The Phylogeny of the Gasterosteidae: Combining Behavioral and Morphological Data Sets

Abstract

Genealogical relationships among the five genera of stick-the subject of intense experimental and theoretical inleback (Gasterosteidae: Smegmomorpha) fishes were vestigation for more than 60 years (Pelkwijk and Tinreconstructed based upon 47 behavioral and 89 morpho- bergen, 1937;Wootton, 1976Wootton, , 1984, and references logical characters. Phylogenetic systematic analysis of therein), which has led to the accumulation of an extenthe combined data set produced a single MPT (consis-sive behavioral, morphological, and ecological datatency index of 82.54%) structured (Spinachia(Apeltesbase. The size of the database notwithstanding, at-((Pungitius ؉ Culaea)(Gasterosteus aculeatus ؉ G.tempts to determine the genealogical relationships wheatlandi)))). When analyzed separately, both behavwithin the family have had a volatile history. All auioral and morphological data sets produced the same thors agree that the Gasterosteidae comprises five gentree topology, although the consistency index (excluding era residing in north temperate habitats. Spinachia, uninformative characters) was higher for the behavioral Apeltes, and Culaea are monotypic and geographically (90.70%) than for the morphological (68.13%) characrestricted. Spinachia spinachia inhabits the shallow ters. This total evidence analysis provides a highly robust coastal waters of Western Europe, Apeltes quadracus is estimate of phylogeny for the sticklebacks and reiterates a brackish-water species distributed across the eastern the importance of discarding a priori biases about behavcoast of North America from the Gaspé Basin of Queioral evolution when selecting characters for phylogebec to Virginia, and Culaea inconstans is restricted to netic analysis. ᭧ 2001 The Willi Hennig Society freshwaters in the heartland of North America. Pungitius and Gasterosteus, on the other hand, are both geographically widespread and morphologically variable. Sticklebacks (Gasterosteidae) are small, bony plated At least three species of Pungitius are recognized, P. neoteleost fishes renowned for their intricate courtship pungitius (the ninespine stickleback), P. tymensis (redances and vibrant nuptial coloration. They have been stricted to the islands of Sakhalin and Hokkaido: Takata et al., 1984), and P. hellenicus (restricted to three sites in the Sperchios drainage, Greece: Keivany et al.,

Total evidence phylogeny of Gasterosteidae: combining molecular, morphological and behavioral data

“…Specimens from the Royal Ontario Museum were examined under a Zeiss SV8 stereo-zoom dissecting microscope and character states assigned under the expert guidance of R. Winterbottom (see Appendix 1 for accession numbers). This examination revealed that G. wheatlandi showed the plesiomorphic condition for the following characters listed in McLennan and Mattern (2001): character 6 (broad horizontal median plate on median ethmoid) and character 47 (posterolateral lamina over scapular foramen in cleithra absent). Gasterosteus wheatlandi had the apomorphic condition in two characters, character 10 (interorbital commissures in frontals absent) and character 36 (preoperculo-mandibular canals in dentary absent).…”

“…Bowne did not examine the osteology of Gasterosteus wheatlandi, so this created many missing entries in the data set. In this study we examine the state for four characters that appear to be synapomorphies for Gasterosteus (McLennan and Mattern, 2001), based upon computer extrapolation from missing character codes for G. wheatlandi. Specimens from the Royal Ontario Museum were examined under a Zeiss SV8 stereo-zoom dissecting microscope and character states assigned under the expert guidance of R. Winterbottom (see Appendix 1 for accession numbers).…”

“…Given the importance of these fishes, it is imperative that the ecological, evolutionary and physiological information collected over the years be examined, whenever possible, within a well-supported phylogenetic framework in order to construct rigorous hypotheses of character evolution (see References in Brooks and McLennan, 2002). McLennan and Mattern (2001) presented one such phylogenetic hypothesis. A phylogenetic analysis of 89 morphological and 47 behavioral characters identified a single hypothesis of 189 steps and a consistency index of 0.825 structured (Spinachia (Apeltes (Gasterosteus (Pungitius, Culaea)))) (see McLennan and Mattern, 2001 for a review of systematic studies to that point).…”

“…McLennan and Mattern (2001) presented one such phylogenetic hypothesis. A phylogenetic analysis of 89 morphological and 47 behavioral characters identified a single hypothesis of 189 steps and a consistency index of 0.825 structured (Spinachia (Apeltes (Gasterosteus (Pungitius, Culaea)))) (see McLennan and Mattern, 2001 for a review of systematic studies to that point). Mattern (2004) sequenced 2879 base pairs from five mitochondrial genes, 12S and 16S rRNA, cytochrome b, ATPase 6 and control region.…”

A receiver bias in the origin of three–spined stickleback mate choice

“…Thus, the degree of red coloration of male sticklebacks appears to signal both direct and indirect benefits to females, at least in those populations where correlations exist. The stickleback family (Gasterosteidae) is relatively small, comprising only five genera (McLennan & Mattern 2001). Not all stickleback species exhibit distinctive nuptial coloration, but it is…”