Woodward, S. P. A Manual of the Mollusca; or, Rudimentary Treatise of Recent and Fossil Shells (Weale, 1854).
Smith, C. R., Kukert, H., Wheatcroft, R. A., Jumars, P. A. & Deming, J. W. Vent fauna on whale remains. Nature 341, 27–28 (1989).
Squires, R. L., Goedert, J. L. & Barnes, L. G. Whale carcasses. Nature 349, 574 (1991).
Fujioka, K., Wada, H. & Okano, H. Torishima whale bone deep-sea animal community assemblage—new finding by Shinkai 6500. J. Geogr. 102, 507–517 (1993).
Dell, R. K. New species and records of deep-water Mollusca from off New Zealand. Tuhinga 2, 1–26 (1995).
Smith, C. R. & Baco, A. R. Ecology of whale falls at the deep-sea floor. Oceanogr. Mar. Biol. Ann. Rev. 41, 311–354 (2003).
Smith, C. R., Glover, A. G., Treude, T., Higgs, N. D. & Amon, D. J. Whale-fall ecosystems: recent insights into ecology, paleoecology, and evolution. Annu. Rev. Mar. Sci. 7, 571–596 (2015).
Carney, R. S. Zonation of deep biota on continental margins. Oceanogr. Mar. Biol. Ann. Rev. 43, 211–278 (2005).
Lundsten, L. et al. Time-series analysis of six whale-fall communities in Monterey Canyon, California, USA. Deep Sea Res. 1 Oceanogr. Res. Pap. 57, 1573–1584 (2010).
Dominici, S. et al. Mediterranean fossil whale falls and the adaptation of mollusks to extreme habitats. Geology 37, 815–18 (2009).
Pavlyuk, O. N., Trebukhova, Y. A. & Tarasov, V. G. The impact of implanted whale carcass on nematode communities in shallow water area of Peter the Great Bay (East Sea). Ocean Sci. J. 44, 181–88 (2009).
Danise, S. & Dominici, S. A record of fossil shallow-water whale falls from Italy. Lethaia 47, 229–243 (2014).
Fujiwara, Y. et al. Three-year investigations into sperm whale-fall ecosystems in Japan. Mar. Ecol. 28, 219–232 (2007).
Sumida, P. Y. G. et al. Deep-sea whale fall fauna from the Atlantic resembles that of the Pacific Ocean. Sci. Rep. 6, 22139 (2016).
Munschy, M. The Diamantina Zone as the result of rifting between Australia and Antarctica: geophysical constraints. Earth Planet. Sci. 326, 533–540 (1998).
Royer, J. Y. & Sandwell, D. T. Evolution of the Eastern Indian Ocean since the Late Cretaceous: constraints from geosat altimetry. J. Geophys. Res. Solid Earth 94, 13755–13782 (1989).
Ekdale, E. G., Berta, A. & Deméré, T. A. The comparative osteology of the petrotympanic complex (ear region) of extant baleen whales (Cetacea: Mysticeti). PLoS ONE 6, e21311 (2011).
Lorion, J., Duperron, S., Gros, O., Cruaud, C. & Samadi, S. Several deep-sea mussels and their associated symbionts are able to live both on wood and on whale falls. Proc. R. Soc. B 276, 177–185 (2009).
Watanabe, H., Kojima, S., Fujikura, K., Miyake, H. & Yamamoto, H. in The Vent and Seep Biota (ed. Kiel, S.) Ch. 12 (Springer, 2010).
Payne, C. Y. et al. Xyloplax princealberti (Asteroidea, Echinodermata): a new species that is not always associated with wood falls. Diversity 15, 1212 (2023).
McLeod, C. D. et al. Known and inferred distributions of beaked whale species (Cetacea: Ziphiidae). J. Cetacea. Res. Manag. 7, 271–286 (2006).
Mead, J. G. Beaked whales of the genus Mesoplodon. in Handbook of Marine Mammals, Vol. 4: River Dolphins and the Larger Toothed Whales (eds Ridgway, S. H. & Harrison, R.) 349–430 (Academic Press, 1989).
Bianucci, G., Lambert, O. & Post, K. A high diversity in fossil beaked whales (Mammalia, Odontoceti, Ziphiidae) recovered by trawling from the sea floor off South Africa. Geodiversitas 29, 561–618 (2007).
Kootker, L. M. & Laffoon, J. E. Assessing the preservation of biogenic strontium isotope ratios (87Sr/86Sr) in the pars petrosa ossis temporalis of unburnt human skeletal remains: a case study from Saba. Rapid Commun. Mass Spectrom. 36, e9277 (2022).
McArthur, J. M. et al. in A Geologic Time Scale Vol. 1 (eds Gradstein, F. M. et al.) Ch. 7 (Elsevier, 2020).
Branch, T. A. & Butterworth, D. S. Southern Hemisphere minke whales: standardised abundance estimates from the 1978/79 to 1997/98 IDCR/SOWER surveys. J. Cetacea. Res. Manag. 3, 143–174 (2001).
Murase, H. et al. Review of the assessment of two stocks of Antarctic minke whales (eastern Indian Ocean and western South Pacific). J. Cetacea. Res. Manag. 21, 95–122 (2020).
Friedlaender, A. S. et al. Feeding rates and under-ice foraging strategies of the smallest lunge filter feeder, the Antarctic minke whale (Balaenoptera bonaerensis). J. Exp. Biol. 217, 2851–2854 (2014).
Horwood J. in Encyclopedia of Marine Mammals (eds Würsig, B. et al.) 845–847 (Academic Press, 2018).
Ishii, M. et al. Diving behavior of sei whales Balaenoptera borealis relative to the vertical distribution of their potential prey. Mammal. Study 42, 1–9 (2017).
Jefferson, T. A., Webber, M. A. & Pitman, R. L. Marine Mammals of the World: A Comprehensive Guide to Their Identification (Academic, 2015).
Berta, A. Whales, Dolphins, and Porpoises: A Natural History and Species Guide (Univ. of Chicago Press, 2015).
Schorr, G. S., Falcone, E. A., Moretti, D. J. & Andrews, R. D. First long-term behavioral records from Cuvier’s beaked whales (Ziphius cavirostris) reveal record-breaking dives. PLoS ONE 9, e92633 (2014).
Costa, D. P. in Encyclopedia of Life Sciences 1–7 (Wiley, 2007).
Ponganis, P. J. Diving Physiology of Marine Mammals and Seabirds (Cambridge Univ. Press, 2015).
Falcone, E. A. et al. Diving behaviour of Cuvier’s beaked whales exposed to two types of military sonar. R. Soc. Open Sci. 4, 170629 (2017).
Fernández, A. et al. “Gas and fat embolic syndrome” involving a mass stranding of beaked whales (family Ziphiidae) exposed to anthropogenic sonar signals. Vet. Pathol. 42, 446–457 (2005).
Tyack, P. L., Johnson, M., Soto, N. A., Sturlese, A. & Madsen, P. T. Extreme diving of beaked whales. J. Exp. Biol. 209, 4238–4253 (2006).
Williams, T. M. et al. Sink or swim: strategies for cost-efficient diving by marine mammals. Science 288, 133–136 (2000).
Shipboard Scientific Party. in Proceedings of the Ocean Drilling Program Vol. 121, Initial Reports (eds Peirce, J. et al.) Ch. 13 (Texas A & M Univ. Ocean Drilling Program, 1989).
Li, Z. & Pasteris, J. D. Chemistry of bone mineral, based on the hypermineralized rostrum of the beaked whale Mesoplodon densirostris. Am. Mineral. 99, 645–653 (2014).
Fujikura, K. et al. The deepest chemosynthesis-based community yet discovered from the hadal zone, 7326 m deep, in the Japan Trench. Mar. Ecol. Prog. Ser. 190, 17–26 (1999).
Peng, X. et al. Flourishing chemosynthetic life at the greatest depths of hadal trenches. Nature 645, 679–685 (2025).
Glover, A. G., Goetze, E., Dahlgren, T. G. & Smith, C. R. Morphology, reproductive biology and genetic structure of the whale-fall and hydrothermal vent specialist, Bathykurila guaymasensis Pettibone, 1989 (Annelida: Polynoidae). Mar. Ecol. 26, 223–234 (2005).
Wiklund, H. et al. Systematics and biodiversity of Ophryotrocha (Annelida, Dorvilleidae) with descriptions of six new species from deep-sea whale-fall and wood-fall habitats in the north-east Pacific. Syst. Biodivers. 10, 243–259 (2012).
Lorion, J. et al. Adaptive radiation of chemosymbiotic deep-sea mussels. Proc. R. Soc. B 280, 20131243 (2013).
Thubaut, J., Puillandre, N., Faure, B., Cruaud, C. & Samadi, S. The contrasted evolutionary fates of deep-sea chemosynthetic mussels (Bivalvia, Bathymodiolinae). Ecol. Evol. 3, 4748–4766 (2013).
Carroll, E. L. et al. Speciation in the deep: genomics and morphology reveal a new species of beaked whale Mesoplodon eueu. Proc. R. Soc. B. 288, 20211213 (2021).
Bianucci, G., Miján, I., Lambert, O., Post, K. & Mateus, O. Bizarre fossil beaked whales (Odontoceti, Ziphiidae) fished from the Atlantic Ocean floor off the Iberian Peninsula. Geodiversitas 35, 105–153 (2013).
Lambert, O., de Muizon, C. & Bianucci, G. Neogene and Quaternary fossil remains of beaked whales (Cetacea, Odontoceti, Ziphiidae) from deep-sea deposits off Crozet and Kerguelen islands, Southern Ocean. Geodiversitas 40, 135–160 (2018).
Ryan, W. B. F. et al. Global multi-resolution topography (GMRT) synthesis data set. Geochem. Geophys. Geosyst. 10, Q03014 (2009).
Balech, B., Sandioniggi, A., Marzano, M., Pesole, G. & Santamaria, M. MetaCOXI: an integrated collection of metazoan mitochondrial cytochrome oxidase subunit-I DNA sequences. Database 2022, baab084 (2022).
Huang, Y., Gilna, P. & Li, W. Identification of ribosomal RNA genes in metagenomic fragments. Bioinformatics 25, 1338–1340 (2009).
Chaumeil, P. A., Mussig, A. J., Hugenholtz, P. & Parks, D. H. GTDB-Tk: a toolkit to classify genomes with the Genome Taxonomy Database. Bioinformatics 36, 1925–1927 (2019).
Parks, D. H. et al. GTDB: an ongoing census of bacterial and archaeal diversity through a phylogenetically consistent, rank normalized and complete genome-based taxonomy. Nucleic Acids Res. 50, D785–D794 (2022).
Wei, X., Xu, Y., Zhang, C., Zhao, J. & Feng, Y. Petrology and Sr-Nd isotopic disequilibrium of the Xiaohaizi Intrusion, NW China: genesis of layered intrusions in the Tarim Large Igneous Province. J. Petrol. 55, 2567–2598 (2014).
Weis, D. et al. High-precision isotopic characterization of USGS reference materials by TIMS and MC-ICP-MS. Geochem. Geophys. Geosyst. 7, Q08006 (2006).
McArthur, J. M., Millar, I. L., Drury, A. J. & Wagner, D. Strontium-isotope stratigraphy: methodology, standard values (SRM987, EN-1, E&A), a new Neogene curve of 87Sr/86Sr against time, its implications for astrochronology ([I]ODP Sites 1146, 1264, U1337, and U1338), and its application to ODP Site 758 (Indian summer monsoon) and IODP Site 1120 (a new age-model). Palaeogeogr. Palaeoclimatol. Palaeoecol. 669, 112907 (2025).
Swofford, D. L. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4 (Sinauer, 2002).
Bianucci, G. et al. A new Late Miocene beaked whale (Cetacea, Odontoceti) from the Pisco Formation, and a revised age for the fossil Ziphiidae of Peru. Boll. Soc. Paleontol. Ital. 63, 21–43 (2024).
Goloboff, P. A. Estimating character weights during tree search. Cladistics 9, 83–91 (1993).
Peng, X. et al. A 5.3-million-year-old deep-sea whale necropolis in the Diamantina Zone [DS/OL]. V1. Science Data Bank https://doi.org/10.57760/sciencedb.28830 (2026).
Peng, X. et al. A 5.3-million-year-old deep-sea whale necropolis in the Diamantina Zone. MorphoBank https://doi.org/10.7934/P6064 (2026).
Daly, M. & Gusmão, L. The first sea anemone (Cnidaria, Anthozoa, Actiniaria) from a whale fall. J. Nat. Hist. 41, 1–11 (2007).
Rodríguez, E. et al. Hidden among sea anemones: the first comprehensive phylogenetic reconstruction of the order Actiniaria (Cnidaria, Anthozoa, Hexacorallia) reveals a novel group of hexacorals. PLoS ONE 9, e96998 (2014).
Kongsrud, J. A. & Rapp, H. T. Nicomache (Loxochona) lokii sp. nov. (Annelida: Polychaeta: Maldanidae) from the Loki’s Castle vent field: an important structure builder in an Arctic vent system. Polar Biol. 35, 161–170 (2012).
Nishi, E. & Rouse, G. W. First whale fall chaetopterid; a gigantic new species of Phyllochaetopterus (Chaetopteridae: Annelida) from the deep sea off California. Proc. Biol. Soc. Wash. 126, 287–298 (2014).
Georgieva, M. N. et al. The annelid community of a natural deep-sea whale fall off eastern Australia. Rec. Aust. Mus. 75, 167–213 (2023).
Taviani, M. et al. Whale fall chemosymbiotic communities in a southwest Australian submarine canyon fill a distributional gap. Heliyon 10, e29206 (2024).
Okanishi, M. et al. Large populations of two new species of Ophiambix (Echinodermata, Ophiuroidea) discovered on Japanese hot vents and cold seeps. Raffles Bull. Zool. 68, 196–213 (2020).
Baker, A. N., Rowe, F. W. & Clark, H. E. A new class of Echinodermata from New Zealand. Nature 321, 862–864 (1986).
Rowe, F. W. E., Baker, A. N. & Clark, H. E. S. The morphology, development and taxonomic status of Xyloplax Baker, Rowe and Clark (Echinodermata: Concentricycloidea), with the description of a new species. Proc. R. Soc. Lond. B 233, 431–459 (1988).
Mah, C. L. A new species of Xyloplax (Echinodermata: Asteroidea: Concentricycloidea) from the northeast Pacific: comparative morphology and a reassessment of phylogeny. Invertebr. Biol. 125, 136–153 (2006).
Schnabel, K. E. The Marine Fauna of New Zealand: Squat Lobsters (Crustacea, Decapoda, Chirostyloidea) (NIWA, 2020).
Chen, C. et al. Biological surveys reveal unexpectedly high faunal diversity at Nankai Trough methane seeps. Ecosphere 16, e70451 (2025).
Cheng, J., Yan, H., Hui, M. & Sha, Z. Novel insights into deep-sea hydrothermal vent and cold seep adaptation inferred from comparative transcriptome analysis of a munidopsid squat lobster distributed in both environments. Deep Sea Res. 1 Oceanogr. Res. Pap. 205, 104245 (2024).
Seid, C. A. et al. A faunal inventory of methane seeps on the Pacific margin of Costa Rica. ZooKeys 1222, 1–250 (2025).
Bó, B. D. et al. Methane-powered sea spiders: diverse, epibiotic methanotrophs serve as a source of nutrition for deep-sea methane seep Sericosura. Proc. Natl Acad. Sci. USA 122, e2501422122 (2025).
Zhang, S. & Zhang, S. A new species of the genus Phymorhynchus (Neogastropoda: Raphitomidae) from a hydrothermal vent in the Manus Back-Arc Basin. Zootaxa 4300, 441–444 (2017).
Avila, A. K. et al. Whale falls as chemosynthetic refugia: a perspective from free-living deep-sea nematodes. Front. Mar. Sci. 10, 1111249 (2023).
