Department of Biology

Dr. Andrew Van Zandt Brower


Phone: (615) 898-2064
Fax: (615) 898-5093

Office Location:
Haynes House, Rm. 5

Mail Address:

Department of Biology
P.O. Box 60
Middle Tennessee State University
Murfreesboro, TN 37132


B.A., 1985, Yale College (major in biology)
M.E.S., 1987, Yale School of Forestry and Environmental Studies
(concentrations in conservation biology and museology)
Ph.D., 1994, Cornell University, Section of Ecology and Systematics

My empirical research is focused on the molecular systematics, biogeography and evolution of butterflies, mainly in the family Nymphalidae. I am also interested in theoretical principles and methods of systematic biology and biological nomenclature.

Research Associate, Dept. of Entomology, National Museum of Natural History,
Research Associate, Dept. of Invertebrates, American Museum of Natural History,
Associate Editor, Cladistics
Associate Editor , Systematics and Biodiversity
Editorial Board, Canadian Entomologist
Rhopalocera TWIG leader, NSF ATOL Lepidoptera Grant (Mitter, Regier, Cummings, Parr, Weller, Davis)
Taxonomy Advisor (Rhopalocera), NCBI GenBank

Publications: 103 total authored, co-authored or edited (57 peer-reviewed research publications, 1 book, 45 reviews, comments and published abstracts)
Web publications: ~2300 pages in the Tree of Life
Presentations: 96 total authored or co-authored (80 invited keynotes or lectures, 16 contributed papers)

Selected peer-reviewed publications—
Brower, AVZ. 1994. Rapid morphological radiation and convergence in geographical races of the butterfly, Heliconius erato, inferred from patterns of mitochondrial DNA evolution. Proc. Natl. Acad. Sci. USA 91:6491-6495.
Brower, AVZ. 1994. Phylogeny of Heliconius butterflies inferred from mitochondrial DNA sequences. Mol. Phylogenet. Evol. 3:159-174.
Brower, AVZ. and R DeSalle. 1994. Practical and theoretical considerations for choice of a DNA sequence region in insect molecular systematics, with a short review of published studies using nuclear gene regions. Ann. Entomol. Soc. Amer. 87:702-716.
Brower, AVZ. 1995. Locomotor mimicry in butterflies? A critical review of the evidence. Phil. Trans. Royal Soc. Lond. B 347: 413-425.
Brower, AVZ. 1996. Parallel race formation and the evolution of mimicry in Heliconius butterflies: a phylogenetic hypothesis from mitochondrial DNA sequences. Evolution 50:195-221.
Brower, AVZ. 1996. A new mimetic species of Heliconius (Lepidoptera: Nymphalidae), from southeastern Colombia, revealed by cladistic analysis of mitochondrial DNA sequences. Zool. J. Linn. Soc. 116:317-332.
Brower, AVZ, R DeSalle and AP Vogler. 1996. Gene trees, species trees, and systematics: a cladistic perspective. Annu. Rev. Ecol. Syst. 27:423-450.
Brower, AVZ and VA Schawaroch. 1996. Three steps of homology assessment. Cladistics 12:265-272.
Miller, JS, AVZ Brower and R DeSalle. 1997. Phylogeny of the neotropical moth tribe Josiini (Notodontidae: Dioptinae): Comparing and combining evidence from DNA sequences and morphology. Biol. J. Linn. Soc. 60: 297-316.
Brower, AVZ. 1997. The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): a cladistic review. Zool. J. Linn. Soc. 119: 457-472.
Brower, AVZ and MG Egan. 1997. Cladistics of Heliconius butterflies and relatives (Nymphalidae: Heliconiiti): the phylogenetic position of Eueides based on sequences from mtDNA and a nuclear gene. Proc. Royal Soc. Lond. B 264: 969-977.
DeSalle, R and AVZ Brower. 1997. Process partitions, congruence and the independence of characters: inferring relationships among closely-related Hawaiian Drosophila from multiple gene regions. Syst. Biol. 46: 751-764.
Brower, AVZ and R DeSalle. 1998. Mitochondrial vs. nuclear DNA sequence evolution among nymphalid butterflies: the utility of Wingless as a source of characters for phylogenetic inference. Insect Mol. Biol. 7: 1-10.
Brower, AVZ. 1999. The delimitation of phylogenetic species with DNA sequences: a critique of Davis and Nixon's population aggregation analysis. Syst. Biol. 48: 199-213.
Brower, AVZ. 2000. Evolution is not a necessary assumption of cladistics. Cladistics 16: 143-154.
Campbell, DL, AVZ Brower and NE Pierce. 2000. Molecular evolution of the Wingless gene and its implications for the phylogenetic placement of the butterfly family Riodinidae (Lepidoptera: Papilionoidea). Mol. Biol. Evol.17: 684-696.
Brower, AVZ. 2000. Phylogenetic relationships among the Nymphalidae (Lepidoptera), inferred from partial sequences of the wingless gene. Proc. R. Soc. Lond. B 267: 1201-1211.
Brower, AVZ. 2002. Cladistics, populations and species in geographical space: the case of Heliconius butterflies. Pp.5-16 in R DeSalle, WC Wheeler and G Giribet (eds.), Molecular Systematics and Evolution: theory and practice. Birkhaüser Verlag.
Goldstein, PZ and AVZ Brower. 2002. Molecular systematics and the origin of species: new syntheses or methodological introgressions? Pp. 147-161 in R DeSalle, WC Wheeler and G Giribet (eds.), Molecular Systematics and Evolution: theory and practice. Birkhaüser Verlag.
Brower, AVZ and M Jeansonne. 2004. Geographical populations and "subspecies"; of New World monarch butterflies ( Danaus plexippus (L.) Nymphalidae) share a recent origin and are not phylogenetically distinct. Ann. Entomol. Soc. Amer. 97: 519-523.
Leathers, JW, DD Judd, and AVZ Brower. 2005. A review of the species of the New World braconid genus Cyclaulacidea (Hymenoptera) with key and descriptions of nine new species. J. Hymenoptera Research 14:151-176.
Silva-Brandão, KL, AVL Freitas, AVZ Brower and VN Solferini. 2005. Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF1? genes. Mol. Phylogenet. Evol. 36: 468-483.
Wahlberg, N., MF Braby, AVZ Brower, R de Jong, M-M Lee, S Nylin, NE Pierce, FAH Sperling, R Vila, AD Warren and E Zakharov. 2005. Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proc. R. Soc. Lond. B 272: 1577-1586.
Wahlberg, N, AVZ Brower and S Nylin. 2005. Phylogenetic relationships of tribes and genera in the subfamily Nymphalinae ((Lepidoptera: Nymphalidae) based on three gene sequences. Biol. J. Linn. Soc. 86: 227-251.
Whinnett A, AVZ Brower, M-M Lee, KR Willmott, and J Mallet 2005. The phylogenetic utility of Tektin, a novel region for inferring systematic relationships amongst Lepidoptera. Ann. Entomol. Soc. Amer. 98: 873-886.
Whinnett A, KR Willmott, AVZ Brower, F Simpson, M Zimmermann, G Lamas, and J Mallet. 2005. Mitochondrial DNA provides an insight into the mechanisms driving diversification in the ithomiine butterfly Hyposcada anchiala (Lepidoptera: Nymphalidae, Ithomiinae). European J. Entomol. 102: 633-639.
Brower, AVZ. 2006. The how and why of branch support and partitioned branch support, with a new index to assess partition incongruence. Cladistics 22: 378-386.
Brower, AVZ. 2006. Problems with DNA barcodes for species delimitation: "ten species"; of Astraptes fulgerator reassessed (Lepidoptera: Hesperiidae). Syst. and Biodiv. 4: 127-132.
Brower, AVZ, AVL Freitas, M-M Lee, KL Silva Brandão, A Whinnett and KR Willmott 2006. Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions. Syst. Entomol. 31: 288-301.
Peña, C, N Wahlberg, E Weingartner. U Kodandaramaiah, S Nylin, AVL Freitas and AVZ Brower. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Mol. Phylogenet. Evol. 40: 29-49.
Simonsen, TJ, N Wahlberg, AVZ Brower and R de Jong 2006. Molecules, morphology and fritillaries: a combined approach towards the phylogeny of Argynnini (Lepidoptera: Nymphalidae). Insect Systematics and Evolution 37: 405-418.
Elias, M, RI Hill, KR Willmott, KK Dasmahaptra, AVZ Brower, J Mallet and CD Jiggins. 2007. Limited performance of DNA barcoding in a diverse community of tropical butterflies. Proc. R. Soc London B 274: 2881-2889 (on line: doi:10.1098/rspb2007.1035)
Beltrán, M, CD Jiggins, AVZ Brower, E Bermingham and J Mallet. 2007. Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA sequence data. Biol. J. Linn. Soc. 92:221-239.
Frentiu, FD, GD Bernard, MP Sison-Mangus, AVZ Brower and AD Briscoe. 2007. The role of gene duplication in the evolution of spectral shifts in the long wavelength photopigments of butterflies. Mol. Biol. Evol. 24: 2016-2028.
Warren, AD, J Ogawa and AVZ Brower. 2008. Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). Cladistics 24: 642-676.
Townzen, JS, AVZ Brower and DD Judd. 2008. Identification of mosquito bloodmeals using mitochondrial cytochrome oxidase subunit I and cytochrome b gene sequences. Med. Vet. Entomol. 22: 386-393
Elias, M, M Joran, KR Willmott, V Kaiser, KL Silva-Brandão, C Arias, LM Gomez-Piñeres, S Uribe, AVZ Brower, AVL Freitas and CD Jiggins. 2009. Out of the Andes: patterns of diversification in clearwing butterflies. Mol. Ecol. 18: 1716-1729.
Warren, AD, J Ogawa and AVZ Brower. 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology 34: 467-523.
Brower, AVZ. 2009. Heliconius tristero Brower, 1996 and Heliconius melpomene mocoa Brower, 1996 (Lepidoptera: NYMPHALIDAE ): proposed conservation by suppression of Heliconius melpomene bellula Brown, 1979. Bulletin of Zoological Nominclature 66: 256-260.
Wahlberg, N, J Leneveu, U Kodandaramaiah, C Peña, S Nylin, ALV Freitas and AVZ Brower. 2009. Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary. Proc. R. Soc. London B 276: 4295-4302.
Brower, AVZ. 2010. Stability, replication, pseudoreplication and support. Cladistics 25: 112-113.
Brower, AVZ, N Wahlberg, JR Ogawa, M Bopp
ré and RI Vane-Wright. 2010. Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences. Systematics and Biodiversity 8: 75-89.
Brower, AVZ. 2010. Alleviating the taxonomic impediment of DNA barcoding: names for ten species of "Astraptes fulgerator" (Lepidoptera: Hesperiidae: Eudaminae). Systematics and Biodiversity 8: 485-491.
Brower, AVZ. 2011. Hybrid speciation in Heliconius butterflies: a review and critique of the evidence. Genetica 139: 589-609; online version DOI: 10.1007/s10709-010-9530-4
Rindal, E and AVZ Brower. 2011. Do model-based phylogenetic analyses outperform parsimony? A test with empirical data. Cladistics 27: 331-334; online version DOI:10.1111/j.1096-0031.2010.00342.x
Brower, AVZ. 2011. Repeatability and reality. Cladistics online version DOI: 10.111/j.1096-0031.2011.00354.x

Schuh, RT and AVZ Brower. 2009. Biological Systematics: principles and applications. 2nd edn. Ithaca: Cornell University Press. 311 pp.

Rice Chair of Systematic Entomology, Oregon State University
Assistant Professor (Nov. 1997- June 2006)
Curator of Lepidoptera, Oregon State Arthropod Collection, Oregon State University
(Nov. 1997-June 2006 )
Smithsonian Postdoctoral Fellow, National Museum of Natural History (Mar.-Nov. 1997)
Kalbfleisch Fellow, American Museum of Natural History (May 1996-Feb. 1997 )
Postdoctoral Fellow, American Museum of Natural History (Oct. 1995-April 1996)
NSF Postdoctoral Fellow, American Museum of Natural History (Oct. 1993-Sept. 1995)