Introduction

The Galapagos archipelago is a volcanic island chain located in the Tropical Eastern Pacific, south of the Galapagos Spreading Center and forms part of the western end of the Carnegie Ridge (White et al. 1993). The archipelago originated approximately 10 Myr from the active volcanism of a melting anomaly (or “hotspot”) in the Earth’s upper mantle in the eastward-moving Nazca Plate (Glass et al. 2007). Resulting from this region’s active volcanic history, both the terrestrial and marine environments of the archipelago are defined by complex and diverse lava formations (Sinton et al. 1996, Hoernle et al. 2000), with the seafloor on and around the Galapagos platform being a heterogeneous seascape shaped by seamounts, lava flows, slopes, rift zones and hydrothermal vents (Geist et al. 2008). Therefore, these environments are likely to host rich deep-sea invertebrate communities (McClain and Barry 2010, Souza Júnior et al. 2014). To date, however, relatively little is known about the life and physical environments of these deep ocean seascapes compared to their shallow water counterparts.

The Galapagos Marine Reserve (GMR) covers approximately 138,000 km² and was established in 1998 (Fig. 1) to protect the archipelago’s marine biodiversity by banning large-scale commercial exploitation of marine resources (Henry and Roberts 2017). While deep-sea environments cover by far the largest proportion of the GMR, the technological challenges of studying deep waters mean that most baseline studies on biodiversity are limited to coastal and shallow pelagic ecosystems (Glynn and Wellington 1983, James 1991). However, there are notable historical exceptions. In the late 1890s, the Albatross Expedition pioneered the collection and study of deep-sea fauna in the Tropical Eastern Pacific (Agassiz 1905). Employing dredging methods, benthic sampling was mostly unsuccessful owing to the volcanic nature of the seafloor in the region. Still, some samples were retrieved, but only for soft bottom environments (Bergh and Agassiz 1894, Grove and Lavenberg 1997). Nearly one hundred years later, in 1986, the Harbor Branch Oceanographic Institutes submersible, Johnson SeaLink, collected the first targeted collections at a maximum depth of 915 m (Pomponi et al. 1988), with subsequent expeditions in the following years (Banks et al. 2014). Since these expeditions, research efforts have been made to list and describe the biodiversity of Galapagos’ deep-sea environment (Agassiz 1905, Pomponi et al. 1988, Manning 2016), especially on cnidarians (Cairns 1991, Cairns 2018), chordates (McCosker and Smith 1997, Grove and Lavenberg 1997, Barnett et al. 2006, McCosker et al. 2012), echinoderms (Pawson and Ahearn 2000) and isopods (Faxon 1895, Wetzer 1990). Nonetheless, to date, there are no comprehensive image inventories of the deep-sea macrofauna within this region.

Figure 1.  

Map of the Galapagos Marine Reserve with the EV Nautilus dives locations.

In recent decades, the use of Remotely operated vehicles (ROVs), equipped with effective sampling gear and high-resolution recording technologies, has greatly accelerated exploration and surveying of deep-sea habitats (Danovaro et al. 2014) and the use of such deep-submergence technology has contributed to our knowledge of deep-water decapod crustacea in both the western (Komai and Tsuchida 2014) and southern (Poupin et al. 2012) Pacific. In the same way, the Nautilus expedition conducted in 2015 marks the future of deep-sea exploration in the GMR and we hope that this inventory, based on in-situ images of the Arthropoda, will facilitate the standardisation of the morphospecies and will be useful for targeted specimen collections during future deep-sea studies in the GMR and the Eastern Tropical Pacific.

Being the first organisms to colonise the islands, even before the appearance of macroscopic plants, the terrestrial arthropods from the Galapagos Islands have been a subject of interest for many years (Peck 1990, Peck 1994). By studying these terrestrial organisms, ecologists and geologists gain a better understanding on how the terrestrial environment of the islands came into being at its present state (Peck et al. 1998). There were intensive collections of arthropods in the intertidal and shallow subtidal regions of the Galapagos Islands during 1931-1938, performed by the ship Velero III of the Allan Hancock Foundation, University of Southern California (Meredith and Hancock 1939). These collections resulted in work on brachyurans by Garth (1946), Garth (1991) and records of carideans in various taxonomic publications. James (1991) published a series of accounts of various Galapagos invertebrates but these also were from shallow waters. However, after the initial descriptions in the 1890s of the collections of the Albatross Expediton (Faxon 1893, Faxon 1895), deep-sea marine arthropods of the Galapagos received very little attention, compared to their terrestrial and shallow-water counterparts.

Materials and methods

Analysis

Discussion

Acknowledgements

References

Agassiz A (1905)
Albatross expedition to the eastern Pacific
.
Science
21
(
527
):
178
‑
183
. https://doi.org/10.1126/science.21.527.178
Ahyong ST, O’Meally D (2004)
Phylogeny of the Decapoda Reptantia: resolution using three molecular loci and morphology
.
Raffles Bulletin of Zoology
52
(
2
):
673
‑
693
.
Ahyong ST (2008)
Deepwater crabs from seamounts and chemosynthetic habitats off eastern New Zealand (Crustacea: Decapoda: Brachyura)
.
Zootaxa
1708
(
1
):
1
‑
72
. https://doi.org/10.11646/zootaxa.1708.1.1
Ahyong ST (2009)
The polychelidan lobsters: phylogeny and systematics (Polychelida: Polychelidae)
. In: Martin J, Crandall K, Felder D (Eds)
Decapod crustacean phylogenetics
.
Crustacean Issues
,
18
.
CRC Press
,
Boca Raton
,
360–390
pp.
Ahyong ST, Ng PK (2016)
The species of Mathildella Guinot and Richer de Forges, 1981 and Neopilumnoplax Serène in Guinot, 1969 (Decapoda: Brachyura: Mathildellidae)
.
Journal of Crustacean Biology
36
(
4
):
538
‑
552
. https://doi.org/10.1163/1937240x-00002446
Baba K, Wicksten M (1997)
Janetogalathea, a new genus of squat lobster, with redescription of its type species Galathea californiensis Benedict, 1902 (Anomura: Galatheidae)
.
Crustacean Research
26
:
38
‑
46
. https://doi.org/10.18353/crustacea.26.0_38
Baba K, Wicksten MK (2019)
Chirostyloidean squat lobsters (Crustacea: Decapoda: Anomura) from the Galapagos Islands
.
Zootaxa
4564
(
2
):
391
‑
421
. https://doi.org/10.11646/zootaxa.4564.2.5
Baillon S, Hamel J, Mercier A (2014)
Diversity, distribution and nature of faunal associations with deep-sea pennatulacean Corals in the Northwest Atlantic
.
PLoS ONE
9
(
11
):
e111519
. https://doi.org/10.1371/journal.pone.0111519
Banks S, Acuña D, Calderón R (2014)
Manual de Monitoreo Submareal Ecologico
.
Fundación Charles Darwin
,
Puerto Ayora
. [In
Spanish
].
Barnett LK, Didier DA, Long DJ, Ebert DA (2006)
Hydrolagus mccoskeri sp. nov., a new species of chimaeroid fish from the Galapagos Islands (Holocephali: Chimaeriformes: Chimaeridae)
.
Zootaxa
1328
:
27
‑
38
.
Bergh R, Agassiz A (1894)
Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico and in the Gulf of California, in charge of Alexander Agassiz, carried on by the von Rudolph Bergh.
Museum of Comparative Zoology at Harvard College
https://doi.org/10.5962/bhl.title.13744
Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, Breinholt JW, Bendall M, Palero F, Chan TY, Felder DL, Robles R, Chu KH, Tsang LM, Kim D, Martin JW, Crandall KA (2014)
The Emergence of Lobsters: Phylogenetic Relationships, Morphological Evolution and Divergence Time Comparisons of an Ancient Group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida)
.
Systematic Biology
63
(
4
):
457
‑
479
. https://doi.org/10.1093/sysbio/syu008
Braga-Henriques A, Carreiro-Silva M, Tempera F, Porteiro FM, Jakobsen K, Jakobsen J, Albuquerque M, Santos RS (2011)
Carrying behavior in the deep-sea crab Paromola cuvieri (Northeast Atlantic)
.
Marine Biodiversity
42
(
1
):
37
‑
46
. https://doi.org/10.1007/s12526-011-0090-3
Buhl-Mortensen L, Mortensen PB (2004)
Crustaceans associated with the deep-water gorgonian corals Paragorgia arborea (L., 1758) and Primnoa resedaeformis (Gunn., 1763)
.
Journal of Natural History
38
(
10
):
1233
‑
1247
. https://doi.org/10.1080/0022293031000155205
Cairns S (1991)
A revision of the ahermatypic Scleractinia of the Galapagos and Cocos Islands
.
Smithsonian Contributions to Zoology
504
:
1
‑
32
. https://doi.org/10.5479/si.00810282.504
Cairns S (2018)
Deep-Water octocorals (Cnidaria, Anthozoa) from the Galápagos and Cocos Islands. Part 1: Suborder Calcaxonia
.
ZooKeys
729
:
1
‑
46
. https://doi.org/10.3897/zookeys.729.21779
Capezzuto F, Maiorano P, Panza M, Indennidate A, Sion L, D'Onghia G (2012)
Occurrence and behaviour of Paromola cuvieri (Crustacea, Decapoda) in the Santa Maria di Leuca cold-water coral community (Mediterranean Sea)
.
Deep Sea Research Part I: Oceanographic Research Papers
59
:
1
‑
7
. https://doi.org/10.1016/j.dsr.2011.10.006
Clark MR, Rowden AA, Schlacher T, Williams A, Consalvey M, Stocks KI, Rogers AD, O'Hara TD, White M, Shank TM, Hall-Spencer JM (2010)
The ecology of seamounts: structure, function, and human impacts.
Annual Review of Marine Science
2
:
253
‑
78
. https://doi.org/10.1146/annurev-marine-120308-081109
Danovaro R, Snelgrove PR, Tyler P (2014)
Challenging the paradigms of deep-sea ecology
.
Trends in Ecology & Evolution
29
(
8
):
465
‑
475
. https://doi.org/10.1016/j.tree.2014.06.002
Dixon CJ, Ahyong ST, Schram FR (2003)
A new hypothesis of decapod phylogeny
.
Crustaceana
76
(
8
):
935
‑
975
. https://doi.org/10.1163/156854003771997846
Faxon W (1893)
Reports on the dredging operations off the west coast of central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer "Albatross" during 1891, Lieut. Commander Z.L. Tanner, commanding. VI. Preliminary descriptions of new species of Crustacea.
Bulletin of the Museum of Comparative Zoology at Harvard College
(
24
)
149
‑
220
.
Faxon W (1895)
Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer "Albatross," during 1891, Lieut.-Commander Z.L. Tanner, U.S.N., Commanding. XV. The stalk-eyed Crustacea
.
18
.
Memoirs of the Museum of Comparative Zoology at Harvard College
,
292
pp.
Galil BS (2000)
Crustacea Decapoda: Review of the genera and species of the family Polychelidae Wood-Mason, 1874
. In: Crosnier A (Ed.)
Résultats des Campagnes Musorstom
.
184
,
21
.
Mémoires du Muséum national d’Histoire naturelle
,
Paris
,
285-387
pp.
Garth J (1946)
Littoral brachyuran fauna of the Galapagos Archipelago. Allan Hancock Pacific Expedition
.
University of Southern California Press.
5
(
10
):
1
‑
60
.
Garth J (1991)
Taxonomy, Distribution, and Ecology of Galápagos Brachyura
.
Topics in Geobiology
123
‑
145
. https://doi.org/10.1007/978-1-4899-0646-5_6
Geist D, Diefenbach B, Fornari D, Kurz M, Harpp K, Blusztajn J (2008)
Construction of the Galápagos platform by large submarine volcanic terraces
.
Geochemistry, Geophysics, Geosystems
9
(
3
):
n/a
‑
n/a
. https://doi.org/10.1029/2007gc001795
Glass J, Fornari D, Hall H, Cougan A, Berkenbosch H, Holmes M, White S, La Torre GD (2007)
Submarine volcanic morphology of the western Galápagos based on EM300 bathymetry and MR1 side-scan sonar
.
Geochemistry, Geophysics, Geosystems
8
(
3
):
n/a
‑
n/a
. https://doi.org/10.1029/2006gc001464
Glynn P, Wellington G (1983)
Corals and coral reefs of the Galapagos Islands
.
University of California Press
URL: https://www.ucpress.edu/op/9780520047136/corals-and-coral-reefs-of-the-galapagos-islands [ISBN
9780520047136
]
Grove J, Lavenberg RJ (1997)
The fishes of the Galapagos Islands
.
Stanford University Press, Stanford, California 863 pages
.
Guilloux EL, Hall-Spencer JM, Söffker MK, Olu K (2010)
Association between the squat lobster Gastroptychus formosus and cold-water corals in the North Atlantic
.
Journal of the Marine Biological Association of the United Kingdom
90
(
07
):
1363
‑
1369
. https://doi.org/10.1017/s0025315410000524
Harpp K, Geist D (2002)
Wolf-Darwin lineament and plume-ridge interaction in northern Galápagos
.
Geochemistry, Geophysics, Geosystems
3
(
11
):
1
‑
19
. https://doi.org/10.1029/2002gc000370
Hazlett BA (1981)
The behavioral ecology of hermit crabs
.
Annual Review of Ecology and Systematics
12
(
1
):
1
‑
22
. https://doi.org/10.1146/annurev.es.12.110181.000245
Hendrickx M, Wicksten M (2016)
New records of decapod crustaceans in the eastern Pacific
.
Crustaceana
89
(
5
):
603
‑
610
. https://doi.org/10.1163/15685403-00003541
Henry L, Roberts JM (2017)
Global biodiversity in cold-water coral reef ecosystems
. In: Rossi S, Bramanti L, Gori A, Orejas C (Eds)
Marine Animal Forests
. https://doi.org/10.1007/978-3-319-21012-4_6
Hoernle K, Werner R, Morgan JP, Garbe-Schönberg D, Bryce J, Mrazek J (2000)
Existence of complex spatial zonation in the Galápagos plume
.
Geology
28
(
5
):
435
. https://doi.org/10.1130/0091-7613(2000)282.0.co;2
James M (1991)
Galapagos marine invertebrates: Taxonomy, biogeography, and evolution in Darwin's Islands
.
Topics in Geobiology
474
. https://doi.org/10.1007/978-1-4899-0646-5_16
Jonsson L, Lundälv T, Johannesson K (2001)
Symbiotic associations between anthozoans and crustaceans in a temperate coastal area
.
Marine Ecology Progress Series
209
:
189
‑
195
. https://doi.org/10.3354/meps209189
Komai T, Tsuchida S (2014)
Deep-Sea decapod crustaceans (Caridea, Polychelida, Anomura and Brachyura) collected from the Nikko Seamounts, Mariana Arc, using a remotely operated vehicle “Hyper-Dolphin”
.
Zootaxa
3764
(
3
):
279
‑
316
.
Lemaitre R (1998)
Revisiting Tylaspis anomala Henderson, 1885 (Parapaguridae), with comments on its relationships and evolution
.
Zoosystema
20
:
289
‑
305
.
Manning C (2016)
Some off-shore marine species coming to light in Galapagos, Ecuador
.
Galapagos Research
(
68
)
28
‑
33
. URL: http://aquaticcommons.org/21553/1/GR_68_p28-33.pdf
McClain C, Barry J (2010)
Habitat heterogeneity, disturbance, and productivity work in concert to regulate biodiversity in deep submarine canyons
.
Ecology
91
(
4
):
964
‑
976
. https://doi.org/10.1890/09-0087.1
McCosker J, Smith D (1997)
Two new Indo-Pacific morays of the genus Uropterygius (Anguilliformes: Muraenidae)
.
Bulletin of Marine Science
60
:
1005
‑
1014
.
McCosker J, Long D, Baldwin C (2012)
Description of a new species of deepwater catshark, Bythaelurus giddingsi sp. nov., from the Galápagos Islands (Chondrichthyes: Carcharhiniformes: Scyliorhinidae)
.
Zenodo
https://doi.org/10.5281/ZENODO.280250
Mercier A, Hamel J (2008)
Nature and role of newly described symbiotic associations between a sea anemone and gastropods at bathyal depths in the NW Atlantic
.
Journal of Experimental Marine Biology and Ecology
358
(
1
):
57
‑
69
. https://doi.org/10.1016/j.jembe.2008.01.011
Meredith D, Hancock A (1939)
Voyages of the Velero III: a pictorial version, with historical background of scientific expeditions through tropical seas to equatorial lands aboard M/V Velero III.
Bookhaven Press
https://doi.org/10.5962/bhl.title.10190
Mora C, Tittensor D, Adl S, Simpson AB, Worm B (2011)
How many species are there on Earth and in the Ocean?
PLoS Biology
9
(
8
):
e1001127
. https://doi.org/10.1371/journal.pbio.1001127
Morato Gomes TAF, Pauly D (2004)
Seamounts: biodiversity and fisheries
.
Fisheries Centre. University of British Columbia
https://doi.org/10.14288/1.0074789
Morato T, Watson R, Pitcher T, Pauly D (2006)
Fishing down the deep
.
Fish and Fisheries
7
(
1
):
24
‑
34
. https://doi.org/10.1111/j.1467-2979.2006.00205.x
Muñoz I, García-Isarch E, Sobrino I, Burgos C, Funny R, González-Porto M (2011)
Distribution, abundance and assemblages of decapod crustaceans in waters off Guinea-Bissau (north-west Africa)
.
Journal of the Marine Biological Association of the United Kingdom
92
(
03
):
475
‑
494
. https://doi.org/10.1017/s0025315411001895
Pawson DL, Ahearn C (2000)
Bathyal echinoderms of the Galapagos Islands
. In: Barker M (Ed.)
Echinoderms
.
41-46
pp.
Peck S (1990)
Eyeless arthropods of the Galapagos Islands, Ecuador: Composition and origin of the cryptozoic fauna of a young, tropical, oceanic archipelago
.
Biotropica
22
(
4
):
366
. https://doi.org/10.2307/2388554
Peck S (1994)
Sea-surface (pleuston) transport of insects between islands in the Galápagos archipelago, Ecuador
.
Annals of the Entomological Society of America
87
(
5
):
576
‑
582
. https://doi.org/10.1093/aesa/87.5.576
Peck S, Heraty J, Landry B, Sinclair B (1998)
Introduced insect fauna of an oceanic archipelago: The Galápagos Islands, Ecuador
.
American Entomologist
44
(
4
):
218
‑
237
. https://doi.org/10.1093/ae/44.4.218
Pomponi S, Reed J, Rinehart K (1988)
1986 expedition to Galapagos Islands, Cocos Islands, and Pearl Islands.
Harbor Branch Oceanographic Institution, Fort. Pierce
Poupin J, Corbari L, Perez T, Chevaldonne P (2012)
Deep-water decapod crustaceans studied with a remotely operated vehicle (ROV) in the Marquesas Islands, French Polynesia (Crustacea: Decapoda)
.
Zootaxa
3550
(
1
). https://doi.org/10.11646/zootaxa.3550.1.3
Scholtz G (1995)
Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca)
.
Zoological Journal of the Linnean Society
113
(
3
):
289
‑
328
. https://doi.org/10.1006/zjls.1995.0011
Sigovini M, Keppel E, Tagliapietra D (2016)
Open nomenclature in the biodiversity era
.
Methods in Ecology and Evolution
7
(
10
):
1217
‑
1225
. https://doi.org/10.1111/2041-210x.12594
Sinton C, Christie D, Duncan R (1996)
Geochronology of Galápagos seamounts
.
Journal of Geophysical Research: Solid Earth
101
:
13689
‑
13700
. https://doi.org/10.1029/96jb00642
Smith CR, Grange LJ, Honig DL, Naudts L, Huber B, Guidi L, Domack E (2011)
A large population of king crabs in Palmer Deep on the west Antarctic Peninsula shelf and potential invasive impacts
.
Proceedings of the Royal Society B: Biological Sciences
279
(
1730
):
1017
‑
1026
. https://doi.org/10.1098/rspb.2011.1496
Souza Júnior Md, Ferreira F, de Oliveira V (2014)
Effects of the spatial heterogeneity on the diversity of ecosystems with resource competition
.
Physica A: Statistical Mechanics and its Applications
393
:
312
‑
319
. https://doi.org/10.1016/j.physa.2013.08.045
Wetzer R (1990)
A new species of isopod, Aega (Rhamphion) francoisae (Flabellifera: Aegidae), from the cloaca of an ascidian from the Galápagos Islands.
Proceedings of The Biological Society of Washington
103
:
655
‑
662
.
White W, McBirney A, Duncan R (1993)
Petrology and geochemistry of the Galápagos Islands: Portrait of a pathological mantle plume
.
Journal of Geophysical Research: Solid Earth
98
:
19533
‑
19563
. https://doi.org/10.1029/93jb02018
Wicksten M, Mendez M (1983)
Bathypalaemonella delsolari, a new species of shrimp from Peru (Decapoda, Caridea, Campylonotidae)
.
Crustaceana
45
(
3
):
225
‑
231
. https://doi.org/10.1163/156854083x00271
Wicksten MK (1989)
Ranges of offshore decapod crustaceans in the Eastern Pacific Ocean
.
Transactions of the San Diego Society of Natural History.
21
:
291
‑
316
. https://doi.org/10.5962/bhl.part.24590