Zootaxa, Taxonomy of Serpulidae (Annelida ... - Magnolia Press

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Membranopsis Bush, 1910......................................................................................................................................... 62 Metavermilia Bush, 1905 ........................................................................................................................................... 62 Microprotula Uchida, 1978........................................................................................................................................ 64 Neomicrorbis Rovereto, 1904 .................................................................................................................................... 64 Neovermilia Day, 1961 ............................................................................................................................................... 66 Nogrobs de Montfort, 1808........................................................................................................................................ 68 Omphalopomopsis Saint-Joseph, 1894 ...................................................................................................................... 69 Paraprotis Uchida, 1978 ............................................................................................................................................ 70 Paumotella Chamberlin, 1919 ................................................................................................................................... 71 Placostegus Philippi, 1844 ......................................................................................................................................... 74 Pomatoceros Philippi, 1844 ....................................................................................................................................... 75 Pomatoleios Pixell, 1913 ........................................................................................................................................... 77 Pomatostegus Schmarda, 1861 .................................................................................................................................. 78 Protis Ehlers, 1887 ..................................................................................................................................................... 78 Protula Risso, 1826.................................................................................................................................................... 81 Pseudochitinopoma Zibrowius, 1969a ....................................................................................................................... 83 Pseudovermilia Bush, 1907 ....................................................................................................................................... 85 Pyrgopolon de Montfort, 1808 ................................................................................................................................... 86 Rhodopsis Bush, 1905 ................................................................................................................................................ 88 Salmacina Claparède, 1870........................................................................................................................................ 89 Semivermilia ten Hove, 1975 ..................................................................................................................................... 91 Serpula Linnaeus, 1758 .............................................................................................................................................. 93 Spiraserpula Regenhardt, 1961 ................................................................................................................................. 95 Spirobranchus de Blainville, 1818............................................................................................................................. 96 Tanturia Ben-Eliahu, 1976......................................................................................................................................... 98 Vermiliopsis Saint-Joseph, 1894 .............................................................................................................................. 100 Vitreotubus Zibrowius, 1979b ................................................................................................................................... 103 Invalid genera (as Serpulid) ............................................................................................................................................ 105 Key to serpulid genera (described before 2008) ............................................................................................................. 107 Acknowledgements ......................................................................................................................................................... 109 Glossary ........................................................................................................................................................................... 110 References ....................................................................................................................................................................... 113 Abstract The Serpulidae are a large group of sedentary polychaetes inhabiting calcareous tubes. The relationships within the group are poorly understood and taxonomy of the group is very confused which is a major obstacle to accessing their phylogeny. This review provides up-to-date information on the current state of taxonomy of Serpulidae sensu lato (not including Spirorbinae). The morphology of the group is reviewed with special reference to the features that can provide characters for future phylogenetic analyses. Scanning electron micrographs illustrate the structure of the chaetae and uncini. The list of 46—in our opinion valid—genera is accompanied by detailed generic diagnoses, species composition and distribution (checklist), and remarks on major taxonomic literature. A taxonomic key to the genera and a list of invalid genera with synonymy is also provided. Key words: Annelida, Polychaeta, Serpulidae, taxonomy, morphology, SEM ultrastructure of chaetae and uncini 4 · Zootaxa 2036 © 2009 Magnolia Press TEN HOVE & KUPRIYANOVA
Introduction The family Serpulidae is a discrete group of sedentary calcareous tubeworms within the large clade Sabellida, which shares a presence of radiolar crown and separation of the body into thoracic and abdominal regions, as divergence from the usually rather uniformly segmented motile polychaete form. The views on the relationships within Serpulidae have undergone many changes over the years (see Kupriyanova et al. 2006 for the details). The Serpulidae Rafinesque, 1815 had been traditionally divided into subfamilies Serpulinae Rafinesque, 1815 (although probably in most papers attributed to MacLeay, 1840) and Spirorbinae Chamberlin, 1919 until Rioja (1923) established the subfamily Filograninae. Pillai (1960) included 5 brackish-water serpulid genera in the subfamily Ficopomatinae but ten Hove & Weerdenburg (1978) revised the group and placed all its genera in the genus Ficopomatus. Uchida (1978) created 11 sub-families and numerous new genera, but his scheme, strongly criticized by ten Hove (1984) has not been accepted widely. Pillai (1970) elevated Spirorbinae to the family Spirorbidae, but later a number of authors suggested that Spirorbidae are more closely related to Serpulinae than to Filograninae (ten Hove 1984, Fitzhugh 1989, Smith 1991, Rouse & Fitzhugh 1994) and that the maintenance of the family Spirorbidae is not justified. Recent phylogenetic analyses confirmed the position of Spirorbinae as a subfamily of Serpulidae (Kupriyanova 2003, Kupriyanova et al. 2006, Lehrke et al. 2007, Kupriyanova & Rouse 2008). Ten Hove (1984) regarded the Filograninae as paraphyletic and a morphology-based cladistic analysis of some Serpulidae (Kupriyanova 2003) supported his conclusions. Moreover, the most recent phylogenetic analyses of Serpulidae using 18S ribosomal DNA (Lehrke et al. 2007) and another using combined molecular and morphological data (Kupriyanova et al. 2006) suggested that both traditionally formulated sub-families Serpulinae and Filograninae are not monophyletic. Kupriyanova et al. (2006) refrained from revising the serpulid classification and suggested that a major revision of serpulid taxonomy is needed based on more genera than used in their study. The major obstacle to a comprehensive phylogenetic analysis of the Serpulidae remains the state of its alpha taxonomy. It is almost proverbial to say that serpulid taxonomy is very confused and most currently recognized serpulid genera have long and convoluted taxonomic histories. Within Serpulidae, specific identification has traditionally been based on a combination of characters such as morphology of the operculum and opercular peduncle (if present), degree of development of the collar and thoracic membranes, structure of collar chaetae and tube and, to a lesser degree, structure of chaetae and uncini. Serpulid genera have been described on the basis of unique characters or on unique combinations of characters (even on absence of characters) rather than on presence of shared derived characters. Although traditionally only few characters have been used in serpulid taxonomy, variability of these characters remains largely unstudied. There have been very few reviews of serpulid taxonomy. The very first revision (Mörch 1863) was followed by early reviews by Saint-Joseph (1894), Bush (1905), and Pixell (1912, 1913). Chamberlin (1919) gave a key to the serpulid genera without attempting to revise the family, and so did Southward (1963), half a century later. Fauchald (1977) compiled a list of generic diagnoses and a key to genera for all polychaetes, including serpulids and spirorbids. In addition to the Spirorbidae, he acknowledged 331 species of serpulids, divided into 3 sub-families; the Serpulinae with 44 genera, the Filograninae with 5 genera, and the Ficopomatinae with 5 genera. Of these 54 genera, 22 were monotypic and another 13 had only 2 species. Uchida (1978) provided a systematic review of the group with a description of new species and new genera, but gave no key. He mentioned only 233 species, as compared to the 331 of Fauchald (1977). Of the 61 genera distinguished by Uchida (1978), 26 were monotypic, and 15 had only two species. No attempts to review Serpulidae have been made ever since and now, thirty years later, Fauchald (1977) still remains the most commonly used source of information on the generic composition of serpulids. During the last three decades serpulid taxonomy underwent significant changes, with numerous taxa being synonymized, older diagnoses emended and extended, new species described and about 10 genera added. TAXONOMY OF SERPULIDS: STATE OF AFFAIRS Zootaxa 2036 © 2009 Magnolia Press · 5
Page 1 and 2: ZOOTAXA 2036 Taxonomy of Serpulidae
Page 3: Zootaxa 2036: 1-126 (2009) www.mapr
Page 7 and 8: Morphology The tube Whereas tubes o
Page 9 and 10: FIGURE 2. Variability of serpulid t
Page 11 and 12: large crystals in the structure of
Page 13 and 14: 30A) most probably too are a series
Page 15 and 16: Throughout Spiraserpula, there is a
Page 17 and 18: FIGURE 3. Morphology of serpulid an
Page 19 and 20: FIGURE 5. General morphology of ser
Page 21 and 22: FIGURE 7. Serpulid morphology (cont
Page 23 and 24: occur incidentally in Pomatoceros,
Page 25 and 26: completely rasp-shaped (Knight-Jone
Page 27 and 28: Valid genera with diagnoses and lis
Page 29 and 30: 5. Apomatus globifer Théel, 1878,
Page 31 and 32: FIGURE 10. SEM micrographs of chaet
Page 33 and 34: tonguelets between ventral and late
Page 37 and 38: membrane. Radioles with ocellar clu
Page 39 and 40: Remarks. This relatively poorly kno
Page 41 and 42: 9. Ficopomatus Southern, 1921 (Fig.
Page 43 and 44: Filograna implexa M. Berkeley, 1835
Page 45 and 46: with 12-14 teeth in profile, up to
Page 49 and 50: 14. Galeolaria Lamarck, 1818 (Fig.
Page 51 and 52: operculate H. cancerum Knight-Jones
Page 53 and 54: 14. Hydroides bulbosus ten Hove, 19
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84. Hydroides trilobulus Chen & Wu,
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membrane and stylodes absent. Branc
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Schröder 1971), France (Fauvel 192
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most recent study (Kupriyanova et a
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FIGURE 28. SEM micrographs of chaet
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FIGURE 29. Photos and SEMs of chaet
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absent. Thoracic uncini saw-to-rasp
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Tube white, opaque, circular in cro
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FIGURE 33. Photos (A-E) and a SEM m
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normal radiole and maximally coveri
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31. Pomatoleios Pixell, 1913 (Fig.
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Jirkov (1997) further extended it t
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Paraprotula was based on the absenc
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limbate chaetae. Apomatus chaetae a
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1. Salmacina amphidentata Jones, 19
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7. Semivermilia torulosa (delle Chi
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13. Serpula maorica (Benham, 1927),
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limbate zone and proximal wing (Fig
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46. Vitreotubus Zibrowius, 1979b (F
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forming ventral apron. Collar chaet
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** According to Ippolitov (2007: 26
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38 (37) Thoracic uncini saw shaped
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chaeta (pl. chaetae): chitinous bri
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thoracic membranes: thin folds on b
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Bush, K.J. (1907) Descriptions of t
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Gravier, C. (1905 (1906?)) Sur deux
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Hove, H.A. ten & Wolf, P.S. (1984)
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Marsden, J.R. & Anderson, D.T. (198
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Radl, E. (1912) Neue Lehre vom Zent
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Verrill, A.E. (1873) Report upon th
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