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Kinorhyncha: Cyclorhagida) Martin V ARTICLE IN PRESS Organisms, Diversity & Evolution 8 (2008) 233–246 www.elsevier.de/ode Phylogenetic analysis of the Echinoderidae (Kinorhyncha: Cyclorhagida) Martin V. Sørensen Ancient DNA and Evolution Group, Department of Biology, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen, Denmark Received 10 July 2007; accepted 14 November 2007 Abstract The phylogeny of the kinorhynch family Echinoderidae is analyzed using morphological characters and parsimony as the optimization criterion. Thirty-six characters were coded in a matrix with eleven echinoderid terminals, representing all echinoderid genera and six non-echinoderid outgroup taxa. The ingroup includes Polacanthoderes martinezi and Cephalorhyncha liticola, newly described in a separate paper. The resulting most parsimonious trees support Polacanthoderes as the most basal echinoderid genus, followed by an unresolved clade with non-monophyletic Fissuroderes and monophyletic Cephalorhyncha and Echinoderes. r 2008 Gesellschaft fu¨ r Biologische Systematik. Published by Elsevier GmbH. All rights reserved. Keywords: Cephalorhyncha; Phylogeny; Morphology; Echinoderes; Fissuroderes; Polacanthoderes Introduction Echinodera were treated as synonymous with the name ‘‘Kinorhyncha’’ that had been introduced by Reinhard Even though the kinorhynchs have been known for a (1885, 1887) and now applies to the whole phylum. reasonable amount of time, the group’s internal Present-day kinorhynch taxonomy was founded by relationships remain a puzzle. Only few studies have Zelinka (1896) who discriminated between the two main dealt with kinorhynch phylogeny (Neuhaus 1994, 1995; clades, Homalorhagida and Cyclorhagida. Echinoderi- Adrianov and Malakhov 1996, 1999; Kristensen 2002; dae was assigned to the latter, and considered as closely Neuhaus and Higgins 2002), all of them based on related to Centroderidae Zelinka, 1896. Until the mid theoretical considerations and discussions; thus, kinor- 1920s, more than 70 years after the first kinorhynch had hynch relationships have never been analyzed using been recorded by Dujardin (1851), only few species had modern numerical methods. The present study repre- been formally described. Then the known kinorhynch sents the first attempt to understand kinorhynch biodiversity increased dramatically with the publication phylogeny using morphological data analyzed under a of Zelinka’s (1928) monumental ‘‘Monographie der cladistic approach. The analyses focus on the relation- Echinodera’’, in which he described a total of 48 new ships within the family Echinoderidae. species, and several new genera. In his classification, The family Echinoderidae was originally erected by Zelinka assigned four genera to Echinoderidae. Un- Bu¨ tschli (1876) to contain the single kinorhynch species, fortunately, two of these genera, Habroderes and Echinoderes dujardini Clapare` de, 1863. For some Habroderella, were mostly based on specimens that time, the derived higher-taxon names Echinoderida or have since turned out to be juveniles of Echinoderes,and most of the corresponding names are considered as E-mail address: [email protected]. nomina dubia today. The two species of the fourth genus 1439-6092/$ - see front matter r 2008 Gesellschaft fu¨ r Biologische Systematik. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.ode.2007.11.002 ARTICLE IN PRESS 234 M.V. Sørensen / Organisms, Diversity & Evolution 8 (2008) 233–246 recognized by Zelinka, Echinoderella, have been assigned 1999; and Zelinkaderes brightae Sørensen, Heiner, Ziemer to Echinoderes as well. Hence, the family Echinoderidae & Neuhaus, 2007. The following 11 echinoderid species, became monogeneric again for some time, containing representing all known echinoderid genera, were selected Echinoderes only (for further historic overview of as ingroup taxa: Polacanthoderes martinezi Sørensen, kinorhynch classification, see Higgins 1971). Recently, 2008; Cephalorhyncha liticola Sørensen, 2008; C. asiatica the genera Cephalorhyncha and Fissuroderes have been (Adrianov, 1989); C. nybakkeni (Higgins 1986); Fissu- added to Echinoderidae (Neuhaus and Blasche 2006); roderes higginsi Neuhaus & Blasche, 2006; F. novaezea- thus, the family comprises three genera today. landia Neuhaus & Blasche, 2006; F. papai Neuhaus & In a separate paper in the present issue of ODE Blasche, 2006; F. rangi Neuhaus & Blasche, 2006; (Sørensen 2008), a new echinoderid genus and species, F. thermoi Neuhaus & Blasche, 2006; Echinoderes and a new species of Cephalorhyncha are described. spinifurca Sørensen, Heiner & Ziemer, 2005; and Character states of these new taxa have been included in E. truncatus Higgins, 1983. the present paper’s data matrix, together with selected All character codings are based either on published character states of other known echinoderid species, in descriptions or on personal observations. Documenta- order to analyze the relationships within the diverse tion of most previously unreported character traits is family Echinoderidae. included in the character descriptions below. The main literature sources for character codings per species are as follows. Paracentrophyes quadridentatus: Higgins (1983), Material and methods Neuhaus (1995). Campyloderes macquariae: Higgins (1967), Neuhaus (2004). Antygomonas paulae: Sørensen Morphological traits of 17 terminal taxa were coded (2007). Dracoderes abei: Higgins and Shirayama (1990). in a character state matrix (Table 1). The matrix Dracoderes orientalis: Adrianov and Malakhov (1999). includes 36 characters, 32 of them bistate and 4 Zelinkaderes brightae: Sørensen et al. (2007). Pola- multistate, resulting in a total of 76 character states. canthoderes martinezi: Sørensen (2008). Cephalorhyncha All characters were coded as non-additive, except for liticola: Sørensen (2008). Cephalorhyncha asiatica: character 5 which was ordered. Detailed descriptions of Adrianov (1989), Adrianov and Malakhov (1999), all characters and character states are given below. Neuhaus and Blasche (2006). Cephalorhyncha nybakke- Outgroup taxa selected for the analysis are the ni: Higgins (1986a), Neuhaus and Blasche (2006). All homalorhagid Paracentrophyes quadridentatus (Zelinka, five species of Fissuroderes: Neuhaus and Blasche (2006). 1928) and five cyclorhagid non-echinoderid species: Echinoderes spinifurca: Sørensen et al. (2005). Echino- Antygomonas paulae Sørensen, 2007; Campyloderes deres truncatus: Higgins (1983), Sørensen (2006). macquariae (Johnston, 1938); Dracoderes abei Higgins Most species included in the data matrix were & Shirayama, 1990; D. orientalis Adrianov & Malakhov, examined with a light microscope; all except the species Table 1. Morphological character state matrix for analysis of relationships among selected kinorhynch species Character 000000000111111111122222222223333333 Species 123456789012345678901234567890123456 Paracentrophyes quadridentatus 1000000000––––11–––00–1–––––0––00000 Campyloderes macquariae –1000000101111110001100111100––00000 Antygomonas paulae 02000000111111110001100?11100––00000 Zelinkaderes brightae 02002000110001110001100100000––00000 Dracoderes abei 110100010201110011000–1100100––00000 Dracoderes orientalis 110?00010201110011000–100000?––00000 Polacanthoderes martinezi 020200010–0000001000111000110––01000 Cephalorhyncha liticola 021211?1120000001010111000110––00000 Cephalorhyncha asiatica 020211111200000010101110001111100000 Cephalorhyncha nybakkeni 021?11111200000010101??0001011?00000 Fissuroderes higginsi ?20200011200000010101110001110110111 Fissuroderes novaezealandia ?20?000112000000101011100010101p0ppp Fissuroderes papai ?20?00111200000010101110001111000000 Fissuroderes rangi 020?0001120000001000111000001100p000 Fissuroderes thermoi ?20200011200000010101110001010110111 Echinoderes truncatus 02022001120000001010111000100––00010 Echinoderes spinifurca 02022001120000001010111000100––00000 ? ¼ missing data; – ¼ inapplicable character state; p ¼ character state coded as polymorphic. ARTICLE IN PRESS M.V. Sørensen / Organisms, Diversity & Evolution 8 (2008) 233–246 235 of Fissuroderes, Dracoderes orientalis, Cephalorhyncha 4. Number of trichoscalids: 0 ¼ 14 trichoscalids; 1 ¼ 7 asiatica and C. nybakkeni were examined with SEM as trichoscalids; 2 ¼ 6 trichoscalids. Trichoscalids are well. Specimens of Fissuroderes and C. asiatica were small, modified sensorial head appendages with a loaned from the Museum fu¨ r Naturkunde, Berlin, featherlike appearance (Fig. 2E and F). They are Germany. Specimens of Paracentrophyes quadridentatus located in the posteriormost introvert ring (ring 07), were collected from muddy sediment at 42 m depth but usually they are not attached to the same outside of Kaldbak Fjord on the Faroe Islands radii as the scalids in the six more anterior rings. (6210400100N0614602300W) and kindly provided by Prof. Instead, their exact location is often related to R.M. Kristensen. The author collected specimens of another kind of structures, the trichoscalid plates, Campyloderes macquariae from intertidal aggregations which are small plates attached to the placids of the of the coralline alga Corallina officinalis in Kaldbak neck (Figs. 1A and B, 2F). However, trichoscalid Fjord, Faroe Islands (6210302800N0614904000W). Speci- plates can be missing in some species, e.g. in mens of Dracoderes abei were collected at the species’ Antygomonas paulae (Fig. 2E). In some kinorhynch type locality in Tanabe Bay, Japan (Higgins and species, e.g. in A. paulae, Campyloderes macquariae Shirayama 1990). Detailed information from SEM and
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