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Research Article
Three new deep-sea species of Marphysa (Annelida, Eunicida, Eunicidae) from Papua New Guinea (Bismarck and Solomon seas)
expand article infoNicolas Lavesque‡, Guillemine Daffe‡, Christopher Glasby§|, Stéphane Hourdez, Pat Hutchings|#
‡ Université de Bordeaux, Arcachon, France
§ Museum and Art Gallery Northern Territory, Darwin, Australia
| Australian Museum Research Institute, Australian Museum, Sydney, Australia
¶ Sorbonne Université, Banyuls, France
# Macquarie University, North Ryde, Australia

Corresponding author: Nicolas Lavesque ( nicolas.lavesque@u-bordeaux.fr )

Academic editor: Andrew Davinack
© 2022 Nicolas Lavesque, Guillemine Daffe, Christopher Glasby, Stéphane Hourdez, Pat Hutchings.
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: Lavesque N, Daffe G, Glasby C, Hourdez S, Hutchings P (2022) Three new deep-sea species of Marphysa (Annelida, Eunicida, Eunicidae) from Papua New Guinea (Bismarck and Solomon seas). ZooKeys 1122: 81-105. https://doi.org/10.3897/zookeys.1122.89990
ZooBank: urn:lsid:zoobank.org:pub:0DF18698-843E-4F78-9360-58A46B2010F7
Open Access

Abstract

Three new species of Marphysa Quatrefages, 1866, Marphysa banana sp. nov., Marphysa papuaensis sp. nov., and Marphysa zanolae sp. nov. are described from deep-sea sunken vegetation off Papua New Guinea, using both morphology and molecular data (for two species). With the presence of compound spinigers only and the branchiae present over many chaetigers, Marphysa banana sp. nov. belongs to the group B2. This species is characterised by the presence of eyes, the presence of branchiae starting from chaetiger 20, and by the presence of three types of pectinate chaetae and bidentate subacicular hooks starting from chaetigers 13–52. With the presence of compound falcigers only and the branchiae restricted to a short anterior region, Marphysa papuaensis sp. nov. belongs to the group C1. This species has a bilobed prostomium but no eyes, has branchiae from chaetigers 7 to 14–16 with up to 16 filaments. Marphysa papuaensis sp. nov. is also characterised by the presence of bidentate subacicular hooks from chaetiger 20 and by a single type of pectinate chaetae. Finally, Marphysa zanolae sp. nov. belongs to the group C2, with the presence of compound falcigers only and the branchiae present over many chaetigers. This species is characterised by the absence of eyes, by the presence of branchiae with a single long filament starting from chaetiger 31, by unidentate subacicular hooks starting from chaetiger 28 and finally by one type of pectinate chaetae with very long outer teeth.

Keywords

Bloodworm, COI, deep sea, Marphysa, morphology, polychaete, sunken vegetation, taxonomy

Introduction

Situated in the Coral Triangle, Papua New Guinea is considered a marine biodiversity hotspot and shows a high level of endemism. Although the terrestrial and shallow water fauna is well known, the deep-sea fauna has rarely been studied (Pante et al. 2012). Indeed, the historical expeditions of ‘Siboga’ and the HMS ‘Challenger’, which both sampled the deep sea, did not pay much attention to this area, and only the ‘Galathea’ and RV ‘Vityaz’ expeditions carried out benthic sampling in the deep sea (hadal zone) (Pante et al. 2012; Corbari et al. 2019). Only a few taxonomic studies on polychaetes have been conducted in the region, in coastal habitats (Dahlgren 1996; Rouse 1996; Britayev et al. 1999; Rouse 2012) and hydrothermal vents (Watson 2001; Reuscher et al. 2011) but data on other types of deep-sea habitats are still lacking. Between 2010–2014, the Muséum National d’Histoire Naturelle (MNHN) and the Institut de Recherche pour le Développement (IRD), in collaboration with the University of Papua New Guinea (UPNG) launched four sampling campaigns (BIOPAPUA 2010, PAPUA NIUGINI 2012 MADEEP, and KAVIENG 2014) aiming to explore the deep-sea biodiversity of this region, especially in the Bismarck and Solomon seas (Pante et al. 2012; Corbari et al. 2019).

Marphysa Quatrefages, 1866 is a very speciose genus with 83 accepted species (Read and Fauchald 2022), commonly found from intertidal shores to shallow waters (Glasby and Hutchings 2010). As far as we know, and unlike Eunice the other species-rich genus of the family, Marphysa species are never found in the deep sea. Except for the non-indigenous species Marphysa victori Lavesque, Daffe, Bonifácio & Hutchings, 2017 (Lavesque et al. 2020), most of the species show restricted distributions (Hutchings and Kupriyanova 2018; Lavesque et al. 2019) and are often very restricted to particular habitats (Hutchings and Karageorgopoulos 2003; Glasby and Hutchings 2010; Zanol et al. 2016; Lavesque et al. 2019). Following Fauchald (1970) and Glasby and Hutchings (2010), species of the genus Marphysa can be separated into five artificial groups based on the type of compound chaetae: no compound chaetae present (Group A), only compound spinigers present (Group B), only compound falcigers present (Group C), both compound spinigers and falcigers present (Group D), and compound spinigers only anteriorly and posterior segments only with simple limbate chaetae (Group E). Finally, each group can also then be divided into species having branchiae present over a short anterior region (subdivision 1) or branchiae present over many chaetigers (subdivision 2).

Until now, 15 species of Marphysa have been described from the Central Indo-Pacific Realm (sensu Spalding et al. 2007), two species belonging to Group A (M. fijiensis Molina-Acevedo & Idris, 2021 and M. moribidii Idris, Hutchings & Arshad, 2014), nine species to Group B (M. hongkongensa Wang, Zhang & Qiu, 2018; M. iloiloensis Glasby, Mandario, Burghardt, Kupriyanova, Gunton & Hutchings, 2019; M. maxidenticulata Liu, Hutchings & Kupriyanova, 2018; M. mullawa Hutchings & Karageorgopoulos, 2003; M. multipectinata Liu, Hutchings & Sun, 2017; M. orientalis Treadwell, 1936; M. tribranchiata Liu, Hutchings & Sun, 2017; M. tripectinata Liu, Hutchings & Sun, 2017, and M. bulla Liu, Hutchings & Kupriyanova, 2018, which was recently synonymised with M. victori Lavesque, Daffe, Bonifácio & Hutchings, 2017 (Lavesque et al. 2020); two species belong to Group C (M. bernardi Rullier, 1972 and M. soembaensis Augener, 1933); and only one species belongs to each of groups D (M. digitibranchia Hoagland, 1920) and E (M. fauchaldi Glasby & Hutchings, 2010).

In this study, three new deep-sea species belonging to groups B2 (Marphysa banana sp. nov.), C1 (Marphysa papuaensis sp. nov.), and C2 (Marphysa zanolae sp. nov.) are described using both morphology and molecular data (for two of them). The type specimens were found in deep-sea sunken vegetation (decaying wood or cultivated plants leaves). It is not surprising, as this region is known to accumulate large quantities of decomposing vegetation, transiting from tropical forests to marine canyons (Pante et al. 2012), hosting original and diverse fauna (Samadi et al. 2007).

Materials and methods

Sampling and morphological analyses

Specimens were collected by beam trawl during the MADEEP cruise (see https://expeditions.mnhn.fr/campaign/madeep) and the KAVIENG cruise (https://expeditions.mnhn.fr/campaign/kavieng2014) in May–September 2014, in the Solomon Sea (Fig. 1). All material was sorted on board RV ‘Alis’ and fixed in 80% ethanol. A few parapodia were removed from several specimens for molecular analysis. Specimens were examined under a Nikon SMZ25 stereomicroscope and a Nikon Eclipse Ci microscope, and photographed with a Nikon DS-Ri 2 camera. Measurements were made with the NIS-Elements Analysis software. Drawings were made from pictures using Inkscape software. Width of all specimens was obtained by measuring chaetiger 10 with parapodia. Morphological terminology is based on Fauchald (1992) for general terms, Paxton (2000) for head appendages and Molina-Acevedo and Carrera-Parra (2015) for maxillary apparatus. Terminology of pectinate chaetae follows Glasby et al. (2019), based on a previous study of Molina-Acevedo & Carrera-Parra (2017): isodont means outer teeth much longer than internal teeth, anodont means outer teeth more or less same length as internal teeth, and heterodont when one long and one short (same length as internal teeth) lateral tooth are present. The width of the pectinate blade is wide when ≥ 30 μm and narrow below this. Finally, the size of the internal teeth is long when they measure 12 µm or more and thick when are 2 µm or more, below these values the teeth are defined as short and slender, respectively.

Figure 1. 

Sampling sites of type material of Marphysa banana sp. nov. (black circle), Marphysa papuaensis sp. nov. (black star), and Marphysa zanolae sp. nov. (black square) in Papua New Guinea.

Some parapodia along the body were removed from the type material of each species (see Material examined), dehydrated in ethanol, critical point dried, covered with 20 nm of gold, examined under the scanning electron microscope (JEOL JSM 6480LA) and imaged with a secondary detector at Macquarie University, Sydney, Australia.

The studied material is deposited at the Muséum National d’Histoire Naturelle, Paris (MNHN) and the Australian Museum, Sydney (AM). Additional material is lodged in the collection housed at the Arcachon Marine Station (SMA).

Molecular data and analyses

Extraction of DNA was done with ISOLATE II Genomic DNA kit (BIOLINE) following protocol supplied by the manufacturers. Approximately 600 bp of COI (cytochrome c oxidase subunit I) gene was amplified, using primers polyLCO and polyHCO COI (Carr et al. 2011). PCR (Polymerase Chain Reaction) was performed with Taq DNA Polymerase QIAGEN Kit in 20 μL mixtures containing: 2 μL of 10X CoralLoad PCR Buffer (final concentration of 1X), 1.5 μL of MgCl2 (25 Mm) solution, 1.5 μL of PCR nucleotide mix (final concentration of 0.2 mM each dNTP), 0.4 μl of each primer (final concentration of 0.2 μM), 0.1 μl of Taq DNA Polymerase (5U/μl), 1 μl template DNA and 13.1 μL of nuclease-free water. The temperature profile was as follows 94 °C / 60 s – (94 °C / 40 s – 45 °C / 40 s – 72 °C / 60 s)*5 cycles – (94 °C / 40 s – 51 °C/ 40 s – 72 °C / 60 s)*35 cycles – 72 °C / 300 s – 4 °C. PCR success was verified by electrophoresis in a 1% p/v agarose gel stained with Gelred. Amplified products were sent to Macrogen Company to obtain sequences, using the same set of primers as used for PCR.

Fifty-nine COI sequences were downloaded from GenBank or obtained during this study, fifty-six COI sequences of Marphysa species and three outgroup species from closely related genera in the family Eunicidae (Table 1). During this study, one COI sequence was obtained both for Marphysa papuaensis sp. nov. and Marphysa zanolae sp. nov., but we failed to obtain a sequence for Marphysa banana sp. nov.

Table 1.
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Terminal taxa used in the molecular part of the study (COI gene), with type localities, collection localities, GenBank accession numbers, and references.

Species Type locality Collection locality GenBank accession number Reference
Eunice cf. violaceomaculata Tortugas, Caribbean Carrie Bow Cay, Belize GQ497542 Zanol et al. 2010
Palola viridis Samoa, Pacific Ocean Kosrae, Micronesia GQ497556 Zanol et al. 2010
Leodice rubra Saint Thomas, Caribbean Ceara, Brazil GQ497528 Zanol et al. 2010
M. aegypti Suez Canal, Egypt Suez Canal, Egypt MF196969 Elgetany et al. 2018
M. bifurcata WA, Australia Qld, Australia KX172177 Zanol et al. 2016
M. bifurcata WA, Australia Qld, Australia KX172178 Zanol et al. 2016
M. brevitentaculata Tobago Quintana Roo, Mexico GQ497548 Zanol et al. 2010
M. californica California, USA California, USA GQ497552 Zanol et al. 2010
M. disjuncta California, USA California, USA GQ497549 Zanol et al. 2010
M. chirigota Bay of Cadiz, Spain Bay of Cadiz, Spain MN816442 Martin et al. 2020
M. chirigota Bay of Cadiz, Spain Bay of Cadiz, Spain MN816443 Martin et al. 2020
M. chirigota Bay of Cadiz, Spain Bay of Cadiz, Spain MN816444 Martin et al. 2020
M. fauchaldi NT, Australia NT, Australia KX172165 Zanol et al. 2016
M. gaditana Bay of Cadiz, Spain Bay of Cadiz, Spain MN816441 Martin et al. 2020
M. hongkongensa Hong Kong Hong Kong MH598525 Wang et al. 2018
M. hongkongensa Hong Kong Hong Kong MH598526 Wang et al. 2018
M. iloiloensis Iloilo, Philippines Tigbauan, Philippines MN106279 Glasby et al. 2019
M. iloiloensis Iloilo, Philippines Tigbauan, Philippines MN106280 Glasby et al. 2019
M. iloiloensis Iloilo, Philippines Tigbauan, Philippines MN106281 Glasby et al. 2019
M. kristiani NSW, Australia NSW, Australia KX172160 Zanol et al. 2016
M. kristiani NSW, Australia NSW, Australia KX172161 Zanol et al. 2016
M. kristiani NSW, Australia NSW, Australia KX172162 Zanol et al. 2016
M. kristiani NSW, Australia NSW, Australia KX172158 Zanol et al. 2016
M. madrasi Chennai, India Chennai, India MT813506 Hutchings et al. 2020
M. madrasi Chennai, India Chennai, India MT813507 Hutchings et al. 2020
M. mossambica Mozambique Iloilo, Philippines KX172164 Zanol et al. 2016
M. mullawa Qld, Australia NSW, Australia KX172166 Zanol et al. 2016
M. mullawa Qld, Australia NSW, Australia KX172167 Zanol et al. 2016
M. mullawa Qld, Australia NSW, Australia KX172168 Zanol et al. 2016
M. mullawa Qld, Australia NSW, Australia KX172176 Zanol et al. 2016
M. papuaensis sp. nov. Papua New Guinea Papua New Guinea OP184050 This study
M. pseudosessiloa NSW, Australia NSW, Australia KY605405 Zanol et al. 2010
M. pseudosessiloa NSW, Australia NSW, Australia KY605406 Zanol et al. 2010
M. regalis Bermuda Ceara, Brazil GQ497562 Zanol et al. 2010
M. sanguinea Devon, UK Callot Island, France GQ497547 Zanol et al. 2010
M. sanguinea Devon, UK Cornwall, UK MK541904 Lavesque et al. 2019
M. sanguinea Devon, UK Arcachon Bay, France MK950853 Lavesque et al. 2019
M. sanguinea Devon, UK Brest, France MK967470 Lavesque et al. 2019
M. tripectinata Beihai, China Beihai, China MN106271 Glasby et al. 2019
M. sherlockae Durban, South Africa Strand, South Africa MT840349 Kara et al. 2020
M. sherlockae Durban, South Africa Strand, South Africa MT840350 Kara et al. 2020
M. sherlockae Durban, South Africa Strand, South Africa MT840351 Kara et al. 2020
M. tripectinata Beihai, China Beihai, China MN106272 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106273 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106274 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106275 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106276 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106277 Glasby et al. 2019
M. tripectinata Beihai, China Beihai, China MN106278 Glasby et al. 2019
M. victori Arcachon Bay, France Arcachon Bay, France MG384996 Lavesque et al. 2017
M. victori Arcachon Bay, France Arcachon Bay, France MG384997 Lavesque et al. 2017
M. victori Arcachon Bay, France Arcachon Bay, France MG384998 Lavesque et al. 2017
M. victori Arcachon Bay, France Arcachon Bay, France MG384999 Lavesque et al. 2017
M. victori Arcachon Bay, France Mangoku-ura Inlet, Japan LC467767 Abe et al. 2019
M. victori Arcachon Bay, France Sendai Bay, Japan LC467769 Abe et al. 2019
M. victori Arcachon Bay, France Ena Bay, Japan LC467772 Abe et al. 2019
M. victori Arcachon Bay, France China MT012514 Lavesque et al. 2020
M. viridis Florida, USA Ceara, Brazil GQ497553 Zanol et al. 2010
M. zanolae sp. nov. Papua New Guinea Papua New Guinea OP184049 This study

All COI sequences were aligned in Geneious Prime 2019.0.4 using the MUSCLE plugin and default settings. The AIC and BIC tests in jModeltest 2.2.10 (Darriba et al. 2012) were used to select the GTR + I + G model of molecular evolution as the best evolutionary model for the COI gene alignment. The phylogenetic analysis was performed in MrBayes v. 3.2.6 (Ronquist and Huelsenbeck 2003). The analysis was run for 10 million generations (sampled every 1000), 25% of the generations were discarded as burn-in and the standard deviation of split frequencies decreased below 0.01. FigTree v. 1.4.4 (Rambaut 2007) was used to visualise the majority-rule consensus tree displaying all nodes with a posterior probability > 0.5. Pair-wise Kimura 2-parameter (K2P) genetic distance was performed using MEGA v. 7.0.26.

Taxonomic account

Family Eunicidae Berthold, 1827

Marphysa Quatrefages, 1866

Type species

Nereis sanguinea Montagu, 1813.

Marphysa banana sp. nov.

https://zoobank.org/36B09BD6-2080-4266-9945-98EA1CA40913
Figs 2, 3, 4

Material examined

Holotype : MNHN-IA-2015-1608, complete. Paratypes: AM W.53773, complete; AM W.53774, complete, some parapodia mounted for SEM; MNHN-IA-2021-725, anterior part only. All material collected from South Pacific Ocean, Papua New Guinea, New Ireland, CP4254, -2.483°S, 150.66°E, depth 273–324 m, April 2014.

Description

(based on holotype, with variation in parentheses for paratypes). Preserved specimens strongly iridescent (Fig. 2A, C, D), ~ 230 (220) chaetigers, 112 mm (71–157) long, 3.3 mm (2.1–5.4) width at chaetiger 10, excluding parapodia. Body elongated and tapered gradually at posterior end (Fig. 2B).

Figure 2. 

Marphysa banana sp. nov. holotype MNHN-IA-2015-1608 (B) paratype MNHN-IA-2021-725 (A, C–E): A anterior end, lateral view B pygidium, ventral view C anterior end, dorsal view D anterior end, frontal view E anterior chaetigers, dorso-lateral view.

Prostomium rounded anteriorly with two dorsoventrally flattened buccal lips an anterior notch between them, notch more visible ventrally (Fig. 2A, C, D). Two palps and three antennae slender and tapering, arranged in an arc on posterior margin of prostomium. Antennae more or less smooth, of equal length, longer than palps (same size), shorter (same size) than prostomium (Fig. 2C). Eyes present, one pair, brownish, very faint, present at posterior base between palps and lateral antennae. First peristomial ring ~ 3× longer than second one dorsally (Fig. 2C).

Maxillary apparatus (Fig. 3D, E) partially everted in holotype or paratypes. Formula as follows: MF = 1+1, 5+5, 6+0, 4+9, 1+1. MI ~ 2× longer than maxillary carrier, rectangular anteriorly, triangular posteriorly, with a pair of rounded wings situated at posterolateral margins. MI forceps-like, without attachment lamellae, sub-right-angle falcal arch. Closing system ~ 4× shorter than MI. Ligament between MI and MII golden. MII without attachment lamella, teeth triangular, distributed in less than half of plate length. Ligament between MII and MIII absent (or not sclerotized). MIII, single, longer than left MIV, curved, with equal-sized triangular teeth; short attachment lamella situated in the centre of posterior edge of maxilla, thin, dark. Left MIV short (less than half the size of right MIV) with wide, rounded base, left-most teeth longer than right-most ones; attachment lamella dark, semi-circular. Right MIV long, with teeth triangular, decreasing in size posteriorly; attachment lamella wide, semi-circular, dark. MV, paired, as long as high, with a dorsal curved tooth. Mandibles light brown, concentric stripes not visible; longer than MI; cutting plates whitish, without dorsal teeth (Fig. 3E).

Figure 3. 

Marphysa banana sp. nov. holotype MNHN-IA-2015-1608 (A–C, F–G), paratype MNHN- IA-2021-725 (D, E): A parapodia from anterior chaetiger (chaetiger 29) B parapodia from mid-body (chaetiger 100) C parapodia from posterior chaetiger (chaetiger 190) D maxilla, dorsal view E mandibles, dorsal view F subacicular hook from mid-body (chaetiger 100) G spiniger from chaetiger 29. Abbreviations: MI to MV, maxillae I to V; Ac, aciculae; Bf, branchial filament; Cp, cutting plate; Mc, maxillary carrier; Nc, notopodial cirri; Sah, subacicular hook; Vc, ventral cirri. A–C Sah and Ac are illustrated schematically to indicate position.

First few parapodia located below middle line of body wall, but gradually positioned dorsally to approximately midline in subsequent segments (Fig. 2A). Notopodial cirri slender, tapering, slightly longer than ventral cirri, thinner posteriorly (Fig. 3A–C). Chaetal lobes comprising a low pre-chaetal lip and a globular post-chaetal lobe. Ventral cirri bluntly conical, with rounded tip, shorter than post-chaetal lobes anteriorly, thereafter slightly longer than post-chaetal lobes (Fig. 3A–C). Branchiae pectinate, commencing from chaetiger 20 (18–19) and continuing to near end, very short anteriorly, longer in medium chaetigers but not reaching mid-dorsal line; number of filaments increasing from 1–3 anteriorly to 4–6 in mid-body, decreasing to 3–4 in last several chaetigers (Figs 2E, 3A–C).

Aciculae black with paler blunt tips, approximately three or four per parapodium in anterior chaetigers, one or two per parapodium in middle chaetigers, and one per parapodium in posterior chaetigers. Supra-acicular chaetae with limbate capillaries and pectinates; capillaries present from first chaetiger to near pygidium, numbering up to 20 in anterior chaetigers (Fig. 3A–C).

Pectinate chaetae commencing from approximately chaetiger 20 to near end, three types identified. Type 1 from anterior parapodia to mid-body: isodont-narrow-slender (INS), having ~ 20 short internal teeth, each tooth prolonged by a thin filament (Fig. 4A, B). Types 2 and 3 from posterior parapodia only (Fig. 4C, D): type 2 asymmetrical, anodont-wide-thick (AWT), having ~ 10 thick internal teeth, each tooth prolonged by a thin filament; type 3 asymmetrical, anodont-wide-slender (AWS), having ~ 20 internal teeth, each tooth prolonged by a thin filament (Fig. 4D).

Figure 4. 

SEM images of pectinate chaetae of Marphysa banana sp. nov. paratype AM W.53774 A chaetiger 35 B chaetiger 67 C chaetiger 99 D chaetiger 131. Abbreviations: AWS, anodont-wide-slender; AWT, anodont-wide-thick; INS, isodont-narrow-slender; Fil, filament.

Subacicular chaetae with compound spinigers and subacicular hooks (Fig. 3F). Compound spinigers commencing from first chaetiger to near pygidium, with long, tapered blade. Subacicular hooks transparent, commencing from anterior chaetigers 43–52 (range for type material) to near end and inferior to bundle of spinigers, one per parapodium; much thinner than aciculae; subacicular hooks bidentate (Fig. 3F).

Pygidium round, with crenulated margin, dorsally positioned, with two pairs of tapering pygidial cirri attached at ventral edge, dorsal pair 2–3× length of ventral pair (Fig. 2B).

Etymology

The species name refers to the decomposing banana leaves among which all the specimens were found.

Type locality

South Pacific, Papua New Guinea, New Ireland.

Distribution

Only known from type locality.

Habitat

Between 273 and 324 m, found inside banana leaves that presumably have been entrained from river runoff via coastal waters.

Remarks

With the presence of compound spinigers only and the branchiae present over many chaetigers Marphysa banana sp. nov. belongs to the group B2, also known as the sanguinea-group Quatrefages, 1866. Among the nine species of this group occurring in the Central Indo-Pacific Realm, M. banana sp. nov. is similar to M. hongkongensa, M. iloiloensis, and M. mullawa by the presence of subacicular hooks starting from chaetigers 30–50 and the branchiae commencing from chaetigers 14–20.

However, M. banana sp. nov. differs from M. hongkongensa by the presence of pectinate chaetae starting from around chaetiger 20 instead of starting from the first few chaetigers as found for M. hongkongensa; and by the presence of three different types of pectinate chaetae instead of four types as found in M. hongkongensa. Moreover, M. banana sp. nov. has eyes whereas M. hongkongensa does not have any. The subacicular hooks of M. banana sp. nov. are bidentate while those of M. hongkongensa are unidentate and the maximum number of branchial filaments reaches six for M. banana sp. nov., while it can be ten for M. hongkongensa. Finally, M. hongkongensa lives in the lower intertidal of the Hong Kong region, while M. banana sp. nov. is a deep-sea species found inside banana leaves.

Marphysa banana sp. nov. differs from M. iloiloensis by the presence of four and nine teeth on the maxillary MIV, while M. iloiloensis has three and five teeth respectively. The two species show three different types of pectinate chaetae but not the same ones, as M. banana sp. nov. has INS, AWT and AWS with the first ones starting from chaetiger 20 while M. iloiloensis has INS, IWS and ANT, with first ones commencing from the first few chaetigers. The subacicular hooks are also different as they are bidentate for M. banana sp. nov. and unidentate for M. iloiloensis. Finally, M. iloiloensis lives in the brackish waters of the Philippines region, which is a very different habitat from the deep-sea habitat of M. banana sp. nov.

Marphysa banana sp. nov. differs from M. mullawa by the presence of pectinate chaetae starting from around chaetiger 20 instead of commencing from the first few chaetigers for M. mullawa, and the anterior chaetae numbering ~ 20 internal teeth instead of 10 for M. mullawa. The two species differ by their maxillary formulae, especially for maxillary MII (7+7 for M. mullawa, 5+5 for M. banana sp. nov.) and MIII (8+0 for M. mullawa, 6+0 for M. banana sp. nov.). Another difference concerns the shape of anterior branchiae, which are palmate for M. mullawa and pectinate for M. banana sp. nov. Once again, the two species live in very different habitats as M. mullawa is found in intertidal and shallow waters only, on mud or in seagrass beds.

Marphysa papuaensis sp. nov.

https://zoobank.org/991E2117-4921-4A7F-83B8-8F30EF6CA73C
Figs 5, 6, 7

Material examined

Holotype : MNHN-IA-2015-1559, complete, South Pacific Ocean, Papua New Guinea, New Britain, CP4264, -4.6°S, 152.4°E, depth 430–523 m, April 2014. Paratypes: MNHN-IA-2015-1415, complete, South Pacific Ocean, Papua New Guinea, New Britain, CP4337, -6.083°S, 149.316°E, depth 287–447 m, May 2014; MNHN-IA-2015-1593, anterior part only, South Pacific Ocean, Papua New Guinea, New Britain, CP4329, -6.133°S, 149.166°E, depth 250–500 m, May 2014; AM W.53770, complete (several parapodia mounted for SEM), South Pacific Ocean, Papua New Guinea, New Britain, CP4264, 4.6°S, 152.4°E, depth 430–523 m, April 2014; AM W.53771, anterior part only, mounted for SEM, South Pacific Ocean, Papua New Guinea, New Britain, CP4334, -6.116°S, 149.166°E, depth 430–620 m, May 2014; AM W.53772, complete, gravid, South Pacific Ocean, Papua New Guinea, New Britain, CP4266, -4.6166°S, 152.416°E, depth 575–616 m, April 2014.

Additional material

MNHN-IA-2015-1610, anterior part only, gravid, South Pacific Ocean, Papua New Guinea, New Ireland, CP4260, -2.9°S, 151.1°E, depth 350–847 m, April 2014; MNHN-IA-2015-1949, anterior part only, South Pacific Ocean, Papua New Guinea, New Ireland, CP4434, -2.25°S, 150.8°E, depth 1066–1200 m, August 2014; MNHN-IA-2015-1615, anterior part only, few parapodia used for molecular analysis, South Pacific Ocean, Papua New Guinea, New Hanover, CP4482, -2.683°S, 150.116°E, depth 761–825 m, September 2014.

Description

(based on holotype, with variation in parentheses for paratypes). Specimens strongly iridescent (Fig. 5B), 88 (89) chaetigers, 45 mm (41–80) long, 3.6 mm (2.5–2.8) width at chaetiger 10, excluding parapodia. Body elongated and tapered gradually at posterior end, anteriorly not flattened (Fig. 5A).

Figure 5. 

Marphysa papuaensis sp. nov. holotype MNHN-IA-2015-1559: A entire specimen, lateral view B anterior end, ventral view C anterior end, dorsal view D pygidium, lateral view.

Prostomium bilobed, with buccal lips separated by a ventral notch only (Fig. 5B). Two palps and three antennae slender and tapering, palpophores not visible, arranged in an arc on posterior margin of prostomium. Antennae more or less smooth, of equal length, slightly longer than palps and prostomium (same size) (Figs 5C, 7A). Eyes absent. First peristomial ring ~ 1.8× longer than second one dorsally (Figs 5C, 7A).

Maxillary apparatus (Fig. 6D, E) partially everted in holotype or paratypes. Formula as follows: MF = 1+1, 5(6)+5(6), 7+0, 4(3)+10(11), 1+1. MI ~ 2.5× longer than maxillary carrier, rectangular anteriorly, triangular posteriorly, with a pair of rounded wings situated at posterolateral margins. MI forceps-like, without attachment lamellae, sub-right-angle falcal arch. Closing system ~ 4–5× shorter than MI. Ligament between MI and MII rectangular, dark. MII without attachment lamella, teeth triangular, distributed in less than half of plate length. Ligament between MII and MIII absent (or not sclerotized). MIII, single, longer than left MIV, curved, with equal-sized triangular teeth; short attachment lamella situated in the centre of posterior edge of maxilla, oval, dark. Left MIV short (half the size of right MIV) with wide, rounded base, two left teeth longer than right-most ones; attachment lamella dark, semi-circular. Right MIV long, with teeth triangular, decreasing in size posteriorly; attachment lamella wide, semi-circular, dark. MV, paired, longer than wide, with a long tooth pointed ventrally, and a rounded dorsal margin (Fig. 6D). Mandibles dark with golden tips, with fine concentric stripes visible dorsally and ventrally, same size as MI; cutting plates whitish, with distinct growth rings, with three dorsal teeth (Fig. 6E).

Figure 6. 

Marphysa papuaensis sp. nov. paratypes MNHN-IA-2015-1415 (A–C, F, G), MNHN-IA-2015-1593 (D, E): A parapodia from anterior chaetiger (chaetiger 12) B parapodia from mid-body (chaetiger 36) C parapodia from posterior chaetiger (chaetiger 74) D maxilla, dorsal view E mandibles, dorsal view F compound falcigers from anterior chaetiger (chaetiger 12) G subacicular hook from mid-body (chaetiger 36). Abbreviations: MI to MV, maxillae I to V; Ac, aciculae; Cp, cutting plate; Mc, maxillary carrier; Nc, notopodial cirri; Sah, subacicular hook; Vc, ventral cirri. A–C Sah and Ac are illustrated schematically to indicate position.

Notopodial cirri very long, slender and, tapering (Fig. 6A–C), 2–3× longer than ventral cirri in all chaetigers. Pre-chaetal lobe inconspicuous. Post-chaetal lobe digitiform in the two or three first chaetigers, triangular with tapering tip from chaetiger 4, reducing in size from chaetiger 17, almost inconspicuous from chaetiger 28 (21) (Fig. 6A–C). Ventral cirri (Fig. 6A–C) bluntly conical, with slightly expanded bases and rounded tips from chaetigers 1–22, subconical and thinner thereafter. Branchiae pectinate (Figs 5C, 6A, 7A), starting from chaetiger 7 (7) and continuing for a limited number of segments, until chaetiger 16 (14); with 8–16 long filaments.

Aciculae black with paler blunt tips, 2–4 per parapodium along the body. Supra-acicular chaetae with limbate capillaries and pectinates; capillaries present from first chaetiger to near pygidium, numbering up to 20 in anterior chaetigers. Pectinate chaetae commencing from first few chaetigers to near end, one type identified as heterodont-narrow-slender (HNS; Fig. 7B, D, E), one outer tooth very long (Fig. 7E), having nine or ten short internal teeth, each tooth prolonged by a thin filament.

Figure 7. 

SEM images of Marphysa papuaensis sp. nov. paratypes AM W.53771 (A), AM W.53770 (B–E) A anterior end, dorsal view B parapodia, chaetiger 79 C subacicular hook, chaetiger 79 D pectinate chaetae, chaetiger 79 E pectinate chaetae, chaetiger 41. Abbreviations: HNS, heterodont-narrow-slender; Lot, Long outer tooth.

Subacicular chaetae with compound falcigers and subacicular hooks (Figs 6F, G, 7B, C). Compound falcigers bidentate, with short blade and large teeth, commencing from first chaetiger to near pygidium, with more than 50 chaetae within a parapodium in anterior part, with ~ 10 chaetae in mid-body and ~ 3–5 in last chaetigers (Fig. 6F). Subacicular hooks black with pale yellow tip, commencing from anterior chaetiger 20 (24) to end, most of the body with one hook per parapodia, but some posterior chaetigers with two, subacicular hooks bidentate (Figs 6G, 7B, C).

Pygidium round and crenulated, dorsally positioned, with two pairs of tapering pygidial cirri attached at ventral edge, dorsal pair 2 (3) × length of ventral pair (Fig. 5D).

Etymology

This species name refers to the type locality and geographical distribution of this species.

Type locality

Papua New Guinea, Solomon Sea, New Britain.

Distribution

Papua New Guinea, Solomon Sea (New Britain) and Bismarck Sea (New Ireland).

Habitat

Between 250 and 1200 m, mostly found inside sunken wood.

Remarks

Within the Central Indo-Pacific Realm, a single species with only compound falcigers present and branchiae restricted in a short region (group C1) has been described: M. bernardi Rullier, 1972 (type locality in New Caledonia). However, this species differs from M. papuaensis sp. nov. by the presence of a prostomium that is not bilobed, of antennae that are articulated and the absence of eyes. In contrast, M. papuaensis sp. nov. has smooth antennae, no eyes and a bilobed prostomium. The branchiae of M. bernardi are present from chaetiger 3, instead of chaetiger 7 for M. papuaensis sp. nov. and apparently M. bernardi has no pectinate chaetae, while M. papuaensis sp. nov. has pectinates commencing from first few chaetigers to near end. Finally, M. bernardi was collected in a bay from 7–8 m depth while M. papuaensis sp. nov. occurs only in deep sea, at 1200 m depth.

Marphysa zanolae sp. nov.

https://zoobank.org/EAB90680-0FDD-4B21-8FB9-DD43F378A119
Figs 8, 9

Material examined

Holotype : MNHN-IA-2015-1519, entire, few parapodia used for molecular analysis, South Pacific Ocean, Papua New Guinea, New Ireland, CP4260, -2.9°S, 151.1°E, depth 350–847 m, April 2014. Paratype: MNHN-IA-2015-1607, anterior part only, South Pacific Ocean, Papua New Guinea, New Britain, CP4266, -4.616°S, 152.25°E, depth 575–616 m, April 2014.

Description

(based on holotype, with variation in parentheses for paratype). Preserved specimens 197 (85 ant. part only) chaetigers, 101 mm (36 mm) long, 4.1 mm (2.8 mm) wide at chaetiger 10, excluding parapodia. Body elongated, slightly tapering at posterior end.

Prostomium strongly bilobed with two dorsoventrally flattened buccal lips and an anterior notch between them (Fig. 8B, C). Two palps and three antennae slender and tapering, arranged in an arc on posterior margin of prostomium. Antennae more or less smooth, of equal length, shorter than prostomium (slightly longer for PNG012), slightly longer than palps (palps very short for paratype PNG12, but probably broken) (Fig. 8C). Eyes absent. First peristomial ring approximately the same size as second one dorsally (Fig. 8C).

Figure 8. 

Marphysa zanolae sp. nov. holotype MNHN-IA-2015-1519 (A, C, D), paratype MNHN-IA-2015-1607 (B): A anterior end, lateral view B anterior end, ventral view C anterior end lateral view D pygidium.

Maxillary apparatus yellow to golden brown, partially everted in holotype and paratype. Maxillae with carriers and four paired elements and one single one, formula as follows (Fig. 9F): MF = 1+1, 4+4, 5+0, 3+6, 1+1. MI ~ 2× longer than maxillary carrier, rectangular anteriorly, triangular posteriorly, with a pair of oval wings situated at posterolateral margins. MI forceps-like, without attachment lamellae; well-developed falcal arch. Closing system ~ 4–5× shorter than MI. MII wide, without attachment lamella, teeth triangular, recurved, and distributed in less than half of plate length. Ligament between MII and MIII absent (or not sclerotized). MIII, single, slightly shorter than right MIV, curved forming part of distal arc; with left four teeth recurved, equal-sized and triangular, two right teeth shorter and blunt, without attachment lamella. Left MIV short (half the size of right MIV) with wide, triangular base, left 2 teeth longer than right-most one; attachment lamella dark, semi-circular. Right MIV with teeth triangular, recurved, decreasing in size posteriorly; attachment lamella large, wide, best developed centrally. MV, paired, rectangular (longer than wide), with a broad cutting edge, and no clearly defined teeth (but following tradition to score as 1+1). Mandibles (Fig. 9G) yellow to golden brown, slightly shorter than MI plus carriers; cutting plates whitish, with distinct growth rings.

Figure 9. 

Marphysa zanolae sp. nov. holotype MNHN-IA-2015-1519 (A–E), paratype MNHN-IA-2015-1607 (F, G): A parapodia from anterior body (chaetiger 8) B parapodia from mid-body (chaetiger 31) C parapodia from posterior body D compound falcigers from anterior chaetiger (chaetiger 12) E subacicular hook from mid-body (chaetiger 43) F maxilla, dorsal view G mandibles, dorsal view H pectinate chaeta, chaetiger 48 I pectinate chaeta, chaetiger 33. Abbreviations: MI to MV, maxillae I to V; Ac, aciculae; Bf, branchial filament; Cp, cutting plate; Fil, filament; IWT, isodont-wide-thick; Mc, maxillary carrier; Nc, notopodial cirri; Sah, subacicular hook; Vc, ventral cirri. A–C Sah and Ac are illustrated schematically to indicate position.

First two parapodia located below middle line of body wall, but gradually positioned dorsally to approximately midline in subsequent segments (Fig. 8A). Notopodial cirri with large base and slender, tapering tip from anterior to mid-body chaetigers, digitiform cirri in posterior chaetigers; same size as neuropodial cirri, but shorter than post-chaetal lobe in anterior chaetigers (Fig. 9A–C). Chaetal lobes comprising a low pre-chaetal lip and a large tongue-like post-chaetal lobe from first chaetiger to approximately chaetiger 25, almost inconspicuous thereafter. Ventral cirri bluntly conical until chaetiger 25, digitiform with bulbous base thereafter (Fig. 9A–C).

Branchiae with a long single filament (Figs 8A, 9B), commencing from chaetiger 31 (32) and continuing to mid-body (i.e., chaetiger 118 for holotype).

Aciculae black with paler blunt tips, ~ four per parapodium in anterior chaetigers, two or three per parapodium in middle chaetigers, and one or two per parapodium in posterior chaetigers. Supra-acicular chaetae with limbate capillaries and pectinates; capillaries present from first chaetiger to near pygidium, numbering up to 20 in anterior chaetigers. Pectinate chaetae commencing from first few chaetigers to near end, one type only (Fig. 9H, I), with two or three pectinate chaetae per parapodium in anterior body, up to seven from posterior chaetigers, isodont-wide-thick (IWT) having 11–20 long teeth (Fig. 9H, I).

Subacicular chaetae compound falcigers and subacicular hooks (Fig. 9D, E). Compound falcigers bidentate, with long blades and short teeth, commencing from first chaetiger to near pygidium, with more than 30 chaetae within a parapodium in anterior part, with ~ 20 chaetae in mid-body and ~ 5–7 in last chaetigers (Fig. 9D). Subacicular hooks amber to black, with much paler tip, commencing from anterior chaetiger 28 (chaetiger 30) to near end, one per parapodium in anterior and posterior parts, few chaetigers with two hooks in middle body; slightly thinner than aciculae; subacicular hooks unidentate, with blunt tip (Fig. 9E).

Pygidium round, dorsally positioned, with two pairs of tapering pygidial cirri attached at ventral edge, dorsal pair 3× length of ventral pair (Fig. 8D).

Etymology

This species is dedicated to Joana Zanol for her great contributions to the knowledge of Eunicidae and Marphysa, and her friendship to PH.

Type locality

Solomon Sea, Papua New Guinea, New Britain and New Ireland.

Distribution

Only known from type locality.

Habitat

Between 350 to 616 m depth, among pumice rocks, inside sunken wood.

Remarks

Within the Central Indo-Pacific Realm, only one species having only compound falcigers present and branchiae present in a long region (group C2) occurs: M. soembaensis Augener, 1933 (type locality in Pulau Sumba, South Indonesia). However, this species differs from M. zanolae sp. nov. by the presence of poorly developed branchiae with two or three branchial filaments instead of well-developed branchiae with a single long filament only for M. zanolae sp. nov. These branchiae start from chaetiger 40 for M. soembaensis and from chaetiger 31 for M. zanolae. Moreover, M. soembaensis has bidentate subacicular hooks while they are unidentate for M. zanolae sp. nov. Finally, M. zanolae sp. nov. has pectinate chaetae with very long outer teeth, which are not present in M. soembaensis. The blade of the compound falcigers is very short for M. soembaensis compared to those of M. zanolae sp. nov. Finally, specimens of M. soembaensis were sampled intertidally in a bay in Indonesia while M. zanolae sp. nov. occurs in deep-sea environments in Papua New Guinea.

Genetic data

COI gene was successfully sequenced and published at NCBI GenBank for two species: M. papuaensis sp. nov. and M. zanolae sp. nov. (Table 1, Fig. 10). Unfortunately, despite several attempts, sequences could not be obtained for the third species M. banana sp. nov. The two species M. papuaensis sp. nov. and M. zanolae sp. nov. are very different from all other species of Marphysa for which COI data are available and are relatively close to M. regalis Verrill, 1900 described from Bermuda (Fig. 10). The Pair-wise Kimura 2-parameter (K2P) between M. papuaensis sp. nov. and M. zanolae sp. nov. equal to 18.8% is relatively important and confirms the separation between these two species.

Figure 10. 

Majority-rule consensus tree of Marphysa species from Bayesian analysis using COI. Asterisk indicates posterior probability > 80%. Text in red indicates specimens used in this study.

Acknowledgements

Type material was collected during the two MADEEP and KAVIENG expeditions. The MADEEP deep sea cruise (PIs: Sarah Samadi, Laure Corbari, Karine Olu-Le Roy) took place in April and May 2014 on board RV ‘Alis’ deployed by Institut de Recherche pour le Développement (IRD). It operated under a Memorandum of Understanding with University of Papua New Guinea (UPNG), with a permit delivered by the Papua New Guinea Department of Environment and Conservation (DEC). The PIs acknowledge funding from Agence Nationale de la Recherche (ANR) and the National Science Council of Taiwan (ANR TF-DeepEvo 12 ISV7 005 01) and the CNRS Institut Ecologie et Environnement (INEE) (http://expeditions.mnhn.fr/campaign/madeep).

The KAVIENG 2014 expedition (Principal Investigators: Philippe Bouchet, Jeff Kinch, Claude Payri; https://expeditions.mnhn.fr/campaign/kavieng2014/leg/2) was part of the Our Planet Reviewed expeditions organized jointly by Muséum National d’Histoire Naturelle (MNHN), Pro-Natura International (PNI) and Institut de Recherche pour le Développement (IRD), with support from Papua New Guinea’s National Fisheries Authority. The lagoon survey took place in June, based at the Nago Island Mariculture and Research Facility, and in August on board RV ‘Alis’; the deep-water component, part of Tropical Deep-Sea Benthos, took place in September on board RV ‘Alis’. The organizers acknowledge supporting funding from the Total Foundation, the Laboratoire d’Excellence Diversités Biologiques et Culturelles (LabEx BCDiv, ANR-10-LABX-0003-BCDiv), the Programme Investissement d’Avenir (ANR-11-IDEX-0004-02), the Fonds Pacifique, and CNRS Institut Ecologie et Environnement (INEE). The expedition was endorsed by the New Ireland Provincial Administration. It operated under a Memorandum of Understanding with University of Papua New Guinea (UPNG), with a permit delivered by the Papua New Guinea Department of Environment and Conservation (DEC).

We also want to thank Céline Houbin, Céline Labrune, Jérome Jourde, and Joachim Langeneck for their help in sorting all worms collected during this cruise, during the MNHN workshop, and Sue Lindsay who mounted the parapodia for the SEM and took the images. Finally, we would like to thank Carol Simon for her critical review of the manuscript.

Nicolas Lavesque and Guillemine Daffe have received financial support from the French State in the frame of the “Investments for the future” Programme IdEx Bordeaux, reference ANR-10-IDEX-03-02.

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