Annelids of the eastern Australian abyss collected by the 2017 RV ‘Investigator’ voyage

Laetitia M. Gunton; Elena K. Kupriyanova; Tom Alvestad; Lynda Avery; James A. Blake; Olga Biriukova; Markus Böggemann; Polina Borisova; Nataliya Budaeva; Ingo Burghardt; Maria Capa; Magdalena N. Georgieva; Christopher J. Glasby; Pan-Wen Hsueh; Pat Hutchings; Naoto Jimi; Jon A. Kongsrud; Joachim Langeneck; Karin Meißner; Anna Murray; Mark Nikolic; Hannelore Paxton; Dino Ramos; Anja Schulze; Robert Sobczyk; Charlotte Watson; Helena Wiklund; Robin S. Wilson; Anna Zhadan; Jinghuai Zhang

doi:10.3897/zookeys.1020.57921

Research Article

Annelids of the eastern Australian abyss collected by the 2017 RV ‘Investigator’ voyage

Laetitia M. Gunton^‡, Elena K. Kupriyanova^‡§, Tom Alvestad^|, Lynda Avery^¶, James A. Blake^#, Olga Biriukova^¤, Markus Böggemann^«, Polina Borisova^», Nataliya Budaeva^|», Ingo Burghardt^‡, Maria Capa^˄, Magdalena N. Georgieva^˅, Christopher J. Glasby^¤, Pan-Wen Hsueh^¦, Pat Hutchings^‡§, Naoto Jimi^ˀ, Jon A. Kongsrud^|, Joachim Langeneck^ˁ, Karin Meißner^₵, Anna Murray^‡, Mark Nikolic^ℓ, Hannelore Paxton^‡§, Dino Ramos^˅, Anja Schulze^₰, Robert Sobczyk^₱, Charlotte Watson^¤, Helena Wiklund^˅₳, Robin S. Wilson^ℓ, Anna Zhadan^₴, Jinghuai Zhang^₣

‡ Australian Museum Research Institute, Sydney, Australia

§ Macquarie University, Sydney, Australia

| University of Bergen, Bergen, Norway

¶ Museum Victoria, Melbourne, Australia

# Aquatic Research & Consulting, Duxbury, United States of America

¤ Museum and Art Gallery of the Northern Territory, Darwin, Australia

« University of Vechta, Vechta, Germany

» P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia

˄ University of the Balearic Islands, Palma, Spain

˅ Natural History Museum, London, United Kingdom

¦ National Chung Hsing University, Taichung City, Taiwan

ˀ National Institute of Polar Research, Tachikawa, Japan

ˁ University of Pisa, Pisa, Italy

₵ Forschungsinstitut Senckenberg, Hamburg, Germany

ℓ Museums Victoria, Melbourne, Australia

₰ Texas A&M University, Galveston, United States of America

₱ University of Lodz, Lodz, Poland

₳ University of Gothenburg, Gothenburg, Sweden

₴ Lomonosov Moscow State University, Moscow, Russia

₣ South China Sea Environmental Monitoring Centre, State Oceanic Administration, Guangzhou, China

Corresponding author: Laetitia M. Gunton ( laetitia.gunton@austmus.gov.au )

Academic editor: Greg Rouse

© 2021 Laetitia M. Gunton, Elena K. Kupriyanova, Tom Alvestad, Lynda Avery, James A. Blake, Olga Biriukova, Markus Böggemann, Polina Borisova, Nataliya Budaeva, Ingo Burghardt, Maria Capa, Magdalena N. Georgieva, Christopher J. Glasby, Pan-Wen Hsueh, Pat Hutchings, Naoto Jimi, Jon A. Kongsrud, Joachim Langeneck, Karin Meißner, Anna Murray, Mark Nikolic, Hannelore Paxton, Dino Ramos, Anja Schulze, Robert Sobczyk, Charlotte Watson, Helena Wiklund, Robin S. Wilson, Anna Zhadan, Jinghuai Zhang.

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Gunton LM, Kupriyanova EK, Alvestad T, Avery L, Blake JA, Biriukova O, Böggemann M, Borisova P, Budaeva N, Burghardt I, Capa M, Georgieva MN, Glasby CJ, Hsueh P-W, Hutchings P, Jimi N, Kongsrud JA, Langeneck J, Meißner K, Murray A, Nikolic M, Paxton H, Ramos D, Schulze A, Sobczyk R, Watson C, Wiklund H, Wilson RS, Zhadan A, Zhang J (2021) Annelids of the eastern Australian abyss collected by the 2017 RV ‘Investigator’ voyage. ZooKeys 1020: 1-198. https://doi.org/10.3897/zookeys.1020.57921

ZooBank: urn:lsid:zoobank.org:pub:CC23B8CE-8C8E-473C-BD8C-44E74252A33D

Abstract

In Australia, the deep-water (bathyal and abyssal) benthic invertebrate fauna is poorly known in comparison with that of shallow (subtidal and shelf) habitats. Benthic fauna from the deep eastern Australian margin was sampled systematically for the first time during 2017 RV ‘Investigator’ voyage ‘Sampling the Abyss’. Box core, Brenke sledge, and beam trawl samples were collected at one-degree intervals from Tasmania, 42°S, to southern Queensland, 24°S, from 900 to 4800 m depth. Annelids collected were identified by taxonomic experts on individual families around the world. A complete list of all identified species is presented, accompanied with brief morphological diagnoses, taxonomic remarks, and colour images. A total of more than 6000 annelid specimens consisting of 50 families (47 Polychaeta, one Echiura, two Sipuncula) and 214 species were recovered. Twenty-seven species were given valid names, 45 were assigned the qualifier cf., 87 the qualifier sp., and 55 species were considered new to science. Geographical ranges of 16 morphospecies extended along the eastern Australian margin to the Great Australian Bight, South Australia; however, these ranges need to be confirmed with genetic data. This work providing critical baseline biodiversity data on an important group of benthic invertebrates from a virtually unknown region of the world’s ocean will act as a springboard for future taxonomic and biogeographic studies in the area.

Keywords

Biodiversity, Biogeography, deep sea, Echiura, lower-bathyal, Marine Parks, Polychaeta, Sipuncula, Tasman Sea

Introduction

The deep sea (> 200 m depth) is the least explored environment on our planet, where most species have not been sampled and remain undiscovered. The vast sediments of the deep sea cover approximately 65% of the Earth’s surface, and it is a unique environment characterised by darkness, low temperatures and low currents, high hydrostatic pressure, and well oxygenated oligotrophic waters (Gage and Tyler 1991). Unfortunately, the deep-sea environment is increasingly affected by anthropogenic impact such as overfishing (Bailey et al. 2009), oil and gas exploration and extraction, waste disposal and pollution (reviewed in Glover and Smith 2003; Ramirez-Llodra et al. 2011; Mengerink et al. 2014). Thus, accurate baseline data on species is essential for monitoring, protecting and managing biological communities.

In Australia, the abyssal plain (3000 to 6000 m depth) and deep ocean floor covers ~ 2.8 million km², or 30% of Australia’s marine territory (Heap and Harris 2008). The abyssal plain is a major part of Australia’s ~ 10 million km² Exclusive Economic Zone (EEZ), which is the third largest EEZ in the world. The deeper areas of Australia’s EEZ including Marine Parks (MP) have been extremely poorly sampled for fauna compared with the intertidal and shallow sublittoral waters (Ponder et al. 2002). While the subtidal and shelf fauna of eastern Australian coasts is the best studied in the continent due to research capacity and high population density in the area, the deep-water fauna beyond the narrow shelf was virtually unknown. Until recently more was known about deep-water benthic fauna off the sparsely populated western Australian coast (McEnnulty et al. 2011; Poore et al. 2015) than off eastern Australia.

Earlier sampling of the eastern Australian abyss was performed as part of research expeditions to the area organised by non-Australian institutions. These include expeditions dating back to the H.M.S. ‘Challenger’ expedition (1874, the UK), the ‘Galathea’ expedition (1951–52, Denmark), the research vessel (RV) ‘Dmitry Mendeleev’ (1975–76, USSR), and RV ‘Tangaroa’ voyages (1982, New Zealand) (reviewed in O’Hara 2019). This is because Australia lacked the required capacity to consistently collect biological material from the seafloor at great depths. Surveys of the lower shelf to abyssal depths (200–3150 m) off the south-eastern margin of Australia were conducted from the Australian oceanographic research vessel (ORV) ‘Franklin’ in 1986 and 1988 (Poore et al. 1994) along four transects, but the report focused on isopods only.

A new era for deep-sea biological exploration in Australia began in 2014 with the launch of the Marine National Facility’s RV ‘Investigator’, the first Australian research vessel equipped to routinely perform biological sampling to depths of 5000 m. The systematic biological study of abyssal depths in Australia on board RV ‘Investigator’ started with the Great Australian Bight (GAB) Research Program. This programme conducted six surveys off the southern coastline of Australia during 2013, 2015, and 2017, sampling epifauna from soft substrates, rocky outcrops in canyons and seamounts from depths of 200–5000 m (MacIntosh et al. 2018).

The significant gap in knowledge about the eastern abyss was addressed by the 2017 ‘Sampling the Abyss’ research project supported by the Marine National Facility, the Commonwealth Scientific and Industrial Research Organisation (CSIRO) and Museums Victoria. This was the first dedicated deep-sea cruise to perform a systematic biological survey along the eastern Australian coast, from Tasmania to southern Queensland covering the lower bathyal (~ 2500 m) and abyssal environments (~ 4000 m). This was also the first expedition to collect biological samples from the deeper parts of the eastern network of Australian Marine Parks (O’Hara 2017). The present study focuses on the biodiversity of annelids collected during this voyage.

Annelids occur in all marine environments and they are typically a dominant macrofauna (> 300 µm) taxon in terms of abundance and species diversity in deep-sea soft sediments (Herring 2010; Rex and Etter 2010). They display a diverse range of life history strategies and feeding modes (Jumars et al. 2015) and play important roles in processing and burying organic matter, recycling nutrients and bioturbation of seafloor sediments (Hutchings 1998). More than 2000 annelid species are known from Australia (http://www.ala.org.au), yet only 15 species from six families had been described from below 1000 m and three species from below 2500 m (Table 1), the shallower depth limits of the present study. Of 158,400 records of annelids in Australia 770 are from below 1000 m and only 99 from below 2500 m depth (http://www.ala.org.au), suggesting deep-water biodiversity is severely underestimated.

This study reports an illustrated and annotated preliminary species-level checklist of the annelid fauna collected during the 2017 ‘Sampling the Abyss’ survey along with species diversity and distribution data. Morphospecies are compared with those collected from the GAB sampling programme where possible.

Table 1.

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Annelid species described below 1000 m in Australian waters (roughly corresponding to Exclusive Economic Zone, 12 nautical miles from the coast). Bold font indicates species from eastern Australian margin.

Family	Species	Depth (m)	Type Locality
Polynoidae	Lepidasthenia australiensis (Augener, 1927)	1000	Off eastern Victoria
Sabellidae	Potaspina australiensis Capa, 2007	1000	South of Point Hicks, Victoria
Polynoidae	Brychionoe karenae Hanley & Burke, 1991	1100	Cascade Plateau off Tasmania
Onuphidae	Paradiopatra imajimai Paxton & Budaeva, 2013	1277	Off eastern Victoria
Polynoidae	Lagisca torbeni Kirkegaard, 1995	1320–1340	Great Australian Bight, south of Adelaide
Polynoidae	Harmothoe australis Kirkegaard, 1995	1340	Great Australian Bight, south of Adelaide
Spionidae	Laonice pectinata Greaves, Meißner & Wilson, 2011	1440	Indian Ocean, west of Perth
Onuphidae	Paradiopatra spinosa Paxton & Budaeva, 2013	1600	Bass Canyon
Polynoidae	Eunoe ivantsovi Averincev, 1978	1640	Lord Howe Island Rise
Polynoidae	Eunoe papillaris Averincev, 1978	1800	Off southwestern Tasmania
Nephtyidae	Aglaophamus profundus Rainer & Hutchings, 1977	2195	Off northeastern Tasmania
Polynoidae	Parapolyeunoa flynni (Benham, 1921)	2379	Off Maria Island, Tasmania
Fauveliopsidae	Fauveliopsis challengeriae McIntosh, 1922	3566	South Indian Ocean, midway between Australia and Antarctica
Polynoidae	Eunoe abyssorum McIntosh, 1885	4755	South of Australia
Polynoidae	Polynoe ascidioides McIntosh, 1885 (now considered a nomen dubium)	4755	South of Australia

Materials and methods

Sampling area

The eastern Australian continental shelf is relatively narrow compared with the rest of the continent. The shelf break occurs ~ 15 km from the coast and the foot-of-slope and beginning of the abyssal plain can be as close as 60 km from the coast (Heap and Harris 2008). The eastern margin contains a range of geomorphological features including plateaus, basins, terraces, deeps/holes, and submarine canyons seamounts/guyots (Heap and Harris 2008).

The East Australian Current (EAC) is an important shallow water current carrying ~ 22–27 Sverdrups from north to south along the east coast of Australia. This counter-clockwise southern Pacific gyre circulates shallow water from the Coral Sea along the continental margin until 32–35°S before heading eastward to New Zealand. Part of the EAC is deflected offshore ~ 30°S along the Tasman Front, this divides the warm waters of the Coral Sea and the cooler waters of the Tasman Sea (Rintoul et al. 2017). Deeper currents (> 2000 m) have not been directly measured but have been inferred from sediment deposition and erosion patterns; these currents are thought to be weak with a western boundary undercurrent flowing northwards along the eastern Australian margin and an eastern boundary counter-flow along the eastern margin of the Tasman Sea (Jenkins 1984).

Field collection and processing

Biological samples were collected from 13 sites at one-degree intervals of latitude from 42°S to 24°S along the east coast of Australia from Tasmania to Southern Queensland (Fig. 1, Table 2) on the RV ‘Investigator’ (voyage code IN2017_V03), from 15 May to 16 June 2017. Seven Marine Parks were included (Freycinet MP, Flinders MP, East Gippsland MP, Jervis MP, Hunter MP, Central Eastern MP, and Coral Sea MP). Benthic sampling was conducted at lower bathyal (~ 2500 m) and abyssal (~ 4000 m) depths, with some (seven operations) comparative samples taken at shallower mid-bathyal depths (~ 1000 m). The three types of sampling gear used were beam trawl (35 operations, ops.), Brenke sledge (28 ops.), and box core (8 ops.).

Figure 1.

Map of sampling sites from expedition IN2017_V03 along eastern Australia. Blue dots represent beam trawl, yellow dots Brenke sledge, red dots box core sampling sites, green polygons represent areas of Australian Marine Parks.

Table 2.

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Sample Sites. Beam trawl, Brenke sledge, and box core deployments on RV ‘Investigator’ cruise IN2017_V03 from the Australian eastern lower bathyal and abyssal environment. Abbreviations: Op., operation, BT, CSIRO 4-metre beam trawl; BS, Brenke sledge; BC, box core; *, unfired ; **, aborted.

Op.	Location	Gear	Date	Start latitude and longitude	End latitude and longitude	Trawling distance (km)	Start depth (m)	End depth (m)
004	Freycinet MP	BT	18/05/17	-41.731, 149.120	-41.791, 149.156	7.3	2820	2751
005	Freycinet MP	BS	18/05/17	-41.730, 149.135	-41.753, 149.147	2.8	2789	2779
006	Freycinet MP	BT	18/05/17	-41.626, 149.552	-41.689, 149.584	7.5	4022	4052
007	Freycinet MP	BC	18/05/17	-41.647, 149.570			4030
008	Freycinet MP	BC	19/05/17	-41.647, 149.569			4012
009	Freycinet MP	BS	19/05/17	-41.626, 149.560	-41.662, 149.574	4.2	4021	4035
011	Freycinet MP	BC	19/05/17	-41.721, 149.125			2793
013	Flinders MP	BT	20/05/17	-40.386, 148.928	-40.383, 148.951	2.0	932	1151
014	Flinders MP	BT	20/05/17	-40.464, 149.102	-40.461, 149.147	3.8	2298	2486
015	Flinders MP	BT	20/05/17	-40.473, 149.397	-40.464, 149.426	2.6	4114	4139
016	Flinders MP	BS	21/05/17	-40.463, 149.415	-40.461, 149.364	4.3	4129	4131
017	Flinders MP	BC	21/05/17	-40.460, 149.109		s	2331
022	Bass Strait	BT	22/05/17	-39.462, 149.276	-39.465, 149.242	2.9	2760	2692
023	Bass Strait	BS	22/05/17	-39.462, 149.277	-39.465, 149.246	2.7	2774	2694
027	Bass Strait	BC	22/05/17	-39.462, 149.271			2741
028	Bass Strait	BC	22/05/17	-39.500, 149.535			4147
030	Bass Strait	BT	23/05/17	-39.552, 149.553	-39.496, 149.598	7.3	4197	4133
031	Bass Strait	BS	23/05/17	-39.422, 149.604	-39.391, 149.597	3.5	4150	4170
032	East Gippsland MP	BT	24/05/17	-38.479, 150.185	-38.453, 150.186	2.9	3850	3853
033	East Gippsland MP	BS	24/05/17	-38.521, 150.213	-38.498, 150.207	2.6	4107	4064
035	East Gippsland MP	BT	25/05/17	-37.792, 150.382	-37.818, 150.353	3.9	2338	2581
040	East Gippsland MP	BS	25/05/17	-37.815, 150.373	-37.818, 150.356	1.5	2746	2600
041	off Bermagui	BT	26/05/17	-36.418, 150.800			3980
042	off Bermagui	BS	26/05/17	-36.385, 150.863	-36.434, 150.863	5.4	4744	4716
043	off Bermagui	BT	27/05/17	-36.351, 150.914	-36.384, 150.913	3.7	4800	4800
044	off Bermagui	BT	27/05/17	-36.355, 150.644	-36.315, 150.651	4.5	2821	2687
045	off Bermagui	BS	27/05/17	-36.360, 150.644	-36.323, 150.650	4.1	2835	2739
046	off Bermagui	BC	27/05/17	-36.284, 150.658			2643
053	Jervis MP	BT	28/05/17	-35.114, 151.469	-35.084, 151.441	4.2	3952	4011
054	Jervis MP	BS	28/05/17	-35.117, 151.473	-35.099, 151.455	2.6	4026	3881
055	Jervis MP	BS	28/05/17	-35.335, 151.259	-35.334, 151.219	3.6	2667	2665
056	Jervis MP	BT	29/05/17	-35.333, 151.258	-35.332, 151.214	4.0	2650	2636
57*	Jervis MP	BC
065	off Newcastle	BT	30/05/17	-33.441, 152.702	-33.435, 152.665	3.5	4280	4173
066	off Newcastle	BS	30/05/17	-33.448, 152.733	-33.437, 152.674	5.6	4378	4195
067	off Newcastle	BT	31/05/17	-32.985, 152.952	-33.015, 152.913	4.9	2704	2902
068	off Newcastle	BS	31/05/17	-32.993, 152.957	-33.023, 152.943	3.6	2745	2963
069	Hunter MP	BT	03/06/17	-32.479, 152.994	-32.507, 152.991	3.1	1006	1036
070	Hunter MP	BT	03/06/17	-32.575, 153.162	-32.632, 153.142	6.6	2595	2474
076	Hunter MP	BS	03/06/17	-32.577, 153.161	-32.613, 153.149	4.2	2534	2480
078	Hunter MP	BT	04/06/17	-32.138, 153.527	-32.182, 153.524	4.9	3980	4029
079	Hunter MP	BS	04/06/17	-32.131, 153.527	-32.163, 153.524	3.6		4031
080	Central Eastern MP	BT	05/06/17	-30.099, 153.596	-30.128, 153.571	4.0	1257	1194
086	Central Eastern MP	BT	05/06/17	-30.098, 153.899	-30.119, 153.875	3.3	2429	2518
087	Central Eastern MP	BS	06/06/17	-30.113, 153.898	-30.116, 153.867	3.0	2634	2324
088	Central Eastern MP	BT	06/06/17	-30.264, 153.870	-30.287, 153.830	4.6	4481	4401
089	Central Eastern MP	BS	06/06/17	-30.263, 153.859	-30.289, 153.844	3.2	4436	4414
090	off Byron Bay	BT	07/06/17	-28.677, 154.203	-28.709, 154.190	3.8	2587	2562
096	off Byron Bay	BS	07/06/17	-28.678, 154.204	-28.716, 154.189	4.5	2591	2566
097	off Byron Bay	BT	08/06/17	-28.355, 154.636	-28.414, 154.615	6.9	3762	3803
098	off Byron Bay	BS	08/06/17	-28.371, 154.647	-28.389, 154.612	4.0	3811	3754
099	off Byron Bay	BT	09/06/17	-28.371, 154.649	-28.388, 154.617	3.7	3825	3754
100	off Byron Bay	BT	09/06/17	-28.054, 154.083	-28.097, 154.081	4.8	999	1013
101	off Moreton Bay	BT	09/06/17	-26.946, 153.945	-26.971, 153.951	2.8	2520	2576
102	off Moreton Bay	BT	10/06/17	-27.008, 154.223	-27.049, 154.224	4.6	4274	4264
103	off Moreton Bay	BS	10/06/17	-27.000, 154.223	-27.061, 154.223	6.8	4260	4280
104	off Moreton Bay	BT	10/06/17	-26.961, 153.848	-26.991, 153.847	3.3	1071	1138
109	off Fraser Island	BT	11/06/17	-25.221, 154.164	-25.253, 154.192	4.5	4006	4005
110	off Fraser Island	BS	11/06/17	-25.220, 154.160	-25.261, 154.200	6.1	4005	4010
115	off Fraser Island	BT	11/06/17	-25.325, 154.068	-25.351, 154.076	3.0	2350	2342
118**	off Fraser Island	BS
119	off Fraser Island	BS	12/06/17	-25.206, 153.991	-25.178, 153.979	3.3	2247	2369
121	Coral Sea MP	BT	13/06/17	-23.587, 154.194	-23.617, 154.195	3.3	1013	1093
122	Coral Sea MP	BT	13/06/17	-23.751, 154.639	-23.773, 154.616	3.4	2369	2329
123	Coral Sea MP	BS	13/06/17	-23.749, 154.641	-23.774, 154.617	3.7	2271	2339
128	Coral Sea MP	BT	13/06/17	-23.631, 154.660	-23.659, 154.644	3.5	1770	1761
131	Coral Sea MP	BS	14/06/17	-23.748, 154.643	-23.778, 154.613	4.5	2297	2358
132	Coral Sea MP	BS	14/06/17	-23.756, 154.568	-23.780, 154.540	3.9	2181	2132
134	Coral Sea MP	BS	14/06/17	-23.750, 154.572	-23.774, 154.546	3.8	2093	2156
135	Coral Sea MP	BT	15/06/17	-24.352, 154.291	-24.384, 154.325	5.0	3968	4034

The CSIRO 4 m wide by 0.5 m high beam trawl used to collect megafaunal invertebrates had a net mesh size of 12 mm in the forward section, and 10 mm in the cod end (Lewis 2010). From the time samples were brought aboard the ship to preservation, a ‘cold-chain’ method (Glover et al. 2016) was employed to ensure specimens could be used for future morphological and DNA taxonomy. Substrate and larger specimens from the beam trawl catch were transferred to the wet laboratory on board in containers filled with chilled seawater (5 °C) and material was roughly sorted on ice made from seawater into higher taxonomic categories. Larger clumps of substrate were broken with a hammer and smaller animals were picked from the pieces. Selected specimens were photographed. The majority of the catch was preserved in 95% ethanol, the remainder in 10% buffered formalin, while selected specimens were fixed in RNAlater. When time allowed, for ~ 20% of the catch, tissue samples from selected specimens were taken and fixed in ethanol, while the voucher specimens were fixed in formalin. Larger samples containing numerous annelid tubes were split, half was fixed in ethanol and half in formalin. Operation 100 included a dead pilot whale skull and vertebrae, annelids were picked off the bones and preserved in 95% ethanol.

The Brenke sledge (mesh size 1 mm) was used to collect microbenthic infauna living near the sediment-water interface and more mobile epibenthic fauna (Brenke 2005). Both cod-ends of the Brenke sledge nets were emptied into chilled seawater and visible animals were picked out. The remainder of the sample was elutriated with chilled seawater to separate small animals from sediment, and gently sieved using a 300 µm sieve. Sediment residue was fixed in 95% ethanol and the remaining specimens were hand-picked from residues and sorted under microscopes in the dry lab on board. Selected preserved specimens were also photographed on board. The box core (Hessler and Jumars 1974) was used to collect infaunal invertebrates. Box core sampling was less successful than the trawl and sledge sampling: of five deployments, one deployment resulted in a full sample, the other four resulted in only partial samples or none at all. For successful samples, the top 2 cm sediment layer of the core was elutriated in chilled seawater, sieved using a 300 µm mesh sieve and treated as samples collected by Brenke sledge.

Prior to fixation, all specimens were weighed and registered on board and assigned labels with operation (op) and accession numbers (acc).

Annelid specimens collected during the voyage were shipped to the Australian Museum, Sydney (AM), Museums Victoria, Melbourne (MV), and the Natural History Museum, London (NHMUK) where they were registered and assigned permanent registration numbers of the respective institutions.

Laboratory identification of annelids

At the respective institutions, annelids fixed in formalin were soaked in water, preserved with 80% ethanol and sorted in 80% ethanol, while ethanol-fixed annelids were sorted in 95% ethanol. Mixed lots of annelids were sorted to families at the AM and MV. Annelid families for which no taxonomic expertise is available in Australia (Acrocirridae, some Ampharetidae, Cirratulidae, Dorvilleidae, Flabelligeridae, Glyceridae, Goniadidae, Lumbrineridae, Maldanidae, some Melinnidae, Opheliidae, Orbiniidae, Paraonidae, Scalibregmatidae, Sphaerodoridae, Spionidae, Sternaspidae, Echiura, and Sipuncula) were sent internationally to taxonomic authorities for species-level identification.

All beam trawl specimens were identified. Brenke sledge and box core material was identified past family level when specimens were large enough (considered adult) and/or complete. Annelids were assigned Latin binomial names where possible or determined in open nomenclature following Sigovini et al. (2016). Species were designated cf. qualifiers to the closest morphological match and not the full species name following the reasons stated in Neal et al. (2020); i) specimens were too damaged or incomplete, ii) the original description was not detailed enough, iii) the described species had a type locality from shallow water or in a different region from Australia (another ocean basin). Some deep-sea species are known to be widespread and in these cases we follow the individual authors of each section as to whether the species is thought to be widespread and thus whether cf. is designated or not. It is important to stress that no formal descriptions of the species are given here, only preliminary identifications and diagnoses (a short written description of the species, which allows that species to be distinguish from other species with which it is likely to be confused). Subsequent taxonomic papers will describe the species and compare them genetically.

The matrix of all annelid species-level abundance and presence data (including beam trawl, box core, and Brenke sledge material) from voyage IN2017_V03 was constructed in MS Excel in standardised Darwin Core format.

Results

Taxonomic overview

Family Acoetidae Kinberg, 1856

A. Murray

This family of scale worms is characterised by the presence of internal ‘spinning’ glands which produce fibres used to construct their tough fibrous permanent tubes. These fibres often appear as golden strings emerging from the notopodia. Acoetidae are active carnivores and predators, and most frequently collected by fishers on baited lines, in shallow to deep waters (1–200 m). There are currently nine valid genera with 58 nominal species worldwide (Read and Fauchald 2020). In Australian waters they have been collected rarely, and usually only single individuals are found, often as fragments, possibly due to their ability to actively avoid grabs and dredges by rapidly withdrawing into their tubes when detecting vibration in benthic sediments. The few Acoetidae specimens recorded from Australia have all been collected from shallow waters, with the deepest species being Polyodontes australiensis (McIntosh, 1885) reported from 120 m off Tasmania (as Eupompe australiensis) by Benham (1915), and a specimen of Euarche sp., reported as Eupanthalis sp., from off Cronulla, New South Wales by Hutchings (2000a) from < 100 m depth. In this study we report one species.

Panthalis sp.

Fig. 2A

Diagnosis

One damaged specimen, with 24 anterior segments measuring 1.2 cm long, 0.6 mm wide. Head region badly damaged, but some features recognisable: low rounded ommatophores without necks and colourless, a single long median antenna attached mid-prostomium, longer than prostomium length; lateral antennae and palps missing, however; tentaculophores with a few chaetae, styles missing; elytra present on segments 2, 4, 5, 7 and alternating segments thereafter, delicate, transparent. All chaetae simple. Acicular neurochaetae starting from chaetiger 3, notochaetae absent from chaetiger 4 and on all parapodia thereafter. Notopodia with notoaciculum and spinning glands internally, golden ‘spinning’ fibres emergent from the inner surface of the notopodial bract. Superior group of neurochaetae from chaetiger 9 onwards, of two types: long, with plumose (brush) tips, and shorter chaetae with few whorls of short widely spaced hairs along shafts; middle group of neurochaetae stout, acicular chaetae with hairy aristate tips; inferior group of neurochaetae curved, lanceolate, with many transverse rows of overlapping spines along shaft.

Remarks

This specimen possesses brush-tipped neurochaetae typical of the genera Acoetes and Panthalis, but lacks notochaetae in all middle segments, a feature which distinguishes it as a species of Panthalis. The genus Panthalis has not yet been reported from Australian waters; however, specimens have been collected previously from deep water in the Arafura Sea off Western Australia and Northern Territory (Murray and Hutchings in prep.).

Records

1 specimen. Suppl. material 1: op. 104 (AM).

Figure 2.

Records

6 specimens. Suppl. material 1: ops. 23, 119 (AM).

Flabelligena spp.

Records

6 specimens. Suppl. material 1: ops. 16, 54, 98, 110 (NHMUK).

Swima sp.

Records

2 specimens. Suppl. material 1: op. 134 (NHMUK).

Acrocirridae gen. sp. 1

Incomplete, length ~ 4 mm, width 0.4 mm, 12 chaetigers, 2–3 notochaetae, 2–3 composite neurochaetae. Body papillae short, without sediment particles. Large ventral papillae absent.

Records

1 specimen. Suppl. material 1: op. 79 (AM).

Acrocirridae gen. spp.

Remarks

Samples were identified to family level only or individuals were too fragmented for further analysis.

Records

16 specimens. Suppl. material 1: ops. 31, 54, 55, 56, 76, 87, 96, 98, 119, 134 (AM).

Family Ampharetidae Malmgren, 1866

T. Alvestad, L. M. Gunton

Ampharetidae are tubicolous annelids, with a body divided into a distinct thorax and abdomen, unlike the closely related Terebellidae, species of Ampharetidae are able to fully retract buccal tentacles into the mouth. The family Ampharetidae is composed of 64 accepted genera and > 300 species (Ebbe and Purschke 2021). Thirty-two of these genera are monospecific making the taxonomy complex (Eilertsen et al. 2017). Ampharetids are found from intertidal to abyssal depths (Aguirrezabalaga and Parapar 2014; Bonifácio et al. 2015). Deep-water ampharetids are found in high abundance on abyssal plains (Böggemann 2009) and chemosynthesis-based environments such as hydrothermal vents and cold seeps (Reuscher et al. 2009; Eilertsen et al. 2017). The ampharetid fauna of Australia has been poorly studied. To date, five genera (Amphicteis, Auchenoplax, Phyllamphicteis, Pseudoamphicteis, and Neosabellides) and nine species (Day and Hutchings 1979; Hutchings and Rainer 1979; Hartmann-Schröder 1981; Alvestad and Budaeva 2015) have been recorded from Australian waters. These Australian records are primarily from shallow waters (< 100 m); however, ampharetids are known to be well-represented in deep-sea benthic samples, indicating that a high number of Australian ampharetid species are yet to be described. In this study > 300 ampharetid specimens belonging to more than six species were recovered from the Australian lower bathyal and abyssal environment, at least four are new to science.

Amage sp. nov. 1

Fig. 4A

Diagnosis

Length 12 mm, width 4 mm. Body short, thick, with a short abdomen. Prostomium complex; central part drawn out into two lateral horns, lateral parts form large lobes while front part forms a ‘lip’. No glandular ridges or eyes. Approximately three pairs of branchiae in a transverse line in two widely separate groups. No paleae. Fourteen thoracic segments with notopodia with chaetae. First three pairs of notopodia and chaetae small. Thoracic uncini from segment VI. Eleven thoracic uncinigers. Nine abdominal uncinigers. Abdomen with rudimentary notopodia. Pygidium without lateral cirri.

Records

1 specimen. Suppl. material 1: op. 53 (AM).

Figure 4.

Ampharetidae and Melinnidae. Ampharetidae A Amage sp. nov. 1 B Anobothrus sp. nov. 1, anterior end methyl blue staining C Anobothrus sp. nov. 2 anterior end methyl blue staining. Melinnidae D Melinna cf. armandi and tube (AM W.50354), d dorsal hook. Ampharetidae E Amphicteis sp. (AM W. 50429). Melinnidae F Melinnopsis gardelli anterior dorsal view (AM W.50735) G Melinnopsis sp. nov., lateral view (AM W.50397). Scale bars: 2 mm (A); 1 mm (B, C); 2.5 mm (D); 1 cm (E, G); 1 mm (F).

2 specimens. Suppl. material 1: op. 56 (AM).

Jugamphicteis galatheae Holthe, 2000

Diagnosis

Length 25–40 mm, width 2–3 mm. Prostomium with four curved nuchal arches. Body long tapering towards pygidium. Four pairs of branchiae. Paleae present, long golden extend past rim of prostomium. First abdominal segment with dorsal fan with large median notch.

Remarks

The holotype of Jugamphicteis galatheae was collected from Kermadec Trench in the South Pacific Ocean ~ 4500 m; however, paratypes were recovered from both the Kermadec Trench and off the east coast of South Africa between Cape Town and Durban ~ 5000 m. The species is reported to have a wide distribution, which may indicate a species complex. Due to the matching morphology and close proximity of the specimens from this study to collection location of the holotype, we assign the name Jugamphicteis galatheae.

Records

40 specimens. Suppl. material 1: ops. 6, 15, 30, 32, 43, 53, 65, 86, 121 (AM).

Ampharetidae gen. spp.

Remarks

Beam trawl specimens were incomplete which does not allow further identification, while Brenke sledge samples were identified to family level.

Records

260 specimens. Suppl. material 1: ops. 9, 16, 22, 23, 30, 31, 33, 40, 42, 43, 45, 46, 54, 55, 56, 65, 76, 79, 88, 89, 90, 96, 98, 100, 101, 103, 110, 119, 123, 134 (AM). 7 specimens. Suppl. material 1: ops. 16, 31 (NHMUK).

Family Amphinomidae Lamarck, 1818

L. M. Gunton, D. Ramos, R. S. Wilson

The family Amphinomidae is characterised by simple calcareous chaetae, in some species these chaetae are very fragile breaking off if touched and causing a burning sensation giving them the common name, ‘fireworms’. The family is divided in to two subfamilies, Archinominae Kudenov, 1991 and Amphinominae Lamarck, 1818 based on the presence of accessory dorsal cirrus in the former and absence in the latter. Currently, there are 23 genera containing 148 valid species (Read and Fauchald 2020). Amphinomids occur worldwide from intertidal to abyssal depths, they are predominantly associated with shallow reefs, rocky and soft bottoms of intertidal and continental shelf habitats, comparatively few have been recorded from the deep sea (Rouse and Pleijel 2001), but some are associated with deep-sea chemosynthetic environments (Borda et al. 2012, 2013). There are > 1700 records of amphinomids in Australian waters but only nine species from seven genera have been listed (http://www.ala.org.au). In this study, at least four species were recovered, three may be new to science.

Bathychloeia cf. sibogae Horst, 1910

Diagnosis

Body short, ovate, ~ 8 mm in length. 17–18 chaetigers bearing long (3–44 mm) furcate chaetae. Body pale colour, pair of purple spots dorsal on chaetiger 6, visible under skin. Dark blue-black colouration visible under skin on chaetigers 10–13, dorsal and ventral. Caruncle lobed, extending to chaetiger 3. Branchiae branched, only found on chaetiger 5. Parapodia short, but neuro-and notochaetae well separated, neurochaetae lateral. Notochaetae dorsal (remaining tuft on chaetiger 7). Parapodial cirri on all (?) chaetigers, longer on final five. Chaetae long, bifurcate. No serrations or harpoon chaetae. Neurochaetae shorter than notochaetae. Faint membrane/covering visible over the furcate tips of some chaetae. Pygidium with thick anal cirrus, may be part of a pair.

Remarks

The type locality of Bathychloeia sibogae is in the Banda Sea, Malay Archipelago 1158 m depth. Böggemann (2009) redescribed the species using the type material, material from the Canaries (~ 2800 m) and material from the abyssal SE Atlantic (~5000 m). The species is also recorded from 12 stations (138–2074 m) in the GAB (MacIntosh et al. 2018: additional file 2). Due to the species broad distribution it is highly likely a species complex and thus we assign the name Bathychloeia cf. sibogae.

Records

4 specimens: Suppl. material 1: ops. 96, 102, 103 (AM). 2 specimens Suppl. material 1: op. 110 (NHMUK).

Linopherus sp. 1

Fig. 5A, B

Diagnosis

Prostomium divided into two. Posterior portion pentagonal with medial antennae on posterior edge, flanked laterally by the first chaetiger. Anterior section round with antennae and palps reduced to small bumps anterolaterally and laterally respectively. Body small, slightly wider anteriorly and tapering posteriorly. Eyes absent. First chaetiger reduced, not continuous dorsally. Papilliform notopodial postchaetal lobe present throughout. Bipinnate branchiae present on chaetigers 3–5.

Remarks

Linopherus sp. 1 differs from a second species of Linopherus known from the GAB (MacIntosh et al. 2018: additional file 2) in having branchiae first present on chaetiger 3 rather than chaetiger 4 in GAB specimens.

Records

1 specimen. Suppl. material 1: op. 100 (NHMUK).

Remarks

Specimens range from 0.2 mm to 7 cm in length. Some individuals are badly damaged, with chaetae missing, and other intact specimens show some morphological variability from the original description by Hutchings and McRae (1993) in the longer palps, and some with fewer than 13 pairs of elytra. However, they most resemble L. yarramba, but share some similarities with L. producta Grube, 1877 (which is also considered to be species complex by Hutchings and McRae 1993), such as tuberculate shafts of the harpoon chaetae, but from which it can be distinguished by the fewer pairs of elytra and the absence of nuchal flaps on the prostomium. Widespread around Australia in depths of 60–3950 m, including records from the GAB (MacIntosh et al. 2018: additional file 2).

Records

354 specimens. Suppl. material 1: ops. 4, 13, 15, 22, 23, 30, 31, 32, 33, 35, 42, 43, 44, 53, 56, 65, 70, 86, 115 (AM). 4 specimens. Suppl. material 1: op. 4 (MV).

Laetmonice cf. producta sensu Hutchings & McRae, 1993

Diagnosis

Large-bodied specimen, body shape elongate, > twice as long as maximum width. Dorsal felt of fine notochaetae absent, 18 pairs elytra with purple colouration on inner halves. Prostomium with pair of large ocular peduncles, without eye pigment; facial tubercle well-developed, with long conical papillae; small nuchal flaps present. Palps extending to segment 11, margins finely papillate. Median antenna with ceratophore half the length of the prostomium; antennal ceratostyle longer than prostomium, slender, clavate-tipped, 3 × length of prostomium. Notochaetae of three kinds: ~ 15 smooth, golden, unidentate acicular chaetae; ~ ten long, stout, yellow brown harpoon-like chaetae with 3–5 recurved fangs below tips, shafts smooth or tuberculate; tuft of short, fine mud-covered capillary chaetae ventrally. Neurochaetae in two tiers: superior tier of yellow acicular chaetae with basal spur and subdistal fringe of long hairs and bare tips, inferior tier with numerous golden bipinnate neurochaetae. Ventrum covered with small papillae.

Remarks

This specimen agrees well with the description of other Australian specimens assigned to Laetmonice producta Grube, 1877 by Hutchings & McRae (1993), who stated that the species displayed much morphological variability and had a broad distribution. Grube originally described L. producta from the area of the Kerguelen Islands in the Southern Ocean. McIntosh (1885, 1900) described several “varieties” of L. producta, ranging from the Azores to Antarctic waters, most of which were subsequently raised to species level by Chamberlin (1919). However, Hartman (1965) suggested that the variety represented by Grube’s original species (now known as the subspecies L. producta producta, see Read and Fauchald 2020) was restricted in distribution to the area of Kerguelen Islands, the South Georgia Islands and the Antarctic Peninsula. The records from southeastern Australian waters were assigned to L. producta by Hutchings and McRae (1993) because they were unable to examine the type specimens of McIntosh’s “varieties”, and they suggest that it may possibly belong to another new species. The specimen examined herein differs from L. producta producta Grube, 1877 by the number of elytra (18, not 20 pairs), the lack of eye pigment, smaller nuchal flaps, and the shafts of the harpoon notochaetae which may be smooth or finely granulated. Live specimens are usually pale, with a longitudinal purple stripe mid-dorsally.

Records

1 specimen. Suppl. material 1: op. 14 (AM).

Family Capitellidae Grube, 1862

L. M. Gunton

Capitellids resemble terrestrial earthworms due to their simple cylindrical body shape, lack of head appendages and often reduced parapodia. The family contains 44 genera and ~ 186 species (Magalhães and Blake 2019). Capitellids are common and widespread annelids that occur in every marine habitat from intertidal to hadal depths. In Australia, Capitellidae fauna comprises of at least 37 species in 18 genera (Hutchings 2000b). Unfortunately, the capitellids collected from the present study were damaged. Complete or nearly complete specimens are needed for generic diagnosis, furthermore, no taxonomic expertise in the family was available. Two genera (Capitella and Notomastus) are reported here.

Capitella spp.

Phyllochaetopterus spp.

Records

7 specimens. Suppl. material 1: ops. 6, 41, 53, 78 (AM).

Chaetopteridae gen. spp.

Remarks

The specimens were too fragmented to be identified further.

Records

2 specimens. Suppl. material 1: ops. 88, 134 (AM). Tubes only. Suppl. material 1: ops. 65, 104 (NHMUK).

Family Chrysopetalidae Ehlers, 1864

C. Watson

Chrysopetalids are distinguished by broad notochaetal, leaf-like paleae and/or notochaetal spines in fans covering the dorsum. Chrysopetalidae currently contains 29 genera and ~ 110 species (Watson 2020). There are three subfamilies, paleate Chrysopetalinae, spinous Dysponetinae, and putatively lacking spinous notochaetae or notochaetae Calamyzinae. Abyssal Chrysopetalinae taxa have been described from hydrothermal vents (e.g., Thrausmatos Watson, 2001) and wood falls (e.g., Strepternos Watson Russell, 1991). Dysponetinae taxa have been described from abyssal plains of the East Atlantic (Böggemann 2009). Free-living Calamyzinae taxa occur in seep and whale fall communities, e.g., Micospina auribohnorum Watson et al. 2016 from the East Pacific, and symbiotic Calamyzinae taxa from vent bivalve hosts, e.g., Nautilina calyptogenicola Miura & Laubier, 1989 from the West Pacific. No Calamyzinae taxa have yet been recorded from Australian waters. Chrysopetalinae taxa from southern Australia, shelf to ~1000 m, are currently being described (Watson in prep.). In this abyssal study we report one species, Dysponetus cf. caecus.

Dysponetus cf. caecus (Langerhans, 1880)

Fig. 2B, C

Diagnosis

Prostomium truncate, with three short antennae and two short palps. No eyes. Large mouth cirrus and pair of rod-like stylets. Two pairs of cirriform tentacular cirri on segments 1 and 2; segments one and two fused; dorsal cirri segment 1 very long, notopodia 2 with notochaetae. Mid-body notopodia with two lengths of semi-erect, golden-coloured notochaetal fascicles: short, broad and long, slender, spines with two rows of spinelets; elongate cirrophores and long dorsal styles. Neuropodia with fascicle of very slender falcigerous neurochaetae with very long shafts; ventral cirri longer than neuropodia.

Remarks

Similar to, but differs from abyssal East Atlantic Dysponetus cf. caecus (sensu Böggemann 2009; Watson et al. 2014) in possession of broad short spines with less spinulation. Dysponetus caecus is currently considered a global species complex.

Records

48 specimens. Suppl. material 1: ops. 11, 23, 40, 76, 98, 110, 134 (AM). 2 specimens. Suppl. material 1: op. 100 (NHMUK).

H. Wiklund

Dorvilleids contain some of the smallest described annelids and are the only extant group with ctenognath jaws. The family Dorvilleidae consists of ~ 200 species arranged in 32 genera (Read and Fauchald 2020), with more than a third of the genera containing only one species. The most speciose dorvilleid genus is Ophryotrocha Claparède & Mecznikow, 1869 with ~ 75 described species. The first Ophryotrocha species were described from shallow water, but with advancing deep-sea sampling, now more species are known from the deep sea than shallow water. Worms in the genus are opportunistic feeders and thrive in organically enriched habitats both in deep-sea and shallow waters, for example in polluted harbours, sewer outlets, beneath fish farms and on hydrothermal vents, cold seeps and whale falls. One species of Ophryotrocha has been described from Australian waters, O. shieldsi Paxton & Davey, 2010, that was found in large numbers beneath a shallow fish farm in Tasmania. In this study we report eight species of Ophryotrocha. Some species were far more abundant than others, with Ophryotrocha sp. 2 and Ophryotrocha sp. 3 being the most common. Several of these species are likely new to science.

Remarks. Species were preliminary separated on the basis of the forms of mandibles, shape of head and appendages, and shape of chaetae. The species vary slightly in size, with the smallest species being just 1 mm long (Ophryotrocha sp. 2) and the largest being 3.2 mm long (Ophryotrocha sp. 5).

Ophryotrocha sp. 1

Fig. 7A

Records

55 specimens. Suppl. material 1: op. 100 (NHMUK).

Figure 7.

17 specimens. Suppl. material 1: op. 100 (NHMUK).

Ophryotrocha sp. 7

Records

1 specimen. Suppl. material 1: op. 100 (NHMUK).

Ophryotrocha sp. 8

Records

3 specimens. Suppl. material 1: op. 100 (NHMUK).

Dorvilleidae gen. spp.

Remarks

Specimens from whale fall (op. 100) were too damaged or dried out to be identified. A single specimen provisionally referred to Schistomeringos was recorded from 932 m in the GAB (MacIntosh et al. 2018: additional file 2). Further investigation is needed to understand if any of the GAB species match those found in this study.

Records

75 specimens. Table 1: op. 100 (NHMUK).

Family Eunicidae Berthold, 1827

R. S. Wilson

Along with other members of the order Eunicida, species of Eunicidae possess a ventral muscular pharynx with mineralized or sclerotized jaws. Eunicidae are recognisable by possessing a prostomium with one to three antennae which lack ringed ceratophores. The family consists of 11 extant genera and 453 currently accepted species (Read and Fauchald 2020). They are not dominant members of abyssal benthic communities. Shallow water species from Australia are comparatively well known (Zanol et al. 2020) and historical records include about 32 species from six genera from Australian waters, all from <100 m water depth (http://www.ala.org.au). This study reports two species from depths of ~ 1000–2800 m off the east coast of Australia; neither species is previously known from Australia and both appear to be undescribed.

Eunice sp. nov.

Fig. 8A, B

Diagnosis

No pigmentation on preserved specimens. Prostomium bilobed, slightly notched. Eyes present, behind bases of palps. Prostomial appendages with widest gap separating palps from lateral antennae. Palpostyles, antennal styles, and peristomial cirri with irregular articulations. Peristomial rings distinct dorsally and ventrally but continuous laterally. Maxillae dentition: Mx I left 1, right 1. Mx II left 7, right 6. Mx III left 6. Mx IV left 5, right 9. Mx V left 1, right 1.

Branchiae absent. Lateral black dot between posterior parapodia absent. Dorsal cirri length short, at most as long as two body segments. Dorsal cirri of anterior chaetigers tapering, median chaetigers tapering, posterior chaetigers tapering, smooth, without articulations. Digitiform ventral cirri, basally inflated, commence chaetiger 3.

Pectinate chaetae absent. Compound falcigers present, appendages distally bidentate, hoods without mucros (rounded). Compound spinigers absent. Aciculae dark honey-coloured to black, distally bluntly pointed. Subacicular hooks dark honey-coloured to black, bidentate, distal tooth directed distally, subdistal tooth directed laterally. Subacicular hooks first present from chaetiger 24–27.

Remarks

Although referred to here as ‘Eunice sp.’, the above combination of characters cannot be accommodated in any currently known eunicid genus. The species is here treated as a member of Eunice since that genus remains poorly defined and already contains species of uncertain relationships (Zanol et al. 2020).

Records

4 specimens. Suppl. material 1: ops. 69, 104 (AM).

Figure 8.

Eunicidae A Eunice sp. nov., anterior view (AM W.50148) B Maxillae of Eunice sp. nov. (AM W.50148) C Leodice sp. nov., anterior view (AM W.50146) D Maxillae of Leodice sp. nov. (AM W.50152). Scale bars: 2 mm (A, C); 1 mm (B, D).

Leodice sp. nov.

Fig. 8C, D

Diagnosis

Prostomial lobes frontally rounded, bilobed, slightly notched. Eyes behind bases of palps. Prostomial appendages evenly spaced, with palps slightly thinner than antennae. Antennal styles and palpostyles without articulations. Peristomial rings distinct dorsally and ventrally, continuous laterally. Peristomial (tentacular) cirri present, reach anterior region of peristomium, styles tapering, without articulations.

Maxillae dentition: Mx I left 1, right 1. Mx II left 7, right 9. Mx III left 12 (right absent). Mx IV left 6, right 11. Mx V left 1, right 1.

Branchiae present from chaetiger 4. Branchiae distinctly longer than dorsal cirri. Maximum number of branchial filaments 12–13 (at ~ chaetiger 10–12); five or six anterior chaetigers with single branchial filaments, no posterior chaetigers with single branchial filaments. Branchiae continuing until chaetiger 32–34. Ventral cirri of anterior segments digitiform.

Compound falcigers present, appendages bidentate, hoods without mucros (rounded). Aciculae light yellow, translucent. Subacicular hooks colour light yellow, translucent, bidentate.

Variation

A number of very small specimens (< ~ 1.5 mm maximum width) differ from the above description in having at most three or four branchial filaments, and a single specimen much larger than the remaining material has < 18 branchial filaments. These specimens apparently do not differ in other respects and all are assumed to represent a single species.

Remarks

Laubieriopsis hartmanae (Levenstein, 1970)

Fig. 9A–C

Diagnosis

Prostomium retracted. Body linear, blunt on both ends with 16 chaetigers. Chaetigers 1–4 shorter with 2–3 large acicular chaetae and 2–3 small acicular chaetae per parapodia. Chaetigers 5–16 with one acicular and one capillary per ramus, longest on chaetiger 16. Granular genital papillae on boundary of chaetigers 6 and 7.

Remarks

Similar in appearance to L. brevis from the Atlantic Ocean, but differs in the tips of the aciculars (bidentate in L. brevis) and genital papilla (smooth in L. brevis) (Salazar-Vallejo et al. 2019b). One species of Laubieriopsis was recorded from five stations (932–4068 m) in the GAB (MacIntosh et al. 2018: additional file 2); however, further investigation is required to determine if the species in the present study are the same as at the GAB.

Records

22 specimens. Suppl. material 1: ops. 9, 16, 23, 40, 42, 45, 76, 79, 96, 110, 134 (NHMUK). 2 specimens. Suppl. material 1: op. 79.

Figure 9.

Fauveliopsidae A Laubieriopsis hartmanae B L. hartmanae, chaetae C L. hartmanae, genital papilla D Riseriopsis cf. santosae E Riseriopsis cf. santosae, chaetae. Scale bars: 1 mm (A, D); 50 µm (B); 100 µm (C); 250 µm (E).

Riseriopsis cf. santosae Salazar-Vallejo, Zhadan & Rizzo, 2019b

Records

1 specimen. Suppl. material 1: op. 42 (AM).

Figure 10.

Flabelligeridae A Bradabyssa cf. kirkegaardi (AM W.52573) B Bradabyssa sp.1 (AM W.52551) C Diplocirrus sp. nov. 1 (AM W.52559) D Diplocirrus sp. nov. 2 (AM W.52563) E Diplocirrus sp. nov. 4 (AM W.52551) F Diplocirrus sp. 5 (AM W.52562) G Ilyphagus sp. (AM W.52561) H Flabelligeridae sp. (AM W.52554) I Flabelligeridae sp. Scale bars: 500 µm (A, B, D, E); 1 mm (C, F, G, I); 2 mm (H).

Bradabyssa sp. 1

Fig. 10B

Diagnosis

Incomplete, length 9 mm, width 1 mm, 26 chaetigers, body papillae long, thin, abundant, with mud. Cephalic cage not developed. 2–4 capillary notochaetae, 3–4 anchylosed neurochaetae. Sediment particles present on body surface.

Remarks

This species belongs to group ‘villosa’ described in Salazar-Vallejo (2017).

Records

2 specimens. Suppl. material 1: ops. 16, 31 (AM).

2 specimens. Suppl. material 1: op. 45 (AM).

M. Böggemann, R. Sobczyk

Cylindrical and long-bodied worms, widely distributed in soft bottom sediments from intertidal zone to abyssal depths (Wilson 2000; Böggemann 2002). Glyceridae and Goniadidae constitute the Glycerimorpha group. Glycerids may be easily recognized by having annulated prostomium with two pairs of terminal appendages and four cross-arranged jaws on anterior end of eversible pharynx. The family consist of 46 valid species grouped in three genera (Böggemann 2014a). Sixteen species from three genera have been found in intertidal to upper abyssal zones of Australian waters (Hartman 1964; Day and Hutchings 1979; Hutchings and Murray 1984; Böggemann 2002; Böggemann and Wilson 2003; Böggemann 2015). This study reports at least two glycerid species, one may be new to science.

Glycera cf. capitata Örsted, 1842

Fig. 11C

Diagnosis

Specimens < 7 mm long, 1.2 mm wide. Prostomium with ~ ten rings. Parapodia of mid-body with longer neuropodial than notopodial prechaetal lobes and one rounded postchaetal lobe, dorsal cirri inserted on body wall far above parapodial base. Proboscideal papillae of two types, long digitiform (Fig. 11C) and shorter oval to globular ones. Jaw ailerons with pointed triangular bases. Branchiae absent.

Remarks

The identification to species level is tentative because the type locality of Glycera capitata is in the Atlantic Ocean off Greenland and there are no molecular data.

Records

4 specimens. Suppl. material 1: ops. 79, 96, 98 (AM).

Figure 11.

Glyceridae and Goniadidae. A Glyceridae, Glycera sp. anterior body with everted pharynx, ventrolateral view (AM W.52668) B Goniadidae, Bathyglycinde profunda, whole specimen (AM W.52667) C Glyceridae, Glycera cf. capitata, digitiform proboscideal papilla, lateral view (AM W.52672) D Goniadidae, Bathyglycinde profunda, anterior fragment (AM W.52667) E Bathyglycinde profunda, proboscideal papillae of area II, posterior view (AM W.52669) F Glyceridae, Glycera cf. russa, conical proboscideal papilla with eight ridges, posterior view (AM W.52605). Scale bars: 1 mm (A, B, D); 10 µm (C); 50 µm (E); 10 µm (F).

Glycera cf. russa Grube, 1870

Fig. 11F

Diagnosis

Specimen 74 mm long, 6 mm wide. Prostomium with only nine rings. Postchaetal lobes of mid-parapodia both triangular and similar in length along most of the body, posteriorly more acute. Proboscideal papillae of two types, long conical ones (Fig. 11F) and shorter oval ones, both with 6–10 transverse U-shaped ridges (and V-shaped apically). Jaw ailerons with triangular bases, not pointed nor deeply incised. Branchiae completely absent along body.

As the sister group to glycerids, goniadids are cylindrical, long-bodied annelids. The family is easily distinguished from glycerids by presence of usually one pair of macrognaths and variable number of ventral and dorsal micrognaths instead of two pairs of jaws on the anterior end of pharynx (Böggemann 2005). In addition, some genera have longitudinally arranged rows of V-shaped chevrons on each side of the proboscis. The family consist of 64 valid species in eight genera, two of which are monotypic (Böggemann 2014b). Goniadids are widely distributed from intertidal to abyssal depths (Böggemann 2005, 2009). In Australia, 22 species from six genera have been recorded (Day and Hutchings 1979; Walker-Smith and Wilson 2003; Böggemann 2005, 2015). In this study we report one named goniadid.

Bathyglycinde profunda (Hartman & Fauchald, 1971)

Fig. 11B, D, E

Specimens incomplete. Prostomium subrectangular (broader than long), posteriorly with slight mid-dorsal incision; antennae, presence implied from scars as follows: two lateral and one median antenna situated mid-posteriorly. Eyespots not visible. Palps bi-articulate, palpostyle cylindrical to conical, approximately equal in length to palpophore. Everted proboscis with many cushion-shaped distal papillae. First four segments tentacular, achaetous, bearing eight pairs tentacular cirri, slightly shorter than dorsal cirri of chaetiger 1. Nuchal organs, unpigmented, coalescing mid-dorsally. Parapodia biramous throughout, dorsal cirri slender tapered, ~ equal in length to parapodial lobes, except those on first chaetiger which are many times longer than parapodial lobe; ventral cirri distally attached, slender tapered, ~ half-length parapodial lobes throughout. Notopodia bearing serrated capillaries and one or two smooth spines. Neurochaetae all compound spinigers. Colour in ethanol yellow-white, unpigmented.

Remarks

See above for discussion justifying placement of this material into Neogyptis, tribe Amphidurini Pleijel, Rouse, Sundkvist & Nygren, 2012, and reasons for considering it represents an undescribed species. Neogyptis sp. 2 differs from Neogyptis sp. 1 in having palpostyles approximately equal in length to the palpophore, and lacking brown pigmentation on tentacular segments and the nuchal organs.

Records

2 specimens. Suppl. material 1: ops. 16, 110 (AM).

Parahesiocaeca sp. 1

Diagnosis

Specimens in poor condition, incomplete. Prostomium sub-rectangular, with three antennae, all tapering rapidly and broader at base, extending just beyond tip of palps. Bi-articulated palps, palpostyle cylindrical. Eyes absent. Proboscis with marginal papillae. First two segments tentacular, achaetous, bearing four pairs of tentacular cirri (inferred from stubs). Tentacular and dorsal cirri appearing to be articulated. Parapodia sub-biramous. Neuropodia with long prechaetal lobe, short postchaetal lobe, and short ventral cirrus; all neurochaetae with long-bladed heterogomph falcigers with very fine tips.

Remarks

The present specimens fit the description of the genus Parahesiocaeca Uchida, 2004, although it differs from the description of the only species, P. japonica Uchida, 2004, in lacking eyes, having brown-pigmented nuchal organs, and the heterogomph falcigers having very long blades with very fine tips. The specimens probably represent an undescribed species, but their condition is too poor to describe as a new species.

Records

2 specimens. Suppl. material 1: op 96 (AM).

A. Murray

Lacydoniidae are an uncommon, but widespread group of polychaetes that have affinities with phyllodocids (Rizzo and Magalhães 2019; Rouse and Fauchald 1997). Currently there are 12 described valid species in one genus, Lacydonia (Read and Fauchald 2020). They have been collected worldwide, from intertidal habitats to depths of 5600 m (Rouse and Pleijel 2001). Lacydoniids have previously been recorded from abyssal depths off the west coast of Africa (Böggemann 2009) and in the Campos Basin off Brazil in the South Atlantic Ocean (Rizzo et al. 2016). Although lacydoniids have been recorded from shallow water Antarctic locations (Ehlers 1913; Hartmann-Schröder 1993; Hartmann-Schröder and Rosenfeldt 1988), to date there have not yet been published reports from mainland Australia, with only one record of Lacydoniidae from shallow northern waters in the Timor Sea at 49–62 m depth, as ‘Lacydonia sp.’ (Przeslawski et al. 2018). However, other unreported lacydoniid specimens from shallow waters around Australian coasts (Western Australia, South Australia, Tasmania and Queensland) and deeper waters from Tasmanian seamounts are also held in the Australian Museum collections (Murray, pers. obs.). In this study we report one species.

Lacydonia cf. laureci Laubier, 1975

Diagnosis

Specimen incomplete, ~ 2 mm wide excluding chaetae, 3 mm long for head plus 12 anterior segments. Body dorsoventrally flattened. Prostomium approximately as wide as long, somewhat indented anteriorly, with conspicuous lateral lobes present on posterior margin of prostomium, giving the appearance of a much wider than long prostomium. Eyes absent, median antenna missing. Pair of short digitiform to filiform lateral antennae located in slight incisions mid-prostomium; pair of similar-sized/shaped palps arising ventral to prostomial anterior margin. Faded pale brown pigment present on prostomium, dorsally and ventrally, and dorsally as transverse bands on tentacular segment and some other segments, and as spots on dorsal cirri and parapodia. Tentacular segment short, achaetous, with pair of ventrolateral cirri. Chaetigers 1–3 uniramous, with compound spinigerous chaetae, subsequent parapodia biramous, rami elongate and widely separated, with elongate supracicular lobes. Notochaetae simple capillary chaetae, finely spinulose distally; neurochaetae compound spinigers with heterogomph shaft-heads and long, finely spinulose blades. Dorsal cirri short, thick, digitiform, glandular, inserted basally on first three chaetigers, thereafter medially to distally on notopodia. Ventral cirri of similar size and shape, inserted distally on neuropodia. Posterior segments, pygidium and pygidial cirri, all missing and therefore unknown.

Remarks

Lacydonia laureci Laubier, 1975 is the only currently described species that possesses conspicuous lateral lobes on the posterior margin of the prostomium. This specimen bears some similarity to L. laureci, because of these lobes, as well as the absence of eyes, but there are a few differences also apparent: Rizzo et al. (2016) report that L. laureci possesses capillary notochaetae that have coarse serrations on the distal part of the chaetae, but Böggemann (2009), however, describes fine serrations over the entire length of the notochaetae for that species, whereas this specimen from Australian waters appears to possess fine serrations only on the distal portion of the notochaetae. L. laureci has also been reported from several widely distributed locations in 1001–5497 m depths: the type locality, Matapan Trench, Mediterranean Sea (Laubier 1975); Angola, Cape and Guinea Basin, SE Atlantic Ocean (Böggemann 2009), and the Campos Basin off Brazil, South Atlantic Ocean (Rizzo et al. 2016). Because this specimen consists of an anterior end only, this can only be a tentative identification.

Records

1 specimen (incomplete). Suppl. material 1: op. 110 (AM).

Family Lumbrineridae Schmarda, 1861

P. Borisova, N. Budaeva, D. Ramos

Lumbrineridae is a family of jaw-bearing annelids from the large monophyletic group Eunicida. Lumbrinerids have a simple external morphology, with uniform elongated body, simple uniramous parapodia, and conical prostomium lacking distinct appendages or eyes. No appendages are present on the peristomium consisting of two rings, and only few species have branchiae associated with parapodia. In contrast, the diversity of jaw morphology is remarkable, and morphology of maxillary plates is used as diagnostic characters at genus and species levels. The family comprises 19 genera and ~ 300 species and has world-wide distribution (Carrera-Parra 2006). Lumbrineridae are very common in deep waters being the fifth most diverse annelid family found below 2000 m (Paterson et al. 2009). Australian lumbrinerids are poorly studied with two shallow water species described and one presumably cosmopolitan species reported from the region (Hutchings and Murray 1984; Hartmann-Schröder 1987). In this study we report at least three species from three genera.

Cenogenus sp. nov.

Fig. 13A, B, C

Diagnosis

Body width: 1–2 mm. Prostomium conical, elongated, shorter than peristomium. Nuchal antenna not observed. All parapodia well developed, first four pairs of parapodia smaller than remaining parapodia. Parapodia with inconspicuous prechaetal and postchaetal lobes equal in size. Anterior parapodia with simple digitate elongate (length to width ratio near 2:3) branchia attached dorsally and posteriorly to parapodial lobes, in middle parapodia branchiae decrease in length becoming more rounded.

Anterior parapodia with long fragile limbate chaetae only, middle parapodia with limbate chaetae and simple multidentate hooded hooks with six to seven small teeth and long blade. All chaetae dark, black or dark brown in colour, becoming translucent near tip. Acicula dark, two in median parapodia.

Maxillary apparatus dark, stout, with four pairs of maxillae. Maxillary carriers shorter than MI. MI forceps-like with attachment lamellae, without connecting plates. MII as long as MI, with two large teeth. MIII unidentate, completely pigmented, dark in colour. MIV large, unidentate round-square plates, completely pigmented.

Remarks

The specimen AM W.50137 (op. 33, Suppl. material 1) differs from others by being smaller (width 1 mm, other three specimens near 2 mm wide) and having asymmetrical MII with three teeth on left MII and two teeth on right MII.

Records

4 specimens. Suppl. material 1: ops. 30, 33, 53 (AM).

Figure 13.

Lumbrineridae A Cenogenus sp. nov. (AM W.50139), parapodium from chaetiger 7 B Cenogenus sp. nov., posterior parapodium C Cenogenus sp. nov., maxillae D Eranno sp. nov., AM W.50140, parapodium from chaetiger 11 E Eranno sp. nov., maxillae F Lumbrineris sp., op. 4 (NHMUK), anterior part of the body, dorsal view G Lumbrineris sp., simple hooded hooks. Abbreviations: Mc, maxillary carriers; MI–MIV, maxillary plates. Scale bars: 200 µm (A, B, D); 500 µm (C, E); 5 mm (F); 25 µm (G).

Eranno sp. nov.

Fig. 13D, E

Diagnosis

Prostomium conical, as long as wide, shorter than peristomium. Nuchal antenna absent. All parapodia well developed, first four to five pairs of parapodia smaller than remaining parapodia. Prechaetal lobes inconspicuous in all parapodia, postchaetal lobes auricular in anterior parapodia (1–40), becoming small and rounded posteriorly, always longer than prechaetal lobes.

Anterior parapodia with limbate chaetae and limbate simple hooded hooks. Clear simple multidentate hooded hooks present after chaetiger 25, with seven to eight teeth and long blades, all of similar size. In median chaetigers (27–40) part of limbate chaetae exceedingly longer than in the remaining chaetigers and ~ twice as long as hooded hooks.

Maxillary apparatus dark, with five pairs of maxillae, elongated. Maxillary carriers shorter than MI. MI forceps-like with attachment lamellae, with narrow connecting plates. MII shorter than MI, with five teeth. MIII unidentate, completely pigmented, dark in colour. MIV unidentate, large, completely pigmented. MV reduced to attachment lamella, partly fused with MIV.

Remarks

Limbate simple hooded hooks are not typical for Eranno and similar to those reported for Abyssoninoe, however, maxillary apparatus is of typical Eranno shape with narrow connecting plates and MII significantly shorter than MI. There is not enough information to consider this a new genus, but we suggest it is a new species.

Records

1 specimen. Suppl. material 1: op. 56 (AM).

Lumbrineris sp.

Fig. 13F, G

Diagnosis

Bluntly conical prostomium without appendages. Two peristomial rings of similar sizes. Parapodia uniramous, neuropodia bearing yellow acicula, limbate chaetae compound (until chaetiger 13) and simple (chaetiger 14 onwards) hooded hooks with up to nine teeth. Maxillary apparatus: MII quadridentate, almost as long as MI; MIII and MIV unidentate. Colour in ethanol pale yellow.

Records

1 specimen. op. 4 (NHMUK).

Lumbrineridae gen. spp.

Remarks

Brenke sledge samples were identified to family level.

Records

5 specimens: Suppl. material 1: ops. 16, 31, 43 (AM). 4 specimens. Suppl. material 1: ops. 31, 54, 66, 79 (NHMUK).

Family Maldanidae Malmgren, 1867

J.A. Kongsrud

The family Maldanidae, commonly known as bamboo worms, are infaunal burrowers inhabiting tubes made of sediments consolidated by mucus. The family comprises ~ 240 valid species in 38 genera and five subfamilies (Read and Fauchald 2020). Maldanids are common members of deep-sea soft bottom communities, with ~ 50 species known from deeper than 2000 m depth (Paterson et al. 2009). The family is poorly studied in Australian waters with only ~ 20 species recorded, mostly from coastal areas (http://www.ala.org.au). In this study we report ten species from six genera, with four species likely new to science.

Boguea sp. nov.

Fig. 14A

Diagnosis

Complete specimens with ~ 30 chaetigers, < 20 mm long and 0.2 mm wide. Head rounded without a cephalic plate. Cephalic keel well developed. Nuchal slits curved, parallel on each side of the cephalic keel. Neuropodia with avicular uncini present from chaetiger 5. Avicular uncini in single row in chaetiger 5–8, in double rows from chaetiger 9 and onwards. Pygidium simple, without papillae. Anus terminal. Tube cylindrical, straight, thick and solid, consisting of a thin inner organic layer incrusted with a thick layer of densely packed mud. Ventral glandular pads on anterior part of chaetigers 4–6 with reddish-brown pigmentation.

Remarks

At present, only two species of Boguea have been described, B. enigmatica Hartman, 1945 from North Carolina, USA and B. panwaensis Meyer & Westheide, 1997 from Phuket, Thailand. This is the first record of the genus in the deep sea.

Records

10 specimens. Suppl. material 1: ops. 44, 56 (AM).

Maldanidae gen. spp.

Remarks

Brenke sledge samples were identified to family level or material was too damaged to identify further.

Records

19 specimens. Suppl. material 1: ops. 31, 33, 42, 55, 66, 89, 110 (AM). 9 specimens. Suppl. material 1: ops. 4, 11, 33, 40, 42, 66 (NHMUK).

Family Melinnidae Chamberlin, 1919

T. Alvestad, L. M. Gunton

Melinnidae are tubicolous annelids that often have dorsal hooks. Recently the subfamily Melinninae Chamberlin, 1919 within Ampharetidae was raised to the family Melinnidae (Stiller et al. 2020). The family Melinnidae is composed of five accepted genera and 49 species (Read and Fauchald 2020). Solis-Weiss (1993) suggested that Melinnidae are generally restricted to deeper waters, this appears to only hold true for the genera Melinnopsis McIntosh, 1885 and Melinantipoda Hartman, 1967a of which all species are described from 50–5600 m. Prior to this study, only two species of Melinnidae were reported from Australian waters, Isolda pulchella Müller in Grube 1858 and Isolda warnbroensis Augener, 1914 (Day and Hutchings 1979). Here we report at least five species from two genera. Material from the present study was used to describe two deep-sea Melinnidae species Melinnopsis chadwicki Gunton, Kupriyanova & Alvestad 2020 and Melinnopsis gardelli Gunton, Kupriyanova & Alvestad 2020 , reported here.

Melinna cf. armandi McIntosh, 1885

Fig. 4D, d

Diagnosis

No complete specimens, large, robust worm > 30 mm length, 5 mm width. Abdominal segments very badly preserved and/or missing on all specimens. Body long, widest in postbranchial region. Thorax with 18 chaetigers; neurochaetae as small acicular spines on first four chaetigers and uncini on remaining 14 chaetigers. Prostomium with well-defined anterior and posterior parts, separated by a pair of deep transverse nuchal that almost meeting mid-dorsally. Anterior part distally trilobed. No eyespots. Many smooth buccal tentacles. Chaetiger 1 collar-like, laterally and ventrally encompassing head region; anterior margin not crenulated. Branchiae in two basally fused groups of four. Inner and anterior most branchia of each group only fused at base. Branchiae all long, circular in cross section, tapering evenly to narrow tips. Postbranchial hooks (Fig. 4d) with sharply pointed and gently curved tips. Dorsal end of neurochaetal row on chaetiger 1–3 on elevated lobe. Chaetiger 3 with a few notochaetal capillaries. Chaetiger 4 with small, but well developed notopodia. Serrated brim/fold behind the hooks (dorsal membrane) with ~ 14 equally sized lanceolate projections.

Remarks

Melinna armandi was originally described from west of North Island, New Zealand. Specimens here have 14 lanceolate projections on transverse dorsal membrane whereas M. armandi have eight.

Records

21 specimens. Suppl. material 1: ops. 4, 30 (AM).

Melinnopsis chadwicki Gunton, Kupriyanova & Alvestad, 2020

Diagnosis

Neurochaetae small acicular spines with lanceolate tips on segments 2–5. Neuropodial uncini from chaetiger 5 (segment VI), present in 12 thoracic uncinigers. Postbranchial dorsal membrane low inconspicuous, located on chaetiger 4. Branchiae emerging together on dorsal branchial ridge at level of segments II and III, arranged in two basally fused groups of four. Uncini of thoracic uncinigers with two teeth in one vertical row over rostral tooth, subrostral process and basal prow.

Remarks

Type locality is eastern Australia at 1006–1257 m. For detailed description see Melinnopsis chadwicki Gunton et al. (2020).

Records

27 specimens: Suppl. material 1: ops. 69, 80, 104, 121 (AM).

Melinnopsis gardelli Gunton, Kupriyanova & Alvestad, 2020

Fig. 4F

Diagnosis

Neurochaetae small acicular spines with lanceolate tips on segment II–V. Neuropodial uncini from chaetiger 5 (segment VI), present in 12 thoracic uncinigers. Postbranchial dorsal membrane low inconspicuous, located on chaetiger 4. Branchiae emerging together on dorsal branchial ridge at level of segments II–III, arranged in two basally fused groups of four. Conspicuous stained band immediately behind dorsal fold ending between chaetigers 9 and 10. Uncini of thoracic uncinigers with three teeth in one vertical row over rostral tooth, subrostral process and basal prow.

Remarks

Type locality is eastern Australia at 2520–2821 m. For detailed description see Melinnopsis gardelli Gunton et al. (2020).

Records

62 specimens. Suppl. material 1: ops. 4, 22, 44, 54, 56, 90, 101, 122 (AM).

Melinnopsis spp. nov.

Records

17 specimens. Suppl. material 1: ops. 5, 16, 31, 33, 40, 45, 54, 66, 76, 79, 110, 119 (NHMUK). 65 specimens. Suppl. material 1: ops. 16, 23, 31, 33, 40, 45, 46, 54, 55, 76, 134 (AM).

Micronephthys sp. 1

Diagnosis

Small bodied specimens, 4–8 mm long for < 30 chaetigers. Branchiae (or interramal cirri) absent. Prostomium with straight to slightly convex anterior margin, subpentagonal to round in shape; short, conical antennae inserted on distal margin slightly medial to anterolateral corners; palps single, short, conical, inserted and directed ventrally. Subdermal eyespots not visible. First chaetiger similar in size to following chaetigers, not reduced. Parapodial acicular lobes conical, aciculae with curved tips. Neuropodial superior lobes absent. Parapodial rami with four types of chaetae: barred chaetae present in preacicular position, spinose chaetae present in postacicular position; capillary chaetae present in neuropodia of chaetiger 1; and some finely spinulose (almost smooth) long capillary-like chaetae in postacicular position of notopodia on following chaetigers. Furcate (lyrate) chaetae absent; thick dentate chaetae absent from chaetiger 1. Some specimens with very long spinulose chaetae in mid and posterior chaetigers, so specimens perhaps in swimming phase. Small papilla-like dorsal cirrus present on inner posteroventral face of all notopodia. Dissected pharynx with pair of conical jaws, 20 bifid terminal papillae (none enlarged more than others), plus 20–22 longitudinal rows of subterminal papillae with more than eight long papillae per row, diminishing in size proximally, single elongate middorsal and midventral papillae absent.

Remarks

These specimens agree with the emended diagnosis for the genus by Murray et al. (2015). There are currently 15 valid nominal species of Micronephthys (Read and Fauchald 2020), of which only two species possess the combination of absence of both branchiae and lyrate chaetae (see Dnestrovskaya and Jirkov 2010, 2019): Micronephthys abranchiata Ehlers, 1913 and M. ambrizettana Augener, 1918. These species, however, have fewer papillae in the subterminal rows of pharyngeal papillae (4–6, cf. > 8 for IN2017_V03 specimens), and the latter species also possesses eyespots which the specimens herein do not. Without examination of type material of all Micronephthys species, the identity of specimens described above must remain unknown. Micronephthys MoV6847 (four stations, 203–1521 m) were recorded from the GAB (MacIntosh et al. 2018: additional file 2) and further investigation is required to determine if the species from the GAB are the same as those from the present study.

Records

71 specimens. Suppl. material 1: ops. 9, 16, 23, 27, 31, 33, 40, 54, 55, 79 (AM).

Nephtys cf. paradoxa Malm, 1874

Fig. 2E

Diagnosis

Body stout, wider anteriorly and tapering from middle chaetigers to posterior. Prostomium subrectangular, eyes not visible, pharynx when everted with ten pairs of terminal bifid papillae, and 22 rows of subterminal papillae (rows with 4–6 similar conical papillae) extending only one third of pharyngeal length, median dorsal and ventral papillae not elongated, proximal region smooth. Antennae and palps conical, short, nuchal organs conspicuous, rounded. Parapodia biramous with well-separated rami. Interramal branchiae present, somewhat recurved, from chaetiger 11, becoming foliaceous from ~ chaetiger 16 and appearing membranous from chaetiger 14–16, ciliated and fully developed with membranous expansions (or ‘foliaceous lamellae’) ~ chaetiger 20, becoming small and rudimentary from ~ chaetiger 38 to posterior chaetigers. Acicular lobes obliquely rounded, notopodia with rudimentary preacicular and low postacicular lobes; neuropodia with rudimentary preacicular lobe, postacicular lamella longer than acicular lobe. Chaetae short and ‘spiky’, of three kinds: barred chaetae in preacicular position, spinulose chaetae in postacicular position and capillary chaetae present in neuropodia of chaetiger 1.

Remarks

This species has previously been recorded from off the east Australian coast by Paxton (1974), collected in 1912 from 860 m, and has also been collected from 200 m northeast of Coffs Harbour, NSW in 1993 (unpublished AM records). However, it is a deep-water species widely distributed in the Arctic, the northern Atlantic Ocean, the Mediterranean Sea and the Pacific Ocean (Ravara et al. 2010a), originally described from the North Sea. It is surmised that it may be a species complex and that examination of global material as well as molecular analyses can only resolve its taxonomic status (Ravara et al. 2010b). Therefore, this is a tentative identification.

Records

2 specimens Suppl. material 1: ops. 80, 101 (AM).

Nephtyidae gen. spp.

Remarks

Brenke sledge samples were identified to family level.

Records

7 specimens. ops. 16, 31, 40, 54 (NHMUK).

Family Nereididae Blainville, 1818

D. Ramos, R. S. Wilson

Nereididae are commonly found in intertidal areas worldwide, as a result the family has been extensively studied and used by physiologists for laboratory experiments and as bait by fishermen. They possess an eversible pharynx with one pair of jaws and often have accessory papillae or denticles in a regular pattern. There are 48 currently accepted genera and ~ 708 extant marine species (Read and Fauchald 2020). Abyssal Nereididae faunas are typically dominated by species of Ceratocephale (e.g., see Hilbig 1997; Böggemann 2009). In Australian waters there are 100 species reported from 24 genera (http://www.ala.org.au). In this study we report at least eight species from four genera.

Ceratocephale sp. 1

Diagnosis

Eyes absent; tentacular cirri all very short, longest extending just beyond peristomium; cirrophore of dorsal cirri not significantly expanded; double ventral cirri from chaetiger 3, posterior one twice length of anterior one. Specimens too small to dissect to observe paragnath/papillae arrangement.

Remarks

Specimens do not fit descriptions of any named species. Members of the genus are typically common in deep-sea samples, however, in this study they were rare.

Records

3 specimens. Suppl. material 1: ops. 79, 89, 110 (AM).

Neanthes cf. bassi Wilson, 1984

Diagnosis

Area I = 0–4; II = 6–27; III = 1–14; IV = 1–18 conical paragnaths + 2–7 smooth bars; V = 0–1; VI = 2–16; VII–VIII = 5–30. Dorsal notopodial ligule similar size to acicular ligule throughout (not markedly reduced or expanded on posterior chaetigers). Prechaetal notopodial lobe absent. Neuropodial postchaetal lobe present on chaetigers 1– ~ 12. Ventral neuropodial ligule on posterior chaetigers reduced, up to half length of acicular neuropodial ligule. Notochaetae homogomph spinigers only (homogomph falcigers absent). Neuropodial dorsal fascicle fused falcigers absent.

Remarks

Prior to the collection of the first abyssal samples in Australian waters, Neanthes bassi was only known from 0–147 m. The most similar species is Neanthes tasmani Bakken, 2002 (known from slightly deeper locations, 75–220 m). However, Neanthes specimens from this study and recent GAB voyages are very close to N. bassi, yet they have been recorded from 200–4800 m. This taxon is referred to here as Neanthes cf. bassi, a hypothesis to be tested when molecular data are available. Two species other than N. tasmani are similar enough to be confused with N. bassi (until the pharynx is dissected), Neanthes kerguelensis (which has fewer maxillary ring paragnaths and lacks oral ring paragnaths or has at most VI = 1 and VII–VIII = 8) and Nicon maculata (which is also described here and lacks paragnaths completely yet is otherwise strikingly similar to N. kerguelensis).

This species was also recorded from eight stations (199–4518 m) in the GAB (MacIntosh et al. 2018: additional file 2).

Records

15 specimens. Suppl. material 1: ops. 15, 30, 32, 43, 65 (AM).

Neanthes cricognatha (Ehlers, 1904)

Fig. 15A

Diagnosis

Prostomium with entire anterior margin. Longest tentacular cirri extend back to chaetiger 4. Maxillary ring of pharynx without papillae. Area I = 9–16; II = 22–45; III = 23–45 paragnaths; IV = 29–54; V, VI, VII–VIII forming a dorsally and ventrally continuous ring.

Notopodial prechaetal lobe present, well developed, so that notopodium made up of two ligules and one lobe similar and triangular. Dorsal cirrus length ~ 1 × ventral notopodial ligule at chaetiger 10–20. Neuropodial postchaetal lobe present, at least on some anterior chaetigers. Ventral neuropodial ligule on posterior chaetigers similar to length of acicular neuropodial ligule. Ventral cirri single.

Notopodial homogomph spinigers present; sesquigomph spinigers absent. Notopodial homogomph falcigers absent. Neuropodial dorsal fascicle fused falcigers absent.

Remarks

At 1194–1257 m, this is the deepest record of this species which is widely distributed in Australia and New Zealand from the intertidal to 253 m, suggesting this may be a species complex.

Records

1 specimen. Suppl. material 1: op. 80 (AM).

Figure 15.

Nereididae A Neanthes cricognatha (op. 80) B Neanthes heteroculata, prostomium C Neanthes sp. 1, live specimen D Neanthes sp. 1, prostomium E Neanthes sp. 2, live specimen F Neanthes sp. 2, prostomium G Nereis sp. 1 H Nereis sp. 1, prostomium. Scale bars: 5 mm (A); 1 mm (B, D, F); 10 mm (C, E); 5 mm (G); 0.5 mm (H).

Neanthes heteroculata (Hartmann-Schröder, 1981)

Fig. 15B

Diagnosis

Prostomium slightly wider than long, one anterior pair of very large eyes and one very small pair posteriorly. Jaws with dentate cutting edge, translucent yellow to brown with six teeth. Maxillary ring of pharynx with paragnaths as follows: Area I absent; II absent; III absent; IV 1–3 conical paragnaths; V absent; VI 2 conical paragnaths; VII–VIII 7 conical paragnaths in a ventral band.

Longest tentacular cirri extend back to chaetiger 6. Dorsal notopodial ligule of posterior chaetigers similar to those on anterior chaetigers. Prechaetal notopodial lobe present; smaller than dorsal notopodial ligule on anterior chaetigers, reduced and ultimately absent posteriorly. Acicular process absent. Dorsal cirrus basally attached throughout, ~ 1 × acicular notopodial ligule at chaetiger 10–20. Neuropodial prechaetal and postchaetal lobes absent. Ventral neuropodial ligule of anterior chaetigers present, ~ as long as acicular neuropodial ligule throughout.

Notoaciculae absent from segments 1 and 2. Notopodial homogomph spinigers present. Neurochaetae: dorsal fascicle heterogomph spinigers absent. Neuropodial dorsal fascicle homogomph spinigers present. Neuropodial dorsal fascicle with heterogomph falcigers and homogomph falcigers on anterior chaetigers present. Neurochaetae, ventral fascicle: homogomph spinigers; ventral fascicle heterogomph falcigers.

Remarks

Diagnosis

Eyes absent. Paragnath counts: Area I = 2; II = 14–16; III = 0–3 (unclear, possibly damaged during dissection); IV = 9; V absent; VI = 6; VII–VIII = 30–40. Dorsal notopodial ligule markedly broader and elongate on posterior chaetigers. Prechaetal notopodial lobe absent. Dorsal cirrus length ~ twice acicular notopodial ligule at chaetiger 10–20. Dorsal cirrus on posterior chaetigers terminally attached to dorsal notopodial ligule. Neuropodial prechaetal and postchaetal lobes absent. Ventral neuropodial ligule ~ as long as acicular neuropodial ligule. Notopodial homogomph spinigers present. Notopodial homogomph falcigers present. Notopodial homogomph falciger blades very long. Notopodial homogomph falcigers multidentate, with two or more small lateral teeth, first and subsequent lateral teeth much smaller than terminal tooth. Neurochaetae: dorsal fascicle heterogomph spinigers absent. Neuropodial dorsal fascicle homogomph spinigers and heterogomph falcigers. Neurochaetae, ventral fascicle: heterogomph spinigers and heterogomph falcigers.

Records

1 specimen. Suppl. material 1: op. 100 (NHMUK).

Nicon maculata Kinberg, 1865

Diagnosis

Longest tentacular cirri extending back to chaetiger 5–9. Maxillary and oral rings of pharynx entirely bare of papillae and paragnaths. Dorsal notopodial ligule similar size to acicular ligule throughout (not markedly reduced or expanded on posterior chaetigers). Prechaetal notopodial lobe absent. Neuropodial postchaetal lobe present on chaetigers 1– ~ 12. Ventral neuropodial ligule on posterior chaetigers reduced, up to half length of acicular neuropodial ligule. Notochaetae homogomph spinigers only (homogomph falcigers absent). Neuropodial dorsal fascicle fused falcigers absent.

Remarks

Until the pharynx is dissected and found to be bare, this species is strikingly similar to Neanthes bassi and N. kerguelensis (McIntosh, 1885). Although Nicon maculata does have slightly longer tentacular cirri and the postchaetal neuropodial lobe (which is present in all three species) seems slightly longer and appears articulated or constricted in Nicon maculata where it meets the neuropodial lobe (see further comments for N. bassi, above). These taxa are otherwise very similar, and it is difficult to sustain the placement of Nicon maculata in a different genus as no such revision has yet been undertaken. Hutchings and Reid (1991) were the first to record Nicon maculata from Australia.

This record here at 2687–2821 m is the deepest record of this species that we are aware of. The species is also known from a single specimen (1391 m) in the GAB (MacIntosh et al. 2018: additional file 2) and widely recorded in southeastern Australia, the vicinity of Heard Island, and on the Antarctic continental margin at depths 145–1650 m.

Records

1 specimen. Suppl. material 1: op. 44 (AM).

Nereididae gen. spp.

Remarks

Specimens were too small to dissect to observe paragnath arrangement. Brenke sledge samples were identified to family level.

Records

3 specimens. Suppl. material 1: ops. 33, 54, 128 (NHMUK).

Family Onuphidae Kinberg, 1865

H. Paxton, N. Budaeva

The family Onuphidae, a member of the jaw-bearing order Eunicida, consists of the subfamilies Onuphinae comprising 18 genera and Hyalinoeciinae Paxton, 1986 comprising five genera. Most onuphids are tubicolous; while the Onuphinae are sediment dwellers, well represented in intertidal to shelf depths, the Hyalinoeciinae are often found as epibenthic crawlers in deeper environments, making the onuphids the fourth most diverse polychaete family in the deep sea (Paterson et al. 2009). Onuphinae are much better studied than the Hyalinoeciinae as a result of their depth distribution and the scarcity of bathyal and abyssal sampling (Paxton 2000). In Australian waters there are 31 species reported from 13 genera (http://www.ala.org.au). The present study recovered 161 onuphid specimens, almost exclusively made up of hyalinoeciines. Ten species from four hyalinoeciine genera were identified, the most diverse being Nothria with five species. The only onuphine genus collected was Paradiopatra, with three species reported. A key to genera and their definitions can be found in Paxton (1986).

Anchinothria cf. pycnobranchiata (McIntosh, 1885)

Fig. 16A

Diagnosis

Peristomial cirri present. Parapodia 1 enlarged; parapodia 1–3 with bi- to trilobed prechaetal lobes; subulate ventral cirri on chaetiger 1 and 2. Bidentate simple to pseudocompound hooks on first three parapodia; pectinate chaetae scoop-shaped. Simple branchiae from chaetiger 16–19. Round tubes of inner parchment-like lining and outer muddy layer with embedded foreign objects such as spines or spicules.

Remarks

The species is widely distributed in great depths of southern oceans, perhaps a species complex. It is new to Australian waters, also collected from six stations at the GAB (990–1790 m depth) reported as Anchinothria sp. 1 (MacIntosh et al. 2018: additional file 2).

Records

6 specimens. Suppl. material 1: ops. 56, 69, 115, 122 (AM).

Figure 16.

Onuphidae A Anchinothria cf. pycnobranchiata (op. 56) B Nothria cf. paxtonae (op. 56) C Nothria sp. nov. 1 (AM W.49940) D Nothria sp. nov. 1, tube (AM W.49940) E Nothria sp. nov. 2 (AM W.49933) F Nothria sp. nov. 3 (AM W.49934) G Paradiopatra sp. nov. 2 (AM W.49950). Scale bars: 5 mm (A); 1 mm (B–E, G); 4 mm (F).

Hyalinoecia abranchiata Lechapt, 1997

Diagnosis

Frontal lips fused; palps subulate, with brown median patch. Eyes absent; peristomial cirri absent. Parapodium 1 enlarged; chaetigers 1 and 2 with bidentate simple hooks; limbate and scoop-shaped pectinate chaetae from chaetiger 2. Branchiae absent. Clear, quill-like tubes.

Remarks

The species was originally described from abyssal zones off New Caledonia. It is new to Australian waters.

Records

7 specimens. Suppl. material 1: ops. 80, 128 (AM).

Hyalinoecia sp. nov. 2

Diagnosis

Frontal lips fused; palps subulate, with brown median patch. Eyes absent; peristomial cirri absent. Parapodium 1 enlarged. Only chaetiger 1 with bidentate simple hooks; limbate and scoop-shaped pectinate chaetae from chaetiger 2. Branchiae absent. Clear, quill-like tubes.

Records

43 specimens. Suppl. material 1: ops. 69, 100, 121 (AM).

Hyalinoecia sp. 3

Diagnosis

Frontal lips subulate; palps cirriform. Eyes absent; peristomial cirri absent. Parapodium 1 enlarged, with weakly bidentate simple hooks. Scoop-shaped pectinate chaetae and limbate chaetae from chaetiger 2. Single branchial filament from chaetiger 26. Clear, quill-like tubes.

Remarks

Specimen is perhaps a juvenile Hyalinoecia longibranchiata (McIntosh, 1885).

Records

1 specimen. Suppl. material 1: op. 69 (AM).

40 specimens. Suppl. material 1: ops. 100, 121 (AM).

Nothria sp. nov. 4

Diagnosis

Eyes absent; peristomial cirri present. Chaetiger 1 enlarged; with auricular prechaetal lobes. Simple branchiae from chaetiger 10. Bidentate simple and pseudocompound hooks on chaetiger 1 and 2. Chaetiger 3 with limbate chaetae and ‘scoop-shaped’ pectinate chaetae only. Subacicular hooks from chaetiger 14–16. Flattened tube with transparent inner layer, covered on outside with foraminifera and small shell pieces.

Records

3 specimens. Suppl. material 1: ops. 86, 115 (AM).

Paradiopatra ehlersi (McIntosh, 1885)

Diagnosis

Ceratophores without lateral projections. Peristomial cirri present. Almost unidentate and bidentate pseudocompound hooks with long pointed hoods on first three chaetigers. Subacicular hooks from chaetiger 10. Branchiae with single filaments from chaetiger 17–22, becoming pectinate. Tube with tough lining, outside muddy.

Remarks

This species was previously reported NE of Sydney, collected from 4530 m during RV ‘Galathea’ expedition (Kirkegaard 1994).

Records

3 specimens. Suppl. material 1: ops. 30, 65 (AM).

Paradiopatra sp. nov. 1

Diagnosis

Ceratophores without lateral projections. Peristomial cirri present. Bidentate pseudocompound hooks with long pointed hoods on first three chaetigers. Subacicular hooks from chaetiger 9. Branchiae with single filaments throughout, starting on chaetiger 14–16, filaments becoming very long. Thick mud tube.

Records

11 specimens. Suppl. material 1: ops. 4, 35, 56, 70 (AM).

Paradiopatra sp. nov. 2

Fig. 16G

Diagnosis

Ceratophores with lateral projections. Almost unidentate pseudocompound hooks with long pointed hoods on first three chaetigers. Subacicular hooks from chaetiger 14. Branchiae absent. Mud tube.

Remarks

This species was also recorded from 3794 m at the GAB as Paradiopatra sp. nov.

Records

29 specimens. Suppl. material 1: ops. 33, 53, 54, 56, 78 (AM).

Family Opheliidae Malmgren, 1867

D. Ramos

Opheliids are usually elongate, tapering at both ends, with ventral grooves and reduced parapodial lobes (Magalhães et al. 2019). The family Opheliidae is composed of > 120 species in five genera (Magalhães et al. 2019). A comprehensive taxonomic revision of this family is needed given its confusing taxonomic history and evidence of paraphyly in Ophelina, the most speciose genus (Paul et al. 2010; Wiklund et al. 2019). Travisia was formerly included in Opheliidae, but has since been assigned its own family, which forms a sister group with Scalibregmatidae (Paul et al. 2010). Opheliids occupy intertidal to abyssal sandy and muddy environments (Hutchings 2000c). To date, there have been 30 species reported mostly in the shallow waters of Australia, with all currently accepted genera represented. These include 13 species of Armandia, six species each of Ophelia and Ophelina, one species of Polyophthalmus, and four species of Thoracophelia (Day and Hutchings 1979; Hutchings and Murray 1984; Hartmann-Schröder and Parker 1995; Neave and Glasby 2013; Parapar and Moreira 2015; Moreira and Parapar 2017). In this study we report at least six species from one genus, Ophelina.

Ophelina sp.

Diagnosis

Following synonymy of Ammotrypanella and Ophelina (Blake and Maciolek 2019a). Bluntly conical prostomium with oval palpode having an enlarged base. 32 chaetigers. Chaetiger 24–32 ventrally located and compressed. Chaetigers 1–6 shorter than midbody chaetigers, having more abundant chaetae. Branchial scars present from chaetigers 24–30. Anal funnel damaged, slightly longer than the last two posterior chaetigers, directed dorsally.

Records

15 specimens. Suppl. material 1: ops. 9, 16, 27, 31, 33, 55, 79, 96 (AM). 46 specimens. Suppl. material 1: ops. 9, 16, 31, 33, 45, 54, 76, 79, 98, 110, 134 (NHMUK).

Ophelina cf. cirrosa (Schüller, 2008)

Fig. 17A–C

Diagnosis

Bluntly conical prostomium with slit-like nuchal organs. Eyes absent. Chaetigers 1–9 compressed, having more abundant chaetae. Midbody chaetigers longer than anterior and posterior chaetigers. Ventral and lateral grooves present along entire body. Fan-shaped parapodia. Branchiae from chaetiger 23 to chaetiger 30. Abranchiate chaetigers compressed, with the long chaetae directed dorsally. Anal funnel as long as last six posterior chaetigers, directed dorsally. Colour in ethanol white.

Remarks

Observed specimens resemble Ammotrypanella cirrosa described from the Weddell Sea at 3050 m depth. These also possess an anal funnel with small cirri on the posterior margin but differ in having a large terminal ventral papilla instead of a ventral cirrus in the anal funnel. We followed the synonymy of Ammotrypanella and Ophelina as suggested by Blake and Maciolek (2019a).

Records

76 specimens. Suppl. material 1: ops. 9, 11, 16, 23, 31, 76, 96, 134 (NHMUK).

Figure 17.

Opheliidae A Ophelina cf. cirrosa B Ophelina cf. cirrosa, posterior end C Ophelina cf. cirrosa, anterior end D Ophelina cf. helgolandiae E Ophelina cf. helgolandiae, prostomium F Ophelina cf. bowitzi, branchiae with blister-shaped bumps G Ophelina cf. bowitzi, anal funnel. Scale bars: 3 mm (A); 0.5 mm (B, C); 5 mm (D); 0.25 mm (E, F); 1 mm (G).

1 specimen. Suppl. material 1: op. 79 (AM).

Figure 18.

Orbiniidae A Berkeleyia sp. (op. 79) B Leitoscoloplos cf. abranchiatus (op. 54) C Leitoscoloplos cf. kerguelensis (op. 96) D Leitoscoloplos sp. 1 (op. 30) E Orbiniella cf. aciculata (op. 42) F Protoaricinae gen. sp. (op 100). Scale bars: 500 µm (A, C, E); 2 mm (B, D); 1 mm (F).

Leitoscoloplos cf. abranchiatus (Hartman, 1967a)

Fig. 18B

Diagnosis

All specimens incomplete posteriorly. Longest fragment ~ 15 mm long, 1.3 mm wide, consisting of 41 chaetigerous segments. Thorax slightly flattened, abdomen cylindrical. Prostomium conical; one peristomial ring. Eleven thoracic chaetigers. Abdominal parapodia lateral in anterior abdomen, shifted dorsally on posteriorward segments. Branchiae from chaetiger 24, first short triangle, then becoming longer, strap-like. Thoracic postchaetal lobes conical in both rami; short in anterior thorax, becoming longer in middle and posterior parts; notopodial lobes longer. No subpodial or stomach papillae. Abdominal notopodia digitiform, same length as branchiae or shorter; abdominal neuropodia weakly bilobed, with short round lobes, inner slightly larger. Subpodial flange not developed, no flange papillae. All chaetae crenulated capillaries; forked chaetae not observed, probably broken. Colour in ethanol whitish-brown, with pigment spots on prostomium and in intersegmental furrows, or completely white.

Remarks

Leitoscoloplos abranchiatus was described as entirely lacking branchiae, but all specimens studied were posteriorly incomplete (Hartman 1967a; Mackie 1987; Blake 2017). Specimens described here resemble L. abranchiatus by the number of thoracic chaetigers, shape of thoracic postchaetal lobes, shape and more lateral than dorsal position of abdominal parapodia.

Records

4 specimens. Suppl. material 1: ops. 9, 16, 54 (AM).

Leitoscoloplos cf. kerguelensis (McIntosh, 1885)

Fig. 18C

Diagnosis

Incomplete specimen, 5 mm long, 0.4 mm wide, consisting of 21 chaetigers. Body cylindrical, thoracic segments short, abdominal segments long. Prostomium conical with round tip, one peristomium ring. Nine thoracic chaetigers. Branchiae from chaetiger 17–19, exact position unknown. Thoracic postchaetal lobes short conical in anterior thorax, becoming long and oval in middle and posterior parts; notopodial lobes longer. No subpodial or stomach papillae. Abdominal parapodia small, with short lobes. Notopodia oval, shorter than in thorax; neuropodia weakly bilobed, with short subequal lobes. Chaetae supposedly all crenulated capillaries, but mostly broken; presence of uncini or forked chaetae unknown. Colour in ethanol white.

Remarks

Leitoscoloplos kerguelensis is widespread in Antarctic and subantarctic seas, intertidal to 1400 m; it has 8–10 thoracic chaetigers and branchiae from chaetigers 13–17 (Blake 2017). The specimen studied here has close affinities with this species due to the number of thoracic chaetigers, late beginning of branchiae and the shape of postchaetal lobes. Due to the absence of accurate information on branchiae position, chaetal structure, the large distance from the species area, and bathymetric difference of material collected in this study we suggest the name Leitoscoloplos cf. kerguelensis.

Records

1 specimen. Suppl. material 1: op. 96 (AM).

Leitoscoloplos cf. simplex Blake, 2017

Remarks

Specimens resemble Leitoscoloplos simplex Blake, 2017 from Clarion-Clipperton Fracture Zone.

Records

4 specimens. Suppl. material 1: 79 (AM).

Leitoscoloplos sp. 1

Fig. 18D

Diagnosis

Incomplete specimen, 17 mm long, 1.9 mm wide. Thorax slightly flattened, abdomen cylindrical. First nine segments short, then becoming longer. Fourteen thoracic chaetigers, last one intermediate. Prostomial sharp conical; one peristomial ring. All thoracic segments and at least first three abdominal segments without branchiae. Thoracic postchaetal lobes short conical in both rami, notopodial lobes slightly longer. No subpodial or stomach papillae. Thoracic noto- and neurochaetae long crenulated capillaries. Abdominal region macerated; chaetae broken. Shape of abdominal parapodia and chaetae unknown. Colour in ethanol brown-yellowish.

Remarks

This specimen is similar to L. abranchiatus but has more thoracic chaetigers and shorter thoracic postchaetal lobes. Two OTUs assigned to Leitoscoloplos (four stations, 486–2224 m) were recorded in the GAB (MacIntosh et al. 2018: additional file 2), but further investigation is required to determine if any of the species from this study match those from the GAB.

Records

1 specimen. Suppl. material 1: op. 30 (AM).

? Leitoscoloplos spp.

2 specimens. Suppl. material 1: op. 31 (AM).

Protoaricinae gen. spp.

Records

14 specimens. Suppl. material 1: ops. 43, 44, 56, 65; 78, 99, 135 (AM).

Oweniidae gen. spp.

Remarks

Material is too damaged to be identified further or Brenke sledge material was identified to family level only.

Records

18 specimens. Suppl. material 1: ops. 27, 30, 40, 43, 54, 56, 78, 99 (AM).

Family Paraonidae Cerruti, 1909

J. Langeneck, D. Ramos

Paraonids are small, elongate worms. They have a well-defined prostomium on which some genera have one distinct median antenna. The family Paraonidae includes ~ 140 described species (Blake 2019b). The family is currently divided into eight genera, one of which is further split into four subgenera (Blake 2019b), but molecular data suggest that substantial rearrangements are needed (Langeneck et al. 2019a). Paraonidae typically occur in soft sediments and are especially abundant in shelf to slope depths, where they represent one of the most abundant groups (Blake 2019b), they also show high diversity in bathyal and abyssal environments (Aguirrezabalaga and Gil 2009; Langeneck et al. 2019b). The diversity of Paraonidae is likely largely underestimated even in the best studied geographic areas (Blake 2019b; Langeneck et al. 2019a). In Australian waters this group is poorly known, and published data only refer to shallow environments, whereas the majority of collection data are still unpublished, and presumably a high number of species remains undescribed (Glasby 2000b). This study yielded 23 specimens of Paraonidae belonging to four taxa. Although all specimens could be assigned, at least tentatively, to described species, the frequent occurrence of pseudocryptic taxa in this family and the type localities often far apart from the eastern Australia suggest that they may be undescribed, and that integrative taxonomy is needed to clarify the diversity of this family.

Aricidea sp.

Fig. 19A, B

Diagnosis

Three-lobed prostomium with deep nuchal grooves flanking a cirriform median antenna reaching chaetiger 3. Narrow elongated body. Notopodial postchaetal lobe becoming distinctly cirriform on chaetiger 8. Papilliform neuropodial postchaetal lobe. Cirriform branchiae from chaetigers 4–7. Notopodia with simple capillaries, neuropodia with hooks and capillaries with long arista.

Records

6 specimens. Suppl. material 1: ops. 11, 31, 33, 54 (NHMUK).

Paraonella as currently described is most likely polyphyletic, including species close to Paradoneis Hartman, 1965 and to Paraonis Grube, 1873. The pattern of notopodial lobes observed in this species clearly resembles that occurring in Paradoneis, as in the majority of the known Paraonella species. However, there is the possibility that Paradoneis-like Paraonella also represent separate lineages that independently lost the modified notochaetae.

Records

11 specimens. Suppl. material 1: ops. 33, 54 (AM).

Paraonis cf. quadrilobata (Webster & Benedict, 1887)

Records

6 specimens. Suppl. material 1: ops. 4, 11, 22, 35, 56 (AM).

Figure 20.

Pectinariidae Petta investigatoris (AM W.50672) A ventral view B dorsal view C tube. Petta williamsonae (AM W.50667) D ventral view E anterior end, lateral view. Scale bars: 2 mm (A, B); 5 mm (C); 1 mm (D); 0.5 mm (E).

Petta williamsonae Zhang, Hutchings & Kupriyanova, 2019

Fig. 20D, E

Diagnosis

Cephalic veil completely free from operculum, with smooth or bearing several lappets (slightly raised mounds) anterior margin. Operculum semi-circular with smooth dorsal and lateral margins. Ventral margin of operculum with a transverse row of numerous stout notopodial paleae on each side. Two pairs of comb-like branchiae on segments 3 and 4, consisting of large basal hump and series of well separated free lamellae. Pair of dorso-lateral pads on segment 5. Ventro-lateral lobes smooth on segment 3. Notopodia with paleae on segment 1 and with notochaetae on segments 5–21 (17 pairs). Neuropodia present on segments 8–21, > 14 pairs with transverse tori, each with a row of uncini. Scaphe distinctly separated from posterior segments.

Remarks

Type locality is Bass Strait, eastern Australia, 2760–2692 m.

Records

2 specimens. Suppl. material 1: op. 22 (AM).

Pectinariidae gen. spp.

Remarks

Material is too damaged, no further identification is possible.

Records

2 specimens. Suppl. material 1: op. 11 (AM). 5 specimens. Suppl. material 1: op. 89 (NHMUK).

Family Phyllodocidae Örsted, 1843

D. Ramos, R. S. Wilson

Phyllodocids are commonly known as ‘paddle-worms’ due to their large leaf-like dorsal cirri. There are currently 31 extant genera and 497 accepted species (Read and Fauchald 2020). Benthic Phyllodocidae (pelagic phyllodocids were not sampled in this study) are most common and diverse in shallow waters, especially associated with hard substrates, but some genera inhabit mud flats. However, most genera are also represented in bathyal and abyssal habitats (Blake 1997; Böggemann 2009; Paterson et al. 2009). In Australian waters 15 genera and 30 named species have been reported (http://www.ala.org.au). In this study we report at least five species from three genera.

Clavadoce sp.

Records

1 specimen. Suppl. material 1: op. 69 (AM).

Eumida sp.

Records

36 specimens. Suppl. material 1: ops. 13, 44, 88, 100 (AM).

Eumida cf. angolensis Böggemann, 2009

Fig. 21A

Diagnosis

Prostomium wider than long, with three antennae and two palps. Antennae and palps digitiform. Eyes absent. Tentacular cirri with broad base tapering to a fine tip, four pairs on anterior three segments (1–2–1 arrangement). First tentacular segment dorsally reduced. Everted proboscis barrel-shaped with round terminal papilla. Parapodia uniramous. Chaetae present from segment 2. Dorsal cirri lanceolate, ventral cirri conical, both approximately as long as the neuropodia. Colour in ethanol pale yellow.

Remarks

This specimen closely resembles Eumida angolensis Böggemann, 2009, described from the Angola Basin at depths of 3950–5443 m. Alalykina (2018) reports the presence of Eumida cf. angolensis in the Sea of Okhotsk at 1676–3366 m, which would expand its range to the Pacific Ocean if confirmed. Unfortunately, no descriptions were provided for these specimens that would allow comparisons with the Australian sample. We currently consider this as a different species due to the distance from the type locality and some observed morphological differences such as having 18 instead of 16 terminal proboscideal papilla and bearing more neurochaetae per fascicle.

Records

1 specimen, anterior fragment only. Suppl. material 1: op. 100 (NHMUK).

Sigambra sp. 1

Diagnosis

Small specimen, complete, 47 chaetigers. Pharynx with eight or nine terminal papillae. Dorsal cirri (except for first) slightly longer than ventral cirri. Dorsal hooks starting from chaetiger 3, extending to within a few segments from pygidium; accompanied by one or two small capillary chaetae. Neurochaetae smooth, broad-bladed capillaries of varying lengths. Chaetiger 2 with ventral cirri. Median antenna much longer than laterals, extending back to chaetiger 6; first dorsal cirri cirriform, several times longer than following ones which are slender, foliose. Colour in ethanol white.

Remarks

The specimen is similar to S. magnuncus Paterson & Glover, 2000. It probably represents a new species.

Records

1 specimen. Suppl. material 1: op. 96 (AM).

Pilargidae gen. spp.

Records

1 specimen. Suppl. material 1: op. 134 (AM).

Family Polynoidae Kinberg, 1856

A. Murray, R. S. Wilson

The Polynoidae is the most species-rich of the seven families of Aphroditiformia, commonly known as scale-worms (Read and Fauchald 2020). Although molecular studies have shown the family to be monophyletic, the character supporting the Polynoidae clade (presence of tubercles on the elytra) exhibits many subsequent reversals and there is no morphological diagnosis that separates Polynoidae from other scale worms, nor do all scale-worms have scales (elytra) (Wiklund et al. 2005; Norlinder et al. 2012; Gonzalez et al. 2018). The Polynoidae are widely distributed geographically and ecologically, occurring in all depths from intertidal waters to hadal trenches (Hutchings 2000d; Paterson et al. 2009). There are currently ~ 852 accepted species in 164 genera (Read and Fauchald 2020), some of which appear to be restricted to the deep sea, e.g., species in the subfamily Macellicephalinae (Bonifácio and Menot 2018). Polynoids are one of the dominant epifaunal annelid families in abyssal (> 2000 m) depths (Paterson et al. 2009) but their diversity in Australian waters is poorly known. Records of polynoids from deep water in Australia (> 1000 m) are from McIntosh (1885) who described Eunoe abyssorum from 4755 m depth from 800 km southwest of Victoria, as well as Polynoe ascidioides, which is now considered as nomen dubium (Read and Fauchald 2020); Benham (1921) described Parapolyeunoa flynni as Hololepidella flynni from 2379 m off Tasmania; and Augener (1927) described Lepidasthenia australiensis as Nectochaeta australiensis from 1000 m off the Victorian coast. Averincev (1978) described Eunoe ivantsovi from 1640 m from the Lord Howe Rise, Eunoe papillaris from 1800 m from the GAB, and Harmothoe paxtoni from 1800 m in southern Australian waters, although he also described other more shallowly-collected species, and recorded many polynoid species not previously recorded from deeper waters in the Australian and New Zealand region. More recently, Hanley and Burke (1991) described a new species and genus Brychionoe karenae from 1100 m from the Cascade Plateau in the Tasman Sea, and Kirkegaard (1995) described two new species, Lagisca torbeni from 1320 m and Harmothoe australis from 1340 m, both in the GAB.

From recent sampling voyages by RV ‘Investigator’ to the GAB in 2013–2017, 16 polynoid taxa were distinguished to species level OTUs (MacIntosh et al. 2018), some of which were also found in the 2017 Sampling the ‘Abyss’ voyage (present study material), an additional seven species not previously recorded were also discovered during this latter voyage.

In this study we report at least 11 genera and 15 species, some of which are likely undescribed.

Admetella cf. longipedata (McIntosh, 1885)

Fig. 22A

Diagnosis

Four long-bodied specimens with < ~ 62 segments, at least 24 pairs of elytrophores, all elytrae missing. Pharynx dark purple, with four plate-like falcigerous jaws, and ringed with 21 or 22 pairs of papillae. Prostomiums all badly degraded. Antennal styles all missing, median antenna ceratophore present, inserted in medial notch on prostomium, dorsal to scars of small lateral antennae inserted anteroterminally, antennal scales missing. Cephalic peaks absent. Palps long and robust, with longitudinal rows of very fine papillae. Facial tubercle present, rounded, located dorsal to ridged upper lip, between palp bases. Eyes absent. Low transverse nuchal fold present between first pair of elytrophores. Tentaculophores with chaetae. Parapodia subbiramous, well developed, flattened transversely, elongated (neuropodium 2 × longer than notopodium), with elongated acicular lobes and long cirriform subacicular processes, neuropodia not deeply split dorsally and ventrally, short notopodia arising from anterodorsal faces of parapodia. Neurochaetae flattened iridescent chaetae with pointed bare tips (often split) with rows of fine serrations along one side. Notochaetae all missing or absent.

Remarks

Because of the poor state of prostomiums, it was not possible to determine the presence of antennal scales or sheaths between ceratophores of median and lateral antenna, which A. longipedata possesses. All specimens lacked notochaetae which were assumed to be broken off. In all other features, such as elongate parapodia with long acicular lobes and long cirriform subacicular processes, the position of antennae on prostomium, presence of a nuchal fold, numerous, long, flattened, transparent neurochaetae, > 20 pairs of elytra and > 50 body segments, the specimens resemble Admetella and are closer to A. longipedata than to A. brevis Levenstein, 1978, due to the presence of chaetae on the tentaculophore (see Fauchald 1972). Pettibone’s (1967) illustrations (and part of description) of A. longipedata are incorrect according to Fauchald (1972), who regards that description as belonging to A. hastigerens Chamberlin, 1919.

Records

4 specimens. Suppl. material 1: op. 35 (AM).

Figure 22.

Polynoidae A Admetella cf. longipedata anterior end, dorsal view (AM W.51461) B Austrolaenilla sp. anterior end, dorsal view (AM W.52215) C Bathyeliasona nigra anterior end, dorsal view (AM W.52216) D Eunoe cf. abyssorum anterior end, dorsal view (AM W.51463) E Eunoe cf. opalina dorsal view and elytra (AM W.51464). Scale bars: 1 mm (A, D, E); 3 mm (B); 2 mm (C).

Austrolaenilla sp.

Fig. 22B

Diagnosis

Two complete specimens (AM W.52215; 1.5 cm long, 0.5 cm wide excluding chaetae; AM W.52016: 7 mm long, 3 mm wide excluding chaetae) with 30–35 segments, 14 or 15 pairs of elytrophores, on 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32, all elytra missing. Dorsum with transverse red-brown pigment on every segment to posterior. Head with small cephalic peaks, two pairs of eyes of similar small size, anterior pair situated mediodorsally and level with widest part of prostomium, posterior pair more dorsal. Median antenna with ceratophores inserted in anteromedial notch, style missing, lateral antennae with ceratophores inserted ventral to prostomial lobes, bases touching, with short papillated styles, less than half length of prostomium. Palps long, with rows of small papillae. Tentaculophores with stout notochaetae present. Dorsal cirri present throughout body; dorsal tubercles low, nodular. Parapodia with notopodia shorter than neuropodia, both with elongate acicular lobes, neuropodial acicular lobe with extended papilla-like process. Notochaetae all stout, thicker than neurochaetae, with rows of spines and blunt tips. Neurochaetae all long, fine capillaries, with rows of long slender spines longer than the width of the neurochaeta, and more distally, long fine hairs terminating in a tuft, some with capillary tips and some with truncate tips. No other types of neurochaetae present.

Remarks

The small size of the specimens (30–35 segments, ~ 40+ segments for Austrolaenilla species) indicates that they may be juvenile, but the identification is based on the diagnostic feature for the genus: the presence of neurochaetae much slenderer than notochaetae, and with capillary tips that terminate distally in tufts of fine hairs, which these specimens possess. There are ten currently valid Austrolaenilla species (Read and Fauchald 2020), some of which have been described and reported from southern Antarctic and New Zealand waters, but we are unwilling to attempt a more specific identification due to the small and probably juvenile nature of the specimens.

Records

2 specimens. Suppl. material 1: ops. 4, 42 (AM).

Bathyedithia sp.

Diagnosis

Specimen small-bodied, short, with 23 segments and 10 pairs of elytrophores. All elytra missing. Prostomium bilobed, median and lateral antennae absent, frontal filaments absent, facial tubercle absent. Jaws with lateral serrations, pharynx not everted, thus number of terminal papillae not observed. Palps with small palpophores, palps of similar length to tentacular cirri. Tentaculophores achaetous. Notopodia shorter than neuropodia, with elongate acicular lobes. Notochaetae slenderer than neurochaetae; notochaetae with subdistal rows of fine spines, neurochaetae flattened to concave, serrated along both margins. Ventral cirri from segment 3 inserted medially on neuropodia. Nephridial papillae not observed. Ventral keel absent posteriorly.

Remarks

The specimen is damaged and fragile, and most notochaetae and all elytra are missing. Two genera share the characters which distinguish our specimen (10 pairs of elytra, median and lateral antennae absent): Bathyedithia and Polaruschakov. Both genera also have similar neurochaetae, which are flattened and serrated on both margins. Bathyedithia has seven or nine pairs of terminal papillae on the pharynx compared with Polaruschakov which has 14 pairs (Pettibone 1976) but we have been unable to observe this character on our single fragile and damaged specimen. On the basis of the presence of lateral serrations of the jaws which were observed (and which are absent from Polaruschakov species) we assign this specimen to Bathyedithia.

Records

1 specimen. Suppl. material 1: op. 31 (AM).

Bathyeliasona nigra (Hartman, 1967b)

Fig. 22C

Diagnosis

Specimens short-bodied, complete ones with 18 segments, eight pairs of elytra. Body colouration dark purple-black. Bilobed prostomium tapering anteriorly to subulate frontal filaments (lateral antennae absent). Median antenna on long ceratophore inserted mid-dorsally on prostomium, posterior to frontal filaments. Palps long, tapering, smooth. Tentaculophores with chaetae. Facial tubercle absent. Eyes absent. Dorsal tubercles indistinct/absent, dorsal cirri with papillated styles. Nephridial papillae large, wide, on segments 10–12, but small, cylindrical, on segments 5–9 and 13–18. Parapodia with distally elongate pre-acicular neuro- and notopodial lobes, neuropodial supra-acicular process absent. Notochaetae present, as thick as neurochaetae, stout, with numerous transverse spinous rows and blunt bare tips. Neurochaetae wide, flattened, with serrated lateral margins, tips bluntly pointed, unidentate.

Remarks

Of the four species of Bathyeliasona (Pettibone 1976; Bonifácio and Menot 2018), only Bathyeliasona nigra has 18 segments, tentacular cirri with smooth styles and notochaetae as thick as neurochaetae. Our material conforms closely with the description of Pettibone (1976).

Bathyeliasona nigra was also recorded from the GAB surveys 2013/2015, albeit as ‘Lepidonotinae sp. 2’ (MacIntosh et al. 2018). Most specimens are missing chaetae (broken off), palps and styles of cirri and antenna. This species has been recorded previously from Antarctic waters in depths of ~ 2500 m (Hartman 1967b), as well as from the Indian Ocean, in 3300–4360 m depth (Kirkegaard 1995).

Records

22 specimens. Suppl. material 1: ops. 33, 42, 53, 54, 65, 78, 89 (AM).

Bathypolaria magnicirrata (Neal, Barnich, Wiklund & Glover, 2012)

Diagnosis

Specimens with short bodies, 18–19 segments, nine pairs of small, reduced elytrophores (all elytra missing). Everted pharynx light brown, with seven pairs of similar-sized terminal papillae, two pairs of smooth amber-coloured jaws. Prostomium wider than long, bilobed, eyes absent, median antenna present on small cylindrical ceratophore, style long, tapering. Lateral antennae and frontal filaments absent. Long smooth palps present, inserted ventrolaterally. Tentacular segment fused to prostomium, tentaculophores achaetous, styles filiform, long, ventral style longer than dorsal style. Cirrophores on non-elytrigerous segments more prominent than elytrophores, large, cylindrical anteriorly. Parapodia biramous, elongate with notopodia almost as long as neuropodia, aciculae penetrating epidermis. Ventral cirri inserted subdistally from segment 3. Notopodia with long flattened, wide chaetae with fine serrations along one side, tips pointed, unidentate. Neurochaetae slenderer than notochaetae, flattened and with fine serrations along both sides of each chaeta. Posterior ventral keel present.

Remarks

Austropolaria was originally described as a monotypic genus, based on A. magnicirrata described by Neal et al. (2012) from 1000–1500 m in the Amundsen Sea of the Antarctic region. These specimens agree closely with this species diagnosis including that the pharynx has seven pairs of terminal papillae, nine pairs of reduced elytrophores, large dorsal tubercles, and a posterior ventral keel.

Kolbasova et al. (2020) showed that the monotypic genus Austropolaria is a junior synonym of Bathypolaria Levenstein, 1981 so the combination of the type species Bathypolaria magnicirrata (Neal, Barnich, Wiklund & Glover, 2012) was implicit.

Records

8 specimens. Suppl. material 1: ops. 23, 45, 110 (AM).

Bruunilla sp.

Diagnosis

Remarks

This species is different to all the GAB Harmothoe spp. 1–4, and other Harmothoe species reported from Australian waters.

Records

16 specimens. Suppl. material 1: ops. 5, 9, 11, 14, 40, 45, 55, 56, 70, 76, 86 (AM).

Figure 23.

Polynoidae 2 A Harmothoe sp. 5 anterior end, dorsal view, and elytra (AM W.52581) B Lepidasthenia indet. dorsal view, pharynx extended (AM W.51580) C Macellicephalinae sp. 5 anterior end, dorsal view (AM W.52014) D Polaruschakov sp. dorsal view (AM W.52580). Scale bars: 5 mm.

Harmothoe indet.

Diagnosis

Single specimen, broken, with 37 segments, missing elytrae. Fifteen pairs of elytrophores on 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 26, 29, 32. Palps smooth. Prostomium with prominent cephalic peaks. Two pairs widely spaced large eyes, anterior ones laterally located, posterior ones also laterally placed. Median antenna with large ceratophore, style papillated and longer than palps, lateral antenna short, inserted ventrally, bases separated. Lateral antennae styles papillated, short, approximately half length of prostomium. Prostomium pigmented (brown). Tentacular segment with notochaetae. Dorsal cirri styles papillate, tapering. Notochaetae slightly thicker than neurochaetae. Notochaetal bundles held erect but not meeting dorsally. Notochaetae with rows of spines to tip. Neurochaetae slenderer, with rows of spines, medially inflated, and fine bidentate bare tips (secondary tooth small). Neuropodial acicular lobe with small distal digitiform lobe. Notopodial lobe with elongate acicular lobe.

Remarks

As elytra are all missing, this specimen could not be identified to species. It does not resemble Harmothoe sp. 5 of this survey or any of the Harmothoe species found in the GAB surveys.

Records

1 specimen. Suppl. material 1: op. 121 (AM).

Lepidasthenia indet.

Macellicephalinae gen. sp. 5

Fig. 23C

Diagnosis

Large bodied specimen, complete, 33 mm long, 13 mm wide (including parapodia) for 26 segments. Facial tubercle present. Pharynx not everted, but jaws observed via dissection: two pairs of triangular jaws with four or five teeth per jaw; unknown number of pharyngeal papillae. Prostomium bilobed, lobes rounded anteriorly, and posteriorly, median antenna missing or absent, lateral antennae and frontal filaments absent, eyes absent. Palps smooth, short, with reduced palpophores. Tentacular segment fused to prostomium, with tentaculophores situated lateral to palps, both dorsal and ventral styles similar in length and form to palps, achaetous. Large dorsal papillated swollen structures present on (non-elytrigerous) segments 6, 8, 10, located between dorsum and base of cirrophores, possibly reproductive. Dorsal cirri long, longer than parapodia; ventral cirri on segment 2 larger than following segments, becoming small and filiform from segment 3 to posterior segments, inserted subdistally on neuropodia. Parapodia with notopodia reduced to small elongate acicular lobe on anterior face, with slender notochaetae emerging basally, chaetae with rows of fine spines and filiform tips; neuropodia large with elongate pre-chaetal lobe and rounded postchaetal bract-like lobe, neurochaetae much more stout than notochaetae, golden, lanceolate with rows of spines along two sides. Only six pairs of elytrophores distinct, on segments 2, 4, 5, 7, 9, and 11; thereafter difficult to discern due to swollen dorsal bases of parapodia. Pair of thick elytra present on first chaetiger (segment 2), elongate-reniform, covering dorsum, with some dark pigment spots and small marginal papillae; a single large elytra present on segment 11, thin, translucent, round to oval, without pigment or marginal papillae, not covering dorsum. Dorsal tubercles large, and from segment 12, basal swellings present dorsally on every parapodium, with two pairs of ridges running anteriorly and posteriorly along parapodium from dorsum towards dorsal cirrus; these swollen bases also papillated from segment 21. Ventral keel absent. Pygidium rounded, anus dorsal.

Remarks

Due to the lack of lateral antennae, this specimen is assigned to Macellicephalinae, but does not appear to bear resemblance to any of the 37 currently valid Macellicephalinae genera (Read and Fauchald 2020), due to the combination of the form of the neuropodial lobes and notopodial lobes, lack of branchiae, ridged and swollen dorsal tubercles (similar to those of Lepidonotopodium spp. but otherwise dissimilar to that genus), lack of frontal filaments, and the presence of serrated jaws.

Records

1 specimen. Suppl. material 1: op. 79 (AM).

Macellicephalinae indet.

Diagnosis

Small specimens from 3–8 mm in length, with 12–23 segments, and 6–11 pairs of elytrophores. Most somewhat damaged, thus difficult to identify. Possibly seven morphologically different species. Median antenna present or absent, lateral antennae absent, frontal filaments present or absent, facial tubercle absent or present, palps long or short (one specimen with thick, leaf-shaped palps), tentaculophores achaetous. Parapodia with subequal noto- and neuropodial lobes, or notopodial lobes reduced; notochaetae missing, present or absent, neurochaetae either similar thickness to, more robust than, or thinner than, notochaetae; most neurochaetae distally flattened, concave with rows of serrations along both sides. Dorsal tubercles large, or indistinct. Jaws present; pharynx with terminal papillae. Branchial and reproductive structures absent. Ventral keel absent.

Remarks

Numerous specimens were from Brenke sledge samples, most damaged. Approximately seven species are present.

Records

30 specimens. Suppl. material 1: ops. 16, 23, 31, 33, 40, 54, 79, 87, 89, 96, 110, 134 (AM).

Polaruschakov sp.

Records

> 100 specimens. Suppl. material 1: op. 100 (AM). 12 specimens Suppl. material 1. op. 100 (NHMUK).

Figure 24.

Protodrilidae A Protodrilus cf. puniceus B same, ventral side C same, pygidium. Scale bars: 250 µm.

Family Sabellariidae Johnston, 1865

E. K. Kupriyanova, J. Zhang

Sabellariids are filter feeding annelids, which inhabit tubes made of sand and shell fragments cemented together. The family Sabellariidae is composed of 12 genera and 132 species (Read and Fauchald 2020). Most sabellariids live in intertidal or subtidal habitats, sometimes building reefs, thus the common name ‘honey-comb worms’ (Kirtley 1994; Hutchings et al. 2012). The genera Bathysabellaria, Gesaia, Phalacrostemma and Tetreres are often found in bathyal and abyssal localities (Lechapt and Kirtley 1996; Hutchings et al. 2012). Five genera and 11 species have been recorded from Australian waters, but only Phalacrostemma maloga Hutchings, Capa & Peart, 2012 and Tetreres terribilis Hutchings, Capa & Peart, 2012 are known from bathyal depths (Hutchings et al. 2012). Material from the present study was used to describe new species Gesaia csiro Zhang, Hutchings, Burghardt & Kupriyanova, 2020 and Phalacrostemma timoharai Zhang, Hutchings, Burghardt & Kupriyanova, 2020, while a single specimen of Phalacrostemma was too damaged to be formally described as a new species (Capa and Hutchings 2019c; Zhang et al. 2020).

Gesaia csiro Zhang, Hutchings, Burghardt & Kupriyanova, 2020

Fig. 25A, B

Diagnosis

Operculum completely divided into two elongate free lobes. Twenty-two pairs of outer paleae, their blades with frayed thecae and rolled inward tips. Two pairs of inner opercular paleae on dorsal margin of opercular lobes, with straight cylindrical blades with smooth margins. Six pairs of long conical papillae spirally arranged around opercular lobes. One pair of nuchal hooks without limbation. Three pairs of tentacular filaments along margins of buccal cavity. Buccal flaps absent. One pair of long palps extending beyond operculum. Thoracic segment 1 with neuropodial cirri. Thoracic segment 2 with one pair of triangular lateral lobe. Eleven pairs of dorsal branchiae on chaetigers 2–12. Four parathoracic chaetigers (3–6) bearing notopodia with robust lanceolate chaetae interspersed with fine capillaries and neuropodia bearing thin lanceolate chaetae interspersed with fine capillaries. Cauda long and smooth with three pairs of anal appendages.

Remarks

Type locality is Central Eastern MP, eastern Australia, 4414–4436 m. The genus Gesaia is recorded from eastern Australian waters for the first time. Seven specimens of Gesaia sp. 1 were recorded from three stations in the GAB (932–1836 m) (MacIntosh et al. 2018: additional file 2), molecular data are needed to confirm if the species in this study are the same as from the GAB.

Records

208 specimens. Suppl. material 1: ops. 88, 89, 128 (AM).

Figure 25.

Sabellariidae A Gesaia csiro holotype (AM W.49506), anterior end with tube B G. csiro, entire body, lateral view C Phalacrostemma timoharai holotype (AM W.50674), ventro-lateral view D P. timoharai, lateral view E P. timoharai, tube F P. timoharai, entire body in tube, lateral view. Scale bars: 1 mm (A, B, E, F); 0.5 mm (C, D).

Gesaia sp.

Remarks

The specimen is identified to genus, further investigation required to determine if it is the same species as Gesaia csiro.

Records

1 specimen. Suppl. material 1: op. 86 (NHMUK).

Phalacrostemma timoharai Zhang, Hutchings, Burghardt & Kupriyanova, 2020

Fig. 25C–F

Due to small size of specimens, examination of uncinal teeth was difficult, and a positive identification to genus was not able to be confirmed. Specimens were stained with methyl blue and photographed to show staining pattern. Some characters of the genus Amphicorina could be observed: eight thoracic and at least eight abdominal chaetigers; three pairs of radioles with few pinnules, long ventral pinnular appendages present; glandular girdle on chaetiger 2; thoracic notochaetae either elongate and narrowly-hooded or small thin bayonet chaetae (with paleate chaetae absent entirely), thoracic uncini acicular with rows of teeth over main fang (though presence of one larger proximal tooth was not observable); abdominal neurochaetae narrowly-hooded, uncini rasp-shaped; posterior anal depression absent. The collar features are, however, more consistent with Chone or Jasmineira, though the chaetal features are more consistent with Amphicorina. However, Amphicorina is more typically recorded from shallow and nearshore waters than deep and has not been reported previously from abyssal depths.

Records

6 specimens. Suppl. material 1: ops. 42, 68 (AM).

Figure 26.

Sabellidae, Syllidae, Terebellidae. Sabellidae A Sabellidae sp. 1 (AM W.52133) B Sabellidae sp. 2 (AM W.52214) C Potamethus cf. scotiae (AM W.51585). Syllidae D Anguillosyllis sp. (AM W.52625) E Exogone cf. heterosetosa (AM W.52329) F Terebellidae (sensu stricto) (AM W.50426). Scale bars: 1 mm (A, B, D, E); 5 mm (C, F).

Sabellidae gen. sp. 2

Potamethus cf. scotiae (Pixell, 1913)

Fig. 26C

Diagnosis

Eight thoracic and numerous abdominal segments. Branchial crown with 6–9 pairs radioles. Wide ventral ‘flange’ on ventralmost radioles; ventral sacs large, external to crown; ventral shields present on body. Collar with dorsal lamellae and dorsal pockets, prolonged ventrally with large ventral lappets, oblique laterally. Peristomial ring elongate, exposed above collar. Thoracic notochaetae including superior narrowly-hooded chaetae and two inferior rows of paleate chaetae; thoracic uncini avicular with extremely long handles (> 10 × distance of main fang to breast); companion chaetae present, with similarly long handles. Abdominal neurochaetae of two types: short, broadly-hooded with long tips and longer elongate narrowly-hooded chaetae; abdominal uncini avicular with long handles/shafts, but with breast reduced to narrow swelling at curvature. Ventral surface glandular, thoracic shields prominent. Tubes muddy with fine transverse striations.

Remarks

Four large fragmented specimens were removed from tubes. There are currently no Potamethus species reported from Australia, however, there are museum records of Potamethus collected from deep water east of Tasmania in 1986 (Murray, pers. obs.), and more recently from the GAB surveys in 2015 and 2017 (MacIntosh et al. 2018: additional file 2). Worldwide, there are 11 nominal species, all described from deep waters, and types would need to be examined to determine to which species these specimens belong, or if it is new. Based on descriptions from the literature, the specimens bear greatest resemblance to P. scotiae (Pixell, 1913) from Antarctic waters.

Records

4 specimens. Suppl. material 1: op. 53, 122 (AM).

Potamethus sp.

Remarks

Identification is to genus only, based on presence of dorsal lamellae joining dorsal collar, and presence of extremely long-handled thoracic uncini and companion chaetae, paleate inferior thoracic chaetae, and broadly-hooded superior thoracic chaetae. Possibly is the same as above specimens of Potamethus cf. scotiae, but specimens were small, incomplete and/or degraded too much from poor preservation whilst in their tubes.

Records

2 specimens. Suppl. material 1: op. 53, 121 (AM).

Family Scalibregmatidae Malmgren, 1867

J.A. Blake

Scalibregmatids, sometimes called maggot worms, are characterized by anteriorly swollen, short bodies. They are active burrowers and subsurface deposit feeders, which never form tubes. The family is composed of 16 genera and ~ 72 species (Blake 2015, 2019c; Read and Fauchald 2020). In Australia, six named species in five genera (Asclerocheilus, Hyboscolex, Oligobregma, Pseudoscalibregma and Scalibregma) have been reported (http://www.ala.org.au). In the present study samples contained at least four genera and six species; at least four new to science. New species include Asclerocheilus (one), Scalibregmides (one), Pseudoscalibregma (one) and Oligobregma (one).

Asclerocheilus sp. nov. 1

Diagnosis

Large specimens, heavy yellow spines in noto- and neuropodia of chaetiger 3.

Records

5 specimens. Suppl. material 1: ops. 53, 65 (AM).

Oligobregma sp. nov. 1

Fig. 27D, E

Diagnosis

The most abundant species in the collections. Similar to Oligobregma mucronata Blake, 2015, from upper slope depths, Antarctica.

Records

193 specimens. Suppl. material 1: ops. 5, 9, 16, 23, 31, 33, 54, 79, 89, 103, 110 (AM).

Figure 27.

Scalibregmatidae and Siboglinidae. Siboglinidae A Frenulate tube (op. 11) B Osedax sp. nov., preserved female specimen inside tube C Osedax sp. nov., detail of palps of preserved female specimen. Scalibregmatidae D Oligobregma sp. nov. 1 (AM W.52686) E Oligobregma sp. nov. 1 (AM W.52698). Scale bars: 2 mm (A, B); 500 µm (C); 1 mm (D, E).

Oligobregma spp.

Records

3 specimens. Suppl. material 1: ops. 11, 27 (AM).

Pseudoscalibregma sp. nov. 1

Bathyvermilia challengeri Zibrowius, 1973

Fig. 28A

Diagnosis

Tubes with characteristic sculpture of numerous transverse ridges close to each other.

Remarks

Only empty tubes were collected. The original records of this species came from three RV ‘Challenger’ stations in the North and South Pacific Ocean taken at 4246–5719 m (Zibrowius 1973).

Records

2 tubes. Suppl. material 1: op. 90 (AM).

Records

661 specimens. Suppl. material 1: ops. 4, 14, 56, 78, 97, 99, 102, 109, 128, 135 (AM).

Smooth white tube. Elongated operculum with distinctly differentiated endplate. Peduncle smooth, of same thickness as radioles, annulated distally, separated from operculum by a constriction. Five thoracic chaetigerous segments plus minute simple collar chaetae. Thoracic membranes short, ending at chaetiger 3. Collar high, trilobed, ventral lobe larger than lateral ones. Uncini rasp-shaped with very characteristic for the genus anterior peg made of two or more rounded lobes with shallow incision(s) in between. Apomatus chaetae absent. Abdominal chaetae in posterior chaetigers only, capillary with tip made of two rows of denticles.

Records

3 specimens. Suppl. material 1: ops. 86, 97, 104 (AM).

Protis sp. 1

Diagnosis

Smooth white relatively thick tubes lacking sculpture. Six thoracic chaetigerous segments plus collar chaetae. Operculum, if present, globular transparent on normal pinnulated radioles.

Remarks

SEM and molecular data are needed to confirm this preliminary identification.

Records

48 specimens. Suppl. material 1: ops. 35, 86, 88, 97, 99, 100, 128 (AM).

11 specimens of frenulate tube pieces. Suppl. material 1: op. 11 (NHMUK).

Osedax sp. nov.

Fig. 27B, C

Diagnosis

Siboglinid found colonising fin whale fall. Female living within tube and with ‘root’ structures embedded into bones. Females with crown of four palps fused for much of their length, without obvious pinnules but with distinct blood vessels in live specimens. Trunk short in relation to the length of the palps, ovisac was not observed. Tube: anterior thin, semi-transparent and appearing closed at the tip, posterior tougher and creased. Colour in ethanol pale yellow.

Remarks

Genetic data confirm that these specimens are a new species that falls within the same clade as other nude palp Osedax species (Georgieva et al. in prep) associated with whale falls.

Records

More than 20 specimens. Suppl. material 1: op. 100 (NHMUK) (whale bones, MV).

Family Sigalionidae Kinberg, 1856

A. Murray

Sigalionids are a family of scale worms with elongate, narrow bodies and usually with a larger number of segments than in the Polynoidae. There are currently considered to be five subfamilies: Sigalioninae, Pelogeniinae, Pholoinae, Pisioninae, and Sthenelanellinae (Gonzalez et al. 2018; Eibye-Jacobsen et al. 2020), all characterised by the presence of some kind of compound neurochaetae. There are currently ~ 252 accepted species in 32 genera (Eibye-Jacobsen et al. 2020). They occur worldwide from intertidal to depths of < 4000 m (Pettibone 1989; Wehe 2007; Eibye-Jacobsen et al. 2020) but are rarely present in large numbers. Some Sigalioninae and Pholoinae species are exclusive to the deep sea (Gonzalez et al. 2018; Eibye-Jacobsen et al. 2020), particularly those from the genera Neoleanira, Pholoides and Pholoe (Pettibone 1970; Pettibone 1992; Wehe 2007; Ravara and Cunha 2016). In Australia there are 12 genera and 15 named species recorded (http://www.ala.org.au). In this study we report at least five species (two of which are new) from four genera.

Leanira sp. nov.

Fig. 29A, B

Diagnosis

Mostly incomplete specimens, at least 28 mm long, 3 mm wide, for 70 segments (or 26 mm long, 4.5 mm wide for 40 segments). Prostomium sometimes with dark pigmentation or a few spots. Elytra smooth on surface, small and round anteriorly, becoming larger and more ovate (kidney-shaped) posteriorly, almost covering dorsum, lacking lateral indentations. Median antenna short, subulate, without auricles, lateral antennae located on inner dorsal side of tentacular segment. Eyes absent. Palps long, palpal sheaths present. Labial lobes present on lateral lips, bulbous. Dorsal cirri absent from segment 3. Neurochaetae all compound spinigers with entire tips, canaliculate. Clavate stylodes present on parapodia. Branchiae starting at ~ chaetiger 30.

Remarks

This species has also been collected from depths of < 920 m along the east Australia coast during cruises by the FRV ‘Kapala’ (1980) and RV ‘Franklin’ (1988) (Hutchings et al. in prep.). It bears some resemblance to L. quatrefagesi Kinberg, 1856 (only recorded from intertidal areas in the South Pacific and Southern Oceans) and L. hystricis Ehlers, 1874 (recorded from 900–2600 m in North Atlantic waters) because of the presence of labial lobes, the lack of neurochaetae, the lack of lateral indentations of the elytra and the absence of segmental papillae (Hutchings et al. in prep.).

Records

17 specimens. Suppl. material 1: ops. 22, 23, 40, 45, 54, 56.

Figure 30.

Sphaerodoridae A–C Clavodorum cf. longipes D–G Sphaerephesia sp. nov. 1 (D, stained with methyl blue) H–J Sphaerephesia sp. nov. 3 K, L Sphaerephesia sp. nov. 5 M, N Sphaerodorum sp. Scale bars: 100 µm (A, B, D, E, I–M); 1 mm (G, H); 10 µm (C, N).

Geminofilum sp. 1

Diagnosis

Body short (1.5 mm), sub-cylindrical, strongly converse dorsally (sub-circular in cross section), with a dark purple-brown pigment in preserved specimen, and whitish macrotubercles. Head invaginated, appendices not observed. Macrotubercles sessile, hemispherical, arranged in two transverse rows per segment, with eight and nine tubercles each. Additional dorsal epithelial papillae absent. Scarce ventral papillae in mid-body segments, but not clearly observed. Parapodia with cylindrical ventral cirri, reaching the tip of acicular lobe. One spherical parapodial papillae, at the base of parapodia (?). All chaetae compound (five or six per parapodium), with blades 4–5 × as long as wide in mid-body segments.

Remarks

This is a possible new species, but material is too damaged to confirm.

Records

1 specimen. Suppl. material 1: op. 45 (AM).

2 specimens. Suppl. material 1: op 9, 16 (AM).

Sphaerodoridae gen. spp.

5 specimens. Suppl. material 1: ops. 27, 40, 54, 79 (AM).

Spiophanes cf. viriosus Meißner & Hutchings, 2003

Remarks

The specimen is only a middle fragment in poor condition, without prostomium and posterior end. Due to this we abstain from a more detailed description. However, based on pigment observable in parapodia of the middle body region, chaetal spreaders which are not of the ‘0+1’ type but possibly of the ‘2+3 type’ present in chaetigers 5–7, and glandular openings in chaetiger 8 being absent the fragment might be tentatively referred to Spiophanes viriosus Meißner & Hutchings, 2003.

Spiophanes viriosus was originally described from coastal waters in Queensland, Australia.

Records

1 specimen, middle fragment. Suppl. material 1: op. 11 (AM).

Spionidae gen. spp.

Remarks

Specimens from Brenke sledge samples were incomplete and could not be identified.

Records

24 specimens. Suppl. material 1: ops. 5, 9, 23, 27, 31, 33, 40, 42, 45, 54, 66, 79, 96, 98, 110, 134 (NHMUK).

Family Sternaspidae Carus, 1863

M. Georgieva

Commonly known as mud owls, Sternaspidae are distinctive round-bodied or peanut-shaped worms are easily recognized by their characteristic and often colourful ventro-caudal shield (Drennan et al. 2019). Currently, Sternaspidae is comprised of 42 species in three genera, with the largest genus, Sternaspis Otto, 1820, containing 32 species (Read and Fauchald 2020). They have a global distribution and live buried in soft sediment at depths varying from the intertidal zone to 4400 m. Three named species from two genera Caulleryaspis Sendall & Salazar-Vallejo, 2013 and Sternaspis have been reported from Australian waters (http://www.ala.org.au). In this study, we report two species from the genus Sternaspis.

Sternaspis sp.

Fig. 32A, B

Diagnosis

Body ~ 5 mm long and < 2 mm wide. Segments between introvert and rest of body highly cinched, with body 0.7 mm wide at narrowest point. Differing colouration between introvert and abdomen apparent. Ventro-caudal shield a bright orange colour, ribbed and concentrically ringed.

Remarks

These specimens may also represent Sternaspis cf. annenkovae, but further investigation is required to confirm this.

Records

3 specimens. Suppl. material 1: ops. 35, 40 (AM).

Figure 32.

Sternaspidae. Travisiidae A Sternaspis sp., ventral view (op. 40) B Sternaspis sp., dorsal view (op. 40) C Sternaspis cf. annenkovae, ventral view (op. 40) D Sternaspis cf. annenkovae, ventral view (op. 40) E Travisia sp. 1 (AM W.52547). Scale bars: 1 mm (A, B); 3 mm (C, D); 5 mm (E).

Sternaspis cf. annenkovae Salazar-Vallejo & Buzhinskaja, 2013

Fig. 32C, D

Remarks

Specimens of Pista were in poor condition. They probably represent at least two species, but branchiae and lateral lobes, critical characters to distinguish species, are damaged and in many cases incomplete, including posterior thorax. Molecular data may help in distinguishing between species.

Records

23 specimens. Suppl. material 1: ops. 4, 22, 30, 35, 56, 67 (AM).

Terebellidae gen. nov. sp. nov. 1

Diagnosis

Fourteen pairs of notopodia, neuropodia begining before end of notopodia, abranchiate genus.

Remarks

Potentially new genus and species.

Records

1 specimen. Suppl. material 1: op. 22 (AM).

Terebellidae gen. spp.

Fig. 26F

Remarks

Specimens were too damaged for further identification.

Records

Figure 34.

Echiura whole animals of Bonelliidae spp. A AM W.49516 B AM W.49541 C AM W.49541 D AM W.49663 E AM W.49664 F AM W.49665 G AM W.49666. Scale bars: 10 mm (A, D, E); 2 mm (B); 1 mm (C); 5 mm (F, G).

Class Sipuncula

A. Schulze

Formerly considered a distinct phylum, Sipuncula are now regarded as a branch within the annelid radiation (Weigert et al. 2014; Weigert et al. 2016). Sipunculan classification was recently revised based on phylogenetic and phylogenomic studies (Kawauchi et al. 2012; Lemer et al. 2015). Approximately 150 sipunculan species currently recognized (Cutler 1994; Schulze et al. 2019) are organized into six families. The actual number of species is likely much higher based on recent molecular studies indicating that cryptic species are commonplace (Schulze et al. 2012).

In contrast to the ‘typical’ annelid body plan, the sipunculan body is unsegmented. It consists of a trunk region and a retractable introvert, generally with a crown of tentacles at its anterior end. Recurved, proteinaceous hooks are often present along the introvert. The anus is usually located dorsally at the anterior end of the trunk. Nephridiopores (usually two) open at a similar level as the anus on the ventral side. Internally, one two four introvert retractor muscles present.

Sipunculans are generally cryptic in their lifestyle, but can reach high densities in some habitats. They range from the intertidal zone to depths of > 7000 m (Saiz Salinas et al. 2018). Cutler (1977) who examined the material from the ‘Galathea’ expedition, reported 43 species collected below 400 m depth, some of them from southeastern Australia. Saiz Salinas et al. (2018) reviewed literature on deep-sea sipunculans from below 2000 m and listed 51 species. Few of these represented records from southeastern Australia within that depth range. MacIntosh et al. (2018: additional file 2) listed > 1000 specimens from 25 stations (depth range 388–3884 m) in the GAB; together with the present material these collections, when formally described, will add significantly to knowledge of the diversity of abyssal sipunculans.

Sipuncula fam., gen. spp.

Fig. 35D

Remarks

Many specimens not identified beyond phylum level.

Records

101 specimens. Suppl. material 1: ops. 4, 6, 22, 32, 33, 43, 53, 54, 69, 88, 90, 100, 104 (AM). 2 specimens. Suppl. material 1: ops. 11, 100 (NHMUK).

Figure 35.

Sipuncula A Sipunculus sp. (AM W.49645), lateral view B Golfingia muricaudata (AM W.49656), lateral view C Golfingia muricaudata (AM W.49657) D Sipuncula gen. sp. (op. 35), lateral view E Phascolion lutense (AM W.49601), top: specimen removed from sediment tube; bottom: specimen partially removed from tube; right: sediment tube. Scale bars: 5 mm.

Family Sipunculidae

Relatively large sipunculans (usually > 5 cm), with an introvert shorter than the trunk; no introvert hooks. Tentacles arranged in a circle surrounding the mouth. Body wall with externally visible bands of longitudinal and circular musculature crossing each other, giving the impression of rectangular ‘mini-pillows’. One genus and four species are known from the deep sea (> 2000 m) (Saiz Salinas et al. 2018). In Australian waters 11 species from two genera (Siphonosoma and Sipunculus) have been reported (http://www.ala.org.au). We report at least one species from Sipunculus.

Sipunculus spp.

Fig. 35A

Diagnosis

Four stem-like tentacles, with the ventral pair smaller than the dorsal pair. Nephridiopores located slightly anterior to the anus.

Records

25 specimens. Suppl. material 1: ops. 4, 80, 104 (AM).

Family Golfingiidae Stephen & Edmonds, 1972

Small to large-sized worms (max 200 mm); introvert length similar to trunk length or shorter. Tentacles encircling mouth. Hooks, if present, simple, scattered, not sharply curved and often deciduous. Trunk wall externally smooth or covered with small papillae. The family is well represented in the deep sea (> 2000 m) with 36 species and six genera (Saiz Salinas et al. 2018). In Australian waters six species from two genera (Golfingia and Nephasoma) have been reported (http://www.ala.org.au). We report at least seven species from four genera.

Golfingiidae gen. sp.

Records

1 specimen. Suppl. material 1. op. 42 (AM).

Golfingia sp.

Diagnosis

Small to medium worms (usually < 30 mm). Introvert hooks, if present, small (< 40 μm); nephridial pores anterior to anus. Body wall smooth.

Remarks

Further identification is uncertain.

Records

6 specimens. Suppl. material 1: ops. 6, 30, 45, 101 (AM).

Golfingia (Golfingia) muricaudata (Southern, 1913)

Fig. 35B, C

Diagnosis

Presence of a distinctive caudal appendage (< ~ 30% of the trunk length); introvert shorter than trunk.

Records

34 specimens. Suppl. material 1: ops. 11, 22, 35, 43, 44, 65, 70, 86, 99 (AM).

Phascolion spp.

Diagnosis

Small to medium-sized worms (< ~ 50 mm), commonly inhabiting abandoned gastropod or scaphopod shells, polychaete tubes or foraminiferan tests. Trunk usually with unique holdfast papillae. Single nephridium, usually located posterior to the anus.

Remarks

Specimens are not identified to species.

Records

25 specimens. Suppl. material 1: ops. 31, 44, 56, 100, 121, 128 (AM).

Phascolion (Montuga) lutense Selenka, 1885

Fig. 35E

Diagnosis

Trunk smooth, except for densely packed papillae at the anterior end. No holdfast papillae. Trunk with characteristic grey anterior ‘cap’. Inhabiting soft clay tubes instead of hard shells.

Remarks

This is the first report of this species from Australian waters.

Records

4 specimens. Suppl. material 1: ops. 6, 67 (AM).

Phascolion (Isomya) cf. hedraeum Selenka & De Man, 1883

Diagnosis

Inhabiting sediment tube; round holdfast papillae with hardened borders anteriorly.

Records

1 specimen. Suppl. material 1: op. 89 (AM).

Onchnesoma sp.

Diagnosis

Small worms (< 10 mm). Introvert always longer than trunk. Anus on distal end of introvert. No introvert hooks. Single nephridium.

Remarks

Specimens are not identified to species. This is the first report of this genus from Australian waters.

Records

4 specimens. Suppl. material 1: ops. 31, 119 (AM).

Thysanocardia cf. catharinae (Grube, 1868)

Diagnosis

Up to 70 mm trunk length. Complex tentacular arrangement in ‘festoons’ (double rows extending along the distal introvert) and around the nuchal organ. Tentacles arranged in 14–16 festoons, each with < 40 tentacles.

Remarks

This is the first record of this genus from Australian waters.

Records

9 specimens. Suppl. material 1: ops. 80, 104 (AM).

Family Phascolosomatidae Stephen & Edmonds, 1972

Small to medium-sized worms. Tentacles surrounding the nuchal organ in a semi-circle. Introvert hooks usually present, recurved and organized in circles, typically with internal structures visible under transmitted light. Four introvert retractor muscles. Two genera and eight species are known from the deep sea (> 2000 m) (Saiz Salinas et al. 2018). In Australian waters, 15 species from two genera (Apionsoma and Phascolosoma) have been reported.

? Apionsoma sp.

Diagnosis

Small specimens (usually > 20 mm) with an introvert much longer than trunk. Introvert hooks arranged in rings with accessory basal spinelets. Nephridia bilobed.

Remarks

Identification is uncertain.

Records

15 specimens. Suppl. material 1: po. 6 (AM).

Analysis of annelid biodiversity

We report > 6000 annelid specimens (box core – 57 specimens, Brenke sledge – 2481, beam trawl – 3470) (Suppl. material 1). Of these, 4714 specimens (78.5%) represented 214 putative species from 50 families. The remaining 1294 specimens were too small or damaged to be assigned to a morphospecies at this time and they were assigned to a family or genus only. Small-bodied representatives of eight families, namely Acrocirridae, Cirratulidae, Euphrosinidae, Fabriciidae, Flabelligeridae, Goniadidae, Lacydoniidae, and Pilargidae, were found exclusively in Brenke sledge and box core material. Only 27 species (624 specimens, 13% of the specimens identifiable to morphospecies) were attributed to known taxa and given valid names, while the remaining 187 (4090 specimens, 87% of the specimens identifiable to morphospecies) were determined in open nomenclature following Sigovini et al. (2016) and were assigned qualifiers sp. nov. (55 species, including six already described from this material), cf. (45), or sp. (87) (Suppl. material 1). The 55 species confidently confirmed by taxonomic experts in the respective families to be new (sp. nov.) will be formally described in the near future. The remaining 132 species (or some of them) assigned cf. and sp. qualifiers may also be new to science; however, the specimens require further study (e.g., molecular data or scanning electron microscopy) to determine the status with confidence. Preliminary data also suggest that complete specimens from the families Eunicidae, Orbiniidae, and Terebellidae could be described within three new genera at a later date. Thus, the material can contain a minimum of 55 species new to science and at least three new genera.

Sixteen of the species collected from the eastern abyss were also reported from stations at the GAB. Of these 16, seven (Nereididae: Nicon maculata, Polynoidae: Bathyeliasona nigra, Eunoe abyssorum, Aphroditidae: Aphrodita goolmarris, Laetmonice benthaliana, Laetmonice yarramba, Goniadidae: Bathyglycinde profunda) were named species. The remaining nine undescribed species were confirmed by specialists who had compared both sets of material (restricted to the families Ampharetidae, Aphroditidae, Fabriciidae, Nereididae, Onuphidae, Polynoidae, and Sabellidae).

The most species-rich family was the Polynoidae (> 17 species) followed by Onuphidae (13 species), Serpulidae (12 species), Acrocirridae (10 species) and Maldanidae (10 species). Total species richness of annelids was similar between lower bathyal and abyssal depths (163 and 160 species respectively) and decreased to mid-bathyal depths (70 species). Total species richness was highest at Bass Strait (90 species), Byron Bay (82 species) and Jervis MP (81 species) whilst the Moreton Bay (30 species) and off Fraser Island had the lowest richness (41 species).

Discussion

This is the first comprehensive report of annelids from the eastern Australian lower bathyal and abyssal region. Our results indicate a higher number of families and species (50 and 214 respectively) than was reported from the recent deep-water survey of the Great Australian Bight (GAB) at 200 to 5000 m depth (42 annelid families, 179 species) (MacIntosh et al. 2018), and from the south western continental margin of Western Australia (12 families, 57 species) where samples were collected from lower shelf and upper bathyal depths at 100, 500, and 1000 m depth contours (Poore et al. 2015). Indeed, more specimens were collected from eastern Australia (Annelida n > 6000) than from the GAB (Annelida n = 2364) and western Australia (polychaete n = 660) combined. Unfortunately, these results are not directly comparable as different sampling gear was used in each study (eastern Australia 54 successful samples using box core, Brenke sledge and beam trawl; GAB 304 samples using eight gear types including beam trawl, benthic sledge, multicorer and ROV pushcore; western Australia 135 grab samples). Our study presents the largest dataset to date on deep-water annelids from Australia. As a result, it adds important records of occurrence and distribution of species and provides a springboard for future taxonomic studies.

Prior to the 2017 survey, only eight annelid species were described from the eastern continental margin of Australia from below 1000 m and only two of them (Parapolyeunoa flynni and Aglaophamus profundus) from the targeted depths in this study, below 2000 m (Table 1). The high proportion of undescribed (new to science) species, at least 55 and up to 187 species (< 87% of specimens identified to morphospecies), found in this study is unsurprising given that this was the first systematic survey of the region and so few annelids had been previously described from abyssal depths in Australia. The high number of new annelid species is typical for previously un-sampled deep-sea environments. Recent deep-sea investigations have reported varying percentages of annelid (polychaetes) species new to science, for example, 33% in abyssal SE Atlantic (Böggemann 2009), ~ 40% in NE Pacific (Méndez 2006), > 40% from the slope to upper trench in NW Pacific (Alalykina 2018), 55% on the shelf/slope of the Southern Ocean (Neal et al. 2018), ~ 70% in SW Atlantic (Neal et al. 2020), and even up to 90% (higher than in the present study) in the abyssal Pacific (Glover et al. 2002).

While so far only six species have been formally described from this survey material (pectinariids Petta investigatoris and P. williamsonae, sabellariids Gesaia csiro and Phalacrostemma timoharai and melinnids Melinnopsis gardelli and Melinnopsis chadwicki) taxonomic studies are ongoing. The 24 new species currently being described by co-authors of this study include one acoetid of the genus Panthalis (Murray), one glycerid (Böggemann), two lumbrinerids (Borisova and Budaeva), four maldanids of the genera Boguea, Chirimia and Notoproctus (Kongsrud), four onuphids of the genus Nothria (Paxton and Budaeva), four serpulids of the genera Bathyvermilia, Hyalopomatus and Spirodiscus (Kupriyanova), four sphaerodorids of the genera Geminofilum and Sphaerephesia (Capa), and four scalibregmatids of the genera Asclerocheilus, Oligobregma, Pseudoscalibregma and Scalibregmides (Blake). At least one new species of Siboglinidae (genus Osedax) and several species of Dorvilleidae (genus Ophryotrocha) were recovered from a whale carcass collected from op. 100 (Georgieva). It is anticipated that the large international effort in species identification by taxonomic specialists in each annelid group will eventually result in the description of all the new annelids from the expedition. Once this taxonomic work is complete, we can test relevant biogeographical hypotheses, in particular we will be able to address the question of species connectivity along the eastern continental margin across to the deep the GAB.

Sixteen morphospecies reported from this study were also recorded from the GAB in southern Australia (MacIntosh et al. 2018) suggesting that at least some species ranges can span across both the eastern and southern margin of Australia. This agrees with the traditional view that deep-sea species have larger geographical ranges than shallow-water species (Ekman 1953), a view that is supported by some recent studies (e.g., McClain and Hardy 2010; Higgs and Attrill 2015). Indeed, geographical range is generally thought to increase with increasing depth (e.g., Etter and Rex 1990; Allen and Sanders 1996). Depth has been long known to have a strong structuring influence on deep-sea species distributions (e.g., Ekman 1953; France and Kocher 1996; Zardus et al. 2006). High species turnover occurs between the shelf break and 1000 m marking a transition between shelf and slope fauna, and between 2000 and 3000 m the transition between slope and abyssal fauna (Carney 2005; Brown and Thatje 2014) and thus, boundaries of biogeographical provinces are typically delineated by depth. Modelling shows that at lower bathyal depths (800–3000 m) the southern Australian fauna is distinct from the eastern fauna, while at abyssal depths (3500–6500 m) species are distributed across the Indian Ocean to the eastern margin of Australia (UNESCO 2009; Watling et al. 2013). Thus, we expect abyssal, but not bathyal fauna, to range from eastern Australia to the GAB. Interestingly, when these biogeographical hypotheses are tested against fine-resolution data from ophiuroids, the bathyal zone fauna (0–2000 m) is continuous from southeastern to southwestern Australia (O’Hara et al. 2011) suggesting the predicted biogeographical provinces around Australia may not hold true for all taxa.

Existing molecular data on the annelid material collected during ‘Sampling the Abyss’ voyage support annelid species connectivity along the eastern Australian margin from southern Queensland to Tasmania. Analysis on DNA sequence data confirmed that specimens of pectinariid Petta investigatoris were collected < 491 km apart from Bass Strait to Jervis MP, NSW (Zhang et al. 2019) and the ampharetids Melinnopsis gardelli occurred at Freycinet MP and Coral Sea MP (distance 2064 km), while Melinnopsis chadwicki was found from the Hunter Marine Park to the Coral Sea Marine Park (996 km) (Gunton et al. 2020). The latter two melinnid species were found to have distinct bathymetric ranges: M. gardelli was recorded from 2520–2821 m and M. chadwicki from 1006–1257 m depth (Gunton et al. 2020). While annelid species are found within distinct depth ranges along the eastern Australian margin, whether these species ranges extend to the GAB is less clear due to lack of genetic studies.

From 16 morphospecies reported from the present study and the GAB, the two nominal morphospecies with the widest reported distributions in our data were the aphroditid Laetmonice yarramba Hutchings & McRae, 1993 and the goniadid Bathyglycinde profunda (Hartman and Fauchald 1971). These species were reported from both the northernmost (Coral Sea MP and off Fraser Island) and southernmost (Flinders MP and Freycinet MP) stations.

Laetmonice yarramba, originally described from 60–102 m off the coast of NSW, was reported across a distance of more than 1800 km from Freycinet MP (op. 4) to off Fraser Island (op. 115) with a total depth range of 3868 m (Suppl. material 1). This species was also recorded from 21 stations at the GAB from 189–3884 m (MacIntosh et al. 2018). While the geographic range from eastern Australia to the GAB is likely, the apparent broad bathymetric range from shelf to abyssal depths (60–3884 m) requires further investigation as this putative depth range would extend vertically across shelf, slope and abyssal faunal zones. Genetic evidence suggests some deep-water annelids have ‘broad’ (> 2500 m sensu Glazier and Etter 2014) depth ranges, e.g., the maldanid Nicomache lokii range 3668 m (Eilertsen et al. 2018), but bathymetric ranges of other deep-sea annelid species are more restricted e.g., 1300 m for the spionid Laonice weddellia (see Brasier et al. 2017). According to the census of abyssal polychaetes (> 2000 m), ~ 62–78% of species within a family had bathymetric ranges smaller than 1000 m (Paterson et al. 2009). Thus, the broad (> 3000 m) depth range of L. yarramba along with its reported wide geographic range and morphological variability (Hutchings and McRae 1993 and herein) suggests that the nominal species is a species complex and molecular data are needed to test this hypothesis.

Bathyglycinde profunda (Hartman & Fauchald, 1971) was originally described from the equatorial region off northeast South America (4825 m). This taxon in the present study was recorded from Flinders MP to Coral Sea MP (ops. 16 to 134), a distance of 1920 km and depth range 2093–4280 m. The same morphospecies has been also reported from northwest Atlantic Ocean (2862–5023 m) by Hartman and Fauchald (1971), from the outer shelf and continental slope off Brazil (325–508 m) by Rizzo and Amaral (2004), from the Clarion-Clipperton Zone in the Pacific by Janssen et al. (2019) and the GAB by MacIntosh et al. (2018). Whether the species in this study is the same as the one described from South America requires further genetic evidence as distribution of such a species would fall into multiple biogeographical provinces as defined by UNESCO (2009) and Watling et al. (2013). Recent molecular studies have provided contrasting results on annelid species range size previously based on morphological data. Genetic evidence exists for abyssal annelids with broad geographic ranges (4150 km, see Schüller and Hutchings 2012) and as well as for bipolar distributions of deep-sea species from chemosynthetic environments (16000 km, Georgieva et al. 2015; Eilertsen et al. 2018). Yet, some deep-sea annelids previously believed to have broad distributions based on morphology alone were found to include multiple species as a result of molecular studies (e.g., Stiller et al. 2013). As with L. yarramba, the distribution of B. profunda is likely to be more restricted than morphology suggests, and molecular data are needed to test these broad species ranges.

Once more data on the extent of annelid species range sizes from this study become available, we will be able to not only address the degree of genetic connectivity between the GAB and eastern Australian margin, but also to examine underlying biogeographical the patterns across annelids. In particular, we will be able to address the question whether the tropical to temperate transition between 30–40°S reported for deep-sea ophiuroids (O’Hara et al. 2011, 2014) and megafauna collected during the same cruise ‘Sampling the Abyss’ (O’Hara et al. 2020b) is also observed in annelid fauna.

Conclusions

We report 214 annelid species from the eastern Australian margin, at least 55 of which are new to science. Prior to 2017, only two annelid species were described from the region below 2000 m, consequently this work vastly increases our knowledge of deep-water annelids and provides critical baseline data on an important group of benthic invertebrates from a virtually unknown region of the world’s ocean. This is important as comprehensive taxonomically-consistent deep-water datasets that cover large areas in Australia are rare (O’Hara 2008; Alderslade et al. 2014; Althaus et al. 2017; MacIntosh et al. 2018; Williams et al. 2018; Farrelly and Ahyong 2019; Ekins et al. 2020; O’Hara et al. 2020a, b, c), in particular for smaller benthic invertebrates (Poore et al. 2015), which highlights the uniqueness of this dataset.

The strong feature of this study is that specific annelids groups were sent to taxonomic specialists around the world giving us more confidence in the species identifications. Furthermore, because all material from this study is deposited or will soon be in properly curated museum collections (AM, MV, NHMUK) and a significant proportion has been preserved for molecular studies, these valuable samples are easily accessible and can be used for answering important questions on taxonomy and species ranges along the eastern Australian margin including seven deep-water Marine Parks.

Acknowledgements

The authors wish to thank the CSIRO Marine National Facility (MNF) for its support in the form of sea time on RV ‘Investigator’, support personnel, scientific equipment and data management. All data and samples acquired on the voyage are made publicly available in accordance with MNF Policy. We thank Tim O’Hara for leading the expedition, Karen Gowlett-Holmes for the images of live specimens taken on board the expedition, and all the scientific staff and crew who participated in voyage IN2017_V03. Project funding was provided by the Marine Biodiversity Hub, supported through the Australian Government’s National Environmental Science Program (NESP). Many thanks to collection staff at the Australian Museum for handling the many loan requests. Many thanks to Susan Sweller, Aru Bopiah, Michael Bojko and Nika Mikhin for help with editing images. This project was funded by the Chadwick Biodiversity Fellowship from the Australian Museum, Deep-Sea Biology Society ‘Dive Deeper’ grant, Linnean Society of New South Wales Joyce Vickery grant to LG, as well as Australian Biological Resources Study (ABRS) grant RG18–21 to EK, PH, RW and CG. We also thank Rickard Gardell for his generous private donation to the Australian Museum.

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Appendix 1

Note added in proofs: Microphthalmidae Hartmann-Schröder, 1971 was accepted as a valid family in the World Register of Marine Species on 27 June 2020. [Read G, Fauchald K [Ed.] (2021) World Polychaeta database. Microphthalmidae Hartmann-Schröder, 1971. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=322549 on 2021-01-25].

Supplementary materials

Supplementary material 1

Annelid identifications from IN2017_V03_Darwin Core format

Laetitia M. Gunton, Elena K. Kupriyanova, Tom Alvestad, Lynda Avery, James A. Blake, Olga Biriukova, Markus Böggemann, Polina Borisova, Nataliya Budaeva, Ingo Burghardt, Maria Capa, Magdalena N. Georgieva, Christopher J. Glasby, Pan-Wen Hsueh, Pat Hutchings, Naoto Jimi, Jon A. Kongsrud, Joachim Langeneck, Karin Meißner, Anna Murray, Mark Nikolic, Hannelore Paxton, Dino Ramos, Anja Schulze, Robert Sobczyk, Charlotte Watson, Helena Wiklund, Robin S. Wilson, Anna Zhadan, Jinghuai Zhang

Data type: occurrences

Explanation note: species occurrences along with museum registration numbers.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

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Supplementary material 2

Spionidae characters

Karin Meißner

Data type: morphological

Explanation note: Comparison of morphological characters of Prionospio species from IN2017_V03.

Download file (10.58 kb)