Research

Research  Spiders — Phylogeny of salticid spiders ♦ Biodiversity discovery ♦ Salticid evolution & biogeography ♦ Evolution and behaviour of Habronattus ♦ Other spiders ♦ ♦ Phylogeny — Character evolution ♦ Diversification ♦ Phylogeography ♦ Computation and informatics ♦ Other phylogenetic projects

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Spider Systematics and Evolution

The family of jumping spiders (Salticidae) includes more than 5000 species of varied body forms and habits. Their unique high-resolution eyes permit visually-mediated predatory and courtship behaviour. Their best vertebrate analogs as predators are perhaps the cats — sighting prey from a distance, stalking, and pouncing. In courtship behavior, some groups such as Habronattus rival the birds of paradise in ornaments and behavioural complexity.

Phylogeny & systematics of salticid spiders — The handful of systematists who have worked on salticid phylogenetics over the last century have begun to resolve the outlines of groups, but considerable work remains. A better resolved phylogeny and the corresponding classification of subfamilies and genera would facilitate efforts to discover new species and distinguish among them, and would also open the group to phylogenetically-based studies of evolutionary processes. A long term project of the laboratory is to reconstruct the phylogenetic structure of the family using both molecular and morphological data. Recent sampling from South America, Asia, Africa and Australasia have yielded a reasonably complete outline of the phylogeny, but continued sampling and data from more genes are needed to resolve the phylogeny more completely.

  • Maddison WP. 2015. A phylogenetic classification of jumping spiders (Araneae: Salticidae). Journal of Arachnology 43: 231-292.
  • Ruiz GRS, Maddison WP. 2015. The new Andean jumping spider genus Urupuyu and its placement within a revised classification of the Amycoida (Araneae: Salticidae). Zootaxa 4040: 251–279.
  • Bustamante AA, Maddison WP, Ruiz GRS. 2015. The jumping spider genus Thiodina Simon, 1900 reinterpreted, and revalidation of Colonus F.O.P-Cambridge, 1901 and Nilakantha Peckham & Peckham, 1901 (Araneae: Salticidae: Amycoida). Zootaxa 4012: 181–190.
  • Zhang JX & Maddison WP. 2015. Genera of euophryine jumping spiders (Araneae: Salticidae), with a combined molecular-morphological phylogeny. Zootaxa 3938: 1–147.
  • Maddison WP, Li D, Bodner M, Zhang JX, Xu X, Liu Q, Liu F. 2014. The deep phylogeny of jumping spiders (Araneae, Salticidae). Zookeys 440: 57–87.
  • Zhang JX & Maddison WP. 2013. Molecular phylogeny, divergence times and biogeography of spiders of the subfamily Euophryinae (Araneae: Salticidae). Molecular Phylogenetics and Evolution. 68: 81–92.
  • Bodner MR & Maddison WP. 2012. The biogeography and age of salticid spider radiations (Araneae: Salticidae). Molecular Phylogenetics and Evolution. 65: 213-240.
  • Maddison, W., M. Bodner and K. Needham. 2008. Salticid spider phylogeny revisited, with the discovery of a large Australasian clade (Araneae: Salticidae). Zootaxa. 1893: 49–64.
  • Maddison, W.P., J.X. Zhang, & M.R. Bodner. 2007. A basal phylogenetic placement for the salticid spider Eupoa, with descriptions of two new species (Araneae: Salticidae). Zootaxa 1432: 23–33.
  • Maddison, W.P. and K. Needham. 2006. Lapsiines and hisponines as phylogenetically basal salticid spiders (Araneae: Salticidae). Zootaxa. 1255:37-55.
  • Maddison, W. P. & M. Hedin. 2003. Jumping spider phylogeny (Araneae: Salticidae). Invertebrate Systematics. 17: 529-549.
  • Hedin, M.C. and W.P. Maddison. 2001. A combined molecular approach to phylogeny of the jumping spider subfamily Dendryphantinae (Araneae, Salticidae). Molecular Phylogenetics and Evolution. 18: 386-403.


Biodiversity discovery: taxonomy of salticid spiders — We have discovered only a small fraction of the species on Earth. The field of taxonomy continues the age of discovery by collecting and describing new species. We have contributed to this effort through expeditions to Ecuador, S.E. Asia, Gabon, Papua New Guinea, Borneo, and Mexico. A public lecture describing an expedition and my perspective on biodiversity discovery can be found online.

  • Ruiz GRS, Maddison WP. 2015. The new Andean jumping spider genus Urupuyu and its placement within a revised classification of the Amycoida (Araneae: Salticidae). Zootaxa 4040: 251–279.
  • Maddison WP & Piascik EK. 2014. Jerzego, a new hisponine jumping spider from Borneo (Araneae: Salticidae). Zootaxa 3852: 569–578.
  • Zhang JX & Maddison WP. 2014. Tisaniba, a new genus of marpissoid jumping spiders from Borneo (Araneae: Salticidae). Zootaxa 3852: 252–272.
  • Zhang JX & Maddison WP. 2012. New euophryine jumping spiders from the Southeast Asia and Africa (Araneae: Salticidae: Euophryinae). Zootaxa. 3581: 53–80.
  • Zhang JX & Maddison WP. 2012. New euophryine jumping spiders from Central America and South America (Araneae: Salticidae: Euophryinae). Zootaxa. 3578: 1–35.
  • Zhang JX & Maddison WP. 2012. New euophryine jumping spiders from Papua New Guinea (Araneae: Salticidae: Euophryinae). Zootaxa. 3491: 1–74.
  • Zhang JX & Maddison WP. 2012. New euophryine jumping spiders from the Dominican Republic and Puerto Rico (Araneae: Salticidae: Euophryinae). Zootaxa. 3476: 1–54.
  • Maddison WP. 2012. Five new species of lapsiine jumping spiders from Ecuador (Araneae: Salticidae). Zootaxa. 3424: 51-65.
  • Ruiz G & Maddison WP. 2012. DNA sequences corroborate Soesiladeepakius as a non-salticoid genus of jumping spiders: placement with lapsiines, phylogeny and description of six new species (Araneae, Salticidae). Zoological Journal of the Linnean Society. 165: 274–295.
  • Maddison, W.P. 2009. New cocalodine jumping spiders from Papua New Guinea (Araneae: Salticidae: Cocalodinae). Zootaxa. 2021: 1–22.
  • Maddison, W.P., J.X. Zhang, & M.R. Bodner. 2007. A basal phylogenetic placement for the salticid spider Eupoa, with descriptions of two new species (Araneae: Salticidae). Zootaxa 1432: 23–33.
  • Maddison, W.P. 2006. New lapsiine jumping spiders from Ecuador (Araneae: Salticidae). Zootaxa. 1255:17-28.
  • Maddison, W.P. 1996. Pelegrina and other jumping spiders formerly placed in the genus Metaphidippus (Araneae: Salticidae). Bulletin of the Museum of Comparative Zoology. l54(4): 215-368.

Salticid evolution & biogeography — Our molecular phylogeny of salticids has revealed a surprisingly strong biogeographical signal: major clades are mostly restricted to a continental region. The New World tropics are dominated (in species count) by clades that are nearly absent from the Old World. Similarly, there are major clades largely restricted to Australasia, others to Afro-eurasia. Each of these clades shows a diversity of body forms, suggesting replicate ecological radiations have occurred independently on different continents. This provides an opportunity to study adaptive radiation and community assembly on a scale much larger (many hundreds of species and among continents) than well-known island examples such as Anolis lizards. Our first question will be whether consistency or contingency dominates: do faunas on different continents converge to the same constellation of “ecomorphs”?

  • Bodner MR & Maddison WP. 2012. The biogeography and age of salticid spider radiations (Araneae: Salticidae). Molecular Phylogenetics and Evolution. 65: 213-240.
  • Maddison, W., M. Bodner and K. Needham. 2008. Salticid spider phylogeny revisited, with the discovery of a large Australasian clade (Araneae: Salticidae). Zootaxa. 1893: 49–64.
  • Maddison, W. P. & M. Hedin. 2003. Jumping spider phylogeny (Araneae: Salticidae). Invertebrate Systematics. 17: 529-549.

Evolution and behaviour of Habronattus — One genus of salticids, Habronattus, has been a special focus of our work. The approximately 100 described species, primarily North American, include many with complex courtship ornaments and behaviours. A few are shown here. At the fine scale, we have examined the effect of sexual selection on differentiation among isolated montane populations of H. pugillis. At the broad scale, we have reconstructed phylogeny among species. Our eventual goal is to use the phylogeny to answer questions of behavioural evolution (e.g., why do alternating asymmetrical motions arise frequently? why are visual signals synchronized to sound production?) and of chromosome evolution (e.g., what forces affect repeated evolution of X-autosomal fusions?). This project is done in collaboration with Damian Elias (University of California, Berkeley) and Marshal Hedin (San Diego State University).

  • Blackburn GS & Maddison WP. 2015. Insights to mating strategies of Habronattus americanus jumping spiders from natural behaviour and staged interactions in the wild. Behaviour 152: 1169-1186.
  • Blackburn, GS & Maddison WP. 2014. Stark sexual display divergence among jumping spider populations in the face of gene flow. Molecular Ecology 23: 5208-5223.
  • Maddison WP & Leduc-Robert G. 2013. Multiple origins of sex chromosome fusions correlated with chiasma localization in Habronattus jumping spiders (Araneae: Salticidae). Evolution. 67: 2258–2272.
  • Elias DO, Maddison WP, Peckmezian C, Cirard MB, & Mason AC. 2012. Orchestrating the score: Complex multimodal courtship in the H. coecatus group of Habronattus jumping spiders (Araneae: Salticidae). Biological Journal of the Linnaean Society. 105: 522–547.
  • Elias, D.O., E.A. Hebets, R.R. Hoy, W.P. Maddison & A.C. Mason. 2007. Regional song differences in sky-island populations of the jumping spider Habronattus pugillis Griswold. Journal of Arachnology. 34 (3): 545-556.
  • Hebets, E.A. and W.P. Maddison. 2005. Xenophilic mating preferences among populations of the jumping spider Habronattus pugillis Griswold. Behavioural Ecology 16:981–988.
  • Maddison, W.P. and M.C. Hedin. 2003. Phylogeny of Habronattus jumping spiders (Araneae: Salticidae), with consideration of genitalic and courtship evolution. Systematic Entomology 28:1-21.
  • Masta, S. and W.P. Maddison. 2002. Sexual selection driving diversification in jumping spiders. Proceedings of the National Academy of Sciences of the United States of America 99(7):4442-4447.
  • Maddison, W.P. and M.M. McMahon. 2000. Divergence and reticulation among montane populations of the jumping spider Habronattus pugillis Griswold. Systematic Biology 49:400-421.
  • Maddison, W.P. and G.E. Stratton. 1988. Sound production and associated morphology in male jumping spiders of the Habronattus agilis species group (Araneae: Salticidae). J. Arachnology 16: 199-211.
  • Maddison, W.P. 1982. XXXY sex chromosomes in males of the jumping spider genus Pellenes (Araneae: Salticidae). Chromosoma (Berlin) 85: 23-37.

Other spider projects — Mostly on social spiders, in collaboration with Leticia Avilés and Ingi Agnarsson.

  • Agnarsson I, Avilés L, & Maddison WP. 2013. Loss of genetic variability in social spiders: genetic and phylogenetic consequences of population subdivision and inbreeding. Journal of Evolutionary Biology. 26: 27-37.
  • Agnarsson I, Maddison WP, & Avilés L. 2010. Complete separation along matrilines in a social spider metapopulation inferred from hypervariable mitochondrial DNA region. Molecular Ecology. 19: 3052-3063
  • Muster C, Maddison WP, Uhlmann S, Berendonk TU, & Vogler AP. 2009. Arctic‐Alpine Distributions—Metapopulations on a Continental Scale? American Naturalist 173: 313–326
  • Agnarsson I, Maddison WP, & Avilés L. 2007. The phylogeny of the social Anelosimus spiders (Araneae: Theridiidae) inferred from six molecular loci and morphology. Molecular Phylogenetics and Evolution. 43: 833-851.
  • Agnarsson I, Avilés L, Coddington JA, & Maddison WP. 2006. Sociality in theridiid spiders — Repeated origins of an evolutionary dead-end. Evolution. 60: 2342-2351
  • Avilés L, Maddison WP, & Agnarsson I. 2006. A new independently-derived social spider with explosive colony proliferation and a female size dimorphism. Biotropica 38: 743-753.
  • Avilés L, Maddison WP, Salazar P, Estévez G, Tufiño P, & Cañas G.. 2001. Arañas sociales de la Amazonía ecuatoriana, con notas sobre seis especies sociales no descritas previamente. [Social spiders of the Ecuadorian Amazonia, with notes on previously undescribed social species.] Rev. Chil. Hist. Nat., 74:619-638.
  • Avilés L & Maddison WP. 1991. When is the sex ratio biased in social spiders?: Chromosome studies of embryos and male meiosis in Anelosimus species (Araneae: Theridiidae). J. Arachnology 19: 126-135.

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Phylogenetic Theory and Programming

Phylogeny, the broad-scale genetic history of life, is a central historical framework by which we can make interpretations about evolutionary processes. Repeated patterns of change among species can suggest underlying evolutionary forces and constraints. Abundant sequence data has allowed the historical approach to be brought to finer scale work examining genealogical history of individual gene loci within or among populations.

Character evolution— How knowledge of phylogeny can be used to infer the ancestral states of characters and correlations among characters. Selected papers are:

  • Maddison WP & FitzJohn RG. 2015. The unsolved challenge to phylogenetic correlation tests for categorical characters. Systematic Biology 64: 127–136.
  • Maddison, W.P. 2006. Confounding asymmetries in evolutionary diversification and character change. Evolution 60: 1743-1746.
  • Maddison, W.P. 2000. Testing character correlation using pairwise comparisons on a phylogeny. Journal of Theoretical Biology. 202: 195-204.
  • Maddison, W.P. 1995. Calculating the probability distributions of ancestral states reconstructed by parsimony on phylogenetic trees. Systematic Biology. 44:474-481.
  • Maddison, W.P. 1991. Squared-change parsimony reconstructions of ancestral states for continuous-valued characters on a phylogenetic tree. Systematic Zoology. 40: 304-314.
  • Maddison, W.P. 1990. A method for testing the correlated evolution of two binary characters: are gains or losses concentrated on certain branches of a phylogenetic tree?. Evolution 44: 539-557.
  • Maddison, W.P., M.J. Donoghue and D.R. Maddison. 1984. Outgroup analysis and parsimony. Syst. Zool. 33: 83-103.

Diversification— Methods to understand speciation and extinction using phylogeny.

  • Maddison WP & FitzJohn RG. 2015. The unsolved challenge to phylogenetic correlation tests for categorical characters. Systematic Biology 64: 127–136.
  • Davis MP, Midford PE, & Maddison WP. 2013. Exploring power and parameter estimation of the BiSSE method for analyzing species diversification. BMC Evolutionary Biology. 13:38.
  • FitzJohn RG, Maddison WP, & Otto SP. 2009. Estimating trait-dependent speciation and extinction rates from incomplete phylogenies. Systematic Biology. 58: 595-611.
  • Maddison, W.P., P.E. Midford & S.P. Otto. 2007. Estimating a binary character’s effect on speciation and extinction. Systematic Biology. 56:701-710
  • Maddison, W.P. 2006. Confounding asymmetries in evolutionary diversification and character change. Evolution 60: 1743-1746.
  • Agnarsson, I., L. Avilés, J.A. Coddington and W.P.Maddison. 2006. Sociality in theridiid spiders — Repeated origins of an evolutionary dead-end. Evolution. 60: 2342-2351.

Phylogeography — The “ultrastructure” of species phylogenies, and the histories of individual gene loci.

  • Maddison, W.P. and L.L. Knowles. 2006. Inferring phylogeny despite incomplete lineage sorting. Systematic Biology. 55(1):21–30.
  • Knowles, L.L. and W.P. Maddison. 2002. Statistical phylogeography. Molecular Ecology. 11:2623-2635.
  • Maddison, W.P. 1997. Gene trees in species trees. Systematic Biology 46:523-536.
  • Maddison, W.P. 1995. Phylogenetic histories within and among species. In: Experimental and molecular approaches to plant biosystematics (Hoch, P. C., and A. G. Stevenson, editors). Monographs in Systematics. Missouri Botanical Garden, St. Louis. 53: 273-287. [pdf]
  • Slatkin, M. and W. P. Maddison. 1989. A cladistic measure of gene flow inferred from the phylogeny of alleles. Genetics 123: 603-613.

Computation and informatics — Theoretical work introducing new methods is not fully available to the biological community until delivered as tools. In some cases, however, tools go far beyond simply completing basic calculations. Visualizations and exploratory data analysis tools can provoke imaginations and help shape the very concepts of the field. These principles have guided the the first three of the following projects, done in collaboration with David Maddison.

Mesquite project — an open-source project, actively in development. Its goal is to stimulate novel evolutionary analyses by permitting easy building and combining of modules. Its analyses include ancestral state reconstruction with parsimony and likelihood, tests of character correlation, diversification models, tree simulations and comparisons, morphometrics, and coalescence modelling.

MacClade — a widely-used program for exploring phylogenetic trees interactively. Recent versions have sophisticated DNA sequence editing capabilities.

  • Maddison DR & Maddison WP. 2000. MacClade version 4: Analysis of phylogeny and character evolution. Sinauer Associates, Sunderland Massachusetts. 
  • Maddison WP & Maddison DR. 1992. MacClade version 3: Analysis of phylogeny and character evolution. 398 pp (book) + 900K (computer program). Sinauer Associates, Sunderland Massachusetts.
  • Maddison WP & Maddison DR. 1987. MacClade version 2.1 (Grand Prize Winner of Apple Computer’s Wheels for the Mind competition).
  • Maddison WP 1986. MacClade version 1.0 (An interactive computer program for reconstructing phylogeny and analyzing character evolution, distributed by the author; reviewed by W.L. Fink, Science 234: 1135-1139, 28 November 1986).

Tree of Life web project — a collaborative presentation of Life’s phylogeny. Described here: Maddison, D.R., K. Schulz, & W.P. Maddison. 2007. The Tree of Life Web Project. Zootaxa. 1168: 19-40

CIPRes — Ciber-infrastructure for phylogenetic research. A large multi-institution collaboration between computer science and biology to enhance computational resources for phylogenetic analyses.

Other phylogenetic projects — The following are writings making general commentary on phylogenetic biology. The MacClade book has several chapters that amount to a partial review of phylogenetics as a field.

  • Mooers, A.Ø., D.P. Faith and W.P.Maddison. 2008. Converting endangered species categories to probabilities of extinction for phylogenetic conservation prioritization. PLoS ONE 3(11): e3700.
  • Maddison, W. and T.M. Pérez. 2001. Biodiversidad y lecciones de la historia. In: Enfoques contemporáneos para el estudio de la biodiversidad (Hernández, H.M., A. García Aldrete, F. Álvarez and M. Ulloa, editors). Instituto de Biología, UNAM, Mexico. Pp. 201-220. [pdf]
  • Maddison, W.P. 1996. Molecular approaches and the growth of phylogenetic biology. In: Molecular zoology: Advances, strategies, and protocols (J.D. Ferraris and S.R. Palumbi, eds.). Pages 47­63. Wiley-Liss, New York. [pdf]
  • Maddison, W.P. and D.R. Maddison. 1992. MacClade version 3: Analysis of phylogeny and character evolution. 398 pp (book) + 900K (computer program). Sinauer Associates, Sunderland Massachusetts.
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Recent Posts

2017 Expedition to México

Undescribed blue-legged Mexigonus from the mountains of Oaxaca

In mid 2017 we undertook a spectacularly-successful expedition to the hills and mountains of Oaxaca and Chiapas in Mexico (and a bit also to the state of Queretaro). I blogged live from the field to my personal blog, and have compiled all the posts into a report on the expedition. Enjoy!

  1. MacClade’s 30th Anniversary Leave a reply
  2. BIOL 336: Fundamentals of Evolutionary Biology Leave a reply
  3. 27000 images of jumping spiders Leave a reply
  4. A new phylogenetic classification of jumping spiders Leave a reply
  5. The new Andean jumping spider genus Urupuyu and a revised classification of the Amycoida Leave a reply
  6. Paper published: A reinterpretation of Thiodina 1 Reply
  7. Mesquite 3.04 released Leave a reply
  8. PechaKucha: Can spiders inspire art and design? 1 Reply
  9. Lorna Levi Leave a reply