Tod W. Reeder

Associate Professor

Ph.D. University of Texas at Austin, 1993

treeder@sunstroke.sdsu.edu

Primary Affiliation: Evolutionary Biology Program Area

Secondary Affiliation: Ecology Program Area

Last Modified:18 October, 2007

Research Interests:

My general research interests are in the population and evolutionary biology of amphibians and reptiles. Much of my research has involved the use of molecular and morphological data to address questions pertaining to the evolution and phylogenetic relationships of squamate reptiles (lizards and snakes). More recently, a lot of the research in my lab has been aimed more at the population level in an attempt to address questions at the interface of ecology and evolutionary biology (e.g., phylogeography, gene flow, speciation-species limits). Below are some of the specific research projects that I am participating in:

Diversification and Evolution of the Herpetofauna of the Southwest and Mexico:

I have conducted extensive phylogenetic studies on phrynosomatid lizards, particularly the genera Phrynosoma (horned lizards) and Sceloporus (fence and spiny lizards). Research on Sceloporus continues, with efforts underway to infer the phylogenetic relationships within the jarrovi complex/torquatus species group and species limits within the widespread "S. undulatus". The work on S. "undulatus" is currently a collaborative effort with Adam Leache (now in doctoral program at UC-Berkeley). Within S. "undulatus", we are concentrating on western populations and trying to better understand what is happening at apparent contact zones between major mitochondrial haplotype clades. We are in the process of fine scale mapping of the distributions of mtDNA haplotypes and developing microsatellite loci to gain a nuclear perspective of these contact zones.

For the past several years, I have been investigating the evolution of Aspidoscelis (="Cnemidophorus"; whiptails). In collaboration with Jay Cole (American Museum of Natural History) and Herb Dessauer (LSU Medical Center), we completed a higher-level phylogenetic study of whiptails and their close relatives (Reeder et al., 2002; see Publications). The goal of this project was to rigorously test Cnemidophorus (sensu lato) monophyly, the monophyly of the described species groups, and the interrelationships among these groups. This phylogenetic study was based on mitochondrial 12S and 16S rDNA, allozymes, and morphology. This extensive data base is currently being augmented with additional DNA sequence data from the mitochondrial and nuclear genomes, and additional "Ameiva" and South American "Cnemidophorus" species in order to further resolve some of the more weakly supported relationships among the neotropical cnemidophorines.

The current focus of my research on Aspidoscelis (=North American "Cnemidophorus") is to further investigate the evolution of the sexlineata species group. Using mitochondrial and nuclear DNA sequences, this work is focusing on inferring the phylogenetic relationships between the bisexual species and re-examining the origins of the unisexual lineages. Also, gene flow and species limits problems are being addressed within the C. burti, C. costatus, and C. gularis complexes. This work on the sexlineata group is part of a collaboration with Dr. Adrian Nieto (Universidad Nacional Autonoma de Mexico) and is currently supported by NSF grant DEB-0108484.

Higher-level squamate phylogeny:

I have been collecting morphological and molecular data in order to infer the phylogenetic relationships among the ~40 squamate families. To date, the molecular data primarily consists of mitochondrial DNA sequences from the 12S and 16S rRNA genes and the ND4 protein coding gene. More recntly, nuclear gene sequences (i.e., c-myc) have been collected by graduate student Bill McJilton. This research has emphasized inferring the relationships among the "lizard" families, as well as determining the phylogenetic placement of the major limbless squamate clades (snakes, amphisbaenians, and dibamids). There are also several extinct squamate taxa known only from the fossil record, with their phylogenetic placement also uncertain. In collaboration with Michael Lee (South Australia Museum), we are currently compiling this extensive molecular and morphological data base for a combined analysis squamate phylogeny.

Skink phylogeny and the evolution of limb reduction:

Phylogeny of the Sphenomorphus group.--The Sphenomorphus group is cosmopolitan and contains most of the diversity of lygosomine skinks. Most of the species diversity is found within Australia. The Australian Sphenomorphus group is a diverse skink clade (200+ species), composed of many taxa (e.g., Anomalopus, Lerista) possessing limbed and limbless species, as well as many species exhibiting various intermediate conditions towards limblessness. Therefore, this is an ideal group for which to study the evolution of limb reduction from a phylogenetic perspective. Many changes in morphology and life history are postulated to be correlated with limb reduction (e.g., body elongation precedes limb reduction, loss of external ear, reproductive mode). The first objective of this project is to infer a phylogeny for most of the members of the Australian Sphenomorphus group. Finally, within the context of a phylogeny, modern comparative methods can be used to test various hypotheses involving the evolution of limb reduction. Parts of this research was supported by NSF grant DEB-9707428. While much of my attention has been on the Australian subgroup, I have more recently starting examining more non-Australian species in order to more rigorously test the monophyly of the Australia group and to get a more global perspective on the evolution of this group.

Phylogenetic relationships and species limits within "Glaphyromorphus." This paraphyletic assemblage is a member of the Australian Sphenomorphus group. Most of the species occur in northern Australia, with a single species ("G". gracilipes) occurring in southwest Australia. The southwestern species appears to be more closely related to Hemiergis (Jennifer Reichet, grad student). However, even the northern assemblage does not appear to be monophyletic. All species are being examined in an attempt to determine the clades within this assemblage and determine which non-"Glaphyromorphus" taxa are their closest relatives. Also, the widespread "G". isolepis is likely a species complex. Fieldwork in the Kimberley (NW Australia) was conducted in January '00 in order to begin the dense sampling across it's range that will be needed in order to further examine this problem.

In collaboration with John Wiens (Stony Brook University), we have recently begun research to resolve the basal relationships within the Scincidae. Most of this work involves inferring the phylogeny among "scincines" and determining which other major non-"scincine" lineages (e.g., Acontias, lygosomines) have been derived from this basal assemblage. Also of interest is the fact that many "scincine" groups exhibit limb reduction. Thus, the resulting phylogeny will compliment our ongoing studies of limb reduction in other squamate clades.

Information for Prospective Graduate Students:

I am interested in highly motivated and independent graduate students who wish to conduct research addressing questions in the population and evolutionary biology of amphibians and reptiles, particularly if the questions compliment ongoing research in the lab. For evolutionary or systematic projects, students may employ molecular and/or morphological techniques for data collection. For population biology oriented problems, the student should be interested in using molecular markers to address their questions.

M.S. Students.—Students wishing to obtain an M.S. degree should apply to do so through the Evolutionary Biology Program Area.

Ph.D. Students.—The Department of Biology at SDSU and the University of California, Davis jointly offers a Ph.D in Ecology. I am interested in having doctoral students through this program, if they are interested in conducting molecular ecology/population biology oriented projects. Our department currently does not have a Ph.D. program in evolutionary biology. However, the Evolutionary Biology Program Area is currently in the process of establishing a joint program with the University of California, Riverside. It is hoped that this program will be ready to admit students within a couple years (target date of Fall 2009).

Publications:

Wiens, J. J., C. A. Kuczynski, S. A. Smith, D. Mulcahy, J. W. Sites, T. M. Townsend, and T. W. Reeder. Branch length, support, and congruence: Testing the phylogenomic approach with 20 nuclear loci in snakes. Systematic Biology, submitted.

Zaldivar-Riverón, A., A. Nieto Montes de Oca, N. Manríquez-Morán, and T. W. Reeder. Support of Anelytropsis (Reptilia: Squamata) within Dibamidae based on 16S rDNA sequence evidence. Journal of Herpetology, submitted.

Wood, D. A., R. N. Fisher, and T. W. Reeder. Novel patterns of historical isolation, dispersal, and secondary contact across Baja California in the rosy boa (Lichanura trivirgata). Molecular Phylogenetics and Evolution, submitted.

Townsend, T. M., E. R. Alegre, S. T. Kelley, J. J. Wiens, and T. W. Reeder. Rapid development of multiple nuclear loci for phylogenetic analysis using genomic resources: An example from the squamate reptile Tree of Life project. Molecular Phylogenetics and Evolution, accepted.

de Quieroz, K. and T. W. Reeder. 2007. Squamata – Lizards. In B. I. Crother (ed.), Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, pp. #-#. SSAR Herpetological Circular #, in press.

Crawford, N. G., J. Zalidvar-Rae, C. Hagen, A. Schable, E. B. Rosenblum, M. G. Ritchie T. W. Reeder, J. A. Graves, and T. C. Glenn. 2007. Thirteen polymorphic microsatellite DNA loci from whiptails of the genus Aspidoscelis and related cnemidophorine lizards. Molecular Ecology Notes, in press.

Wiens, J. J., C. A. Kuczynski, W. E. Duellman, and T. W. Reeder. 2007. Loss and re-evolution of complex life cycles in marsupial frogs: Does ancestral trait reconstruction mislead? Evolution 61:1886–1899.

Smith, S. A., A. Nieto Montes de Oca, T. W. Reeder, and J. J. Wiens. 2007. A phylogenetic perspective on elevational species richness patterns in Middle American treefrogs: why so few species in lowland tropical rainforests? Evolution 61:1188– 1207.

Wiens, J. J., C. H. Graham, D. S. Moen, S. A. Smith, and T. W. Reeder. 2006. Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity. American Naturalist 168:579–596.

Wiens, J. J., M. C. Brandley, and T. W. Reeder. 2006. Why does a trait evolve multiple times within a clade? Repeated evolution of snake-like body form in squamate reptiles. Evolution 60:123–141.

Wiens, J. J., T. W. Reeder, J. W. Fetzner, C. L. Parkinson, and W. E. Duellman. 2005. Hylid frog phylogeny and sampling strategies for speciose clades. Systematic Biology 54:778–807.

Reeder, T. W., and C. J. Cole. 2005. Letter to the Editor: Aspidoscelis versus Cnemidophorus as a genus of whiptail lizards in North America. Herpetological Review 36:233–234.

Brandley, M. C., A. Schmitz, and T. W. Reeder. 2005. Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards. Systematic Biology 54:373–390.

Fyler, C. A., T. W. Reeder, A. Berta, G. Antonelis, A. Aguilar, and E. Androukaki. 2005. Historical biogeography and phylogeny of monachine seals (Pinnipedia: Phocidae) based on mitochondrial and nuclear DNA data. Journal of Biogeography 32:1267–1279.

Rychel, A. L., T. W. Reeder, and A. Berta. 2005. Response to Ulfur Arnason: Where’s the beef? Molecular Phylogenetics and Evolution 35:311–312.

Schmitz, A., M. C. Brandley, P. Mausfeld, M. Vences, F. Glaw, R. A. Nussbaum, and T. W. Reeder. 2005. Opening the black box: Phylogenetics and morphological evolution of the Malagasy fossorial lizards of the subfamily “Scincinae”. Molecular Phylogenetics andEvolution 34:118–133.

Gergus, E., T. W. Reeder, and B. K. Sullivan. 2004. Geographic variation in Hyla wrightorum: Advertisement calls, allozymes, mtDNA, and morphology. Copeia 2004:758–769.

Rychel, A. L., T. W. Reeder, and A. Berta. 2004. Phylogeny of mysticete whales based on mitochondrial and nuclear data. Molecular Phylogenetics and Evolution 32:892–901.

Lee, M. S. Y., T. W. Reeder, J. B. Slowinski, and R. Lawson. 2004. Resolving reptile relationships: Molecular and morphological markers. In: Cracraft, J. and M. J. Donoghue (Eds.), Assembling the Tree of Life. Oxford University Press, NY, pp. 451–467.

Crother, B. I., J. Boundy, J. A. Camp bell , K. de Queiroz, D. R. Frost, D. M. Green, R. Highton, J. B. Iverson, P. A. Meylan, T. W. Reeder, M. E. Seidel, J. W. Sites, Jr., T. W. Taggart, S. G. Tilley, and D. B. Wake. 2003. Scientific and standard English names of amphibians and reptiles of North America north of Mexico : An update. Herpetological Review 34:196–203.

Reeder,. T. W. 2003. A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): A Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Molecular Phylogentics and Evolution 27:384-397. PDF file

Richmond, J. Q., and T. W. Reeder. 2002. Evidence for parallel ecological speciation in scincid lizards of the Eumeces skiltonianus species group (Squamata: Scincidae). Evolution 56:1498-1513. PDF file

Leache, A. D., and T. W. Reeder. 2002. Molecular systematics of the Eastern Fence Lizard (Sceloporus undulatus): A comparison of parsimony, likelihood, and Bayesian approaches. Systematic Biology 51:44-68. PDF file

Reeder, T. W., C. J. Cole, and H. C. Dessauer. 2002. Phylogenetic relationships of whiptail lizards of the genus Cnemidophorus (Squamata: Teiidae): A test of monophyly, reevaluation of karyotypic evolution, and review of hybrid origins. American Museum Novitates 3365:1-61. data set PDF file

Reeder, T. W., and R. R. Montanucci. 2001. A phylogenetic analysis of the horned lizards (Phrynosomatidae: Phrynosoma): Evidence from mitochondrial DNA and morphology. Copeia 2001:309-323. PDF file Phrynosoma data set

Crother, B. I., J. Boundy, J. A. Campbell, K. de Queiroz, D. R. Frost, R. Highton, J. B. Iverson, P. A. Meylan, T. W. Reeder, M. E. Seidel, J. W. Sites, Jr., T. W. Taggart, S. G. Tilley, and D. B. Wake. 2000. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding the confidence in our understanding. Herpetological Circular No. 29, pp.1-82. PDF file

Wiens, J. J., T. W. Reeder, and A. Nieto Montes De Oca, 1999. Molecular phylogenetics and evolution of sexual dichromatism among populations of the Yarrow's spiny lizard (Sceloporus jarrovii). Evolution 53:1884-1897. PDF file 1) S. jarrovii range map figure, 2) S. jarrovii phylogeny, 3) S. jarrovi data set

Wiens, J. J., and T. W. Reeder. 1997. Phylogeny of the spiny lizards (Sceloporus) based on molecular and morphological evidence. Herpetological Monographs 11:1-101. Sceloporus data set

Dessauer, H. C., T. W. Reeder, C. J. Cole, and A. Knight. 1996. Rapid screening of DNA diversity using dot-blot technology and allele-specific oligonucleotides: Maternity of hybrids and unisexual clones of hybrid origin (Lizards, Cnemidophorus). Molecular Phylogenetics and Evolution 6:366-372. PDF file

Reeder, T. W., and J. J. Wiens. 1996. Evolution of the lizard family Phrynosomatidae as inferred from diverse types of data. Herpetological Monographs 10:43-84.

Reeder, T. W. 1996. A new species of Pholidobolus (Squamata: Gymnophthalmidae) from the Huancabamba Depression of northern Peru. Herpetologica 52:282-289.

Wiens, J. J., and T. W. Reeder. 1995. Combining data sets with different numbers of taxa for phylogenetic analysis. Systematic Biology 44:548-558. PDF file

Reeder, T. W. 1995. Phylogenetic relationships among phrynosomatid lizards as inferred from mitochondrial ribosomal DNA sequences: Substitutional bias and the information content of transitions relative to transversions. Molecular Phylogenetics and Evolution 4: 203-222. PDF file

Simmons, A. D., J. L. Longmire, T. W. Reeder, H. A. Wichman, and R. J. Baker. 1992. Restriction fragment lenght polymorphisms in heterochromatic DNA distinguish chromosomal races of Peromyscus leucopus. Molecular Ecology 1:251-254.

Reeder, T. W. 1990. Eumeces managuae. Catalogue of American Amphibians and Reptiles (467):1-2.

Wichman, H. A., C. T. Payne, and T. W. Reeder. 1990. Intrageneric variation in repetitive sequences isolated by phylogenetic screening of mammalian genomes. In: "Molecular Evolution" (M.T. Clegg, and S. J. O'Brien, Eds.), pp. 153-160, Alan R. Liss, New York.

Current M.S. Graduate Students:

Michael Anguiana, started 1/04. “Effects of fragmentation on snake communities and spatial ecology of the California kingsnake (Lampropeltis getula californiae).” Co-advised with Dr. Jay Diffendorfer.

Ben Lowe, started 1/05. Phylogeography of the Long-tailed Brush Lizard (Urosaurus graciosus).

Anny Peralta, started 1/07. Project not completely defined as yet, but will involve evolutionary biology questions of Mexican amphibians and/or reptiles.

Angela Marion, started 8/07

Current Ph.D. Graduate Students:

Dean Leavitt, started 8/04. SDSU/University of California-Davis joint doctoral program in Ecology.

Current Undergraduate Students in the Lab:

Andrew Schlossman

Aleili Carmachi

The Dearly Departed (aka Former M.S. Graduate Students):

Adam Leache, 5/00. "Phylogenetic relationships within the Sceloporus undulatus species group (Squamata: Phrynosomatidae) based on mitochondrial DNA sequence data." Presently, University of California-Berkeley PhD student.

William McJilton, 5/00. "Phylogenetic analysis of Amniota using the single copy nuclear genes c-myc and c-mos." Presently, biology instructor at San Diego Community College.

Jonathan Richmond, 5/00. "Evolution of the Eumeces skiltonianus species group and intraspecific phylogeography of Eumeces gilberti." Obtained PhD at University of Connecticut (2006). Presently, post-doctoral associate at Cornell University.

Brian Yang, 5/01. "Molecular phylogenetics and evolution of the Malagasy-Comoroan phyllodactyle geckos, Genus Paroedura." Presently, Environmental Management Specialist, Department of Parks and Recreation, County of San Diego.

Jennifer Reichert, 12/01. "Molecular systematics of the limb-reduced Australian scincid lizards of the Genus Hemiergis." Presently, SDSU/UCSD joint PhD student.

Darren Burton, 5/02 (co-advised). "Phylogeny, evolution of morphology, and taxonomic implications for the genus Dudley (Crassulaceae) based on nrDNA ITS sequence data." Presently, full-time employee in biotech.

Dustin Wood, 5/02. "Intraspecific phylogeny of the rosy boa (Charina trivirgata): Implications for phylogeography, taxonomy, and conservation.". Presently, United States Geological Survey (USGS).

Amanda Rychel, 12/02 (co-advised). "Molecular phylogeny of the Mysticeti (Mammalia: Cetacea) based on mitochondrial and nuclear sequence data." Presently, University of Washington PhD student.

Lars Bell, 5/03. "Maternal ancestry and phylogeography of the unisexual Plateau Striped Whiptail (Aspidoscelis velox)."

Carrie Fyler, 12/03 (co-advised). “Molecular phylogeny of monachine seals (Pinnipedia: Phocidae) with implications for their origin and diversification.” Presently, University of Connecticut PhD student.

Andy Pece, 5/04. “Phylogeography of the Sidewinder (Crotalus cerastes), with implications for the historical biogeography of southwestern North American deserts.”

Matt Brandley, 5/05. “Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards.” Presently, University of California-Berkeley PhD student.

Nathan Marshal, 12/06. “Phylogeography of the Western Whiptail (Aspidoscelis tigris): “The phylogenetic affinities of A. t.maxima and its implications for the historical biogeography of southern Baja California.”

Nick Crawford, 8/07 "Microsatellites in cnemidophorine lizards: Their utility in investigating the landscape genetics of the Plateau Striped Whiptail (Aspidoscelis velox complex)." Presently, Boston University PhD student.

Mersee Madison-Villar, 8/07. Fine scale population genetic structure of the Western Spadefoot (Spea hammondi) in southern California. Presently, University of Texas-Arlington PhD student.

Courses Taught:

Natural History of Vertebrates (Biol 472)

Molecular Methods in Ecology & Evolution (Biol 596)

Phylogenetic Systematics (Biol 740)

Herpetology (Biol 523)

Organismal Biology (Biol 201B; vertebrate section)

Evolution (Bio 319; undergraduate non-majors)

Seminars Taught:

Molecular Systematics (Bio 770)

Systematics and Evolution (Bio 770)

Conservation Genetics (Bio 600)