Revision of the Australian Sabellariidae (Polychaeta)

Zootaxa 3306: 1–60 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

ISSN 1175-5326 (print edition)

Monograph

ZOOTAXA ISSN 1175-5334 (online edition)

ZOOTAXA 3306

Revision of the Australian Sabellariidae (Polychaeta) and description of eight new species PAT HUTCHINGS, MARÍA CAPA & RACHAEL PEART The Australian Museum, 6 College Street, Sydney, NSW 2010 Australia

Magnolia Press Auckland, New Zealand

Accepted by A. Nygren: 6 Mar. 2012; published: 9 May 2012

PAT HUTCHINGS, MARÍA CAPA & RACHAEL PEART Revision of the Australian Sabellariidae (Polychaeta) and description of eight new species (Zootaxa 3306) 60 pp.; 30 cm. 9 May 2012 ISBN 978-1-86977-899-6 (paperback) ISBN 978-1-86977-900-9 (Online edition)

FIRST PUBLISHED IN 2012 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2012 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326

(Print edition)

ISSN 1175-5334

(Online edition)

2 · Zootaxa 3306 © 2012 Magnolia Press

HUTCHINGS ET AL.

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Key to all genera worldwide and Australian species of Sabellariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Taxonomic account . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Idanthyrsus Kinberg, 1867 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Idanthyrsus australiensis (Haswell, 1883) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Idanthyrsus nesos n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Idanthyrsus willora n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Lygdamis Kinberg, 1867 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Lygdamis augeneri Kirtley, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Lygdamis giardi (McIntosh, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Lygdamis wambiri n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Phalacrostemma Marenzeller, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Phalacrostemma maloga n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Sabellaria Lamarck, 1812 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Sabellaria kooraltha n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Sabellaria lungalla n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Sabellaria pyramis n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Tetreres Caullery, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Tetreres terribilis n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Phylogenetic analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 List of tables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 About the authors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

Abstract The family Sabellariidae is represented in Australian waters by eleven species belonging to five genera, including eight of them new to Science: Idanthyrsus nesos n. sp., I. willora n. sp., Lygdamis wambiri n. sp., Phalacrostemma maloga n. sp., Sabellaria kooraltha n. sp., S. lungalla n. sp., S. pyramis n. sp., and Tetreres terribilis n. sp. Three genera, Phalocrostemma, Sabellaria and Tetreres, are newly recorded from Australia. Descriptions of all species are given, acompanied by detailed illustrations, including drawings, photographs and scanning electron micrographs, tables summarising specific diagnostic characters of all species of each genus, and distribution maps within Australia. A key to all genera worldwide and Australian species is given. Maximum parsimony analyses based on morphological features have been performed to assess the position of the new species and relationships with other Sabellariidae. For this purpose, the Australian species, the type species of each genus and some others species, representing the variability within each group, were incorporated into the analyses. Some of the species were recovered within monophyletic genera (Phalacrostemma and Tetreres), while others have been considered as member of previously recognised genera (Idanthyrsus, Lygdamis and Sabellaria) have not been assessed as monophyletic. Key words: Sabellariidae, Polychaeta, systematics, Australia, new species, new records

Resumen La familia Sabellariidae está representada en la costa Australiana por once especies pertenecientes a cinco géneros, ocho de las cuales son nuevas para la Ciencia: Idanthyrsus nesos n. sp., I. willora n. sp., Lygdamis wambiri n. sp., Phalacrostemma maloga n. sp., Sabellaria kooraltha n. sp., S. lungalla n. sp., S. pyramis n. sp., and Tetreres terribilis n. sp. Tres géneros, Phalocrostemma, Sabellaria y Tetreres, son citados por primera vez en Australia. Todas las especies han sido descritas y se acompañan de ilustraciones detalladas, incluyendo dibujos, fotografías y micrografías electrónicas de barrido, tablas donde se resumen los caracteres específicos de todas las especies de cada género y los mapas de distribución de la especies en Australia. Se presentan claves dicotómicas de todos los géneros y de las especies australianas. Se han realizado análisis de máxima parsimonia basados en caracteres morfológicos para comprobar la posición de las nuevas especies y las relaciones filogenéticas con otros Sabellariidae. Para ello, las especies australianas junto con las especies tipo y otras especies representantes de la variabilidad morfologica de cada grupo han sido incorporadas en dichos análisis. Algunas especies han sido recuperadas dentro de grupos monofiléticos

HUTCHINGS, CAPA AND PEART, AUSTRALIAN SABELLARIIDAE

Zootaxa 3306 © 2012 Magnolia Press ·

3

(Phalacrostemma y Tetreres), mientras que otras se han considerado como miembros de géneros previamente reconocidos (Idanthyrsus, Lygdamis y Sabellaria) a pesar de no haber sido estos recuperados como monofiléticos. Palabras clave: Sabellariidae, Polychaeta, sistemática, Australia, especies nuevas, nuevas citas.

Introduction Sabellariids are an easily recognised group of polychaetes with a well constructed tube of cemented sand grains and an operculum with rows of paleae which almost completely occupy and seal the entrance of the tube. Before the present study, the group was poorly known in Australia, and only four species in two genera were recorded (Day and Hutchings 1979). In Australia, sabellariids appear not to commonly form extensive reefs as found in other parts of the world (e.g., Kirtley 1994), and typically appear isolated or in small colonies. They have been collected from the intertidal to depths of 500 m worldwide but because deep water off Australia have been poorly sampled, we expect that the diversity will increase with further sampling. The number of sabellariid species reported worldwide is of 126, accommodated in 13 genera (Kirtley 1994; Lechapt and Gruet 1993; Lechapt and Kirtley 1996, 1998; Nishi and Kirtley 1999; Nishi et al. 2004; Bailey-Brock et al. 2007; Nishi et al. 2010; Santos et al. 2011). Kirtley (1994) undertook a major and most recent revision of the family, describing 43 new species genera from all over the world and erecting four genera. He established two subfamilies, although he did not perform any phylogenetic analyses to validate these taxonomic emendations. Recently, after the first phylogenetic analysis performed on the group (Capa et al. 2012), most genera established or emended by Kirtley (1994) were recovered monophyletic but subfamilies were found to be paraphyletic. The aim of the present study was to review the collections in Australian Museums, to describe the species present around the Australian coastline and to provide identification keys and complete illustrations for their characterization. Phylogenetic analyses were performed to assess the position of these taxa within the sabellariid tree and their relationships, especially after the finding that some of them showed a combination of generic features.

Materials and methods Sabellariid collections from several localities around the Australian coasts, housed in the main Australian museums were examined. Type and additional material of similar species were also studied for comparison. Material examined is listed in a clockwise direction around Australia, starting from the type locality. The number of specimens examined is shown after the locality data and before the registration number. Gravid individuals were selected as type material, when possible, because sabellariids exhibit considerable morphological changes after settlement, such as the elongation of the trunk, the loss of provisional chaetae, the rotation of the opercular lobes and their tentacular filaments (Wilson 1975: Figs 1–5). Presence or absence of coelomic oocytes was noted, and this is indicated in the Material Examined section with a ‘*’. Total length, including paleae and cauda and maximum width (anterior segments) were recorded for type material of each species and ranges given for the additional material examined. Descriptions are based on type material but in cases of damaged structures, additional information is given based on material from type locality and nearby areas. Features such as the ultrastructure and types of paleae and chaetae along the body were examined using scanning electronic micrographs (SEM) obtained by a Zeiss EVO LS15 SEM with a Robinson Backscatter detector. In addition, line drawings are provided as well as light microscopy photographs taken with a Leica MZ16 microscope and Spot flex 15.2 camera attached, to illustrate diagnostic characters. The descriptions were generated using DELTA System (DEscription Language for TAxonomy) to provide standardised species and generic descriptions (Dallwitz 1980; Dallwitz et al. 1993) and then edited. A dichotomic key was also produced with DELTA. Genera and species are listed alphabetically and synonymies included are those for which we have examined material, so for a complete list of synonymies see Kirtley (1994).


HUTCHINGS ET AL.

A total of 35 species of sabellariids and 31 characters (described in Capa et al. 2012) were incorporated into the analysis. Maximum parsimony analyses were performed following Capa et al. (2012) methods: heuristic searches using 10,000 replicates of random taxon addition and tree bisection-reconnection branch swapping algorithm, saving ten trees per replicate using TNT 1.1 (Goloboff et al. 2008a), with characters unordered and implementing implied weighting (Goloboff 1993, 1995, Goloboff et al. 2008b, Capa et al. 2012), varying k values from 3 to 20. Nodal support, estimated by 1,000 jackknife replicates using TBR, in TNT 1.1 (Goloboff et al. 2008a). The following abbreviations are used in the text: LACM-AHF—Los Angeles County Museum, Los Angeles, formerly The Allan Hancock Foundation; AM—Australian Museum, Sydney; BMNH—The Natural History Museum, London; MV—Museum Victoria, Melbourne; NTM—Museum and Art Gallery of the Northern Territory, Darwin; QM—Queensland Museum, Brisbane; SAM—South Australian Museum, Adelaide; WAM— Western Australian Museum, Perth; ZMH—Zoologisches Museum of Hamburg. For each genus we provide a table listing the diagnostic characters of all the described species (Tables 1–5), in many cases species are known only from the original description and/or lack information on informative characters. They are based on the original descriptions supplemented by other additional records from the type localities. When information on the variation of characters is available this is also noted.

Key to all genera worldwide and Australian species of Sabellariidae 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15. 16 -

Opercular lobes completely fused to each other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Opercular lobes partially fused at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Opercular lobes completely separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Outer row of opercular paleae with flattened blades. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phalacrostemma maloga n. sp. Outer row of opercular paleae with cylindrical blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Tentacular filaments simple, resembling long opercular papillae. Four parathoracic segments. . . . . . . . . . . . . Bathysabellaria* Tentacular filaments compound (numerous and arranged in transverse rows). Three parathoracic segments . . . Neosabellaria* Outer paleae with straight blades. Tentacular filaments simple, resembling long opercular papillae. Palps longer than operculum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetreres terribilis n. sp. Outer paleae geniculate, with shaft and blades forming an angle. Tentacular filaments compound (numerous and arranged in transverse rows). Palps shorter than half of length of operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gunnarea* Opercular disc (distal end) oblique to longitudinal axis (with a slope) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Opercular disc perpendicular to longitudinal axis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Outer paleae with straight,flattened blades and with large, pointed denticles on margins. Three parathoracic segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Idanthyrsus 11 Outer paleae with straight, flattened blades and smooth margins. Four parathoracic segments. . . . . . . . . . . . . . . . .Lygdamis 13 Blade of inner paleae cylindrical, resembling spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Blades of inner paleae different (flat or concave) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Nuchal spines straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mariansabellaria* Nuchal spines with bent tips (hooks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gesaia* Buccal flaps present; outer paleae arranged in spirals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phragmatopoma* Buccal flaps absent; outer paleae arranged in semicircles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Outer paleae with flat blades and smooth margins. Nuchal spines if present, straight and without limbations . . . . .Sabellaria15 Outer paleae with flat blades and large pointed denticles. Nuchal spines with bent tips (hooks) and limbation of convex side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paraidanthyrsus* Nuchal hooks with wide limbation, two pairs of lateral lobes on chaetiger 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. nesos n. sp. Nuchal hooks without wide limbation, three or four pairs of lateral lobes on chaetiger 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 15–16 pairs of outer paleae with 12–14 denticles along each margin, three pairs of lateral lobes on segment 2. . I. australiensis 17 pairs of outer paleae with 16 reflected denticles along each margin, four pairs of lateral lobes on segment 2 .I. willora n. sp. Outer paleae with finely denticulate distal tips. Branchiae present on posterior abdominal segments . . . . . . . L. wambiri n. sp. Outer paleae with smooth distal tips. Branchiae absent from posterior abdominal segments . . . . . . . . . . . . . . . . . . . . . . . . . . 14 42–44 pairs of outer paleae and 30–36 pairs of inner paleae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. augeneri 16–20 pairs of outer paleae and 10–14 pairs of inner paleae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. giardi Nuchal spines absent, one kind of middle paleae, triangular cone shaped. Branchiae absent on posterior abdominal segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. pyramis n. sp. Nuchal spines present, one kind of middle paleae, geniculate. Branchiae present on posterior abdominal segments . . . . . . . . 16 22–23 pairs of outer paleae with an elongate finely serrated distal plume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lungalla n. sp. 12 pairs of outer paleae, with large central distal smooth tip with two lateral teeth on either side, one kind of middle paleae 12– 14 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. kooraltha n. sp.

*Genera not known from Australian waters



5

Taxonomic account Idanthyrsus Kinberg, 1867 Idanthyrsus Kinberg, 1867: 350. Kirtley, 1994: 84. Capa et al., 2012: 261. Type species. Idanthyrsus macropaleus Schmarda, 1861 (after Kirtley 1994). Type locality. Valparaiso Harbour, Chile.

Diagnosis. Outer paleae straight (no angle between shaft and blade), flat and with pointed denticles along lateral margins and distal tips. Three thoracic segments and, if confirmed in all species of the genus, only fine lanceolate chaetae on neuropodia of parathoracic segments. Remarks. Before the present study there were 19 accepted species in the genus Idanthyrsus (Kirtley 1994; Nishi and Kirtley 1999). We have synonymised I. glaessneri with I. australiensis and described an additional two new species increasing the total number of valid species to 20 (see Table 1).

Idanthyrsus australiensis (Haswell, 1883) Figures 1A, B, 2–5, 6A, Table 1 Sabellaria (Hermella) australiensis Haswell, 1883: 634, pl. 12; figs 7–11. Sabellaria australiensis.— McIntosh, 1885: 416. Sabellaria sexhamata.— Collin, 1902: 742; not Grube, 1878: 219. Sabellaria (Pallasia) pennata.— Augener, 1914: 79; not Peters, 1854. Pallasia pennata.— Fauvel, 1917: 262. Augener, 1922: 33; 1926: 253; not Peters, 1854: 613. Idanthyrsus pennatus. — Dakin, Bennett and Pope, 1953: 153; not Peters, 1854: 613. Idanthyrsus glaessneri.— Kirtley, 1994: 97, fig. 6.7.

Material examined. TYPE ?: Sabellaria (Hermella) australiensis Haswell, 1883, BMNH 1882.2.22.71*, 47 mm in length excluding cauda, 6 mm in width (anteriorly), 47 chaetigers (in 2 parts, anterior fragment with 30 chaetigers and posterior with 17 chaetigers). Queensland: Thursday Island, 10°35'S 142°13'E, 7.8–12.3 m, collected by HMS Alert. Additional material examined. Queensland: Cape Yorke Peninsula, south of Fly Point, Putta Putta Beach, 10°45'S 142°36'E, 11.vii.1976, 1, AM W26926, on semi-exposed boulder bank; Thursday Island, 10°35'S 142°13'E, 4, AM W7087*, 1, AM W26683 (mounted for SEM), 1, AM W26684 (mounted for SEM), AM W26851; Gladstone, Calliope River, 23°55'S 151°10'E, 1, AM W199314*; Gladstone Harbour, 23°51'S 151°16'E, 06.xi.1975, 1, QM G10564; Gladstone, Calliope River, 23°55'S 151°10'E, 5; Gladstone Harbour, 23°51'S 151°16'E, 06.xi.1975, 3, QM G10565*; Mary River, 26°23' S 152°45' E, 31.xii.1971, low tide, 1, AM W26928; Hervey Bay, Point Vernon, 25°15'S 152°49'E, iv.1972, 1, AM W201700*; Maroochydore, Alexandra Headland, 26°39'S 153°06'E, 01.iv.1972, 1, AM W26929; Caloundra, 26°48'S 153°08'E, viii.1945, 4, QM G4043, from south beach rocks; 2, QM G4044; 1938, 1, QM G212812; Moreton Bay, Peel Island, Horseshoe Bay, 27°30'S 153°21'E, 18.xii.1977, 1, QM G212813*, from under rocks submerged at low tide. New South Wales: Lennox Heads, 28°48'S 153°36'E, 27.iii.1972, 1, AM W26971; south of Clarence River mouth, Minnie Water, 29°47'S 153°18'E, 2–3.xi.1963, 3, AM W26895; 25.ii.1971, 1, AM W26901, from shore reef, in crevices, covered in coralline algae, 1, AM W26900, from shore reef, underneath boulders in rockpools; Wilson Headland, 29°50'S 153°17'E, 12.v.1984, 6, AM W26857; off Coffs Harbour, Arrawarra Head, 30°04'S 153°12'E, 14.x.1974, 11, AM W9196*, cemented together forming small colony; Coffs Harbour, SW face of South Solitary Island, 30°12'S 153°16'E, 19.v.1972, 7.6–10.7 m, 5, AM W26917; 17.v.1973, 15.2 m, 3, AM W26916, from small boulders covered in ascidians; Coffs Harbour, west side of Solitary Island, 30°12'S 153°16'E, 18.v.1972, 1, AM W26923; Toowoon Bay, near Tuggerah Lake, 33°22'S 151°30'E, 13.iv.1986, 0.6 m, 3, AM W26858, from rocks loosely piled, but cemented together, covered in Homosira banksii; off Avalon, 33° 38'S, 151° 20'E, 31.i.1973, 36.6 m, 1, AM W26849, from reef flat; Long Reef, 33°45'S 151°19'E, 16.xi.1970, 3, AM W4524* (mounted for SEM), from on underside of rocks, in sandy, firm tubes; Long Reef, north side of reef, 33°45'S 151°19'E, 23.ii.1971, 1, AM W26902; v.1967, 2, MV F78871*; Port Jackson, 33°50'S 151°16'E, 10.i.1983, intertidal, 1, AM W198478*; North Head, 33°50'S 151°18'E, 26.v.1972, 28.9 m, 1, AM W6963*; 13.xii.1972, 28.9 m, 1, AM W26925; Maroubra,


HUTCHINGS ET AL.

33°57'06"S 151°17'37"E, 01.xii.1954, 1, AM W1575*; Kurnell, Potter Point, 34°03'S 151°13'E, 27.ix.1989, low tide, 4, AM W26863, from on and under rocks; Port Hacking, near Jibbon Head, 34°05'S 151°10'E, 23 m, 1, AM W16351*, from reef; north of Wollongong, Bellambi Reef, 34°26'S 150°53'E, 14.xi.1971, 1, AM W26968, from rock pool, in holdfast of seaweed; Shellharbour, 34°35'S 150°52'E, between tide marks, 5, AM W26903; Shellharbour, near bluestone quarry, 34°35'S 150°52'E, 05.ix.1971, 7.6–9.1 m, 3, AM W26974; Gerroa, Blackhead, 34°46'S 150°49'E, 25.ix.1982, mid tide, 1, AM W198479*; north of Broulee, 35°51'S 150°11'E, i.1971, 1, MV F78866*; near Jervis Bay, Wreck Bay, Cemetry Point, 35°11'S 150°38'E, 27.ii.1976, intertidal, 1, AM W26927, on coralline algae; Twofold Bay, Murrumbulga Point, 37°05'S 149°54'E, 09.x.1984, intertidal, 1, AM W199879*, from gravel and algal washings on rock platform; 27.iii.1985, intertidal, 1, AM W199889*, cryptic fauna and rock platform; 25.vi.1985, subtidal, 2, AM W199880*, crevice fauna and amongst Diopatra tubes on rock platform; Twofold Bay, Red Point, 37°05'S 149°54'E, 20.v.1995, 1, AM W26907; Munganno Point, 37°05'S 149°54'E, 10.x.1984, 6 m, 10, AM W199892* from erect bryozoan colony on wharf pile. Victoria: South tip of Mallacoota, Bastion Point, 37°34'S 149°46'E, 1, AM W26525 (mounted for SEM); Western Port, Merricks, 38°24'S 145°07'E, 1969, 1, MV F78856*; Bass Strait, 5 km south of Point Reginald, 38°48'S 143°17'E, xi.1981, 47 m, 1, MV F78883*; Western Port, Somers, 38°24'S 145°10'E, 2, MV F78853*; Flinders Ocean Beach, 38°29'S 145°02'E, 17.xii.1969, 1, MV F78872*; Venus Bay, west of Eagles Nest, 38°40'S 145°41'E, 01.ii.1966, 1, MV F78868*; Western Port, near Shoreham, Honeysuckle Point, 38°26'S 145°04'E, 14.xi.1970, 1, MV F78864*; Flinders, south reef, 38°29'S 145°02'E, 13.xi.1965, 1, MV F78863*; Airey's Inlet, 38°28'S 144°06'E, 31.i.1967, 1, MV F78869. South Australia: Sellicks Beach, 35°20'S 138°27'E, 16.iii.1979, 3, AM W198439*, York Peninsula, Gleeson's Landing, 34°35'S 137°43'E, 27.xi.1985, intertidal, 1, SAM E3093, from under rocks; 3, SAM E3094, from under rocks; Aldinga Bay, 35°21'S 138°25'E, 12.x.1977, low tide, 1, AM W14055*, under rocks. Western Australia: King George Sound, Mistaken Island, 35°04'S 117°56'E, 1.i.1988, 1, AM W26894, crevice fauna; Cape Naturaliste, Bunker Bay, 33°32'S 115°02'E, 30.i.1972, low tide, 6, WAM V156, from amongst metamorphic rocks; Bunker Bay, 33°31' 5.48"S 115°03 25.56'E, 30.i.72, low tide, many, WAM V284; 2 km off North Point*, 32°01'S 115°30'E, 11.i.1991, 30 m, 1, AM W26908, from reef and coral bommie surrounded by algal beds; Rottnest Island, Strickland Bay, 32°00'S 115°30'E, 20.i.1991, 1, AM W26904, from reef edge, rock and algae; Sandy Cape, Point Peron, 32°01'S 115°30'E, 1946, 2, AM W26856, 1, AM W26855; 2 km south of Cape Peron, reef west of Groyne, 32°01'S 115°30'E, 26.xii.1983, 3, AM W26862, from sponges and gorgonaceans from cove on reef; Point Clone, 32°01'S 115°30'E, 29.xi.1945, 1, AM W26854; Warnborough Sound, 32°20'S 115°43'E, 21.iii.1993, 5 m, 1, AM W26909, from Posidonia sp. beds; Fremantle, South Mole, 32°03'S 115°44'E, 1, AM W26861; Cockburn Sound, 32°01'S 115°45'E, 15.ii.1966, 1, WAM V3814; Cottesloe Reef, near Groyne, 31°59'S 115°45'E, 29.iii.1973, 2, WAM V311, from under stones and in dead shells; Cottesloe Beach, 31°59'S 115°45'E, 08.i.1988, intertidal, 2, AM W26864; 29.iii.1973, 2, WAM 72–74, from under stones and in dead shells; Kendrew Island, Dampier Archipelago, 20°29'S 116°32'E, 03.iv.1987, 10 m, 1, AM W26911, crevice fauna; Finucane Island, 20°18'S 118°33'E, 20.vi. 1970, AM W26914; Roly Rocks, 20°33'S 116°32'E, 03.iv.1987, 10 m, 1, AM W26910; Port Hedland, Finucane Island, 20°18'S 118°33'E, 21.vi.1970, 1, AM W26912; Broome, Gantheame Point, 17°59'S 122°11'E, 04.vii.1970, ELWM spring, 2, AM W26859; 15.iii.1987, 12–18 m, 2, NTM W4916*, from rocky reef; 15.iii.1987, LWS, 1, NTM W4548*, from rock slabs in silt; Broome, Cable Beach, 17°57'S 122°12'E, 29.ix.1984, LWS, 1, NTM W2280*, from under stones; 02.xii.1981, 3 m, 2, MV F78852; Riddell Point, Broome, 17°56.30'S 122°12.33'E, 19.viii.1982, 1 WAM 1890. Northern Territory: Port Essington, Sandy Island, 11°06'S 132°18'E, 02.v.1982, 14 m, 1, NTM W1122*, from muddy bottom; 1, NTM W1124*, from muddy bottom; Port Essington, Burford, 11°29'S 131°57'E, 13.x.1981, 1, NTM W1120*, from reef flat; Cobourg Peninsula, Burford Island, 11°29'S 131°57'E, 13.x.1981, 1, NTM W25*; Charles Point, Beagle Gulf, 12˚18.96'S 130˚40.74'E, 23 m, 13.x.1993, NTM W23480. Other species examined for comparison. HOLOTYPE of Idanthyrsus glaessneri Kirtley, 1994, LACM AHF N6277, sand flat at low tide, Lakes Entrance, Victoria, 37.8°S 147.9° E, coll. B. Dew, 1.3 cm in length, 3.42 mm in width (maximum width of thorax without chaetae). Description. Type flesh coloured, with dark brown blotches on sides of operculum and anterior end and darkly pigmented cauda. Body compact and robust (Fig. 1A). Operculum with completely separate lobes, longer than wide (Figs 1A, 3B), with distal end sloped posteriorly (oblique to longitudinal axis). Fourteen pairs of paleae in outer row, with cylindrical shaft (Fig. 3A, C), and flat, straight blade, ornamented with alternate, sharp-tipped, similar sized and slightly curved denticles on margins. Inner row with nine pairs of cylindrical and smooth paleae,



7

tapering evenly to acute tip (Figs 1A, B, 3A, D). Opercular papillae, 14 pairs, peripheral to outer row of paleae on each lobe (Fig. 3A). Three pairs of nuchal spines with bent tips (hooks) on each side, concave margins smooth,

FIGURE 1. Line drawings of Idanthyrsus australiensis type (BMNH 1882.2.22.71) A. Lateral view. B. Anterior dorsal view. Idanthyrsus nesos n. sp., Holotype (WAM V7850) C. Entire animal anterior end dorsal and twisting to ventral side posteriorly. D. Dorsal view. Idanthyrsus willora n. sp., Holotype (WAM V7852) E. Entire animal anterior end dorsal and twisting to ventral side posteriorly. F. Anterior end, ventral view. Scales A, B, C, E, F =5 mm, D = 1 mm.


HUTCHINGS ET AL.

FIGURE 2. Photos of Idanthyrsus australiensis (one specimen from lot AM W91996) A. Anterior end, ventral view. B. Detail of opercular structures, ventral view. C. Midventral ridge and medial organ. D. Anterior end, lateral view. E. Detail of opercular structures.lateral view. F. Anterior end, dorsal view. G. Opercular paleae and nuchal spines (hooks). H. Posterior end, lateral view. Abbreviations: b2, branchia segment 2; bo, buccal organ; ca, cauda; cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1; chn2, chaetae neuropodium segment 2; es, eyespots; ip; inner paleae; li, lips; ll2, lateral lamellae segment 2; mo; medial organ; mr, median ridge; ns, nuchal spines (hooks); op, outer paleae; tf, tentacular filaments.



9

FIGURE 3. SEM of Idanthyrsus australiensis from type locality, Thursday Island (AM W6683) A. Anterior end, dorsal view. B. Anterior end, ventral view. C. Outer paleae. D. Inner paleae. E. Neurochaetae of segment 1. F. Lanceolate and capillary notochaetae from parathoracic segment. G. Abdominal neurochaetae. H. Posterior abdominal neurochaetae in detail. I. Anterior abdominal uncini. Scales A, D = 200 µm, B. = 400 µm, C =30 µm, E, G, I. 20 µm, F, H. = 10 µm.

limbation absent (Figs 1B, 3A). Tentacular filaments, arranged in nine rows (Fig. 3B). Median ridge and median organ present, extending to dorsal edge of junction of opercular lobes, raised and pigmented (illustrated in non type material, Fig. 2B, C). Numerous eyespots present on both sides of median ridge. Palps about half length of operculum. Segment 1 (chaetiger 1), with neuropodial lobe on either side of U-shaped buccal organ (illustrated in non type material Fig. 2B); capillary neurochaetae present, flattened and with finely denticulated margins (Fig. 3E). Segment 2 (chaetiger 2) with three pairs of triangular lateral lobes, connecting dorsal branchiae to neuropodia; with similar neurochaetae as those on chaetiger 1. Neuropodia of chaetiger 1 and 2 not vertically aligned, those of chaetiger 2 situated more laterally (Fig. 1B). Fourty six pairs of dorsal branchiae present from chaetiger 2, continuing along entire length of body but diminishing in size in posterior segments, some represented only by scars (Fig. 1A). Branchiae conical, with transverse rows of cilia and blunt tips (Fig. 1A, B). Segments 3–5 (parathoracic) with two types of chaetae arranged transversely, eight, straight, flattened, lanceolate, some with frayed twisted tips, majority represented only by stumps and eight fine, smooth capillaries, inserted between lanceolate chaetae (Fig. 3F). Segments 3–5 with lancelolate neurochaetae arranged in two tiers of different lengths (longer about twice the length of shorter, as illustrated in non type material Fig. 4I). Notopodial parathoracic


HUTCHINGS ET AL.

chaetae larger and more robust than neuropodial. Abdominal region with 42 chaetigers. Notopodia as transverse tori with uncini, decreasing in number posteriorly. Each uncinus with two vertical rows of teeth, each with about eight teeth (Fig. 3I). Neuropodia with capillaries becoming longer posteriorly, with thin and flattened blades ornamented with thecal laminar extensions distally forming oblique rows and splayed tips (Fig. 3G, H). Cauda smooth, about a quarter of abdominal length (Fig. 1A). Variation. As the type material and material from the type locality is in relatively poor condition we provide here some supplementary information based on the material from Arrawarra, northern NSW (Fig. 2A–H). The median ridge and median organ (Fig. 2C) have a line of eye spots on both sides. The lobes of chaetiger 1 have fine capillary neurochaetae laterally displaced in relationship to those of chaetiger 2 (Fig. 2E). Chaetiger 2 with three pairs of triangular lateral lobes which connect the branchiae to the neuropodia (Fig. 2D). The triangular shaped branchiae (Fig. 2F) continue all along the abdomen. Parathoracic neuropodia were examined under SEM on material from Long Reef, NSW showing two tiers of lanceolate neurochaetae present (Fig. 4I). A large number of specimens were examined from several localities, including sexually mature individuals, that vary in length (4–60 mm), width (1–6 mm), number of chaetigers (25–52), number of outer opercular paleae (8–28 pairs), inner opercular paleae (6–17 pairs), opercular papillae (5–16 pairs), nuchal hooks (2–3 pairs), number of rows of tentacular filaments (8–11), number of branchiae (14–46 pairs), number of teeth on abdominal uncini (8–10 ) and number of abdominal chaetigers (15–49). These variations can be mainly attributed to size of the specimens, with larger animals tending to have more paleae, papillae, uncinial teeth and abdominal chaetigers. As minor differences in structure of paleae and nuchal hooks have been used previously to differentiate species, we illustrate these structures from material collected from Long Reef, Sydney, NSW (Fig. 4A–M) and from Mallacoota, Victoria (Fig. 5A–G) the type locality of Idanthyrsus glaessneri Kirtley, 1994, in addition to material from type locality (Fig. 3A–I). The outer paleae of specimens examined from each of these three localities vary slightly in terms of length of the denticles on margins, and we suggest that this represents a gradual gradation rather than separate species and represent a difficult character to measure (Figs 3C, 4C, 5C). Lanceolate parathoracic notochaetae appear to be contractile within parapodia (compare Fig. 4H and Fig. 5E), and we suggest that these parapodia are highly mobile. Neurochaetae of the first two segments are missing from one side on holotype, but on non type material from type locality (eg. AM W26683, AM W26684) they are present on both sides. Abdominal neurochaetae are similar in these three distant localities (Figs 3G, H, 4J, K, 5F). Divergence of the opercular lobes varies and we suggest this may be a function of size and fixation. Remarks. The diagnostic characters of I. australiensis are the combination of the absence of a limbation on the nuchal spines, the presence of three lateral lobes on segment 2, and the presence of branchiae on posterior segments. Kirtley (1994) primarily used the shape and thecal development of the outer and inner paleae to distinguish between species, but in most cases intraspecific variation was not considered. We have tried to group this variation observed in some groups regarding the arrangement, shape, relative length and relative numbers of denticles on both sides of outer paleae (Table 1). The structure of the notopodial chaetae of the parathoracic segments also differs between I. australiensis (Figs 3F, 4H) and I. nesos n. sp. (see this paper), in the former the capillaries imbetween the lanceolate chaetae are thin whereas in the new species the capillaries have a wider distal end, resembling lanceolate chaetae (Fig. 8F, G). However, for virtually all described species these parathoracic notochaetae are poorly if at all described and comparison cannot be made at this stage, and this is also true for the parathoracic neurochaetae and therefore these characters are not given in Table 1, but we suggest they may provide useful characters to distinguish between species. Idanthyrsus glaessneri Kirtley, 1994 from Lakes Entrance, Victoria, Australia, was briefly described and only the outer and inner opercular paleae illustrated. In his key to species of Idanthyrsus, Kirtley (1994) separated this species from I. australiensis due to the length of denticles (without sharp tipped denticles in I. glaesneri and sharp in I. australiensis), ornamentation of the thecae (irregular wavy lines across blade in I. glaesneri) and overall shape of paleae (strongly bent in I. glaesneri and slightly sigmoidal in I. australiensis). After examination of the holotype of I. glaesneri, which has been partially dried at some stage, is posteriorly incomplete, and has lost the majority of the outer row of paleae, we would describe the outer opercular paleae as with alternate denticles, slightly curved, with a ratio of 1:1. This together with other diagnostic features (presence of three lateral lobes on segment 2 and the presence of branchiae on posterior segments) appear to fall within the range observed for I. australiensis and we have therefore synonymised the two species.



11

FIGURE 4. SEM of Idanthyrsus australiensis from Long Reef, Sydney (AM W4524) A. Anterior end ventral view. B. Anterior end, dorsal view. C. Outer paleae. D. Inner paleae, E. Nuchal spines (hooks) and opercular papillae. F. Neurochaetae from segment 1. G. Anterior segments (thorax and parathorax) latero-ventral view. H. Lanceolate notochaetae of parathorax. I. Lanceolate neurochaetae of parathorax. J. Abdominal neurochaetae. K. Abdominal neurochaetae in detail. L. Abdominal uncini, lateral view. M. Abominal uncini, frontal view. Scales A, B, G =1mm, C = 200 µm, D, F, H =100 µm, E, I, J, K = 30 µm, L, M = 10 µm.


HUTCHINGS ET AL.

FIGURE 5. SEM of Idanthyrsus australiensis from Mallacoota (AM W26525)—type locality of Idanthyrsus glaessneri Kirtley, 1994. A. Anterior end, ventral view. B. Anterior end dorsal view. C. Outer and inner paleae. D. Neurochaetae of segment 1. E. Lanceolate and capillary notochaetae from parathorax. F. Anterior abdominal neurochaetae, in detail. G. Posterior abdominal uncini. Scales A, B = 200 µm, C = 100 µm, D = 30 µm, E. = 40 µm, F, G = 10 µm.

Examining a large amount of material of I. australiensis from many localities (Fig. 6A) has allowed us to assess the variability of characters such as the ornamentation of the paleae which Kirtley (1994) had largely relied upon to distinguish species and we have concluded that additional characters such as numbers and distribution of denticles on the outer paleae, number and shape of nuchal spines, numbers of lateral lobes on segment 2 and absence or presence of branchiae in posterior abdominal segments are useful to distinguish between species (see Table 1). In summary, we suggest that I. australiensis exhibits some variation in a suite of characters but no discrete geographical patterns can be detected along the distribution range of this species (Fig. 6A). Distribution. Australia wide from NE Queensland to NW Western Australia, except Tasmania (Fig. 6A). Habitat. Found from low intertidal to depths of 50 m attached to solid substrate, sometimes occurring as crevice fauna. Specimens typically form small colonies of 6–10 individuals, although specimens from Bunker Bay, in Western Australia, were found in larger colonies, consisting of several hundreds of specimens.

Idanthyrsus nesos n. sp. Figures 1C–D, 6A, 7, 8, Table 1 Material examined. HOLOTYPE: 1, WAM V7850*, Dolly Beach, Christmas Island, Indian Ocean, 10°31'S 105°41'E, 27–28 xi. 1969, intertidal reef and pools, 18.5 mm in length, 1 mm in width, 33 chaetigers. PARATYPES: 12, WAM V7851*, same locality as holotype, 8–20 mm in length without cauda, 1 mm in width, 20–29 chaetigers; same locality as holotype, AM W37778 (mounted for SEM on two pins).



13

FIGURE 6. Map illustrating the distributions of Australian species of Sabellariidae, A. Idanthyrsus australiensis, Idanthyrsus nesos n. sp., Idanthyrsus willora n. sp., Phalacrostemma maloga n. sp., Sabellaria pyramis n. sp. B. Lygdamis augeneri, Lygdamis giardi, Lygdamis wambiri n. sp., Sabellaria kooraltha n. sp., Sabellaria lungulla n. sp. and Tetreres terribilis n. sp.

Description. Holotype thin (Fig. 1C), body with slightly pigmented cauda, abdominal segments dorsally ridged, pigmented and glandular, ventrum with scattered pigmentation. Operculum with elongate, partially fused lobes; with distal end sloped posteriorly (oblique to longitudinal axis). Outer paleae, 24 pairs (Figs 1D, 7A, 8A, B), with cylindrical shaft and ornamented thecae as compact rings with irregular edges, restricted to proximal third of length of paleae (Fig. 8C) and flattened straight blades with alternate, mostly straight sharp-tipped denticles, increasing in length along shaft towards tip (Fig. 8C). Inner row with 15 pairs of cylindrical paleae tapering evenly to acute tip (Fig. 8D); ornamented distally with compact narrow ring thecae (Fig. 8D). Outer paleae longer than inner paleae (Fig. 8B). Opercular papillae, five pairs, reduced, stout, inserted external to outer row of paleae on each lobe (Fig. 8B). One pair of stout and flattened nuchal spines present, strongly recurved tips (Figs 1D, 7C, 8B), and concave margins with wide limbation. Tentacular filaments compound, arranged in 13 horizontal rows (Fig.


HUTCHINGS ET AL.

7A, B). Median ridge and median organ present at dorsal junction of opercular lobes, with eyespots present on their margins. Palps about one fifth of length of opercular lobes. Segment 1 (chaetiger 1) with rounded lobe-shaped neuropodia (Fig. 7A) on either side of U-shaped buccal organ, with fine capillary neurochaetae, flattened, with finely denticulated margins, widest at base and tapering towards fine tip (Fig. 8E). Segment 2 (chaetiger 2) with two pairs of poorly developed triangular lobes with blunt tips, dorsal one larger than ventral, connecting branchiae to neuropodia (Fig. 1D), and with similar chaetae as those of chaetiger 1. Neuropodia of chaetiger 1 and 2 not vertically aligned, those of chaetiger 2 situated more laterally (Fig. 1D). Twenty one pairs of dorsal branchiae present from chaetiger 2, absent from posterior abdominal segments. Branchiae with narrow base and weakly ridged, tapering gently to rounded tip, decreasing in size posteriorly. Segments 3–5 (parathoracic) with two types of notochaetae arranged transversely, five, large, flattened, lanceolate with frayed tips, with upturned, curved margins (Fig. 8F, G); and thinner chaetae with straight tips, theca with expanded margins, interspersed with lanceolate (Fig. 8F, G). Notochaetae appear to be highly retractile (Fig. 8F). Segments 3–5 (parathoracic) with two tiers of lanceolate neurochaetae; seven, thick and seven thinner ones with finely denticulated margins. Neurochaetae less robust than notochaetae. Abdominal region with 27 chaetigers. Notopodia as transverse tori, with number of uncini per torus decreasing posteriorly. Notopodial tori becoming increasingly erect and decreasing in width posteriorly. Each uncinus with two vertical rows of teeth, each row with seven to eight teeth (Fig. 8I). Neuropodia with capillaries becoming longer posteriorly, with thin and flattened blades ornamented with thecal laminar extensions distally forming oblique rows with elongate frayed tips (Fig. 8H), similar on all abdominal chaetigers. Cauda, smooth, equivalent in length to 15 chaetigers (Fig. 7D).

FIGURE 7. Photos of Idanthyrsus nesos n. sp. holotype (WAM V7850) A. Anterior end, ventral view. B. Same, lateral view. C. Same, dorsal view. D. Posterior end, lateral view. Abbreviations: b2, branchia segment 2; ca, cauda; cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1; ns, nuchal spines (hooks); op, outer paleae; ot, tentacular filaments.

Variations. The paratypes, including sexually mature female specimens, vary from 8–20 mm in length without cauda, no variation in width, with 20–29 segments, 12–24 pairs of outer paleae, 10–15 pairs of inner paleae; 4–9 pairs of opercular papillae; 9–19 pairs of dorsal branchiae; and 15–24 abdominal chaetigers. These variations can be attributed to size of specimens, with larger animals tending to have more paleae, papillae, uncinial teeth and abdominal chaetigers than smaller ones. Remarks. Idanthyrsus nesos n. sp. is the only species described in the genus with the following combination of characters (see Table 1): presence of one pair of limbated nuchal hooks (shared with I. bihamatus (Caullery,



15

1944), I. cretus Chamberlin, 1919, and presumably I. bicornis (Schmarda, 1861), I. kornickeri Kirtley, 1994, I. luciae (Rochebrune, 1882), I. manningi Kirtley, 1994, and I. valentenei Kirtley, 1994), presence of two lateral lobes on segment 2 (shared with I. bihamatus) and presence of fine denticles (20–25) along the edges of the outer paleae, increasing in length towards the tip. Idanthyrsus nesos n. sp., was found at Christmas Island, an isolated location on the Indian Ocean and can be distinguished from I. australiensis, the most common species on the mainland, by the number and shape of nuchal spines (one pair of flattened hooks with large limbation in I. nesos n. sp., and 2–3 pairs of cylindrical hooks lacking limbations in I. australiensis) and the number of pairs of lateral lobes on segment 2 (two in I. nesos n. sp., and three in I. australiensis) and by the distribution of branchiae (restricted to anterior abdominal segments in I. nesos n. sp., and present on all abdominal segments in I. australiensis).

FIGURE 8. SEM of Idanthyrsus nesos n. sp. (AM W37778) A. Anterior end, ventral view. B. Same dorsal view. C. Outer paleae. D. Inner paleae. E. Neurochaetae of segment 1. F. Lanceolate notochaetae, of two sizes from parathoracic segments. G. Same but in detail. H. Anterior abdominal neurochaetae, in detail. I. Posterior abdominal uncini. Scales A = 200 µm, B = 100 µm, C, D, E = 30 µm, F = 20 µm, G, H = 10 µm, I = 2 µm.


HUTCHINGS ET AL.



17

As summarised in Table 1, other species reported from the Indo-Pacific differ from I. nesos n. sp., in one or more of the diagnostic features, although it should be noted that information is missing for many species. For example, I. albigens (Ehlers, 1908), described from deepwater off Diego Garcia,has two pairs of nuchal hooks whereas I. nesos n. sp., has one pair of these spines. Idanthyrsus bicornis (Schmarda, 1861), described from intertidal areas of Sri Lanka, has outer paleae with more than 30 denticles with an asymmetrical arrangement on both margins and with maximum length at mid paleae, (see Fig. 6.4.1 in Kirtley 1994), in contrast to the 20–25 alternate denticles increasing in length towards the tip present in I. nesos n. sp. Idanthyrsus bihamatus (Caullery, 1944), known only from deep water in Indonesia, has outer paleae with denticles similar in size or slightly longer at mid length of paleae while in I. nesos n. sp., denticles increasing in length towards the tip (see Table 1 for comparison of other features). Idanthyrsus bicornis (Schmarda, 1861), described from intertidal areas of Sri Lanka, has outer paleae with more than 30 denticles which increase in size towards the tip and arising at 45˚ (see Fig. 6.4.1 in Kirtley 1994, not illustrated by Schmarda and also no other diagnostic characters are given) which differs from the new species and I. bihamatus (Caullery, 1944) known only from deep water in Indonesia which has outer paleae with smooth shafts and numerous denticles of uniform size along shaft and arising at 45˚ whereas I. nesos n. sp., has straight denticles increasing in length along the strongly denticulate shaft, although they share many other characters (see Table 1). Idanthyrsus nesos n. sp., differs from I. okinawaensis Nishi & Kirtley, 1999, described from shallow waters in Japan, in the number of pairs of opercular papillae, 30 in I. okinawaensis and less than ten in the new species,and the number and shape of nuchal hooks, two pairs and without a limbation in I. okinawaensis and one pair of limbated hooks in I. nesos n. sp. Distribution. Christmas Island, Indian Ocean. Habitat. Intertidal, found in small colonies. Etymology. The specific name nesos is a Greek word for island and refers to the isolated type locality of this species, Christmas Island.

Idanthyrsus willora n. sp. Figures 1E–F, 6A, 9, 10, Table 1 Material examined. HOLOTYPE: 1, WAM V7852*, West Australia: Quondong, 17°34'S 122°09'E, 01.vii.1975, collected beneath intertidal rock slabs, 45 mm in length, 5 mm in width, 48 chaetigers. PARATYPES: 1, WAM V7853*, 32 mm in length, 5 mm in width, 47 chaetigers, 1, AM W37779 (mounted for SEM on two pins), both paratypes from same sample as holotype. Additional material examined. Western Australia: Dampier Archipelago Kendrew Isl. 20°28'30"S 116°325'12"E 7.v.1973, 8 m, 1, WAM V3812; transect N, Kendrew Isl. 20°28'28"S 116°25'12"E 11.ii.1973, 1–2 m, 1 WAM V899; Lady Nova Flat, NE Rosemary Isl. 2.xi.1971, intertidal on oyster, 3, WAM V677; northeast side of Cape Naturaliste, 33°32'S 115°00"E, 26.xii.1958, low tide, 2, WAM V855, under stones; Port Denison, 32°10'51"S 115°45'12"E, 27.x.1973, 9.7–10.6 m, 1, WAM V626, on pieces of reef; Near Little Isl. off Sorento, 31°34'36"S 115°44.22"E, vi.1974, 3 m on bivalve shells, 1, WAM V3820; Cottesloe Reef, 32°35'39"S 115°58.29"E, 14.vi.1973, intertidal on Ninella mollusc, 3, WAM V313. Northern Territory: Trepang Bay, 11°13'S 131°57'E, 15.x.1981, 5 m, 1, NTM W26*, from edge of reef. Description. Holotype robust, compact (Fig. 1E), with dark pigment patches throughout ventrum, on abdominal tori and caudas. Operculum partially fused lobes (Figs 9A, B, 10F), with an outer row of 17 pairs of paleae, with cylindrical and smooth shafts, flat and curved blades, with sharp-tipped denticles present on anterior half of shaft (Fig. 10C), length of which declines towards to tip, more splayed basally (Figs 1F, 9A, 10C). Inner row with 13 pairs of cylindrical paleae, slender, smooth, tapering gently with slightly curved tips (Fig. 10C, D). Opercular papillae, 12 pairs, inserted externally to outer paleae on each lobe (Fig. 1E). Four pairs of nuchal spines (hooks) on each side (Figs 1F, 9A), long shafted, and cylindrical, with subacute tips, lacking limbation; longest ones with more rounded tips, weakly recurved (Figs 1E, F, 9D, 10B). Tentacular filaments compound, arranged in 20 transverse, long rows (Fig. 9A, B). Small median organ present at dorsal juncture of opercular lobes (Figs 1F, 9A, 10A). Eyespots along sides of median ridge. Palps tapered, about one third length of opercular lobes. Segment 1 (chaetiger 1) with lobe-shaped neuropodia (Fig. 9C), with thin and flattened capillary neurochaetae, with


HUTCHINGS ET AL.

serrations on their margins, decreasing in size towards fine tips (Fig. 10E). Segment 2 (chaetiger 2) with four pairs of large triangular-shaped lobes, connecting branchiae to neuropodia (Figs 1F, 9B, C). Neuropodia of chaetiger 1 and 2 not vertically aligned, those of chaetiger 2 situated more laterally (Fig. 9C), neurochaetae similar in structure. Fourty-five pairs of dorsal branchiae present from chaetiger 2, wide based (Fig. 9C), not meeting mid dorsally, continuing along entire length of body, but diminishing in size posteriorly so by posterior segments very small (Figs 1E, 9E). Segments 3–5 (parathoracic) with two types of notochaetae arranged in two rows, 11 lanceolate slightly curved forming a shallow scoop with slightly twisted frayed tips and blades and 11 short, thinner lanceolate chaetae inserted between each of larger ones (Fig. 10F). Segments 3–5, with lanceolate neurochaetae arranged in two tiers and significantly different in width, largest ones with smooth blades and smaller ones with textured blades. Parathoracic notochaetae more robust than parathoracic neurochaetae. Abdominal region with 43

FIGURE 9. Photos of Idanthyrsus willora n. sp. holotype (WAM V7852) A. Anterior end, latero-ventral view. B. Same, laterodorsal view. C. Detail of structures in anterior segments, lateral view. D. Detail of operculum, paleae and nuchal spines. E. Posterior end, lateral view. Abbreviations: b2, branchia segment 2; ca, cauda; cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1;ip; inner paleae; ll2, lateral lamellae segment 2; ns, nuchal spines (hooks); mo, median organ; op, outer paleae; tf, tentacular filaments.



19

chaetigers. Notopodia as transverse tori, with uncini, numbers per torus decreasing posteriorly. Each uncinus with two rows of vertical teeth, each with eight teeth (Fig. 10H). Neuropodia with capillaries becoming longer posteriorly, with thin, flattened blades ornamented with thecal laminar extensions, distally forming oblique rows with elongate thin tips (Fig. 10G), similar in structure on all abdominal chaetigers. Cauda smooth about one third the length of abdomen (Figs 1E, 9E).

FIGURE 10. SEM of Idanthyrsus willora n. sp. (AM W37779) A. Anterior end ventral view. B. Anterior end dorsal. C. Outer paleae. D. Inner paleae. E. Neurochaeta of segment 1. F. Lanceolate and capillary notochaetae from parathorax. G. Anterior abdominal neurochaetae, in detail. H. Uncini from mid abdominal. Scales A, B = 200 µm, C, D = 100 µm, E, F=30 µm, G = 20 µm, H = 10µm.


HUTCHINGS ET AL.

Variation. Material examined, including sexually mature females, varies from 10–35 mm in length without cauda, 2–6 mm in width, with 36–48 chaetigers, 14–18 pairs of outer paleae, 8–15 pairs of inner paleae, 8–17 pairs of opercular papillae, 1–4 pairs of nuchal hooks, 12–20 rows of compound tentacular filaments and 29–45 pairs of dorsal branchiae. These variations can be mainly attributed to size of the specimens, with larger animals tending to have more paleae, papillae, tentacular filaments and dorsal branchiae than smaller ones. Remarks. Idanthyrsus willora is described as a new species because of the presence of four pairs of lateral lobes on segment two, which have irregular margins. The number of lateral lobes is a feature only shared with I. saxicavus (Baird, 1863) although in the latter species these lobes are long and digitiform (Kirtley 1994: Fig. 6.15.1). These two species are also distinguished by the shape of outer paleae increasing in size slightly towards tips in the new species, while in I. saxicavus they are similar in size all along the paleae. Idanthyrsus willora n. sp., can be distinguished from the other Australian species by several features. Idanthyrsus willora has straight denticles increasing in length towards the tip on the outer paleae while there are curved denticles similar in length along the blade in I. australiensis. The number and shape of lateral lobes on segment 2 also varies, (four large triangular ones in I. willora n. sp., two poorly developed in I. nesos n. sp., and three triangular ones in I. australiensis). Idanthyrsus willora n. sp., also differs from I. nesos n. sp., in having branchiae continuing to the far posterior segments. Idanthyrsus willora n. sp., can be separated from other species recorded from the Indo-Pacific by the absence of branchiae on posterior segments, whereas they are present in L. boninensis Nishi & Kirtley, 1999, described from Japan. Another Japanese species, I. okudai Kirtley, 1994, can be separated from I. willora n. sp., based on the number of pairs of lobes on segment 2 (four in I. willora n. sp., and only two in I. okudai). Two other species from from Sri Lanka, I. kornickeri Kirtley, 1994 and I. bicornis (Schmarda 1861), both poorly described (see Table 1), are distinguished from I. willora n. sp., in the shape and arrangement of the denticles on the outer opercular paleae which are curved and similar in length along the blade in I. kornickeri, in contrast to those from I. willora n. sp., which are straight and increase in length towards the tip. Idanthyrsus bicornis has these denticles arranged opposite to each other along the shaft, and in I. willora they are arranged in an alternate pattern on both margins. More over, the nuchal hooks in I. bicornis are limbate whereas they are not in I. willora n.sp. Distribution. Western Australia and Northern Territory. Habitat. Specimens found in the intertidal but there is no information available as to whether this species is gregarious or occurs as single individuals, since specimens examined had all been removed from their tubes. Some of the records indicate they were attached to mollusc shells. Etymology. The specific name willora is the Aboriginal word for the type locality Quondong (Endacott 1973).

Lygdamis Kinberg, 1867 Lygdamis.—Kirtley, 1994: 116. Capa et al. 2012: 261–263. Type species. Lygdamis indicus Kinberg, 1867 by monotypy. Type locality. Bangka Straits, Java, Indonesia.

Diagnosis. Operculum with separated lobes and distal end sloped posteriorly. Opercular paleae straight and smooth, the outer ones flattened and the inner ones cylindrical, resembling spines. Remarks. The monophyly of Lygdamis was not assessed in the phylogenetic analyses undertaken by Capa et al. (2012) but all species currently assigned to the genus conform to the above diagnosis. The genus is known from 19 species (Kirtley 1994; Lechapt & Kirtley 1998; Nishi & Kirtley 1999) including the new species described here. Kirtley (1994) suggested that the shape and colouration of the median organ are useful characters especially in live material, but after fixation and preservation they become so distorted as to loose their usefulness.

Lygdamis augeneri Kirtley, 1994 Figures 6B, 11A–D, 12–13, Table 2 Eupallasia giardi. — Augener, 1927: 236, fig. 14a–c; 1934: 92; not McIntosh, 1885. Pallasia giardi. — Ehlers, 1920: 64; not McIntosh, 1885. Lygdamis augeneri. — Kirtley, 1994: 119–120, figs 7.1.1–2.



21

Material examined. HOLOTYPE: As Eupallasia giardi Augener, 1927 (identified by Kirtley, 1994) off Disaster Bay, New South Wales: Australia: 37°05'S 150°05'E; 01.x.1914; 53–73 m; ZMH V-9569; Th. Mortensen's Endeavour, 20 mm in length (including damaged posterior end), 2 mm in width. PARATYPES: 3, from same sample as holotype.

FIGURE 11. Line drawings of Lygdamis augeneri A. Anterior end, ventro-lateral view, paratype (ZMH V9569). B. Anterior end lateral, paratype (ZMH V9569). C. Posterior end, lateral view AM W27564. Lygdamis giardi D. Anterior end, dorsal view, holotype (BMNH 1885. 12.1.6). E. Entire body, lateral view (AM W6191). Lygdamis wambiri n. sp., holotype (QM G10505) F. Anterior ventral view. G. Lateral dorsal view. Scales A, C= 3 mm, B, G = 5 mm, D, F, =4 mm, E = 6 mm.


HUTCHINGS ET AL.

FIGURE 12. Photos of Lygdamis augeneri (AM W27564) A. Anterior end, ventral view. B. Detail of opercular structures, ventral view. C. Anterior end lateral view. D. Operculum, thoracic and parathoracic segments, latero-dorsal view. E. Detail of structures, anterior end, dorsal view. F. Anterior end, dorsal view. Abbreviations: bo, buccal organ; cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1; ip; inner paleae; li, lips; ll2, lateral lamellae segment 2; ns, nuchal spines; opa, opercular papillae; tf, tentacular filaments; pa: palps.

Additional material examined. New South Wales: (Tasman Sea) off The 156°10'40.8"E, 02.v.1989, 133 m, 1, AM W27564*, 16 mm long, 1 mm wide, 24 (mounted for SEM). Description. Holotype flesh coloured, no pigmentation present. Body elongate Operculum with completely separated lobes, with distal end sloping; longer than wide


Entrance, 33°14'12.6"S segments; AM W27565 and slender (Fig. 11A). (Fig. 11A, B). Length of


23

operculum more than one third the length of the body (Fig. 11B, C). Paleae displayed in two rows, outer row with 42 pairs of paleae, with flattened shafts, smooth margins with tips curved outwards (Fig. 13C); inner row with 30 pairs of paleae, cylindrical and tapering slowly to blunted tips (Fig. 13C). Opercular papillae, 12 pairs, peripheral to outer row of paleae on each lobe, elongate, increasing in length towards nuchal spines (Figs 12C, 13B), pair adjacent to nuchal spines about one quarter length of outer paleae. One pair of nuchal spines, strongly recurved, margins smooth, with pointed tips (Figs 11A, 12D, 13B). Tentacular filaments compound arranged in 19 horizontal rows. Eye spots present along sides of median ridge. Pair of elongate grooved palps with crinkled margins on either side of median ridge, about one quarter length of opercular lobes. Segment 1 with lobe-shaped neuropodia (Fig. 12B–E), capillaries absent. Segment 2 with three triangular lateral lobes on each side of body (Fig. 12E), with fine capillaries arranged in compact fascicle, inserted in neuropodia, posterior to U-shaped buccal organ (Fig. 12B). Eight pairs of dorsal branchiae present from segment 2, with narrow base and tapering gently to a fine tip, slightly ridged, size decreasing posteriorly, not meeting mid dorsally (Fig. 11A). Segments 3–6 (parathoracic) with notochaetae arranged in two transverse rows, six long lanceolate, with tapering frayed tips, blades slightly concave and textured, and six capillaries inserted imbetween (Fig. 13D). Segments 3–6 with two types of neurochaetae, six lanceolate, with frayed twisted tips, and compact thecae and fine capillaries with thin flattened blades with short narrow thecae, alternating with lanceolate (Fig. 13E). Parathoracic notochaetae more robust than neurochaetae. Abdominal region with 19 chaetigers. Notopodia as transverse tori, with number of uncini decreasing posteriorly. Each uncinus with two vertical rows of eight teeth (Fig. 13F). Neuropodia with capillaries arranged in discrete fascicle, with thin, flattened blades with short, narrow thecal rings. Cauda smooth, about length of last five chaetigers (Fig. 11C).

FIGURE 13. SEM of Lygdamis augeneri (AM W27464) A. Anterior end, dorsal view. B. Nuchal spines (hooks) and palps. C. Outer paleae. D. Lanceolate and capillary notochaetae, parathoracic region. E. Lanceolate and capillary neurochaetae, parathoracic region F. Abdominal uncini. Scales A = 200 µm, B = 20, C, D = 20 µm, D = 100 µm, E, F = 10 µm.

Variation. Material examined, including sexually mature specimens, varies from 15–18 mm in length without cauda, 2–3 mm in width, with 19–23 chaetigers, 42–44 pairs of outer paleae, 30–36 pairs of inner paleae, 6–8 teeth on each row of uncini and 15–19 pairs of tentacular filaments. Pigmentation present along the rows of some tentacular filaments. We suggest that these variations are to some extent size dependent and pigmentation may be related to methods of preservation and subsequent storage.


HUTCHINGS ET AL.



25

Remarks. Kirtley (1994) described this species based on material examined by Augener (1927) who had identified it as L. giardi McIntosh, 1885. The two species differ not only in the range of the number of pairs of outer paleae, 42–44 in L. augeneri and 16–20 in L. giardi, but also in the shape of the outer paleae. I. augeneri has outer paleae with curved flattened tips (Fig. 13C) whereas those of L. giardi have asymmetrical flattened pointed tips (Fig. 15C). Distribution. South-eastern Australia. Habitat. Shelf depths of 53–133 m.

Lygdamis giardi (McIntosh, 1885) Figures 6B, 11 D, E, 14, 15, Table 2 Sabellaria (Pallasia) giardi McIntosh, 1885: 421, pl. 26, figs 13–15, pl. 47, fig. 7, pl. 26A, figs 13–15. Pallasia giardi.— McIntosh, 1922: 64; Caullery, 1913: 200. Lygdamis giardi.— Johansson, 1927, 86; Imajima & Hartman, 1964: 324; Kirtley, 1994: 126.

Material examined. HOLOTYPE: as Sabellaria (Pallasia) giardi McIntosh 1885; off Port Jackson, New South Wales: 33°51'S 151°22'E; 03.vi.1874; 64 m, BMNH 1885.12.1.6. Specimen described as 7 mm in length, 2 mm in width and damaged, but currently in worse condition and only anterior end and posterior chaetigers are in vial. Additional material examined. New South Wales: Sydney, Malabar, 33°49'30"S 151°21'36"E, 27.ii.1973, 66 m, 8, AM W6191*; 22.v.1972, 28 m, 6, AM W6185*; North Head, 33°49'25"S 151°20'43"E, 27.ii.1973, 63 m, 81, AM W26682 (mounted for SEM); 7, AM W6192*, 2 km east of Long Bay, 33°58'S 151°15'E, 66 m, 2, AM W6188*, 5, AM W6189*; off Maroubra, 33°57'06"S 151°17'37"E, 20.vii.1972, 58 m, 1, AM W6190*; Tasman Sea, off Sydney, 33°53'06"S 151°21'54"E, 30.iii.1972, 76.8 m, 2, AM W27553*, from gritty calcareous mud; off Eden, 36°39'30"S 150°37'48"E, 01.iv.1972, 86 m, 1, AM W27554, from muddy quartz sand; off Broken Bay, 33°37'–39'S 152°04–02'E, 10.xii.1980, 1, AM W27555*; Sydney, off North Head, 33°48'46"S 151°20'59"E, 60 m, 13.iv.1989, 1, AM W20677*, from sandy substratum; 15.ii.1973, 66 m, 2, AM W27556*; Malabar, 33°58'S 151°19'E, 1973, 1, AM W27557*; 2, AM W27558*; 45 m, 1, AM W27559; 17.v.1972, 69 m, 3, AM W27560*; 22.v.1972, 75 m, 1, AM W27561; 02.i.1973, 54.9 – 60.3 m, 4, AM W27562*; off Tuggerah Lakes, 33°20'S 151°39'24"E, 11.iii.1972, 60 m, 5, AM W27563. Tasmania, Bass Strait: King Island, 35 km northeast of Cape Wickham, 39°16'00"S 144°05'24"E, 23.xi.1981, 82 m, 1, MV F78887*, from sandy shell. Northern Territory: 11 km, NE of Cape Ford, Anson Bay, Beagle Gulf, 13˚25'02"S 129˚55'.98"E, 16 m, 01.x.1993, NTM 23481. Description. As the holotype is incomplete and cut into portions, this description is largely based on material collected from off the NSW coast in the region of the type locality. Body robust, preserved material pale cream, paleae golden with dark brown pigmentation on opercular lobes and tentacular filaments in some specimens. Operculum completely divided into two free long lobes with distal ends oblique (Figs 11E, 15B). Operculum with two rows of paleae, outer row with 20 pairs of paleae, cylindrical at base and flattened, with nearly symmetrical pointed tips, with smooth margins (Fig. 15C); inner row with ten pairs of paleae, smooth cylindrical shafts with blunt tips (Fig. 15D). Opercular papillae, 11 pairs, peripheral to outer paleae of each lobe (Figs 11D, E, 14E, F). One pair of nuchal spines on each side (Figs 11D, 14F), stout, strongly recurved without sharp limbations on concave margin. Tentacular filaments compound, arranged in 15–19 rows (Fig. 14C). Median organ present, with eyespots on either side. Pair of ringed palps, less than quarter length of opercular lobes (Fig. 14B). Segment 1 with lobe shaped neuropodia (Fig. 14A), chaetae absent. Segment 2 with three pairs of triangular shaped lateral lobes, connecting branchiae to neuropodia (Figs 11E, 14D). Fourteen pairs of branchiae present from segment 2 (Fig. 14D) not extending to posterior segments. Branchiae, taper gently and evenly to subacute tip (Fig. 14D). Segments 3–6 (parathoracic), with two types of notochaetae arranged transversely, 7–8 lanceolate, with frayed twisted tips, interspersed with 7–8 capillaries (Fig. 15E). Segments 3–6 with two types of neurochaetae arranged transversely, five lanceolate and five thinner capillaries with compact thecae alternating with lanceolate (Fig. 15F). Abdominal notopodia with uncini decreasing in number posteriorly, each uncinus with two vertical rows each with about 7–9 teeth (Fig. 15H). Neuropodia with capillaries, ornamented with irregular thecal laminar extensions (Fig. 15G), similar in structure on all abdominal chaetigers. Cauda smooth about the length of five posterior abdomen segments (Figs 11E, 14H).


HUTCHINGS ET AL.

FIGURE 14. Photos of Lygdamis giardi A. Anterior segments, ventral view (NTM W1121). B. Same with opened opercular lobes (NTM W1121). C. Operculum, lateral side (AM W6965). D. Anterior segments, lateral view (NTM W1121). E. Detail of opercular paleae and papillae (AM W6965). F. Base of operculum, dorsal view (NTM W1121). G. Parathoracic neurochaetae (NTM W1121). H. Posterior end, lateral view (NTM W1121). Abbreviations: b2, branchia segment 2; bo, buccal organ; ca, cauda; cn1, cirrus neuropodia segment 1; li, lips; ll2, lateral lamellae segment 2; ns, nuchal spines ; op, outer paleae; opa, opercular papillae; tf, tentacular filaments; pa, palps.



27

FIGURE 15. SEM of Lygdamis giardi A. Anterior end, ventral view (AM W6192). B. Anterior end, dorsal view (AM W6192). C. Outer paleae (AM W26682). D. Inner and outer paleae (AM W26682). E. Lanceolate-and capillary notochaetae of parathorax (AM W6192). F. Lanceolate and capillary neurochaetae of parathorax (AM W6192). G. Anterior abdominal neurochaetae (AM W26682). H. Abdominal uncini (AM W6192). Scales A, B = 300 µm, C = 30 µm, D = 40 µm, E = 60µm, F = 6 µm, G, H = 3 µm.

Variations. Material examined, including sexually mature specimens, varies from 12–43 mm in length (without cauda), 2–5 mm in width, with 17–32 chaetigers, 16–20 pairs of outer paleae, 12–14 pairs of inner paleae, 8–12 pairs of opercular papillae, 17–19 pairs of tentacular filaments and 9–10 pairs of dorsal branchiae.


HUTCHINGS ET AL.

Remarks. McIntosh (1885) dissected the holotype and cut up the body into sections and not all are still present in the vial, hence we have also illustrated a complete specimen from the same location (AM W6191). The diagnostic characters of L.giardi are the presence of 16–20 pairs of outer paleae which are straight, nearly symmetrical with distally smooth margins and pointed tips, 15–19 rows of tentacular filaments and three pairs of lateral lobes on segment 2. Lygdamis giardi can be separated from L. augeneri by the number of outer paleae, 16– 20 and 42–44 respectively and by the structure of their tips L. giardi having flattened nearly symmetrical tips whereas L. augeneri has sharply bent outer paleae with broad, blunt flattened tips (see Fig. 15C and 13C). Distribution. Lygdamis giardi has a discontinuous distribution around Australia, and this may represent a true record or reflect the lack of collecting on the shelf around Australia; currently reported from New South Wales, Tasmania and Northern Territory. This species has also been recorded from the Moluccas Islands, Indonesia (Caullery, 1913) although this material has not been checked. Habitat. Inshore at depths of 16– 86 m, in sediments ranging from coarse sand to muddy sand. Most records of solitary individuals but one sample of 81 individuals suggesting they can form small colonies.

Lygdamis wambiri n. sp. Figures 6B, 11F, G, 16, 17, Table 2. Material examined. HOLOTYPE: QM G10505*, 26+ mm in length (incomplete), 3 mm in width, 26+ segments (incomplete). PARATYPES: QM G10338*, 14+ mm in length (incomplete), 2 mm in width, 24+ segments (incomplete), plus one specimen mounted for SEM. All material from Queensland: Moreton Bay, Peel Island, 0.8 km southeast of South West Rocks, 27°30'S, 153°21'E, from shell, grit and sand, 7.62 m, collected vi.1970, 7 m, and iii.1970. Additional material examined. ?Northern Territory: New Year Island, 10°55'S 133°02'E, 14.x.1982, LWM, 1, NTM W1121*, from coarse sandy bottom.

FIGURE 16. Photos of Lygdamis wambiri n. sp. holotype (QM G10505) A. Anterior end, lateral view. B. Same, dorsal view. C. Base of operculum and anterior segments, dorsal view. Abbreviations: b2, branchia segment 2; bo, buccal organ; cn1, cirrus neuropodia segment 1; ip, inner paleae; ll2, lateral lamellae segment 2; ns, nuchal spines; op, outer paleae; opa, opercular papillae; tf, tentacular filaments.



29

Description. Holotype, stout, colourless with golden yellow paleae and dark brown nuchal hooks.Operculum completely divided into two free divergent lobes, longer than wide (Figs 11F, H, 16A, B) with distal end sloping with two rows of paleae (Fig. 17B). Outer row with 29 paleae, blades finely textured, colourless, margins straight, symmetrical, tapering to acute tip with small distal denticles (Fig. 17B, C), inner row with 12 pairs of spines, yellow, smooth and acute tipped (Fig. 17D); outer paleae longer than inner (Fig. 17B) but thinner. Opercular papillae, 18 pairs peripheral to outer row of paleae on each lobe, about one third length of outer paleae (Figs 11G, 16C, 17B). One pair of nuchal hooks present, strongly recurved, margins smooth, convave with pointed tips (Figs 11F, 16B, C). Tentacular filaments compound, arranged in about 30 horizontal rows (Fig. 16A, B). Medial organ present at dorsal base of opercular lobes (Figs 11F, 16A, 17A), palps shorter than a third of opercular lobes. Eyespots absent. Segment 1 with lobe shaped neuropodia (Fig. 16A), chaetae absent. Segment 2 with three pairs of triangular lateral lobes (Figs 11F, G, 16B), with fine capillaries with plumose margins and extended fine tips, in compact fascicle, inserted posteriorly to U-shaped buccal organ. Twelve pairs of branchiae present from segment 2. Branchiae with broad base, tapering gently to rounded tip (Figs 11G, 16B), from chaetiger 7 onwards, triangular, larger in size in anterior abdominal segments, absent from posterior segments of incomplete type material. Segment 3–6 (parathoracic) with chaetae inserted in two transverse rows with eight lanceolate chaetae with frayed twisted tips (Fig. 17E), and fine capillaries with compact thecae interspersed between them (Fig. 17E), we suspect that the lanceolate chaetae can retract but not the capillaries. Segments 3–6 with two types of neurochaetae arranged in oblique transverse row, five lanceolate with fine capillaries. Abdominal region with 20 chaetigers, but holotype is incomplete lacking cauda. Notopodia as transverse tori, with numbers of uncini per row decreasing posteriorly, uncini with two vertical rows of teeth, each row with eight teeth. Neuropodia with capillaries arranged in discrete fascicles with compact theca, distal margins becoming more elongate towards tip (Fig. 17F).

FIGURE 17. SEM of Lygdamis wambiri n. sp. (QM G 10338) A. Anterior end ventral view. B. Anterior end dorsal view. C. Outer paleae. D. Inner paleae. E. Lanceolate and capillary notochaetae of parathorax. F. Abdominal neurochaetae. Scales A, B = 200 µm, C, D = 30 µm, E = 20 µm, F = 6 µm.

Variation. Paratypes exhibit the following variation, number of rows of tentacular filaments (15–30), outer paleae (27–29 pairs), inner paleae (12–16 pairs). A specimen from Northern Territory is 25 mm in length, 4 mm in width and has 32 pairs of outer paleae, 20 pairs of inner paleae, one pair of hooks, 26 pairs of opercular papillae and parathoracic segments with 6–7 notopodial lanceolate chaetae. This specimen has considerably more inner


HUTCHINGS ET AL.

paleae than the holotype which has 12 pairs and the paratype mounted for SEM has 13. At this stage we are referring the Northern Territory material to the species, Lygdamis wambiri n. sp., as it certainly does not represent either of the other two species of Lygadamis recorded from Australia, but additional material from this locality should be examined to determine if this is correct or whether material from the Northern Territory represents another undescribed species of the genus. Remarks. Lygdamis wambiri n. sp., is distinguished from other species in the genus by a combination of characters: 27–29 pairs of outer paleae, 12–16 pairs of inner paleae, three pairs of lateral lobes on segment 2 and having straight markedly asymmetrical outer paleae with distally denticulate margins (Table 2). Lygdamis wambiri n. sp., can be distinguished from the other species of the genus recorded from Australia by the shape of the distal margins of the outer paleae, in L. wambiri n. sp., they are finely denticulate whereas they are smooth in the other Australian species. Lygdamis wambiri n. sp., has short inflated lanceolate notochaetae on parathoracic segments, whereas the two Australian species have longer less inflated lanceolate notochaetae compared to those of the other two species. Lygdamis wambiri n. sp., has branchiae present on posterior segments whereas L. augeneri and L. giardi lack them posteriorly. The only other species which appears to have distal margins of the outer paleae denticulate is L. robinsi Jeldes and Lefevre, 1959 described from off Angola, but this species is poorly known with regard to other characters. Lygdamis wambiri n. sp., can be separated from other species described from the Indo-Pacific by the number of opercular papillae: L. bhaudi Kirtley, 1994, described from Madagascar, has 3–4 pairs whereas L.wambiri has 12–17. Lygdamis curvatus (Johansson, 1922) and L. japonicas Nishi & Kirtley, 1944, both described from Japan, have smooth distal tips of outer paleae in contrast to the denticulate tips of L. wambiri n. sp. Lygdamis ehlersi Caullery, 1913, described from Indonesia has only five pairs of outer paleae in contrast to the 27–29 pairs present in the new species. Distribution. Moreton Bay, Queensland, with a possible record from Northern Territory. Habitat. Inshore in sheltered shallow waters (around 7 m). Etymology. The specific name wambiri is an Aboriginal word for the coast (Endacott 1973) and refers to the extensive coast line of Moreton Bay a shallow protected bay, the type locality of the species.

Phalacrostemma Marenzeller, 1895 Phalacrostemma Marenzeller, 1895:19; Kirtley, 1994: 147; Capa et al., 2012: 267–268. Type species. Phalacrostemma cidariophilum Marenzeller, 1895, by monotypy. Type locality. Off Pelagossa Island, Adriatic Sea.

Diagnosis. Buccal flaps present, arrangement of outer paleae in a spiral on each lobe, in most species. Remarks. Kirtley (1994) recognised 10 species within the genus and four species have been described since then (Lechapt & Kirtley 1998 and present paper). This represents the first record of the genus from Australia and as this genus currently is only known from deep water, this is not surprising given the limited offshore sampling carried out in Australia. As can be seen from Table 3, many species of the genus are extremely poorly known and have been separated based solely on paleal features. A complete revision of the genus is therefore urgently needed.

Phalacrostemma maloga n. sp. Figures 6A, 18, 19, 20, Table 3 Material examined. HOLOTYPE: AM W27566*, New South Wales: Tasman Sea, off Newcastle, 33°30S 152°06'E, iv.1971, 548.6 m, in 2 pieces 10+-5+ mm in length, 2 mm in width, 15-4 chaetigers, attached to mollusc shell. PARATYPES: Tasman Sea, off Newcastle, 32°30'S 152°06'E, iv.1971, 548.6 m, 1, AM W27567; 457.2– 502.9 m, 6, AM W27568 * attached to mollusc shell; 33° 40' S 152° 45' E, 15 vi.1971, 6, AM W27569; 33° 30'S 152° 06' E, iv.1971, 1, AM W26681 (mounted for SEM). Additional material examined. MV F78859, Stn S05/84 29 from Tasman Sea, eastern slope 70 km S of Gabo Island, Victoria (1 specimen).



31

FIGURE 18. Line drawings of Phalacrostemma maloga n. sp. A. lateral dorsal view, entire animal, paratype (AM W27567). B. Anterior end, dorsal view, holotype (AM W27566). C. Anterior end, anterior plan view semi-diagrammatic (paleae shown as “scars” only) 1 mm, (AM W27568). Scales A = 4 mm, B = 2 mm, C = 1 mm.

Description. Holotype with body stout, tapering to elongated abdomen, reddish brown with darker areas along noto- and neuropodia. Operculum completely divided into two free short lobes with distal end perpendicular to longitudinal axis (Figs 18A, B, 19A–D). Operculum with outer row of 12 pairs of paleae, arranged spirally (Fig. 18C); paleae yellow as simple spines with acute tips and compact thecae with margins straight and not expanding (Figs 19E, 20B, C); inner row with two pairs of short spines, yellow and tapering tip (Fig. 18C). Ten pairs of robust and tapering opercular papillae, peripheral to outer paleae (Figs 18C, 19B, C), almost as long as outer paleae. Five


HUTCHINGS ET AL.

FIGURE 19. Photos of Phalacrostemma maloga n. sp. A. Anterior segments, ventral view. B. Same, different specimen. C. Same, left lateral view. D. Same, right lateral view. E. Opercular paleae. Abbreviations: b2: branchiae segment 2; bf: buccal flaps; bo, buccal organ; chn1, chaetae neuropodia segment 1; cn1, cirrus neuropodia segment 1; ll2, lateral lamellae segment 2; ns, nuchal spine; op, outer paleae; opa, opercular papillae; tf, tentacular filaments; pa, palps. Holotype AM W27566.

pairs of flattened nuchal hooks, with limbation present on concave side (Figs 18B, C, 19D, 20B). Medial organ present at dorsal junction of opercular lobes (Fig. 18C, see Capa et al., 2012). Eyes not observed. Three pairs of simple tentacular filaments (Fig. 19B). A pair of buccal flaps below tentacular filaments (Fig. 19A). Palps shorter



33

than length of operculum (Fig. 19B). Segment 1 (chaetiger 1) with long and tapering neuropodial lobe (Fig. 19A) and capillary neurochaetae (Fig. 19B, C). Segment 2 (chaetiger 2) with two triangular lateral lobes connecting branchiae to neuropodia (Figs 18B, 19C). Eight pairs of dorsal tapering branchiae present from chaetiger 2, not meeting mid-dorsally (Fig. 18B), reducing in size posteriorly but present on posterior segments. Segments 3–6 (parathoracic) with two types of notochaetae inserted in transverse rows with two types of notochaetae; eight lanceolate and fine capillaries interspersed (Fig. 20D, E). Segments 3–6 with two types of neurochaetae arranged transversely, four lanceolate and fine short capillaries interspersed (Fig. 20F). Notopodial lanceolate much stouter than those in neuropodia (Fig. 20D–F). Abdominal notopodia as erect expanded tori, with uncini decreasing in numbers posteriorly. Each uncinus with three vertical rows of teeth imbricated (Fig. 20H). Neuropodia with long fine capillaries with compact thecae arranged in discrete fascicles in two tiers (Fig. 20G). Cauda smooth and about one third length of abdomen (Fig. 18A).

FIGURE 20. SEM of Phalacrostemma maloga n. sp. (AM W26681). A. Anterior end ventral view. B. Anterior end dorsal view. C. Outer and inner paleae. D. Lanceolate and capillary notochaetae of parathorax, anterior end, dorsal view. E. Same, in detail. F. Lanceolate and capillary neurochaetae of parathorax. G. Posterior abdominal neurochaetae. H. Posterior abdominal uncini. Scales A, B = 100 µm, C = 20 µm, D, F = 10 µm, E, G = 3 µm, H = 2 µm.


HUTCHINGS ET AL.



35

Variations. The paratypes including sexually mature females, vary from 7–18 mm in length (without cauda) none of which is complete, 2 mm in width with 12–22 chaetigers, 12–16 pairs of outer paleae, 2–3 pairs of inner paleae, 7–13 pairs of opercular papillae, and with 3–9 pairs of dorsal branchiae. All these variations could be size dependent. None of the material is well preserved and fragile. Remarks. Phalacrostemma maloga n. sp., is characterised by a unique combination of: 12–16 pairs of outer paleae with straight edges and without expanded thecae, five pairs of nuchal spines, three pairs of tentacular filaments, buccal flaps present and with some abdominal uncini provided with three vertical rows of teeth. Other species without expanded thecae on the outer paleae (see Table 3) include P. setosa Treadwell, 1906 described from the Caribbean which has irregular edges and P. cidariophilum Marezeller, 1895 described from the Northern Adriatic Sea, which has straight edges but lacks tentacular filaments. The closest species geographically is P. tenue Lechapt & Kirtley, 1998, described from deep water off New Caledonia but this has only two pairs of nuchal spines whereas P. maloga n. sp., has five pairs. Most species of this genus, which typically occur in deep waters (see Table 3), are poorly known. We suggest that the number of pairs of lateral lobes on segment 2, the presence of buccal flaps and the number of pairs of tentacular filaments are useful characters to distinguish between species. The dentition of the abdominal uncini, with three rows of teeth instead of the two rows typical of other sabellariids, may be a synapomorphy for the species. But it could have also been overlooked in other species of the genus and therefore needs further study. Distribution. South-eastern Australia, type known only from type locality and from a single record from south of Gabo Island, off the coast of Victoria.. Habitat. Continental shelf in depths of 457–548 m and attached to mollusc shells in soft sediment substrates forming small colonies. Etymology. The specific name maloga is an Aboriginal word for deep water (Endacott 1973) and refers to the habitat of this species.

Sabellaria Lamarck, 1812 Sabellaria Lamarck, 1812: 96; Kirtley, 1994: 45–46; Nishi et al., 2010: 7 ; Capa et al., 2012: 268–269. Type species. Sabella alveolata Linnaeus, 1767, by monotypy. Type locality. Europe, Hartman (1959).

Diagnosis. Operculum completely divided in two symmetrical lobes, inner paleae arranged in two rows (with mid and inner paleae) and three parathoracic segments. Remarks. The monophyly of Sabellaria has not being assessed in the phylogenetic analysis undertaken by Capa et al. (2012) and the above diagnosis includes characters that are shared by all species in the genus. The total number of species of Sabellaria described to date (including those in Nishi et al. 2010, Santos et al. 2011 and herein) are 39. Features such as the shape and ornamentation of paleae and nuchal spines (as in Nishi et al. 2010, Santos et al. 2011) together with some others have been used for the comparison of the species herein (Table 4). According to Kirtley (1994), all species within the genus are gregarious and capable of forming large colonies and reefs along shores where suitable hydrodynamic and sedimentary conditions occur but the new species described below are known from only a few solitary individuals. First record of the genus in Australian waters.

Sabellaria kooraltha n. sp. Figures 6B, 21, 22, 23, Table 4 Material examined. HOLOTYPE: 1, AM W27333 *, 18 mm long, 2 mm wide, 22 chaetigers. PARATYPE: AM W27195*, 19 mm long, 3 mm wide, 22 chaetigers, from Albany Passage, Queensland, Australia, 10°45'S 142°37'E, 10 m; paratype AM W37820.001 (mounted for SEM) Shell Island, East Arm, Darwin Harbour, Northern Territory, St. RH88/37, 12˚ 29.4''S 130˚ 53.2''E, rocky intertidal, 18 iii. Mar 1988. Additional material examined. Northern Territory: West of Elizabeth Reef, Cape Hotham, Beagle Gulf, 12˚4.02'S 131˚20.04'E, 21 m 29.x.1993, 1, NTM W23479, 09.x.93, 1, NTM W23477; 2 nm E of Quail Isl., Bynoe


HUTCHINGS ET AL.

Harbour, Beagle Gulf, 12˚31.02'S 130˚28.98'E, 28 m, 6.x.1993, 1, NTM W23478; East Arm, Shell Island, Darwin Harbour, 12°29'S 130°53'E, 18.iii.1988, low water, rocky intertidal, 1, AM W37820 *, (mounted for SEM); Darwin Harbour, 12˚32.72'S 130˚51.32'E, DW 105A, 7 m, coll. 24.iii.1994, 1, NTM W15033; 12˚32.72'S 130˚51.32'E, DW1054, 7 m, coll. 24.iii.1994, 1, NTM W15031; 12˚36.17'S 130˚46.87'E, DW 76A, 4 m, coll. 17.iii.1994, 1, NTM W15023; 12°32.9'S 130°50.02E, 1, D103A, 8 m, coll. 15.vi.1993, 1, NTM W10042; 12°31.98'S 130°50.09'E, DH 100A, 24.iii.1994, 15 m, coll. 24.iii.94, 1, NTM W15030; 12˚29.27'S 130˚49.48'E, DW 204A, 14 m, coll. 25.iii.1994, fine & coarse sand, 1, NTM W15026; 12˚30.07'S 130˚55.88'E, DW 187A, 4 m, coll. 22.iii.1994, 2, NTM W15027; 12˚26.93'S 130˚46.5'E, DW 544, 14 m, coll. 16.iii.1994, 1, NTM W15032; 12˚34'S 130˚52.22'E, DW 116A, 12 m, coll. 18.iii.1994, 1, NTM W15024; 12˚34'S 130˚52.22'E, DW 116A, 18.iii.1994, 12 m, 1, NTM W15035, 12˚31.68'S 130˚51.3'E, DW 101A, 8 m, coll. 24.iii.1994, 1, NTM W15028; 12˚31.7'S 130˚51.28'E, 8 m, 14.vii.1993, 1, NTM W23482; 12˚31'54''S 130˚50'25''E, 24.iii.1994, 1, NTM W15029; 12˚27'57''S 130˚46'30''E , 08.vi.1994, 1, NTM W10046; 12˚28'57''S 130˚50'25''E, 17.iii.1994, 1, NTM W15034; Melville Bay, Gove, 12˚11.7'S 136˚41.3'E, 8.vii.1991,7.5 m, 1, NTM W16880, 1, NTM W16887, 1, NTM W16890; Melville Bay, Gove, 12˚12'31''S 136˚40'30''E, 9.vi.1991, 1, NTM W16882, 1, NTM W6881; 12˚10'31''S 136˚39'E, Nov 1991–March 1992, 1, NTM W08265; NW of Granite Islet, Melville Bay, Gove, 12˚14'S 136˚40'E, 22.iii.1992, 12.9 m, 1, NTM W16891; Bing Bong, McArthur River, Gulf of Carpentaria, 15˚37'S 136˚23'E, ix. 1992–iii.1993, 8 m, 1, NTM W007737, 1, NTM W16899. Description. Holotype with speckled faint brown pigment on ridged cilated dorsum (Fig. 21C), rectangular pigment patches across ventrum of chaetiger 5 (3rd parathoracic). Operculum similar in width to length, with lobes completely separated, and three rows of golden paleae arranged in concentric semicircles (Figs 21B, 23A). Outer row with 12 geniculate paleae with flattened blades, smooth lateral margins and serrated distal margins with a middle tapering denticle occupying half of distal end which has marked surface ornamentation (Figs 21B, 22G, 23D, E, 22D, E). Twelve pairs of middle geniculate paleae, with smooth concave shafts and tapering blades pointing towards to outer edge of operculum (Figs 21B, 22H, 23C, F). Fourteen pairs of inner paleae, strongly geniculated, concave blades, similar to middle paleae but wider, shorter and pointing inwards to center of operculum (Fig. 21B). Outer paleae longer than those of inner rows. Opercular papillae, 19 pairs, short, peripheral to outer paleae on each lobe (Figs. 21B, C, 22D, 23B). Three pairs of nuchal spines present (four in paratype), stout, flattened, slightly curved, with smooth blunt tips (Figs 21B, 22I, 23G). Tentacular filaments, compound, arranged in 14 horizontal rows (Fig. 22A). Median ridge and median organ present. Eyespots present. Palps slightly longer than opercular lobes. Segment 1 (chaetiger 1) with lobe-shaped neuropodia, with capillary neurochaetae, on either side of U-shaped buccal organ (Fig. 22B). Segment 2 (chaetiger 2) with one elongate triangular shaped lateral lobe, connecting branchiae to neuropodia, with fine capillary neurochaetae (Figs 21A, C, 22E, F). Neuropodia of chaetigers 1 and 2 not vertically aligned, those of chaetiger 2 situated more laterally (Figs 21B, 23H). Sixteen pairs of dorsal branchiae present from segment 2 (Figs 22F, 23H), continuing along entire length of body, with broad base and strongly ciliated ridges and slightly pigmented, all detached except one, which tapers distally with filiform tip, more remain on paratype (Figs 21A, 23H). Segments 3–5 (parathoracic) with two types of notochaetae arranged transversely, seven (many represented only by stumps) lanceolate with frayed tips and short, flattened smooth shafts with strongly frayed tips, and fine short capillaries with expanded theca inserted between lanceolate (Fig. 23I). Segments 3–5 with two types of neurochaetae arranged in compact fascicle, with seven lanceolate and seven fine capillaries (Fig. 23H). Parathoracic notopodia larger than neuropodia (Fig. 22J). Abdominal region with 16 chaetigers. Notopodia as elongate, erect tori, with long-handled uncini, numbers per torus decreasing posteriorly, each uncinus with two vertical rows, each with seven teeth. Neuropodia becoming considerably elongated posteriorly (Fig. 22I) with ventral bundle of capillaries (Fig. 23J). Cauda smooth and more than half length of abdomen (Figs 22I,K, 23L). Paratype a gravid female with eggs released from between posterior abdominal notopodia. Variations. Material examined varies from 4–12 mm in length, 1–2 mm in width with 12–22 chaetigers, but much of the material is posteriorly incomplete, 12–16 pairs of outer paleae, 10–12 pairs of middle paleae, 16–19 pairs of opercular papillae and 3–4 pairs of nuchal spines. Remarks. Sabellaria kooraltha n. sp., is characterised by the following combination of characters: 12–16 pairs of outer paleae with a smooth median distal spike which has marked thecal bands, and two smaller, smooth basal, lateral spines (Fig. 23D, E) and 11–12 pairs of middle paleae which are geniculate and with a broad based triangular shaped central tooth (Fig. 23F), one pair of lateral lobes on chaetiger 2 and branchiae absent from



37

posterior abdominal segments. This species can easily be separated from the other species of Sabellaria recorded from Australia by the structure of the outer paleae. Sabellaria kooraltha n. sp., has a smooth midline tooth or spike although the thecae are very conspicuous on the outer paleae (Fig. 23D, E) whereas Sabellaria lungalla n. sp., (this paper) has a finely serrated distal plume on the outer paleae (Figs 24G, 25C).

FIGURE 21. Line drawings of Sabellaria kooraltha n. sp., (Holotype AM W27333, A & B; NTM W4798, C). A. Dorso-lateral view. B. Anterior end, dorsal view. C. Lateral view of entire animal. Scales A, B = 3 mm, C = 2 mm.


HUTCHINGS ET AL.

FIGURE 22. Photos of Sabellaria kooraltha n. sp. A. Whole specimen, ventral view. B. Anterior end, ventral view. C. Detail of the mouth and associated structures. D. Same, with opened opercular lobes. E. Anterior end, lateral view. F. Lateral lobes on segment 2. G. Outer opercular paleae. H. Mid-row of inner opercular paleae. I: Nuchal spines. J. Parathoracic segments. K. Posterior abdominal segments and cauda. Abbreviations:b2, branchia on segment 2; bo, buccal organ; ca, cauda;chn2; chaetae neuropodium segment 2; cn1 cirrus neuropodium segment 1; ll2, lateral lamellae segment 2; mp, inner paleae, mid row; ne parathoracic neuropodia; no, parathoracic notopodia; op, outer paleae; tf, tentacular filaments; pa, palps. A, B, C, E, F, G, H, Paratype AM W27333, D, I, Holotype AM W27195.



39

FIGURE 23. SEM of Sabellaria kooraltha n. sp. (AM W37820.001). A. Anterior end lateral view. B. Anterior end, ventral view. C. Looking down onto opercular crown. D. Outer paleae. E. Close up of median tapering denticle. F. Middle paleae. G. Nuchal spines. H. lateral view of parathoracic segments. I. Lanceolate and capillary notochaetae of parathorax. J. Anterior abdominal neurochaetae. K. Anterior abdominal uncini. L. Cauda. Scales A, B, C, H = 200 µm, D, F, G, L = 100 µm, E, J = 10 µm, I = 20 µm; K = 2 µm.


HUTCHINGS ET AL.



41


HUTCHINGS ET AL.



43

With regard to other Indo-Pacific species which have one kind of middle paleae, both S. gilchristi (McIntosh, 1925) described from South Africa, and S. javanica Augener, 1934, described from Indonesia, have outer paleae with midline plume whereas S. kooraltha n. sp., has a smooth distal spike. The value of SEM is highlighted for the characterisation of this species although the strongly ornamented midline plume of the outer paleae can be seen under high magnification using a compound microscope. Details of the other described species of Sabellaria are given in Table 4. Distribution. North-eastern Australia, from Albany Passage, just south of Cape York in North Queensland and Darwin Harbour, Northern Territory. Habitat. No information on habitat is available for the type locality, but co-ordinates suggest a depth of 10 m and collected by dredging. Non type material collected from muddy substrates from low water mark to 15 m. Etymology. The specific name kooraltha is an Aboriginal word for spotted (Endacott 1973) and refers to the pigmented nature of the dorsum.

Sabellaria lungalla n. sp. Figs 6B, 24, 25, Table 4 Material examined. HOLOTYPE: NTM W16893, Northern Territory: Bing Bong, McAthur River, 15°23' S 136°30.35'E, 17.iii.1993, complete except for cauda, 3 mm in length, 1 mm in width, 12 chaetigers. PARATYPES: 2, NTM W16896), Drimmie Arm, Melville Bay, Gove, 12°14'S 136°42'E, 4 mm in length, 1.5 mm in width for 14 chaetigers, 3.45 mm in length 1.00 mm in width for 14 chaetigers; 2, NTM W10043, Darwin Harbour, D92A, 12°29.52' S 130°50.18' E, 7 m, 14.vii.1993, 7 m, 3 mm in length, 1 mm in width, 18 chaetigers, 5 mm in length, 1 mm in width, 16 segments. Additional material examined. Northern Territory: Bing Bong, McArthur River, Gulf of Carpentaria, 15°36.72'S 136°23.45'E, 18.iii.1993, 1, AM W37780 (mounted for SEM); Darwin Harbour, 12°29'S 130°50.18E, 14.vi.1993, 1, NTM W10043; 12°32.72'S 130°51.32'E, 24.iii.1994, 1, NTM W24725; 12°32.9'S 130°50.02'E, 15.vi.1993, 1, NTM W10042; 1, NTM W16889 . Majority of this material is posteriorly incomplete. Western Australia: Mermaid Sound, Dampier Archipelago, 20°52'S 116°44'E, WAM V3816, 1981. Description. Holotype with short compact body, colourless except for scattered pigment around base of opercular lobes (Fig. 24A). Operculum short, similar in width to length, with lobes completely separated along its length. Paleae arranged in three semicircular concentric rows (Fig. 25A, B, D). Outer row with 22 pairs of geniculate golden paleae with flattened blades, smooth lateral margins and distal tip with midline plume and 5–7 smaller denticles on each side (Figs 24F, 25A–C). Middle row with 10–11 pairs of paleae, short, geniculate, with broad and concave blades directed outwards and tapering tips and inner row with 11 pairs of similar in shape paleae, slightly smaller arranged in between the middle paleae directed inwards, broadly flattened, geniculate, yellow-brown, smooth, pointed tips (Fig. 25A, D). Outer paleae 5–6 times longer than inner and middle rows, middle paleae not covering the inner opercular paleae (Fig. 25A). Sixteen pairs of short opercular papillae (Figs 24C, D, 25B) peripheral to outer paleae. Three pairs of nuchal spines present, slightly curved inwards with blunt tips (Figs 24F, 25E). Tentacular filaments compound (branching) arranged in eight rows (Fig. 24B,C). Palps slightly longer than length of operculum. Median organ present at dorsal junction of lobes of operculum, with eye spots on lateral margins. Segment 1 (chaetiger 1) with one pair of neuropodial cirri with small tuft of fine capillary chaetae on either side of U-shaped buccal organ (Fig. 24B). Segment 2 (chaetiger 2) with one pair of elongated lateral lobes, connecting branchiae to neuropodia (Fig. 24C, D). Neuropodia of segments 1 and 2 not vertically aligned, those of segment 2 situated more laterally, neurochaetae similar in structure on both segments. Five to six pairs of branchiae present (difficult to count as holotype slightly damaged). Branchiae narrow based, strongly ciliated, long and tapering to fine tip. Segment 3–5 (parathoracic) with two types of notochaetae arranged transversely (Fig. 25H); six lanceolate chaetae, tapering to an elongate frayed tip (Fig. 25H) and fine capillaries with frayed margins inserted between lanceolate ones (Fig. 25H). Segments 3–5 with two types of neurochaetae arranged in compact lateral fascicle, with about six to eight lanceolate chaetae plus fine capillaries. Parathoracic notopodia more robust that neuropodia (Fig. 24A). Abdominal region with 12 chaetigers. Notopodia as transverse tori, with long handled uncini, numbers per torus decreasing posteriorly, each uncinus with two vertical rows, each row with six to eight major teeth (Fig. 25J), similar throughout abdomen. Neuropodia with ventral bundle of fine


HUTCHINGS ET AL.

notochaetal capillaries with extended fine tips, shafts composed of stacks of thecae with margins extending to fine filamentous tip (Fig. 25I), similar throughout abdomen. Cauda smooth and about half of length of abdomen on paratype.

FIGURE 24. Photos of Sabellaria lungulla n. sp. holotype (NTM W16896) A. Whole animal, lateral view view. B. Anterior end, ventral view. C. Detail of the mouth and associated structures. D. Anterior end, lateral view showing details of lateral lobes. E. Same, showing details of paleae and tentacular filaments. F. Detail of nuchal spines, paleae and oral papillae. Abbreviations: bo, buccal organ; ca, cauda; chn2, chaetae neuropodium segment 2; ll2, lateral lamellae segment 2; ns, nuchal spines (hooks); op, outer paleae; opa, oral papillae; tft, tentacular filaments.

Variation. Material examined varies from 3–5 mm in length, without cauda, 1–2 in width with 13–18 chaetigers, 22–23 pairs of outer paleae, 10–11 pairs of inner paleae, 11–12 pairs of middle paleae and 16–23 pairs of opercular papillae. Remarks. Sabellaria lungalla n. sp., is characterised by the distal margin of the outer paleae being serrated with a midline plume, one kind of middle paleae which are geniculate with smooth pointed tips and inner paleae which are asymmetrical with short blunt tips, three pairs of nuchal spines and one pair of lateral lobes on segment 2, which no other species share (Table 4). Sabellaria lungalla n. sp., can be separated from S. kooraltha n. sp., by the shape of the distal margin of the outer paleae: S. lungalla n. sp., has an elongate finely serrated distal plume (Fig. 25C) whereas S. kooraltha n. sp., HUTCHINGS, CAPA AND PEART, AUSTRALIAN SABELLARIIDAE


45

has a large central distal tooth which is smooth with two lateral teeth on either side (Fig. 23D, E). The other Australian species S. pyramis n. sp., differs from S. lungalla n. sp., by the number and shape of the middle paleae (Fig. 26E, F). Sabellaria lungalla n. sp., belongs to a group of Sabellaria with only one kind of middle paleae (Table 4), and of those occurring in the Indo-Pacific they can be separated from S. lungalla n. sp., on the basis on the dentition of the single kind of mid-paleae. Sabellaria gilchristi (McIntosh, 1925) has only two pairs of teeth on the distal margin of the outer paleae Sabellaria javanica Augener, 1934 described from Indonesia, has no teeth on the lateral margins and S. tottoriensis Nishi et al., 2004, described from Japan, has only three teeth whereas S. lungalla n. sp., has 7–8 pairs of teeth. Distribution. Gulf of Carpentaria, Northern Australia, and Dampier Archipelago, Western Australia. Habitat. Intertidal in sheltered bays from intertidal to 15 m. Etymology. The specific name lungalla refers to an Aboriginal word for a lagoon in the Northern Territory and this species is found in inshore shallow bays.

FIGURE 25. SEM of Sabellaria lungalla n. sp. (AM W37780). A. Anterior end, ventral view. B. Anterior end, dorsal view. C. Outer opercular paleae. D. Middle row of inner opercular paleae. E. Nuchal spines. F. Anterior lateral view showing tentacular filaments. G. Lanceolate and capillary neurochaetae of parathoracic segments. H. Lanceolate and capillary notochaetae of parathoracic segments. I. Anterior abdominal neurochaetae. J. Anterior abdominal uncini. Scales A, B = 30 µm, C, D, E = 10 µm, F = 3 µm, G = 1 µm.


HUTCHINGS ET AL.

Sabellaria pyramis n. sp. Fig. 6A, 26, Table 4 Material examined. Holotype: NTM W10045, 6 mm long, 1 mm wide, posteriorly incomplete with 17 chaetigers, paratypes: 1 NTM W023787, 10 mm long, 1 mm wide, complete with 28 chaetigers, plus cauda, operculum incomplete; 1, NTM W10044, 8 mm long, 1 mm wide, incomplete. All type material from Northern Territory: Darwin Harbour, Holotype and NTM W023787 from St. D24A, 12°25.05'S 130°48.58'E, 8 m, 7.i.1993 and NTM W10044, from St D98A, 12°31.9'S 130°50.43'E, 18 m, coll 14.vii.1993.

FIGURE 26. Photos of Sabellaria pyramis n. sp., A. Anterior end, ventral view. B. Same, lateral view. C. Same, dorsal view. D. Outer and middle row of inner opercular paleae. E. Middle row of inner opercular paleae. F. Same, different angle. G. Anterior segments, ventral view. H. Same, dorsal view. Abbreviations: b2, branchia segment 2; bo, buccal organ; cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodia segment 1; ip, inner paleae; ll2, lateral lamellae segment 2; mp, inner paleae, mid row; op, outer paleae; tf, tentacular filaments. Holotype: NTM W10045.

Description. Holotype pale yellow with scattered brown pigmentation on dorsum of last parathoracic segments and at base of tentacular filaments. Operculum longer than wide, with completely separated lobes. Paleae arranged in three semicircular concentric rows; outer row with 18 pairs, clear and golden paleae with broad



47

flattened smooth blade, distally serrated with fine extended midline plume (majority broken). Middle row with one kind of paleae, five pairs, stout, robust, shaft forming a four sided pyramid with blunt tips (Fig. 26D). Inner row with nine, geniculate with symmetrical blunt tips directed inwards paleae. Middle paleae longest, outer paleae cover bases of middle, inner shortest. Opercular papillae 12 pairs, shorter than wide, cylindrical (Fig. 26B). Nuchal spines not observed. Tentacular filaments compound, long, arranged in five transverse rows (Fig. 26A). Median organ and median ridge present, with eye spots on both margins. Segment 1 (chaetiger 1) with one pair of elongate neuropodial cirri, with long capillaries (Fig. 26G). Segment 2 (chaetiger 2) with one pair of lateral lobes connecting branchiae to neuropodia (Fig. 26H). Neurochaetae similar on both segments. Six pairs of branchiae, elongate, strongly ridged, narrow based, tapering to fine tip, not meeting mid dorsally. Segments 3–5 (parathoracic) with two types of notochaetae arranged transversely, eight lanceolate chaetae with fine capillaries imbetween. Segments 3–5 with two types of neurochaetae, 3–4 lanceolate and capillaries imbetween. Abdomen of 23 segments with short neuropodia with capillary chaetae and expanded rectangular notopodia with terminal transverse torus with long handled uncini with 6–8 teeth arranged vertically, similar throughout abdomen. Cauda smooth and equal in length to last ten segments. Variations. The two paratypes vary in the number of pairs of middle paleae (4–5), inner paleae (9–10) and branchiae (5–6) present, and both are incomplete posteriorly and the opercular lobes are slightly damaged. Remarks. Sabellaria pyramis n. sp., differs from other species of the genus in the particular pyramidal shape of the middle paleae, unique in the genus. All other described species of Sabellaria have geniculate middle paleae. Even though the genus Sabellaria was not recovered as mophyletic in Capa et al. (2012) or the present study (Fig. 28C) we have positioned this species within this genus since it shares the combination of features of the rest of the species (operculum completely divided in two symmetrical lobes, inner paleae arranged in two rows, with mid and inner paleae, and three parathoracic segments) together with the shape of outer paleae provided with a distal plume and denticles on both sides. Distribution. Darwin Harbour, Northern Territory, known only from type locality. Habitat. Muddy sediments in depths of 8–18 m. Etymology. The specific name of pyramis refers to the pyramidal shape of the middle paleae.

Tetreres Caullery, 1913 Tetreres Kirtley, 1994: 188. Capa et al., 2012: 269. Type species. Hermella varians Treadwell, 1901, designation by Kirtley, 1994. Type locality: Mayagüez Harbour, Puerto Rico.

Diagnosis. Characterised by the arrangement of the inner paleae in a short ventral line on each inner margin of the opercular lobes. Nuchal hooks are large and with broadened shafts, unique among sabellariids. Remarks. This genus currently contains ten species plus the one described in this paper, with most known from deep water (188–4825 m). Traditional diagnostic characters of these species, based almost entirely on paleal and nuchal hook characters (as in Kirtley, 1994) are listed in Table 5.

Tetreres terribilis n. sp. Figs 6B, 27, Table 5 Material examined. Holotype: NTM W4994*, FRV Soela, cruise 06/85 st. 5: 17.xi.1985, S of Saumarez Reef, Marian Plateau, 22˚40.0'S, 154˚05.5'E to 22˚42.5'S, 154˚05.9'E, 416–419 m, collected with a lobster trawl. Specimen posteriorly incomplete with 15 chaetigers, 35 mm in length, 6 mm in width. Description. Holotype robust, laterally flattened body, preserved animal pale yellow colourless except for dark brown nuchal hooks. Operculum with two partially fused lobes, as long as wide with deep indentation on ventral margin, laterally compressed (Fig. 27A). Outer paleae, 40 pairs, arranged in semicircles, stout, golden, flattened, blades with three transverse thecal bands and asymmetrical pointed tips (Fig. 27A). Inner row of paleae with three pairs of paleae arranged as a short ventral line on each inner margin of the opercular lobes, about one quarter length of outer paleae. One pair of flat and strongly curved nuchal spines without limbation (Fig. 25C). Opercular


HUTCHINGS ET AL.

papillae, 14 pairs, triangular, almost as long as outer paleae (Fig. 27A, B). Dorsal papilla, digitiform, located between nuchal spines (Fig. 27C). Palps, thick, rounded, with crenulated margins about one third longer than operculum. Median and median ridge present. Eye spots not observed, as difficult to separate lobes. Tentacular filaments, simple, nine pairs. Segment 1 (chaetiger 1) with neuropodial cirri on either side of buccal organ, elongate and tapering and a fascicle of capillaries. Segment 2 (chaetiger 2) with well fascicle of neuropodial capillaries, three elongate narrow lateral lobes present. Branchiae from segment 2, extending to chaetiger 13, last two chaetigers lacking branchiae, long thin flattened, meeting mid dorsally. Segments 3–6 (parathoracic) with two types of notochaetae inserted transversely, eight stout lanceolate with frayed tips and much shorter finer capillaries inserted in between. First parathoracic notopodia very small about one quarter size of subsequent ones which increase in size posteriorly. Notochaetae consist of two rows of five large lanceolate chaetae and short fine capillaries respectively. Segments 3–6 (parathoracic segments) with only one kind of neurochaetae, six capillaries. Abdomen incomplete. Abdominal neuropodia with compact fascicle of long capillaries arranged in two tiers with expanded fine tips, thecae expanded and margins appear denticulate. Notopodia erect expanded tori with transverse row of uncini with two rows of teeth, each row with 7–8 teeth. Cauda missing in holotype.

FIGURE 27. Photos of Tetreres terribilis n. sp. A. Anterior end, lateral view. B. Detail of the opercular papillae and outer opercular paleae. C. Nuchal spines. D. Cirrus and chaetae of neuropodia of segment 1. E. Lanceolate and capillary notochaetae of parathorax. F. Capillary neurochaetae of parathorax. Abbreviations: cn1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1; dpa, dorsal papilla; ns, nuchal spines (hooks); op, outer paleae; opa, oral papillae; tf, tentacular filaments; pa, palps. Holotype: NTM W4994.



49


HUTCHINGS ET AL.

Remarks. Tetreres terribilis n. sp., is characterised by the presence of outer paleae with flattened blades symmetrical tips and transverse thecal bands a feature only shared with T. perryi (Kirtley, 1994), however it should be noted that the latter species is poorly described with only an illustration of the outer paleae and a comment that the nuchal looks are large with sharply recurvedd tips (Table 5). These two species are distinguished by the shape of the tip of these outer paleae with a pointed tip in the new species and a long spike in T. perryi, and the presence of an internal fusiform outline in T. terribilis n. sp., absent in T. perryi (Table 5). The only species described from nearby localities to Marian Plateau is T. robustus Lechapt & Kirtley, 1998 at depths of 440–450 m off New Caledonia. This species was described as having a larger number of outer paleae (70–80 pairs) but with a smooth surface, lacking thecal bands but they can be separated by the number of lateral lobes on segment 2, being three in T. terribilis n. sp. and four in T. robustus. Distribution. Known only from type locality, Marion Plateau in depths of 416–419 m. Habitat. Substrate unknown, appears to be a solitary species. Etymology. The name of this species terribilis refers to the large size of the species and which is much larger than any other material examined and if complete would represent a very scary animal.

Phylogenetic analysis Parsimony analysis of the morphological dataset resulted in 460 most parsimonious trees (L: 81, CI: 0.53, RI: 85) where relationships are not assessed among Sabellariidae and only the monophyly of Bathysabellaria (JK: 67), Phalacrostemma (JK: 83) and Teteres (JK: 71) are indicated (Fig. 28A). Reruning analyses and varying the concavity values (k) between 3 and 20 resulted in more resolved topologies as a result of the increasing weight of characters with less homoplasy (Fig. 28B, C). Monophyly is assessed for Phalacrostemma, Gessaia, Bathysabellaria, Mariansabellaria and Tetreres regardless of k values (Fig. 28B, C) and Idanthyrsus was also recovered monophyletic with lower concavity values (k: 3–6, Fig. 28B). The new species described herein have been considered as members of those genera recovered as monophyletic as in the case of Phalacrostemma maloga n. sp., and Tetreres terribilis n. sp., and for those retrieved paraphyletic we have followed the traditional taxonomic approach of considering them as belonging to those genera when sharing with type species diagnostic features and a unique combination of characters with other members considered in that group.

Discussion Kirtley (1994) conducted an extensive world revision of the family examing types that allowed him to describe many new species and erect four new genera. New species have been described since, but it has only been recently that the first phylogenetic hypothesis of the group has been proposed (Capa et al. 2012). These results showed that the two subfamilies erected by Kirtley (1994) Also, after their analyses, the monophyly of most genera was assessed but others were recovered as paraphyletic. In the present study some of the new species have been recovered in genera shown to be monophyletic but this is not the case for species belonging to genera such as Sabellaria, Lygdamis and Idanthyrsus. Nevertherless, I. nesos n. sp., I. willora n. sp., L. wambiri n. sp., S. kooraltha n. sp., S. lungalla n. sp., and S. pyramis n. sp., have been considered as part of these genera since they share most diagnostic features with other members of these groups. A revision of these groups and further analyses should be made to asses relationships within those groups. Kirtley (1994) also provided a list of characters which he regarded as useful in defining genera. These included the degree of fusion of the opercular lobes, the arrangement and orientation of the paleae and their ornamentation, presence or absence of nuchal spines or hooks and their shape and presence or absence of external marginal limbations, type of tentacular filaments simple or compound, presence of median cirrus, and the type of oral tentacles. Subsequent studies, in which additional new species of Sabellariidae have been described, have tended to follow Kirtley (1994) although the quality of illustrations has improved with some providing SEM photos.



51

FIGURE 28. Trees resulting from parsimony analyses of Sabellariidae rooted with Spionidae: A. Strict consensus of 460 trees after analyses based on 31 morphological features with Jackknife support values; B. Strict consensus of 137 most parsimonious trees after maximum parsimony analysis implementing implied weighting (constant of concavity k = 3–6); C. Strict consensus of 134 most parsimonious trees implementing implied weighting (constant of concavity k = 10-20).


HUTCHINGS ET AL.

TABLE 6. Character matrix of morphological features (based in Capa et al. 2012). ‘‘?’’ stands for unknown and ‘‘-’’ for inapplicable states. 1-10

11-20

21-30

31

Spionidae

-------0--

----------

---0-1-

-

Bathysabellaria neocaledoniens

0000001101

10000---00

110--01001

1

Bathysabellaria spinifera

0101001101

10000---00

110--01201

1

Gesaia elegans

2000001100

20000---10

2110-10111

1

Gesaia fossae

2000001100

20000---10

2110-20011

1

Gunnarea gaimardii

1101100001

00100---01

10---01110

0

Idanthyrsus macropaleus

2010101100

01100---20

2110-00210

?

Idanthyrsus australiensis

2010101100

01100---20

2110-00210

2

Idanthyrsus nesos n. sp.

2011101?00

01100---20

2111100110

2

Idanthyrsus willora n. sp.

201?101100

01100---20

2110-00310

2

Lygdamis indicus

2010101100

00000---20

2110-01211

0

Lygdamis augeneri

2010101100

00000---20

2110-00211

0

Lygdamis giardi

2010101100

00000---20

2110-00211

0

Lygdamis wambiri n. sp.

2011101100

00000---20

2110-00211

0

Mariansabellaria norvegicus

2000002000

20000---10

210--00101

1

Mariansabellaria harrisae

2000002000

20000---10

210--00001

1

Neosabellaria cementum

0101100001

1011111101

10---01110

0

Neosabellaria vitiensis

0101100001

1011111101

10---01110

0

Neosabellaria uschakovi

0101100001

1011111101

10---0?110

0

Paraidanthyrsus quadricornis

2101101001

01100---01

0111001110

0

Phalacrostemma cidarophillum

2100011110

20000---10

2111121111

0

Phalacrostemma profunda

2100011110

20000---10

2111121111

0

Phalacrostemma maloga n. sp.

2100011110

20000---10

2111101111

0

Phragmatopoma caudata

0001101001

0011101201

30---01110

0

Phragmatopoma californica

0001101001

0011101201

30---01110

0

Sabellaria alveolata

2101101101

0010111101

10---01010

0

Sabellaria ishikawai

2101101101

0011111101

10---01010

0

Sabellaria isumiensis

2101101101

0011111101

110--01010

0

Sabellaria kooraltha n. sp.

2101101101

0010111101

110--01010

0

Sabellaria lungalla n. sp.

2101101101

0010111101

110--01010

0

Sabellaria pyramis n. sp.

2000101?01

0011111301

100--01010

0

Tetreres varians

1000002000

00000---30

2110-01301

1

Tetreres superbus

1000002000

00000---30

2110-01301

1

Tetreres robustus

1000002000

00000---30

2110-01401

1

Tetreres terribilis n. sp.

1000002000

00000---30

2110-01201

1



53

While attempting to assign the large collection of Australian sabellariids to species, we examined a selection of material under the SEM and we found that this provides very detailed images of the paleae, chaetae and uncini which facilitates the comparison of these structures between taxa, although we do suggest that they may exhibit some wear and tear. Using SEM allowed us to record, for the first time in Sabellariidae, the presence of three rows of teeth in the abdominal uncini of Phalacrostemma and the two rows of lanceolate chaetae in the parathoracic segments of Idanythrysus. The differences between species related to the shape of paleae, spines, chaetae, papillae, etc, are difficult to quantify and somewhat subjective. So we investigated the value of other characters which are easier to quantify and compare. Some features found to be useful in distinguishing between species and that do not vary with the size of the individual are the number of lateral lobes on segment 2, the number of parathoracic notoand neurochaetae, the distribution of the branchiae in posterior segments, and the presence or absence of the median ridge, median organ and dorsal papillae. We also describe a type of middle paleae which have not been described for any other species of Sabellaria in S. pyramis n. sp. The single individual of T. terribilis n. sp., collected from the Marion Plateau also exhibits a character which has not previously been recorded for this genus, the presence of a dorsal papillae. So these two new species highlight that additional morphological characters are still being documented for the family. Many of these characters were also used in the phylogenetic analyses carried out by Capa et al. (2012). So while paleae and nuchal hooks/spines are useful, they are not the only ones and the additional characters suggested above should be included in any future descriptions of species of sabellariids. As shown in the Tables 1–5, many of the species are poorly known and need to be redescribed from either the type or in the case of species for which types are no longer extant from material collected from the type locality. Prior to this study, only four species in two genera were known from Australia, a number that has increased to 11 species in five genera. It is difficult to compare this diversity with other areas in the Indo-Pacific as no other comprehensive revisions have been undertaken yet. Individual species have been described from specific sites in Indonesia, Sri Lanka, Solomons, French Polynesia, Japan, deep water off New Caledonia and Fiji (for details see Tables 1–5 and Brock et al. 2007; Lechapt and Kirtley 1996; Nishi et al. 2004, 2010) however none of these studies reviewed the diversity of the family in those localities We predict that the diversity of Australian sabellariids will increase with additional collecting especially in deeper waters. Given that very limited sampling has been undertaken offshore, these samples yielded two new species in genera not previously known from Australia (Phalacrostemma and Tetreres). Of the 11 species, only one, I. australiensis appears to be widespread around the country in warm and temperate environments, others have restricted distributions and some are known only from a single locality. For example, S. pyramis n. sp., was only collected from two stations in Darwin Harbour, an area which has been extensively sampled. Other sites in the harbour with similar sediment characteristics lacked this species, so other factors must control the distribution of this species. An interesting feature which seems to characterise the Australian sabellariid fauna is that the species do not appear to form massive reefs which are characteristic of the family in other parts of the world (Kirtley 1994 and refs therein) and while some form small colonies others appear to occur as isolated individuals. While sabellariids have been recorded from most areas of Australia as shown in Fig. 6, no records currently exist from the southern coast from Adelaide west to Esperance although sampling has occurred in this region and exposed coasts are available. Similarly the Queensland coast has no records but this may be related to the fact that the coast consists of protected waters often lined with mangroves. The offshore islands and reefs within the Great Barrier Reef lagoon also seem to lack this family. Localised intensive collecting has been undertaken in this region by us and so we are confident that such habitats are not suitable for the family. Much remains to be learnt as to the habitat requirements of this family but presumably all species require some hard substrate on which to settle, although this may only need to be a shell, such as Idanthyrsus willora n. sp., which allows them to occur on muddy substrates.

Acknowledgements We should like to thank Anna Murray for the line drawings, Mick Ashcroft for preparing the maps, Sue Lindsey for the SEM photos and Kate Attwood for preparing the material for SEM. The following collection managers, loaned us their collections of sabellariids, including Chris Rowley, Museum of Victoria, Mal Bryant, Queensland


HUTCHINGS ET AL.

Museum, Gavin Dally, Museum and Art Gallery of Northern Territory, Liz Turner, Tasmanian Museum, Andrew Hosie, Western Australian Museum and Thierry Laperousaz, South Australian Museum. ABRS provided support to PH which provided a salary for MC and RP.

References Allen, E.J. (1904) Pallasia murata n. sp., a new British Sabellarian. Journal of Marine Biological Association of the United Kingdom, 7, 299–304. Annenkova, N. (1925) Beiträge zur Kenntnis die Polychaete-Fauna Russlands. Nauka for Akademiya Nauk SSR: Leningrad, 1925, 125–126. Augener, H. (1906) Reports on the Results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico and the Caribbean Sea, and on the east coast of the United States, 1877–1880, by the U.S.S. Coast Survey Steamer Blake. Westindische Polychaeten. Bulletin of Museum of Comparative Zoology, Harvard, 43, 91–196. Augener, H. (1914) Polychaeta II. Sedentaria. pp. 1–172 in Michaelsen, W. & Hartmeyer, R. (eds). Die Fauna SüdwestAustraliens. Jena: Fischer, 5(5), 1–170. Augener, H. (1918) Polychaeta. Beiträge zur Kenntnis des Meeresfauna West-Afrikas. Herausgegeben von W. Michaelsen, Hamburg, 2(2), 67–625. Augener, H. (1922) Results of Dr. E. Mjoberg’s Swedish Scientific expeditions to Australia, 1910–13. Polychaeten. Kunglinga Svenska Vetenskapsakademiens Handligar, Stockholm, 63(6), 1–49. Augener, H. (1927) Polychaeten von Sűdost – und Sűd-Australien. Papers from Dr Th. Mortensen’s Pacific Expedition 1914–16. No 38. Videnskabelige Meddelelser fra Dansk naturhistorisk Førenig i København, 83, 71–275. Augener, H. (1934) Polychaeten aus den zoologischen Museen von Leiden und Amsterdam, part IV. Zoologisische Mededelingen Rijksmuseum van natuurlijke histoire Leyden, 17, 67–160. Bailey-Brock, J.H., Kirtley, D.W., Nishi, E. & Pohler, S.M.J. (2007) Neosabellaria vitiensis n. sp. (Annelida: Polychaeta: Sabellariidae), from Shallow Water of Suva Harbor, Fiji Pacific. Pacific Science, 61(3), 399–406. Baird, W. (1863) Descriptions of several new species of worms belonging to the annelida Errantia or Tubicola of MilneEdwards. Proceedings of the Zoological Society of London, 24, 106–110. Bhaud, M.R. & Fernandez-Alamo, M.A. (2001) First description of the larvae of Idanthyrsus (Sabellariidae, Polychaeta) from the Gulf of California and Bahia de Banderas, Mexico. Bulletin of Marine Science, 68, 221–232. Caline, B., Gruet, Y. & Legendre, C. (1992) The sabellariid reefs in the Bay of Mont Saint-Michel, France. Ecology, geomorphology, sedimentology, and geologic implications. Contributions to Marine Science (Florida), 1(1), 1–156. Capa, M,. Hutchings, P.A. & Peart, R. (2012) Systematic revision of Sabellariidae Johnston, 1865 (Polychaeta) and relationships with other polychaetes. Zoological Journal of the Linnean Society, London, 164, 245–284. Carrasco, F.D. & Bustos, R. (1981) Dos especies de poliquetos de la familia Sabellariidae Sabellaria minuta sp. n.e Idanthrysus armatus Kinberg epizoos de Crustaceas Decapodos. Boletin de la Sociedad de Biologia Concepcion, Chile, 51, 167–173. Caullery, M. (1913) Sur le genre Pallasia Qfg. et la region prostomiale des Sabellariens. Bulletin de la Société Zoologique de France, 38, 198–203. Caullery, M. (1944) Polychètes Sédentaire de l'Expédition du Siboga: Ariciidae, Spionidae, Chaetopteridae, Chlorhaemidae, Opheliidae, Oweniidae, Sabellariidae, Sternaspidae, Amphictenidae, Ampharetidae, Terebellidae. Siboga-Expeditie Uitkomsten op Zoologisch, Bonatisch, Oceanographisch en Geologisch gebied verzameld in Nederlandsch Oost–Indië, 1899–1900 XXIV 2, bis,1–204. Chamberlin, R.V. (1919) New Polychaetous annelids from Laguna Beach, California. Journal of Entomology and Zoology, Pomona College, 11, 1–23. Collin, A. (1902) Verzeichniss der von Prof. R. Semon bei Amboina und Thursday Island gesammelten Polychäten.Zoologische Forschungsreisen in australien und dem malayischen Archipel. ausgeführt in den Jahren 1891–1893 von Prof. Dr. Richard Semon ; mit Unterstützung des Herrn Dr. Paul von Ritter. G. Fischer (Jena), 5 (2), 739–742. Dakin, W.J., Bennett, I. & Pope, E.C. (1953) Australian Seashores. Sydney: Angus & Robertson 1–372 pp. Dales, R.P. (1952) The development and structure of the anterior region of the Sabellariidae with special reference to Phragmatopoma californica. Quarterly Journal of the Microscopical Society, London, 93, 435–452. Dallwitz, M.J. (1980) A general system for coding taxonomic descriptions. Taxon, 29, 41–46. Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1993) ‘User’s guide to the DELTA System: a General System for Processing Taxonomic Descriptions.’ 4th ed. (CSIRO Division of Entomology: Canberra). Day, J.H. & Hutchings, P.A. (1979) An Annotated Checklist of Australian and New Zealand Polychaeta and Myzostomidae. Records of the Australian Museum, 32, 80–161. De Silva, P.H.D.H. (1961) Contribution to the knowledge of the polychaete fauna of Ceylon. Spolia Zeylandica, National



55

Museum of Ceylon, 29(2), 164–194. Dubois, S., Retiere, C. & Olivier, F. (2002) Biodiversity associated with Sabellaria alveolata (Polychaeta: Sabellariidae reefs: Effects of human disturbances. Journal of the Marine Biological Association of the United Kingdom, 82(5), 817–826. Eckelbarger, K.J. (1976) Larval development and population aspects of the reef-building polychaete Phragmatopoma lapidosa from the east coast of Florida. Bulletin of Marine Science, 26, 117–132. Ehlers, E. (1908) Wissenschaftliche Ergenbnisse der Deutschen Tiefsee_Expedition auf dem Dampfer Valdivia 1898–99. Im Auftrage des Reichsamptes des Innern, herausgegben von Carl Chun, Professor der Zoology in Leipzig, Leiter der Expedition. Die Bodenassigen Anneliden aus den Sammlungen der Deutsche Tiefsee-Expedition. Jena, Gustav Fischer, 16(1), 1–168. Ehlers, E. (1920) Polychaeten von Java und Amboina. Ein Beitrag zur Kentniss der malaiischen Strandfauna. Abhandlungen der Konigliche Gesellscahft der Wissenschaften zu Gottingen, Mathematisch-Physicalische Klasse, new series, 10(7), 1–73. Endacott, S.J. (1973) Australian Aboriginal Words and Place Names. 10th ed. (Acacia Press: Victoria.) Ehlers, E. (1904) Neuseeländische Anneliden. Abhandlungen der Königleichel Gesellschaft der Wissenschaften zu Göttingen. Mathematisch –Physikalische Klasse Neue Folge, 3(1), 1–79. Fauchald, K. (1972) Benthic polychaetous annelids from deep water off western Mexico and adjacent areas in the eastern Pacific Ocean. Allan Hancock Foundation Monographs in Marine Biology, Los Angeles, University of Southern California Press, 7, 1–575. Fauchald, K. (1977) The Polychaete worms. Definitions and keys to the orders, families and genera. Natural History Museum of Los Angeles County, Science Series, 28, 1–188. Fauvel, P. (1914) Annélides polychètes de San Thome (Golfe de Guinée) recuillis par M. Ch. Gravier. Archives de Zoologie Expérimentale et Générale, 54, 105–155. Fauvel, P. (1917) Annélides polychètes de l'Australie meridionale. Archives de Zoologie Expérimentale et Générale, 56, 159–278. Fauvel, P. (1928) Annélides polychètes nouvelle de l’Indie, part 2. Bulletin du Muséum National d’Histoire Naturelle, Paris, 34(2), 159–165. Fitzhugh, K. (1989) A systematic revision of the sabellidae-caobangiidae-sabellongidae complex (Annelida Polychaeta). Bulletin of the American Museum of Natural History, 192, 1–104. Fournier, J., Etienne, S., & Le Cam, J.B. (2010) Inter- and intraspecific variability in the chemical composition of the mineral phase of cements from several tube-building polychaetes. Geobios (Villeurbanne), 43 (2), 191–200. Gaikwad, U.D. (1990) Occurrence of Sabellaria simplex Day, Colonies and observations on the Building Behavior in Captivity. Current Science, Bangalor, 58(11), 642–643. Gibbs, P.E. (1971) The Polychaete Worms of the Solomons Islands. Bulletin of the British Museum (Natural History), 21(5), 101–211. Goloboff, P.A. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91. Goloboff, P.A. (1995) Parsimony and weighting: a reply to Turner and Zandee. Cladistics, 11, 91–104. Goloboff, P.A, Farris, J.S., & Nixon K. (2008a) TNT, a free program for phylogenetic analysis. Cladistics, 24: 774–786. Goloboff, P.A., Carpenter J.M, Arias J.S, Miranda Esquivel, D.R. (2008b) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 1–16. Gravier, C. (1906) Un Sabellarien vivant sur un Brachiopde (Kingena alcocki) Joubin. Bulletin du Muséum National d’Histoire Naturelle, Paris, 12(7), 540–543. Gravier, C. (1908) Sur un type nouveau d'Annélide polychète. Comptes Rendus (Hebdomadaires) des Séances de l'Academie des Sciences. Série D. Sciences Naturelles, 146, 144–146. Grube, A-E. (1870) Ueber die Goldkrönchen (Sabellarien oder Hermellen). Jahresbericht der Schlesische Gesellschaft für vaterländische Kultur, Breslau, 47, 69–70. Grube, A-E. (1878) Annulata Semperiana. Beitrage zur Kenntnis der Anneliden-fauna der Philippinen nach den von Herrn Prof. Semper mitgebrachten Sammlungen. Memoires de l’Academie imperial des Sciences de St.Petersbourge, serie 7, 25(8), 1–300. Hansen, G.A. (1882) Recherches sur les Annélides recueillies par M. le Professeur Eduard van Beneden pendant son voyage au Brésil et à la Plata. Memoires de L’Academie Royale de Science de Belge, Bruxelles, 44(3), 1–29. Hartman O. (1944) Polychaetous annelids. Part VI. Paraonidae, Magelonidae, Longosomidae, Ctenodrilidae, and Sabellariidae. Allan Hancock Pacific Expeditions, 10(3), 311–389. Hasan, G.A. (1960) Recherches sur les Annélides recueillies par M. le Professeur Eduard van Beneden pendent sone voyage au Brésil et à la Plata. Memoires de l’Academie Royale de Science de Belge, Bruxelles, 44(3), 1–29. Haswell, W.A. (1883) On some new Australian tubicolous annelids. Proceedings of the Linnean Society of New South Wales, 7, 633–638. Imajima, M. (2006) Polychaetous annelids from Sagami Bay and the Sagami Sea, central Japan. Memoirs of the National Science Museum (Tokyo), 40, 317–408.


HUTCHINGS ET AL.

Imajima, M. & Hartman, O. (1964) The polychaetous Annelids of Japan, Part II. Allan Hancock Foundation Occasional Paper, Los Angeles, University of Southern California Press, 26, 1–452. Jeldes, F., Lefevre, S. (1959) Annélides Polychètes non pelagiques. Second Note.Polychètes sèdentaires. In Expedition Océanographique Belge les eaux côtières de L’Atlantique Sud (1948–1949). Institute Royale des Sciences Naturelles Belge, Bruxelles, 4(5), 1–40. Johansson, K.E. (1922) On some new Tubiculous Annelids from Japan, the Bonin Islands and the Antarctic. Arkiv för Zoologi, Stockholm, 15(2), 1–11. Johansson, K.E. (1927) Beitrage zur Kenntnis des Polychaeten-Familien Hermellidae, Sabellidae und Serpulidae. Zoologiska Bidrag från Uppsala, 11, 1–184. Johnston, G. (1865) A catalogue of the British non-parasitical worms in the collection of the British Museum. British Museum, London, 366 pp. Kinberg, J.G.H. (1867) Annulata nova. Öfversigt af Königlich Vetenskapsakademiens förhandlingar, Stockholm, 23(9), 337–357. Kirtley, D.W. (1994) A review and taxonomic revision of the family Sabellariidae Johnston, 1865 (Annelida; Polychaeta). Sabecon Press Science Series, 1, 1–223. Lana, P.C. & Gruet, Y. (1989) Sabellaria wilsoni sp.n. (Polychaeta, Sabellariidae) from the Southeast Coast of Brazil. Zoologica Scripta, 18(2), 239–244. Lamarck J.Bd. (1812) Extrait du cours de zoologie du Muséum d'Histoire Naturelle, sur les animaux sans vertèbres; présentant la distribution et la classification de ces animaux, les caractères des principales divisions, et une simple liste des genres; a l'usage de ceux qui suivent ce cours: Paris, D'Hautel et Gabon., 127 pp. Lechapt, J-P. & Gruet, Y (1993) Bathysabellaria neocaledoniensis, a new genus and species of Sabellariidae (Annelida, Polychaeta) from bathyal zones off New Caledonia (southwest Pacific Ocean). Zoologica Scripta, 22(3), 243–247. Lechapt, J-P. & Kirtley, D.W. (1996) Bathysabellaria spinifera (Polychaeta: Sabellariidae) a new species from deep water off New Caledonia, southwest Pacific Ocean. Proceedings of the Biological Society of Washington, 109(3), 560–574. Lechapt, J-P. & Kirtley, D.W. (1998) New species of bathyal and abyssal Sabellariidae (Annelida: Polychaeta) from near New Caledonia (southwest Pacific Ocean). Proceedings of the Biological Society of Washington, 111(4), 807–822. Leuckart, R. (1849) Zur Kenntnis der Fauna von Island. Archiv für Naturgeschichte, Berlin, 15(1), 149–209. Linnaeus C. Von. (1767) Systema naturae sive regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Laurentii Salvii, Holmiae. Stockholm, 12th edition, 1, 533–1327. Marenzeller E. von, (1895) Phalacrostemme cidariophilum, eine neue Gattung und Art der Hermelliden. Anzeiger, Akademie der Wissenschaften, Wien, 32, 191–192. McCarthy, D.A.; Kramer, P., Price, J. R. (2008) The ecological importance of a recently discovered intertidal sabellariid reef in St. Croix, U.S. Virgin Islands. Caribbean Journal of Science, 44 (2), 223–227. McIntosh, W.C. (1885) Report on the Annelida Polychaeta collected by H.M.S. 'Challenger' during the years 1873–76. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 12, 1–554. McIntosh, W.C. (1922) A monograph of the British marine annelids. Polychaeta: Hermellidae to Sabellidae. London, Ray Society, 4, 1–250. McIntosh, W.C. (1925) A Second Contribution to the Marine Polychaetes of Africa. Reports of the Union of South Africa Marine Biological Survey, Cape Town, 4, 1–93. Monro, C.C.A. (1933) The Polychaeta Sedentaria Collected by Dr C. Crossland at Colon in the Panama Region, and in the Galapagos Islands during the S.Y.St George. Proceedings of the Zoological Society of London, 2(4), 1039–1092. Nishi, E., Bailey-Brock, J.H., Santos, A.S., Tachikawa, H. & Kupriyanova, E.K. (2010) Sabellaria isumiensis n. sp. (Annelida: Polychaeta: Sabellariidae) from shallow waters off Onjuku, Boso Peninsula, Japan, and re-descriptions of three Indo-West Pacific sabellariid species. Zootaxa, 2680, 1–25. Nishi, E., Kato, T. & Hayashi, I. (2004) Sabellaria tottoriensis n. sp. (Annelida: Polychaeta: Sabellariidae) from shallow water off Tottori, the Sea of Japan. Zoological Science, 21, 211–217. Nishi, E. & Kirtley, D.W. (1999) Three new species of Sabellariidae (Polychaeta) from Japan. Natural History Research, 5(2), 93–105. Nishi, E. & Núñez, J. (1999) A new species of shallow water Sabellariidae (Annelida: Polychaeta) from Madeira Island, Portugal, and Canary Islands, Spain. Arquipélago. Life and Marine Sciences, 17A, 37–42. Orrhage, L. (1978) On the structure and evolution of the anterior end of the Sabellariidae (Polychaeta Sedentaria). With some remarks on the general organisation of the polychaete brain. Zoologische Jahrbuecher, Abteilung fűr Anatomie und Ontogenie der Tiere, 100, 343–374. Peters, F.A. (1854) Naturwissenschaftliche Reise nach Mosambique in 1842 bis 1848 ausgefurt. Akademie der Wissenschaften zu Berlin, Monatsberichte, 1854, 610–614. Pohler, S.M.L. (2004) The sabellariid worm colonies of Suva Lagoon, Fiji. South Pacific Journal of Natural Science, 22, 36–42. Quatrefages, A. de. (1849) Études sur les types inférieurs de l'embranchement des Annéles. Mémoire sur la famille des



57

Chlorhemiens, Chloraema Nob. Annales des Sciences Naturelles, Paris, 3(12), 277–306. Rathke, H. (1837) Beitrage zur Fauna der Krym. Mémoires des savantes étranges. Mémoires de l’Academie impériale des Sciences, St.-Petersbourg, 3, 407–429. Rèaumer, R. (1817) Mémoires sur les differents manières dont plusiers espèces d’animaux de mer s’attachent au sable, aux pierres, et les unes aux autres. Mémoires de l’Academie des Sciences, Paris, 1711, 130. Rochebrune, M. (1882) Diagnose d’especies nouvelles pour la faune de l’Archipel du Cap Vert. Annulata. Bulletin de la Société Philomathiques, Paris, série 7, 6, 25–26. Rouse, G.W. & Pleijel, F. (2001). Polychaetes. Oxford, Oxford University Press. Santos, A.S., Riul, P., Claudia, A., Brasil, C.D.S. & Christoffersen, M.L. (2011) Encrusting Sabellariidae (Annelida: Polychaeta) in rhodolith beds, with description of a new species of Sabellaria from the Brazilian coast. Journal of the Marine Biological Association of the United Kingdom, 91(2), 425–438. Schmarda, L.K. (1861). Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise un die Erdr 1853 bis 1857. Erster Band (zweite halfte) Turbellarian, Rotatorien un Anneliden. Wilhelm Engelmann. Leipzig. 161 pp. Sepulveda, R.D., Moreno, R.A. & Carrasco, F.D. (2003) Macroinvertebrate diversity associated to reefs of Phragmatopoma moerchi Kinberg, 1867 (Polychaeta: Sabellariidae) in the intertidal rocky shore at Cocholgue, Chile. Gayana, 67(1), 45–54. Treadwell, A.L. (1901) The Polychaetous annelids of Puerto Rico. Bulletin of the United States Fish Commission, 20, 181–210. Treadwell, A.L. (1906) Polychaetous Annelids of the Hawaiian Islands Collected by the Steamer Albatross in 1902. Bulletin of the United States Fish Commission, 23, 1145–1181. Treadwell, A.L. (1926) Contributions to the biology of the Philippine Archipelago and adjacent regions. Additions to the polychaetous annelids collected by the United States Fisheries steamer 'Albatross', 1907–1910, including one new genus and three new species. Bulletin of the United States National Museum, 100, 183–193. Verrill, A.E. (1873) Report on the Invertebrate Animals of Vineyard Sound and the Adjacent Waters, with an Account of the Physical Characters of the Region. Report for 1871–1872. United States Fish Commission, 72, 295–778. Wilson, D.P. (1977) The Distribution, Settlement and Development of the Sabellarian Polychaete Lygdamis muratus (Allen) near Plymouth. Journal of the Marine Biological Association of the United Kingdom, 57, 761–792.


HUTCHINGS ET AL.

List of Tables TABLE 1. Major distinguishing characters of species of Idanthyrsus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 TABLE 2. Major distinguishing characters of species of Lygdamis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 TABLE 3. Major distinguishing characters of species of Phalacrostemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 TABLE 4. Major distinguishing characters of species of Sabellaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 TABLE 5. Major distinguishing characters of species of Tetreres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 TABLE 6. Character matrix of morphological features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53



59

About the authors Dr Pat Hutchings, is a Senior Principal Research Scientist at the Australian Museum, Sydney and is interested in the phylogeny and systematics of Australian polychaetes, and has spent decades collecting material from around the Australian coast. Dr Maria Capa, has been working as a postdoctoral researcher for the last 6 years at the Australian Museum, undertaking revisions of several groups of polychaetes, describing the Australian fauna and assessing their evolutionary relationships at different hierarchical levels. Dr Rachael Peart was employed at the Australian Museum on this project as a technical officer, but her research interests are amphipods.


HUTCHINGS ET AL.