Abstract
The sighting of giant bivalves and tubeworms at the Rose Garden vent field on the Galápagos Rift in 1977 marked the discovery of hydrothermal vents, a turning point for modern biology. The following decade saw a flurry of taxonomic descriptions of vent endemic species from the first vents. With the finding of high-temperature “black smokers” on the East Pacific Rise, exploration shifted away from Galápagos. A faunal list of Galápagos vents with 65 species was published in 1991, then updated to 74 species in 2006. Since then, few expeditions returned to the Galápagos Rift. Here, we revisited several Galápagos vents including recently confirmed high-temperature sites and inactive sulfide mounds. From our collecting efforts and observations, as well as revisions from the literature, we update the faunal list to 92 species including 15 new records, restricted to obvious vent associates. Accurate regional faunal lists are important for understanding the biogeography of vent fauna, and our list will also be valuable for setting management strategies.
Introduction
The discovery of hydrothermal vents themselves on the eastern Galápagos Rift in 1977 was not really a surprise, as geologists had predicted their presence from the missing heat measured near ridge axes (Sclater and Klitgord 1973) and warm buoyant plumes collected by a towed vehicle (Weiss et al. 1977). But nobody was prepared for the first contact with its bizarre inhabitants at a supposedly nutrient-deficient deep seabed two and a half kilometers below the surface—dense aggregations of giant clams and mussels, worms living in meter-long white tubes with red plumes swaying in shimmering water, and all other animals living with them (Corliss et al. 1979). Starting with the description of the giant vent clam Turneroconcha magnifica and the bythograeid vent crab Bythograea thermydron (Boss and Turner 1980; Williams 1980) followed by the giant tubeworm Riftia pachyptila and the discovery of chemosymbiosis (Cavanaugh et al. 1981; Jones 1981), biologists began to tease apart the taxonomic affinities and evolutionary origins of these creatures (Hessler and Smithey 1983).
Within a decade of its discovery, nearly all animals found in the original diffuse flow Galápagos Rift vents, now known as the Rose Garden vent field, were described. An early faunal list in 1991 included 65 species (Tunnicliffe 1991, 1992), and another in 2006 listed 74 (Desbruyères et al. 2006). With the discovery of “black smoker” chimneys spewing out high-temperature fluids in other systems such as East Pacific Rise (EPR) and Juan de Fuca Ridge (Tunnicliffe et al. 1985; Desbruyères and Laubier 1986), exploration shifted away from Galápagos Rift where vigorous venting was apparently lacking. An expedition in 2002 found that Rose Garden had been buried under fresh basaltic lava flows, and the communities had been largely wiped out except some recent settlers on nearby low-temperature venting from cracks in an area named Rosebud (Shank et al. 2003). The 2002 expedition also found signals for more vents east of Rose Garden on the eastern rift, plus a vent at the western Galápagos Rift near the Galápagos Islands. Between 2005 and 2006, towed-camera surveys in the western Galápagos Rift confirmed the first high-temperature chimneys (Haymon et al. 2008). Though these sites provide likely grounds for new records and subsequent research cruises have visited some of these areas using underwater vehicles (Shank et al. 2012; Raineault et al. 2016), no faunal updates have been published to date.
From October to November 2023, we were able to revisit the Galápagos Rift vents on-board the Schmidt Ocean Institute’s R/V Falkor (too) during the research cruise FKt231024. One aim of the cruise was to investigate the distributions of animal communities associated with both active and inactive vents. Here, we revise the faunal list of Galápagos Rift hydrothermal vents based on the literature since the last compilation (Desbruyères et al. 2006) plus new findings from our research expedition, in order to present all reliable distribution records from vents in this region.
Materials and methods
During R/V Falkor (too) cruise Fkt231024, we visited several hydrothermal vent fields on the Galápagos Rift using the remotely operated vehicle (ROV) SuBastian. These included Rose Garden/Rosebud (0.81° N, 86.22° W, 2450–2550 m deep; dive #603) and Tempus Fugit (0.77° N, 85.91–93° W, 2500–2560 m; dive #606–607, 609) on the eastern Galápagos Rift (Shank et al. 2012; Raineault et al. 2016); as well as Iguanas-Pinguinos (2.10° N, 91.89–94° W, 1650–1700 m; dive #611–613) and Tortugas, a newly-discovered active vent field on the western edge of the East Los Huellos Caldera (0.95° N, 90.53–56° W, 1500–1600 m; dive #614) on the western Galápagos Rift (Haymon et al. 2008). An overview map of the study area is presented in Fig. 1.
We used video and screengrabs from a 4K ultra-HD video camera (SULIS Subsea Z70; resolution 3840 × 2160 pixels) on the ROV SuBastian for seafloor imaging, which allowed up to 12 × zoom for close-up observations of even smaller animals. Animals were collected using either a seven-function manipulator arm (Schilling Robotics TITAN 4) or a suction sampler mounted on ROV SuBastian. Upon recovery on-board, animals were sorted in cold (4 °C) seawater, cleaned with a brush, and photographed using a Canon EOS 5Ds R digital single-lens reflex camera equipped with a Canon EF 100 mm F2.8L MACRO IS USM macro lens. Most new records are based on collected specimens, but some larger fauna were identified using close-up imagery. For peltospirid gastropods, which exhibited considerable morphological variability compared to specimens known from the EPR (McLean 1989a), the barcoding fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene was amplified and sequenced using the universal primer pairs HCO2198-LCO1490 (Folmer et al. 1994) following a published protocol (Chen et al. 2018) to confirm their identities. The new sequences were deposited on GenBank (PP000825-PP000827) and were compared with existing EPR sequences using the search function and the built-in pairwise distance calculator of NCBI BLAST.
Previously published faunal lists for the Galápagos Rift vents (Tunnicliffe 1991, 1992; Desbruyères et al. 2006) were examined for taxonomic status using both the World Register of Marine Species (WoRMS Editorial Board 2023) and primary literature. We aimed to remove erroneous records and to ensure the list only includes those species that rely strongly on the vent environment. To eliminate erroneous records, occurrence records at the Galápagos Rift were checked against the original descriptions and subsequent works on each species; geographic distribution of those species found in other hydrothermal systems were also recorded (see Table 1). New species described since the publication of the previous lists were checked in Google Scholar using search terms “Galapagos AND hydrothermal AND new species”. New records from our present study are added to this “historical” list.
Results and discussion
Overview of vent fields visited
In the eastern Galápagos Rift, the Rose Garden/Rosebud area was covered by fresh basaltic lava flow and devoid of living vent fauna. This confirms the finding from a 2011 cruise that another eruption event between 2005 and 2011 had eliminated fauna in this field (Shank et al. 2012). Furthermore, we revisited a serpulid worm colony found in 2015 (Raineault et al. 2016) in case living vent fauna persisted (0.8049° N, 86.2194° W, 2447 m deep), but only found decaying serpulid tubes and dissolving mussel shell debris. As such, venting at Rose Garden has likely ceased—although we did not visit the location of the East of Eden field. In the nearby Tempus Fugit field (Raineault et al. 2016), we found that venting at the previously known main diffuse flow site (0.7700° N, 85.9114° W, 2561 m deep) had waned, with few living vesicomyid clams and Riftia tubeworms. Nevertheless, we found a new diffuse flow vent nearby (0.7712° N, 85.9236° W, 2602 m deep; “Walking Dead” vent). We also revisited the active chimney (Raineault et al. 2016) at the western end of Tempus Fugit (0.7712° N, 85.9332° W, 2514 m deep; “Zombie” vent) and confirmed high-temperature (>200 °C; measured with the ROV temperature probe) venting there. A number of dead spires or inactive mounds were found around the Zombie vent and were also surveyed.
Shifting to the western Galápagos Rift, we revisited all three vent sites in the Iguanas-Pinguinos vent field (Haymon et al. 2008; Raineault et al. 2016), including Iguanas East (2.0992° N, 91.9053° W, 1670 m), Iguanas West (2.1050° N, 91.9378° W, 1670 m), and Pinguinos (2.0993° N, 91.9052° W, 1670 m). We confirmed chimney structures associated with vigorous venting of high-temperature fluid at all three locations. At East Los Huellos Caldera, where only plume signals were known (Haymon et al. 2008), we discovered active venting associated with chemosynthetic communities. This included both diffuse venting areas dominated by mussels and vesicomyid clams (0.9546° N, 90.5566° W, 1590 m) and active chimney complexes with high-temperature (>250° C) venting (0.9543° N, 90.5613°W, 1520 m).
Revising the existing faunal list
The faunal list of the Galápagos Rift vents in Desbruyères et al. (2006) included a total of 74 species. Of these, two orbiniid annelid species including Orbiniella aciculata and Scoloplos ehlersi were erroneously included in the list, as the author clearly states these were collected from box cores deployed near the Galápagos Rift but were not from the vent community (Blake 1985). Here, we further remove the lysianassoid amphipod Abyssorchomene abyssorum on the grounds that it is a globally distributed deep-sea species found in non-chemosynthetic seafloor and that its vent record is based on a single specimen that may have been a by-catch (Barnard and Ingram 1990). Similarly, we took out the abyssal grenadier Coryphaenoides armatus since it is merely an occasional visitor to vents from the surrounding deep sea. Though there are two species of dubious taxonomic status—the hesionid polychaete Nereimyra alvinae with poorly preserved types (Pleijel et al. 2012) and the crab Bythograea intermedia described from megalopa and juveniles only (de Saint Laurent 1988)—we have kept them, pending future taxonomic revision. The melanodrymid snail Melanodrymia sp. and the raphitomid snail Nepotilla sp. were initially reported in a conference abstract (Gustafson 1991) and then included in a gastropod faunal list by taxonomic experts (Warén and Bouchet 1993). Though their species-level identification remains unclear, they remain on the list pending more taxonomic information. Moalic et al. (2012) supplemented the list in Desbruyères et al. (2006) to report 83 taxa. In addition to the annelids above, occasional visitors, and a double record of Thermichthys hollisi, we removed species we could not verify such as polychaetes only known from Juan de Fuca/Gorda Ridges, Bythograea microps, and Aphotopontius acanthinus. There remained 74 species.
Since the 2006 list was published, three additional species have been recorded from Galápagos Rift in the published literature. The first is the squat lobster Munidopsis recta Baba, 2005, that was confirmed as a Galápagos record by Jones and Macpherson (2007) using COI sequencing. The second species is the Pompeii worm Alvinella pompejana, visually confirmed from Tempus Fugit vent field in 2010, but not sampled (Raineault et al. 2016). The third species is Lepetodrilus aff. tevnianus Galápagos sensu Matabos and Jollivet (2019), morphologically resembling Lepetodrilus tevnianus found on the EPR vents but is a genetically distinct lineage considered to represent an undescribed species (Matabos and Jollivet 2019). Altogether, these bring the historical species occurrence record to 77 species.
New records
From our observations and collections during the 2023 cruise, we encountered a total of 15 species that are clearly associated with the chemosynthetic ecosystem and not previously recorded from Galápagos Rift vents. Table 1 lists our updated full faunal list comprising 92 species, with our new records shown in bold. Figure 2 presents key in situ screengrabs including records based on species clearly identifiable from imagery, while Fig. 3 shows photographs of specimens collected. In the following paragraphs, we provide more details on our newly recorded species.
The alvinellid worm Alvinella caudata (Figs. 2a and 3k) was seen on active chimney walls at Tempus Fugit, Iguanas-Pinguinos, and Tortugas. It co-occurred with A. pompejana, and both were collected together at Tempus Fugit; we show a specimen photo of A. pompejana (Fig. 3l) since this is the first time a specimen was collected from Galápagos Rift and serves as a confirmation of the previous record (Raineault et al. 2016). Also found on the same habitat was the hesionid worm Hesiolyra bergi (Figs. 2d and 3j), which occurred in aggregations on the chimneys; individuals were sometimes seen going into tubes of Alvinella worms. We give a tentative identification of Eunice cf. pulvinopalpata (Fauchald 1982) to a eunicid worm seen in the same habitat, but not collected (Fig. 2d). The bythograeid crab Cyanagraea praedator (Fig. 2b) was common on the active chimneys too, readily identified from images by the well-developed eye-stalk sockets and their large size (de Saint Laurent 1984). The association between Cyanagraea and Alvinella is also known from EPR vents, where the former is a predator of the latter (Desbruyères et al. 2006).
We also collected three species of peltospirid gastropods from the high-temperature Zombie vent at Tempus Fugit, including Nodopelta heminoda (Fig. 3d), Peltospira delicata (Fig. 3b), and Peltospira operculata (Fig. 3e). Although only one damaged specimen of N. heminoda could be collected, several individuals were seen near Alvinella tubes (Fig. 2c); the COI sequence of the collected specimen (GenBank PP000825) matched an existing mitogenome of the same species (GenBank BioProject PRJNA927338) with a pairwise identity of 99.82%. Peltospira operculata is also recorded based on a single young specimen (Fig. 3e) still displaying strongly ribbed shell sculpture that fades in adults (McLean 1989a). The spacing of its ribbing is wider than typical specimens from the EPR (McLean 1989a) but its COI sequence (GenBank PP000826) was closely comparable to five existing sequences (GenBank GU984275-GU984279) with pairwise identities between 99.19% and 99.68%, indicating this spacing is intraspecific variation. Peltospira delicata, recorded based on two collected adult specimens, was unusual in lacking clear spiral ridges on the body whorl (McLean 1989a). The COI sequence of the ethanol-preserved specimen (GenBank PP000827) matched an existing sequence of P. delicata (GenBank AY923931) with a pairwise identity of 99.85%. Though having a smooth adult shell is reminiscent of Peltospira operculata, all other external morphological features of our specimens such as the overall weaker coiling and the lack of operculum agree with identification as P. delicata (McLean 1989a; Warén and Bouchet 2001). This is indicative of a wider range of phenotypic variability in this species than previously known. Peltospirid snails were not seen on active chimneys in the western Galápagos Rift, but as we did not sample those chimneys for animals, we may have missed them on video due to their small size.
At both diffuse flow areas and active chimney walls in Iguanas-Pinguinos and Tortugas, we saw bouquets of the tubeworm Tevnia jerichonana (Fig. 2e), and a single juvenile specimen (Fig. 3g) was collected from the Zombie vent at Tempus Fugit where no adults could be seen. Only at the diffuse flow site at Tortugas did we see a cluster of Oasisia alvinae. The only tubeworm known from previous explorations in the eastern Galápagos Rift was Riftia pachyptila (Corliss et al. 1979; Jones 1981; Raineault et al. 2016), which also occurred in both diffuse flow sites in Tempus Fugit (but in lower abundance than previous expeditions due to waning activity there). Conversely, at the western Galápagos Rift, we did not see any sign of Riftia. At Tortugas, we found two species of the pycnogonid genus Sericosura in abundance around diffuse flows (Fig. 2f). One species with seven-segmented palps was readily identifiable as Sericosura cyrtoma (Fig. 3h), but the other (Fig. 3i) with nine-segmented palps did not match any described eastern Pacific congeners (Child 1987; Child and Segonzac 1996; Wang et al. 2013) and may represent an undescribed species. Though we did not find pycnogonids in the eastern Galápagos Rift, a previous cruise reported seeing pycnogonids there (Raineault et al. 2016), likely also Sericosura.
The raphitomid snail Phymorhynchus was often seen in the periphery zone of all vent fields we visited. Initially, Phymorhynchus from the Galápagos Rift was considered to be conspecific with those on the EPR (Warén and Bouchet 1989), but this distribution record was not mentioned when P. major was formally named based on only EPR material (Warén and Bouchet 2001). Here, we collected a specimen (Fig. 3a) and confirm the presence of P. major in the Galápagos. Though not seen on our expedition, we note that a recent expedition also on R/V Falkor (too) (Fkt230812) encountered dense coverage of a vent barnacle tentatively identified as Eochionelasmus cf. paquensis (Hiromi K. Watanabe, pers. comm.) at a vent site named Sendero del Cangrejo (2.53° N, 94.33° W, 2490 m deep). This species is added to our list based on imagery shown on an openly available YouTube stream of ROV SuBastian dive #573 at this site (Schmidt Ocean Institute 2023).
We also saw several individuals of Lebbeus co-occurring with Alvinocaris lusca on the vent periphery only in the West Iguanas vent (Fig. 2g). Though Lebbeus was not collected, our imagery provided sufficient resolution for its tentative identification as L. laurentae based on external morphology (Komai et al. 2012). Numerous individuals of the hydrozoan Candelabrum were seen also near the periphery of West Iguanas (Fig. 2h). The collected individual (Fig. 3f) was morphologically similar to Candelabrum phrygium which has a pan-arctic distribution and also known from Mid-Atlantic Ridge vents (Segonzac and Vervoort 1995). As Galápagos is far from its known range, we consider it likely to be a distinct species and tentatively identified it as C. cf. phrygium. Further away from high-temperature venting, we found many individuals of the true limpet Neolepetopsis densata on inactive chimneys near Zombie vent near Tempus Fugit (Fig. 3c). Although Gustafson and Lutz (1994) published a record for N. densata from an inactive mound on the Galápagos Rift, the validity of this was questionable as the figure captions listed the illustrated specimens as from Galápagos but the same figures were cited in the main text as specimens from 9 to 10° N on the EPR. Our present finding serves to confirm their Galápagos record. To our knowledge, this is the only Galápagos Rift vent-specific species likely restricted to inactive chimneys, a distribution pattern typical for genus Neolepetopsis (McLean 1990; Chen et al. 2021).
During our exploration, we also saw a number of animals typical of non-chemosynthetic seafloor environments within proximity to vents, such as the Pacific white skate Bathyraja spinosissima known to incubate egg cases at Galápagos Rift vents (Salinas-de-León et al. 2018), the octopus Graneledone (likely an undescribed species, Janet Voight pers. comm.) (Desbruyères et al. 2006), and some encrusting demosponges. We did not include them in our list due to the likely incidental nature of their presence in or near the chemosynthetic ecosystem.
We note that a limitation of our study is that some new records such as the tubeworm Oasisia alvinae or the vent crab Cyanagraea praedator were not collected and identified based on imagery data only, precluding future genetic studies. Though these species are easily identified from external morphology based on our current understanding with just one species in their respective genera in the eastern Pacific vents, we cannot rule out the presence of cryptic species specific to the Galápagos Rift. Previous studies in annelids and gastropods have highlighted the presence of genetic barriers and population subdivisions between the Galápagos Rift and the EPR (Hurtado et al. 2004; Matabos and Jollivet 2019). This includes cryptic species that are separated across the two ridge systems; for example, the limpet Lepetodrilus elevatus is known to consist of at least four cryptic genetic lineages across the eastern Pacific vents that are tentatively treated as one species (Matabos and Jollivet 2019).
Forty-five years after the discoveries at Rose Garden, there still are vent communities within 35 km of the original site. As the most easterly extension of the east/southeast Pacific biogeographic region, these vents may be both a population sink and a source of novel genetic diversity. Given the location in international waters near the large Galápagos mound sulfide deposits, consideration of protections such as Ecologically and Biologically Sensitive Area (EBSA) designation is warranted. The abundant active and inactive chimneys of the Iguanas-Pinguinos and Tortugas sites are testament to long-term hydrothermalism that has supported vent communities and diversification of the fauna. Currently, at least 14 species (not including nomen dubium) are known only from the Galápagos Rift (Table 1), an endemism proportion of 15% and another five species whose endemism is uncertain. While not high compared to the endemism among western Pacific vent systems (Tunnicliffe et al. 2023), as there are no geographic barriers separating the Rift from EPR, specific environmental conditions may foster the endemics. For example, sustained venting over numerous large chimneys may foster population maintenance compared to the high turnover at EPR vents (Gollner et al. 2017). Further collecting and molecular work is needed to investigate the biogeographic relationships between Galápagos Rift and the EPR in finer detail. Accurate species lists and occurrence data can reveal key processes driving the biogeographic patterns and evolution of hydrothermal vent fauna in general (Giguère and Tunnicliffe 2021; Brunner et al. 2022).
Conclusions
We revised the existing faunal list of Galápagos Rift vents and added 15 new records based on our observations and specimens collected, bringing the total to 92 species. Of these species, 14 are only known from Galápagos Rift. Though only based on qualitative observations, our results suggest some differences in fauna composition of vents at eastern vs western Galápagos Rift, warranting future research. Diversity data provide important grounds for constructing management strategies and spatial planning, especially with the growing interests for deep-sea mineral resources. As the Galápagos Rift is partially included in the Galápagos Marine Reserve, our updated species list will also be useful for conservation and marine spatial planning in this world heritage site. On the one hand, we increase considerably the number of vent species (especially those living on high-temperature chimneys) living within the Reserve, including those lacking any formal protection on the extensive East Pacific Rise south of Mexican waters. On the other hand, we show that Galápagos Rift vents host several endemic species. This is further supplemented by cryptic lineages and genetic diversities not found outside Galápagos due to isolation from the EPR at least for some gastropods (Matabos and Jollivet 2019), likely also true for some newly recorded species herein. Altogether, our results highlight the Galápagos vents as a candidate for focused conservation efforts.
References
Anderson ME (1989) Review of the eelpout genus Pachycara Zugmayer, 1911 (Teleostei: Zoarcidae), with descriptions of six new species. Proc Calif Acad Sci 46:221–242
Barnard JL, Ingram CL (1990) Lysianassoid Amphipoda (Crustacea) from deep-sea thermal vents. Smithson Contrib Zool 499:1–80
Blake JA (1985) Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I: Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae and Maldanidae. Bull Biol Soc Wash 6:67–101
Borda E, Kudenov JD, Chevaldonné P et al (2013) Cryptic species of Archinome (Annelida: Amphinomida) from vents and seeps. Proc R Soc B Biol Sci 280:20131876. https://doi.org/10.1098/rspb.2013.1876
Boss KJ, Turner RD (1980) The giant white calm from the Galapagos Rift, Calyptogena magnifica species novum. Malacologia 20:161–194
Brunner O, Chen C, Giguère T et al (2022) Species assemblage networks identify regional connectivity pathways among hydrothermal vents in the Northwest Pacific. Ecol Evol 12:e9612. https://doi.org/10.1002/ece3.9612
Burreson EM (1981) A new deep-sea leech, Bathybdella sawyeri, n. gen., n. sp., from thermal vent areas on the Galápagos Rift. Proc Biol Soc Wash 94:483–491
Cavanaugh CM, Gardiner SL, Jones ML, Jannasch HW, Waterbury JB (1981) Prokaryotic cells in the hydrothermal vent tube worm Riftia pachyptila Jones: possible chemoautotrophic symbionts. Science 213:340–342
Chen C, Ohara Y, Watanabe HK (2018) A very deep Provanna (Gastropoda: Abyssochrysoidea) discovered from the Shinkai Seep Field, Southern Mariana Forearc. J Mar Biol Assoc UK 98:439–447. https://doi.org/10.1017/S0025315416001648
Chen C, Zhou Y, Watanabe HK, Zhang R, Wang C (2021) Neolepetopsid true limpets (Gastropoda: Patellogastropoda) from Indian Ocean hot vents shed light on relationships among genera. Zool J Linn Soc 194:276–296. https://doi.org/10.1093/zoolinnean/zlab081
Child CA (1987) Ammothea verenae and Sericosura venticola, two new hydrothermal vent associated pycnogonids from the Northeast Pacific. Proc Biol Soc Wash 100:892–901
Child CA, Segonzac M (1996) Sericosura heteroscela and S. cyrtoma, new species, and other Pycnogonida from Atlantic and Pacific hydrothermal wents, with notes on habitat and environment. Proc Biol Soc Wash 109:664–676
Cohen DM, Rosenblatt RH, Moser HG (1990) Biology and description of a bythitid fish from deep-sea thermal vents in the tropical eastern Pacific. Deep Sea Res A Oceanogr Res Pap 37:267–283. https://doi.org/10.1016/0198-0149(90)90127-H
Conroy-Dalton S, Huys R (1999) New genus of Aegisthidae (Copepoda: Harpacticoida) from hydrothermal vents on the Galapagos Rift. J Crustac Biol 19:408–431
Corliss JB, Dymond J, Gordon LI et al (1979) Submarine thermal springs on the Galápagos Rift. Science 203:1073–1083. https://doi.org/10.1126/science.203.4385.1073
de Saint Laurent LM (1984) Crustacés Décapodes d’un site hydrothermal actif de la dorsale du Pacifique oriental (13° Nord), en provenance de la campagne française Biocyatherm. CR Acad Sci 299:355–360
de Saint Laurent LM (1988) Les mégalopes et jeunes stades crabe de trois espèces du genre Bythograea Williams, 1980 (Crustacea Decapoda Brachyura). Oceanol Acta Spec 8:99–107
Desbruyères D, Laubier L (1982) Paralvinella grasslei, new genus, new species of Alvinellinae (Polychaeta: Ampharetidae) from the Galapagos Rift geothermal vents. Proc Biol Soc Wash 95:484–494
Desbruyères D, Laubier L (1986) Les Alvinellidae, une famille nouvelle d’annélides polychètes inféodées aux sources hydrothermales sous-marines: systématique, biologie et écologie. Can J Zool 64:2227–2245
Desbruyères D, Segonzac M, Bright M (2006) Handbook of deep-sea hydrothermal vent fauna (2nd eds.). Denisia 18:1–544
Fauchald K (1982) A eunicid polychaete from a white smoker. Proc Biol Soc Wash 95:781–787
Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Mol Mar Biol Biotechnol 3:294–299
Giguère TN, Tunnicliffe V (2021) Beta diversity differs among hydrothermal vent systems: implications for conservation. PLoS ONE 16:e0256637. https://doi.org/10.1371/journal.pone.0256637
Gollner S, Kaiser S, Menzel L et al (2017) Resilience of benthic deep-sea fauna to mining activities. Mar Environ Res 129:76–101. https://doi.org/10.1016/j.marenvres.2017.04.010
Guinot D, Hurtado LA (2003) Two new species of hydrothermal vent crabs of the genus Bythograea from the southern East Pacific Rise and from the Galapagos Rift (Crustacea Decapoda Brachyura Bythograeidae). Comptes Rendus Biologies 326:423–439
Gustafson RG (1991) Distribution of molluscan morphospecies at deep-sea hydrothermal vents and cold-water seep areas. Abstracts of the 6th Deep-sea Biology Symposium, Copenhagen, 1991 1
Gustafson RG, Lutz RA (1994) Molluscan life history traits and deep-sea hydrothermal vents and cold methane/sulphide seeps. In: Young CM, Eckelbarger KJ (eds) Reproduction, larval biology, and recruitment of the deep-sea benthos. Columbia University Press, New York, pp 76–97
Haymon RM, White SM, Baker ET et al (2008) High-resolution surveys along the hot spot–affected Gálapagos Spreading Center: 3. Black smoker discoveries and the implications for geological controls on hydrothermal activity. Geochem Geophys Geosyst 9. https://doi.org/10.1029/2008GC002114
Hessler RR (1984) Dahlella caldariensis, new genus, new species: a leptostracan (Crustacea, Malacostraca) from deep-sea hydrothermal vents. J Crustac Biol 4:655–664. https://doi.org/10.2307/1548079
Hessler RR, Smithey WM (1983) The distribution and community structure of megafauna at the Galapagos Rift hydrothermal vents. In: Rona PA, Boström K, Laubier L, Smith KL (eds) Hydrothermal processes at seafloor spreading centers. Springer, Boston, pp 735–770
Humes AG (1987) Copepods from deep-sea hydrothermal vents. Bull Mar Sci 41:645–788
Humes AG (1988a) Oncaea praeclara n. sp. (Copepoda: Poecilostomatoida) from deep-sea hydrothermal vents in the eastern Pacific. J Plankton Res 10:475–485
Humes AG (1988b) Hyalopontius boxshalli, new species (Copepoda: Siphonostomatoida), from a deep-sea hydrothermal vent at the Galapagos Rift. Proc Biol Soc Wash 101:825–831
Humes AG (1989) Rhogobius pressulus n. sp. (Copepoda, Siphonostomatoida) from a deep-sea hydrothermal vent at the Galapagos Rift. Pac Sci 43:27–31
Humes AG (1990) Aphotopontius probolus, sp. nov., and records of other siphonostomatoid copepods from deep-sea vents in the eastern Pacific. Sci Mar 54:145–154
Humes AG, Dojiri M (1980) A siphonostome copepod associated with a vestimentiferan from the Galapagos Rift and the East Pacific Rise. Proc Biol Soc Wash 93:697–707
Humes AG, Segonzac M (1998) Copepoda from deep-sea hydrothermal sites and cold seeps: description of a new species of Aphotopontius from the East Pacific Rise and general distribution. Cah Biol Mar 39:51–62
Hurtado LA, Lutz RA, Vrijenhoek RC (2004) Distinct patterns of genetic differentiation among annelids of eastern Pacific hydrothermal vents. Mol Ecol 13:2603–2615. https://doi.org/10.1111/j.1365-294X.2004.02287.x
Jones ML (1981) Riftia pachyptila, new genus, new species, the vestimentiferan worm from the Galápagos Rift geothermal vents (Pogonophora). Proc Biol Soc Wash 93:1295–1313
Jones WJ, Macpherson E (2007) Molecular phylogeny of the East Pacific squat lobsters of the genus Munidopsis (Decapoda: Galatheidae) with the descriptions of seven new species. J Crustac Biol 27:477–501. https://doi.org/10.1651/s-2791.1
Kenk VC, Wilson BR (1985) A new mussel (Bivalvia, Mytilidae) from hydrothermal vents, in the Galapagos Rift zone. Malacologia 26:253–271
Komai T, Tsuchida S, Michel S (2012) Records of species of the hippolytid genus Lebbeus White, 1847 (Crustacea: Decapoda: Caridea) from hydrothermal vents in the Pacific Ocean, with descriptions of three new species. Zootaxa 3241:35–63. https://doi.org/10.11646/zootaxa.3241.1.2
Krantz GW (1982) A new species of Copidognathus Trouessart (Acari: Actinedida: Halacaridae) from the Galapagos Rift. Can J Zool 60:1728–1731. https://doi.org/10.1139/z82-225
Maciolek NJ (1981) Spionidae (Annelida: Polychaeta) from the Galapagos Rift geothermal vents. Proc Biol Soc Wash 94:826–837
Matabos M, Jollivet D (2019) Revisiting the Lepetodrilus elevatus species complex (Vetigastropoda: Lepetodrilidae), using samples from the Galápagos and Guaymas hydrothermal vent systems. J Molluscan Stud 85:154–165. https://doi.org/10.1093/mollus/eyy061
McLean JH (1981) The Galapagos Rift limpet Neomphalus: relevance to understanding the evolution of a major Paleozoic-Mesozoic radiation. Malacologia 21:291–336
McLean JH (1988) New archaeogastropod limpets from hydrothermal vents; superfamily lepetodrilacea I. Systematic descriptions. Phil Trans R Soc Lond B Biol Sci 319:1–32. https://doi.org/10.1098/rstb.1988.0031
McLean JH (1989a) New archaeogastropod limpets from hydrothermal vents: new family Peltospiridae, new superfamily Peltospiracea. Zoolog Scr 18:49–66. https://doi.org/10.1111/j.1463-6409.1989.tb00123.x
McLean JH (1989b) New slit-limpets (Scissurellacea and Fissurellacea) from hydrothermal vents. Part 1. Systematic descriptions and comparisons based on shell and radular characters. Contributions in Science 407:1–29. https://doi.org/10.5962/p.208131
McLean JH (1990) Neolepetopsidae, a new docoglossate limpet family from hydrothermal vents and its relevance to patellogastropod evolution. J Zool 222:485–528. https://doi.org/10.1111/j.1469-7998.1990.tb04047.x
Moalic Y, Desbruyères D, Duarte CM et al (2012) Biogeography revisited with network theory: retracing the history of hydrothermal vent communities. Syst Biol 61:127–127. https://doi.org/10.1093/sysbio/syr088
Pettibone MH (1984a) A new scale-worm commensal with deep-sea mussels on the Galapagos hydrothermal vent (Polychaeta: Polynoidae). Proc Biol Soc Wash 97:226–239
Pettibone MH (1984b) Two new species of Lepidonotopodium (Polychaeta: Polynoidae: Lepidonotopodinae) from hydrothermal vents off the Galapagos and East Pacific Rise at 21 N. Proc Biol Soc Wash 97:849–863
Pettibone MH (1985a) Additional branchiate scale-worms (Polychaeta: Polynoidae) from Galapagos hydrothermal vent and rift-area off western Mexico at 21 N. Proc Biol Soc Wash 98:447–469
Pettibone MH (1985b) New genera and species of deep-sea Macellicephalinae and Harmothoinae (Polychaeta: Polynoidae) from the hydrothermal rift areas off the Galapagos and western Mexico at 21°N and from Santa Catalina Channel. Proc Biol Soc Wash 98:740–757
Pettibone MH (1990) New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from the Axial Seamount Caldera of the Juan de Fuca ridge in the northeast Pacific Ocean off northern California. Proc Biol Soc Wash 103:825–838
Pleijel F, Rouse GW, Nygren A (2012) A revision of Nereimyra (Psamathini, Hesionidae, Aciculata, Annelida). Zool J Linn Soc 164:36–51. https://doi.org/10.1111/j.1096-3642.2011.00756.x
Pugh PR, Marshall NB (1983) Benthic siphonophores: a review of the family Rhodaliidae (Siphonophora, Physonectae). Phil Trans R Soc Lond B Biol Sci 301:165–300. https://doi.org/10.1098/rstb.1983.0025
Raineault N, Ballard R, Mayer L et al (2016) Exploration of hydrothermal vents along the Galápagos Spreading Center. Oceanography 29:35–37
Rosenblatt RH, Cohen DM (1986) Fishes living in deepsea thermal vents in the tropical eastern Pacific, with descriptions of a new genus and two new species of eelpouts (Zoarcidae). Trans San Diego Soc Nat Hist 21:71–79
Salinas-de-León P, Phillips B, Ebert D et al (2018) Deep-sea hydrothermal vents as natural egg-case incubators at the Galapagos Rift. Sci Rep 8:1788. https://doi.org/10.1038/s41598-018-20046-4
Schein-Fatton E (1985) Découverte sur la ride du Pacifique oriental à 13°N d’un Pectinidae (Bivalvia, Pteriomorphia) d’affinités paléozoïques. CR Acad Sci Paris Ser 3(301):491–496
Scheltema A (2000) Two new hydrothermal vent species, Helicoradomenia bisquama and Helicoradomenia acredema, from the eastern Pacific Ocean (Mollusca, Aplacophora). Argonauta 14:15–25
Schmidt Ocean Institute (2023) Sendero del Cangrejo | SOI Divestream 573 YouTube, Schmidt Ocean Channel, https://youtu.be/5rlniuALVnM?t=23684
Sclater JG, Klitgord KD (1973) A detailed heat flow, topographic, and magnetic survey across the Galapagos Spreading Center at 86°W. J Geophys Res 1896–1977(78):6951–6975. https://doi.org/10.1029/JB078i029p06951
Segonzac M, Vervoort W (1995) First record of the genus Candelabrum (Cnidaria, Hydrozoa, Athecata) from the Mid-Atlantic Ridge: a description of a new species and a review of the genus. Bull Mus National Hist Nat 17:31–63. https://doi.org/10.5962/p.290312
Shank T, Fornari D, Yoerger D et al (2003) Deep submergence synergy: Alvin and ABE explore the Galapagos Rift at 86°W. EOS Trans Am Geophys Union 84:425–433. https://doi.org/10.1029/2003EO410001
Shank T, Baker E, Embley R et al (2012) Exploration of the deepwater Galápagos region. Oceanography 25:50–51
Ten Hove HA, Zibrowius H (1986) Laminatubus alvini gen. et sp. n. and Protis hydrothermica sp. n. (Polychaeta, Serpulidae) from the bathyal hydrothermal vent communities in the eastern Pacific. Zoolog Scr 15:21–31
Tunnicliffe V (1991) The biology of hydrothermal vents: ecology and evolution. Mar Biol Oceanogr Annu Rev 29:319–407
Tunnicliffe V (1992) The nature and origin of the modern hydrothermal vent fauna. Palaios 7:338–350. https://doi.org/10.2307/3514820
Tunnicliffe V, Juniper SK, Burgh MED (1985) The hydrothermal vent community on Axial Seamount, Juan de Fuca Ridge. Bull Biol Soc Wash 6:453–464
Tunnicliffe V, Chen C, Giguère T et al (2023) Hydrothermal vent fauna of the western Pacific Ocean: distribution patterns and biogeographic networks. Divers Distrib. https://doi.org/10.1111/ddi.13794
Tyler PA, Pendlebury S, Mills SW et al (2008) Reproduction of gastropods from vents on the East Pacific Rise and the Mid-Atlantic Ridge. J Shellfish Res 27(107–118):112
Wang J, Lin R, Bamber R, Dingyong H (2013) Two new species of Sericosura Fry & Hedgpeth, 1969 (Arthropoda: Pycnogonida: Ammotheidae) from a hydrothermal vent on the East Pacific Rise. Zootaxa 3669:165–171. https://doi.org/10.11646/zootaxa.3669.2.8
Warén A, Bouchet P (1986) Four new species of Provanna Dall (Prosobranchia, Cerithiacea?) from East Pacific hydrothermal sites. Zoolog Scr 15:157–164. https://doi.org/10.1111/j.1463-6409.1986.tb00218.x
Warén A, Bouchet P (1989) New gastropods from East Pacific hydrothermal vents. Zoolog Scr 18:67–102. https://doi.org/10.1111/j.1463-6409.1989.tb00124.x
Warén A, Bouchet P (1993) New records, species, genera, and a new family of gastropods from hydrothermal vents and hydrocarbon seeps. Zoolog Scr 22:1–90. https://doi.org/10.1111/j.1463-6409.1993.tb00342.x
Warén A, Bouchet P (2001) Gastropoda and Monoplacophora from hydrothermal vents and seeps; new taxa and records. The Veliger 44:116–231
Weiss RF, Lonsdale P, Lupton JE, Bainbridge AE, Craig H (1977) Hydrothermal plumes in the Galapagos Rift. Nature 267:600–603. https://doi.org/10.1038/267600a0
Williams AB (1980) A new crab family from the vicinity of submarine thermal vents on the Galapagos Rift (Crustacea: Decapoda: Brachyura). Proc Biol Soc Wash 93:443–472
Williams AB, Chace FAJ (1982) A new caridean shrimp of the family Bresiliidae from thermal vents of the Galapagos Rift. J Crustac Biol 2:136–147
Woodwick K, Sensenbaugh T (1985) Saxipendium coronatum, new genus, new species (Hemichordata: Enteropneusta): the unusual spaghetti worms of the Galápagos Rift hydrothermal vents. Proc Biol Soc Wash 98:351–365
WoRMS Editorial Board (2023) World register of marine species. Available from https://www.marinespecies.org/ at VLIZ. Accessed 14 Nov 2023. https://doi.org/10.14284/14170
Zottoli R (1983) Amphisamytha galapagensis a new species of ampharetid polychaete from the vicinity of abyssal hydrothermal vents in the Galapagos Rift, and the role of this species in rift ecosystems. Proc Biol Soc Wash 96:379–391
Acknowledgements
We thank the captain and crew of R/V Falkor (too) during the research cruise Fkt231024 (“Project Zombie: Bringing dead vents to life – Ultra fine-scale seafloor mapping”), as well as the ROV SuBastian team for their immense support. To clarify, (We Were Never Promised A) Rose Garden on this cruise. Viola Watts (University of Victoria), Janet Voight (The Field Museum), Hiromi K. Watanabe (JAMSTEC), and Anders Warén (Swedish Museum of Natural History) are thanked for helping with identifications and historical records. Ana-Belén Yánez Suárez (Marine Institute of Memorial University of Newfoundland) and Stuart Banks (Charles Darwin Research Station) are gratefully acknowledged for facilitating permissions to carry out research within the Galápagos Marine Reserve in Ecuador. Miwako Tsuda (JAMSTEC) helped us with molecular barcoding. The cruise FKt231024 was supported and authorized by the Galápagos National Park Directorate, the Instituto Oceanográfico y Antártico de la Armada de Ecuador (INOCAR) and facilitated by the Charles Darwin Foundation Deep-Ocean Research Program under permit number PC 51-23. This publication is contribution number 2602 of the Charles Darwin Foundation for the Galápagos Islands. We also thank our on-board Ecuadorian observers, Diego Bermeo (Galápagos National Park) and Richard Porfirio Narea Ortega (INOCAR). We recognize the extensive support from the Schmidt Ocean Institute for this expedition and associated logistics. The vent field name “Tortugas” is a reference to the Galápagos giant tortoise (tortugas gigantes de las islas Galápagos, Chelonoidis niger) and was decided in consortia among the scientific party including discussions with the observers. Comments and edits from two anonymous reviewers helped improve an earlier version of this paper.
Funding
The research cruise FKt231024 on-board R/V Falkor (too) was funded by the Schmidt Ocean Institute. VT and JWJ participation funded by Natural Sciences and Engineering Research Council of Canada.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Conflict of interest
The authors declare no competing interests..
Ethical approval
Study species were invertebrates and no experimental manipulation was undertaken on live animals in this study.
Sampling and field studies
All necessary permits for sampling and observation have been obtained by the authors where applicable. Permission for ROV dives inside the Galápagos Marine Reserve in Ecuador was granted through a partnership between Schmidt Ocean Institute and the Charles Darwin Foundation (MAATE-DPNG/DGA-2023-1449-O).
Data availability
Data relevant to our conclusions are included in the main text. Specimens collected within the Galápagos Marine Reserve are vouchered in the Charles Darwin Research Station on Santa Cruz Island (https://www.darwinfoundation.org/en/about/cdrs). All ROV SuBastian dives during cruise FKt231024 are available online through the Schmidt Ocean Institute YouTube page: https://www.youtube.com/@SchmidtOcean/streams.
Author contribution
CC and VT conceived the project. CC and VT identified animal species and updated the faunal list. CC undertook photography and made the figures. JWJ secured funding, planned and led the expedition, and provided map of the research area. CC drafted the original manuscript which was critically revised by JWJ and VT. All authors agreed to the submission of the final version.
Additional information
Communicated by S. Gollner
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Chen, C., Jamieson, J.W. & Tunnicliffe, V. Hydrothermal vent fauna of the Galápagos Rift: updated species list with new records. Mar. Biodivers. 54, 16 (2024). https://doi.org/10.1007/s12526-024-01408-w
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s12526-024-01408-w