The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2010 26: 57–67 New records and a new species of Hermundura Müller, 1858, the senior synonym of Loandalia Monro, 1936 (Annelida: Phyllodocida: Pilargidae) from northern Australia and New Guinea Christopher J. Glasby1 and Shona A. Hocknull2 1 Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin NT 0801, AUSTRALIA chris.glasby@nt.gov.au 2 Benthic Australia, PO Box 5354, Manly QLD 4197, AUSTRALIA shona.hocknull@benthicaustralia.com ABSTRACT The Principle of Priority (ICZN 1999) is applied in resurrecting the generic name Hermundura Müller, 1858 for pilargid polychaetes (Annelida) formerly referred to as Loandalia Monro, 1936 and Parandalia Emerson & Fauchald, 1971. All available specimens of Hermundura from northern Australia and New Guinea are described based on material held in the collections of Australia’s natural history museums. Our study recognises two species, H. gladstonensis (Marks & Hocknull, 2006) comb. nov. and a new species, H. philipi sp. nov., from near Mornington Island in the southern Gulf of Carpentaria. The former species is redescribed, and new information on intraspeciic variability is provided, including the presence of hardened, cuticular structures and a muscle band (sphincter) of the anterior alimentary canal. Analysis of these structures has led to a re-evaluation of the so-called ‘jaws’ of the closely related monotypic genus Talehsapia Fauvel, 1932, and the conclusion that they are homologous with the pharyngeal sphincter of Hermundura. Talehsapia therefore is also synonymised with Hermundura. The genus Hermundura now contains 17 species including the type species, Hermundura tricuspis Müller, 1858, H. philipi sp. nov., and ifteen others, which are all new name combinations, viz. Hermundura annandalei (Fauvel, 1932), H. aberrans (Monro, 1936), H. fauveli (Berkeley & Berkeley, 1941), H. americana (Hartman, 1947), H. gracilis (Hartmann-Schröder, 1959), H. indica (Thomas, 1963), H. ocularis (Emerson & Fauchald, 1971), H. maculata (Intes & Le Loeuff, 1975), H. bennei (Solís-Weiss, 1983), H. riojai (Salazar-Vallejo, 1986), H. vivianneae (Salazar-Vallejo & Reyes Barragán, 1986), H. evelinae (León-González, 1991), H. salazarvallejoi (León-González, 1991), H. fredrayorum (Marks & Hocknull, 2006) and H. gladstonensis (Marks & Hocknull, 2006). The genus Hermundura appears to be restricted to the tropics and subtropics, both in Australia and globally. Keywords: Polychaeta, Pilargidae, systematics, taxonomy, redescription, synonymy, new combination. (1990) recommended the continuing use of the junior synonym name (Parandalia), the name Hermundura was ignored. However, under current ICZN (1999) rules, in which strict publication criteria need to be applied in order to suppress a senior synonym, we consider it appropriate to abide by the Principle of Priority and resurrect the senior synonym name for this taxon. Emerson & Fauchald (1971) re-examined the type species of Loandalia, L. aberrans, and concluded that, as its name implies, it has several unique features that set it apart from other synelmines, most importantly the absence of notopodial spines (and notoacicula) and the presence of branchiae. They restricted the concept of the genus to include only the type species and moved the other species in the genus to Parandalia. Salazar-Vallejo (1998) demonstrated that the so-called branchiae were in fact enlarged nephridial papillae and, in his opinion, the presence of nephridial papillae could not be used as a feature to separate Loandalia from Parandalia. Further, because of contradictory information over whether notopodial spines INTRODUCTION The concept and validity of Loandalia Monro, 1936, a genus of pilargid polychaetes, has been in lux since it was originally described by Monro (1936) based on L. aberrans from a specimen from the continental shelf off Angola, southwest Africa. Monro was unable to assign it unequivocally to a family, although he favoured the Hesionidae. Later, Loandalia was assigned to the Pilargidae by Hartman (1947), and it was treated as an established member of that family in the revision by Pettibone (1966). Pettibone regarded the monotypic Hermundura Müller, 1858 as a questionable synonym of Loandalia. Later, Salazar-Vallejo (1990) redescribed the type species of Hermundura, H. tricuspis (as Parandalia tricuspis), designated a neotype, and established unequivocally that it represented a senior synonym of Parandalia. When Salazar-Vallejo (1998) syonymised Parandalia with Loandalia, Hermundura became the senior synonym of both Parandalia and Loandalia, but because Salazar-Vallejo 57 C. J. Glasby and S. A. Hocknull are present in the type – observed by Monro, but not by Emerson & Fauchald (1971) – this feature could not be used to diagnose the genus. We follow Salazar-Vallejo’s (1998) interpretation of the type species, and his conclusion that Parandalia is a junior synonym of Loandalia. This action also addresses the concern of Licher & Westheide (1994) that monophyly of Parandalia was questionable. The monotypic genus Talehsapia Fauvel, 1932 was also regarded as a junior synonym of Loandalia, albeit questionably (Pettibone 1966). The taxonomic history of Talehsapia and its single species T. annandalei Fauvel, 1932 is summarised by Salazar-Vallejo et al. (2001), who also redescribed specimens from the type locality. They concluded that the genus should be maintained and it can be recognised by its completely fused palps and the presence of denticulate bands, which originally were interpreted as jaws. However, we demonstrate herein using a large range of differently preserved specimens of H. gladstonensis, that these two features should not be used to distinguish T. annandalei from other Hermundura. Thus Talehsapia also becomes a junior synonym of Hermundura. Four species of Hermundura are currently known from the Indo-west Paciic, H. annandalei comb. nov. (Thailand), H. indica (Thomas, 1963) comb. nov. (India), and two species from Australia, H. fredrayorum (Marks & Hocknull, 2006) comb. nov., from the type locality Moreton Bay near Brisbane, and H. gladstonensis (Marks & Hocknull, 2006) comb. nov. from the type locality Gladstone, NE Queensland (Table 1). In this paper we describe all available Hermundura specimens from northern Australia and New Guinea based on material held in the collections of Australia’s natural history museums. (formerly AHF) – Los Angeles County Museum of Natural History, Los Angeles; LIPI – Pusat Penelitian Oseanograi, Lembaga llmu Pengetahuan, Jakarta; MEU – Marine Ecology Unit, formerly belonging to NTM; NTM – Museum and Art Gallery, Northern Territory (formerly Northern Territory Museum), Darwin; QM – Queensland Museum, Brisbane. MORPHOLOGY OF HERMUNDURA Body. Hermundura species have a more or less uniformly cylindrical body except for the anterior 5–10 segments which are inlated, and the posterior body may be slightly lattened. The body surface has a smooth to aereolated texture, not shining or papillated like other pilargids. Pigmented subdermal glands may be present on the lateral body and the pygidium (as a pair of spots) (Fig. 1). Head. The prostomium is reduced and indistinct, rectangular or ovate in shape, and together with the reduced palps the head appears bluntly triangular in shape. The palps are biarticulated (comprising a palpophore and a palpostyle) and separated from each other (though they may appear to be medially fused when the head is contracted). At least two species, H. tricuspis Müller, 1858 and H. annandalei comb. nov., are reported in the literature to have fused palps but this needs to be conirmed based on fresh specimens. Palpostyles maybe button-like or rod-like in shape and occur singly or in a pair within each palpophore (see Marks & Hocknull 2006). A palpal papilla is absent. Nuchal organs are not apparent. The antennae are absent. The eyes, if present, are subdermal, lying on the brain. The brain is often visible dorsally through the body wall as an elongate, tripartite structure projecting posteriorly from the prostomium and extending through the irst few anterior chaetigers; the midbrain often has paired subdermal eyespots. The irst annulus (referred to in recent literature as both the peristomium or segment 1) lacks parapodia or tentacular cirri, and it may be set off from the prostomium or fused with it. Parapodia. The parapodia in Hermundura are ‘subbiramous’, with the anterior one or two chaetigers lacking notospines, and dorsal cirri are absent in all species except H. aberrans comb. nov. The chaetal lobe is rectangular, rounded distally and does not project far from the body wall. The ventral cirri are subdistal and start on chaetigers 3–8, thereafter they are present on all chaetigers. The start of the ventral cirri is a useful character to separate species. Nephridial papillae may be present on the posteroventral margin of the neuropodia of posterior chaetigers (only in H. aberrans comb. nov. and H. maculata comb. nov.). Chaetae. The notochaetae include straight notospines (rarely absent, H. aberrans comb. nov.), which start on chaetigers 2–10, and a few accompanying capillaries. The irst few notospines may be overlooked because they have not fully emerged from the parapodium, but they can be seen on slide-mounted specimens under a compound microscope. The irst occurrence of notospines appears not to be body- MATERIALS AND METHODS All specimens were ixed in 10% formaldehyde-seawater and preserved in 70% ethanol. Observations were made using a Nikon SMZ 1500 stereomicroscope and a Nikon Eclipse 80i compound microscope with Nomarsky optics; photographs were made on both microscopes using a Qimaging Micropublisher 5.0 RTV digital camera; stacked images were combined with Helicon Focus software (version 4.77) and line drawings were compiled from digital images using a Wacom Intuos 4 drawing pad with Adobe Illustrator CS2. Generic and species level descriptions were constructed using the DELTA system, Descriptive Language for Taxonomy, a standardised format for coding taxonomic descriptions (Dallwitz 1980; Dallwitz & Paine 1986; http://www.biodiversity.uno.edu.delta). Features included in the generic description were not repeated at the species level and vice versa. The key was constructed manually. The distribution maps were made using PanMap software (Dipenbroek et al. 2000). Abbrevations. The following abbreviations are used in this paper: AM – Australian Museum, Sydney; LACM 58 Hermundura species from northern Australia and New Guinea On the anterior margin of the muscular pharynx a narrow light-refractive band of soft tissue encircles the opening. It appears to be homologous with the ‘denticulated bands’ that Salazar-Vallejo et al. (2001) found in Talehsapia. The intestinal caeca are absent. size dependent as in other pilargids (e.g. Sigambra), and is therefore a good species-level character. The neurochaetae are distinctive, hirsute capillaries that have a whorl of hairs along a variable portion of the chaeta. The notoaciculae are absent, although some earlier authors have interpreted the single notospine as an emergent notoacicula (e.g. Hartman (1947)); a single neuroacicula is present, and the tip may be tapered or knobbed. Pygidium. The pygidium is spoon-like with a dorsal concavity, and ovate to rounded. It bears 2, 3 or 5 anal cirri. Alimentary canal. The anterior-most part of the alimentary canal consists of a short, non-muscular buccal tube which extends from the mouth posteriorly to a muscular pharynx. The buccal tube, which is exposed when the pharynx is partially everted, may carry one or a few hardened denticles which appear to be derived from the cuticle (Fig. 2A), however these features of the buccal tube appear not to be consistent within a species and therefore are given no taxonomic importance herein. Occasionally the denticles appear to be located within the pharynx itself (Figs 2C, 3D), nevertheless they resemble the denticles of the buccal tube rather than the ‘denticulated bands’ described by Salazar-Vallejo et al. (2001). When the proboscis is fully everted, a ring of up to 10 papillae can be seen at the junction between the buccal tube and the muscular pharynx. SYSTEMATICS Pilargidae Saint-Joseph, 1899 Synelminae Sallazar-Vallejo, 1986 Hermundura Müller, 1858 Hermundura Müller, 1858: 216. Gender feminine. Type species, by monotypy, Hermundura tricuspis. Recent, Santa Catarina Island, Brazil. Talehsapia Fauvel, 1932: 251–252. Gender feminine. Type species, by monotypy, Talehsapia annandalei. Recent, Songkhla Lagoon, Thailand. New synonymy. Loandalia Monro, 1936: 193. Gender feminine. Type species, by original designation, Loandalia aberrans. Recent, St Paul de Loanda, Angola. Parandalia Emerson & Fauchald, 1971: 19. Gender feminine. Type-species, by original designation, Parandalia ocularis. Recent, Santa Barbara Channel, California, USA. Table 1: Currently accepted Hermundura species worldwide, arranged chronologically, together with the original name and author, type locality and depth, and nomenclatural/taxonomic comments. Original combination Authority Hermundura tricuspis Müller, 1858 Talehsapia annandalei Fauvel, 1932 Loandalia aberrans Loandalia fauveli Monro, 1936 Berkeley & Berkeley, 1941 Hartman, 1947 Loandalia americana Loandalia gracilis Hartmann-Schröder, 1959 Thomas, 1963 Emerson & Fauchald, 1971 Loandalia maculata Intes & Le Loeuff, 1975 Parandalia bennei Solís-Weiss, 1983 Loandalia riojai Salazar-Vallejo, 1986 Parandalia vivianneae Salazar-Vallejo & Reyes- Barragán, 1986 Parandalia evelinae León-González, 1991 Loandalia salazarvallejoi León-González, 1991 Loandalia fredrayorum Marks & Hocknull, 2006 Loandalia gladstonensis Marks & Hocknull, 2006 Hermundura philipi sp. nov. Glasby & Hocknull, 2010 Loandalia indica Parandalia ocularis Type locality, depth Nomenclatural/taxonomic comments São Antonio de Lisboa, Santa Catarina Island, Neotype designated by Brazil; intertidal Salazar-Vallejo (1990) Songkhla Lagoon,Thailand; no depth stated Redescribed by SalazarVallejo et al. (2001) Off St Paul de Loanda, Angola, 64-65 m Newport Bay, California, USA; intertidal Biloxi, Mississippi, USA; intertidal La Herradura, El Salvador; intertidal Arabian Sea, off west coast of India; 15-18 m Santa Barbara Channel, off southern California; 42-46 m Off Ivory Coast, west Africa; 15-100 m Mazatlan Bay, Paciic coast of Mexico; 3.5-25 m Bahía de Manzanillo, Colima, Mexico; 30-80 m Laguna La Mancha, Veracruz, Gulf of Mexico; shallow water Baha California Sur, Mexico, west coast; 106 m Baha California Sur, Mexico, west coast; 80 m Moreton Bay, Queensland, Australia; 3-11 m Gladstone Harbour, Queensland, Australia; intertidal to 5 m North of Mornington Island, Gulf of Carpentaria, Australia; 35-45 m 59 Synonym of H. fauveli according to Petttibone (1966) Synonym of H. fauveli according to Petttibone (1966) C. J. Glasby and S. A. Hocknull Diagnosis. Body long, slender, subcylindrical, inlated anteriorly; surface smooth to aereolated. Prostomium and palps reduced; combined structure bluntly triangular. Palps biarticulated, separated from each other; palpostyles single or double; palpal papilla absent. Antennae absent. Prostomial eyes absent. Brain longer than wide, divided into fore-, mid-, and hind-brain; forebrain bifurcate, divided anteriorly into 2 lobes, hind brain with 2 lobes; ocular spots present on mid- and hind-brain. First annulus lacks parapodia or tentacular cirri. Parapodia with notopodia reduced, notoaciculae absent and dorsal cirri usually absent. Ventral cirri present, subdistal. Notochaetae including emergent straight notospines and few capillaries. Neurochaetae hirsute capillaries. Neuroaciculae present. Pygidium spoon-shaped plaque (dorsal concavity), bearing 2, 3 or 5 anal cirri. Ring of distal papillae between buccal tube and pharynx; cuticular denticles of various shapes may be present on buccal tube or anterior pharynx, pharynx with a sphincter on anterior margin. Intestinal caecae absent. Nephridial papillae present or absent. Remarks. In 1990 Salazar-Vallejo redescribed the only species of Hermundura, H. tricuspis (as Parandalia tricuspis) and designated a neotype. Later (1998) he synonymised Parandalia with Loandalia. Therefore, Hermundura became the oldest available name for the taxon. Salazar-Vallejo (1990) stated that the name Hermundura “has not been used formally for a long time” and should it be adopted… “confusion would result”. However, under the present ICZN Code (1999) the younger name Loandalia can only be used by successfully arguing the case for ‘prevailing usage’ (ICZN 1999: Article 23.9.1.2). However, Loandalia is a rarely reported taxon and it would be dificult to satisfy the publication criteria set out under Article 23.9.1.2, especially because other names such as Parandalia and Talehsapia have also been used for members of the taxon in the last 50 years. The issue would almost certainly need to be referred to the Commission for a ruling. Therefore, the simplest and most effective option is to apply the Principle of Priority strictly and use the oldest available generic name, which is Hermundura, for the taxon. We have expanded the diagnosis of Hermundura slightly to include the newly synonymised Talehsapia Fauvel, 1932. Talehsapia was differentiated from other synelmine pilargids by its completely fused palps and the presence of pharnygeal denticulate bands (Salazar-Vallejo et al. 2001). However, examination of a large number of specimens of Hermundura during this present study have shown that members of this genus also have palps that are very short and retractile, and that in the retracted state the palps often appear to be fused (Fig. 3A,B) as in Talehsapia. Further, although not mentioned by SalazarVallejo et al. (2001), the palps of Talehsapia appear to be bi-articulated with minute biid palpostyles (see his ig. 1c), which is a condition only known in some species of Hermundura. Finally, Hermundura species also have pharyngeal structures resembling denticulate bands. As Salazar-Vallejo et al. (2001) correctly pointed out, the bands are not true jaws as described by Fauvel (1932) because they are not a solid structure nor do they have cusps or any other features normally associated with jaws. He observed that the denticulate bands are symmetrical but discontinuous laterally, and ‘rugose’ or ‘granular’ due to the presence of ‘many tiny denticles’. Bands of the same form (Fig. 2D,E) and texture (Fig. 2F) were also observed in our material of Hermundura, although the degree of granularity was generally less in our material. We believe that the granularity represents the minute protruberances (e.g. papillae) in the overlying tissue, because the graininess in our specimens extends beyond the limits of the bands (Fig. 2F). The bands appear to represent a sphincter because of their position at the anterior opening of the pharynx. Hermundura gladstonensis (Marks & Hocknull, 2006) comb. nov. (Figs 1, 2A–F, 3A–N, 5A) Loandalia gladstonensis Marks & Hocknull, 2006, 65–66, ig. 3a–d; Hocknull & Glasby 2009: 544 (in part). Material examined. H olotype – QM G222948, Gladstone Harbour, 23˚51.0’S, 151˚37.0’E, coll. S.A. Marks, 26 July 2002. Additional material: AUSTRALIA: Western Australia: Port George, Kimberley coast, NTM W.23378, 15˚23’S, 124˚42’E, coll. J. Jelbart. Northern Territory: Darwin Harbour, approx. 12˚30’S, 130˚47’E, 1–21 m, coll. MEU, July 1993/March 1994, NTM W.10296, NTM W.10297, NTM W.10305, NTM W.10310, NTM W.10298, NTM W.10299, NTM W.10302, NTM W.10307, NTM W.10304, NTM W.10308, NTM W.10300, NTM W.10306, NTM W.10301, NTM W.10309, NTM W.10303, NTM W.13853, NTM W.13844, NTM W.13849, NTM W.13848, NTM W.13868, NTM W.13867, NTM W.13862, NTM W.13864, NTM W.13874, NTM W.13846, NTM W.13845, NTM W.13847, NTM W.13851, NTM W.13843, NTM W.13850, NTM W.13859, NTM W.13860, NTM W.13878, NTM W.13863, NTM W.13870, NTM W.13854, NTM W.13861, NTM W.13856, NTM W.13852, NTM W.13857, NTM W.13876, NTM W.13858, NTM W.13871, NTM W.13879, NTM W.13865, NTM W.13877, NTM W.13869, NTM W.13872, NTM W.13855, NTM W.13866, NTM W.13873, NTM W.13875; opposite Port, Wickham Point, NTM W.23368, 12˚31.107’S, 130˚52.485’E, coll. K. Metcalfe, Dry Season 2005; Bleesers Creek, NTM W.23369, 12˚27.5713’S, 130˚54.7586’E, coll. M. Neave, 16 April 2007; Hudson Creek, NTM W.22284 (includes DNA sample), 12˚28.93’S, 130˚55.6’E, coll. M. Neave, 1 May 2007; Charles Darwin National Park, NTM W.22277 (includes DNA sample), 12˚27.43’S, 130˚52.01’E, coll. M. Neave, 20 April 2007, NTM W.22304, 12˚27.43’S, 130˚52.01’E, coll. M. Neave, 20 August 2007, NTM W.22318, 12˚27.43’S, 130˚52.01’E, coll. M. Neave, 2 August 2007; Kitchener Bay, NTM W.21120, 12˚28.09’S, 60 Hermundura species from northern Australia and New Guinea 1992–March 1993, NTM W.7800. Queensland: Burnett River Estuary, NTM W19341, 24 46’S, 152 25’E, coll. P. Crosser, November 1987; AM W.13200, Gladstone, Auckland Creek, St. 4, coll. P. Saenger, August 1976; AM W13568, Gladstone, Calliope River, St. 2, coll. P. Saenger, May 1976. INDONESIA: Papua, Timika, approx. 5˚S, 137˚E, NTM W.23370, coll. LIPI, 9 August 2003, NTM W.23371, coll. LIPI, 28 June 2004, NTM W.23372, coll. LIPI, June 2004, NTM W.23373, coll. LIPI, 28 June 2004, NTM W.23374, coll. LIPI, 28 June 2004, NTM W.23375, coll. LIPI, 9 November 2009, NTM W.23376, coll. LIPI, 24 June 2004, NTM W.23377, coll. LIPI, 24 October 2002, NTM W.23379, NTM W.23380, NTM W.23381, coll. LIPI, 1 August 2002, NTM W.23382, coll. LIPI 28 June 2004, NTM W.23383, coll. LIPI, 28 June 2004, NTM W.23384, coll. LIPI, 28 June 2004, NTM W.23385, coll. LIPI 28 July 2004, NTM W.23386, coll. 28 June 2004, NTM W.23387, coll. LIPI 1 July 2004, NTM W.023388, coll. 1 July 2004, NTM W.23389, coll. LIPI, coll. LIPI, 1 July 2004, NTM W.23390, coll. LIPI, 29 July 2004, NTM W.23391, coll. LIPI, 1 July 2004, NTM W.23392, coll. LIPI 28 June 2004, NTM W.23393, coll. LIPI, NTM W.23394, coll. LIPI, August 2004, NTM W.23395, coll. LIPI 19 November 2009; Digul, approx. 7˚S, 138˚E, NTM W.23396, coll. LIPI, October 2002, NTM W.23397, coll. LIPI, 22 October 2002, NTM W.23398, coll. LIPI, 21 October 2002, NTM W.23399, coll. LIPI, 22 October 2002, NTM W.23400, coll. LIPI, 17 October 2002, NTM W.23401, coll. LIPI, October 2002. Diagnosis. Palps with biid, divergent palpostyles; buccal tube with 2–5 peg-like structures, honey-coloured to clear, or lacking any apparent structures; dorsal cirri absent; ventral cirri present from chaetiger 5; notopodial spines present from chaetiger 2, initially small and embedded, large and emergent from chaetiger 7–12; 5–9 neuropodial chaetae; neuroaciculae more or less tapered, but may have coloured distal knobs in mid-anterior body; pygidium bearing 3 anal cirri, middle one centrally positioned on plaque. Description. Material examined ranges in size from 3.7–60 mm long, 0.4–1.5 mm wide (maximum width excluding parapodia), bearing 30–130 chaetigers. Brown pigment patches on lateral body and paired circular-shaped patches on pygidium (fading over time in alcohol). Body surface areolated; anterior body region (chaetigers 4–5) inlated. Brain with 2 pairs of pigment spots – one pair on mid-brain, other pair on posterior lobes; reddish-brown to black. Palps biarticulated, separate from each other; palpostyles rod-like, biid. Palpophores not fused. Buccal tube with 4–5 peg- or jaw-shaped denticles, or absent; honey-coloured to clear. Pharynx with distal ring of 10 papillae (rarely 8); all papillae more or less same size. Notopodial lobes low (absent on irst 4–6 chaetigers). Notochaetae comprising spines and capillaries; spines present from chaetiger 2, but not fully emergent until Fig. 1. Hermundura gladstonensis, entire animal approx. 12 mm long, lateral view, head facing downwards. Non-type specimen NTM W.23395. Photograph: C. Glasby. 130˚50.478’E, 3.3 m, coll. C. Clark, 23 April 2007, NTM W.19239, 12˚28.11’S, 130˚50.83’E, 1 m, URS consultants, 22 September 2004. Melville Bay, Gove, approx. 12˚12’S, 136˚42’E, 5–18 m, coll. J.R. Hanley, July 1991/March 1992, NTM W.16824, NTM W.16825, NTM W.16828, NTM W.16830, NTM W.16831, NTM W.16832, NTM W.16833, NTM W.16834, NTM W.16836, NTM W.16842, NTM W.16846, NTM W.16848, NTM W.16849, NTM W.16850, NTM W.16852, NTM W.16853, NTM W.16855, NTM W.16856, NTM W.8229, NTM W.16826; NTM W.16840, NTM W.16841, NTM W.16843, NTM W.16851, NTM W.16844, NTM W.16837, NTM W.16835, NTM W.16838, NTM W.16839, NTM W.16845, NTM W.16847, NTM W.16854, NTM W.16823, NTM W.16827, NTM W.16829; Mangrove Creek, 12˚12’S, 136 43’E, coll. NT Fisheries Department, 22 August 1971 NTM W.592; Bing Bong, McArthur River, 15 37’S, 136 23’E, coll. MEU, September 61 C. J. Glasby and S. A. Hocknull Fig. 2. Hermundura gladstonensis, structures associated with anterior alimentary canal including buccal tube (A–C) and pharynx (D–F): A, jaw-like denticle (NTM W.23395); B, conical denticle (NTM W.23392); C, sickle-shaped denticle (NTM W.23386); D, undissected pharynx, anterior end, showing laterally well-developed musculature resembling jaws (NTM W.22304); E, dissected pharynx, anterior end, showing transverse sphincter and longitudinal muscle sutures (NTM W.22318); F, dissected pharynx showing close up of sphincter (NTM W.22318). Scale bars: A–E, 0.1 mm; F, 20 μm chaetigers 7–8, some with reddish-brown colouring distally (may also fade in alcohol); spines well developed until last chaetiger. Notopodial capillary chaetae present from chaetiger 2. Notoaciculae absent. Neuropodial lobe digitiform, very low on irst 1–3 chaetigers, gradually inceasing in size to maximum length in mid body. Ventral cirri present from chaetiger 5. Neurochaetae capillary-like, with numerous transverse rows of slender teeth over short portion of chaeta, 5–9 per fascicle. Neuroaciculae present, distally knobbed or tapered (knob-tipped types sometimes present on chaetigers 8–22). Paired lateral anal cirri present, papilliform. Mid-anal cirri present, papilliform, centrally positioned on plate. Anal opening dorsal. Remarks. The large amount of material available for study, covering a wide geographic range, has allowed us to document the intraspeciic variability of this common 62 Hermundura species from northern Australia and New Guinea Fig. 3. Hermundura gladstonensis, A, anterior end, dorsal view (NTM W.22318); B, anterior end, ventral view (NTM W.22318); C, anterior end, dorsal view, specimen partially cleared with glycerol to show buccal denticles (NTM W.23373); D, pharynx dissected from body showing denticles (NTM W.23386); E, pygidium dorsal view (NTM W.22318); F, pygidium ventral view (NTM W.22318); G, parapodium of chaetiger 2 (NTM W.22318); H, parapodium of chaetiger 60 (NTM W.22318); I, parapodium of chaetiger 17 (NTM W.23373); J, parapodium of chaetiger 13 (NTM W.23373); K, neuroacicula, anterior chaetiger (NTM W.22318); L, M, neuroacicula, anterior chaetiger (NTM W.23373); N, hirsute neurochaeta, anterior chaetiger (NTM W.22318). Scale bars: A, B: 0.5 mm; C–F, 0.2 mm; G–J, 0.1 mm; K–M, 40 μm; N, 25 μm. 63 C. J. Glasby and S. A. Hocknull species, which until now was only known from the type description and the brief account of the forms from the Gulf of Carpentaria by Hocknull & Glasby (2009) (which is actually a composite of two species – H. gladstonesis and H. philipi sp. nov.). Two of the most interesting variable features are the presence of cuticularised denticles on the buccal tube and the pigmented knobbed-tipped neuroaciculae. The cuticularised denticles may be elongate and resemble serrated jaws (Fig. 2A), or be conical to sickle-shaped (Figs 2B,C; 3C,D); the denticles numbered from 1 to 4 and were sometimes arranged 2 ventrally and 2 dorsally. Their occurrence within the taxon appears likely to be size-related (i.e. more common in larger worms) rather than geographic- or sex-related. Hardened structures of the buccal tube have not been reported previously for the family, although similar structures of the pharynx were thought to exist (see Remarks for genus). Similarly, variability in the presence or absence of knob-tipped neuroaciculae could not be easily explained. They were observed only in parapodia of the irst 20 or so chaetigers; the neuroaciculae of some individials were always tapered (Fig. 3K), whilst those of others showed various degrees of knobbiness (Fig. 3 L, M). In addition, the presence of knobbed-tipped neuroaciculae appeared to correspond with a reddish pigment spot at the tip of the same neuropodium – whether the tip itself was red or it was the tissue of the distal neuropodium could not be established. Knob-like protruberances have also been reported by Salazar-Vallejo (1990) on the distal region of the notopodial spines of the type species (H. tricuspis) raising the possibility that they represent aberrations common to members of the genus. Habitat. Hermundura gladstonensis occurs from the lower intertidal zone to about 20 m in soft sediments (for the northern Australia material). It is not present in the higher intertidal zone under the mangrove canopy in Darwin Harbour (CJG pers. obs.). Distribution. Sahul shelf including northern Australia and eastern Indonesia; also coastal north-eastern Australia, excluding the Great Barrier Reef (Fig. 5A). Fig. 4. Hermundura philipi sp. nov. Holotype, A, anterior end, dorsal view; B, anterior end, ventral view; C, parapodia from chaetiger 16; D, pygidium dorsal view; E, pygidium ventral view; F, Neuroacicula, anterior chaetiger; G, hirsute neurochaeta, anterior chaetiger. Scale bars: A, B, 0.2 mm; C, 0.05 mm; D–E, 0.1 mm; F–G, 15 μm. 64 Hermundura species from northern Australia and New Guinea Hermundura philipi sp. nov. (Fig. 4A–G, 5B) Loandalia gladstonensis Marks & Hocknull, 2006, 65–66, ig. 3a–d; Hocknull & Glasby 2009: 544 (in part). Material examined. H olotype – QM G230618, Queensland, Gulf of Carpentaria, north of Mornington Island, coll. CSIRO R.V. Southern Surveyor, 15˚59.4486’S, 139˚53.346’E, 37.8 m, 2 March 2005. paratypes – collection details as for holotype, 1(QM G230616), 15˚48.4818’S 139˚47.4612’E, 42.6 m, coll. 1 March 2005; 3(QM G230619), 16˚0.5412’S, 139˚45.5016’E, 39.8 m, coll. 3 March 2005; 1(QM G230620), 16˚1.3464’S, 139˚36.0294’E, 39.8 m, coll. 3 March 2005; 1(QM G230624), 15˚48.5046’S, 139˚45.5112’E, 43.6m, coll. 4 March 2005; 1(NTM W.21979), 15˚56.5596’S, 139˚41.109’E, coll. 2 March 2005; 1(NTM W.21980), 15˚55.4136’S, 139˚53.7162’E, coll. 2 March 2005; 1(NTM W.21981), 16˚0.5676’S, 139˚37.4982’E, coll. 3 March 2005. Non-type material. QUEENSLAND: Gulf of Carpentaria, north of Mornington Island, coll. CSIRO R.V. Southern Surveyor, QM G230611, 15˚58.3164’S, 139˚39.3606’E, 41.4 m, coll. 4 March 2005; QM G230612, 16˚9.5718’S, 139˚44.4606’E, 35.4 m, coll. 28 February 2005; QM G230613, 15˚55.4622’S 139˚28.497’E, 43 m, coll. 27 February 2005; QM G230614, 15˚58.488’S, 139˚40.4964’E, 41.4 m, coll. 27 February 2005; QM G230615, 15˚59.5026’S, 139˚41.4948’E, 40.6 m, coll. 1 March 2005; QM G230617, 15˚58.9056’S, 139˚44.0076’E, 39.8 m, coll. 2 March 2005; QM G230621, 16˚0.5676’S, 139˚37.4982’E, 41.6 m, coll. 3 March 2005; QM G230622, 16˚7.9314’S, 139˚46.479’E, 37.8 m, coll. 4 March 2005; QM G230623, 15˚58.3164’S, 139˚39.3606’E, 41.4 m, coll. 4 March 2005; QM G230625, 15˚59.6574’S, 139˚39.2496’E, 41 m, coll. 4 March 2005; QM G230626, 16˚9.4224’S, 139˚39.5118’E, 34.6 m, coll. 4 March 2005; QM G230627, 15˚59.4978’S, 139˚42.498’E, 39.8 m, coll. 4 March 2005; QM G230628, 16˚0.6588’S, 139˚35.2044’E, 41.8 m, coll. 5 March 2005; QM G230629, 15˚58.5228’S, 139˚44.4756’E, 40.4 m, coll. 5 March 2005; QM G230630, 15˚57.4908’S, 139˚36.5082’E, 41.8 m, coll. 5 March 2005; QMG 230631, 16˚1.3656’S, 139˚46.4376’E, 39.4 m, coll. 5 March 2005; QM G230632, 16˚1.6434’S, 139˚39.7044’E, 41.4 m, coll. 6 March 2005. Diagnosis. Body with constriction after anterior inlated segment (between chaetigers 6–7); palps with simple, button-like palpostyle; pharynx and buccal tube unarmed; dorsal cirri absent; ventral cirri present from chaetiger 7; notopodial spines present from chaetiger 5, initially small and embedded, large and emergent from chaetiger 7; 5 or 6 neuropodial chaetae, with large proportion of chaetae hirsute; neuroaciculae of mid-body tapered; pygidium bearing 3 anal cirri, middle one located centrally on plaque. Description. Holotype 6.0 mm long, 0.45 mm wide (max. width including parapodia) for 35 chaetigers. Paratypes range in size from 2.2–10.0 mm long, 0.40–0.50 mm wide, with 21–55 chaetigers. Brown pigment A B Fig. 5. A, Distribution of Hermundura gladstonensis; B, distribution of H. philipi sp. nov. patches on lateral body and paired circular-shaped patches on pydidium. Body surface areolated anteriorly, smooth posteriorly; anterior body region (chaetigers 1–6) inlated, with slight constriction between chaetigers 6–7 (Fig. 4A,B). Brain with 3 pairs pigment spots; 2 pairs on mid-brain and 1 pair on hind-brain. Palps biarticulated, separate from each other; palpostyles button-like; simple. Buccal tube with tooth-like structures not observed. Notopodial lobes low (absent on irst 6 chaetigers). Notochaetae comprising spines and capillaries; spines present from chaetigers 5–6 very small and not emergent initially; from chaetiger 7 onward, large fully developed emergent spine to last chaetiger (Fig. 3C). Notopodial capillary chaetae present from chaetiger 2, 0–3 per fascicle. Notoaciculae absent. Neuropodial lobe digitiform. Ventral cirri present from chaetiger 7. Neurochaetae capillarylike, with numerous transverse rows of slender teeth covering large portion of chaeta, 5–6 per fascicle (Fig. 4G). Neuroaciculae present, tapering to a slightly expanded tip (Fig. 4F). Pygidium plate-like (Fig. 4D). Paired lateral anal cirri present, papilliform; mid-ventral anal cirri present, positioned centrally on plate (Fig. 4E). Anal opening dorsal. Intraspecific variation. The largest non-type specimen was 16 mm long, 0.75 mm wide for 59 chaetigers (incomplete). Specimen QM G230627 was encased in a hyaline tube, similar to those of species of Phyllochaetopterus and Spiochaetopterus (Chaetopteridae). 65 C. J. Glasby and S. A. Hocknull It is therefore possible that H. philipi is commensal with a chaetopterid, especially because other Hermundura species have also been found associated with Phyllochaetopterus tubes (Berkeley & Berkeley 1941; Thomas, 1963), and other pilargids are known to be commensal with chaetopterids, for example, Pilargis berkeleyae is known to be associated with Chaetopterus tubes (Britaev 1993). Etymology. The new species is named in honour of Philip Marks, the junior author’s father. Remarks. The brief account of Hermundura gladstonensis by Hocknull & Glasby (2009) is a composite of two species, H. gladstonesis and H. philipi sp. nov. The new species appears to be closest in morphology to H. fredrayorum and H. indica (Thomas, 1963), which also have ventral cirri starting on chaetiger 7. Hermundura fredrayorum differs in being a much larger species (181 mm long, 2.55 mm wide with chaetae), with its notopodial spines starting on chaetiger 9, in having a greater number of both notopodial capillaries (2–4) and neuropodial chaetae (20–24), and in lacking a mid-ventral anal cirrus. Hermundura indica differs in having the irst few chaetigers biramous, notopodial spines present from chaetiger 7, and having 5 anal cirri. The new species may be distinguished from all other species in the Indo-west Paciic region using the key below. Habitat. Hermundura philipi occurs in about 35–45 m in soft sediments. Distribution. This species is presently known only from the southern Gulf of Carpentaria (Fig. 5B). Key to Hermundura species of Australia and the Indowest Paciic 1. Ventral cirri present from chaetiger 4; palpostyles biid .................................................... H. annandalei — Ventral cirri present from chaetiger 5; palpostyle biid, divergent......................................... H. gladstonensis — Ventral cirri present from chaetiger 7; palpostyles simple ..................................................................... 2 2. — — Notopodial spines present from chaetiger 5, initially small and embedded, large and emergent from chaetiger 7; pygidial plate with 3 anal cirri .............. ...................................................... H. philipi sp. nov. Notopodial spines present from chaetiger 7; pygidial plate with 5 anal cirri ................................ H. indica Notopodial spines present from chaetiger 9, initially small and embedded, large and emergent from midbody; pygidial plate with 2 anal cirri.................. ..........................................................H. fredrayorum ACKNOWLEDGEMENTS For access to specimens used in this study, we thank Stephen Keable (AM), Inayat al Hakim (LIPI) and Mal Bryant (QM). We thank Hannelore Paxton and Robin Wilson for constructive comments in review and Richard Willan with advice on interpretation of the Code. This study was funded by grant number 208-35 of the Australian Biological Resources Study, Canberra. The authors gratefully acknowledge this support. DISCUSSION The expanded deinition of Hermundura containing both Talehsapia and Parandalia now gives the genus a total of 17 species worldwide (Table 1). Three species of Hermundura occur in the northern Australian region. Although H. gladstonensis and H. philipi potentially overlap in distribution in the southern Gulf of Carpentaria, it appears that they are disjunct based on habitat preference. Hermundura gladstonensis is restricted to coastal areas down to 20 m, whereas H. philipi occurs in waters from 35–45 deep. This pattern of habitat partitioning agrees with the general pattern identiied for the northern Australia Pilargidae (Hocknull & Glasby 2009), viz. an inshore component (e.g. H. gladstonensis, H. fredrayorum), a shallow shelf component (H. philipi), and a deep shelf component (includes other pilargids, but no species of Hermundura in this category). Interestingly, Hermundura fredrayorum was not reported in this study. It appears to have a limited distribution in the subtropical east coast of Australia. The three species may be distinguished from each other and from remaining Indowest Paciic Hermundura species using the following key. REFERENCES Berkeley, E. & Berkeley, C. 1941. 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