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A whale fall occurs when the carcass of a whale has fallen onto the ocean floor, typically at a depth greater than 1,000 m (3,300 ft), putting them in the bathyal or abyssal zones.[1] On the sea floor, these carcasses can create complex localized ecosystems that supply sustenance to deep-sea organisms for decades.[1] In some circumstances, particularly in cases with lower water temperatures, they can be found at much shallower depths, with at least one natural instance recorded at 150 m (500 ft) and multiple experimental instances in the range of 30-382 m (100-1300 ft).[1] Whale falls were first observed in the late 1970s with the development of deep-sea robotic exploration.[2] Since then, several natural and experimental whale falls have been monitored[1][3] through the use of observations from submersibles and remotely operated underwater vehicles (ROVs) in order to understand patterns of ecological succession on the deep seafloor.[4]

Deep sea whale falls are thought to be hotspots of adaptive radiation for specialized fauna.[1] Organisms that have been observed at deep-sea whale fall sites include chordates, arthropods, cnidarians, echinoderms, mollusks, nematodes, and annelids.[1][5] octopus, giant isopods, squat lobsters, polychaetes, prawns, shrimp, lobsters, hagfish, Osedax, crabs, sea cucumbers, and sleeper sharks. New species have been discovered, including some potentially specializing in whale falls.[1] It has been postulated that whale falls generate biodiversity by providing evolutionary stepping stones for multiple lineages to move and adapt to new environmentally-challenging habitats.[1] Researchers estimate that 690,000 carcasses/skeletons of the nine largest whale species are in one of the four stages of succession at any one time.[6] This estimate implies an average spacing of 12 km (7.5 mi) and as little as 5 km (3.1 mi) along migration routes. They hypothesize that this distance is short enough to allow larvae to disperse/migrate from one to another.[6]

Whale falls are able to occur in the deep open ocean due to cold temperatures and high hydrostatic pressures. In the coastal ocean, a higher incidence of predators as well as warmer waters hasten the decomposition of whale carcasses.[1] Carcasses may also float due to decompositional gases, keeping the carcass at the surface.[7] The bodies of most great whales (which includes sperm whales and many species of baleen whale[8]) are slightly denser than the surrounding seawater, and only become positively buoyant when the lungs are filled with air.[9] When the lungs deflate, the whale carcasses can reach the seafloor quickly and relatively intact due to a lack of significant whale fall scavengers in the water column.[1] Once in the deep-sea, cold temperatures slow decomposition rates, and high hydrostatic pressures increase gas solubility, allowing whale falls to remain intact and sink to even greater depths.

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Ecology

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A whale bone being recovered from the Santa Catalina Basin floor five years after experimental emplacement. The bone surface contains patches of white bacterial mats and a squat lobster. Hydroids have sprouted on the loop of yellow line attached to the bone.[10]

Whale falls are distributed heterogeneously throughout space and time, with a concentration along migration routes.[11] There is much faunal overlap in these whale falls across oceans. Mussels and vesicomyid clams belong to groups that harbor chemosynthetic bacteria, which can draw energy from inorganic chemicals, such as sulfur. Before their presence was discovered at whale falls, the only known habitats of these groups were sunken wood and hydrothermal vents. Similarly, lucinid clams were previously only known to inhabit carbon seeps and anoxic seafloor sediments.[12] Osedax, a genus of deep-sea polychaete worms, acts as an ecosystem engineer by excreting acid to erode whale bones and absorbing the nutrients trapped within.[13] This enhances biodiversity in the deep sea by increasing the water diffusion into the matrix of bones and facilitating colonization of the bone matrix by rarer species.[14] Members of Osedax have more dramatic effects in juvenile skeletons, which are not as well-calcified as adult skeletons.[15]

At whale fall sites it is common to see between three and five trophic levels present, with two main nutritional sources constituting the base of the food web. Adult whale carcasses can house up to five trophic levels, whereas juveniles more typically have three.[15]

Recent studies also show a possible trend of "dual niche partitioning", in which scavengers tend to reach peak densities on the carcass during the day and predators are more present during the night, reducing competition between the two trophic groups.[16] There is also a possible trend in tidal patterns and species occurrence, indicating that tides play a role in niche partitioning as well.[16]

Similar ecosystems exist when other large volumes of nutrient-rich material fall to the sea floor. Sunken beds of kelp create kelp falls, and large trees can sink to create wood falls. In more recent years, shipwrecks have also provided bases for deepwater communities. In ecosystems formed following a whale fall event, there are four stages of ecological succession.[13]

Biodiversity

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Many different taxa are known to interact with and inhabit whale falls, including multiple newly discovered species.[1] At the base of these ecosystems is the microbial community.[1] Sulfur-oxidizing, sulfate-reducing, and methanogenic microbes are the most prevalent types found on whale falls.[1] Among the sulfate-reducing bacteria, Desulfobacteraceae and Desulfobulbaceae are the most common, while Methanomicrobiales and Methanosarcinales are the most common among the methanogenic archaea.[1] Though chemosynthetic, and specifically chemolithoautotrophic, microorganisms are significant to the ecology of whale falls, these ecosystems are typically first inhabited by heterotrophic microbes such as actinomycetes, which break down collagen, and sulfate reducers.[1] The presence of such heterotrophic microbes paves the way for the chemosynthetic organisms, which then form bacterial mats that provide for larger organisms, such as certain annelid species.[1]

Chordate scavengers are also early inhabitants of whale falls.[1] Some of these relatively large scavengers that have been recorded include hagfish, sleeper sharks, and various bony fish species such as blob sculpin, Dover sole, and snubnose eelpout.[3] Many crustacean species can also be found on whale falls, including tanner and Galatheid crabs.[3] Another common crustacean inhabitant of whale falls is amphipods, which often show up in relatively high concentrations.[3]

Whale falls also house cnidarians, echinoderms, and mollusks.[5] Sea anemones, brittle stars, and sea urchins in particular have been recorded at whale fall sites.[5] Additionally, there are many species of bivalve, including members of Mytilidae and Vesicomyidae, and of marine gastropods, including members of the bone-eating genus Rubyspira.[1] Marine nematodes in the genera Halomonyhystera, Anticoma, and Theristus have also been recorded, though research on them is less extensive than other whale fall taxa.[17]

Of all taxa observed at whale falls, annelids have received the most research focus. Though marine leeches have been observed at whale falls,[3] polychaetas tend to be the focus of much of the annelid research on whale falls. This is in part due to the number of new polychaeta species discovered in these ecosystems.[1] Two common genera are Ophryotrocha, which displays adaptive radiation on whalefalls, and the genus Osedax, which are specialists that burrow into bones.[1] Members of Osedax can be found on whale falls across the globe,[1] though different species have been discovered on Atlantic whale falls than on Pacific whale falls[5].

Ecosystem stages

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There are four stages of decomposition associated with a whale fall.[13] These stages vary in duration and overlap with one other with the size of the carcass, water depth, and other environmental variables, such as tidal flow.[11] Large, intact whale falls appear to pass through the four decomposition stages, while the stages on smaller or partial carcasses may be truncated.[18] Smaller cetaceans, such as porpoises and dolphins, do not undergo the same ecological succession stages due to their small size and lower lipid content.[18] Researchers believe the presence of Osedax worms may also be a contributing factor in the observed successional differences.[19]

Stage 1

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The initial period begins with "mobile scavengers" such as hagfish and sleeper sharks actively consuming soft tissue from the carcass. Consumption can be at a rate of 40–60 kilograms (88–132 lb) per day.[12] This stage typically lasts months up to 1.5 years.[16]

Stage 2

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The second stage introduces the "enrichment opportunists". These are animals which colonize the bones and surrounding sediments that have been contaminated with organic matter from the carcass and any other tissue left by the scavengers.[12] This stage can last months up to 4.5 years.[16]

Stage 3

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In the third stage, sulfophilic bacteria anaerobically break down the lipids embedded in the bones. Instead of oxygen, they reduce dissolved sulfate (SO2−
4) and excrete hydrogen sulfide. Due to the toxicity of H
2S, only resistant chemosynthetic bacteria survive. The bacterial mats provide nourishment for mussels, clams, limpets and sea snails. As whale bones are rich in lipids, representing 4–6% of its body weight, the final digestion stage can last between 50 and possibly 100 years.[12]

Stage 4

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Some scientists postulate a fourth stage of ecological succession at whale fall sites, called the "reef stage".[13] A whale fall enters this stage once the organic compounds have been exhausted and only minerals remain in the bones, which provide a hard substrate for suspension and filter feeders.[18]

Methanogenesis

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A process called methanogenesis can also occur around whale falls. Archaea that produce methane can be abundant in anoxic sediment, but are typically not found in co-occurrence with the sulfur reducing bacteria found at whale falls. Whale falls do however support both sulfur reducing bacteria and methane producing archaea, leading to the conclusion that the area is not electron donor limited or there is minimal or no competition for suitable substrate.[20] Concentration gradients of both sulfide and methane can be found around whale falls, with the highest concentration coming within one meter of the carcass, which is several orders of magnitude higher than the surrounding sediment concentrations. Methanogenesis appears to only occur in sediments as opposed to sulfur reduction, which occurs both in sediments and on the bones of the carcass.[20] The addition of sulfur reduction in both sediments and high lipid whale bones is a key factor for why whale falls are able to sustain deep-sea communities for extended periods of time.

References

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  1. ^ a b c d e f g h i j k l m n o p q r s t u Smith, Craig R.; Glover, Adrian G.; Treude, Tina; Higgs, Nicholas D.; Amon, Diva J. (2015). "Whale-Fall Ecosystems: Recent Insights into Ecology, Paleoecology, and Evolution". Annual Review of Marine Science. 7 (1): 571–596. Bibcode:2015ARMS....7..571S. doi:10.1146/annurev-marine-010213-135144. PMID 25251277. S2CID 43201905.
  2. ^ Vetter, Tom (2015). 30,000 Leagues Undersea: True Tales of a Submariner and Deep Submergence Pilot. Tom Vetter Books, LLC. ISBN 978-1-941160-10-7.[permanent dead link]
  3. ^ a b c d e Lundsten, Lonny; Schlining, Kyra L.; Frasier, Kaitlin; Johnson, Shannon B.; Kuhnz, Linda A.; Harvey, Julio B. J.; Clague, Gillian; Vrijenhoek, Robert C. (2010-12-01). "Time-series analysis of six whale-fall communities in Monterey Canyon, California, USA". Deep Sea Research Part I: Oceanographic Research Papers. 57 (12): 1573–1584. Bibcode:2010DSRI...57.1573L. doi:10.1016/j.dsr.2010.09.003. ISSN 0967-0637.
  4. ^ Aguzzi, J.; Fanelli, E.; Ciuffardi, T.; Schirone, A.; De Leo, F. C.; Doya, C.; Kawato, M.; Miyazaki, M.; Furushima, Y.; Costa, C.; Fujiwara, Y. (2018-07-24). "Faunal activity rhythms influencing early community succession of an implanted whale carcass offshore Sagami Bay, Japan". Scientific Reports. 8 (1): 11163. Bibcode:2018NatSR...811163A. doi:10.1038/s41598-018-29431-5. ISSN 2045-2322. PMC 6057991. PMID 30042515.
  5. ^ a b c d Sumida, Paulo Y. G.; Alfaro-Lucas, Joan M.; Shimabukuro, Mauricio; Kitazato, Hiroshi; Perez, Jose A. A.; Soares-Gomes, Abilio; Toyofuku, Takashi; Lima, Andre O. S.; Ara, Koichi; Fujiwara, Yoshihiro (2016-02-24). "Deep-sea whale fall fauna from the Atlantic resembles that of the Pacific Ocean". Scientific Reports. 6 (1): 22139. doi:10.1038/srep22139. ISSN 2045-2322. PMC 4764926. PMID 26907101.{{cite journal}}: CS1 maint: PMC format (link)
  6. ^ a b Little, C. T. (2010). "Life at the Bottom: The Prolific Afterlife of Whales". Scientific American. 302 (2): 78–82, 84. doi:10.1038/scientificamerican0210-78. PMID 20128227.
  7. ^ Allison, Peter A.; Smith, Craig R.; Kukert, Helmut; Deming, Jody W.; Bennett, Bruce A. (1991). "Deep-water taphonomy of vertebrate carcasses: a whale skeleton in the bathyal Santa Catalina Basin". Paleobiology. 17 (1): 78–89. doi:10.1017/S0094837300010368. JSTOR 2400991. S2CID 129439319.
  8. ^ Baldanza, Angela; Bizzarri, Roberto; Famiani, Federico; Garassino, Alessandro; Pasini, Giovanni; Cherin, Marco; Rosatini, Francesco (2018). "The early Pleistocene whale-fall community of Bargiano (Umbria, Central Italy): Paleoecological insights from benthic foraminifera and brachyuran crabs". Palaeontologia Electronica. 21 (16): 1–27. doi:10.26879/779. ISSN 1094-8074.
  9. ^ Reisdorf, Achim G.; Bux, Roman; Wyler, Daniel; Benecke, Mark; Klug, Christian; Maisch, Michael W.; Fornaro, Peter; Wetzel, Andreas (2012-03-01). "Float, explode or sink: postmortem fate of lung-breathing marine vertebrates" (PDF). Palaeobiodiversity and Palaeoenvironments. 92 (1): 67–81. doi:10.1007/s12549-011-0067-z. ISSN 1867-1608. S2CID 129712910.
  10. ^ Russo, Julie Zeidner (24 August 2004). "This Whale's (After) Life". NOAA's Undersea Research Program. NOAA. Archived from the original on 3 November 2017. Retrieved 13 November 2010.
  11. ^ a b Allison, Peter A.; Smith, Craig R.; Kukert, Helmut; Deming, Jody W.; Bennett, Bruce A. (1991). "Deep-water taphonomy of vertebrate carcasses: a whale skeleton in the bathyal Santa Catalina Basin". Paleobiology. 17 (1): 78–89. doi:10.1017/S0094837300010368. JSTOR 2400991. S2CID 129439319.
  12. ^ a b c d Little, C. T. (2010). "Life at the Bottom: The Prolific Afterlife of Whales". Scientific American. 302 (2): 78–82, 84. doi:10.1038/scientificamerican0210-78. PMID 20128227.
  13. ^ a b c d Smith, Craig R.; Glover, Adrian G.; Treude, Tina; Higgs, Nicholas D.; Amon, Diva J. (2015). "Whale-Fall Ecosystems: Recent Insights into Ecology, Paleoecology, and Evolution". Annual Review of Marine Science. 7 (1): 571–596. Bibcode:2015ARMS....7..571S. doi:10.1146/annurev-marine-010213-135144. PMID 25251277. S2CID 43201905.
  14. ^ Alfaro-Lucas, Joan M.; Shimabukuro, Maurício; Ferreira, Giulia D.; Kitazato, Hiroshi; Fujiwara, Yoshihiro; Sumida, Paulo Y. G. (2017-12-01). "Bone-eating Osedax worms (Annelida: Siboglinidae) regulate biodiversity of deep-sea whale-fall communities". Deep Sea Research Part II: Topical Studies in Oceanography. Geo and bio-diversity in the South West Atlantic deep sea: the Iatá-piúna expedition with the manned submersible Shinkai 6500. 146: 4–12. Bibcode:2017DSRII.146....4A. doi:10.1016/j.dsr2.2017.04.011. ISSN 0967-0645.
  15. ^ a b Alfaro-Lucas, Joan M.; Shimabukuro, Maurício; Ogata, Isabella V.; Fujiwara, Yoshihiro; Sumida, Paulo Y. G. (2018-05-28). "Trophic structure and chemosynthesis contributions to heterotrophic fauna inhabiting an abyssal whale carcass". Marine Ecology Progress Series. 596: 1–12. Bibcode:2018MEPS..596....1A. doi:10.3354/meps12617. ISSN 0171-8630.
  16. ^ a b c d Aguzzi, J.; Fanelli, E.; Ciuffardi, T.; Schirone, A.; De Leo, F. C.; Doya, C.; Kawato, M.; Miyazaki, M.; Furushima, Y.; Costa, C.; Fujiwara, Y. (2018-07-24). "Faunal activity rhythms influencing early community succession of an implanted whale carcass offshore Sagami Bay, Japan". Scientific Reports. 8 (1): 11163. Bibcode:2018NatSR...811163A. doi:10.1038/s41598-018-29431-5. ISSN 2045-2322. PMC 6057991. PMID 30042515.
  17. ^ Avila, Ana K. F.; Shimabukuro, Maurício; Couto, Daniel M.; Alfaro-Lucas, Joan M.; Sumida, Paulo Y. G.; Gallucci, Fabiane (2023-05-02). "Whale falls as chemosynthetic refugia: a perspective from free-living deep-sea nematodes". Frontiers in Marine Science. 10. doi:10.3389/fmars.2023.1111249. ISSN 2296-7745.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  18. ^ a b c Hilario, Ana; Cunha, Marina R.; Génio, Luciana; Marçal, Ana Raquel; Ravara, Ascensão; Rodrigues, Clara F.; Wiklund, Helena (2015). "First clues on the ecology of whale falls in the deep Atlantic Ocean: results from an experiment using cow carcasses". Marine Ecology. 36 (S1): 82–90. Bibcode:2015MarEc..36...82H. doi:10.1111/maec.12246. ISSN 1439-0485.
  19. ^ Bryner, Jeanna (21 September 2009). "New Worm Species Discovered on Dead Whales". Live Science.
  20. ^ a b Treude, Tina; Smith, Craig R.; Wenzhöfer, Frank; Carney, Erin; Bernardino, Angelo F.; Hannides, Angelos K.; Krüger, Martin; Boetius, Antje (2009). "Biogeochemistry of a deep-sea whale fall: sulfate reduction, sulfide efflux and methanogenesis". Marine Ecology Progress Series. 382: 1–21. Bibcode:2009MEPS..382....1T. doi:10.3354/meps07972. hdl:21.11116/0000-0001-CC9B-B. JSTOR 24873149.
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