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Zootaxa 1119: 59–68 (2006) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 1119 Copyright © 2006 Magnolia Press ISSN 1175-5334 (online edition)

New of Loandalia (Polychaeta: ) from Queensland, Australia

SHONA MARKS1 & SCOTT HOCKNULL2 1 S. A. Marks. CSIRO Marine Research, PO Box 120, Cleveland QLD 4163. [email protected]. 2 S. A. Hocknull. Queensland Museum, 122 Gerler Rd Hendra QLD 4711. [email protected]

Abstract

Two new species of Loandalia are described from Queensland, Australia. Loandalia fredrayorum sp. nov. is described from Moreton Bay, south eastern Queensland and is distinguished from all other species of Loandalia by the presence of singular palpostyles; uniramous parapodia at chaetiger 1; an emergent notopodial spine at chaetiger 9; neurochaetae numbering 20–24; ventral cirri begin on chaetiger 7 and the pygidium with two lateral papillae-like anal cirri. Loandalia gladstonensis sp. nov. is described from Gladstone Harbour, central eastern Queensland and is distinguished from all other species of Loandalia by the presence of bifid palpostyles; chaetiger 1 uniramous with remaining chaetigers biramous; an emergent notopodial spine from chaetiger 7–8; ventral cirri present from chaetiger 5 and neurochaetae numbering 5–6.

Key words: Loandalia fredrayorum sp. nov., Loandalia gladstonensis sp.nov., Pilargidae, Queensland, Australia, new species, systematics.

Introduction

Saint-Joseph (1899) established the Pilargidae for the new species Pilargis verrucosa Saint-Joseph. Prior to this, pilargids had been placed in several different families including the , and (Licher & Westheide 1994). Recent cladistic analyses of the firmly recognise Pilargidae as a distinct clade (Glasby 1993; Pleijel & Dahlgren 1998). Hutchings and Johnson (2003) and Licher and Westheide (1994) recognise twelve pilargid genera including Loandalia Monro, 1936 and Parandalia Emerson & Fauchald, 1971, however, Salazar-Vallejo (1998) considers Parandalia to be a junior synonym of Loandalia due to an absence of autapomorphic characteristics. Talehsapia Fauvel, 1932 and Loandalia share the closest morphological features, including a nearly complete

Accepted by G. Rouse: 2 Dec. 2005; published: 30 Jan. 2006 59 ZOOTAXA absence of all cephalic appendages and a strong reduction of the dorsal and ventral cirri. 1119 Talehsapia is distinguished from Loandalia by the presence of jaws. Licher and Westheide (1994) suggest that the reduced features seen throughout Pilargidae, especially in Loandalia, are due to a progenetic retention of juvenile morphology, from an ancestral hesionid. The three genera Sigambra, Ancistrosyllis and Synelmis, listed by Hartmann-Schröder (1959) and Hutchings and Murray (1984) have been reported from Australia. This has recently been amended to include undescribed species of Cabira, Litocorsa, Loandalia, Parandalia and Pilargis by Glasby (2000). Fifteen Loandalia species are recognized worldwide with most species occurring in the Central and North American regions, in particular, the Gulf of Mexico (Fig 1). Loandalia fauveli and L. tricuspis have the widest distribution of all Loandalia species. Loandalia fauveli is found farthest north to Alaska (Banse & Hobson 1974) and south to Chile (Rozbaczylo & Quiroga 2000). Loandalia tricuspis is found farthest south to Uruguay and north to the Gulf of Mexico (Salazar-Vallejo & Orensanz 1991). Loandalia indica, L. aberrans and L. maculata are all found outside this western hemisphere distribution, occurring respectively in the Arabian Sea west of India (Thomas 1963); coastal Angola (Monro 1936); and off the Ivory Coast (Intes & Le Loeuff 1975). The first reports of two new species from the South Western Pacific of Australia are presented herein.

FIGURE 1. World distribution map of Loandalia species. L. aberrans, solid triangle; L. americana, solid diamond; L. bennei, left-hand open triangle; L. evelinae, open triangle; L. fauveli, solid circle; L. fredrayorum sp. nov. solid star; L. gladstonensis sp. nov. open star; L. gracilis, open square; L. indica, open diamond; L. maculata, solid cross; L. ocularis, open circle; L. riojai, solid cross in circle; L. salazarvallejoi, solid hexagon; L. tricuspis, solid square; L. vivianneae, upside- down solid triangle.

60 © 2006 Magnolia Press MARKS & HOCKNULL Loandalia species occur in a variety of habitats. Loandalia americana was reported by ZOOTAXA Hartman (1947) and L. bennei by Solís-Weiss (1983) from shallow intertidal areas and 1119 sandy beaches. Loandalia vivianneae was reported by Salazar-Vallejo & Reyes-Barragán (1990) from seagrass beds and mangrove forests, whilst L. indica and L. ocularis have been found in estuaries and mud flats (Thomas 1963; Emerson & Fauchald 1971). Species of Loandalia have been found at varying depths, up to 106 m reported for L. evelinae by León-González (1991). Specimens referred to here as Loandalia fredrayorum sp. nov. and L. gladstonensis sp. nov. are deposited in the Queensland Museum and are given the collection prefix QMC G. Types species of Loandalia are in the Natural History Museum London collections, prefix NHM.

Systematics

Order Phyllodocida Dales

Family Pilargidae Saint-Joseph

Genus Loandalia Monro

Type Species: Loandalia aberrans Monro, 1936. Type number NHM 1936-2-8-3376. [Angola, Africa]

Generic composition: L. americana Hartman (1947), [Gulf of Mexico]; L. bennei (Solís-Weiss 1983), [Mazatlan Bay, Mexico]; L. evelinae (León-González 1991), [Bahia of California]; L. fauveli Berkeley & Berkeley (1941), [Southern California]; L. fredrayorum sp. nov., [Moreton Bay, Australia]; L. gladstonensis sp. nov., [Gladstone Harbour, Australia]; L. gracilis Hartmann-Schröder (1959), [El Salvador]; L. indica Thomas (1963), [Arabian Sea]; L. maculata Intes & Le Loeuff (1975), [Ivory Coast]; L. ocularis Emerson & Fauchald (1971), [Santa Barbara, California]; L. riojai Salazar-Vallejo (1986), [Bahia of Manzanillo, Mexico]; L .salazarvallejoi León-González (1991), [Western Coast of Baja California, Mexico]; L. tricuspis Müller (1858), [Brazil]; L. vivianneae (Salazar-Vallejo & Reyes-Barragán 1990), [Eastern Mexico]. Diagnosis: Body cylindrical. Prostomium wider then long and reduced; biarticulated palps reduced. Median and lateral antennae absent; peristomial cirri absent. Jaws absent. Emergent notopodia spine present, straight; capillary notochaetae present, beside spine; notoacicula absent. Dorsal cirri absent (except L. aberrans). Ventral cirri present.

A NEW LOANDALIA © 2006 Magnolia Press 61 ZOOTAXA Loandalia fredrayorum sp. nov. 1119 Fig. 2, a–d.

Etymology: For Frederick Marks and Raymond Fitzgerald. Material: Holotype. QMC G223179, Loandalia fredrayorum sp. nov. [Middle Banks Moreton Bay, S27°12’, E153°19’, collected July/ August 1982]. Paratype. QMC G223180 [Middle Banks Moreton Bay, S27°12’, E153°19’, July/ August 1982]. Habitat is a submerged sandbank with a depth of 3–11 m. Diagnosis: Singular palpostyles. Parapodia uniramous on chaetiger 1, thereafter biramous; notopodial spine emergent at chaetiger 9; notopodial chaetae numbering 2–4; neurochaetae numbering 20–24; ventral cirri from chaetiger 7; dorsally positioned anus with two lateral papillae-like anal cirri.

FIGURE 2. Loandalia fredrayorum sp. nov. Holotype QMC G223179 a) Anterior region, dorsal view. Scale = 1mm. b) Parapodia of chaetiger 21, anterior view right side, Scale = 0.2mm. c) Posterior region, dorsal view, Scale = 0.5mm. d) Posterior region, ventral view, Scale = 0.5mm.

Description: Pilargid of large size, length 181mm, width 1.85mm (without chaetae), 2.55 (with chaetae); 218 chaetigers. Body of preserved individuals creamy/white to reddish/brown.

62 © 2006 Magnolia Press MARKS & HOCKNULL Body corpulent; posterior segments slightly dorso-ventrally flattened. Chaetiger 1–6 ZOOTAXA inflated, with segmentation inconspicuous, surface areolated; lateral-grooves extend from 1119 first parapodia to pygidium (Fig 2, a). Prostomium discrete; palpophores divided; palpostyles are singular and papillae-like. Pharynx eversible; eye spots absent. First parapodium uniramous with remaining chaetigers biramous; notopodial lobes present at chaetiger 2, fully developed by chaetiger 6; 2–4 notochaetae present from chaetiger 2; notopodial spines weakly emerging at chaetiger 9, fully emergent at chaetiger 46. Neuropodial lobe larger than notopodia, reduced in chaetiger 1–4; simple curved neurochaetae with numerous whorled teeth; Neurochaetae in 5–6 bundles of 4, totalling 20–24 neurochaetae per parapodia; emergent neuroacicula from chaetiger 1, positioned anterior to chaetae. Ventral cirri appears from chaetiger 7, fully developed on chaetiger 8– 9 (Fig 2, b). Two most posterior segments possesses emergent notopodial spines and neuroacicula, chaetae are absent, with considerably reduced parapodia lobes. Pygidium as an anal plate, 2 lateral anal cirri present, papillae-like, mid ventral anal cirrus not visible; ventral surface of pygidium plate has ‘m’ like groove (Fig 2, d); anus is positioned dorsally (Fig 2, c). Remarks: Loandalia fredrayorum differs from all other species of Loandalia by possessing the highest neurochaetae count, 20–24. L. fredrayorum differs from L. aberrans, L. gracilis, L. indica and L. ocularis by possessing uniramous parapodia only on chaetiger 1, rest biramous. Differs further from L. aberrans by the absence of dorsal cirri. L. fredrayorum differs from L. bennei, L. evelinae, L. riojai and L. gladstonensis sp. nov. by possessing singular palpostyles. Differing further from L. bennei, L. evelinae and L. gladstonensis by possessing notopodial spines emergent at chaetiger 9. L. fredrayorum differs from L. tricuspis and L. vivianneae by possessing ventral cirri starting at chaetiger 7 versus chaetiger 4; and notopodial spines starting at chaetiger 9 versus chaetiger 7. L. fredrayorum differs from L. americana, L. evelinae and L. maculata by possessing ventral cirri starting from chaetiger 7 versus chaetiger 6. Further differing from L. americana by possessing an emergent notopodial spine from chaetiger 9 versus chaetiger 2. L. fredrayorum differs further from L. fauveli by possessing a notopodial spine at chaetiger 9 versus chaetiger 8. L. fredrayorum differs further from L. maculata by possessing a notopodial spine at chaetiger 9 versus chaetiger 6. L. fredrayorum differs from L. salazarvallejoi by possessing anal cirri that are papillae-like versus elongate (longer than pygidium). Distinguishing features summarised in table 1.

A NEW LOANDALIA © 2006 Magnolia Press 63 ZOOTAXA 1119

64 © 2006 Magnolia Press MARKS & HOCKNULL Loandalia gladstonensis sp. nov. ZOOTAXA Fig 3, a–d. 1119

Etymology: Gladstone Harbour, type locality. Type Material: Holotype. QMC G222948, Loandalia gladstonensis sp. nov. [Gladstone Harbour; station GH144, S23°57’976”, E151°20’908”; depth 0–5m]. Paratypes. QMC G222949, [Gladstone Harbour; station GH148, S23°50’894”, E151°13’811”; intertidal]. QMC G222950, [Gladstone Harbour; station GH038, S23°47’162”, E151°13’276”; intertidal]. QMC G222951, [Gladstone Harbour; station GH140, S23°51’170”, E151°18’892”; depth 0–5m]. QMC G222952, [Gladstone Harbour; station GH016, S23°44’431”, E151°08’132”; intertidal]. QMC G222953, [Gladstone Harbour, station GH014; S23°43’562”, E151°13’276”; depth 0–5m]. Habitat is intertidal or at depths of 0–5m, usually with sand or muddy substrates. Diagnosis: Bifid palpostyles; chaetigers 1–5 inflated; pharynx with distal ring of papillae (2 groups of 5); chaetiger 1 uniramous with remaining chaetigers biramous; single notochaeta present; straight emergent spine from chaetiger 7–8; ventral cirri present from chaetiger 5; 5–6 neurochaetae; posterior most segment possesses emergent notopodial spines and neuroacicula, with reduced parapodial lobes. Description: Pilargids of small size, length 30–62mm, with 79–114 chaetigers. Body of preserved individuals creamy/white to reddish/brown. Body slender with chaetiger 1–5 inflated; segmentation inconspicuous. Posterior segments slightly dorso-ventrally flattened. Prostomium discrete; palpophores divided; palpostyles papilla-like and bifid. Pharynx eversible with distal ring of 10 papillae, 5 dorsal and 5 ventral to pharyngeal slit; papillose band on pharynx, proximal to prostomium; pharyngeal slit vertical. Eye spots present, subdermal at chaetiger 1 (Fig 3, a). First parapodium uniramous with remaining chaetigers biramous; first neuropodia positioned dorsally compared to remaining chaetigers. Notopodial lobes fully developed by chaetiger 4; a single notochaetae present from chaetiger 2; notopodial spines emerging at chaetiger 7–8, increasing in length in posterior most 10 segments. Neuropodial lobe larger than notopodia, reduced in chaetiger 1–2; neurochaetae capillaries with numerous whorled teeth; 5–6 neurochaetae present per parapodia; emergent acicular from chaetiger 2. Ventral cirri from chaetiger 5(Fig 3, b). Posterior most segment possesses emergent notopodial spines and neuroacicula, with considerably reduced parapodial lobes. Pygidium as an anal plate, dorsally concave with dorsal terminating anus, positioned at base of concave plate; 3 anal cirri present, papillae- like, 2 laterally and 1 mid ventrally positioned (Fig 3, c&d). Remarks: Loandalia gladstonensis differs from all species of Loandalia (except L. bennei, L.

A NEW LOANDALIA © 2006 Magnolia Press 65 ZOOTAXA evelinae and L. riojai) by possessing bifid palpostyles. L. bennei, L. evelinae and L. riojai 1119 possess bifid palpostyles. L. gladstonensis further differs from L. gracilis and L. indica by possessing uniramous parapodia. L. gladstonensis differs from L. riojai, L. indica, L. ocularis, L. fauveli and L. americana by possessing a higher number of pharyngeal papillae (10). L. gladstonensis differs from L. evelinae, L. bennei and L. riojai by possessing a large rounded plate as the pygidium with papillae-like anal cirri. L. gladstonensis differs from L. evelinae, L. ocularis and L. vivianneae by the presence of emergent spines and neuroacicula on the last segment. L. gladstonensis differs from L. aberrans by lacking dorsal cirri. L. salazarvallejoi and L. fredrayorum differ from L. gladstonensis by possessing emergent notopodial spines starting at chaetiger 9 and 10 respectively; higher neurochaetae counts, 10–12 and 20–24 respectively. L. gladstonensis differs from L. maculata and L. tricuspis by possessing ventral cirri starting at chaetiger 5, versus chaetiger 4 in L. tricuspis and chaetiger 6 in L. maculata. Distinguishing features summarised in table 1.

FIGURE 3. Loandalia gladstonensis sp. nov. Holotype QMC G222948 a) Anterior region, dorsal view. Scale = 0.5mm. b) Parapodia of chaetiger 25, anterior view, left side. Not to scale. c) Posterior region, dorsal view. Scale = 0.2mm. d) Posterior region, ventral view. Scale = 0.2mm.

66 © 2006 Magnolia Press MARKS & HOCKNULL Acknowledgements ZOOTAXA 1119 We thank Dr David Currie, Damon Shearer, Vikki Shearer and Kirsty Small of the Centre for Environmental Management, Central Queensland University and Dr Mel Bryant (Queensland Museum) for the use of equipment, their assistance and the opportunity to describe these specimens. We thank the Editor and all the reviewers for their valuable time and suggestions for the manuscript. Robin Wilson, Ted Wassenberg, Roland Pitcher, Philip Marks, Wendy Marks and Aston Marks are thanked for their support and assistance.

References

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