
0186275
Grant No.
(N/A)
Cumulative Award Amt.
(N/A)
Proposal No.
(N/A)
Multistate No.
(N/A)
Project Start Date
Oct 1, 2005
Project End Date
Sep 30, 2011
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
Identification of scale insects requires expert knowledge and technical skills. Also many genera are poorly studied with inadequate literature for identification. For example, in the family Margarodidae, Icerya species and Matsucoccus species can be pests of agricultural crops and forest pine trees, respectively, and yet neither group has been studied by modern systematic methods. Taxonomic and phylogenetic work on scale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. In addition, such research provides a solid framework for studying the comparative biology of scale insects, and will assist in determining the native areas of new pest species. This project aims to provide taxonomic and phylogenetic data on scale insects to allow accurate identification, improved classification and accessible biological information. Research on mealybugs (family Pseudococcidae) is especially important because this large scale insect family (about 2,000 species) includes many significant crop pests. Taxonomic research on mealybug and other scale insect groups (such as margarodids) that include pest or potential pest species, will have direct benefit to applied entomologists who attempt to identify, quarantine or control scale pests, and indirectly will benefit everyone who uses Californian agricultural, horticultural or forestry products.
Animal Health Component
10%
Research Effort Categories
Basic
90%
Applied
10%
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
21131101130100%
Goals / Objectives
1. Phylogenetic analysis of molecular and morphological data to estimate evolutionary relationships of scale insects (superfamily Coccoidea).
2. Preparation of taxonomic revisions of scale insect genera, especially of the families Coccidae, Eriococcidae, Margarodidae, and Pseudococcidae.
3. Provision of taxonomic and biological information on scale insects found in California.
4. Provision of accurate and rapid identifications of scale insect specimens.
Project Methods
The research will be conducted using traditional taxonomic methods of field collection, microscope slide-mounting and examination of specimens, followed by preparation of illustrated descriptions and keys, combined with phylogenetic (=cladistic) analysis of morphological and molecular data. Additional morphological, locality and host-plant data will be obtained from museum specimens.

Progress 10/01/05 to 09/30/11

Outputs
OUTPUTS: Dr. Gullan has completed taxonomic and phylogenetic work that is essential for the accurate identification of scale insects (Coccoidea), including pests and potential pests, and especially mealybugs (family Pseudococcidae) and lac scale insects (family Kerriidae). The work has provided modern identification tools as well as biological information, and allows the association of biological data with names. In addition, the research provides a solid framework for studying the comparative biology of scale insects, and the worldwide perspective will assist in determining the native areas of new pest species as they enter the USA. The results of the research have been disseminated at conferences, during seminars and workshops, and via peer-reviewed publications. The latter are distributed as PDFs to colleagues who are involved in pest diagnostics or research on scale insects (Coccoidea). Dr. Gullan also answered many email enquiries about scale insects and frequently sent PDFs of her publications, as well as performing specialist identifications of scale insects based on her taxonomic research. The most significant outputs for the entire project (since year 2000) have been almost 50 publications on scale insects, mostly in peer-reviewed journals. The most significant dissemination activities have been (1) the distribution of the above articles to end-users, (2) seminars particularly to the pest diagnostic staff at the California Department of Food & Agriculture in Sacramento, and (3) the provision of species identifications for hundreds of scale insect samples submitted by staff at state and federal agencies, universities and by members of the public PARTICIPANTS: The individuals who were project collaborators in 2010-2011 (listed alphabetically): Ms. S.I. Han, UC Davis, USA Dr. N.B. Hardy, University of New Mexico, Albuquerque, USA Dr. C.J. Hodgson, The National Museum of Wales, Cardiff, UK Dr. M.B. Kaydan, Faculty of Agriculture, Yuzuncu Yıl University, Van, Turkey Dr. T. Kondo, CORPOICA, Palmira, Valle, Colombia Dr. I.M. Millar, Plant Protection Research Institute, Pretoria, South Africa Dr. D.J. Williams, The Natural History Museum, London Drs Hardy, Hodgson, Kondo, Millar and Williams collaborated with Dr. Gullan on various scale insect taxonomic research projects. Drs Gullan and Kaydan continued their research on the Ferrisia mealybug project (not yet published). Sarah Han is a UC Davis undergraduate student who was trained by Dr. Gullan and worked part-time in Dr. Gullan's lab from 2009-2010. TARGET AUDIENCES: Target audiences for these taxonomic research projects include quarantine officers and pest diagnostic specialists, as well as government, university and museum researchers. PROJECT MODIFICATIONS: Not relevant to this project.

Impacts
Dr. Gullan and her collaborators have produced diverse publications that report on the systematics and biology of Coccoidea (the scale insects), a group that includes numerous plant pests. As part of this Hatch project, we have provided tools to allow users to identify these insects. Accurate identifications and biological information on scale insects are essential to a diverse range of other activities, including pest diagnostics and research ranging from very applied to very basic. Some of the outcomes of the research have been the fruitful redirection of searches for biological control agents of pest scale insects (e.g. the lobate lac insect, Paratachardina pseudolobata), and improved diagnostic capability resulting from the ability to recognize species complexes of pest scale insects (such as that associated with the striped mealybug Ferrisia virgata). Statistics are not available on how often and who accesses our publications, especially as Dr. Gullan made it easy for other people to obtain PDFs (since wherever possible she paid journals for open access and put publications on the departmental home page). Thus it is impossible to quantify the impacts of our systematic findings and products, although their value is very long-term. Certainly the research outcomes will be used for many decades. Resources essential for the research included well-trained taxonomic collaborators (Dr. Gullan trained Drs Hardy, Kaydan and Kondo), well-curated collections (such as the collection of Bohart Museum of Entomology), high quality microscopes (to examine the diagnostic morphology of the insects), and facilities for DNA extraction and sequencing (to provide DNA data to assist with species delimitation and study of insect relationships).

Publications

  • Hodgson, C.J., Millar, I.M. and Gullan, P.J. (2011) Cissococcus Cockerell (Hemiptera: Coccoidea: Coccidae), a unique gall-inducing soft scale genus on Vitaceae from South Africa, with description of a new species. Zootaxa. 2996: 1-32.
  • Hardy, N.B., Beardsley, J.W. and Gullan, P.J. (2011) Uncovering diversity of Australian Eucalyptus-constrained felt scales (Hemiptera: Coccoidea: Eriococcidae). Systematic Entomology 36: 497-528.
  • Kondo, T. and Gullan, P.J. (2011) Taxonomic review of the genus Tachardiella Cockerell (Hemiptera: Kerriidae), with a key to species of lac insects recorded from the New World. Neotropical Entomology 40: 345-367.
  • Williams, D.J., Gullan, P.J., Miller, D.R., Matile-Ferrero, D., and Han, S.I. (2011) A study of the scale insect genera Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea: Putoidae) and Ceroputo Sulc (Pseudococcidae) with a comparison to Phenacoccus Cockerell (Pseudococcidae). Zootaxa 2802: 1-22.


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

Outputs
OUTPUTS: The results of our research are disseminated at conferences (e.g. the XIIth International Symposium of Scale Insect Studies, held in April 2010 in Crete), in seminars (e.g. to the Plant Pest Diagnostic Center at CDFA, Sacramento, CA), and via peer-reviewed publications. The latter are distributed as PDFs to colleagues who are involved in pest diagnostics or research on scale insects (Coccoidea). Dr. Gullan also answered many email inquiries about scale insects and frequently sent PDFs of her publications, as well as performing specialist identifications of scale insects based on her taxonomic research. Since the last report, three important articles (Gruwell et al. 2010; Gullan et al. 2010; Kondo et al. 2010) have been published on scale insects, and 3 conference talks on scale insects presented. PARTICIPANTS: Dr. K. Dittmar, Dept of Biological Sciences, SUNY at Buffalo, New York, USA Dr. M.E. Gruwell, Penn State Erie School of Science, Erie, PA, USA Dr. N.B. Hardy, Queensland Primary Industries & Fisheries. Brisbane, Australia Dr. M.B. Kaydan, Faculty of Agriculture, Yuzuncu Yıl University, Van, Turkey Dr. T. Kondo, CORPOICA, Palmira, Valle, Colombia Dr. R.W. Pemberton, USDA-ARS, Florida, USA TARGET AUDIENCES: Target audiences for the Ferrisia and Paratachardina projects include quarantine officers and pest diagnostic specialists. Our research publications have alerted these people to the need to conduct identifications carefully in light of the cryptic species diversity that we have identified for these pest groups. PROJECT MODIFICATIONS: Not relevant to this project.

Impacts
Dr. Gullan and her collaborators produce diverse publications that report on the systematics and biology of Coccoidea (the scale insects), a group that includes numerous plant pests. As part of my Hatch project, we provide tools to allow users to identify these insects. Accurate identifications and biological information on scale insects are essential to a diverse range of other activities, including pest diagnostics and research ranging from very applied to very basic. I do not have statistics on know how often and who accesses our publications, especially as I make it easy for other people to obtain PDFs (since wherever possible I pay journals for open access and put publications on my departmental home page). Thus it is impossible to quantify the impacts of our systematic findings and products, although their value is very long-term. For example, in 2008 we established new classifications for two very important scale insect pest groups, the mealybugs (family Pseudococcidae) and the Iceryini or cottony-cushion scales (family Monophlebidae), and these products will be used and cited for decades into the future. During 2010 we continued to work on the taxonomy and relationships of the striped mealybug complex, Ferrisia species, and this information will be important to allow accurate identification of these pests, which frequently turn up as quarantine incursions into the USA from central and South America. We also continued to study the pestiferous lobate lac scale with the aim of determining its native range in order to find natural enemies for biological control. Resources essential for my research include well-trained taxonomic collaborators (I trained Drs, Hardy, Kaydan and Kondo), well-curated collections (such as the collections of Bohart Museum of Entomology), high quality microscopes (to examine the diagnostic morphology of the insects), and facilities for DNA extraction and sequencing (to provide DNA data to assist with species delimitation and study of insect relationships).

Publications

  • Kondo, T., Gullan, P.J. and Pemberton, R.W. (2010) A new species of Paratachardina Balachowsky (Hemiptera: Coccoidea: Kerriidae) related to the lobate lac scale, P. pseudolobata Kondo & Gullan. Journal of Asia-Pacific Entomology 14:141-146
  • Gruwell, M.E., Hardy, N.B., Gullan, P.J. and Dittmar, K. (2010) Evolutionary relationships among primary endosymbionts of the mealybug subfamily Phenacoccinae (Hemiptera: Coccoidea: Pseudococcidae). Applied and Environmental Microbiology 76:(22)7521-7525
  • Gullan, P.J., Kaydan, M.B. and Hardy, N.B. (2010) Molecular phylogeny and species recognition in the mealybug genus Ferrisia Fullaway (Hemiptera: Pseudococcidae). Systematic Entomology 35:329-339


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

Outputs
OUTPUTS: The results of our research are disseminated at conferences but largely via peer-reviewed publications and we have several in press papers, including a major work on the mealybug genus Ferrisia in the journal Systematic Entomology. I also answered email enquiries about scale insects and frequently sent PDFs of our publications, as well as performing specialist identifications of scale insects based on our group's taxonomic research. Since the last report, we have published two review articles (Gullan & Martin, 2009; Kondo et al. 2008) on scale insects, and one conference paper on lac scale insects. PARTICIPANTS: The individuals were project collaborators in 2009 (listed alphabetically): Dr. T.M. Collins, Florida International University, Miami, Florida, USA Dr. N.B. Hardy, Queensland Primary Industries & Fisheries. Brisbane, Australia Dr. M.B. Kaydan, Faculty of Agriculture, Yuzuncu Yıl University, Van, Turkey Dr. T. Kondo, CORPOICA, Colombia Dr. R.W. Pemberton, USDA-ARS, Florida, USA Drs. Collins, Kondo and Pemberton were collaborators on lac insect research with the aim of determining the native range of the pestiferous lobate lac scale. Drs. Hardy and Kaydan collaborated on the Ferrisia mealybug project. Research conducted with these collaborators is ongoing. TARGET AUDIENCES: Not relevant to this project. PROJECT MODIFICATIONS: Not relevant to this project.

Impacts
My lab provides diverse publications that report on the systematics and biology of Coccoidea (the scale insects), a group that includes numerous plant pests. As part of our project, we provide tools to allow users to identify these insects. Accurate identifications and biological information on scale insects are essential to a diverse range of other activities, including research ranging from very applied to very basic. I don't know how often and who accesses our publications, especially as I make it easy for other people to obtain PDFs (since wherever possible I pay journals for open access and put publications on my departmental home page). Thus it is impossible to find out the impacts of our systematic findings and products, although their value is very long-term. For example, in 2008 we established new classifications for two very important scale insect pest groups, the mealybugs (family Pseudococcidae) and the Iceryini or cottony-cushion scales (family Monophlebidae), and these products will be used and cited for decades into the future. During 2009 we worked on the taxonomy and relationships of the striped mealybug complex, Ferrisia species, and this information will be important to allow accurate identification of these pests, which frequently turn up as quarantine incursions. We also continued to study the pestiferous lobate lac scale with the aim of determining its native range in order to find natural enemies for biological control.

Publications

  • Gullan, P.J. and Martin, J.H. (2009) Sternorrhyncha (jumping plant-lice, whiteflies, aphids, and scale insects). pp. 957 to 967. In: Resh, V.H. & Carde, R.T. (eds.) Encyclopedia of Insects. 2nd edn. Elsevier, San Diego.
  • Gullan, P.J. and Kondo, T. (2009) The morphology of lac insects (Hemiptera: Coccoidea: Kerriidae). pp. 63 to 70. In: Branco, M., Franco, J.C. & Hodgson, C. (Eds) Proceedings of the XI International Symposium of Scale Insect Studies, Oeiras, Portugal, 24 to 27 September 2007. ISA Press, Lisbon, Portugal.
  • Kondo, T., Gullan, P.J. and Williams, D.J. (2008) The study of scale insects (Hemiptera: Sternorrhyncha: Coccoidea). Revista Corpoica - Ciencia y Tecnologia Agropecuaria 9(2):55 to 61.


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

Outputs
OUTPUTS: Results of my group's research has been disseminated largely via peer-reviewed publications in the following journals: Annals of the Entomological Society of America, Biological Control, Bulletin of Zoological Nomenclature, Invertebrate Systematics, Journal of Natural History, Systematic Entomology, and Zootaxa. A large identification guide to iceryine scale insects (a group with many pests) was published in the online journal Zootaxa, and I paid for open access; Dr. Zhang, the editor of Zootaxa, told me that this publication was the most frequently downloaded work from Zootaxa in the months after its publication in early 2008. In addition, my former students and I have presented research findings at scientific meetings, such as the Entomological Society of America and the International Congress of Entomology. I also answer email enquiries about scale insects and frequently send PDFs of our publications, as well as performing specialist identifications of scale insects based on our group's systematic research. PARTICIPANTS: The following three individuals, who are all former members of the Gullan research group, worked on this project in some way in 2008: Dr. Takumasa Kondo - postdoctoral researcher; now employed in agricultural research in Colombia. Dr. Nathaniel Hardy - PhD student and then short-term postdoctoral researcher; now employed at the Queensland Department of Primary Industries & Fisheries in Brisbane, Australia, as an insect taxonomist. Dr. Corinne Unruh - former PhD student (completed in 2007, but worked on publications after submission of her dissertation); now married and living overseas. Major collaborators in 2008: Dr. L.G. Cook, School of Integrative Biology, University of Queensland, Australia. Mrs. R.C. Henderson, Landcare Research, New Zealand. C.M. Oliveira, Embrapa Cerrados, Brazil. Drs. S. Schroer and R.W. Pemberton, USDA, Florida. Dr. D.J. Williams, The Natural History Museum, London TARGET AUDIENCES: Not relevant to this project. PROJECT MODIFICATIONS: Not relevant to this project.

Impacts
My group provides diverse publications that report on the systematics and biology of Coccoidea (the scale insects), a group that includes numerous plant pests. As part of our project, we provide tools to allow users to identify these insects. It is difficult to know what the many users of our systematic (taxonomic and phylogenetic) research use it to generate, but accurate identifications and biological information on scale insects are essential to a diverse range of other activities, including research ranging from very applied to very basic. We are unable to know how often and who accesses our publications, especially as I make it easy for other people to obtain PDFs (since wherever possible I pay journals for open access and put publications on my departmental home page). Thus it is impossible to find out the impacts of my group's systematic findings and products, although their value is very long-term. For example, we have established new classifications for two very important scale insect pest groups, the mealybugs (family Pseudococcidae) and the Iceryini or cottony-cushion scales (family Monophlebidae), and these products will be used and cited for decades into the future.

Publications

  • Oliveira, C.M., Frizzas, M.R., Fialho, J.F. and Gullan, P.J. 2008. Biology of Protortonia navesi (Hemiptera: Monophlebidae), a new cassava pest in Brazil, with notes on its behavior in the field. Annals of the Entomological Society of America 101:779-785.
  • Schroer, S., Pemberton, R.W., Cook, L.G., Kondo, T. and Gullan, P.J. 2008. The genetic diversity, relationships, and potential for biological control of the lobate lac scale, Paratachardina pseudolobata Kondo & Gullan (Hemiptera: Coccoidea: Kerriidae). Biological Control 46:256-266.
  • Unruh, C.M. and Gullan P. J. 2008. Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing re-interpretation of morphology and a revised generic classification. Systematic Entomology 33:8-50.
  • Unruh, C.M. and Gullan, P.J. 2008. Identification guide to species in the scale insect tribe Iceryini (Coccoidea: Monophlebidae). Zootaxa 1803:1-106.
  • Williams, D.J. and Gullan, P.J. 2008. A revision of the Neotropical scale insect genus Protortonia Townsend (Hemiptera: Coccoidea: Monophlebidae: Llaveiini), Journal of Natural History 42:77-128.
  • Hardy, N.B. and Gullan, P.J. 2008. Case number 3424. Pseudococcini Cockerell, 1905 (insecta, Hemiptera): proposed precedence over Sphaerococcini Cockerell, 1899. Bulletin of Zoological Nomenclature 64(1):1-3.
  • Hardy, N.B., Gullan, P.J. & Hodgson, C.J. 2008. A subfamily-level classification of mealybugs (Hemiptera: Pseudococcidae) based on integrated molecular and morphological data. Systematic Entomology 33:51-71.
  • Hardy, N.B., Gullan, P.J., Henderson, R.C. and Cook, L.G. 2008. Relationships among scale insects (Hemiptera: Coccoidea: Eriococcidae) of southern beech, Nothofagus (Nothofagaceae), with the first descriptions of Australian species of the Nothofagus-feeding genus Madarococcus Hoy. Invertebrate Systematics 22:365-405.


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

Outputs
Major research priorities for the lab in 2007 centered on phylogenetic and taxonomic studies of monophlebid scale insects (family Monophlebidae, tribes Iceryini and Llaveiini), felt scales (family Eriococcidae), mealybugs (family Pseudococcidae), and lac scales (family Kerriidae). I completed a co-authored review article on the phylogeny and classification of the world families of scale insects (Gullan and Cook 2007). In collaboration with my former Ph.D. student Corinne Unruh, I completed a revised, worldwide classification of iceryine scale insects based on molecular and morphological data (to be published in the journal Systematic Entomology and Zootaxa in 2008). We re-organized the 80 named species of iceryines into five redefined genera and provided an identification guide to allow recognition of the genera and all known species. Ph.D. student Nathaniel Hardy and I published two papers describing new species of gall-inducing felt scale from Australia (Hardy and Gullan 2007 c & d). In addition, Mr. Hardy and I have completed a revised classification of the mealybugs, family Pseudococcidae (to be published in the journal Systematic Entomology in 2008). I also worked in collaboration with my NSF-funded postdoctoral researcher, Dr. Takumasa Kondo, on the identity and relationships of the lobate lac scale, which has been introduced to Florida and the Caribbean region. Morphological data allowed taxonomic description of this new species as Paratachardina pseudolobata (Kondo & Gullan 2007b), and molecular data showed it to be more closely related to species of Paratachardina from India than from Australia (manuscript submitted to Biological Control). I completed a taxonomic revision (in press in The Journal of Natural History) of the Neotropical genus Protortonia (Monophlebidae) completed in collaboration with Dr. Douglas Williams (Natural History Museum, London), and published a third collaborative study with Brazilian entomologists on pest scale insects from Brazil (Culik et al. 2007a). During 2007, I made species determinations/identifications for over 30 scale insect samples, mostly for scientists from universities in the USA and government agencies such as CDFA and USDA. The preparation of microscope slide-mounts and identification of each specimen/sample is extremely time consuming and requires both technical skill and expert knowledge. Approximately 200 specimens of scale insects (mostly as microscope slide-mounts) have been added to the Coccoidea Collection of the Bohart Museum of Entomology, UCD.

Impacts
Our lab's taxonomic and phylogenetic work on scale insects has provided new information and identification tools for several groups of important plant pests. In particular, our research on lac scales, iceryines and mealybugs will be useful to applied entomologists who attempt to identify, quarantine or control scale pests. Our work on the lobate lac scale, P. pseudolobata, is especially significant because the recognition of this pest species as distinct from all previously known species has explained the failure of a USDA-ARS biological control program that used natural enemies of an Indian species, P. lobata, which had been assumed to be the same as the pest. A number of species of iceryine scales also are serious plant pests, including a species newly introduced to the southeastern USA, which probably will spread. Our work on iceryine classification allowed the accurate identification of this pest species.

Publications

  • Gullan, P.J. and Cook, L.G. 2007. Phylogeny and higher classification of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea). In Zhang, Z.-Q. & Shear, W.A. (Eds) Linnaeus Tercentenary: Progress in Invertebrate Taxonomy. Zootaxa, 1668: 413-425.
  • Culik, M.P., Martins, D.S., Ventura, J.A., Peronti, A.L.B.G., Gullan, P.J. and Kondo, T. 2007(a). Coccidae, Pseudococcidae, Ortheziidae, and Monophlebidae (Hemiptera: Coccoidea) of Esperito Santo, Brazil. Biota Neotropica 7(3): 1-5.
  • Kondo, T. and Gullan, P.J. 2007(b). Taxonomic review of the lac insect genus Paratachardina Balachowsky (Hemiptera: Coccoidea: Kerriidae), with a revised key to genera of Kerriidae and description of two new species. Zootaxa 1617: 1-41.
  • Hardy, N.B. and Gullan, P.J. 2007(c). A new species of Phacelococcus Miller (Hemiptera: Coccoidea: Eriococcidae) on Bursaria (Pittosporaceae). The Australian Entomologist 34: 85-91.
  • Hardy, N.B. and Gullan, P.J. 2007(d) A new genus and four new species of felt scales (Hemiptera: Coccoidea: Eriococcidae) on Eucalyptus in southeastern Australia. Australian Journal of Entomology 46: 106-120.


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

Outputs
My major research priorities for 2006 centered on phylogenetic and taxonomic studies of felt scales (family Eriococcidae), margarodoid scales (families Margarodidae, Matsucoccidae and Monophlebidae), and lac scale insects (family Kerriidae). I have been working in collaboration with my NSF-funded postdoctoral researcher, Dr. Takumasa Kondo, on the identity and relationships of a new pest lac scale insect found in Florida, the Bahamas and Christmas Island. We have used morphological and molecular data to show that this species is new to science, and thus we have been able to explain the failure of a USDA-ARS biological control program that used natural enemies of an Indian species, Paratachardina lobata, that was believed to be the same as the pest. Currently we have two papers in preparation on the identity, relationships, taxonomy and possible native origin of this new pest and its relatives. I completed a molecular phylogenetic study with my former Masters student, Janie Booth, on the pine-feeding genus Matsucoccus in North America. One significant finding was that three of the recognized pestiferous Matsucoccus species were just different names for the same species (Booth & Gullan 2006b). I have collaborated with my Ph.D. student Corinne Unruh to establish the identity of several ground pearl species in the genus Heteromrgarodes (Margarodidae) found on plant roots in the western USA, and with Dr. Douglas Williams (Natural History Museum, London) on a newly recognized pest Icerya attacking grasses in Micronesia (Williams et al. 2006a) and on a taxonomic revision of the Neotropical genus Protortonia (Monophlebidae). I have conducted felt scale research with a number of collaborators including Dr. Kondo, Dr. Lyn Cook, and my Ph.D. student Nathaniel Hardy. This year I published one co-authored paper (Kondo et al. 2006d) on South American eriococcid species, and another (Gullan et al. 2006) on an African eriococcid genus. Mr. Hardy and I also have been studying felt scales that attack eucalypt trees and other felt scales that feed on southern beech trees (Nothofagus). In addition, I completed a collaborative study with Brazilian entomologists on pest mealybugs (family Pseudococcidae) from Brazil (Culik et al. 2006c). Thus during 2006, my research has covered a diversity of scale insect groups, spanning genera from six families (Eriococcidae, Kerriidae, Margarodidae, Matsucoccidae, Monophlebidae and Pseudococcidae).

In addition, species determinations were made of over 40 scale insect samples, mostly for scientists from UCD, other universities and government agencies such as CDFA and USDA, but also for several pest advisors. Microscope slide preparation and identification of each specimen/sample takes a total of 3-5 hours because of the technical procedures involved in working with very small, soft-bodied insects. Approximately 300 specimens of scale insects (mostly as microscope slide-mounts) have been added to the Coccoidea Collection of the Bohart Museum of Entomology, UCD.

Impacts
Taxonomic and phylogenetic work on scale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. In addition, it provides a solid framework for studying the comparative biology of scale insects, and will assist in determining the native areas of new pest species. My research on lac scale insects (family Kerriidae) is especially important because several of these insects are spreading around the world and causing extensive damage, and even death, to woody native plants. Results of my research will directly benefit applied entomologists who attempt to identify, quarantine or control scale pests, and indirectly will benefit everyone who uses California agricultural and horticultural products.

Publications

  • Booth, J.M. and P.J. Gullan. 2006.(b) Synonymy of three pestiferous Matsucoccus scale insects (Coccoidea: Margarodidae) based on morphological and molecular evidence. Proceedings of the Entomological Society of Washington 108: 749-760.
  • Culik, M.P., D. dos S. Martins, and P.J. Gullan. 2006.(c) First records of two mealybug species (Hemiptera: Pseudococcidae) in Brazil and new potential pests of Carica papaya and Coffea canephora. Journal of Insect Science (23): 1-6.
  • Kondo, T., N. Hardy, L. Cook, and P. Gullan. 2006.(d) Descriptions of two new genera and species of Eriococcidae (Hemiptera: Coccoidea) from southern South America Zootaxa 1349: 19-36.
  • Gullan, P.J., J.H. Giliomee, C.J. Hodgson, and L.G. Cook. 2006. The systematics and biology of the South African gall-inducing scale insect Calycicoccus merwei Brain (Hemiptera: Coccoidea: Eriococcidae). African Entomology 14: 13-33.
  • Williams, D.J., P.J. Gullan. 2006.(a) Englberger, K. and Moore, A. Report on the scale insect Icerya imperatae Rao (Hemiptera: Coccoidea: Margarodidae) seriously infesting grasses in the Republic of Palau. Micronesica 38(2): 267-272.


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

Outputs
My major research priorities for 2005 centered on phylogenetic and taxonomic studies of mealybugs (family Pseudococcidae) and their relationship to endosymbiont bacteria (important to the health of the mealybugs). I concluded a fruitful collaboration with Dr. Paul Baumann (Department of Microbiology, UCD) who retired in August 2005. My collaborator Dr. Doug Downie and I showed that mealybugs have cospeciated with their bacterial endosymbionts (Downie & Gullan, 2005). I have been working with Nate Hardy (formerly my PGR and now my graduate student) to develop another mealybug data set, derived from the morphological features of the adult female and first-instar mealybugs. We will use this independent data set to test the utility of the subfamily and tribal groups suggested by anlaysis of DNA data (published by Downie & Gullan, 2004). In addition, I have collaborated with Brazilian entomologists on two studies on pest mealybugs from Brazil (Culik & Gullan, 2005; Kondo et al., 2005c), and I have co-authored another three taxonomic papers (Kondo & Gullan, 2005; Kondo et al., 2005a,b) with my NSF-funded postdoctoral researcher, Dr. Takumasa Kondo. During 2005, my taxonomic research has covered a diversity of scale insect groups, spanning genera from four families (Coccidae, Eriococcidae, Kerriidae and Pseudococcidae).

A minor research focus for 2005 has been the evolution and biology of gall-inducing scale insects, mainly in the family Eriococidae. I worked with my Australian collaborator, Dr. Lyn Cook (Australian National University) and my graduate student, Nate Hardy, on a paper on the phylogeny, biogeography and taxonomy of Australasian eriococcids, including several galling species, associated with the tree genus Nothofagus.

In addition, species determinations were made of over 45 scale insect samples, mostly for scientists from UCD, other universities and government agencies, but also for several pest advisors. Microscope slide preparation and identification of each specimen/sample takes a total of at least 3 hours because of the technical procedures involved. Approximatley 250 specimens of scale insects (mostly as microscope slide-mounts) have been added to the Coccoidea Collection of the Bohart Museum of Entomology, UCD.

Impacts
Taxonomic and phylogenetic work on mealybugs and other sale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. In addition, it provides a solid framework for studying the comparative biology of scale insects, and will assist in determining the native areas of new pest species. Research on mealybugs (family Pseudococcidae) is especially important because this large family (about 2,000 species) includes many significant crop pests. Results of this research will directly benefit applied entomologists who attempt to identify, quarantine or control scale pests, and indirectly will benefit everyone who uses California agricultural and horticultural products.

Publications

  • Kondo, T., M.L. Williams, and P.J. Gullan. 2005a. Taxonomic review of the genus Xenolecanium Takahashi and description of the new genus Takahashilecanium Kondo (Hemiptera: Coccidae; Coccinae, Paralecaniini). Entomological Science 8:109-120.
  • Kondo, T. and P.J. Gullan. 2005. A new lac insect from Columbia, with revised keys to lac insect genera and to species of Austrotacharidiella Chamberlin (Hemiptera: Coccoidea: Kerriidae). Neotropical Entomology 34:401-407.
  • Kondo, T., M.L. Williams, and P.J. Gullan. 2005b. Redescription of Octolecanium perconvexum (Cockerell), new genus and new combination, with description of new species from Guatemala (Hemiptera: Coccoidea: Coccidae). TIP Revista Especializada en Ciencias Quimico-Biologicas 8:5-11.
  • Kondo, T., P.J. Gullan, J.A. Ventura, and M. Culik. 2005c. Taxonomy and biology of the mealybug genus Plotococcus Miller & Denno (Hemiptera: Pseudococcidae) in Brazil, with descriptions of two new species. Studies on Neotropical Fauna and Environment 40:[in press].


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

Outputs
For 2004, research on the systematics and evolution of scale insects (order Hemiptera, superfamily Coccoidea) has centered on mealybugs (family Pseudococcidae) and felt scales (family Eriococcidae). A significant publication on the phylogeny and classification of mealybugs (Downie & Gullan, 2004) reported the first analysis of mealybug relationships based on molecular data. Representative species, including some important pests, from each of the large mealybug genera were included in the data set. Our phylogenetic hypotheses suggest that genera such as Dysmicoccus, Peudococcus, Nipaecoccus and Paracoccus do not form natural (monophyletic) groups and that the generic concepts and classification will need to be revised. Our study also provided evidence that all mealybug species and genera can be grouped into three monophyletic subfamilies, the Pseudococcinae, Phenacoccinae and Rhizoecinae. The greatest diversity of pest species is found in the Pseudococcinae, whereas most root-feeding pest mealybugs belong to the Rhizoecinae. All previous hypotheses of pseudococcid relationships have been based on intuitive assessment of morphological features of adult female mealybugs, rather than explicit analytical methods, and no accepted subfamily classification for mealybugs existed prior to our work. We are testing and extending our hypotheses by acquiring molecular data from additional genera and by including morphological data. In addition, we have provided survey information for the mealybug species present in ornamental crops in California nurseries (Laflin et al. 2004). Research on felt scales, or eriococcids, has resulted in one publication (Cook and Gullan, 2004) on the diversity and relationships of gall-inducing species, many of which distort the foliage of eucalypts and related plants. A substantial review chapter (60 pages) on gall-inducing scale insects (Gullan et al., 2004) provides a comprehensiveness synthesis of information, including some previously unpublished data, on the biology, taxonomy and diversity of scale insects that control and manipulate the growth of their host plants to induce plant deformations.

In addition, species determinations were made of over 30 scale insect samples, mostly for scientists from UCD, other universities and government agencies, but also for several pest advisors. Microscope slide preparation and identification of each specimen/sample takes a total of at least 3 hours because of the technical procedures involved. Additionally, approximately 300 specimens of scale insects (mostly as microscope slide-mounts) have been added to the Coccoidea Collection of the Bohart Museum of Entomology, UCD.

Impacts
Taxonomic and phylogenetic work on mealybugs and other scale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. In addition, it provides a solid framework for studying the comparative biology of scale insects, and will assist in determining the native areas of new pest species. Research on mealybugs (family Pseudococcidae) is especially important because this large family (about 2,000 species) includes many significant crop pests. Results of this research will directly benefit applied entomologists who attempt to identify, quarantine or control scale pests, and indirectly will benefit everyone who uses Californian agricultural and horticultural products.

Publications

  • Downie, D.A. and Gullan, P.J. (2004) Phylogenetic analysis of mealybugs (Hemiptera: Pseudococcidae) based on DNA sequences from three nuclear genes, and a review of the higher classification. Systematic Entomology 29: 238-259.
  • Laflin, H.M., Gullan, P.J. and Parrella, M.P. (2004) Mealybug species (Pseudococcidae) found on ornamental crops in California nursery production. Proceedings of the Entomological Society of Washington 106: 475-477.
  • Gullan, P.J., Miller, D.R. and Cook. L.G. (2004) Gall-inducing scale insects (Hemiptera: Sternorrhyncha: Coccoidea). Chapter 7, pp. 159-229. In: A. Raman, C.W. Schaefer and T.M. Withers (eds), Biology, Ecology, and Evolution of Gall-Inducing Arthropods. Oxford & IBH Publishing.
  • Cook, L.G. and Gullan, P.J. (2004) Gall induction has evolved multiple times among the eriococcid scale insects (Sternorrhyncha: Coccoidea: Eriococcidae). Biological Journal of the Linnean Society 83: 441-452.


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

Outputs
For 2003, research on the systematics and evolution of scale insects (order Hemiptera, superfamily Coccoidea) has centered on mealybugs (family Pseudococcidae), with one paper on Ferrisia mealybugs published and a major paper on the phylogeny and classification of mealybugs recently accepted for publication (not listed below). This research aimed to facilitate identification, improve classifications and provide biological data on this important pest group. A new pest species of Ferrisia, F. gilli Gullan, was described from pistachio trees in central California and its biology and pest status were reported. All life stages were described and illustrated, and molecular data were used to support the status of this species as new and distinct from other known species. Research on the genus Ferrisia continues because our research has shown that another pest, the striped mealybug Ferrisia virgata, is actually a species complex. An important review chapter on scale insects, aphids, jumping plant lice and whiteflies, collectively called Sternorrhyncha, was published in the Encyclopedia of Insects. This review deals with the appearance, classification, biology and pest status of these agriculturally significant insects. A third paper (Thao et al. 2003) documented the chlamydia bacteria associated with the European elm scale Eriococcus spurius (common in California on elm trees) and other related insects. In addition, species determinations were made of a number of scale insect samples, mostly for scientists from UCD, other universities and government agencies. Microscope slide preparation and identification of each specimen/sample takes a total of about 3 hours because of the technical procedures involved. Additional specimens of scale insects (mostly as microscope slide-mounts) have been added to the Coccoidea Collection of the Bohart Museum of Entomology, UCD.

Impacts
Taxonomic and phylogenetic work on scale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. Research on mealybugs (family Pseudococcidae) is especially important because this large family (about 2,000 species) includes many significant pests of agriculture.

Publications

  • Gullan, P.J., Downie, D.A. and S.A. Steffan (2003) A new pest species of the mealybug genus Ferrisia Fullaway (Hemiptera: Pseudococcidae) from the USA. Annals of the Entomological Society of America 96: 723-737.
  • Thao, M.L., Baumann, L., Hess, J.M., Falk, B.W., Ng, J.C.K., Gullan, P.J. and Baumann, P. (2003) Phylogenetic evidence for two new insect-associated chlamydia of the family Simkaniaceae. Current Microbiology 47: 46-50.
  • Gullan, P.J. and Martin, J.H. (2003) Sternorrhyncha (jumping plant-lice, whiteflies, aphids and scale insects). pp. 1079-1089. In V.H. Resh and R.T. Carde (eds), Encyclopedia of Insects. Academic Press, Amsterdam.


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

Outputs
Active research on the systematics and evolution of scale insects (order Hemiptera, superfamily Coccoidea) continued during 2002, with the aims of facilitating identification, improving classifications and providing biological data on this important pest group. One major paper (Cook et al. 2002) on the relationships and classification of scale insect families was published (listed below), and another significant paper (Thao et al. 2002) was published on mealybugs (family Pseudococcidae) and their associated endosymbiont bacteria. The latter research demonstrated the coevolution of the primary endosymbionts with their mealybug hosts and the multiple infection of the primary with the secondary endosymbionts. The presence of the primary bacteria is essential to the health of the host mealybugs. Taxonomic and phylogenetic research also continued on the mealybug genus Ferrisia. The new pest Ferrisia species that was recognized last year from pistachio trees in southern California is being formally described in all life stages and molecular data have been acquired to support the status of this species as new and distinct from described species. In July 2002, a visit was made to the Coccoidea Collection of the US National Museum in Beltsville, Maryland, to study museum specimens of other Ferrisia species for comparative purposes. Also determinations were made of a number of scale insect samples, mostly for scientists from UCD, other universities and government agencies. Slide preparation and identification of each specimen/sample takes a total of about 3 hours because of the technical procedures involved. Additional photographs of scale insects have been taken for the digital database of images of live Coccoidea.

Impacts
Taxonomic and phylogenetic work on scale insects is essential for the accurate identification of pests and potential pests, and also allows the association of biological data with names. Research on mealybugs (family Pseudococcidae) is especially important because this large family (about 2,000 species) includes many significant pests of agriculture.

Publications

  • Cook, L.G., Gullan, P.J. and Trueman, H.E. 2002. A preliminary phylogeny of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea) based on nuclear small-subunit ribosomal DNA. Molecular Phylogenetics & Evolution 25: 43-52.
  • Thao, M.L., Gullan, P.J. and Baumann, P. 2002. Secondary endosymbionts infect the primary endosymbionts of mealybugs multiple times and coevolve with their hosts. Applied and Environmental Microbiology 68(7): 3190-3197.
  • Gullan, P.J. and Cook, L.G. 2002. Phenacoleachia Cockerell (Hemiptera: Coccoidea: Phenacoleachiidae) re-visited. Bollettino di Zoologia agraria e di Bachicoltura, series II, 33(3): 163-173.
  • Cook, L.G. and Gullan, P.J. 2002. Longevity and reproduction in Apiomorpha Rubsaamen (Hemiptera: Sternorrhyncha: Coccoidea). Bollettino di Zoologia agraria e di Bachicoltura, series II, 33(3): 259-265.


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

Outputs
This project deals with the systematics and evolution of scale insects (order Hemiptera, superfamily Coccoidea), with the aims of facilitating accurate identification, improving classification and providing biological data on scale insects. Progress for the year 2001 consisted of taxonomic and phylogenetic research on scale insects, especially mealybugs, and curatorial and identification work. A project on the molecular phylogeny of scale insect families was completed in collaboration with Dr. L.G. Cook (A.N.U., Australia) and the manuscript was submitted for publication. A large number of fresh mealybug samples were collected for a molecular phylogenetic study being conducted in collaboration with Dr. P.A. Baumann (Microbiology, UCD). Research on the mealybug genus Ferrisia continued with the acquisition of additional samples and the recognition of a new species affected pistachio trees in southern California. A major part of the scale insect slide collection of the Bohart Museum of Entomology has been re-housed in modern slide cabinets in my laboratory and I have updated the names of the species and genera for all families except the Asterolecaniidae (pit scales) and Diaspididae (armored scales). I also hosted the visit of two world authorities on scale insect taxonomy, Drs D.R. Miller and D.J. Williams (visit funded by a NSF curation grant to Dr. L.S. Kimsey), who assisted me with the identification of accession material in the slide collection. Additional photographs of scale insects have been taken for the digital database of images of live Coccoidea. Also I have made determinations of a number of scale insect samples, mostly mealybugs, for scientists from UCD, other universities and government agencies. Slide preparation and identification of each sample takes a total of about 3 hours because of the technical procedures involved. Four papers were published from my research on scale insects.

Impacts
The recognition of a previously undescribed Ferrisia mealybug infesting pistachio trees will aid applied studies on this pest, which previously has been confused with other Ferrisia species. My re-curation and digital imaging work will facilitate future identification of scale insects, including pest species. One of my recent papers, Gullan and Sjarda (2001), presented the first phylogenetic study of the family Margarodidae, which has a controversial classification and includes a number of important pest species including the cottony cushion scale.

Publications

  • Gullan, P.J. and Sjaarda, A.W. 2001. Trans-Tasman Platycoelostoma Morrison (Hemiptera: Coccoidea: Margarodidae) on endemic Cupressaceae, and the phylogenetic history of margarodids. Systematic Entomology. 26: 1-22.
  • Cook, L.G. and Gullan, P.J. 2001. Are the enlarged ducts of Eriococcus (Coccoidea: Eriococcidae) plesiomorphic? Entomologica 33 (1999): 59-66.
  • Gullan, P.J. and Cook, L.G. 2001. Are cochineal insects (Coccoidea, Dactylopius) eriococcids? Entomologica 33 (1999): 91-99.
  • Gullan, P.J. 2001. Why the taxon Homoptera does not exist. Entomologica 33 (1999): 101-104.


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

Outputs
This project commenced in October 2000 and builds on previous research conducted by Dr. Gullan in Australia. Research on the taxonomy and phylogeny of scale insects (order Hemiptera, superfamily Coccoidea) is in progress with the broad aims of facilitating accurate identification, improved classification and access to biological information. Since October 2000, the following results have been achieved or studies are in progress: Taxonomic research on Australian felt scales (family Eriococcidae) and soft scales (family coccidae) is underway in collaboration, respectively, with one of my Ph.D. students (Ms. L. G. Cook, whose thesis is on eriococcids) and an overseas colleague (Dr. C. J. Hodgson, the world's leading expert on soft scales, based at the National Museum of Wales). Several scale insect samples have been collected for future molecular analyses. These samples will provide DNA sequence data that will be used to estimate evolutionary relationships of scale insects. This is an ongoing collaborative project. An illustrated guide was published to allow the identification of the immature (nymphal) specimens of mealybug species found on citrus in Australia. These species are potential pests in countries, such as the USA, where Australian citrus is imported. Specimens of two species of FERRISIA mealybugs were borrowed from R. J. Gill at the Plant Pest Diagnostic Center of the California Department of Food and Agriculture and sorted in preparation for a study of the immature stages and taxonomic variability of these pests. Re-curation of slide-mounted specimens of scale insects from the Bohart Museum of Entomology is in progress. This work involves re-housing thousands of glass slide-mounts in new metal storage cabinets and updating the names of the species and genera for each family of Coccoidea. A well-curated collection is essential to tool aiding the accurate and rapid identification of scale insect specimens. The production of a digital database of photographic images of live scale insects has commenced. These images will aid future identification of Californian scale insects.

Impacts
The part of this research dealing with immature mealybugs will assist US quarantine entomologists to identify mealybug pests imported accidentally with Australian citrus fruit. The re-curation work and the digital images will aid future identification of scale insects.

Publications

  • Cook, L. G. and Gullan, P. J. 2000. First-instar morphology and sexual dimorphism in the gall-inducing scale insect APIOMORPHA Rubsaamen (Hemiptera: Coccoidea: Eriococcidae). Journal of Natural History. 34: 879-894.
  • Gullan, P. J. 2000. Identification of the immature instars of mealybugs (Hemiptera: Pseudococcidae) found on citrus in Australia. Australian Journal of Entomology 39:160-166.