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Home , Bismarck Sea, Cataegis, Eunicidae, Fauna of New Guinea, Pectinodonta, Philippines

Exploration of deep- of Eric Pante, Laure Corbari, Justine Thubaut, Tin- Chan, Ralph Mana, Marie-Catherine Boisselier, Philippe Bouchet, Sarah Samadi

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Eric Pante, Laure Corbari, Justine Thubaut, Tin-Yam Chan, Ralph Mana, et al.. Exploration of the deep-sea fauna of . Oceanography, Oceanography Society, 2012, 25 (3), pp.214-225. ￿10.5670/oceanog.2012.65￿. ￿hal-00873493￿

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CITATION Pante, E., L. Corbari, J. Thubaut, T.-Y. Chan, R. Mana, M.-C. Boisselier, P. Bouchet, and S. Samadi. 2012. Exploration of the deep-sea fauna of Papua New Guinea. Oceanography 25(3):214–225, http://dx.doi.org/10.5670/oceanog.2012.65.

DOI http://dx.doi.org/10.5670/oceanog.2012.65

COPYRIGHT This article has been published inOceanography , Volume 25, Number 3, a quarterly journal of The Oceanography Society. Copyright 2012 by The Oceanography Society. All rights reserved.

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LEFT | A freshly collected specimen of the spider crab Oxypleurodon christiani, newly described from the BioPapua col- lections (Richer de Forges and Corbari, 2012). TOP AND BOTTOM | Specimens of king crabs preliminar- ily attributed to the Paralomis.

Exploration of the Deep-Sea Fauna of Papua New Guinea

By Eric Pante, Laure Corbari, Justine Thubaut, t I N -Ya m C h a n , Ralph Mana, Marie-Catherine Boisselier, Philippe Bouchet, and Sarah Samadi

214 Oceanography | Vol. 25, No. 3 Abstract. Little is known of New Guinea’s deep benthic communities. In Vityaz expeditions collected biological fall 2010, the Muséum d’Histoire naturelle, Institut de Recherche pour le benthic samples in the , namely Développement, and University of Papua New Guinea spearheaded an international the hadal part of the Trench three-leg cruise, BioPapua, aimed at exploring the deep waters of eastern Papua in the (Belyaev, 1972). New Guinea and its satellite . Special attention was given to faunal In fall 2010, the Muséum national assemblages associated with sunken wood and decomposing vegetation as well as d’Histoire naturelle (MNHN) and seamount summits and slopes. In this article, we review the information available the Institut de Recherche pour le on the deep ecosystems of Papua New Guinea and summarize preliminary results Développement (IRD; France), in col- of the BioPapua cruise. laboration with the University of Papua New Guinea (UPNG), put together New Guinea and 1975). recently, research conducted a team of international scientists to Our Understanding in New Guinea by Vojtech Novotný explore and describe the deep benthic of and collaborators on plant-insect host fauna of the Bismarck and Solomon . The of New Guinea has a spe- specificity has bearing on global bio- The research cruise BioPapua mobilized cial place in the heart of biologists. diversity numbers (e.g., Novotný et al., 11 scientists from six countries, who Biodiversity studies in New Guinea and 2002; Hamilton et al., 2010). And occupied 156 stations within 2–9°S and its satellite islands have helped shape then, of course, there is Tim Flannery’s 144–155°E (Figure 1). In this article, some of the pillars of evolutionary biol- immensely popular Throwim Way Leg, a we review the available information on ogy and ecological theory. In the late travelogue that perpetuates our percep- the deep-sea environments of Papua 1850s, New Guinea was the eastern- tion of New Guinea as a frontier of bio- New Guinea (PNG), and describe the most of Alfred Wallace’s destinations diversity exploration (Flannery, 1998). efforts deployed for the BioPapua cruise. during his exploration of the Malay The faunal diversity of the island’s terres- , and his comparative obser- trial, shallow marine, and environ- The Deep-Sea Environments vations nurtured his theories on island ments is well known; the island is nestled of Papua New Guinea and the mechanisms pro- in the middle of the , a PNG is situated in one of the most geo- moting evolution (Wallace, 1869, 1876). zone delineated by , Bali, and the logically active in the , In the 1920s, ornithologist that is home to about where all possible types of plate bound- accumulated data on New Guinea three quarters of the world’s coral species aries can be observed (Tregoning et al., that would later appear in his Systematics (Veron et al., 2009). 2000). Most of the tectonic energy results and the Origin of Species, a landmark in In contrast, there is a surprising lack from the collision of the Indo-Australian the development of the modern synthe- of information on the deep-sea ecosys- and Pacific Plates, which entrap the sis (Mayr, 1942). Similarly, data from tems surrounding New Guinea and its Bismarck, Solomon, and Woodlark New Guinean diversity found their satellite islands. Symptomatically, the Plates (e.g., Tregoning et al., 2000; Hall, place in MacArthur and Wilson’s theory Challenger expedition sailed almost 2002). To the north, the Manus Basin of island biogeography (MacArthur and directly from Torres Strait to Wilson, 1967). Jared Diamond’s observa- and the ; though it visited the The authors wish to dedicate this feature tions of the in the Bismarck Sea to Christian Késiano Fitialeata, who and its satellite islands nourished his from March 4–10, 1875, only a single tragically passed away during the first leg work on the structure of ecological com- deepwater haul was made in 150 fath- of BioPapua. Christian had been a crew munities and fed the “Single Large Or oms; likewise, the Dutch Siboga expedi- member on R/V Alis for many years, and Several Small [SLOSS]” debate on the tion (1899–1900) did not sample east of was instrumental in the success of many application of the principles of bioge- Halmahera. In fact, of the “historical” cruises of the Tropical Deep-Sea Benthos ography in reserve design and conser- exploring expeditions, only the Danish program. He was a great companion at vation biology (Diamond, 1972, 1973, Galathea (1950–1952) and the Russian sea and on land, and is sorely missed.

Oceanography | 2012 215 vent endemic, a very meager inventory for an (EEZ) 2 Manus totaling over 2 million km of benthic surface below 100 m depth (Vanden Limana Kaia Mata na Taru Seamount Seamount Berghe, 2007; Edward Vanden Berghe, OBIS, pers. comm., 2011). The Bismarck Sea of Bismark (Manus Basin) Rabaul and Solomon Seas host many under- water features such as seamounts, of New Britain Bougainville Sanguma which only a very few (e.g., Franklin and Seamount Papua New Guinea Lae Edison Seamounts) have been sampled, mapped, or even visited. Allain et al. (2008) recently reviewed the seamounts Solomon Sea of the southwestern Pacific by screening (Woodlark Basin) Moresby and cross checking 20 existing data sets, and listed 91 underwater features for the sunken wood fauna EEZ of PNG. No information is avail- cold seep fauna other benthic fauna able for 69 of those features. The 22 other

Figure 1. Map of Papua New Guinea and its satellite islands. Dots, color-coded based on faunal type, features were further classified into sea- represent sampling stations. The inset provides a regional map. A marine geological map of PNG can be mounts (11), drowned atolls and banks found in Tregoning et al. (2000). (7 and 1), ridges (2), and a plateau. This lack of information on PNG seamounts recently formed (< 4 million years ago) German, Austrian, and French teams is reflected by the fact that no seamount in the Bismarck Sea. This sedimentary (Auzende et al., 2000, and references site is available on the Seamount Online basin is characterized by several active therein). These efforts mostly focused or Seamount Catalogue databases, not accretion zones (the Manus Spreading on the Pacmanus and Vienna Woods even the few that have been sampled. Center, the South Eastern Rift, and vent fields in the Bismarck Sea, and on the Western Rift zones) and is rapidly Franklin Seamount in the Solomon Sea. New Guinean Deep Benthic spreading. To the south, the young The geological complexity and the Fauna in a Regional Woodlark Basin (formed < 2.5 million rapidity of change observed in the Biogeographic Context years ago) produces new oceanic crust, are of particular biogeographic interest, The current body of knowledge on while the Solomon Plate subducts both in terms of biological colonization New Guinean deep-sea benthic fauna beneath the Bismarck Plate, forming the and diversity of habitat (e.g., hydro- comes almost exclusively from the study New Britain chain. thermal vent fields, river sedimentary of highly specialized hydrothermal Most of the biological deep-sea sci- cones, seamounts, slopes and can- vent , the distribution of which entific exploration done to date in the yons, sedimentary ). Significant strictly depends on the location of vent PNG region stemmed from geological efforts have been made to describe fields. Hydrothermal vent animals might research. PNG’s strong tectonic activity the endemic hydrothermal-vent fauna therefore not be good models to repre- is particularly conducive to hydrother- (e.g., Desbruyères et al., 2006, and ref- sent the biology of deep New Guinean mal venting, which was first revealed in erences therein), but the general deep fauna. Data on other benthic groups, the mid- to late (Both et al., 1986; benthic fauna have yet to be described. however, are sorely lacking, and do Tufar 1990; Lisitsyn et al., 1993). A series For example, the Biogeographic not allow us to understand patterns of international research efforts fol- Information System (OBIS) database of faunal connectivity across the deep lowed, with cruises involving American, contains information on 34 species of waters of the western Canadian, Australian, Japanese, Russian, deepwater , 30 of which are at this time. This section summarizes

216 Oceanography | Vol. 25, No. 3 current knowledge on the biogeography strongly affected their evolutionary Other models suggest of deep-sea benthic organisms in the history and current distribution. This a dichotomy between northern and western Pacific Ocean. fauna may therefore not be represen- southern biogeographic . In a Studies of the global biogeogra- tative of the overall biogeography of large-scale analysis of bio- phy of hydrothermal vent fauna place deep-sea benthic fauna from this area. geographic patterns, Macpherson et al. New Guinean assemblages within Unfortunately, data from other deep-sea (2010) established that the fauna of the a Southwestern Pacific . In organisms are very scarce. The known (, , and the analysis of Bachraty et al. (2009), biogeography of the deep-sea coral the Solomon Islands) was significantly the Mariana back-arc basin, Loihi genus Chrysogorgia well exemplifies the different from the species assemblages Seamount offshore , seamounts lack of information on nonvent, deep found in eastern , of the Kermadec Arc in , benthic fauna (Figure 2; Pante et al., (Banda and Celebes Seas), and the and the Central Indian Ridge delimit 2012). Chrysogorgia shows sampling gaps Philippines. No data were available for this region. Previous studies offer dif- around PNG indicative of our state of PNG. Similar biogeographic studies ferent boundaries for this province knowledge on the New Guinean deep-sea focusing on deep-sea bathymodioline (e.g., Van Dover et al., 2002; Tyler and fauna. Out of 65 species (and variations), support the same overall geo- Young, 2003) but show faunal affini- 21 were originally described from loca- graphic separations, with assem- ties between New Guinean vents and tions in or on the immediate periphery blages being very similar among New the Mariana, North , and Lau back- of the Coral Triangle. Although five spe- Caledonia, Vanuatu, and the Solomon arc basins. These affinities, however, cies from the were Islands, and different from northern are relatively weak, as most vents are also observed in the Northeast Pacific locations such as the Philippines (Lorion characterized by high at the (Hawaiian Archipelago), only two species et al., 2010; Lorion and Samadi, 2010). species level. In a multivariate commu- were found to occur both east and west Again, no data were available for PNG. nity analysis, Desbruyères et al. (2006) of the island of New Guinea. Although Puillandre et al. (2010), questioning found that the highest similarity level these results might simply be a sampling how modes of larval dispersal relate between vents of the Manus basin and artifact or the result of incomplete taxo- to geographical fragmentation in the other Southwest Pacific locations was nomic knowledge, they might also reflect gastropod Bathytoma in the western 24% at the species level and 46% at the actual biogeographic patterns. Pacific, also underlined the sore lack of genus level, and linked the New Guinean vents to the communities of the North Eric Pante was a PhD candidate at the University of at Lafayette (USA) during Fiji and Lau back-arc basins. In fact, the BioPapua cruise, and is now a postdoctoral researcher at the Laboratoire Littoral, Vrijenhoek (2010) and Van Dover (2011) Environnement et Sociétés (LIENSs), Université de La Rochelle, La Rochelle, France. recently proposed a biogeographic Laure Corbari is Assistant Professor, Muséum national d’Histoire naturelle, Département model in which New Guinean vents Systématique et Evolution, Paris, France. Justine Thubaut is PhD Candidate, Muséum constitute the northwestern end of the national d’Histoire naturelle, Département Systématique et Evolution, Paris, France. range of a distinct Southwestern Pacific Tin-Yam Chan is Professor, Institute of and Center of Excellence for Marine Province. The New Guinean back-arc Bioenvironment and Biotechnology, National Taiwan University, Taiwan. Ralph Mana is basins are particularly isolated within Professor, School of Natural and Physical Sciences, University of Papua New Guinea, Port the mosaic of western Pacific spreading Moresby, Papua New Guinea. Marie-Catherine Boisselier is a researcher at the Centre centers, as about 1,500 km separate the National de la Recherche Scientifique, working at the Muséum national d’Histoire naturelle, Woodlark Basin from North Fiji and Département Systématique et Evolution, Paris, France. Philippe Bouchet is Professor, Manus from the Mariana Basin (Hessler Muséum national d’Histoire naturelle, Département Systématique et Evolution, Paris, and Lonsdale, 1991). France. Sarah Samadi ([email protected]) is a researcher at the Institut de Recherche pour The geographic isolation of PNG le Développement, working at the Muséum national d’Histoire naturelle, Département hydrothermal-vent organisms may have Systématique et Evolution, Paris, France.

Oceanography | September 2012 217 specimens from New Guinea. the historical and contemporary mecha- conducted at over 5,000 stations between It is clear from hydrothermal vent nisms leading to the faunal discontinui- 100 and 1,500 m depth. Through taxo- community data available today, as well ties within the region? nomic networking with an international as from the biogeographic patterns of team of more than 200 scientists, over , galatheid crabs, and bathymo- BioPapua and the Tropical 2,000 new species have been described dioline mussels, that the deep waters of Deep-Sea Benthos Program based on material from the TDSB pro- PNG are located at the confluence of dif- BioPapua is part of a long-term research gram (Bouchet et al., 2008). ferent biogeographic regions. The foun- endeavor called the Tropical Deep- TDSB cruises, because of their robust- dations of our exploration of PNG deep Sea Benthos (TDSB) program. Started ness and relative technical simplicity, benthic ecosystems are therefore both in the 1980s by the IRD (formerly often represent the first-ever exploration discovery- and hypothesis-driven, as we Office de la Recherche Scientifique et of the deep-sea benthos in remote areas aim to document the deepwater benthic Technique Outre-Mer [ORSTOM]) and such as , the Solomon Islands, organisms present in nonhydrothermal the MNHN, its goal is to explore the or Vanuatu. BioPapua, as most TDSB ecosystems and to understand their his- deep benthic ecosystems of the tropical cruises, relied on the New Caledonia- torical and contemporary biogeography Indo-West Pacific, with special emphasis based R/V Alis, a 28 m long ship with in the context of the tropical Western on macroinvertebrates. Since the late high maneuverability. The combined use Pacific. Are organisms from this region , more than 60 TDSB cruises have of a beam trawl and a dredge maximizes more closely related to southern, north- been conducted from Taiwan to the the types of substrates and faunal diver- ern, or western species pools? What are Marquesas Islands, and sampling was sity that can be collected (Figure 3).

BioPapua Research Themes and Preliminary Results The first purpose of BioPapua, as for previous TDSB cruises, was to explore remote and uncharted and describe its deep benthic fauna. Beyond this effort of discovery, BioPapua was designed to sample specimens from targeted habitats and specific Faunal divide questions on the evolution of deep- between SE and NW water . We outline research deep-sea provinces? themes and preliminary results below.

Sunken Wood and the Evolution of Chemosynthetic Organisms Since their discovery in 1977 on the Galápagos Spreading Center (Lonsdale, 1977), hydrothermal vents have been the focus of tremendous research interest. Although significant efforts were put forth to character- ize this environment and elucidate the Figure 2. Geographic distribution of 634 records of the octocoral genus Chrysogorgia. Inset: one of the 56 of Chrysogorgia sampled during BioPapua, with associated mechanisms involved in its function- squat lobster (data from Watling et al., 2011). ing and maintenance, much less is

218 Oceanography | Vol. 25, No. 3 known about the evolutionary origins diversity (morphological, genetic, and a unique fauna, composed of mollusks of its fauna. Phylogenetic studies of ecological) associated with organic falls, (wood-boring bivalves and mytilids, the deep-sea mussels of the subfamily (2) to determine how organisms rely on , gastropods; see Warén, 2011), Bathymodiolinae show that species from organic remains, and (3) to investigate crustaceans (isopods, amphipods, bar- vents, seeps, and deep-sea organic falls the historical and ecological factors that nacles, and decapods), echinoderms (vegetation and whale carcasses) form may have influenced the diversification (sea urchins, sea stars, and ophiuroids), monophyletic clades, suggesting that of this fauna. (19 families identified so these deep-sea bivalves may have used Among the 156 stations sampled far), and sipunculids (see Figure B1 in organic falls as a transitional habitat during BioPapua, 84 contained sunken Box 1), a composition that was con- from shallow habitats to deep vents vegetation. A wide array of bays and sistent with previous findings from and seeps (Distel et al., 2000). Recent submarine canyons was sampled, other locations in the Southwest Pacific results suggest that the evolutionary from the mouth of the River to (Samadi et al., 2010). For example, the history of deep-sea mussels associated Mambare Bay on the eastern coast of crustaceans collected on plant remains with organic falls is completely tangled the island of New Guinea, and on the belong to a few genera repeatedly found with that of vent and seep mussels, with eastern and western coasts of New on such material throughout the Indo- scenarios probably more complex than Britain. Trawling revealed a wealth of Pacific (for decapods: the galatheids the “wooden steps” hypothesis (Lorion decomposing wild and cultivated plants Munida and Munidopsis, the hermit et al., 2010). Moreover, geography, rather (tree trunks, branches and leaves, sugar crabs Xylopagurus and Pylocheles, and than habitat specificity or chemosym- cane, coco, and nypa nuts). Decaying the thalassinids Callianassa; Samadi biotic requirements, seems to signifi- vegetation was found accumulating at et al., 2010). Most of the taxa, however, cantly structure diversification patterns the bottom of submarine canyons. As probably belong to new species. Author in sunken wood-associated mytilids expected, these substrates were host to Thubaut and colleagues included (Lorion et al., 2009, 2010). There are still large gaps in our under- standing of the series of evolutionary a b events that lead to the diversification of vent, seep, and organic fall fauna. First, sampling plays a crucial role in com- parative phylogenetic inference, and most studies to date are biased toward a more robust sampling of vents and seeps. Incomplete sampling can lead to phylogenies that do not accurately reflect evolutionary history. There is c d therefore a need to thoroughly charac- terize both vent/seep and organic fall biodiversity. One objective of BioPapua was to provide new material that may promote understanding of the evo- lutionary origins of the crustaceans and mollusks inhabiting deep-sea chemosynthetic environments. To Figure 3. Sampling strategy employed during BioPapua and other Tropical Deep-Sea Benthos cruises. fill the identified sampling gaps, the (a) Beam trawl. (b) Warén dredge. (c–d) Raw samples that illustrate the abundance of fauna collected main objectives are (1) to describe the and the diversity of substrates.

Oceanography | September 2012 219 Box 1. Vegetation-Associated Fauna

TheChallenger expedition (1872–1876) casually documented the bacterial ectosymbiosis have been confirmed for several specimens, and presence of animals on plant remains at the deep seafloor, but the molecular analyses revealed unexpectedly high diversity (recent work importance of plant material in the deep sea was not underscored until of author Corbari). Wood-dependent diets and bacterial interactions Wolff (1979). Until recently, plant-associated organisms were mainly have already been demonstrated for different wood-associated organ- looked at as zoological and/or ecological curiosities and thus (apart from isms (sea urchins, Becker et al., 2009; Pectinodonta, Zbinden et al., 2010; taxonomic consideration) only anecdotally studied. During previous Munidopsidae, Hoyoux et al., 2009). Future work will focus on placing TDSB cruises in the Solomon Islands, it became clear that the slopes of trophic relationships in an evolutionary context. oceanic islands within the region accumulate large amounts of decom- Animals are not the only active decomposers of plant remains. Wood posing vegetation. Organic debris was associated with a rich and original decay also results from the metabolic activities of bacteria (Waterbury fauna (Samadi et al., 2007a; Warén, 2011), and its discovery catalyzed new et al., 1983; Distel and Roberts, 1997) and fungi (Ray, 1959; Ray and research interests for the TDSB team (Samadi et al., 2010). Subsequent Stuntz, 1959). Although shallow marine fungi are common and diverse cruises in the Solomon Islands and Vanuatu specifically targeted sunken (550 species described, at least 1,500 expected to exist), deep-sea fungi vegetation to further characterize this fauna. PNG and its satellite islands appear to be rare (four species described from > 1,600 m) and are still appear to be a very promising location for such studies. Their lush coastal little studied (Dupont et al., 2009, and references therein). For example, tropical forests and large rivers transition into bays and canyons that may Oceanitis scuticella, originally described in 1977, was not seen until 2004, accumulate sunken vegetation. Figure B1 shows some of the diversity when specimens were collected in Vanuatu and the Solomon Islands found on plant remains. Adipicola longissima (Figure B1 c) is a mussel during TDSB cruises (Dupont et al., 2009). Alisea longicolla, a new genus found mainly in association with nuts. In contrast with and species described from material collected alongside O. scuticella dur- other sunken wood mussels that have extensive geographic distributions ing a 2005 TDSB cruise (Dupont et al., 2009), is another example of how in the Pacific,Adipicola longissima is restricted to the areas were nypa rare and understudied taxa are brought to light by the exploratory TDSB palms grow (mainly the Philippines, Indonesia, PNG, and the northern cruises. During BioPapua, deep-sea fungi were sampled at seven stations area of the Solomon Islands). Chitons revealed unsuspected diversity, deeper than 700 m. Samples include a new occurrence for O. scuticella and a wealth of undescribed species (Figure B1 a–f; Nicolas Puillandre, and also a new occurrence (at 700–1,150 m depth) of an undescribed Muséum national d’Histoire naturelle, pers. comm., 2012). The amphipod morpho-species that was previously known from only a few specimens genus Bathyceradocus (Maeridae; Figure B1 i) was represented by eight collected in 2007 off the Solomon Islands (Figure B1 g). species, four of them new. A wood-based diet and the presence of

a b c d

e f g h

i j k l

Figure B1. Deck photos of vegetation-associated fauna. (a) , Ferreiraella sp. (b) Cataegidae, Cataegis sp. (c) Mytilidae, Adipicola longissima nested in nypa palm . (d) Fissurellidae, Puncturella sp. (e) Pectinodontidae, Pectinodonta sp. (f) Chiton Leptochiton sp. (g) Undescribed fungi. (h) Munidopsidae Galacantha sp. (i) Maeridae, 220 Oceanography | Vol. 25, No. 3 Bathyceradocus sp. (j) ?Echinidae. (k) Polynoidae, Harmothoe sp. (l) Eunicidae, sp. samples collected during BioPapua features that could significantly hin- productivity) and geographical locations into a larger study of the biogeography der faunal dispersal (e.g., Clark et al., (e.g., latitude and isolation from conti- and integrative biology of Adipicola 2010). The extent of faunal connectiv- nental margins). iwaotakii, a small bathymodioline mus- ity among seamounts and seamount One of the goals of BioPapua was sel associated with sunken wood. This chains is, however, still poorly known. to help fill the geographical gap in our group has found high genetic diversity Beyond theoretical inquiries, there is understanding of Pacific seamount (seven mitochondrial COI haplotypes), a strong practical need to understand communities. Three of the seamounts a signal of population expansion, and seamount ecosystems in order to man- catalogued by Allain et al. (2008) were a closer biogeographic relationship age fragile fisheries resources and sampled during BioPapua. Two are to the Solomon Islands, Vanuatu, and associated fauna. Seamounts aggregate located in the Bismarck Sea, between New Caledonia than to and the commercially important species and the Western Rift Philippines. Among the 26 putative and are hotspots of pelagic and ben- accretion zone (3.05°S, 147.53°E and species of wood-associated mussels thic biodiversity (Samadi et al., 2007b; 3.23°S, 147.33°E, five stations sampled). recognized by Lorion et al. (2010) for Morato et al., 2010). The impact of deep- The first seamount, Limana Kaia, the Southwest Pacific, 11 were collected sea trawling on developing seamount appeared poor in benthic fauna, as our during BioPapua. None of the nine puta- fisheries has been shown to be cata- trawl mostly brought back bare rocks. tive species found in the Philippines and strophic and long lasting (Althaus et al., This negative result might be explained Japan were collected in PNG; the wood- 2009; Williams et al., 2010); therefore, by limited and/or unsuccessful sampling associated mussel fauna of PNG has there is no more critical time to study on rugged terrain. The second seamount, therefore stronger affinities with south- seamount ecosystems. Mata Na Taru, is more interesting from ern locations such as New Caledonia. The scarcity of data on seamount a biological standpoint, as we were able fauna, however, prevents us from con- to collect a diverse fauna. The third sea- Connectivity and Endemism ceptualizing the processes that shape mount (Sanguma, 5.53°S, 154°E, seven of Seamount Fauna and maintain seamount faunal assem- stations sampled) is located east of the Describing biodiversity from under- blages. The obstacles to describing north end of Bougainville Island, on the water mountains or seamounts the biodiversity and biogeography of edge of the Solomon Trench, an active (e.g., Stocks, 2009) has become one of seamount fauna are (1) a lack of geo- zone of of the Solomon Plate the major themes of deep-sea explora- graphically comprehensive sampling, under the Pacific Plate. This seamount, tion (e.g., special issue on “Mountains (2) the paucity of model systems allow- sampled at depths between 369 and in the Sea,” Oceanography vol. 23, March ing faunal comparisons across seamount 768 m, revealed a wealth of filter-feeding 2010, http://www.tos.org/oceanography/ chains, , and depth regimes, and invertebrates, such as hydroids, sea pens, archive/23-1.html). Seamounts are (3) the lack of integration of genetic octocorals, and sea anemones, as well as among the most ubiquitous underwater information into the estimation of ende- a rich fauna. features in the ocean (Wessel et al., 2010) mism levels. Indeed, since the inception Forty-four live specimens of Nassaria and may constitute one of the larg- of seamount science, many paradigms were collected during BioPapua, includ- est marine biomes, with a cumulative for the ecology of seamount fauna have ing seven on Sanguma Seamount. This surface area exceeding 17 million km2 been proposed, but few are strongly sup- neogastropod, with a nonplanktotrophic, (Etnoyer et al., 2010; Yesson et al., 2011). ported by empirical evidence (Rowden poorly dispersive larval stage, is one of Seamounts constitute an ideal system et al., 2010). There is therefore a strong the model systems used to investigate for the study of deep-sea biogeogra- need for the seamount research com- faunal connectivity patterns between phy as they are prevalent across ocean munity to sample across the diversity slope and seamount environments in basins, are patches of hard substrata of seamount environments (e.g., depth New Caledonia (Castelin, 2010; Castelin scattered on vast sedimentary plains, range, steepness, geological origin, et al., 2010). Among the 250 individuals and are characterized by hydrographic habitat composition, and water-column previously analyzed (collected during

Oceanography | September 2012 221 seven cruises representing approximately common among BioPapua stations, all species (based on DNA barcoding 800 sampling stations), 11 putative the others being rare) were detected, using mtMutS; McFadden et al., 2010; molecular species were detected, of and only two of these are shared with Pante and Watling, 2011) were detected, which four were found exclusively in the the rich New Caledonian fauna (Magalie 25% of them new to our collections Philippines (111 stations), and seven in Castelin, Muséum national d’Histoire (Pante et al., 2012) and most being rare. New Caledonia (600 stations). Among naturelle, pers. comm., 2012). This species pool represents 35% of the 44 BioPapua specimens of Nassaria, Among octocorals, Chrysogorgia the diversity found in the Pacific, and nine putative molecular species (one was particularly diverse, as 12 putative 27% of the total haplotypic richness of

Box 2. Biodiversity Highlights

One main focus of TDSB cruises is to explore a b c and document deep-sea biodiversity. While searching for accumulations of sunken vegeta- tion at the mouth of the Sepik River (03°53’S, 144°41’E), we sampled organisms typically found at cold seeps and hydrothermal vents, among them beard worms (Siboglinidae: Escarpia sp. and Lamellibrachia sp.) and two d e undescribed species of bathymodioline mus- sels (Gigantidas sp.; Figure B2 e–f), one of which associated with a polynoid scale worm (Branchipolynoe aff. pettiboneae; Figure B2 g). Several undescribed species of Gigantidas were already known from nearby seep localities f g h (Kyuno et al., 2009) off New Guinea. On the last leg of BioPapua, we sampled the extraordinary copepod Cardiodectes sp. (Figure B2 i; Jean-Lou Justine, Muséum national d’Histoire naturelle, pers. comm., 2012) on the eye of its fish host, and eulimid gastropods neatly attached on their urchin host (Figure B2 c). Species of the i j k Eulimidae are all parasites of echinoderms, and much work remains to be done to understand patterns of host specificity and echinoderm- mollusk co-evolution. Another noteworthy find was a specimen ofBayerotrochus (Figure B2 a). This grazing mollusk of the fam- ily Pleurotomariidae has a continuous Figure B2. Biodiversity examples: (a) Pleurotomariidae, Bayerotrochus sp. (b) Amphinomidae, Chloeia sp. record since the . As for many other (c) Eulimidae, Pelseneeria sp. on sea-urchin. (d) Nassaridae, Nassarius sp. (e) Mytlidae, Gigantidas sp. nov 1. groups, pleurotomariids had never before been (f–g) Mytilidae, Gigantidas sp. nov 2 in association with Branchipolynoe sp. nov. (h) Buccinidae, Nassaria sp. (i) Parasitic copepod Pennellidae, Cardiodectes sp. (j) Deep-sea fish, ?Chauliodidae. (k) Cirripedia, Lepadidae. recorded from the vicinity of New Guinea (Harasewych, 2002). This specimen probably belongs to an undescribed species (Patrick Anseeuw, Royal Belgian Institute of Natural Sciences, pers. comm., 2012).

222 Oceanography | Vol. 25, No. 3 the genus (although we warn that these results will have to be analyzed using rarefaction analysis). Interestingly, two of these 12 putative species col- lected on Sanguma Seamount were also previously found on New Caledonian seamounts (one on Norfolk Ridge, the other on Loyalty Ridge), suggesting, as for the gastropod Nassaria, broad geographical distribution of these seamount organisms.

Conclusion Very little is known of the biodiversity and biogeography of deep-sea animals from PNG. The data available, however, suggest that deep New Guinean fauna is at the confluence of significantly differ- ent biogeographic regions, and is more closely related to the southern than the northern Pacific fauna. Preliminary data suggest that deep New Guinean fauna is highly diversified. As an example, a rapid assessment of decapod crustacean bio- Figure 4. New and undescribed crustacean species. (a–b) Family Solenoceridae, Maximiliaeus diversity revealed more than 500 species odoceros (newly described by Chan, 2012). (c) Family Epialtidae, Oxypleurodon stimpsoni. (d) Family Goneplacidae, Thyraplax sp. (e) Family Paguridae, Bathypaguropsis sp. (f) Family Munidopsidae, collected, including four new genera (in Munidopsis sp. Photos: Tin-Yam Chan the families Solenoceridae, Galatheidae, Callianassidae and Parapaguridae; Figure 4) and about 15% new species. ecosystems are still poorly known. We and Anders Warén (Swedish Museum We expect that further study of the therefore hope that our geographically of Natural History). We warmly material collected during BioPapua will comprehensive sampling program will thank our contacts at the University shed light on large-scale biogeographic provide a baseline for the biodiversity of Papua New Guinea and the PNG patterns in the tropical deep sea of the of the deep benthic organisms of the National Research Institute, particularly Western Pacific Ocean. While we are Bismarck and Solomon Seas. Jim Robins who was instrumental in just beginning to fathom the amplitude obtaining research permits, and Edward of biodiversity in deep New Guinean Acknowledgements Vanden Berghe and Gilberto Marani for waters, mining exploration and exploita- The authors warmly thank the crew of their help extracting data from the OBIS tion of seafloor massive sulfides (SMS) R/V Alis, who made the TDSB cruise database. We thank Nicolas Puillandre, is flourishing in the region (Hoagland series a great success. We thank the Magalie Castelin, Stéphane Hourdez, et al., 2010; Van Dover, 2011). The participants of the BioPapua cruise: Jean-Lou Justine, Patrick Anseeuw, exploitation of metals from SMS depos- Jesse Apone, Edwin Sohun, and Alfred and James Albert for sharing data and its is an emerging industry, and adverse Ko’ou (UPNG); Julie Ponsard (University providing specimen identifications. environmental effects are hard to pre- of Liège); Mao-Ying Lee and Ruei Yi Lee Comments from Amélia Viricel and dict, as hydrothermal vent and adjacent (National Taiwan Ocean University); three anonymous reviewers significantly

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