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Portraits of Evolution: Studies of Coloration in Hawaiian Spiders Author(S): GEOFF S

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Portraits of Evolution: Studies of Coloration in Hawaiian Spiders Author(S): GEOFF S Portraits of Evolution: Studies of Coloration in Hawaiian Spiders Author(s): GEOFF S. OXFORD and ROSEMARY G. GILLESPIE Source: BioScience, 51(7):521-528. 2001. Published By: American Institute of Biological Sciences DOI: http://dx.doi.org/10.1641/0006-3568(2001)051[0521:POESOC]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1641/0006-3568%282001%29051%5B0521%3APOESOC %5D2.0.CO%3B2 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Articles Portraits of Evolution: Studies of Coloration in Hawaiian Spiders GEOFF S. OXFORD AND ROSEMARY G. GILLESPIE olor variation, when genetically determined, Cprovides a visual tool with which to investigate natural THE DISCRETE COLOR POLYMORPHISMS selection. Here we examine how color variation in two spi- der systems in the Hawaiian Islands can be used to understand IN SPIDERS ALLOW THE STUDY OF evolutionary phenomena at both population and species lev- els. First, the happy-face spider (Theridion grallator) shows a EVOLUTION “IN ACTION” number of discrete color morphs, the frequencies of which are very similar among populations, although the genetic basis for them differs between islands. These observations permit Transient polymorphisms are those that are not maintained intriguing insights into the evolution and maintenance of color in a balanced state over extended time periods. Perhaps the polymorphism. Second, we consider the adaptive radiation of best-known situation is when one color morph replaces an- Hawaiian Tetragnatha spiders. Here the spectacularly diverse other over time (which may be comparatively short, de- array of color forms is found within closely related species on pending on the relative advantage of the favored allele) as a each island, with similar sets of colors evolving indepen- result of directional selection. In other situations, one ances- dently in each locale, suggesting selection is acting to main- tral color morph can give rise to two or more descendant color tain such diversity. morphs as a result of disruptive selection, a phenomenon well Genetically determined color variation provides an im- known in adaptive radiations. For example, populations may mediately tangible link between a genotype and an exter- diverge in allopatry (that is, when geographically separate from nally expressed phenotype and, as a result, has long been each other), with disruptive selection acting on color variants used to probe evolutionary questions. In insects generally, and when the two ecologically similar color forms regain sympa- in Lepidoptera in particular, studies of color and pattern try (overlap geographically), as has been observed for mor- have contributed major advances to our understanding of a phological characters in Anolis lizards (Losos et al. 1998) and number of key evolutionary phenomena. Classical studies in- cichlid fish (Ruber et al. 1999). Alternatively, selection on a sin- clude those on the nature and evolution of dominance (Ford gle population for adaptation to different habitats (or mi- 1975), the power and mode of action of natural selection (Ket- croniches within a habitat) within a particular locality can lead tlewell 1973), the evolution of mimetic resemblances (Turner 1970), and the developmental rules underlying differences in patterns within and among species (Nijhout 1991). Geoff Oxford (e-mail: [email protected]) is on the faculty in the Color polymorphism is variation that is determined ge- Department of Biology, University of York, York YO10 5YW, UK, and netically at a small number of major loci (Ford 1940) and is has been utilizing spiders as model organisms for many years. With classically divided into two categories: balanced and tran- them he has studied, among other things, the determination of sient. In balanced polymorphisms, morph frequencies can re- sperm priority patterns, speciation and hybridization, and the main constant over long periods of time, usually as a result evolutionary significance of color. Rosemary Gillespie (e-mail: of an equilibrium between advantages and disadvantages of [email protected]) currently holds the Schlinger Chair of the different morphs. Indeed, balanced polymorphisms often Systematics and is director of the Essig Museum of Entomology at transcend species boundaries, for example, the color and the University of California, Berkeley, CA 94720. She focuses on pattern variation in the spiders Enoplognatha ovata and E. la- patterns of diversification on remote oceanic islands of the Pacific. timana (Oxford and Reillo 1993). © 2001 American Institute of Biological Sciences. July 2001 / Vol. 51 No. 7 • BioScience 521 Articles to polymorphism (Mather 1955). If disruptive selection is the basis for many of the advances in our understanding of strong, the subpopulations of the different habitat-specific ecological genetics made during the 20th century. The Hawai- color morphs may cease to exchange genes and evolve sym- ian Islands offer an ideal opportunity for studying the main- patrically into separate species (Thoday and Gibson 1962). In tenance and adaptive significance of color variation within spi- either case, color polymorphism is implicated in the process, der populations, as well as the interaction between genetic drift and its existence may be identified by examining the evolu- and natural selection. These aspects have been studied in tionary end products. some detail in the endemic Hawaiian happy-face spider Consider coloration in spiders. Like many other arthropods, Theridion grallator (Araneae, Theridiidae). The species occurs spiders display a wide range of colors and patterns that vary on the islands of Oahu (largest population to date known from both inter- and intraspecifically. As a group they have been Mt. Kaala summit, elevation 1220 m), Molokai (largest pop- relatively neglected, until recently, as possible models with ulation in Kamakou Preserve, elevation 1110 m), Maui (largest which to study evolutionary processes. However, investigations population in Waikamoi Preserve, Haleakala, elevation 1360 of spider coloration have a long and distinguished history. For m; other populations on Puu Kukui, West Maui, elevation 1387 example, one of the very early studies of sexual selection was m, and in Auwahi on the south slope of Haleakala, elevation based on the sexual dimorphism for color and pattern ex- 1370 m), and Hawaii (populations in the Kohala mountains, hibited by many jumping spiders (Salticidae; Peckham and 1152 m; the Saddle, 1600 m; and Kilauea, 1190 m). These are Peckham 1889, 1890). This work served to underline the all sites of native Hawaiian wet-to-mesic forest, with a canopy principles espoused in Charles Darwin’s book on the subject generally dominated by Metrosideros polymorpha (Myrtaceae) published just a few years earlier (Darwin 1871). Perhaps and Acacia koa (Leguminosae), and the subcanopy by Brous- the most common explanation for many of the color patterns saisia arguta (Saxifragaceae), Clermontia arborescens (Cam- exhibited by spiders is for avoidance of predation (Foelix panulaceae), Cheirodendron trigynum (Araliaceae), Coprosma 1982). Spiders can use color to hide from (crypsis) or confuse sp. (Rubiaceae), Ilex anomolum (Aquifoliaceae), Myrsine spp. predators (Oxford and Gillespie 1998); here color is generally (Myrsinaceae), and Pelea spp. (Rutaceae). Populations of T. associated with a specific habitat. Therefore, the existence of grallator are patchily distributed, and at most sites they occur discrete color polymorphisms makes it possible to elucidate at very low densities. the selective forces acting on color and hence the evolution The spiders build very small, flimsy webs consisting of a of ecological affinity. scanty, two-dimensional layer of silk covering the entire un- Two different selective forces, selection for balanced poly- derside of a leaf. During the day, the spiders are usually tightly morphism and disruptive selection acting on transient poly- appressed to the underside of the leaf, but at night they of- morphisms, are well illustrated by spiders in the Hawaiian Is- ten hang well below the leaf from silk threads. They capture lands. Islands have long been recognized for their importance insects on the underside of a leaf and feed in situ. The most in studies of the genetic differentiation of populations and the common prey items are small adult flies of the families processes involved in speciation (Darwin 1859). Remote Dolichopodidae and Drosophilidae (Gon 1985). Upon mat- oceanic archipelagos, in particular the Hawaiian Islands, pro- uration, a male is often found sharing a leaf with a penulti- vide opportunities to observe the effects of colonization,
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