Spivak Lab Publications

Studies on Social Immunity

Goblirsch M, Warner JF, Sommerfeldt BA, Spivak M. 2020. Social fever or general immune response?  Revisiting an example of social immunity in honey bees. Insects 11: 528  doi:10.3390/insects11080528

Spivak M, Goblirsch, M, Simone-Finstrom M.  2019 Social-medication in bees: The line between individual and social regulation. Curr. Opinion Insect Science 33:49-55. https://doi.org/10.1016/j.cois.2019.02.009   (If you cannot access this, email for a copy) 

Evans J.D., Spivak M. 2010. Socialized Medicine: Individual and communal disease barriers in honey bees. J. Invert. Pathol.103:S62-S72.

Wilson-Rich N, Spivak M, Fefferman NH, Starks, PT. 2009. Genetic, individual, and group facilitation of disease resistance in insect societies. Annu. Rev. Entomol. 54: 405-23.


Studies on Propolis

Shanahan M,  Simone-Finstrom M, Tokarz P , Rinkevich F , Read Q, Spivak M. Thinking inside the box: building beehives that stimulate propolis collection and support honey bee health. PlosOne e0291744 https://doi.org/10.1371/journal.pone.0291744

Shanahan M, Spivak M. 2021. Resin use by stingless bees:  A review. Insects.  12, 719. https://doi.org/10.3390/insects12080719

Dalenberg H, Maes P, Mott B, Anderson KE, Spivak M. 2020.  Propolis envelope promotes beneficial bacteria in the honey bee (Apis mellifera) mouthpart microbiome.  Insects 11, 453. doi:10.3390/insects1107/0453

Saelao P, Borba RS, Ricigliano V, Spivak M, Simone-Finstrom M. 2020. Honeybee microbiome is stabilized in the presence of propolis.  Biology Letters 16: 202003. doi.org/10.1098/rsbl.2020.0003

Spivak M, Simone-Finstrom M. 2019. Propolis. In: Starr C. (eds) Encyclopedia of Social Insects. Springer, Cham. https://doi.org/10.1007/978-3-319-90306-4_134-1

Borba RS, Spivak M. 2017.  Propolis envelope in Apis mellifera colonies supports honey bees against the pathogen, Paenibacillus larvae.  Sci. Reports 7:11429.  DOI:10.1038/s41598-017-11689-w

Simone-Finstrom M, Borba RS, Wilson M, Spivak M. 2017 Propolis counteracts some threats to honey bee health. Review.  Insects 8: 46:  DOI:10.3390/insects8020046

Wilson MB, Pawlus AD, Brinkman D, Gardner G, Hegeman AD, Spivak M, Cohen JD.  2017. 3-acyl dihydroflavonols from poplar resins collected by honey bees are active against the bee pathogens Paenibacillus larvae and Ascosphaera apis. Phytochemistry  138: 83-92.

Bankova V, Bertelli D, Borba R, Conti BJ, da Silva Cunha IB, Danert C,  Eberlin MN , Falcao SI, Isla MI, Moreno MIN, Papotti G,  Popova M, Santiago KB, Salas, A, Sawaya ACHF,  Schwab NV, Sforcin JM, Simone-Finstrom M, Spivak M, Trusheva B, Vilas-Boas M, Wilson M. Zampini C.  2016. Standard methods for Apis mellifera propolis research.   J. Apicul. Res. DOI: 10.1080/00218839.2016.1222661

Borba RS Klyczek KK, Mogen KL, Spivak M. 2015  Seasonal benefits of a natural propolis envelope to honey bee immunity and colony healthJ. Exp. Biol. doi:10.1242/jeb.127324

Wilson MB, Brinkman D, Spivak M, Gardner G, Cohen JD. 2014.  Regional variation in composition and antimicrobial activity of US propolis against Paenibacillus larvae and Ascosphaera apisJ. Invertebr. Pathol.  234: 44-50.

Wilson MB, Spivak M, Hegeman AD, Rendahl A, Cohen JD. 2013. Metabolomics reveals the origins of antimicrobial resins collected by honey bees. PLoSOne 8(10): e77512. doi:10.1371 /journal.pone.0077512

Simone-Finstrom, MD, Spivak M. 2012. Increased resin collection after parasite challenge: A case of self-medication in honey bees? PLoSOne 7(3): e34601. doi:10.1371/journal.pone.0034601

Simone M, Gardner J, Spivak M. 2010. Tactile learning in resin foraging honeybees. Behav. Ecol. Sociobiol. 64: 1609-1617

Simone M, Spivak M. 2010. Propolis and bee health: the natural history and significance of resin use by honey bees. Special Issue on Bee Health: Apidologie. 41: 295–311

Simone M, Evans J, Spivak M. 2009. Resin collection and social immunity. Evolution 63: 3016-30222

Gekker, G., Hu, S., Spivak, M., Lokensgard, J.R., Peterson, P.K. 2005. Anti-HIV-1 activity of propolis in CD4+ lymphocyte and microglial cell cultures. J. Ethnopharmacology 102: 158-163.

Bastos, EMA, Simone, M, Jorge, D , Soares, AEE, Spivak, M. 2007. In vitro study of the antimicrobial activity of Brazilian and Minnesota, USA propolis against Paenibacillus larvae. J. Invertebr. Pathol. online

Landscapes and Bees

Evans E, Ascher J, Cariveau DP, Spivak M.  2023.  A century of change for prairie bees and their floral associationsPrairie Naturalist 1:78-102 

Carr-Markell MK, Spivak M. 2020. External validation of the new calibration for mapping honey bee waggle dances.  Animal Behaviour. https://doi.org/10.1016/j.anbehav.2020.12.006

Spivak M, Cariveau DP. 2020. Flowers as parasite transmission hubs. Nat Ecol Evol. https://doi.org/10.1038/s41559-020-1200-z

Carr-Markell MK, Demler CM, Cuvillon MJ, Schurch R, Spivak, M. 2020. Do honey bee (Apis mellifera) foragers recruit their nestmates to native forbs in reconstructed prairie habitats? PlosOne. 15(2): e0228169. https://doi.org/10.1371/journal.pone.0228169

Lane I, Watkins E, Spivak M. 2019. Turfgrass species affect the establishment and bloom of Kura clover (Trifolium ambiguum) in lawns. Hortscience 54(5):824–828. https://doi.org/10.21273/HORTSCI13779-18 (If you cannot access this email for a copy)

Ramer H, Nelson K, Spivak M, Watkins E, Wolfin J, Pulscher M.  2019. Exploring park visitor perceptions of 'flowering bee lawns' in neighborhood parks in Minneapolis, MN, US.  Landscape and Urban Planning.189: 117-128. https://doi.org/10.1016/j.landurbplan.2019.04.015  (If you cannot access this email for a copy)

Ostiguy N, Drummond FA, Aronstein K, Eitzer B, Ellis JD, Spivak M, Sheppard WS.  2019. Pesticide exposure to honey bees in a four-year nationwide study.  Insects, 10, 13, doi:10.3390/insects10010013.

Evans E, Smart M, Cariveau D, Spivak M. 2018. Wild, native bees and managed honey bees benefit from similar agricultural land uses. Agriculture, Ecosystems and Environment. 268: 162-170.

Smart MD, Cornman RS, Iwanowicz DD, McDermott-Kubeczko M, Pettis JS, Spivak MS, Otto CRV.  2017 A comparison of honey bee-collected pollen from working agricultural lands using light microscopy and ITS metabarcoding. J. Environ. Entomol. dx.doi.org/10.1093/ee/nvw159

Wu-Smart J, Spivak M. 2017. Effects of neonicotinoid imidacloprid exposure on bumble bee (Bombus impatiens Cresson) queen survival and nest initiation. J. Environ. Entomol. **:1-8. doi:10.1093/ee/nvx175

Wu-Smart J, Spivak M. 2016.  Sub-lethal effects of dietary neonicotinoid insecticide exposure on honey bee queen fecundity and colony development. Nature Sci. Rep. 6, 32108; doi: 10.1038/srep32108

Smart M.  Pettis JF, Euliss N, Spivak M. 2016. Land use in the Northern Great Plains region of the U.S. influences the survival and productivity of honey bee colonies. Agriculture, Ecosystems, and Environment 230: 139-149.  doi.org/10.1016/j.agee.2016.05.030  

Smart M, Pettis JF, Rice RD, Browning Z, Spivak M.  2016. Linking measures of honey bee colony and individual bee health to survival among commercial apiaries surrounded by varying agricultural land use.  PLOS One 11(3); doi:10.1371/journal.pone.0152685

Bee Health and Survivorship

Nikulin SL, Hesket PJ, Mckeown DA, Spivak, M, Schroeder D. 2024. A semi-automated and high-throughput approach for the detection of honey bee viruses in bee samples. PLOS One 19(3): e0297623 https://doi.org/10.1371/journal.pone.0297623

Kulhanek K, Steinhauer N, Wilkes J, Wilson M, Spivak M, Sagili RR, et al. (2021) Survey-derived best management practices for backyard beekeepers improve colony health and reduce mortality. PLoS ONE 16(1): e0245490. https://doi.org/10.1371/journal.pone.0245490

Lee, KV, Goblirsch M, McDermott E, Tarpy DR, Spivak M.  2019. Is the brood patterns within a honey bee colony a reliable indicator of queen quality? Insects 10, 12; doi:10.3390/insects10010012

Spivak, M, Browning Z, Goblirsch M, Lee K, Otto C, Smart M, Wu-Smart J.  In review. 2017  Why does bee health matter?  The science surrounding honey bee health concerns and what we can do about it.  Council for Agricultural Science and Technology (CAST) Commentary, QTA2017. CAST, Ames, Iowa.

Lee K, Steinhauer N, Travis DA, Meixner MD, Deen J, vanEngelsdorp D.  2015.  Honey bee surveillance: a tool for understanding and improving honey bee healthCurrent Opinion in Insect Science 10:37-44

Lee KV, Steinhauer N, Rennich K, Wilson ME, Tarpy DR, Caron DM, Rose R, Delaplane KS, Baylis, K, Lenkerich EJ, Pettis J, Skinner JA, Wilkes JR, Sagili R, vanEngelsdorp D. 2015.  A national survey of managed honey bee 2013-2014 annual colony losses in the USA. Apidologie. 46:292-305

Chen YP,  Pettis JS, Corona M, Chen WP, Li CJ, Spivak M et al.   2014.  Israeli Acute Paralysis Virus: Epidemiology, Pathogenesis and Implications for Honey Bee HealthPlos Pathogens DOI: 10.1371/journal.ppat.1004261

Gobrlirsch M, Huang Z, Spivak M. In Press 2013. Physiological and behavioral changes in honey bees (Apis mellifera L.) induced by Nosema ceranae infection. PLoSOne

Lee KV, Moon RD, Burkness EC, Hutchison WD, Spivak M. 2010. Practical sampling plans for Varroa destructor (Acari: Varroidae) in Apis mellifera (Hymenoptera: Apidae) colonies and apiaries. J. Econ. Entomol. 103:1039–1050. doi:10.1603/EC10037.

Wild and Native Bees

Evans E, Smart M, Cariveau D, Spivak M.  2018. Wild, native bees and managed honey bees benefit from similar agricultural land uses. Agriculture, Ecosystems and Environment. 268: 162-170.

Gardner J, Spivak M. 2014.  A survey and historical comparison of the Megachilidae (Hymenoptera: Apoidea) of Itasca State Park, MinnesotaAnn. Ent. Soc. Amer. 107(5): 983-993.

Evans, E., Spivak, M. 2006. Effects of honey bee (Hymenoptera: Apidae) and bumblebee (Hymenoptera: Apidae) presence on cranberry (Ericales: Ericaceae) pollination. J. Econ. Entomol. 99: 614-620.

Evans, E., I. Burns, and M. Spivak. 2007. Befriending Bumble Bees: a Practical Guide to Raising Local Bumble Bees. University of MN Extension, MN.

Honey Bee Cell Line

Goblirsch MJ, Spivak MS, Kurtti TJ. 2013.  A cell line resource derived from Honey bee (Apis mellifera) embryonic tissue.  PLoS One 8(7) e69831.

Hygienic Behavior as a Mechanism of Resistance to Diseases and Mites

Wagoner K, Spivak M, Hefetz A, Reams R, Rueppell O. 2019.  Stock-specific chemical brood signals are induced by Varroa and Deformed Wing Virus, and elicit hygienic response in the honey bee. Sci Reports. 9:8753. doi.org/10.1038/s41598-019-45008-2

Wagoner K, Spivak M, Rueppell O. 2018. Brood affects hygienic behavior in the honey bee (Hymenoptera: Apidae).  J. Econ. Entomol.  toy266, https://doi.org/10.1093/jee/toy266

Scannapieco A, Lanzavecchia SB, Parreno MA, Liendo MC, Cladera JL, Spivak M, Palacio MA. 2016. Individual precocity, temporal persistence and task-specialization of hygienic bees from selected colonies of Apis mellifera.  J. Apicultural Science 60: 49-60 DOI: 10.1515/JAS-2016-0006

Palacio MA, Rodriguez E, Goncalves L, Bedascarrasbure E, Spivak M. 2010. Hygienic behaviors of honey bees in response to brood experimentally pin-killed or infected with Ascosphaera apis. Apidologie 41: 602-612

Swanson J, Torto B, Kells S, Mesce K, Tumlinson J, Spivak M. 2009. Volatile compounds from chalkbrood Ascosphaera apis infected larvae elict honey bee (Apis mellifera) hygienic behavior. J. Chem. Ecol. 35: 11088-1116

Wilson-Rich N, Spivak M, Fefferman NH, Starks, PT. 2009 Genetic, individual, and group facilitation of disease resistance in insect societies. Annu. Rev. Entomol. 54: 405-23.

Ibrahim, A., Reuter, GS, Spivak, M. 2007. Field trial of honey bee colonies bred for mechanisms of resistance against Varroa destructor. Apidologie 38: 67-76.

Ibrahim, A., Spivak, M. 2006. The relationship between hygienic behavior and suppression of mite reproduction as honey bee mechanisms of resistance to Varroa destructor. Apidologie. 37: 31-40.

Mondragon, L., Spivak, M., Vandame, R. 2005. A multifactorial study of the resistance of Africanized and hybrid honey bees Apis mellifera to the mite Varroa destructor over one year in Mexico. Apidologie. 36: 345-358.

Flores, JM, Spivak, M., Guiterrez, I. 2005. Spores of Ascosphaera apis contained in wax foundation can pass on chalkbrood in honey bees. Veterinary Microbiology. 108: 141-144.

Spivak, M., Reuter, G.S. 2005 A Sustainable Approach to Controlling Honey Bee Diseases and Varroa Mites. Sustainable Agriculture Research and Education (SARE) Fact Sheet.

Spivak, M., Reuter, G.S. 2001 Resistance to American foulbrood disease by honey bee colonies, Apis mellifera, bred for hygienic behavior. Apidologie 32: 555-565.

Spivak, M., Reuter, G. S. 2001. Varroa jacobsoni infestation in untreated honey bee (Hymenoptera: Apidae) colonies selected for hygienic behavior. J. Econ. Entomol 94(1): 326-331.

Spivak, M., Gilliam, M. 1998. Hygienic behaviour of honey bees and its application for control of brood diseases and varroa mites. Part I: Hygienic behaviour and resistance to American foulbrood. Bee World 79:124-134. Part II: Studies on hygienic behaviour since the Rothenbuhler era. Bee World 79: 165-182.

Spivak, M., Downey , D. 1998. Field assays for hygienic behavior in honey bees (Apidae: Hymenoptera). J. Econ. Entomol. 91(1): 64-70.

Spivak, M., Reuter, G.S. 1998. Performance of hygienic honey bee colonies in a commercial apiary. Apidologie. 29: 291-302.

Spivak, M. 1996. Honey bee hygienic behavior and defense against Varroa jacobsoni. Apidologie 27: 245-260.

Spivak M, Gilliam M. 1993. Facultative expression of hygienic behaviour of honey bees in relation to disease resistance. J. Apicultural Res. 32(3/4): 147-157.

Neuroethology of Honey Bee Hygenic Behavior

Wilson-Rich N, Spivak M, Fefferman NH, Starks, PT. 2009 Genetic, individual, and group facilitation of disease resistance in insect societies. Annu. Rev. Entomol. 54: 405-23.

Arathi, H.S., Ho, G., Spivak M. 2006. Inefficient task partitioning among nonhgienic honeybees, Apis mellifera l., and implications for disease transmission. Animal Behaviour 72: 431-438.

Goode, K, Huber, Z, Mesce, KA, Spivak, M. 2005. The relationships of honey bee (Apis mellifera) behaviors in the context of octopamine neuromodulation: hygienic behavior is independent of sucrose responsiveness and foraging ontogeny. Hormones and Behavior. 49: 391-397.

Gramacho, KP and Spivak M. 2003. Differences in olfactory sensitivity and behavioral responses among honey bees bred for hygienic behavior. Behav. Ecol. Sociobiol. 54: 472-479.

Spivak, M., Masterman, R., Ross, R., Mesce, KA. 2003. Hygienic behavior in the honey bee (Apis mellifera L.) and the modulatory role of octopamine. J. Neurobiol. 55: 341-354.

Lapidge, K., Oldroyd, B., Spivak, M. 2002. Seven suggestive quantitative trait loci influence hygienic behavior of honey bees. Naturwissenschaften 89: 565-568.

Masterman, R. Ross, R., Mesce, K., Spivak, M. 2001. Olfactory and behavioral response thresholds to odors of diseased brood differ between hygienic and non-hygienic honey bees (Apis mellifera L.) J. Comp Physiol. A 187: 441-452.

Arathi, H.S., Spivak, M. 2001 Influence of colony genotypic composition on the performance of hygienic behavior in the honey bee (Apis mellifera L). Animal Behavior 62: 57-66.

Arathi, H.S., Burns, I. , Spivak, M. 2000. Ethology of hygienic behaviour in the honey bee, Apis mellifera (Hymenoptera: Apidae): Behavioural repertoire of hygienic bees. Ethology 106: 1-15.

Masterman, R., Smith, B. Spivak, M. 2000. M. Evaluation of brood odor discrimination abilities in honey bees (Apis mellifera L.) using proboscis extension reflex conditioning. J. Insect Behav. 13(1): 87-101.

Boecking, O., Spivak, M. 1999. Behavioral defenses of honey bees against Varroa jacobsoni Oud. Apidologie 30: 141-158.

Spivak, M. 1996. Honey bee hygienic behavior and defense against Varroa jacobsoni. Apidologie 27: 245-260.

Spivak, M., Gilliam, M. 1993. Facultative expression of hygienic behaviour of honey bees in relation to disease resistance. J. of Apicultural Res. 32(3/4): 147-157.

Of Interest to Beekeepers

Spivak, M, Browning Z, Goblirsch M, Lee K, Otto C, Smart M, Wu-Smart J.  In review.  Why does bee health matter?  The science surrounding honey bee health concerns and what we can do about it.  Council for Agricultural Science and Technology (CAST) Commentary, QTA2017. CAST, Ames, Iowa.

Spivak, M. 2015.  Hygienic Behavior. Science Buzz, American Beekeeping Federation newsletter, May 2015.

Spivak, M. 2014.  What will happen if the bees disappear? May 17,  http://www.cnn.com/2014/05/17/opinion/spivak-loss-of-bees/

Spivak, M.  2014.  Bothered by Bees or Wasps (.pdf)? 

Koch, R.L. and M. Spivak. 2013. Protect pollinators while trying to protect your crops. Minnesota Crop News. August 5, 2013. 

The Benefits of Propolis, Marla Spivak, PhD, www.bee-craft.com

Boots on the Ground, MEA McNeil

Getting Bees Back on their Own Six Feet, MEA McNeil

Lee KV, Moon RD, Burkness EC, Hutchison WD, Spivak M. 2010. Practical sampling plans for Varroa destructor (Acari: Varroidae) in Apis mellifera (Hymenoptera: Apidae) colonies and apiaries. J. Econ. Entomol. 103:1039–1050. doi:10.1603/EC10037.

The plight of bees August 2010

The future of the MN Hygienic stock of bees is in good hands! October 2009

Socialized Medicine: Individual and communal disease barriers in honey bees June 2009

New Direction for the Minnesota Hygienic Line of Bees Dec 2008

Conserving Pollinators: A Primer for Gardeners March 2009

Bee Health: Putting Control in Last Place. American Bee Journal November 2008

Why are honey bees collapsing? Mar 2007

A Sustainable Approach to Controlling Honey Bee Diseases and Varroa Mites

In the News: Popular Press

TED talk, Why Bees are disappearing, Marla Spivak PhD,

Secrets of the Hive; Propolis might have life-saving properties, By Sara Specht, Solutions, Winter 2008

The Bee Queen, Minnesota Magazine, Nov 2005

The Life-giving Secret of Bees, The Rake, Sept 2005

Minnesota Bee Research and Extension (.pdf), Bee Culture, Jan 2004

Older Studies on Africanized Bees by Spivak

Spivak M. 1992. The relative success of Africanized and European honey bees over a range of life-zones in Costa Rica. J. Applied Ecology 29(19):150-163.

Spivak M, Erickson E Jr. 1992. Do measurements of worker cell size reliably distinguish Africanized from European honey bees (Apis mellifera L.)? Am. Bee J. 132(4): 252-255.

Smith R-K, Spivak M,Taylor OR Jr, Bennett C, Smith ML. 1992. Chemotaxonomy of Africanized bees. Part 3: Identification of Africanized honey bee queens. Bee Science 2:93-105.

Spivak M, Castro AL, Segreda F, Ramirez W. 1989. Honey production by Africanized and European Honey Bees in Costa Rica. Apidologie 20:207-220.

Taylor OR Spivak M. 1984. Climatic limits of tropical African honeybees in the Americas. Bee World 65(1):38-47.

Spivak M, Fletcher DJC, Breed MD, editors. 1991. The "African" Honey Bee. Westview Press, Boulder, Colorado. 450 pp.

Spivak M. 1991. The Africanization process in Costa Rica. In: The "African" Honey Bee. ed. M. Spivak, D.J.C. Fletcher, M.D. Breed, pp. 130-148. Westview Press, Boulder, Colorado.

Spivak M, Ranker T, Taylor OR, Taylor W, Davis L. 1988. Discrimination of Africanized honey bees using cell size, behavior, morphometrics and a newly discovered isozyme polymorphism. In: Africanized Honey Bees and Bee Mites. ed. R. Page, G. Needham, M. Delfinado-Baker, C.E. Bowman, pp. 313-324. Ellis Horwood Limited, Chichester, England.

A Collection of Other Spivak Publications

Spivak M, Boecking O. 2001. Honey bee resistance to Varroa jacobsoni mites. In: Mites of the Honey Bee ( T Webster & K Delaplane, eds). pp. 205-227. Dadant & Sons, Hamilton, IL

Otis GW, Taylor OR, Jr, Spivak M, Winston ML, Katz SJ, Kukuk PF. 1999. Timing of mating flights of neotropical African and European honey bee queens and drones (Hymenoptera: Apidae) in eastern Venezuela. Univ. Kansas Nat. Hist. Mus. Spec. Publ. 24: 90-96.

Calderone NW, Wilson WT, Spivak M. 1997. Evaluation of plant extracts for control of the parasitic mites Varroa jacobsoni (Acari: Varroidae) and Acarapis woodi (Acari: Tarsonemideae) in colonies of the Western honey bee, Apis mellifera (Hymenoptera: Apidae) J. Econ. Entomol. 90: 1080-1086.

Spivak M. 1996. Honey bee hygienic behavior and defense against Varroa jacobsoni. Apidologie 27: 245-260.

Calderone NW, Spivak M. 1995. Plant extracts for control of the parasitic mite, Varroa jacobsoni (Acari: Varroidae), in colonies of the Western honey bee, Apis mellifera (Hymenoptera: Apidae). J. Econ. Entomol. 88(5): 1211-1215.

Smith R-K, Spivak M, Taylor OR, Bennett C, Smith ML. 1993. Maturation of tergal gland alkene profiles in European honey bee queens Apis mellifera L. J. Chem Ecol. 19: 133-42

DeGrandi-Hoffman G, Spivak M, Martin JE. 1993. The influence of temperature on cuticular color of honey bee (Apis mellifera L.) queens. Apidologie 24: 101-108.

DeGrandi-Hoffman G, Spivak M, Martin JE. 1993. The role of thermoregulation by nestmates on honey bee (Apis mellifera L.) queen development time and color. Ann. Entomol. Soc. Amer. 86(2): 165-172.

Spivak M, Zeltzer A, DeGrandi-Hoffman G, Martin J. 1992. The influence of temperature during pupation on the development time and color of queen bees. Environ. Entomol. 21(2): 364-370.

Spivak M, Conlon D, Bell W. 1991. Wind-guided landing and search behavior in blowflies and fleshflies exploiting a resource patch (Diptera: Calliphoridae, Sarcophagidae). Ann. Entomol. Soc. Am. 84(4):447-452.