Krell Lab / Entomology Program

About Dr. Frank Krell

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Dr. Frank-T. Krell is the curator of entomology in the Department of Zoology of the Denver Museum of Nature & Science. He received his diploma in biology (1992) and his doctorate (1996) from the University of Tübingen in Germany. His postdoctoral research took him to the Ivory Coast in West Africa where he led a project on scarab beetle biodiversity with the University of Würzburg, Germany.

After a short period with the Zoological Research Institute and Museum Alexander Koenig in Bonn, Germany, he became a research entomologist with The Natural History Museum, London, UK, in 2000. Before joining DMNS in January 2007, he was the head of the beetle division in London. Dr. Krell is Editor-in-Chief of the Denver Museum of Nature & Science Annals and commissioner of the International Commission on Zoological Nomenclature.

Dr Krell's specialty is taxonomy, systematics, and ecology of scarab beetles, especially dung beetles. Recording and explaining the high diversity in this insect group, as well as exploring the effects of anthropogenic habitat changes on dung beetle communities, are two main areas of his research. He is also working on fossil scarab faunas and has done fieldwork on all continents except Antarctica, with the main focus on Africa.

Dr. Krell's scientific publications

Dr. Krell's academic CV

Colorado Scarab Survey

The latest catalogue of beetles occurring in Colorado was published in 1902. Baseline information on the biodiversity of Colorado's beetle fauna is rather insufficient, considering the ongoing population growth in the State, habitat degradation and climate change. In 2007, Dr. Krell and his team at the Denver Museum of Nature & Science have started a statewide collecting program of scarab beetles in Colorado, the Colorado Scarab Survey. Aim of the Survey is to produce a monograph of Colorado scarabs with keys, illustrations, natural history information, and distribution maps for all species, complemented by a regularly updated webpage. The recently published compilation of the scattered literature records on the Colorado scarab fauna will serve as the baseline information for the Survey.

Westcliffe then and now

Change of the beetle fauna of Custer County, Colorado, during the last hundred years

The English naturalist Theodore Dru Alison Cockerell (1866-1948) came to Westcliffe in Custer County in July 1887 to cure a mild tuberculosis. He ended up staying for three years and during this period comprehensively collected and later published the beetle fauna of the area around Westcliffe. This is likely to be the only published comprehensive beetle fauna from Colorado of the 19th century and is a suitable baseline for determining faunal change during the last 120 years. In 2012, we started a collecting program in Custer County to compare the current fauna with Cockerell's records. Despite mining and ranching having certainly affected landscape and associated biota, Custer County's low human population size and consequently limited development and degradation make Custer County a model for the study of climate driven faunal change. We will be collecting in Custer County for at least five more years to gain comparable data of the present fauna.

Human Land Use and Dung Beetles

Almost every spot on earth shows signs of human influence, modification, or destruction. Human land use, be it agriculture, forestry, or urbanization, almost always leads to a reduction of the number of species that can persist in such areas. As a species-rich group of insects, dung beetles are suitable to study effects of land use, because many dung beetle species react early to changes in vegetation cover, microclimate, or provision of food resources. Since 1995 my teams of students and myself have studied the ecology of dung beetles and the influence of human land use on dung beetle communities. In West and East Africa, in Ivory Coast and Kenya, we investigated effects of agricultural practices, pasture, burning, logging, and urbanization. Our main sites in the Ivory Coast were situated in the Parc National de la Comoé, in the village of Bringakro, and in Abidjan. in collaboration with the Centre Suisse de Recherches Scientifiques and the Université d'Abobo-Adjamé. In Kenya, we worked in and around Kakamega Forest, in collaboration with ICIPE and the National Museums of Kenya. Although we are not continuing overseas fieldwork at the moment, we are busy at the Denver Museum of Nature & Science working on the thousands of samples and hundreds of thousands of specimens collected in Africa and will continue publishing the results in the years to come.

Here in Colorado, our Scarab Survey, launched in 2007, will provide baseline data on the current distribution on scarab beetles and compile historic data which will allow us to determine possible effects of human land use in the past and faunal changes in the future.

Guardian Article about my work in West Africa (Dec. 19, 2002)

Fossil Scarabaeoids

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More information will be posted soon.

Biodiversity Exploration / Taxonomy

Work in preparation

Description of a new Hemicyrthus species from New Caledonia (with J. Theuerkauf)

Description of several new Onthophagus species from Ivory Coast.

More information will be posted soon.

Nomenclatural Research

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frank krell at the sherborn meeting in london, oct. 2011

Zoological nomenclature is the scientific naming of animals. It is regulated by a set of rules, the International Code of Zoological Nomenclature. Without a regulated internationally accepted naming system, scientific communication would be seriously hampered. Because of its long history and multicultural character, the same animal species is sometimes known under different names, and the same scientific name is sometimes applied to several species. Nomenclatural research aims at resolving such confusion. The end goal is to have one scientific name as a unique identifier for one taxonomic species.

Nomenclatural research is not scientific research. It is historical research. It is often criticized as a waste of time that could better be spent in discovering and describing biodiversity. Why do I do nomenclatural research? I see it as a welcome compensation for my biological research. I am interested in history, I enjoy old books, and I love to learn about how our predecessors worked and discovered the world. By working on nomenclatural problems, you learn a lot about the history of biology.

Current projects

Clarifying the nomenclature of Protaetia speciosissima/aeruginosa, the largest European flower chafer.

Clarifying the nomenclature of European Trichius species.

Preserving the name Scarabaeus and the current meaning of Scarabaeidae.

ZooBank Development

Every year, over 15,000 new animal species are described. The descriptions are published in a bewildering number of journals and books, in many languages and from all countries. In 2005, the Secretariat and Commissioners of the International Commission on Zoological Nomenclature, together with a group of international zoologists proposed "ZooBank", a central, web-based, freely accessible registry of zoological names and nomenclatural acts. ZooBank will make information about scientific animal names and their nomenclatural status more accessible, and ultimately will provide an authoritative reference of availablilty and spelling of those names. Dr. Frank Krell is the chair of the ZooBank Committee, a group of 30 scientists from 14 countries in charge of developing and implementing ZooBank.

The ZooBank Committee is drafting policies on procedures of registration and determining how ZooBank operates. The procedures how ZooBank policies are developed can be found here or here.

An integral part of drafting ZooBank Policies is the public review. Policies drafted by the ZooBank Committee will be subject to public review of 30 days before the ZooBank Committee will vote on them. The will be posted on several listservers, the ICZN webpage, and on this webpage below.

ZooBank Policies for public review

ZooBank Policy on substitute names (public review from 4 September 2011 to 3 October 2011), Word document, pdf

Apart from listservers (iczn-list, zoobank-list, TAXACOM), the policy can be discussed on the ICZN webpage.

Lepidopterists Conference 2012

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The Denver Museum of Nature & Science has hosted the combined Annual Meeting of the Lepidopterists' Society and the Societas Europaea Entomologica from 23 to 29 July 2012.


24 August: Click here! Group photo of the conference available

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and another photo - and the last photo

List of attendees

The photos are large. "Print Picture" from your browser is likely to print only a part of the picture. Please download the photos and print from Windows Picture and Fax Viewer or another photo software.

26 July:

Additional copies of the Program and Proceedings volume can be purchased for $10 here.

23 July: Click here! Corrections and Additions to Program

Amy Chu designed the conference logo. A great Thank You to her!

6 July: Click here! Program and Proceedings online!

Bison Beetle Project

In ecosystems with large mammals dung beetles are a crucial component of the nutrient cycle by being responsible for breaking up dung and reintegrating dung portions into the soil. The dung beetle fauna of the North American plains is currently rather poor, heavily invaded by European species and doesn't seem capable of dung recycling at a significant scale. When 30-60 million bison had roamed the Great Plains and produced at least 100-200 billion dung pats per year, a more abundant dung beetle fauna can be assumed. Since bison are lacking much of the medical treatment, such as antihelminthic and anti­biotic drugs that cattle get administered, bison dung is likely to be more suitable to maintain an abun­dant dung beetle fauna. If the presence of bison in an area results in a recovering dung beetle fauna, bison herds can have positive implications on the maintenance of soil quality in rangelands.

Our study sites are situated in Elbert County in Colorado, on the West Bijou property of the Plains Conservation Center, and on the adjacent Keen Ranch in Arapahoe County.

We will learn from this eight-year project whether the dung beetle fauna changes after re-introduction of bison in a prairie area. We will also determine which dung beetle species occur in the plains of Colorado. Not a single dung beetle species has been recorded from Elbert County in the literature, and very few from the surrounding prairie. We hope to determine similarity or differences in the dung beetle fauna of (intro­duced) cattle and (native) bison and how the dung beetle fauna changes from spring to fall (phenology). The project has already contributed to a study on the common red-winged Aphodius which turned out to be two species. The fieldwork ended in 2015, but the sorting and identification is still ongoing.

The research was supported in 2010 by Prairie Biotic Research Inc. and featured in the inaugural issue of the journal of the Plains Conservation Center in November 2011.

In June 2017, a Netional Geographic video on the Rainbow Scarab, Phanaeus vindex. featured our work.

Research and Department Associates

Barbara Bartell

Barbara Bartell is a Colorado native. She graduated from Arvada High School in 1963 and attended Colorado State University from 1963 to 1965. Butterflies have always fascinated her. As a child living in Evergreen, Colorado, she collected butterflies and began collecting and studying them again in 1968. As a faculty associate with Colorado State University, she does volunteer work with their butterfly collection. Since 2004 she has volunteered in the butterfly collection of the Denver Museum of Nature & Science where she essentially organized the entire collection. She became Department Associate in 2009.


Bartell, B. 2009. Field study of Boloria freija browni and its habitat in Golden Gate Canyon State Park 2008 (Lepidoptera: Nymphalidae). Denver Museum of Nature & Science Technical Report 2009-4, 17 pp. second part

Bartell, B. 2008. Field study of Boloria selene tollandensis and its habitat in Golden Gate Canyon State Park 2007. Denver Museum of Nature & Science Technical Report 2008-8, 18 pp.

David Bettman

David was Department Associate before he was employed in Frank Krell's CSBR and SCAN grants. After the grant money run out in November 2015, David stayed on as a Department Associate.


Todd Gilligan

Todd M. Gilligan, Ph.D., is a Taxonomist with the USDA-APHIS-PPQ-Science & Technology in Fort Collins, Colorado. He currently leads development of molecular-based assays to detect and identify invasive Lepidoptera for the USDA’s Identification Technology Program. He obtained his Ph.D. in 2012 from Colorado State University, where is he is currently a faculty affiliate in the Department of Bioagricultural Sciences and Pest Management. Todd’s primary specialty is the taxonomy and systematics of moths in the family Tortricidae, and he has authored or coauthored more than thirty publications on this group, including three books. He maintains a website dedicated to tortricids ( and hosts the Online World Catalogue of the Tortricidae along with other taxonomic resources. Todd served as President of the Lepidopterists’ Society from 2013–2015 and currently sits on the Board of Directors for the Wedge Entomological Research Foundation. He became DMNS Research Associate in January 2017.


Wright, D.J. & Gilligan, T.M. 2017. Pelochrista Lederer of the Contiguous United States and Canada (Lepidoptera: Tortricidae: Eucosmini). Alamogordo, New Mexico: Wedge Entomological Research Foundation. 376 pp.

Gilligan, T.M., Huemer, P. & Wiesmair, B. 2016. Different continents, same species? Resolving the taxonomy of some Holarctic Ancylis Hübner (Lepidoptera: Tortricidae). Zootaxa 4178: 347–370.

Gilligan, T.M. & Brown, J.W. 2016. A new genus for Tortrix druana Walsingham, 1914 and a new species from the northern Neotropics (Lepidoptera: Tortricidae: Cochylini: Euliina). Journal of the Lepidopterists’ Society 70: 139–144.

Nagoshi, R.N., Gilligan, T.M. & Brambila, J. 2016. Combining Tpi and CO1 genetic markers to discriminate invasive Helicoverpa armigera from local Helicoverpa zea (Lepidoptera: Noctuidae) populations in the southeastern United States. Journal of Economic Entomology, doi:10.1093/jee/tow177

Ledezma, L.A., Barr, N.B., Epstein, M.E. & Gilligan, T.M. 2016. Diagnosis of Lobesia botrana (Lepidoptera: Tortricidae) using real-time PCR. Journal of Economic Entomology. doi:10.1093/jee/tow111

Tembrock, L.R., McAleer, J.M. & Gilligan, T.M. 2016. A revision of native North American Humulus (Cannabaceae). Journal of the Botanical Research Institute of Texas 10: 11–30.

Gilligan, T.M., Tembrock, L.R., Farris, R.E., Barr, N.B., van der Straten, M.J., van de Vossenberg, B.T.L.H. & Metz-Verschure, E. 2015. A multiplex real-time PCR assay to diagnose and separate Helicoverpa armigera and H. zea (Lepidoptera: Noctuidae) in the New World. PLoS ONE 10(11): e0142912. doi:10.1371/journal.pone.0142912

Wright, D.J. & Gilligan, T.M. 2015. Eucosma Hübner of the Contiguous United States and Canada (Lepidoptera: Tortricidae: Eucosmini). Alamogordo, New Mexico: Wedge Entomological Research Foundation. 256 pp.

Gilligan, T.M. & Passoa, S.C. 2014. LepIntercept, An identification resource for intercepted Lepidoptera larvae. Fort Collins, CO: Identification Technology Program (ITP), USDA-APHIS-PPQ-S&T. [CD-ROM; accessible at].

Gilligan, T.M., Wright, D.J., Munz, J., Yakobson, K. & Simmons, M.P. 2014. Molecular phylogeny and revised classification of Eucosma Hübner and related genera (Lepidoptera: Tortricidae: Eucosmini). Systematic Entomology 39: 49–67.

Gilligan, T.M. & Brown, J.W. 2014. A new name for the western spruce budworm (Lepidoptera: Tortricidae)? The Canadian Entomologist 146: 583–589.

Gilligan, T.M., Baixeras, J., Brown, J.W. & Tuck, K.R. 2014. [email protected]: Online World Catalogue of the Tortricidae (Ver. 3.0),].

Harrison, T.L., Gibson, L.D. & Gilligan, T.M. 2014. A new species of Grapholita Treitschke (Lepidoptera: Tortricidae) from the midwestern USA. Zootaxa 3755: 287–294.

Gilligan, T.M. & Wright, D.J. 2013. Revised World Catalogue of Eucopina, Eucosma, Pelochrista, and Phaneta (Lepidoptera: Tortricidae: Eucosmini). Zootaxa 3746: 301–337.

Gilligan, T.M. & Wright, D.J. 2013. The type species of Eucosma Hübner (Lepidoptera: Tortricidae: Eucosmini). Zootaxa 3630: 489–504.

Epstein, M.E., Weller, S.J., Moon, R.D., Hutchison, W.D., Mattson, W.J., MacRae, I.V. & Gilligan, T.M. 2013. William E. Miller (1930-2013). Journal of the Lepidopterists’ Society 67: 234–239.

Gilligan, T.M. & Epstein, M.E. 2012. TortAI, Tortricids of Agricultural Importance to the United States (Lepidoptera: Tortricidae). Fort Collins, CO: Identification Technology Program (ITP), USDA-APHIS-PPQ-CPHST. [CD-ROM; accessible at].

Valdez Barillas, J.R., Quinn, C.F., Freeman, J.L., Lindblom, S.D., Fakra, S.C., Marcus, M.A., Gilligan, T.M., Alford, E.R., Wangeline, A.L. & Pilon-Smits, E. 2012. Selenium distribution and speciation in hyperaccumulator Astragalus bisulcatus and associated ecological partners. Plant Physiology 159: 1834–1844.

Gilligan, T.M., Brown, J.W. & Hoddle, M.S. 2011. A new avocado pest in Central America (Lepidoptera: Tortricidae) with a key to Lepidoptera larvae threatening avocados in California. Zootaxa 3137: 31–45.

Gilligan, T.M., Epstein, M.E., Passoa, S.C., Powell, J.A., Sage, O.C. & Brown, J.W. 2011. Discovery of Lobesia botrana ([Denis & Schiffermüller]) in California: an invasive species new to North America (Lepidoptera: Tortricidae). Proceedings of the Entomological Society of Washington 113: 14–30.

Barr, N.B., Ledezma, L.A., Farris, R.E., Epstein M.E. & Gilligan, T.M. 2011. A multiplex real-time polymerase chain reaction assay to diagnose Epiphyas postvittana (Lepidoptera: Tortricidae). Journal of Economic Entomology 104: 1706–1719.

Tooman, L.K., Rose, C.J., Carraher, C., Suckling, D.M., Paquette, S.R., Ledezma, L.A., Gilligan, T.M., Epstein, M., Barr, N.B. & Newcomb, R.D. 2011. Patterns of mitochondrial haplotype diversity in the invasive pest Epiphyas postvittana (Lepidoptera: Tortricidae). Journal of Economic Entomology 104: 920–932.

Gilligan, T.M., Epstein, M.E. & Hoffman, K.M. 2011. Discovery of false codling moth, Thaumatotibia leucotreta (Meyrick), in California (Lepidoptera: Tortricidae). Proceedings of the Entomological Society of Washington 113: 426–435.

Brown, J.W., Epstein, M.E., Gilligan, T.M., Passoa, S. & Powell, J.A. 2010. Biology, identification, and history of the light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae: Archipini) in California: an example of the importance of local faunal surveys to document the establishment of exotic insects. American Entomologist 56: 34–43.

Wright, D.J. & Gilligan, T.M. 2010. Two new species of Phaneta Stephens from Western United States (Tortricidae). Journal of the Lepidopterists’ Society 64: 103–107.

Barr, N.B., Ledezma, L.A., Vasquez, J.D., Epstein, M.E., Kerr, P.H., Kinnee, S,, Sage, O. & Gilligan, T.M. 2009. Molecular identification of the light brown apple moth (Lepidoptera: Tortricidae) in California using a polymerase chain reaction assay of the internal transcribed spacer 2 locus. Journal of Economic Entomology 102: 2333–2342.

Gilligan, T.M. & Epstein, M.E. 2009. LBAM ID: Tools for diagnosing light brown apple moth and related western U.S. leafrollers (Tortricidae: Archipini). CD-ROM. Raleigh, NC: Center for Plant Health Science and Technology, USDA-APHIS-PPQ.

Gilligan, T.M., Harrison, T. & Gibson, L.D. 2009. Rediscovery and redescription of Hystrichophora loricana (Grote) (Tortricidae: Olethreutinae). Zootaxa 2117: 65–68.

Gilligan, T.M., Wright, D.J. & Gibson, L.D. 2008. Olethreutine moths of the midwestern United States: an identification guide. Columbus, OH: Ohio Biological Survey. 334 pp.

Gilligan, T.M. & Wenzel, J.W. 2008. Extreme intraspecific variation in Hystrichophora (Lepidoptera: Tortricidae) genitalia — questioning the lock-and-key hypothesis. Annales Zoologici Fennici 45: 465–477.

Gilligan, T.M., Passoa, S. & Harrison. T. 2007. A bibliography of the works of Annette F. Braun. Journal of the Lepidopterists’ Society 61: 113–115.


Charles Harp

Chuck became Research Associate early in 2010.

More info coming soon.


Pogue, M.G., Ouellette, G.D. & Harp, C.E. 2013. A revision of the Schinia volupia (Fitch) species complex (Lepidoptera: Noctuidae: Heliothinae). Zootaxa 3718: 157-191.

Pogue, M.G. & Harp, C.E. 2005. Systematics of Schinia chrysellus (Grote) complex: revised status of Schinia alencis (Harvey) with a description of two new species (Lepidoptera: Noctuidae: Heliothinae). Zootaxa 898: 1-35.

Pogue, M.G. & Harp, C.E. 2004. A review of the Schinia tertia (Grote) species complex (Lepidoptera: Noctuidae: Heliothinae). Zootaxa 473: 1-32.

Pogue, M.G. & Harp, C.E. 2003c. A review of the Schinia regia (Strecker) species complex with a description of a new species (Lepidoptera: Noctuidae: Heliothinae).  Journal of the Lepidopterists' Society 57: 197-203.

Pogue, M.G. & Harp, C.E. 2003b. Systematics of Schinia cupes (Grote) complex: revised status of Schinia crotchii (Hy. Edwards) (Lepidoptera: Noctuidae: Heliothinae). Zootaxa 294: 1-16.

Pogue, M.G. & Harp, C.E. 2003a. Revised status of Schinia unimacula Smith including morphological comparisons with Schinia obliqua Smith (Lepidoptera: Noctuidae: Heliothinae). Zootaxa 226: 1-8.


Paul Opler

Paul became Research Associate in 2011.

More info coming soon.


David Steinmann

David Steinmann is a biospeleologist, or cave biologist, who researches cave life in search of new species. David received a B.A. in Physics and Biology from the University of Colorado at Boulder in 1990 and is a volunteer firefighter in Boulder County. David studies caves throughout Colorado, which has the highest elevation and coldest caves in the continental United States. He began volunteering with the Zoology Department in 2002 and became an Associate in 2009. His research requires climbing, crawling, squeezing, and rappelling into total darkness, where he looks for cave-adapted life forms that are often eyeless and albino. So far, David has discovered over 100 new species and several new genera, with three new species named after him. Colorado's cave invertebrates are being studied to find new and rare species, provide insight into cave fauna evolution and biodiversity, and to help protect cave life.

David also works as a wetlands biologist and rare plant specialist. He has managed over 500 wetland delineation and environmental assessment projects, including local government work, United States Forest Service contracts, ski area expansions, recreational trails, hospital complexes, highways, and public schools. Several new plant species were recently identified while he was out exploring for caves. He is an active member of the National Speleogical Society, Society of Wetland Scientists, and Trout Unlimited. David is often found underground searching for new species, with his caving headlamp providing the illumination. He became a Department Associate in 2009 and a Research Associate in April 2012.

Steamboat Magazine story about Dave's (and others') finds in Sulphur Cave (Holiday 2009)

National Geographic story about Dave's discovery of a new cavernicolous pseudoscorpion in Colorado (Feb. 4, 2011)

9 News story about Dave's discoveries in Colorado caves (Feb. 13, 2011)

Denver Post article about Dave's discovery of extremophile worms in Sulphur Cave (May 10, 2016)


Fend, S.V., Liu, Y., Steinmann, D., Giere, O., Barton, H.A., Luiszer,  F. & Erséus, C. 2016. Limnodrilus sulphurensis n. sp., from a sulfur cave in Colorado, USA, with notes on the morphologically similar L. profundicola (Clitellata, Naididae, Tubificinae). Zootaxa 4066: 451–468.

Giere, O., Wirkner, C.S., Steinmann, D., Fend, S. & Hoeger, U. 2016. Structural and physiological characteristics of Limnodrilus sulphurensis (Oligochaeta, Annelida) thriving in high sulphide conditions. Hydrobiologia, online. DOI 10.1007/s10750-016-3023-0

Giere, O., Steinmann, D., Scholz, S., Wirkner, C.S., Hellmann, N. & Höger, U. 2015. Life in a toxic environment - and exceptional new "sulphur worm" oif the genus Limnodrilus. ISAO 2015, 13th International Symposium on Aquatic Oligochaeta, Brno, Czech Republic, 7–11 September 2015: 25.

Shear, W.A. & Steinmann, D.B. 2013. Cave millipedes of the United States. XIII. A new, troglobiotic species of Austrotyla from Colorado (Diplopoda, Chordeumatida, Conotylidae). Zootaxa 3745: 486-490.

Steinmann, D.B. 2012. Cave Creatures of Glenwood Caverns. Boulder: Professional Wetlands Consulting. 32 pp.

Derkarabetian, S., Steinmann, D.B. & Hedin, M. 2010. Repeated and time-correlated morphological convergence in cave-dwelling harvestmen (Opiliones, Laniatores) from montane western North America. PLoS ONE 5 (5): e10388 (13 pp.)

Steinmann, D.B. 2010. Most Colorado Caves Have Fleas. Rocky Mountain Caving. Winter Edition: 27.

Steinmann, D.B. 2010. Three New Species of Cave Springtail Discovered in Colorado. Rocky Mountain Caving, Spring Edition: 15, 23.

Davis, D.G. & Steinmann, D.B. 2009. Investigating Canyon Cavern. Rocky Mountain Caving, Winter Edition: 24.

Steinmann, D.B. 2009. In the Cave of the Frogs. Rocky Mountain Caving, Autumn Edition: 12-14.

Steinmann, D.B. 2008. Thoughts of a Six-Year Caver. Rocky Mountain Caving, Autumn Edition: 12.

Steinmann, D.B. 2007. New Genus of Cave Collembola Discovered on the White River Plateau. Rocky Mountain Caving, Summer Edition: 28.

Steinmann, D.B. 2006. Salamanders in a Colorado Cave. Rocky Mountain Caving, Summer Edition: 10.

Pomorski, R.J. & Steinmann, D. 2004. Four new genera of the North American Hymenaphorurini (Collembola: Onychiuridae) with a description of new species and key to World genera of the tribe. Insect Systematics and Evolution 35: 15-27.

Steinmann, D.B. 2003. Looking for Life in Glenwood Caverns. Rocky Mountain Caving, Spring Edition: 12-14.

Steinmann, D.B. 2000. Cave Creatures and Cave Ecosystems of Colorado. Rocky Mountain Caving, Spring Edition: 19-20. (Reprinted in the National Speleological Society Speleodigest 2000)

Steinmann, D.B. & Windell, J.T. 1990. Application of an Index of Biotic Integrity to the Boulder Creek Nonpoint Source Pollution Control Phase 1 Demonstration Project. The Journal of the Colorado-Wyoming Academy of Science.


Ken van der Laan

Dr. van der Laan is documenting the distribution of dung beetle species in Colorado life zones. He has been setting up and collecting from dung beetle traps and fecal droppings along Bear Creek, Jefferson County, Red Rocks Denver Mountain Park next to the I-70 bison enclosure, above and below timberline on Mt. Evans, and at the Prairie Conservation Center in Aurora.

He has found that there are dung beetles that are specific to particular life zones, while other species are more general in distribution. Dung beetle species evidence seasonality in their presence. The predominant species in all zones are native, however, there is a puzzling pattern in the presence of introduced European species. "Non-dung" beetles have also been collected adjacent to and in the traps. Those species distributions through space and time are also being analyzed.

Dr. van der Laan has been a native of Colorado since 1939. The trapping system is labor intensive and requires lots of bending over. Thus, the aid of able bodied volunteers has and will be most welcome. Ken occasionally springs for lunch for "his" volunteers.

Ken went to South High School and University of Denver (1961). He completed his doctorate in zoology at the University of California-Berkeley (1971). His publications – now ancient – concern a terrestrial snail species from coastal California and the environmental impact on invertebrates in an Illinois riparian ecosystem. He did extensive work on southern and central California coast hermit crabs and used to know the names of every West Coast gastropod worthy of becoming shelter for the crabs. Yes, even the itty bitty ones.

Dr. van der Laan retired as an educator after 35+ years, teaching every age level except elementary school. He taught biology, invertebrate zoology, animal ecology, marine biology, biostatistics, and human anatomy and physiology. He most recently taught at Golden High School. Ken became Research Associate in 2011.


Kuris, A.M., van der Laan, K. & Stouder, D.J. 1980. Graphical models for intraspecific and interspecific dominance among hermit crabs. Bulletin of the Ecological Society of America 61:136. [Abstract]

Stouder, D.J., van der Laan, K. & Kuris, A.M. 1980. Preferences for shell size and shape by three species of intertidal hermit crabs from California. Bulletin of the Ecological Society of America 61:136. [Abstract]

van der Laan, K., Kuris, A.M. & Stouder, D.J. 1980. Hermit crabs: An experimental field study of interspecific competition and shell availability. Bulletin of the Ecological Society of America 61:136. [Abstract]

van der Laan, K.L. 1980. Terrestrial pulmonate reproduction: seasonal and annual variation and environmental factors in Helminthoglypta arrosa (Binney) (Pulmonata:Helicidae). The Veliger 23 (1): 48-54.

Uetz, G.W., van der Laan, K.L., Summers, G.F., Gibson, P.A.K. & Getz, L.L. 1979. The effects of flooding on flood plain arthropod distribution, abundance and community structure. American Midland Naturalist 101 (2): 286-299.

van der Laan, K.L. & Kuris, A. 1979. Resource movement and interspecific competition in field populations of intertidal hermit crabs. Bulletin of the Ecological Society of America 60: 118. [Abstract]

Uetz, G.W., van der Laan, K.L., Summers, G.F., Gibson, P.K. & Getz, L.L. 1975. The effects of flooding on flood-plain invertebrate distribution, abundance, and community structure. Pp. 289-318 in: Bell, D.T. & Johnson, F.L.: The Upper Sangamon River Basin: Final Report for the Springer-Sangamon Environmental Research Program. Department of Forestry and the Illinois Agricultural Experiment Station, University of Illinois.

van der Laan, K.L. 1975. Feeding preferences in the land snail, Helminthoglypta arrosa (Binney) (Pulmonata:Helicidae). The Veliger 17 (4): 354-359.

van der Laan, K.L. 1975. Aestivation in the land snail, Helminthoglypta arrosa (Binney) (Pulmonata:Helicidae). The Veliger 17 (4): 360-368.

van der Laan, K.L. 1973. The role of the flooding regime on the invertebrate fauna of the Sangamon River flood plains. Preliminary results and conclusions. Pp. 104-125 in: Bell, D.T.: Annual Report - FY73 for the Springer-Sangamon Environmental Research Program. Department of Forestry, University of Illinois, Urbana.


Michael J. Weissmann

Michael J. Weissmann, Ph.D., is a co-founder and former curator of the Butterfly Pavilion and Insect Center in Westminster, Colorado. He has consulted with a variety of similar facilities since then. He was the initial Executive Director of the International Association of Butterfly Exhibitors and Suppliers, a butterfly house industry 501(c)(6) trade association, and currently serves on several IABES committees. Working with former Kallima Consultants partner, Richard Cowan, they created London Pupae Supplies of Los Angeles (now LPS LLC), which has become the largest importer of live tropical butterfly pupae in North America, now operated exclusively by Mr. Cowan. Scientifically, Dr. Weissmann has published a variety of technical articles about insect taxonomy, biology, and behavior, and has co-authored popular guides to insects of regional parks and monuments. Current entomological research is focused on Colorado mosquitoes and the West Nile Virus they vector, seasonally working as Surveillance Manager for Colorado Mosquito Control, Inc. He has also been presenting insect-related educational programs for more than two decades to audiences of all ages, including the current award-winning incarnation, "Dr. Mike's Bag of Bugs." He earned B.A. and M.A. degrees in biology at the University of Colorado at Boulder and is currently an adjunct curator at the University of Colorado Museum. He received his doctorate in entomology from Colorado State University and continues to serve as an affiliate faculty member.


Weissmann, M.J. 2009. Arthropod urban legends and myths. 2009 Invertebrates in Education and Conservation Conference Proceedings. Tucson, AZ: Sonoran Arthropod Studies Institute. Pp. 153-161.

Kent, R., Juliusson, L., M. Weissmann, Evans, S. & Komar, N. 2009. Seasonal blood-feeding behavior of Culex tarsalis (Diptera: Culicidae) in Weld County, Colorado, 2007. Journal of Medical Entomology 46 (2): 380-390.

Bennett, J.K., Hickman, A.D., Kline, M.A., McGinnis, M.W. & Weissmann, M.J. 2005.  New state record for the Asian Tiger Mosquito, Aedes albopictus (Skusse). Journal of the American Mosquito Control Association 21 (4): 341-343.

Weissmann, M.J. 2004. Butterfly farming and habitat conservation: Myths and realities. 2004 Invertebrates in Captivity Conference Proceedings. Tucson, AZ: Sonoran Arthropod Studies Institute. Pp. 1-5.

Doyle, M., McGinnis, M. & Weissmann, M.J. 2004. West Nile Virus on Colorado's front range: The impact of Culex tarsalis. Abstract and Paper for the American Mosquito Control Association Annual Meeting, Savanah, GA.

Penn, L.D. & Weissmann, M.J. 2002. Light at the end of the tunnel: Significant progress in rearing techniques for North American fireflies Photuris sp. and Photinus sp. (Coleoptera: Lampyridae). 2002 Invertebrates in Captivity Conference Proceedings. Tucson, AZ: Sonoran Arthropod Studies Institute. Pp. 1-5.

Weissmann, M.J. & Kondratieff, B.C. 1999. An inventory of arthropod fauna at Great Sand Dunes National Monument, Colorado. Pp. 69-80 in: Byers, G.W., Hagen, R.H. & Brooks, R.W. (eds.): Entomological Contributions in Memory of Byron A. Alexander. University of Kansas Natural History Museum Special Publication 24.Lawrence, KS: Natural History Museum, The University of Kansas.

Weissmann, M.J. 1999. Arthropod life at Great Sand Dunes National Monument. Pp. 238-242 in: Schenk C.J. (ed.): Hydrologic, Geologic, and Biologic Research at Great Sand Dunes National Monument, Colorado: Proceedings National Park Service Research Symposium No. 1. National Park Service, USGS.

Weissmann, M.J. 1999. Invertebrate exhibits for the 21st century: Crawling out from behind the glass. 1999 Invertebrates in Captivity Conference Proceedings. Tucson, AZ: Sonoran Arthropod Studies Institute. Pp. 148-152.

Weissmann, M.J. & Kondratieff, B.C. 1999. Two new species of Amblyderus (Coleoptera: Anthicidae) from Great Sand Dunes National Monument, Colorado. Entomological News 110 (3): 137-143.

Weissmann, M.J. 1999. The great pollinators. Mountain, Plain and Garden: The Magazine of Denver Botanic Gardens 56 (1): 4-13.

Weissmann, M.J. 1998. The complex journey of an exotic butterfly chrysalis from the farm to the exhibit. Proceedings of the 1998 Western Regional Conference of the American Zoo and Aquarium Association, Monterey, CA.

Weissmann, M.J. 1997. Natural history of the giant sand treader camel cricket, Daihinibaenetes giganteus Tinkham (Orthoptera: Rhaphidophoridae). Journal of Orthoptera Research 6: 33-48.

Weissmann, M.J. & Brinkmann. G. 1997. Tropical horticultural collections at the Butterfly Pavilion and Insect Center. Public Garden (Journal of the American Association of Botanical Gardens and Arboreta) 12 (4): 22-23.

Weissmann, M.J. 1997. Exhibiting live butterflies. HMS Beagle Log 2(2): 26.

Weissmann, M.J. 1996. "T-T-T-T-T-Touch ME:" Hands-on education at the Insect Center touch cart. 1996 Invertebrates in Captivity Conference Proceedings. Tucson, AZ: Sonoran Arthropod Studies Institute, pp. 107-109.

Kondratieff, B.C., Weissmann, M.J. & Takumi, R.L. 1996. Kingdom of the Small:Common insects and other arthropods of Dinosaur National Monument. Vernal, UT: Dinosaur Nature Association. 27 pp.

Kinsey, F.M. & Weissmann, M.J. 1995. Gardening for butterflies. Connections 5(5): 21-23.

Weissmann, M.J. 1995. The Butterfly Pavilion and Insect Center: The dream becomes reality. 1995 Invertebrates in Captivity Conference Proceedings. Tuczon, AZ: Sonoran Arthropod Studies Institute,  pp. 8-11.

Weissmann, M.J., Lederhouse, R.C. & Elia, F.C. 1995. Butterfly gardening and butterfly houses and their influence on conservation in North America. Chapter 32. Pp. 393-400 in: Scriber, J.M., Tsubaki, Y. & Lederhouse, R.C. (eds.): Swallowtail Butterflies: Their Ecology & Evolutionary Biology. Gainesville, FL: Scientific Publishers.

Weissmann, M.J. 1995. Natural history of the giant sand treader camel cricket, Daihinibaenetes giganteus Tinkham (Orthoptera: Rhaphidophoridae), at Great Sand Dunes National Monument, Colorado. Ph.D. Dissertation, Department of Entomology, Colorado State University. 132 pp.

Weissmann, M.J., Clement, L.P. & Kondratieff, B.C. 1993. Insects and other arthropods of Great Sand Dunes National Monument. Tucson, AZ: Southwest Parks and Monuments Association. 23 pp.

Weissmann, M.J. &. Leatherman, D.A. 1992. Range extension of the northern true katydid, Pterophylla camellifolia (F.) (Orthoptera: Tettigoniidae: Pseudophyllinae) into eastern Colorado. Journal of the Kansas Entomological Society 65 (4): 448-449.

Kondratieff, B.C., Weissmann, M.J. & Walter, D.E. 1990. A guide to the common insects and other arthropods of Colorado National Monument. Fruita, CO: Colorado National Monument Association publication. 23 pp. [recipient of 1991-92 Award of Honorable Mention in Publication in the National Park Service Cooperation Association Publications Competition]

Lanham, U.N. & Weissmann, M.J. 1988. Scaphandrena and Elandrena (Hymenoptera: Andrenidae). Pan-Pacific Entomologist 64 (2): 183-184.

Weissmann, M.J. 1988. Introduction to the University Museum. Chapter IX. Pp.73-76 in: Bushnell, J., Davis, J. & Grant, M.: Biological Science in the Laboratory. Part II, 3rd edition. Kendall/Hunt Publ., Dubuque.

Weissmann, M.J. 1986. Biology of the water scorpion, Ranatra fusca Palisot de Beauvois in Colorado, with notes on mosquito control capability. M.A. Thesis, University of Colorado. 74 pp.

The DMNS Entomology Program preserves and develops a world collection of over 1 million insects, mainly Coleoptera and Lepidoptera, with a geographical focus on the western United States and Africa. The entomology collection also contains the terrestrial (incl. freshwater) non-arachnid non-mollusc invertebrates.

Frank Krell's scientific publications

Frank Krell's academic CV

ORCID 0000-0001-6065-0812

Researcher ID A-8633-2009

Zoology Collections webpage

Database of the insect collection (work in progress)

Citizen Science project: Japanese Beetle Survey



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