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Ted Schultz, Curator of Hymenoptera (Ants)

  • Phone:   202-633-0999
  • Fax:   202-786-3141
  • E-mail Address:

  • USPS Mailing Address:

    Smithsonian Institution
    PO Box 37012, MRC 188
    Washington, DC 20013-7012

  • Shipping Address:
    Smithsonian Institution
    National Museum of Natural History
    10th & Constitution NW
    Washington, DC 20560-0188

  • Education:

    Ph.D. Cornell University
    B.A. University of California, Berkeley

Ted Schultz

Research Interests:

  • Evolution and systematics of ants, especially the fungus-growing ants (Myrmicinae: Attini). 
  • Phylogenetic analysis using morphological and molecular characters. 
  • Historical ecology and evolution of the fungus-growing behavior. 
  • Theory and methodology of phylogenetic analysis, especially the reconstruction of ancestral character states. 
  • Quantitative methods for assessing ant biodiversity.

Research in the Smithsonian AntLab largely focuses on the systematics, phylogenetics, and biodiversity of ants (family Formicidae), (although AntLab researchers have conducted projects on bees, termites, dragonflies, spiders, and beetles).

Two main themes run through most of our research:

  • the evolution, phylogenetics, and systematics of all ants, with special emphasis on the fungus-growing ants (tribe Attini, subfamily Myrmicinae);
  • the quantitative sampling of leaf-litter ants of South America with particular focus on the ants of the Guiana Shield (a collaboration with John LaPolla, Jeffrey Sosa-Calvo, Brad Wright, and Ted Suman).

Current AntLab personnel include Museum Specialist Eugenia Okonski; Smithsonian Postdoctoral Fellows Bernice DeMarco and Fred Larabee; graduate students Gabi Camacho and Rick Gawne; and a number of Research Associates including Jeffrey Sosa-Calvo, Natasha Mehdiabadi, Dietrich Gotzek, Christian Rabeling, Leeanne Alonso, John LaPolla, Scott Solomon, and Ted Suman; and contract technician Brad Wright. Honorary lifetime AntLab personnel include Faridah Dahlan, Matt Kweskin, Paul Armstrong, Hong Zhao, and Michael Lloyd. (see more at SI AntLab)

Here in the Department of Entomology in the Smithsonian Institution's National Museum of Natural History, TRS is the "contact person" for the following groups of aculeate Hymenoptera: Chrysidoidea (Bethylidae, Chrysididae, Dryinidae, Embolemidae, Plumariidae, Sclerogibbidae, Scolebythidae) and Vespoidea excluding Vespidae (Bradynobaenidae, Formicidae, Mutillidae, Pompilidae, Rhopalosomatidae, Sapygidae, Scoliidae, Sierolomorphidae, Tiphiidae).



Branstetter, M.G., Ješovnik, A., Sosa-Calvo, J., Lloyd, M.W., Faircloth, B.C., Brady, S.G., and Schultz, T.R. 2017. Dry habitats were crucibles of domestication in the evolution of agriculture in ants. Proceedings of the Royal Society, B. 284: 20170095. doi: 10.1098/rspb.2017.0095

Ješovnik A, Sosa-Calvo J, Lloyd M, Branstetter M, Fernandez F & Schultz TR (2016): Phylogenomic species delimitation and host-symbiont coevolution in the fungus-farming ant genus Sericomyrmex Mayr (Hymenoptera: Formicidae): Ultraconserved elements (UCEs) resolve a recent radiation. Systematic Entomology.
doi: 10.1111/syen.12228    

Sosa-Calvo J, Ješovnik A, Lopes C, Rodrigues A, Rabeling C, Bacci M Jr, Vasconcelos HL & Schultz TR (2017): Biology of the relict fungus-farming ant Apterostigma megacephala Lattke, including descriptions of the male, gyne, and larva. Insectes Sociaux. Link:

Ward, P.S., Brady, S.G., Fisher, B.L., Schultz, T.R. 2016. Phylogenetic classifications are informative, stable and pragmatic: the case for monophyletic taxa. Insectes Sociaux 63:489–492. doi:10.1007/s00040-016-0516-9 

Ješovnik A, González VL, Schultz TR (2016) Phylogenomics and Divergence Dating of Fungus-Farming Ants (Hymenoptera: Formicidae) of the Genera Sericomyrmex and Apterostigma. PLoS ONE 11(7): e0151059. doi:10.1371/journal. pone.0151059

Nygaard, S., Hu, H., Li, C., Schiøtt, M., Chen, Z., Yang, Z., Xie, Q., Ma, C., Deng, Y., Dikow, R., Rabeling, C., Nash, D.R., Wcislo, W.T., Brady, S.G., Schultz, T.R., Zhang, G., Boomsma, J.J. 2016. Reciprocal genomic evolution in the ant-fungus agricultural symbiosis. Nature Communications 7:12233. doi:10.1038/ncomms12233

Fernandes, I.O., Souza, J.L.P., Fernández C., F., Delabie, J.H.C. and Schultz, T.R. 2015. A new species of Simopelta (Hymenoptera: Formicidae: Ponerinae) from Brazil and Costa Rica.  Zootaxa, 3956(2): 295-300. 

Sosa-Calvo, J., Jesovnik, A., Okonski, E. and Schultz, T.R. 2015. Locating, collecting, and maintaining colonies of fungus-farming ants (Hymenoptera: Myrmicinae: Attini). Sociobiology, 62(2) 

Blaimer, B.B., Brady, S.G., Schultz, T.R., Lloyd, M.W., Fisher, B.L., Ward, P.S. 2015. Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants. BMC Evolutionary Biology 15:271. doi: 10.1186/s12862-015-0552-5

Rabeling, C., Schultz, T.R., Bacci, Jr., M., Bollazzi, M. 2015. Acromyrmex charruanus: a new inquiline social parasite species of leaf-cutting ants. Insectes Sociaux doi 10.1007/s00040-015-0406-6

Schultz, T.R., Sosa-Calvo, J., Brady, S.G., Lopes, C.T., Mueller, U.G., Bacci, M. & Vasconcelos, H.L. 2015. The Most Relictual Fungus-Farming Ant Species Cultivates the Most Recently Evolved and Highly Domesticated Fungal Symbiont Species. The American Naturalist, 185(5): 693-703. doi:10.1086/680501

Blaimer, B.B., Brady, S.G., Schultz, T.R. and Fisher, B.L. 2015. Functional and phylogenetic approaches reveal the evolution of diversity in a hyper diverse biota. Ecography, doi:10.1111/ecog.01370

Ward, P.S., Brady, S.G., Fisher, B.L. and Schultz, T.R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade. Systematic Entomology, 40(1): 61-81. doi:10.1111/syen.12090

Masiulionis, V.E., Rabeling, C., De Fine Licht, H.H., Schultz, T.R., Bacci, M., Bezerra, C.M.S. and Pagnocca, F.C. 2014. A Brazilian Population of the Asexual Fungus-Growing Ant, Mycocepurus smithii (Formicidae, Myrmicinae, Attini), Cultivates Fungal Symbionts with Gongylidia-Like Structures. Plos One, 9(8): 1-8. doi:10.1371/journal.pone.0103800

Rabeling, Christian, Schultz, Ted R, Pierce, Naomi E and Bacci Jr., Maurício. 2014. A Social Parasite Evolved Reproductive Isolation from Its Fungus-Growing Ant Host in Sympatry. Current Biology, 24(17): 2047-2052. doi:10.1016/j.cub.2014.07.048

Brady, S.G., Fisher, B.L., Schultz, T.R., Ward, P.S. 2014. The rise of the army ants: Diversification of the specialized predatory doryline ants. BMC Evolutionary Biology 14:e93. doi:10.1186/1471-2148-14-93

Sosa-Calvo J, Schultz TR, Brandão CRF, Klingenberg C, Feitosa RM, Rabeling C, Bacci Jr. M, Lopes CT and Vasconcelos HL. (2013) Cyatta abscondita: Taxonomy, Evolution, and Natural History of a New Fungus-Farming Ant Genus from Brazil. PLoS ONE 8(11): e80498. doi:10.1371/journal.pone.0080498

pdf Jesovnik, A., Sosa-Calvo, J.; Lopes, C., Vasconcelos, H., Schultz, T. R. 2013. Nest architecture, fungus gardens, queen, males and larvae of the fungus-growing ant Mycetagroicus inflatus Brandão & Mayhé-Nunes. Insectes Sociaux 60(4): 531-542.

Adams, R.M.M., Jones, T.H., Jeter, A.W., De, F.L., Schultz, T.R. and Nash, D.R. 2012. A comparative study of exocrine gland chemistry in Trachymyrmex and Sericomyrmex fungus-growing ants. Biochemical systematics and ecology, 40: 91-97. doi:10.1016/j.bse.2011.10.011

Mehdiabadi, N.J., Mueller, U.G., Brady, S.G., Himler, A.G., Schultz, T.R. 2012. Symbiont fidelity and the origin of species in fungus-growing ants. Nature Communications, 3 doi:10.1038/ncomms1844

Jones, Tappey H., Adams, Rachelle M. M., Spande, Thomas F., Garraffo, H. M., Kaneko, Tetsuo and Schultz, Ted R. 2012. Histrionicotoxin Alkaloids Finally Detected in an Ant. Journal of natural products, doi:10.1021/np300485v

Rey, O., Estoup, A., Vonshak, M., Loiseau, A., Blanchet, S., Calcaterra, L., Chifflet, L., Rossi, J.-P., Kergoat, G. J., Foucaud, J., Orivel, J., Leponce, M., Schultz, T. and Facon, B. 2012. Where do adaptive shifts occur during invasion? A multidisciplinary approach to unravelling cold adaptation in a tropical ant species invading the Mediterranean area. Ecology Letters, 15(11): 1266-1275. doi:10.1111/j.1461-0248.2012.01849.x

Rabeling, C., Gonzales, O., Schultz, T.R., Bacci, M., Garcia, M.V.B., Verhaagh, M., Ishak, H.D. and Mueller, U.G. 2011. Cryptic sexual populations account for genetic diversity and ecological success in a widely distributed, asexual fungus-growing ant. Proceedings of the National Academy of Sciences of the United States of America, 108(30): 12366-12371. doi:10.1073/pnas.1105467108

Solomon, S. E., Lopes, C.T., Mueller, U.G., Rodrigues, A., Sosa-Calvo, J., Schultz, T.R. and Vasconcelos, H.L. 2011. Nesting biology and fungiculture of the fungus-growing ant, Mycetagroicus cerradensis: New light on the origin of higher-attine agriculture. Journal of Insect Science, 11(12): 1-14.

Mehdiabadi, N. and Schultz, T.R. 2010. Natural history and phylogeny of the fungus-farming ants (Hymenoptera: Formicidae: Myrmicinae: Attini). Myrmecological News, 13: 37-55.

Sosa-Calvo, J., T.R. Schultz, and J.S. LaPolla. 2010. A review of the dacetine ants of Guyana (Formicidae: Myrmicinae). Journal of Hymenoptera Research, 19(1): 12-43. pdf

Ward, P.S., S.G. Brady, B.L. Fisher, and T.R. Schultz.  2010.  Phylogeny and biogeography of dolichoderine ants: Effects of data partitioning and relict taxa on historical inference.  Systematic Biology. 59:342-362. pdf

pdf Sosa-Calvo, J. and T.R. Schultz. 2010. Three remarkable new fungus-growing ant species of the genus Myrmicocrypta (Hymenoptera: Formicidae), with a reassessment of the characters that define the genus and its position within the Attini. Ann. Entomol. Soc. Amer. 103(2): 181-195.

Schultz, T. R. 2010. Ants in museums. In: Ant Ecology (L. Lach, C. Parr, and K. Abbott, eds.). Oxford University Press, New York.

pdf Sosa-Calvo, J., S. G. Brady, and T. R. Schultz.  2009.  The gyne of the enigmatic fungus-farming ant species Mycetosoritis explicata.  Journal of Hymenoptera Research 18: 113-120.

PDF Schultz, T. R. and Brady, S. G. 2008 Major evlolutionary transitions in ant agriculture. Proceedings of the National Academy of Sciences, U.S.A. 105:5435-5440. (Cover article) PNAS Early Edition version published 24 March 2008. (With accompanying "Commentary": U.G. Mueller and C. Rabeling.  2008.  “A breakthrough innovation in animal evolution.”  Proceedings of the National Academy of Sciences 105: 5287-5288.)
Supporting Information:

Schultz, T.R. and J. Sosa-Calvo. 2008. Ants of southern Guyana – A preliminary report (pp. 31-32) and Preliminary list of ant genera collected in the Konashen COCA, October 2006, Appendix 1 (p. 72).  In A Rapid Biological Assessment of the Konashen Community Owned Conservation Area, Southern Guyana: RAP Bulletin of Biological Assessment 51.  (L. Alonso, J. McCullough, P. Naskrecki, E. Alexander, and H.E. Wright, eds.).  Washington, DC: Conservation International.  [Technical report.]

PDF LaPolla, J.S., T. Suman, J. Sosa-Calvo, and T.R. Schultz.  2007.  Leaf litter ants of Guyana.  Biodiversity and Conservation 16: 491-510.

PDF Schultz, T.R.  2007. The fungus-growing ant genus Apterostigma in Dominican amber. In: Advances in Ant Systematics (Hymenoptera: Formicidae): Homage to E.O. Wilson – 50 Years of Contributions (R.R. Snelling, B.L. Fisher, and P.S. Ward, Eds.).  Memoirs of the American Entomological Institute 80: 425-436.

Taerum, S.J., M.J. Cafaro, A.E.F. Little, T.R. Schultz, and C.R. Currie.  2007.  Low host-pathogen specificity in the leaf-cutting ant-microbe symbiosis.  Proceedings of the Royal Society (London), Series B, 274: 1971-1978.

PDF Brady, S. G., T. R. Schultz, B. L. Fisher & P. S. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences 103: 18172-18177. (With accompanying "Commentary": Ross H. Crozier.  2006.  “Charting uncertainty about ant origins.”  Proceedings of the National Academy of Sciences 103: 18029-18030.) 
Supplemental online material:

LaPolla, J.S., T.R. Schultz, K.M. Kjer and J.F. Bischoff. 2006. Phylogenetic position of the ant genus Acropyga Roger (Hymenoptera: Formicidae) and the evolution of trophophoresy.  Insect Systematics and Evolution 37: 197-212.

Sosa-Calvo, J., S. O. Shatuuck, & T. R. Schultz. 2006. Dacetine ants of Panama: New records and description of a new species (Formicidae: Myrmicinae: Dacetini). Proceedings of the Entomological Society of Washington 108: 814-821.

PDF Mueller, U. G, N. M. Gerardo, D. K. Aanen, D. L. Six, & T. R. Schultz. 2005. The evolution of agriculture in insects. Annual Review of Ecology, Evolution, and Systematics 36: 563-595.

Schultz, T. R., U. G. Mueller, C. R. Currie, & S. A. Rehner. 2005. Reciprocal illumination: A comparison of agriculture in humans and ants. Pp. 149-190. In: Fernando Vega and Meredith Blackwell (eds.). Ecological and Evolutionary Advances in Insect-Fungal Associations. Oxford University Press, New York.

PDF Ward, P. S., Brady, S. G., Fisher, B. L. & Schultz, T. R. 2005. Assembling the ant "Tree of Life" (Hymenoptera: Formicidae). Myrmecologische Nachrichten 7: 87-90.

PDF Solomon, S. E., U. G. Mueller, T. R. Schultz, C. R. Currie, S.L. Price, A. C. Oliveira da Silva-Pinhati, M. Bacci Jr., & H. L.Vasconcelos. 2004. Nesting biology of the fungus growing ants Mycetarotes Emery (Attini, Formicidae). Insectes Sociaux 51: 333-338.

PDF Villesen, P., U. G. Mueller, T. R. Schultz, R. M. M. Adams, & M. C. Bouck. 2004. Evolution of ant-cultivar specialization and cultivar switching in Apterostigma fungus-growing ants. Evolution 58: 2252-2265.

PDF Currie, C. R., B. Wong, A. E. Stuart, T. R. Schultz, S. A. Rehner, U. G. Mueller, G. H. Sung, J. W. Spatafora, and N. A. Straus. 2003. Ancient tripartite coevolution in the attine ant-microbe symbiosis. Science 299: 386-388. (With accompanying "News" article: E. Pennisi.  2003.  "On ant farm, a threesome co-evolves."  Science 299: 325.) 
Supplemental online material:

Price, S. L., T. Murakami, U. G. Mueller, T. R. Schultz, and C. R. Currie. 2003. Recent Findings in Fungus-Growing Ants: Evolution, Ecology, and Behavior of a Complex Microbial Symbosis. Pp. in: Genes, Behaviors, and Evolution of Social Insects (T. Kikuchi, N. Azuma, and S. Higashi, Eds.), Hokkaido University Press, Sapporo, Japan. [Book chapter.]

PDF Schultz, T. R. 2003. "Hyperdiversity Up Close." Review of Pheidole in the New World by E.O. Wilson.  Science 300: 57-58.  [Book review.]

Bang, R., T. R. Schultz, and R. DeSalle. 2002. Development, homology, and systematics. In: DeSalle, R., G. Giribet and W. Wheeler (eds.). Molecular Systematics and Evolution: Theory and Practice. Birkhauser, Basel.

Hoyt, Erich and T. R. Schultz (Editors). 2002. Insect Lives: Stories of Mystery and Romance From a Hidden World. (Trade paperback edition.)  Cambridge, Massachusetts: Harvard University Press.  [Book.]

PDF Schultz, T. R., S. A. Solomon, U. G. Mueller, P. Villesen, J. J. Boomsma, R. M. M. Adams, & B. Norden. 2002. Cryptic Speciation in the Fungus-Growing Ants Cyphomyrmex longiscapus Weber and Cyphomyrmex muelleri Schultz and Solomon, new Species (Formicidae: Attini). Insectes Sociaux 49: 331-343.

PDF Vilesen, P., T. Murakami, T. R. Schultz, & J. J. Boomsma. 2002. Identifying the transition between single and multiple mating of queens in fungus-growing ants. Proceedings of the Royal Society (London), Series B 269: 1541-1548.

PDF Artiss, T., T. R. Schultz, D. A. Polhemus, & C. Simon. 2001. Molecular phylogenetic analysis of the dragonfly genera Libellula, Ladona and Plathemis (Odonata: Libellulidae) based on mitochondrial cytochrome oxidase I and 16S rRNA sequence data.  Molecular Phylogenetics and Evolution 18: 348-361.

PDF Mueller, U. G., T. R. Schultz, C. R. Currie, R. M. M. Adams, & D. Malloch. 2001. The origin of the attine ant-fungus mutualism.  Quarterly Review of Biology 76: 169-197.

PDF Sallum, M.A.M., T. R. Schultz, P. G. Foster, K. Aronstein, R. A. Wirtz, & R. C. Wilkerson. 2001. Phylogeny of Anophelinae (Diptera: Culicidae) based on mitochondrial and nuclear ribosomal DNA sequences.  Systematic Entomology 27: 361-382.

Schultz, T. R., M. S. Engel, and J. S. Ascher. 2001. Evidence for the origin of eusociality in the corbiculate bees (Hymoptera: Apidae). Journal of the Kansas Entomological Society 74:10-16.

Schultz, T. R., R. B. Root, and T. Eisner. 2001. "William L. Brown, Jr.: June 1, 1922 - March 30, 1997." Memorial Statements, Cornell University Faculty 1996-97: 12-16. Ithaca, New York: Office of the Dean of the Faculty, Cornell University [Obituary.]

PDF Adams, R. M. M., U. G. Mueller, T. R. Schultz, and B. Norden. 2000. Agro-predation: Usurpation of attine fungus gardens by Megalomyrmex ants.  Naturwissenschaften 87: 549-554.

PDF Agosti, D., J. Majer, L. Alonso, and T. R. Schultz (Editors). 2000. Sampling Ground-dwelling Ants: Case Studies from the Worlds' Rain Forests. Perth, Australia: Curtin School of Environmental Biology Bulletin No. 18.  75 pp.  [Edited volume.]

Agosti, D., J. Majer, L. Alonso, and T. R. Schultz. 2000. The biodiversity challenge (Preface).  Pp. xvii-xix in: Ants: Standard Methods for Measuring and Monitoring Biodiversity.  (D. Agosti, J. Majer, L. Alonso, and T.R. Schultz, eds.).  Washington, D.C.: Smithsonian Institution Press.  [Book preface.]

Agosti, D., J. Majer, L. Alonso, and T. R. Schultz (Editors). 2000. Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, D.C.: Smithsonian Institution Press.  280 pp.  [Book.]

PDF Sallum, M. A. M., T. R. Schultz, and R. C. Wilkerson. 2000. Phylogeny of Anophelinae (Diptera Culicidae) based on morphological characters. Annals of the Entomological Society of America 93: 745-775.

PDF Schultz, T. R. 2000. “In search of ant ancestors.”  “Commentary.”  Proceedings of the National Academy of Sciences 26:14028-14029. [Article.]

Schultz, T. R. and L. E. Alonso. 2000. Glossary.  Pp. 223-229 in: Ants: Standard Methods for Measuring and Monitoring Biodiversity.  (D. Agosti, J. Majer, L. Alonso, and T.R. Schultz, eds.).  Washington, D.C.: Smithsonian Institution Press.  [Book glossary.]

Schultz, T. R. and T. P. McGlynn. 2000. The interactions of ants with other organisms.  Pp. 35-44 in: Ants: Standard Methods for Measuring and Monitoring Biodiversity.  (D. Agosti, J. Majer, L. Alonso, and T.R. Schultz, eds.).  Washington, D.C.: Smithsonian Institution Press.  [Book chapter.]

Hoyt, Erich and T. R. Schultz (Editors). 1999. Insect Lives, Stories of Mystery and Romance from a Hidden World. New York: John Wiley and Sons. [Book.]

PDF Schultz, T. R. 1999. "Ants, Plants, and Antibiotics." "News and Views." Nature 398:747-748. [Article.]

PDF Schultz, T. R. and G. A. Churchill. 1999. The role of subjectivity in reconstructing ancestral character states: A Bayesian approach to unknown rates, states, and transformation asymmetries.  Systematic Biology 48: 651-664.

Schultz, T. R., M. S. Engel, and M. A. Prentice. 1999. Resolving conflict between morphological and molecular evidence for the origin of eusociality in the "corbiculate" bees (Hymenoptera: Apidae): A hypoth­esis-testing approach.  In: Entomological Contributions in Memory of Byron A. Alexander (G.W. Byers, R.H. Hagen, and R.W. Brooks, eds.), University of Kansas Natural History Museum Special Publication 24: 125-138.

Adams, R. M., U. G. Mueller, and T. R. Schultz. 1998. Nest Usurpation and Fungivory in Megalomyrmex sp. Nov. (silvestri species group): Lestobiotic or Predatory Relations with Fungus-Growing Ants? [Abstract]. In: M. P. Schwarz and K.l Hogendoorn, eds (eds.). Social Insects at the Turn of the Millenium: Proceedings of the XIII International Congress of IUSSI, Adelaide, Australia, 29 December 1998-3 January 1999. Flinders University Press, Adelaide, Australia.

PDF Mueller, U.G., S.A. Rehner, and T.R. Schultz.  1998.  The evolution of agriculture in ants.  Science 281: 2034-2038.  (With accompanying "Commentary": Jared Diamond.  1998.  “Ants, crops, and history.”  Science 281: 1974-1975.) 

PDF Schultz, T. R., D. Bekkevold and J. J. Boomsma. 1998. Acromyrmex insinuator new species: An incipient social parasite of fungus-growing ants. Insectes Sociaux 45:457-471.

PDF Wetterer, J. K., T. R. Schultz, and R. Meier. 1998. Phylogeny of fungus-growing ants (tribe Attini) based on mtDNA sequence and morphology.  Molecular Phylogenetics and Evolution 9: 42-47.

Engel, M. S. and T. R. Schultz 1997. Phylogeny and behavior in honey bees (Hymenoptera: Apidae). Annals of the Entomological Society of America 90:43-53.

Meier, R. and T. R. Schultz. 1996. Pilzzucht und Blattschneiden bei Ameisen: Präadaptationen und evolutive Trends. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 35:57-76.

PDF Schultz, T. R., R. B. Cocroft and G. A. Churchill. 1996. The reconstruction of ancestral character states. Evolution 50:504-511.

Schultz, T.R.  1995 ("1994").  Identification Guide to the Ant Genera of the World by Barry Bolton.  Psyche 101: 203-208.  [Book review.]

Schultz, T. R. and R. Meier. 1995. A phylogenetic analysis of the fungus-growing ants (Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae. Systematic Entomology 20:337-370.

PDF Chapela, I. H., S. A. Rehner, T. R. Schultz and U. G. Mueller. 1994. Evolutionary history of the symbiosis between fungus-growing ants and their fungi.  Science 266: 1691-1694.

PDF Hinkle, G., J. K. Wetterer, T. R. Schultz and M. L. Sogin. 1994. Phylogeny of the attine ant fungi based on analysis of small subunit ribosomal RNA gene sequences. Science 266:1695-1697.

Meier, R. & T. R. Schultz. 1994. The fungus-growing ants are monophyletic: A cladistic analysis of the Attini (Formicidae) based on a comparative SEM study of larvae. Verhandlungen der Deutschen Zoologischen Gesellschaft 87:230.

Schultz, T. R. 1993. Stalking the wild attine. Notes from Underground (Cambridge, Mass.: Museum of Comparative Zoology, Harvard University) 8:7-10. [Article.]


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