Cah. Biol. Mar. (2005) 46 : 97-104 Composition and endemism of the deep-sea hydrothermal vent fauna Torben WOLFF Zoological Museum, University of Copenhagen, Universitetsparken 15, DK- 2100 Copenhagen Ø, Denmark Fax: +45-35321010, e-mail: [email protected] Abstract: Based on enumerations from 1997/1998 and personal records, an up-to-date survey of the hydrothermal vent fauna is presented with number of species, genera and families of various taxa. The number of recorded species is 712 and includes species known exclusively from vents (71%), also occurring in cold seeps and/or whale falls (5%), also recorded from non-vent/non-seep environments (9%) or mainly referred to genus only (15%). The number of genera is 373, of fami- lies 185. Molluscs, arthropods and polychaetes are the prevailing groups (36, 34 and 18%, respectively). Prosobranch gas- tropods number almost three fourths of 257 mollusc species, copepods more than one third of 225 crustaceans and phyl- lodocid polychaetes more than half of 129 polychaetes. Calculation of vent endemism (after deduction of 103 species of doubtful status) shows values far exceeding those found elsewhere in the marine environment. On species level it is 83.4%, with maximum values found in cirripeds (100%), copepods (98%), prosobranch gastropods (89%), and polychaetes (86%). The percentage of endemic genera is 45. All six cirriped and most vestimentiferan genera are endemic, and other high val- ues are within copepods (66%), gastropods (58%), polychaetes (51%) and decapod crustaceans (43%). Fourteen among the families (8%) are found in vents exclusively, eight also in seeps. Résumé : Composition et endémisme de la faune des sources hydrothermales profondes. Fondée sur les listes publiées en 1997 et 1998 et sur des données personnelles, une liste mise à jour de la faune des sources hydrothermales est présentée. Le nombre total d’espèces signalées est 712, nombre qui inclut les espèces exclusivement inféodées aux sources hydrother- males (71%), celles également présentes dans les zones de suintements froids ou sur les carcasses de baleines (5%), celles également trouvées dans d’autres types de milieux (9%), enfin celles identifiées seulement au niveau générique (15%). Le nombre total de genres est de 373, et celui des familles de 185. Mollusques, Arthropodes et Polychètes sont les groupes pré- dominants (respectivement 36, 34 et 18% du total). Le nombre de Gastéropodes prosobranches représente à peu près les trois quarts des 257 espèces de Mollusques, les Copépodes plus d’un tiers des 225 Crustacés et les Phyllodocidés plus de la moitié des 129 Polychètes. Une fois soustraites les 103 espèces de statut douteux, le taux d’endémisme de la faune des sources hydrothermales atteint des valeurs qui excèdent toutes celles trouvées dans d’autres environnements marins. Le taux d’endémisme au niveau spécifique est de 83 %, avec des valeurs maximales chez les Cirripèdes (100%), les Copépodes (98%), les Gastéropodes (89%) et les Polychètes (86%). Le pourcentage de genres endémiques est de 45%. Les six genres de Cirripèdes et la plupart des genres de Vestimentifères sont endémiques; d’autres valeurs élevées ont été trouvées chez les Copépodes (66%), les Gastéropodes (58%), les Polychètes (51%) et les Décapodes (43%). Quatorze familles (8%) sont endémiques des sources hydrothermales. Keywords: Hydrothermal vent fauna, Endemism, Cold seeps, Whale carcasses Reçu le 3 mai 2004 ; accepté après révision le 16 mars 2005. Received 3 May 2004; accepted in revised form 16 March 2005. 98 ENDEMISM OF THE HYDROTHERMAL VENT FAUNA Introduction and/or associated with whale carcasses, 3) species also known from other (non-vent) environments, and 4) species After the discovery in the late 1970’s of the fascinating of doubtful status (mostly referred to genus only and not hydrothermal vent fauna, being totally independent of sun recorded as being new species). Of a total 712 vent species energy, the first fauna enumeration was made by Hessler & registered, the four categories number 508, 35, 66, and 103 Smithey (1983) who very preliminarily listed 22 taxa, with species – or 71% , 5%, 9% and 15%, respectively. feeding types, mobility and locality in relation to venting The main reasons for the apparently significant increase activity. in total number of species since the list in Desbruyères & Based on about ten overview articles and many specific Segonzac (1997) and Tunnicliffe et al. (1998), from ca. 440 papers, I published an article for Scandinavian readers on to 712 species, are: 1) about 100 new records since the vent fauna (Wolff, 1985a). This included the first exten- 1997/98, 2) omission in the two lists of several species sive list of all registered vent animals, with their distribu- which, when published, had been stated as true vent forms tion at Galapagos and 13° and 21°North on the East Pacific (mainly some sponges, cnidarians, amphipods, tanaids and Rise (EPR); a paper, including the same list, was published isopods), 3) deliberate omission of all fishes and in English (Wolff, 1985b). The number of species listed, cephalopods, 4) omission of a fair number of species which identified to species or not, was 93, and the genera num- had been recorded as new but yet undescribed species, 5) bered 66. The list included details on feeding types and almost total omission of a large number of species (now mobility, the occurrence at vents, as well as data on more than 100) which were referred to species only (inclu- endemism of species and genera. Also in 1985, a list con- sion of the latter records involves that the total number is on taining 58 described species was published by Newman. the large size, since some, but probably not many, will Since 1985, I have continued to keep a detailed list of eventually be referred to already known vent species). new records of vent animals. This was done in close contact The total number of genera is 373 of which 151 are with Verena Tunnicliffe who was similarly engaged. Her found exclusively in vents. A total of 185 families are re- first published list of species (Tunnicliffe, 1991) numbered presented, with 14 in vents only and another eight also in 236 species and 135 genera. The two later lists, by seeps/whale falls (Table 4). Desbruyères & Segonzac (1997) and Tunnicliffe et al. (1998), are almost identical and record ca. 440 species Dominating groups (exclusive of fish and cephalopods) and ca. 285 genera. The molluscs, the arthropods and the polychaetes are the In continuation of my vent article in Danish (Wolff, three prevailing groups, amounting to 36.1%, 34.3% and 1985a), I recently published an up-to-date survey of new 18.1% of all vent animals (Fig. 1). discoveries in the hydrothermal vents (Wolff, 2002). It also There may be two reasons for the particular success of included chapters on the composition of animals, based on these three groups. First, it is an advantage to find protection a circumstantial list of the number of species, genera and inside a shell, an integument or a tube, respectively, against families of the various animal groups, the number of the continuous precipitation of particles in the ‘dirty’ vent species at major deep-sea vent sites, details about the environment. Especially the water inside polychaete tubes endemism, and conservation. A version in English of most may act as a buffer against rapid changes in temperature or of these items is the background for the present paper. chemical properties. Moreover, these three dominating groups are exactly those which also in other marine environ- Composition ments are able to adjust themselves to extreme conditions regarding temperature, salinity, deficiency of oxygen, etc., in opposition to more stenotopic groups like sponges, cnida- Number of species, genera and families rians, echinoderms and ascidians. Table 1 is based on my list of vent taxa up to January 2005. Molluscs: more than one fourth of all vent animals are The enumeration has been supplemented with information snails (Table 1). About 70% of these belong to the Archaeogastropoda, many of which are limpet-shaped. on special groups provided by several colleagues (see Apart from the two genera, Alviniconcha and Ifremeria Acknowledgements) and was coordinated with the lists of which, with their symbiotic bacteria, are dominating in primarily described species in Gebruk (2002). most of the West Pacific back-arc spreading centres, the Table 1 presents the total number of species, genera and others are of moderate or small size (< 1 cm). families of animal groups of the deep-sea hydrothermal More than one third of the gastropod species belong to macrofauna, with percentage of the number of species for three families (number of species in parenthesis): each of the taxa listed. Also included are 1) species exclu- Provannidae (24, inclusive three also in seeps), sively in vents, 2) species also represented in cold seeps Peltospiridae (22), and Lepetodrilidae (21, one also in T. WOLFF 99 Table 1. Number of macrofauna families, genera and species represented exclusively in hydrothermal vents or also known from cold seeps and/or whale carcasses or elsewhere in the deep sea. Meiofauna nematodes are not considered. Only vent fishes living inside active fields are included (about 90 other fish species hovering in the close surroundings, with occasional visits to the vent fields, are not consi- dered). Tableau 1. Nombre de familles, genres et espèces de macrofaune signalés exclusivement au niveau des sources hydrothermales ou également au niveau des suintements froids et des carcasses de baleines ou dans d’autres environnements profonds. Les Nématodes de la méiofaune ne sont pas pris en compte. Seuls les poissons vivant au sein des sources actives sont inclus (environ 90 autres espèces fréquentant les environs et visiteurs occasionnels ne sont pas pris en compte). Animal Groups Genera Species Fami- All gene- Vent All % of Vent Also at cold Also Dubious3 lies ra ende- species all ende- seeps/ outside mic1 vent mic2 whale vents species falls Porifera, sponges 5 6 0 11 1.6 0 0 3 8 Cnidaria 12 22 2 26 3.7 7 0 10 9 Hydrozoa, hydroids, siphonophores.