Ecology

Andersson, M. Sexual selection (Princeton University Press, 2019).
Google Scholar 
Davy, A. J. & Smith, H. Life-history variation and environment. In Plant Population Ecology (eds Davy, A. J., Hutchings, M. J. & Watkinson, A. R.) 1–22. 28th Symposium of the British Ecological Society, Sussex, 1987 (Blackwell, 1988).Wilson, K. L., De Gisi, J., Cahill, C. L., Barker, O. E. & Post, J. R. Life-history variation along environmental and harvest clines of a northern freshwater fish: plasticity and adaptation. J. Anim. Ecol. 88, 717–733 (2019).PubMed 

Google Scholar 
Laiolo, P. & Obeso, J. R. Life-history responses to the altitudinal gradient. In High Mountain Conservation in a Changing World (eds Catalan, J. et al.) 253–283 (Springer, 2017).
Google Scholar 
Schwarz, R. & Meiri, S. The fast-slow life-history continuum in insular lizards: a comparison between species with invariant and variable clutch sizes. J. Biogeogr. 44, 2808–2815 (2017).
Google Scholar 
Holm, S. et al. Size-related life-history traits in geometrid moths: a comparison of a temperate and a tropical community. Ecol. Entomol. 44, 711–716 (2019).
Google Scholar 
Ferguson, G. W. & Fox, S. F. Annual variation of survival advantage of large juvenile side-blotched lizards, Uta stansburiana: Its causes and evolutionary significance. Evolution 38, 342–349 (1984).PubMed 

Google Scholar 
Madsen, T., Ujvari, B., Shine, R. & Olsson, M. Rain, rats and pythons: Climate-driven population dynamics of predators and prey in tropical Australia. Austral Ecol. 31, 30–37 (2006).
Google Scholar 
Brown, G. P. & Shine, R. Rain, prey and predators: climatically driven shifts in frog abundance modify reproductive allometry in a tropical snake. Oecologia 154, 361–368 (2007).ADS 
PubMed 

Google Scholar 
James, C. & Shine, R. Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone. Oecologia 75, 307–316 (1988).ADS 
PubMed 

Google Scholar 
Mesquita, D. O. et al. Life-history patterns of lizards of the world. Am. Nat. 187, 689–705 (2016).PubMed 

Google Scholar 
Meiri, S. et al. The global diversity and distribution of lizard clutch sizes. Glob. Ecol. Biogeogr. 29, 1515–1530 (2020).
Google Scholar 
Andrews, R. M. Growth rate in island and mainland anoline lizards. Copeia 1976, 477–482 (1976).
Google Scholar 
Jessop, T. S. et al. Maximum body size among insular Komodo dragon populations covaries with large prey density. Oikos 112, 422–429 (2006).
Google Scholar 
Van Buskirk, J. & Crowder, L. B. Life-history variation in marine turtles. Copeia 1994, 66–81 (1994).
Google Scholar 
Broderick, A. C., Godley, B. J. & Hays, G. C. Trophic status drives interannual variability in nesting numbers of marine turtles. Proc. R. Soc. B 268, 1481–1487 (2001).CAS 
PubMed 
PubMed Central 

Google Scholar 
Broderick, A. C., Glen, F., Godley, B. J. & Hays, G. C. Variation in reproductive output of marine turtles. J. Exp. Mar. Biol. Ecol. 288, 95–109 (2003).
Google Scholar 
Pike, D. A. Climate influences the global distribution of sea turtle nesting. Glob. Ecol. Biogeogr. 22, 555–566 (2013).
Google Scholar 
Ujvari, B., Shine, R., Luiselli, L. & Madsen, T. Climate-induced reaction norms for life-history traits in pythons. Ecology 92, 1858–1864 (2011).PubMed 

Google Scholar 
Shine, R., Shine, T. G., Brown, G. P. & Goiran, C. Life history traits of the sea snake Emydocephalus annulatus, based on a 17-yr study. Coral Reefs 39, 1407–1414 (2020).
Google Scholar 
Shine, R., Brown, G. P. & Goiran, C. Population dynamics of the sea snake Emydocephalus annulatus (Elapidae, Hydrophiinae). Sci. Rep. 11, 20701 (2021). ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Goiran, C., Dubey, S. & Shine, R. Effects of season, sex and body size on the feeding ecology of turtle-headed sea snakes (Emydocephalus annulatus) on IndoPacific inshore coral reefs. Coral Reefs 32, 527–538 (2013).ADS 

Google Scholar 
Ineich, I. & Laboute, P. Les Serpents Marins de Nouvelle-Calédonie (IRD éditions, 2002).Udyawer, V., Goiran, C. & Shine, R. Peaceful coexistence between people and deadly wildlife: Why are recreational users of the ocean so rarely bitten by sea snakes? People Nat. 3, 335–346 (2021).
Google Scholar 
Goiran, C., Brown, G. P. & Shine, R. The behaviour of sea snakes (Emydocephalus annulatus) shifts with the tides. Sci. Rep. 10, 1–8 (2020).
Google Scholar 
Lukoschek, V. & Shine, R. Sea snakes rarely venture far from home. Ecol. Evol. 2, 1113–1121 (2012).PubMed 
PubMed Central 

Google Scholar 
Shine, R., Goiran, C., Shine, T., Fauvel, T. & Brischoux, F. Phenotypic divergence between seasnake (Emydocephalus annulatus) populations from adjacent bays of the New Caledonian lagoon. Biol. J. Linn. Soc. 107, 824–832 (2012).
Google Scholar 
Avolio, C., Shine, R. & Pile, A. J. The adaptive significance of sexually dimorphic scale rugosity in sea snakes. Am. Nat. 167, 728–738 (2006).PubMed 

Google Scholar 
White, G. C. & Burnham, K. P. Program MARK: Survival estimation from populations of marked animals. Bird Study 46, S120–S139 (1999).
Google Scholar 
Russell, B. C., Anderson, G. R. V. & Talbot, F. H. Seasonality and recruitment of coral reef fishes. Mar. Freshw. Res. 28, 521–528 (1977).
Google Scholar 
Shine, R., Bonnet, X., Elphick, M. J. & Barrott, E. G. A novel foraging mode in snakes: browsing by the sea snake Emydocephalus annulatus (Serpentes, Hydrophiidae). Funct. Ecol. 18, 16–24 (2004).
Google Scholar 
Calow, P. Adaptive aspects of energy allocation. In Fish Energetics (eds Tytler, P. & Calow, P.) 13–31 (Springer, 1985).
Google Scholar 
Lenormand, T. Gene flow and the limits to natural selection. Trends Ecol. Evol. 17, 183–189 (2002).
Google Scholar 
Bronikowski, A. M. Experimental evidence for the adaptive evolution of growth rate in the garter snake Thamnophis elegans. Evolution 54, 1760–1767 (2000).CAS 
PubMed 

Google Scholar 
Cook, T. R., Bonnet, X., Fauvel, T., Shine, R. & Brischoux, F. Foraging behaviour and energy budgets of sea snakes from New Caledonia: Insights from implanted data-loggers. J. Zool. 298, 82–93 (2016).
Google Scholar 
Bonnet, X., Brischoux, F., Briand, M. & Shine, R. Plasticity matches phenotype to local conditions despite genetic homogeneity across 13 snake populations. Proc. R. Soc. B 288, 20202916 (2021).PubMed 
PubMed Central 

Google Scholar 
Lukoschek, V., Waycott, M. & Marsh, H. Phylogeography of the olive sea snake, Aipysurus laevis (Hydrophiinae) indicates Pleistocene range expansion around northern Australia but low contemporary gene flow. Mol. Ecol. 16, 3406–3422 (2007).CAS 
PubMed 

Google Scholar 
Nitschke, C. R., Hourston, M., Udyawer, V. & Sanders, K. L. Rates of population differentiation and speciation are decoupled in sea snakes. Biol. Lett. 14, 20180563 (2018).PubMed 
PubMed Central 

Google Scholar 
Heatwole, H., Grech, A., Monahan, J. F., King, S. & Marsh, H. Thermal biology of sea snakes and sea kraits. Integr. Comp. Biol. 52, 257–273 (2012).PubMed 

Google Scholar 
Brischoux, F., Rolland, V., Bonnet, X., Caillaud, M. & Shine, R. Effects of oceanic salinity on body condition in sea snakes. Integr. Comp. Biol. 52, 235–244 (2012).PubMed 

Google Scholar 
Bonnet, X. et al. Spatial variation in age structure among populations of a colonial marine snake: the influence of ectothermy. J. Anim. Ecol. 84, 925–933 (2015).PubMed 

Google Scholar 
Heatwole, H. Sea Snakes 2nd edn. (Krieger Publishing, 1999).
Google Scholar 
Blouin-Demers, G. & Weatherhead, P. J. Thermal ecology of black rat snakes (Elaphe obsoleta) in a thermally challenging environment. Ecology 82, 3025–3043 (2001).
Google Scholar 
Goiran, C., Brown, G. P. & Shine, R. Niche partitioning within a population of sea snakes is constrained by ambient thermal homogeneity and small prey size. Biol. J. Linn. Soc. 129, 644–651 (2020).
Google Scholar 
Lowe, J. R. et al. Regional versus latitudinal variation in the life-history traits and demographic rates of a reef fish, Centropyge bispinosa, in the Coral Sea and Great Barrier Reef Marine Parks, Australia. J. Fish Biol. 99, 1602–1612. https://doi.org/10.1111/jfb.14865 (2021).PubMed 

Google Scholar 
Gust, N., Choat, J. & Ackerman, J. Demographic plasticity in tropical reef fishes. Mar. Biol. 140, 1039–1051 (2002).
Google Scholar 
Kingsford, M. J., Welch, D. & O’Callaghan, M. Latitudinal and cross-shelf patterns of size, age, growth, and mortality of a tropical damselfish Acanthochromis polyacanthus on the Great Barrier Reef. Diversity 11, 67 (2019).
Google Scholar  More

Global patterns of ARG distributionWe used a set of 4572 metagenomic samples to illustrate the global patterns of ARG distribution (Supplementary Data 1). These samples were collected from six types of habitats: air, aquatic, terrestrial, engineered, humans and other hosts (Fig. 1a and Supplementary Data 1). From these samples, we identified a total of 2561 ARGs that conferred resistance to 24 drug classes of antibiotics based on the Comprehensive Antibiotic Research Database (CARD). Of these, 2401 were genes conferring resistance to only one drug class, and 160 conferred resistances to multiple drug classes (Supplementary Data 2). Twenty-five ARGs were found in more than 75% samples, however, the frequency of most ARGs (2313/2561) were More

Alves, J. A. et al. Costs, benefits, and fitness consequences of different migratory strategies. Ecology 94(1), 11–17 (2013).PubMed 

Google Scholar 
Alexander, R. M. When is migration worthwhile for animals that walk, swim or fly?. J. Avian Biol. 29(4), 387–394 (1998).
Google Scholar 
Wikelski, M. et al. Costs of migration in free-flying songbirds. Nature 423, 704 (2003).ADS 
CAS 
PubMed 

Google Scholar 
Alerstam, T., Hedenström, A. & Åkesson, S. Long-distance migration: evolution and determinants. Oikos 103(2), 247–260 (2003).
Google Scholar 
Alerstam, T. Optimal bird migration revisited. J. Ornithol. 152, 5–23 (2011).
Google Scholar 
Hedenstrom, A. & Alerstam, T. Optimum fuel loads in migratory birds: Distinguishing between time and energy minimization. J. Theor. Biol. 189, 227–234 (1997).ADS 
CAS 
PubMed 

Google Scholar 
Åkesson, S. & Helm, B. Endogenous programs and flexibility in bird migration. Front. Ecol. Evol. 8, 78 (2020).
Google Scholar 
Jiguet, F. et al. Desert crossing strategies of migrant songbirds vary between and within species. Sci. Rep. 9(1), 20248 (2019).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Senner, N. R., Morbey, Y. E. & Sandercock, B. K. Editorial: Flexibility in the migration strategies of animals. Front. Ecol. Evol. 8, 111 (2020).
Google Scholar 
Mellone, U., López-López, P., Limiñana, R., Piasevoli, G. & Urios, V. The trans-equatorial loop migration system of Eleonora’s falcon: Differences in migration patterns between age classes, regions and seasons. J. Avian Biol. 44, 417–426 (2013).
Google Scholar 
Chevallier, D. et al. Influence of weather conditions on the flight of migrating black storks. Proc. Biol. Sci. 277(1695), 2755–2764 (2010).CAS 
PubMed 
PubMed Central 

Google Scholar 
Verhelst, B., Jansen, J. & Vansteelant, W. South West Georgia: An important bottleneck for raptor migration during autumn. Ardea 99, 137–146 (2011).
Google Scholar 
Klaassen, R. H. G., Strandberg, R., Hake, M. & Alerstam, T. Flexibility in daily travel routines causes regional variation in bird migration speed. Behav. Ecol. Sociobiol. 62(9), 1427–1432 (2008).
Google Scholar 
Alerstam, T. Detours in bird migration. J. Theor. Biol. 209(3), 319–331 (2001).ADS 
CAS 
PubMed 

Google Scholar 
Alerstam, T. & Hedenström, A. The development of bird migration theory. J. Avian Biol. 29(4), 343–369 (1998).
Google Scholar 
Liechti, F., Klaassen, M. & Bruderer, B. Predicting migratory flight altitudes by physiological migration models. Auk 117, 205–214 (2000).
Google Scholar 
Senner, N. R. et al. High-altitude shorebird migration in the absence of topographical barriers: Avoiding high air temperatures and searching for profitable winds. Proc. Biol. Sci. 2018, 285 (1881).
Google Scholar 
Norevik, G., Akesson, S., Andersson, A., Backman, J. & Hedenstrom, A. Flight altitude dynamics of migrating European nightjars across regions and seasons. J. Exp. Biol. 224(20), jeb242836 (2021).PubMed 
PubMed Central 

Google Scholar 
Hadjikyriakou, T. G., Nwankwo, E. C., Virani, M. Z. & Kirschel, A. N. G. Habitat availability influences migration speed, refueling patterns and seasonal flyways of a fly-and-forage migrant. Mov. Ecol. 8, 10 (2020).PubMed 
PubMed Central 

Google Scholar 
Strandberg, R., Klaassen, R. H. G., Olofsson, P. & Alerstam, T. Daily travel schedules of adult Eurasian Hobbies Falco subbuteo—Variability in flight hours and migration speed along the route. Ardea 97(3), 287–295 (2009).
Google Scholar 
Strandberg, R. & Alerstam, T. The strategy of fly-and-forage migration, illustrated for the osprey (Pandion haliaetus). Behav. Ecol. Sociobiol. 61(12), 1865–1875 (2007).
Google Scholar 
McKinnon, E. A. & Love, O. P. Ten years tracking the migrations of small landbirds: Lessons learned in the golden age of bio-logging. Auk 135(4), 834–856 (2018).
Google Scholar 
Backman, J. et al. Actogram analysis of free-flying migratory birds: New perspectives based on acceleration logging. J. Comp. Physiol. A Neuroethol. Sens. Neural Behav. Physiol. 203(6–7), 543–564 (2017).PubMed 
PubMed Central 

Google Scholar 
Evens, R., Beenaerts, N., Witters, N. & Artois, T. Study on the foraging behaviour of the European nightjar Caprimulgus europaeus reveals the need for a change in conservation strategy in Belgium. J. Avian Biol. 48(9), 1238–1245 (2017).
Google Scholar 
Evens, R. et al. Lunar synchronization of daily activity patterns in a crepuscular avian insectivore. Ecol. Evol. 10(14), 7106–7116 (2020).PubMed 
PubMed Central 

Google Scholar 
Liechti, F. et al. Miniaturized multi-sensor loggers provide new insight into year-round flight behaviour of small trans-Sahara avian migrants. Mov. Ecol. 6, 19 (2018).PubMed 
PubMed Central 

Google Scholar 
Liechti, F., Witvliet, W., Weber, R. & Bachler, E. First evidence of a 200-day non-stop flight in a bird. Nat. Commun. 4, 2554 (2013).ADS 
PubMed 

Google Scholar 
Dhanjal-Adams, K. L. PAMLr: Suite of functions for manipulating pressure, activity, magnetism and light data in R. (2020).Lisovski, S. et al. Light-level geolocator analyses: A user’s guide. J. Anim. Ecol. 89(1), 221–236 (2020).PubMed 

Google Scholar 
Lisovski, S. et al. Geolocation by light: Accuracy and precision affected by environmental factors. Methods Ecol. Evol. 3(3), 603–612 (2012).
Google Scholar 
Wotherspoon, S., Sumner, M., Lisovski, S. SGAT-Package: Solar/Satellite Geolocation for Animal Tracking. (2021). R package version 0.1.3. GitHub Repository.Bauer, R. RchivalTag: Analyzing Archival Tagging Data. R package version 0.1.2. (2020).Sjöberg, S. et al. Barometer logging reveals new dimensions of individual songbird migration. J. Avian Biol. 49(9), e01821 (2018).
Google Scholar 
Evens, R. et al. Migratory pathways, stopover zones and wintering destinations of Western European Nightjars Caprimulgus europaeus. Ibis 159(3), 680–686 (2017).
Google Scholar 
Becker, J. J. et al. Global bathymetry and elevation data at 30 arc seconds resolution: SRTM30_PLUS. Mar. Geodesy 32(4), 355–371 (2009).
Google Scholar 
Ricketts, T. H. Terrestrial Ecoregions of North America: A Conservation Assessment (Island Press, 1999).
Google Scholar 
Olson, D. M. et al. Terrestrial ecoregions of the world: A new map of life on earth. Bioscience 51(11), 933–938 (2001).
Google Scholar 
QGIS-Development-Team: QGIS Geographic Information System. Open Source Geospatial Foundation (2021).Vansteelant, W. M. G., Gangoso, L., Bouten, W., Viana, D. S. & Figuerola, J. Adaptive drift and barrier-avoidance by a fly-forage migrant along a climate-driven flyway. Mov. Ecol. 9(1), 37 (2021).PubMed 
PubMed Central 

Google Scholar 
Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9(2), 378–400 (2017).
Google Scholar 
Hartig, F. DHARMa: Residual Diagnostics for Hierarchical Multi-Level/Mixed) Regression Models. R package version 0.3.3.0. (2020).Lenth, R. emmeans: Estimated Marginal Means, aka Least-Squares Means. R package version 1.5.1. (2020).Akesson, S., Bianco, G. & Hedenstrom, A. Negotiating an ecological barrier: Crossing the Sahara in relation to winds by common swifts. Philos. Trans. R. Soc. Lond. B Biol. Sci. 371(1704), 20150393 (2016).PubMed 
PubMed Central 

Google Scholar 
Strandberg, R., Klaassen, R. H. G., Hake, M., Olofsson, P. & Alerstam, T. Converging migration routes of Eurasian Hobbies Falco subbuteo crossing the African equatorial rain forest. Proc. R. Soc. B 276, 727–733 (2009).PubMed 

Google Scholar 
Rodriguez-Ruiz, J. et al. Disentangling migratory routes and wintering grounds of Iberian near-threatened European Rollers Coracias garrulus. PLoS ONE 9(12), e115615 (2014).ADS 
PubMed 
PubMed Central 

Google Scholar 
Vickery, J. A. et al. The decline of Afro-Palaearctic migrants and an assessment of potential causes. Ibis 156, 1–22 (2014).
Google Scholar 
Evens, R. et al. Proximity of breeding and foraging areas affects foraging effort of a crepuscular, insectivorous bird. Sci. Rep. 8(1), 3008 (2018).ADS 
PubMed 
PubMed Central 

Google Scholar 
Conring, C. M., Brautigam, K., Grisham, B. A., Collins, D. P. & Conway, W. C. Identifying the migratory strategy of the Lower Colorado River Valley population of Greater Sandhill Cranes. Avian Conserv. Ecol. 14(1), 11 (2019).
Google Scholar 
Imlay, T. L., Saldanha, S. & Taylor, P. D. The fall migratory movements of Bank Swallows, Riparia riparia: Fly-and-forage migration?. Avian Conserv. Ecol. 15(1), 2 (2020).
Google Scholar 
Piersma, T. Hop, skip, or jump? Constraints on migration of arctic waders by feeding, fattening, and flight speed. Limosa 60, 185–194 (1987).
Google Scholar 
Warnock, N. Stopping vs. staging: The difference between a hop and a jump. J. Avian Biol. 41(6), 621–626 (2010).
Google Scholar 
Gomez, C. et al. Fuel loads acquired at a stopover site influence the pace of intercontinental migration in a boreal songbird. Sci. Rep. 7(1), 3405 (2017).ADS 
PubMed 
PubMed Central 

Google Scholar 
Tottrup, A. P. et al. The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration. Proc. Biol. Sci. 279(1730), 1008–1016 (2012).PubMed 

Google Scholar 
Lisovski, S. et al. Inherent limits of light-level geolocation may lead to over-interpretation. Curr. Biol. 28(3), R99–R100 (2018).CAS 
PubMed 

Google Scholar 
Buler, J. J., Moore, F. R. & Woltmann, S. A multi-scale examination of stopover habitat use by birds. Ecology 88(7), 1789–1802 (2007).PubMed 

Google Scholar 
Loon, A. V. et al. Migratory stopover timing is predicted by breeding latitude, not habitat quality, in a long-distance migratory songbird. J. Ornithol. 158(3), 745–752 (2017).
Google Scholar 
Norevik, G. et al. Wind-associated detours promote seasonal migratory connectivity in a flapping flying long-distance avian migrant. J. Anim. Ecol. 89(2), 635–646 (2020).PubMed 

Google Scholar 
Norevik, G., Åkesson, S. & Hedenström, A. Migration strategies and annual space-use in an Afro-Palaearctic aerial insectivore—The European nightjar Caprimulgus europaeus. J. Avian Biol. 48(5), 738–747 (2017).
Google Scholar 
Cresswell, B. & Edwards, D. Geolocators reveal wintering areas of European Nightjar (Caprimulgus europaeus). Bird Study 60(1), 77–86 (2013).
Google Scholar 
Jacobsen, L. B. et al. Annual spatiotemporal migration schedules in three larger insectivorous birds: European nightjar, common swift and common cuckoo. Anim. Biotelem. 5(1), 1–11 (2017).
Google Scholar 
Liechti, F. & Bruderer, B. The relevance of wind for optimal migration theory. J. Avian Biol. 29(4), 561–568 (1998).
Google Scholar 
Schmaljohann, H., Bruderer, B. & Liechti, F. Sustained bird flights occur at temperatures far beyond expected limits. Anim. Behav. 76(4), 1133–1138 (2008).
Google Scholar 
Schmaljohann, H., Liechti, F. & Bruderer, B. Trans-Sahara migrants select flight altitudes to minimize energy costs rather than water loss. Behav. Ecol. Sociobiol. 63(11), 1609–1619 (2009).
Google Scholar 
Sjöberg, S. et al. Extreme altitudes during diurnal flights in a nocturnal songbird migrant. Science 372, 646–648 (2021).ADS 
PubMed 

Google Scholar 
Bruderer, B., Peter, D. & Korner-Nievergelt, F. Vertical distribution of bird migration between the Baltic Sea and the Sahara. J. Ornithol. 159(2), 315–336 (2018).
Google Scholar  More

This is the first study analyzing mandible shape in both mink species and, together with a previous study on their cranial shape38, it has revealed how small morphological differences in highly similar species can lead to substantial biomechanical differences (see breakdown below). As with cranial shape, mandible shape in minks is influenced by the complex interaction of size and sexual dimorphism both at the inter- and intraspecific levels. However, while in cranial shape both species had divergent shape allometries and parallel interspecific sexual allometries, the opposite was true for mandible shape.Differences in mandible shape between European and American mink were summarized by PC1 (Fig. 2, Fig. S1) and can be mainly related to muscle size and jaw biomechanics (i.e., in-levers and out-levers). The relatively taller and slightly wider coronoid process of European minks suggests a relatively larger temporalis muscle, while the anteriorly expanded masseteric fossa of American mink is indicative of a relatively larger masseter complex17,22,25. The relatively enlarged angular process of European mink provides a larger attachment area for the superficial masseter, with both mink species having a distinctive fossa on the lateral side of the angular process where this muscle attaches. This angular fossa is not present in European polecats (Gálvez-López, pers. obs.), part of the sister clade to European mink41.Regarding jaw biomechanics, the particular morphology of the American mink illustrates the compromise between maximizing both bite force efficiency and increased gape. The MAs for all masticatory muscles were higher in European mink due to their relatively longer in-levers (and also shorter out-levers if measured on PC1 configurations), with the exception of the MA of the deep masseter which was considerably higher in American mink (Table S2; Fig. 1D). These findings indicate that American mink exhibit features that allow them to produce larger forces at wide gape, which is particularly useful for holding and killing terrestrial vertebrates22,42. In agreement with this, a short moment arm of the superficial masseter (as observed in American mink) has been associated with increased gape in other mammals43. It is also worth noting that low MAs for the posterior temporalis and superficial masseter have also been associated with fish capture, as they indicate a relatively longer mandible relative to the muscle in-levers, which in turn allows the mouth to close faster when trying to catch elusive prey underwater21. In contrast, the characteristic features of European mink are indicative of stronger bites at the carnassials, which would allow them to cut through relatively tougher tissues and also to crush harder objects (e.g. shells of aquatic prey). Favoring carnassial over anterior bites could also be advantageous to feeding on fish. Mink catch fish underwater by grabbing them by the fins or back with their anterior teeth, and then dragging them to the surface where they are processed using cheek (carnassial) bites (Gálvez-López, pers. obs.).In our previous study on cranial shape in mink38, morphological differences between both species indicated relatively larger muscle volumes overall in the American mink (temporalis: more developed sagittal and nuchal crests, narrower braincase; masseter: longer and more curved zygomatic arches, larger infratemporal fossa), which suggested that bite forces both at the anterior dentition and at the carnassials were larger in this species. However, when combined with the MA results from this study on mandible shape, the relationship between muscle volume and force production becomes less straightforward. In the case of the European mink, the relatively smaller temporalis has a larger attachment site on the mandible (i.e., a broader and taller coronoid) and becomes more efficient (i.e., has higher MAs) due to the relatively longer in-lever. Similarly, in the American mink the effective length of the superficial masseter is increased by the marked curvature of the zygomatic arches, which mitigates the dorsal displacement of the angular process. However, the efficiency of the relatively larger temporalis is diminished by a smaller coronoid (i.e., reduced attachment area and shorter in-levers). The remaining differences in cranial morphology align with differences in mandible shape. Namely, the relatively broader zygomatic arches of the European mink support a strong superficial masseter, while the larger infratemporal fossae of American mink account for their enlarged deep masseter. On a final note, another finding common to both cranial and mandible shape was the relatively larger crushing dentition of American mink.Thus, after combining the results of cranial and mandible shape, it appears that, while the characteristic features of European mink indeed allow stronger carnassial bites, American mink present morphological indicators of both strong killing bites at wide gapes and powerful carnassial bites with a marked crushing component.The allometric effect on mandible size common to both species was represented by PC2 (Fig. 2, Fig. S3), which complements the common allometric trend recovered for both mink species in cranial shape38. The relative expansion of the masseteric fossa and the angular process with increasing size suggests that larger mink present a larger masseter complex. However, most of the allometric shape changes are related to muscle in-levers and out-levers. With increasing size, the length of both the out-lever at the anterior teeth and the in-levers of its related muscles (anterior temporalis, deep masseter) increases (Table S2), but the in-levers scale faster than the out-lever (Table S2). Thus, the mechanical advantages of both muscles at the anterior teeth also increase with size (Table S2), indicating that larger mink have markedly stronger and more efficient killing bites (particularly true for the deep masseter, which also becomes larger with size). This, together with their relatively larger anterior dentition (both in the mandible and the cranium) and taller anterior corpus, can be related to feeding on larger prey as size increases (i.e., stronger bites to perforate tougher skulls and hold onto stronger struggling prey, which would also require more robust teeth and corpora to resist the stresses placed on them). Similar features have been described for felids18, which also kill prey in this way22,32.Note, however, that one of the shape changes along PC2 does not accurately reflect the common allometric pattern: the lever arm of the superficial masseter, which slightly decreases along PC2 (Fig. 2; Table S2) and results in a decrease of the mechanical advantage of the superficial masseter and hence bite force at the carnassials along this axis (Table S2). In contrast, this lever arm significantly increases with size in the original specimens (Table S2), in agreement with the common allometric trend in cranial shape suggesting stronger bites at all teeth with increasing size38. A likely explanation for this phenomenon is that the common allometric trend is being confounded with interspecific shape differences, as American mink have significantly shorter superficial masseter in-levers than European mink (Fig. 1F; Table S2) yet their males are significantly larger than all other specimens (Fig. 1A). As mentioned above, the relative decrease in MA might reflect the trade-off between producing strong bite forces at the anterior teeth and having a wider gape to capture larger prey43, both of which are heavily supported by other morphological features in this common allometric trend.Sexual dimorphism in mandible shape was significant both within each species, and when grouping sexes from both species together. In her study of Palearctic mustelids, Romaniuk28 also found evidence for interspecific sexual dimorphism in mandible shape, but within species it was only significant for the Siberian weasel (Mustela sibirica). The different results for the European mink in that study might be related to its smaller sample. Note, however, that Hernández-Romero et al.40 did not find evidence for sexual dimorphism in mandible shape within Neotropical otters (Lontra longicaudis) even though their sample sizes were equivalent to those in the present study.Overall, the results of the present study reveal that mandible shape differences between males and females are the consequence of a complex interaction between sex and size at both inter- and intraspecific levels. For instance, each sex in each species has a mandible shape significantly different from each other (Table 1), but allometric shape changes within each of them are similar (except maybe female American mink; Fig. S5A). Additionally, while trajectory analysis indicates that the degree of sexual dimorphism in mandible shape is similar within each species, the specific differences between sexes are different in each species (i.e., same magnitude, different orientation; Table 2, Fig. S5B). While at the interspecific level, male and female mandible shapes change differently with increasing size even though the change per unit size is similar in both sexes (Tables 1, 2; Fig. S5C,D), and some of the allometric changes are common to both species and sexes (see section above; PC2 in Fig. 2). Finally, another set of shape changes related to sexual dimorphism and common to both species are those related to sexual dimorphism in mandible size, illustrated by PC3 (Figs. 2, Fig. S4).Shape changes related to sexual dimorphism in size are represented along PC3 and can be related to an overall increase in bite force (i.e., at all teeth), as higher scores on this axis correspond to increased muscle attachment areas and longer in-levers (taller and wider coronoid, anteriorly expanded masseteric fossa, ventrally expanded angular process), shorter out-levers (particularly at the anterior teeth), and a more robust corpus (dorsoventrally and mediolaterally expanded). This interpretation of shape changes along PC3 is supported by the results of the ANOVAs on the lever arms and MAs measured on the PC3 configurations (Table S2). These variables were only related to sex and size, with female mink having longer out-levers and male mink presenting longer in-levers and higher MAs, while out-levers decreased with increasing size and in-levers and MAs increased in both sexes (no significant interaction between sex and size indicates parallel allometric trajectories in both sexes). This trend is consistent with the common sexual allometry described for cranial shape, which suggested that larger males have bigger masticatory muscles than smaller females and thus produce higher bite forces38. Additionally, even though the relative length of the toothrow decreases, the size of the canine markedly increases and there is no change in molar size or the relative proportions in its shearing and crushing regions. Although this might be interpreted as reinforcing the canines to cope with killing larger prey while maintaining an otherwise similar dietary regime20, it is worth noting that larger canines have been long described as a feature of sexual size dimorphism in mustelids19,44,45.In terms of interspecific differences in sexual allometry, with increasing size the following shape changes were observed in females but not in males (Fig. S5C): a dorsoventrally more robust corpus, a ventral expansion of the angular process, longer in-levers for all masticatory muscles, larger incisors, and an increase in the shearing portion of m1 relative to the crushing portion. Most of these shape changes are similar to those described for PC3, which suggests that the female interspecific allometry bridges the bite force gap caused by sexual dimorphism in size. The changes to the female dentition suggest a shift in diet from crushing tough food items (e.g. aquatic invertebrates) towards slicing meat, which makes sense since these changes occur simultaneously with the common allometric trend (related to improved capabilities for killing larger vertebrate prey). However, as noted earlier, the increased shearing component is also advantageous for a piscivorous diet. Shape changes in male mandibles not observed in females seem to emphasize the common allometric trend (i.e., stronger killing bite at larger gapes) (Fig. S5D): a wider coronoid process for more muscle attachment, a dorsally displaced angular process to allow wider gapes, and mediolateral expansion of the corpus to increase its strength. Regarding their dentition, the opposite trend to females was observed (i.e., slightly smaller anterior teeth and a longer crushing molar portion), suggesting a larger durophagous component in the diet of larger males.As expected, variation in mandible shape could be linked to potential dietary differences between European and American mink, and also between sexes. In summary, the results of the present study show that:

American mink are better equipped for preying on terrestrial vertebrates, as they can achieve relatively larger gapes and their mandibles are able to produce larger forces during the killing bite (i.e., at the anterior teeth and with an open mouth).

European mink, on the other hand, can produce relatively stronger bites at the carnassials, suggesting that they rely more on tougher prey and/or fish.

Regardless of species and sex, morphological features in larger mink demonstrate increased capabilities for feeding on larger terrestrial prey (stronger killing bites and more robust anterior teeth and corpora to resist the stresses caused by struggling prey).

Due to their larger size, male mink of both species have stronger bites than females at both the anterior teeth and the carnassials. However, with increasing size, females bridge the gap by developing relatively stronger bites overall while shifting their diet from tougher or harder prey (probably aquatic invertebrates) towards less mechanically demanding food items (e.g. terrestrial vertebrates and/or fish). In contrast, increasing size in males leads to even more specialization towards feeding on larger terrestrial prey while tough items become more relevant in their diets (probably crushing bones of small prey).

These findings confirm our original predictions based on previous results on cranial shape differences, but do they agree with observed dietary preferences in minks? Diet studies in American mink are numerous, and provide a wide picture of seasonal and regional variation8,11 as well as intraspecific dietary competition6,7,12. However, studies on European mink diet are scarcer9,14, particularly those comparing the sexes13. Additionally, a few studies have compared diets of sympatric European and American mink10,15. All these studies can be summarized as: A, male American mink favor medium-sized mammals and birds usually heavier than themselves; B, female American mink favor aquatic prey, but are displaced towards small mammals and birds when seasonal changes in prey availability shift the males’ diet towards aquatic prey; C, European mink favor aquatic prey, particularly fish and crayfish; but D, they are displaced towards amphibians and small mammals when sympatric with American mink. From these, our results on mandible shape variation support A and somewhat B and C, but provide no information on the interspecific competition scenario or on potential seasonal or local dietary differences. Additionally, there is no information on size-related dietary changes in either species that could validate our findings on sexual allometry in mandible shape. Thus, while mandible shape is very useful for identifying broad dietary indicators even between highly similar species, its ability to provide accurate information on their potential prey is limited.As a final note on mink diets, our previous study on cranial shape38, suggested a gradient in muscle force (and potential dietary range) from female European mink to male American mink. Based on those results and studies on social interactions between and within species35,46, we hypothesized that competition between both mink species could be displacing female European mink towards narrower and poorer diets, which could affect their survivability and ability to successfully reproduce. Fortunately, the results of the present study not only propose that there might be less overlap in diets between species and sexes than suggested by dietary studies7,10,13,15, but also indicate that dietary competition seems to be higher for small terrestrial vertebrates, not aquatic prey (on which female European mink are particularly well equipped to feed). More

Lees, A. C., Attwood, S., Barlow, J. & Phalan, B. Biodiversity scientists must fight the creeping rise of extinction denial. Nat. Ecol. Evol. 4, 1440–1443 (2020).PubMed 

Google Scholar 
Díaz, S. et al. Pervasive human-driven decline of life on Earth points to the need for transformative change. Science 366, eaax3100 (2019).
Google Scholar 
Driscoll, D. A. et al. A biodiversity-crisis hierarchy to evaluate and refine conservation indicators. Nat. Ecol. Evol. 2, 775–781 (2018).PubMed 

Google Scholar 
Jackson, J. B. et al. Historical overfishing and the recent collapse of coastal ecosystems. Science 293, 629–637 (2001).CAS 
PubMed 

Google Scholar 
McCauley, D. J. et al. Marine defaunation: Animal loss in the global ocean. Science 347, 6219 (2015).
Google Scholar 
Sala, E. & Knowlton, N. Global marine biodiversity trends. Annu. Rev. Environ. Resour. 31, 93–122 (2006).
Google Scholar 
Worm, B. et al. Impacts of biodiversity loss on ocean ecosystem services. Science 314, 787–790 (2006).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Beaumont, N. et al. Identification, definition and quantification of goods and services provided by marine biodiversity: Implications for the ecosystem approach. Mar. Pollut. Bull. 54, 253–265 (2007).CAS 
PubMed 

Google Scholar 
Hooper, D. U. et al. Effects of biodiversity on ecosystem functioning: A consensus of current knowledge. Ecol. Monogr. 75, 3–35 (2005).
Google Scholar 
Turpie, J. K. The existence value of biodiversity in South Africa: How interest, experience, knowledge, income and perceived level of threat influence local willingness to pay. Ecol. Econ. 46, 199–216 (2003).
Google Scholar 
Ruiz-Frau, A., Hinz, H., Edwards-Jones, G. & Kaiser, M. Spatially explicit economic assessment of cultural ecosystem services: Non-extractive recreational uses of the coastal environment related to marine biodiversity. Mar. Policy 38, 90–98 (2013).
Google Scholar 
Thrush, S. F., Gray, J. S., Hewitt, J. E. & Ugland, K. I. Predicting the effects of habitat homogenization on marine biodiversity. Ecol. Appl. 16, 1636–1642 (2006).PubMed 

Google Scholar 
Gillies, C. L. et al. Australian shellfish ecosystems: Past distribution, current status and future direction. PLoS ONE 13, e0190914 (2018).PubMed 
PubMed Central 

Google Scholar 
Commito, J. A., Como, S., Grupe, B. M. & Dow, W. E. Species diversity in the soft-bottom intertidal zone: Biogenic structure, sediment, and macrofauna across mussel bed spatial scales. J. Exp. Mar. Biol. Ecol. 366, 70–81 (2008).
Google Scholar 
Tokeshi, M. Species Coexistence: Ecological and Evolutionary Perspectives (Wiley, Hoboken, 2009).
Google Scholar 
Paul, L. J. A history of the Firth of Thames dredge fishrey for mussels: Use and abuse of a coastal resource. Report No. 94, (Wellington, New Zealand, 2012).Enderlein, P. & Wahl, M. Dominance of blue mussels versus consumer-mediated enhancement of benthic diversity. J. Sea Res. 51, 145–155 (2004).ADS 

Google Scholar 
Lejart, M. & Hily, C. Differential response of benthic macrofauna to the formation of novel oyster reefs (Crassostrea gigas, Thunberg) on soft and rocky substrate in the intertidal of the Bay of Brest, France. J. Sea Res. 65, 84–93 (2011).ADS 

Google Scholar 
Norling, P. & Kautsky, N. Patches of the mussel Mytilus sp. are islands of high biodiversity in subtidal sediment habitats in the Baltic Sea. Aquat. Biol. 4, 75–87 (2008).
Google Scholar 
Norling, P., Lindegarth, M., Lindegarth, S. & Strand, Å. Effects of live and post-mortem shell structures of invasive Pacific oysters and native blue mussels on macrofauna and fish. Mar. Ecol. Prog. Ser. 518, 123–138 (2015).ADS 

Google Scholar 
McLeod, I., Parsons, D., Morrison, M., Van Dijken, S. & Taylor, R. Mussel reefs on soft sediments: A severely reduced but important habitat for macroinvertebrates and fishes in New Zealand. N. Z. J. Mar. Freshw. Res. 48, 48–59 (2014).CAS 

Google Scholar 
Seitz, R. D., Wennhage, H., Bergström, U., Lipcius, R. N. & Ysebaert, T. Ecological value of coastal habitats for commercially and ecologically important species. ICES J. Mar. Sci. 71, 648–665 (2014).
Google Scholar 
zu Ermgassen, P. S., Grabowski, J. H., Gair, J. R. & Powers, S. P. Quantifying fish and mobile invertebrate production from a threatened nursery habitat. J. Appl. Ecol. 53, 596–606 (2016).
Google Scholar 
Grabowski, J. H. The influence of trophic interactions, habitat complexity, and landscape setting on community dynamics and restoration of oyster reefs. Ph.D., The University of North Carolina at Chapel Hill (2002).Harding, J. M., Allen, D. M., Haffey, E. R. & Hoffman, K. M. Site fidelity of oyster reef blennies and gobies in saltmarsh tidal creeks. Estuaries Coasts 43, 409–423 (2020).CAS 

Google Scholar 
Parsons, D. et al. Snapper (Chrysophrys auratus): A review of life history and key vulnerabilities in New Zealand. N. Z. J. Mar. Freshw. Res. 48, 256–283 (2014).
Google Scholar 
Callier, M. D., Richard, M., McKindsey, C. W., Archambault, P. & Desrosiers, G. Responses of benthic macrofauna and biogeochemical fluxes to various levels of mussel biodeposition: An in situ “benthocosm” experiment. Mar. Pollut. Bull. 58, 1544–1553. https://doi.org/10.1016/j.marpolbul.2009.05.010 (2009).CAS 
Article 
PubMed 

Google Scholar 
Ysebaert, T., Hart, M. & Herman, P. M. Impacts of bottom and suspended cultures of mussels Mytilus spp. on the surrounding sedimentary environment and macrobenthic biodiversity. Helgol. Mar. Res. 63, 59–74 (2009).ADS 

Google Scholar 
Sea, M. A., Thrush, S. F. & Hillman, J. R. Environmental predictors of sediment denitrification rates within restored green-lipped mussel (Perna canaliculus) beds. Mar. Ecol. Prog. Ser. 667, 1–13 (2021).ADS 
CAS 

Google Scholar 
Hillman, J. R., O’Meara, T. A., Lohrer, A. M., & Thrush, S. F. Influence of restored mussel reefs on denitrification in
marine sediments. J. Sea Res. 175, 102099 (2021).
Google Scholar 
Bacheler, N. M. et al. Comparison of trap and underwater video gears for indexing reef fish presence and abundance in the southeast United States. Fish. Res. 143, 81–88 (2013).
Google Scholar 
Wells, R. D., Boswell, K. M., Cowan, J. H. Jr. & Patterson, W. F. III. Size selectivity of sampling gears targeting red snapper in the northern Gulf of Mexico. Fish. Res. 89, 294–299 (2008).
Google Scholar 
Emslie, M. J., Cheal, A. J., MacNeil, M. A., Miller, I. R. & Sweatman, H. P. Reef fish communities are spooked by scuba surveys and may take hours to recover. PeerJ 6, e4886 (2018).PubMed 
PubMed Central 

Google Scholar 
Piggott, C. V., Depczynski, M., Gagliano, M. & Langlois, T. J. Remote video methods for studying juvenile fish populations in challenging environments. J. Exp. Mar. Biol. Ecol. 532, 151454 (2020).
Google Scholar 
Dean, W. E. Determination of carbonate and organic matter in calcareous sediments and sedimentary rocks by loss on ignition: Comparison with other methods. J. Sediment. Res. 44, 242–248 (1974).CAS 

Google Scholar 
Lorenzen, C. J. Determination of chlorophyll and pheo-pigments: Spectrophotometric equations. Limnol. Oceanogr. 12, 343–346 (1967).ADS 
CAS 

Google Scholar 
Anderson, M. J. A new method for non-parametric multivariate analysis of variance. Austral Ecol. 26, 32–46 (2001).
Google Scholar 
McArdle, B. H. & Anderson, M. J. Fitting multivariate models to community data: A comment on distance-based redundancy analysis. Ecology 82, 290–297 (2001).
Google Scholar 
Clarke, K. R. & Gorley, R. N. PRIMER v7: User Manual/Tutorial (2015).Anderson, M. J., Gorley, R. N. & Clarke, K. R. PERMANOVA+ for PRIMER: Guide to software and statistical methods (2008).R: A language and environment for statistical computing (R Foundation for Statistical Computing, Vienna, Austria, 2021).Saier, B. Subtidal and intertidal mussel beds (Mytilus edulis L.) in the Wadden Sea: Diversity differences of associated epifauna. Helgol. Mar. Res. 56, 44–50 (2002).ADS 

Google Scholar 
Peterson, C. H., Grabowski, J. H. & Powers, S. P. Estimated enhancement of fish production resulting from restoring oyster reef habitat: Quantitative valuation. Mar. Ecol. Prog. Ser. 264, 249–264 (2003).ADS 

Google Scholar 
Gutiérrez, J. L., Jones, C. G., Strayer, D. L. & Iribarne, O. O. Mollusks as ecosystem engineers: The role of shell production in aquatic habitats. Oikos 101, 79–90 (2003).
Google Scholar 
Norkko, A., Hewitt, J. E., Thrush, S. F. & Funnell, T. Benthic-pelagic coupling and suspension-feeding bivalves: Linking site-specific sediment flux and biodeposition to benthic community structure. Limnol. Oceanogr. 46, 2067–2072 (2001).ADS 

Google Scholar 
Russell, B. The food and feeding habits of rocky reef fish of north-eastern New Zealand. N. Z. J. Mar. Freshw. Res. 17, 121–145 (1983).
Google Scholar 
Gillies, C., Creighton, C. & McLeod, I. Shellfish reef habitats: A synopsis to underpin the repair and conservation of Australia’s environmentally, socially and economically important bays and estuaries. Report to the National Environmental Science Programme, Marine Biodiversity Hub, Centre for Tropical Water and Aquatic Ecosystem Research (TropWATER) Publication, James Cook University, Townsville, Qld, Australia (2015).Lenihan, H. S. et al. Cascading of habitat degradation: Oyster reefs invaded by refugee fishes escaping stress. Ecol. Appl. 11, 764–782 (2001).
Google Scholar 
Connell, S. & Jones, G. The influence of habitat complexity on postrecruitment processes in a temperate reef fish population. J. Exp. Mar. Biol. Ecol. 151, 271–294 (1991).
Google Scholar 
Usmar, N. Ontogeny and Ecology of Snapper (Pagrus auratus) in an estuary, the Mahurangi Harbour (University of Auckland, 2009).
Google Scholar 
Willis, T. J. & Anderson, M. J. Structure of cryptic reef fish assemblages: Relationships with habitat characteristics and predator density. Mar. Ecol. Prog. Ser. 257, 209–221 (2003).ADS 

Google Scholar 
Thompson, S. Homing in a territorial reef fish. Copeia 1983, 832–834 (1983).
Google Scholar 
Thrush, S. F., Schultz, D., Hewitt, J. E. & Talley, D. Habitat structure in soft-sediment environments and abundance of juvenile snapper Pagrus auratus. Mar. Ecol. Prog. Ser. 245, 273–280 (2002).ADS 

Google Scholar 
Pickering, H. & Whitmarsh, D. Artificial reefs and fisheries exploitation: A review of the ‘attraction versus production’debate, the influence of design and its significance for policy. Fish. Res. 31, 39–59 (1997).
Google Scholar 
Karp, M. A., Seitz, R. D. & Fabrizio, M. C. Faunal communities on restored oyster reefs: Effects of habitat complexity and environmental conditions. Mar. Ecol. Prog. Ser. 590, 35–51 (2018).ADS 

Google Scholar 
Hanke, M. H., Posey, M. H. & Alphin, T. D. The effects of intertidal oyster reef habitat characteristics on faunal utilization. Mar. Ecol. Prog. Ser. 581, 57–70 (2017).ADS 

Google Scholar 
Cranfield, H., Rowden, A., Smith, D., Gordon, D. & Michael, K. Macrofaunal assemblages of benthic habitat of different complexity and the proposition of a model of biogenic reef habitat regeneration in Foveaux Strait, New Zealand. J. Sea Res. 52, 109–125 (2004).ADS 

Google Scholar 
Norling, P. & Kautsky, N. Structural and functional effects of Mytilus edulis on diversity of associated species and ecosystem functioning. Mar. Ecol. Prog. Ser. 351, 163–175 (2007).ADS 

Google Scholar 
Jaunatre, R. et al. New synthetic indicators to assess community resilience and restoration success. Ecol. Indicators 29, 468–477 (2013).
Google Scholar 
O’Meara, T. A., Hewitt, J. E., Thrush, S. F., Douglas, E. J. & Lohrer, A. M. Denitrification and the role of macrofauna across estuarine gradients in nutrient and sediment loading. Estuaries Coasts 43, 1394–1405. https://doi.org/10.1007/s12237-020-00728-x (2020).CAS 
Article 

Google Scholar 
McCann, L. D. Oligochaete influence on settlement, growth and reproduction in a surface-deposit-feeding polychaete. J. Exp. Mar. Biol. Ecol. 131, 233–253 (1989).
Google Scholar 
Hope, J. A., Paterson, D. M. & Thrush, S. F. The role of microphytobenthos in soft-sediment ecological networks and their contribution to the delivery of multiple ecosystem services. J. Ecol. 108, 815–830 (2020).
Google Scholar 
Christianen, M. J. et al. Benthic primary producers are key to sustain the Wadden Sea food web: Stable carbon isotope analysis at landscape scale. Ecology 98, 1498–1512 (2017).CAS 
PubMed 

Google Scholar 
Commito, J. A. & Dankers, N. M. Dynamics of spatial and temporal complexity in European and North American soft-bottom mussel beds. In Ecological Comparisons of Sedimentary Shores, 39–59 (Springer, Berlin, 2001).Arribas, L. P., Donnarumma, L., Palomo, M. G. & Scrosati, R. A. Intertidal mussels as ecosystem engineers: Their associated invertebrate biodiversity under contrasting wave exposures. Mar. Biodivers. 44, 203–211 (2014).
Google Scholar 
Walles, B., Salvador de Paiva, J., van Prooijen, B. C., Ysebaert, T. & Smaal, A. C. The ecosystem engineer Crassostrea gigas affects tidal flat morphology beyond the boundary of their reef structures. Estuaries Coasts 38, 941–950 (2015).
Google Scholar 
Tsuchiya, M. & Nishihira, M. Islands of Mytilus edulis as a habitat for small intertidal animals: Effect of Mytilus age structure on the species composition of the associated fauna and community organization. Mar. Ecol. Prog. Ser. 31, 171–178 (1986).ADS 

Google Scholar 
Craeymeersch, J. A. & Jansen, H. M. Bivalve assemblages as hotspots for biodiversity. In Goods and Services of Marine Bivalves, 275–294 (Springer, Cham, 2019).Buschbaum, C. et al. Mytilid mussels: Global habitat engineers in coastal sediments. Helgol. Mar. Res. 63, 47–58 (2009).ADS 

Google Scholar  More

Wernberg, T., Krumhansl, K. A., Filbee-Dexter, K. & Pedersen, M. Status and trends for the world’s kelp forests. In World Seas: An Environmental Evaluation (ed. Sheppard, C.) 57–78 (Elsevier, 2019).Chapter 

Google Scholar 
Smale, D. A. et al. Marine heatwaves threaten global biodiversity and the provision of ecosystem services. Nat. Clim. Change 9, 306–312. https://doi.org/10.1038/s41558-019-0412-1 (2019).ADS 
Article 

Google Scholar 
Wernberg, T. et al. Climate-driven regime shift of a temperate marine ecosystem. Science 353, 169–172. https://doi.org/10.1126/science.aad8745 (2016).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Coleman, M. A., Minne, A. J. P., Vranken, S. & Wernberg, T. Genetic tropicalisation following a marine heatwave. Sci. Rep. UK 10, 12726. https://doi.org/10.1038/s41598-020-69665-w (2020).ADS 
CAS 
Article 

Google Scholar 
Krumhansl, K. A. et al. Global patterns of kelp forest change over the past half-century. Proc. Natl. Acad. Sci. 113, 13785–13790. https://doi.org/10.1073/pnas.1606102113 (2016).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Arafeh-Dalmau, N. et al. Extreme marine heatwaves alter kelp forest community near its equatorward distribution limit. Front. Mar. Sci. https://doi.org/10.3389/fmars.2019.00499 (2019).Article 

Google Scholar 
Tanaka, K., Taino, S., Haraguchi, H., Prendergast, G. & Hiraoka, M. Warming off southwestern Japan linked to distributional shifts of subtidal canopy-forming seaweeds. Ecol. Evol. 2, 2854–2865. https://doi.org/10.1002/ece3.391 (2012).Article 
PubMed 
PubMed Central 

Google Scholar 
Wernberg, T. et al. Genetic diversity and kelp forest vulnerability to climatic stress. Sci. Rep. UK 8, 1851. https://doi.org/10.1038/s41598-018-20009-9 (2018).ADS 
CAS 
Article 

Google Scholar 
Graham, M. H., Kinlan, B. P., Druehl, L. D., Garske, L. E. & Banks, S. Deep-water kelp refugia as potential hotspots of tropical marine diversity and productivity. Proc. Natl. Acad. Sci. 104, 16576. https://doi.org/10.1073/pnas.0704778104 (2007).ADS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Marzinelli, E. M. et al. Large-scale geographic variation in distribution and abundance of Australian deep-water kelp forests. PLoS ONE 10, e0118390. https://doi.org/10.1371/journal.pone.0118390 (2015).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Varela, R., Rodríguez-Díaz, L., de Castro, M. & Gómez-Gesteira, M. Influence of Eastern Upwelling systems on marine heatwaves occurrence. Glob. Planet Change 196, 103379. https://doi.org/10.1016/j.gloplacha.2020.103379 (2021).Article 

Google Scholar 
Assis, J. et al. Deep reefs are climatic refugia for genetic diversity of marine forests. J. Biogeogr. 43, 833–844. https://doi.org/10.1111/jbi.12677 (2016).Article 

Google Scholar 
Lourenço, C. R. et al. Upwelling areas as climate change refugia for the distribution and genetic diversity of a marine macroalga. J. Biogeogr. 43, 1595–1607. https://doi.org/10.1111/jbi.12744 (2016).Article 

Google Scholar 
Vranken, S. et al. Genotype-environment mismatch of kelp forests under climate change. Mol. Ecol. 30, 3730–3746. https://doi.org/10.1111/mec.15993 (2021).Article 
PubMed 

Google Scholar 
Wood, G. et al. Genomic vulnerability of a dominant seaweed points to future-proofing pathways for Australia’s underwater forests. Glob. Change Biol. 27, 2200–2212. https://doi.org/10.1111/gcb.15534 (2021).ADS 
Article 

Google Scholar 
Wernberg, T. et al. Biology and ecology of the globally significant kelp Ecklonia radiata. Oceanogr. Mar. Biol.: An Annu. Rev. 57, 265–324 (2019).Article 

Google Scholar 
Durrant, H. M. S., Barrett, N. S., Edgar, G. J., Coleman, M. A. & Burridge, C. P. Shallow phylogeographic histories of key species in a biodiversity hotspot. Phycologia 54, 556–565. https://doi.org/10.2216/15-24.1 (2015).Article 

Google Scholar 
Rothman, M. D. et al. A molecular investigation of the genus Ecklonia (Phaeophyceae, Laminariales) with special focus on the southern hemisphere. J. Phycol. 51, 236–246. https://doi.org/10.1111/jpy.12264 (2015).CAS 
Article 
PubMed 

Google Scholar 
Starko, S. et al. A comprehensive kelp phylogeny sheds light on the evolution of an ecosystem. Mol. Phylogenetics Evol. 136, 138–150. https://doi.org/10.1016/j.ympev.2019.04.012 (2019).Article 

Google Scholar 
Shepherd, S. A. & Edgar, G. J. In (eds Shepherd, S. A. & Edgar, G. J.) (CSIRO Publishing, 2013).Guiry, M. D. et al. AlgaeBase: An on-line resource for algae. Cryptogam. Algol. 35, 105–115, 111 (2014).Barratt, L., Ormond, R. F. G. & Wrathall, T. J. Ecological studies of southern Oman kelp communities. Part 1. Ecology and productivity of the sublittoral algae Ecklonia radiata and Sargassopsis zanardinii (Council for the conservation of the environment and water resources, and regional organisation for the protection of the marine environment, Muscat and Kuwait, 1986).Barratt, L. et al. An ecological study of the rocky shores on the south coast of Oman. Report of IUCN to UNEP’s regional seas programme, Vol. 104 (Tropical Marine Research Unit, York, 1984).Klaus, R. & Turner, J. R. The marine biotopes of the Socotra Archipelago. Fauna Arab. 20, 45–116 (2004).
Google Scholar 
Claereboudt, M. R. Oman. In World Seas: An Environmental Evaluation, (ed. Sheppard, C.) 25–47 (Academic Press, 2019).Savidge, G., Lennon, H. J. & Matthews, A. D. A shore based survey of oceanographic variables in the Dhofar region of southern Oman, August–October 1985. In Ecological Studies of Southern Oman Kelp Communities. Summary Report, 4–21. ROPME/GC-6/001 (1988).Hatcher, B. G., Kirkman, H. & Wood, W. F. Growth of the kelp Ecklonia-radiata near the northern limit of its range in Western-Australia. Mar. Biol. 95, 63–73. https://doi.org/10.1007/Bf00447486 (1987).Article 

Google Scholar 
Veenhof, R. et al. Kelp gametophytes in changing oceans. Oceanogr. Mar. Biol. Annu. Rev. 60 (in press).Goes, J. I., Thoppil, P. G., Gomes, H. D. R. & Fasullo, J. T. Warming of the Eurasian landmass is making the Arabian sea more productive. Science 308, 545. https://doi.org/10.1126/science.1106610 (2005).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Roxy, M. K. et al. A reduction in marine primary productivity driven by rapid warming over the tropical Indian Ocean. Geophys. Res. Lett. 43, 826–833. https://doi.org/10.1002/2015GL066979 (2016).ADS 
Article 

Google Scholar 
Watanabe, T. K., Watanabe, T., Yamazaki, A., Pfeiffer, M. & Claereboudt, M. R. Oman coral δ18O seawater record suggests that Western Indian Ocean upwelling uncouples from the Indian Ocean Dipole during the global-warming hiatus. Sci. Rep. UK 9, 1887. https://doi.org/10.1038/s41598-018-38429-y (2019).ADS 
CAS 
Article 

Google Scholar 
Watanabe, T. K. et al. Past summer upwelling events in the Gulf of Oman derived from a coral geochemical record. Sci. Rep. UK 7, 4568. https://doi.org/10.1038/s41598-017-04865-5 (2017).ADS 
CAS 
Article 

Google Scholar 
Edwards, M. & Estes, J. A. Catastrophe, recovery and range limitation in NE Pacific kelp forests: a large-scale perspective. Mar. Ecol. Prog. Ser. 320, 79–87 (2006).ADS 
Article 

Google Scholar 
Glynn, P. W. Monsoonal upwelling and episodic Acanthaster predation as probable controls of coral reef distribution and community structure in Oman, Indian Ocean. Atoll Res. Bull. 379, 1–66 (1993).Article 

Google Scholar 
Hiscock, S., Barratt, L. & Ormond, R. The marine algae of Dhofar, Oman-an upwelling system in the Arabian Sea. Br. Phycol. J. 19, 194 (1984).Article 

Google Scholar 
Kirkman, H. The 1st year in the life-history and the survival of the juvenile marine macrophyte, Ecklonia-radiata (Turn) J Agardh. J. Exp. Mar. Biol. Ecol. 55, 243–254. https://doi.org/10.1016/0022-0981(81)90115-5 (1981).Article 

Google Scholar 
Maeda, T., Kawai, T., Nakaoka, M. & Yotsukura, N. Effective DNA extraction method for fragment analysis using capillary sequencer of the kelp, Saccharina. J. Appl. Phycol. 25, 337–347 (2013).CAS 
Article 

Google Scholar 
Voisin, M., Engel, C. R. & Viard, F. Differential shuffling of native genetic diversity across introduced regions in a brown alga: Aquaculture vs. maritime traffic effects. Proc. Natl. Acad. Sci. USA 102, 5432. https://doi.org/10.1073/pnas.0501754102 (2005).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Lane, C. E., Lindstrom, S. C. & Saunders, G. W. A molecular assessment of northeast Pacific Alaria species (Laminariales, Phaeophyceae) with reference to the utility of DNA barcoding. Mol. Phylogenetics Evol. 44, 634–648 (2007).CAS 
Article 

Google Scholar 
Kearse, M. et al. Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28, 1647–1649. https://doi.org/10.1093/bioinformatics/bts199 (2012).Article 
PubMed 
PubMed Central 

Google Scholar 
Johnson, M. et al. NCBI BLAST: A better web interface. Nucl. Acids Res. 36, W5–W9 (2008).CAS 
Article 

Google Scholar 
Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A. & Minh, B. Q. W-IQ-TREE: A fast online phylogenetic tool for maximum likelihood analysis. Nucl. Acids Res. 44, W232–W235 (2016).CAS 
Article 

Google Scholar 
Kalyaanamoorthy, S., Minh, B. Q., Wong, T. K., Von Haeseler, A. & Jermiin, L. S. ModelFinder: Fast model selection for accurate phylogenetic estimates. Nat. Methods 14, 587–589 (2017).CAS 
Article 

Google Scholar 
Chernomor, O., von Haeseler, A. & Minh, B. Q. Terrace aware data structure for phylogenomic inference from supermatrices. Syst. Biol. 65, 997–1008. https://doi.org/10.1093/sysbio/syw037 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Hoang, D. T., Chernomor, O., Von Haeseler, A., Minh, B. Q. & Vinh, L. S. UFBoot2: Improving the ultrafast bootstrap approximation. Mol. Biol. Evol. 35, 518–522 (2018).CAS 
Article 

Google Scholar 
Rambaut, A. & Drummond, A. FigTree: Tree Figure Drawing Tool, Version 1.2. 2 (Institute of Evolutionary Biology, University of Edinburgh, 2008).
Google Scholar 
Rozas, J. et al. DnaSP 6: DNA sequence polymorphism analysis of large data sets. Mol. Biol. Evol. 34, 3299–3302. https://doi.org/10.1093/molbev/msx248 (2017).CAS 
Article 
PubMed 

Google Scholar 
Clement, M., Posada, D. & Crandall, K. A. TCS: A computer program to estimate gene genealogies. Mol. Ecol. 9, 1657–1659. https://doi.org/10.1046/j.1365-294x.2000.01020.x (2000).CAS 
Article 
PubMed 

Google Scholar 
Leigh, J. W. & Bryant, D. popart: Full-feature software for haplotype network construction. Methods Ecol. Evol. 6, 1110–1116 (2015).Article 

Google Scholar 
Team, R. C. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2019).Vergés, A. et al. Long-term empirical evidence of ocean warming leading to tropicalization of fish communities, increased herbivory, and loss of kelp. Proc. Natl. Acad. Sci. 113, 13791–13796. https://doi.org/10.1073/pnas.1610725113 (2016).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Wood, G. et al. Using genetics to test provenance effects and to optimise seaweed restoration. J. Appl. Ecol. https://doi.org/10.1111/1365-2664.13707 (2020).Article 

Google Scholar 
Wynne, M. J. A checklist of the benthic marine algae of the Northern Arabian Sea coast of the Sultanate of Oman. Bot. Mar. 61, 481–498. https://doi.org/10.1515/bot-2018-0035 (2018).Richards, G. & Wynne, M. J. 57 (2003).Schils, T. Marine Plant Communities of Upwelling Areas Within the Arabian Sea: A Taxonomic, Ecological ABD Biogeographic Case Study on the Marine Flora of the Socotra Archipelago (Yemen) and Masirah Island (Oman). PhD thesis (2002).Schils, T. & Coppejans, E. Phytogeography of upwelling areas in the Arabian Sea. J. Biogeogr. 30, 1339–1356. https://doi.org/10.1046/j.1365-2699.2003.00933.x (2003).Article 

Google Scholar 
Schils, T. & Wilson, S. C. temperature threshold as a biogeographic barrier in northern Indian Ocean Macroalgae. J. Phycol. 42, 749–756. https://doi.org/10.1111/j.1529-8817.2006.00242.x (2006).Article 

Google Scholar 
Wiggert, J. D., Hood, R. R., Banse, K. & Kindle, J. C. Monsoon-driven biogeochemical processes in the Arabian Sea. Prog. Oceanogr. 65, 176–213. https://doi.org/10.1016/j.pocean.2005.03.008 (2005).ADS 
Article 

Google Scholar 
Serisawa, Y., Imoto, Z., Ishikawa, T. & Ohno, M. Decline of the Ecklonia cava population associated with increased seawater temperatures in Tosa Bay, southern Japan. Fish. Sci. 70, 189–191. https://doi.org/10.1111/j.0919-9268.2004.00788.x (2004).CAS 
Article 

Google Scholar 
Nelson, W., Duffy, C., Trnski, T. & Stewart, R. Mesophotic Ecklonia radiata (Laminariales) at Rangitāhua, Kermadec Islands, New Zealand. Phycologia 57, 534–538. https://doi.org/10.2216/18-9.1 (2018).Article 

Google Scholar 
Richmond, S. & Stevens, T. Classifying benthic biotopes on sub-tropical continental shelf reefs: How useful are abiotic surrogates?. Estuar. Coast. Shelf Sci. 138, 79–89. https://doi.org/10.1016/j.ecss.2013.12.012 (2014).ADS 
Article 

Google Scholar 
Davis, T. R., Champion, C. & Coleman, M. A. Climate refugia for kelp within an ocean warming hotspot revealed by stacked species distribution modelling. Mar. Environ. Res. 166, 105267. https://doi.org/10.1016/j.marenvres.2021.105267 (2021).CAS 
Article 
PubMed 

Google Scholar 
Jooste, C. M., Oliver, J., Emami-Khoyi, A. & Teske, P. R. Is the Wild Coast in eastern South Africa a distinct marine bioregion?. Helgol. Mar. Res. 72, 6. https://doi.org/10.1186/s10152-018-0509-3 (2018).Article 

Google Scholar 
Bolton, J. J. et al. Where is the western limit of the tropical Indian Ocean seaweed flora? An analysis of intertidal seaweed biogeography on the east coast of South Africa. Mar. Biol. 144, 51–59. https://doi.org/10.1007/s00227-003-1182-9 (2004).Article 

Google Scholar 
Bolton, J. J. The biogeography of kelps (Laminariales, Phaeophyceae): A global analysis with new insights from recent advances in molecular phylogenetics. Helgol. Mar. Res. 64, 263–279. https://doi.org/10.1007/s10152-010-0211-6 (2010).ADS 
Article 

Google Scholar 
Bolton JJ, De Clerck O, John DM (2003). Seaweed diversity patterns in Sub-Saharan Africa. In Proceedings of the Marine Biodiversity in Sub-Saharan Africa: The Known and the Unknown. (eds. Decker, C. et al. ) Cape Town, South Africa, pp. 229–241 (2003).Wood, M. et al. Zanzibar and Indian Ocean trade in the first millennium CE: The glass bead evidence. Archaeol. Anthropol. Sci. 9, 879–901. https://doi.org/10.1007/s12520-015-0310-z (2017).Article 

Google Scholar 
Pollard, E., Bates, R., Ichumbaki, E. B. & Bita, C. Shipwreck evidence from Kilwa, Tanzania. Int. J. Naut. Archaeol. 45, 352–369. https://doi.org/10.1111/1095-9270.12185 (2016).Article 

Google Scholar 
Staples, M. In Oman. A Maritime History (eds Al Salimi, A. & Staples, E.) Chap. 4, 81–116 (Georg Olms Verlag, 2017).Assis, J. et al. Past climate changes and strong oceanographic barriers structured low-latitude genetic relics for the golden kelp Laminaria ochroleuca. J. Biogeogr. 45, 2326–2336. https://doi.org/10.1111/jbi.13425 (2018).Article 

Google Scholar 
Wade, R. et al. Macroalgal germplasm banking for conservation, food security, and industry. PLoS Biol. 18, e3000641. https://doi.org/10.1371/journal.pbio.3000641 (2020).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Coleman, M. A. et al. Restore or redefine: Future trajectories for restoration. Front. Mar. Sci. https://doi.org/10.3389/fmars.2020.00237 (2020).Article 

Google Scholar  More

Kelley, N. P. & Pyenson, N. D. Evolutionary innovation and ecology in marine tetrapods from the Triassic to the Anthropocene. Science 348, aaa3716 (2015).PubMed 

Google Scholar 
Gutarra, S. & Rahman, I. A. The locomotion of extinct secondarily aquatic tetrapods. Biol. Rev. 97, 67–98 (2022).PubMed 

Google Scholar 
Owen, R. A description of a portion of the skeleton of the Cetiosaurus, a gigantic extinct saurian reptile occurring in the oolitic formations of different portions of England. Proc. Geol. Soc. Lond. 3, 457–462 (1841).
Google Scholar 
Cope, E. On the characters of the skull in the Hadrosauridae. Proc. Natl Acad. Nat. Sci. USA 35, 97–107 (1883).
Google Scholar 
Bidar, A., Demay, L. & Thomel, G. Compsognathus corallestris, une nouvelle espèce de dinosaurien théropode du Portlandien de Canjuers (Sud-Est de la France). Annales Muséum d’Histoire Naturelle de Nice 1, 9–40 (1972).
Google Scholar 
Norell, M. A., Makovicky, P. J. & Currie, P. J. The beaks of ostrich dinosaurs. Nature 412, 873–874 (2001).ADS 
CAS 
PubMed 

Google Scholar 
Tereschenko, V. S. Adaptive features of protoceratopoids (Ornithischia: Neoceratopsia). Paleontol. J. 42, 273–286 (2008).
Google Scholar 
Lee, Y. N. et al. Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus. Nature 515, 257–260 (2014).ADS 
CAS 
PubMed 

Google Scholar 
Ibrahim, N. et al. Semiaquatic adaptations in a giant predatory dinosaur. Science 345, 1613–1616 (2014).ADS 
CAS 
PubMed 

Google Scholar 
Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395–399 (2017).ADS 
CAS 
PubMed 

Google Scholar 
Ibrahim, N. et al. Tail-propelled aquatic locomotion in a theropod dinosaur. Nature 581, 67–70 (2020).ADS 
CAS 
PubMed 

Google Scholar 
Henderson, D. M. A buoyancy, balance and stability challenge to the hypothesis of a semi-aquatic Spinosaurus Stromer, 1915 (Dinosauria: Theropoda). PeerJ 6, e5409 (2018).PubMed 
PubMed Central 

Google Scholar 
Hone, D. W. E. & Holtz, T. R. Jr Evaluating the ecology of Spinosaurus: shoreline generalist or aquatic pursuit specialist? Palaeontol. Electronica 24, a03 (2021).
Google Scholar 
Thewissen, J. G., Cooper, L. N., Clementz, M. T., Bajpai, S. & Tiwari, B. N. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature 450, 1190–1194 (2007).ADS 
CAS 
PubMed 

Google Scholar 
Houssaye, A. Bone histology of aquatic reptiles: what does it tell us about secondary adaptation to an aquatic life? Biol. J. Linn. Soc. 108, 3–21 (2013).
Google Scholar 
Motani, R. et al. A basal ichthyosauriform with a short snout from the Lower Triassic of China. Nature 517, 485–488 (2015).ADS 
CAS 
PubMed 

Google Scholar 
Rauhut, O. W. & Pol, D. Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Sci. Rep. 9, 1–9 (2019).
Google Scholar 
You, H. L. et al. A nearly modern amphibious bird from the Early Cretaceous of northwestern China. Science 312, 1640–1643 (2006).ADS 
CAS 
PubMed 

Google Scholar 
Wilson, L. E. & Chin, K. Comparative osteohistology of Hesperornis with reference to pygoscelid penguins: the effects of climate and behaviour on avian bone microstructure. R. Soc. Open Sci. 1, 140245 (2014).ADS 
PubMed 
PubMed Central 

Google Scholar 
Gatesy, S. M. & Dial, K. P. Locomotor modules and the evolution of avian flight. Evolution 50, 331–340 (1996).PubMed 

Google Scholar 
Amiot, R. et al. Oxygen isotope evidence for semi-aquatic habits among spinosaurid theropods. Geology 38, 139–142 (2010).ADS 
CAS 

Google Scholar 
Hassler, A. et al. Calcium isotopes offer clues on resource partitioning among Cretaceous predatory dinosaurs. Proc. R. Soc. B 285, 20180197 (2018).PubMed 
PubMed Central 

Google Scholar 
Larramendi, A., Paul, G. S. & Hsu, S. Y. A review and reappraisal of the specific gravities of present and past multicellular organisms, with an emphasis on tetrapods. Anat. Rec. 304, 1833–1888 (2021).
Google Scholar 
Charig, A. J. & Milner, A. C. Baryonyx, a remarkable new theropod dinosaur. Nature 324, 359–361 (1986).ADS 
CAS 
PubMed 

Google Scholar 
Schoener, T. W. The newest synthesis: understanding the interplay of evolutionary and ecological dynamics. Science 331, 426–429 (2011).ADS 
CAS 
PubMed 

Google Scholar 
Houssaye, A. “Pachyostosis” in aquatic amniotes: a review. Integr. Zool. 4, 325–340 (2009).PubMed 

Google Scholar 
Houssaye, A., Sander, M. P. & Klein, N. Adaptive patterns in aquatic amniote bone microanatomy—more complex than previously thought. Integr. Comp. Biol. 56, 1349–1369 (2016).PubMed 

Google Scholar 
Quemeneur, S., De Buffrenil, V. & Laurin, M. Microanatomy of the amniote femur and inference of lifestyle in limbed vertebrates. Biol. J. Linn. Soc. 109, 644–655 (2013).
Google Scholar 
Canoville, A., de Buffrénil, V. & Laurin, M. Microanatomical diversity of amniote ribs: an exploratory quantitative study. Biol. J. Linn. Soc. 118, 706–733 (2016).
Google Scholar 
Amson, E., de Muizon, C., Laurin, M., Argot, C. & de Buffrénil, V. Gradual adaptation of bone structure to aquatic lifestyle in extinct sloths from Peru. Proc. R. Soc. B 281, 20140192 (2014).PubMed 
PubMed Central 

Google Scholar 
Grafen, A. The phylogenetic regression. Philos. Trans. R. Soc. B 326, 119–157 (1989).ADS 
CAS 

Google Scholar 
Liem, K. F. Adaptive significance of intra-and interspecific differences in the feeding repertoires of cichlid fishes. Am. Zool. 20, 295–314 (1980).
Google Scholar 
Turner, A. H., Pol, D., Clarke, J. A., Erickson, G. M. & Norell, M. A. A basal dromaeosaurid and size evolution preceding avian flight. Science 317, 1378–1381 (2007).ADS 
CAS 
PubMed 

Google Scholar 
Voeten, D. F. et al. Wing bone geometry reveals active flight in Archaeopteryx. Nat. Commun. 9, 1319 (2018).
Google Scholar 
Houssaye, A., Martin, F., Boisserie, J. R. & Lihoreau, F. Paleoecological inferences from long bone microanatomical specializations in Hippopotamoidea (Mammalia, Artiodactyla). J. Mamm. Evol. 28, 847–870 (2021).
Google Scholar 
Amson, E. & Bibi, F. Differing effects of size and lifestyle on bone structure in mammals. BMC Biol. 19, 87 (2021).CAS 
PubMed 
PubMed Central 

Google Scholar 
Malafaia, E. et al. A new spinosaurid theropod (Dinosauria: Megalosauroidea) from the upper Barremian of Vallibona, Spain: Implications for spinosaurid diversity in the Early Cretaceous of the Iberian Peninsula. Cret. Res. 106, 104221 (2020).
Google Scholar 
Sereno, P. C. et al. A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids. Science 282, 1298–1302 (1998).ADS 
CAS 
PubMed 

Google Scholar 
Aureliano, T. et al. Semi-aquatic adaptations in a spinosaur from the Lower Cretaceous of Brazil. Cret. Res. 90, 283–295 (2018).
Google Scholar 
Barker, C. T. et al. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Sci. Rep. 11, 19340 (2021).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Taquet, P. Géologie et Paléontologie du Gisement de Gadoufaoua (Aptien du Niger) (Éditions du Centre national de la Recherche Scientifique, 1976).Rayfield, E. J., Milner, A. C., Xuan, V. B. & Young, P. G. Functional morphology of spinosaur ‘crocodile-mimic’ dinosaurs. J. Vertebr. Paleontol. 27, 892–901 (2007).
Google Scholar 
Benson, R. B., Butler, R. J., Carrano, M. T. & O’Connor, P. M. Air‐filled postcranial bones in theropod dinosaurs: physiological implications and the ‘reptile’–bird transition. Biol. Rev. 87, 168–193 (2012).PubMed 

Google Scholar 
Reid, R. E. H. Zonal “growth rings” in dinosaurs. Mod. Geol. 15, 19–48 (1990).
Google Scholar 
Chinsamy, A. & Raath, M. A. Preparation of fossil bone for histological examination. Palaeont. Afr. 29, 39–44 (1992).
Google Scholar 
Griffin, C. T. et al. Assessing ontogenetic maturity in extinct saurian reptiles. Biol. Rev. 96, 470–525 (2021).
Google Scholar 
Carrano, M. T., Benson, R. B. & Sampson, S. D. The phylogeny of Tetanurae (Dinosauria: Theropoda). J. Syst. Palaeontol. 10, 211–300 (2012).
Google Scholar 
Ibrahim, N. et al. Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco. ZooKeys 928, 1–216 (2020).PubMed 
PubMed Central 

Google Scholar 
Smyth, R. S., Ibrahim, N. & Martill, D. M. Sigilmassasaurus is Spinosaurus: a reappraisal of African spinosaurines. Cret. Res. 114, 104520 (2020).
Google Scholar 
Goloboff, P. A., Farris, J. S. & Nixon, K. C. TNT, a free program for phylogenetic analysis. Cladistics 24, 774–786 (2008).
Google Scholar 
Erickson, G. M. Assessing dinosaur growth patterns: a microscopic revolution. Trends Ecol. Evol. 20, 677–684 (2005).PubMed 

Google Scholar 
Hayashi, S. et al. Bone inner structure suggests increasing aquatic adaptations in Desmostylia (Mammalia, Afrotheria). PLoS ONE 8, e59146 (2013).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Straehl, F. R., Scheyer, T. M., Forasiepi, A. M., MacPhee, R. D. E. & Sánchez-Villagra, M. R. Evolutionary patterns of bone histology and bone compactness in xenarthran mammal long bones. PLoS ONE 8, e69275 (2013).ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Houssaye, A., Tafforeau, P., de Muizon, C. & Gingerich, P. D. Transition of Eocene whales from land to sea: evidence from bone microstructure. PLoS ONE 10, e0118409 (2015).PubMed 
PubMed Central 

Google Scholar 
Girondot, M. & Laurin, M. Bone profiler: a tool to quantify, model, and statistically compare bone-section compactness profiles. J. Vertebr. Paleontol. 23, 458–461 (2003).
Google Scholar 
De Ricqlès, A. J., Padian, K., Horner, J. R., Lamm, E. T. & Myhrvold, N. Osteohistology of Confuciusornis sanctus (Theropoda: Aves). Journ. Vertebr. Paleontol. 23, 373–386 (2003).
Google Scholar 
Maddison, W. P. Mesquite: a modular system for evolutionary analysis. Evolution 62, 1103–1118 (2008).
Google Scholar 
Upham, N. S., Esselstyn, J. A. & Jetz, W. Inferring the mammal tree: species-level sets of phylogenies for questions in ecology, evolution, and conservation. PLoS Biol. 17, e3000494 (2019).CAS 
PubMed 
PubMed Central 

Google Scholar 
Simoes, T. R. et al. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature 557, 706–709 (2018).ADS 
CAS 
PubMed 

Google Scholar 
Nesbitt, S. J. et al. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544, 484–487 (2017).ADS 
CAS 
PubMed 

Google Scholar 
Langer, M. C. et al. Untangling the dinosaur family tree. Nature 551, E1–E3 (2017).PubMed 

Google Scholar 
Brusatte, S. L., Lloyd, G. T., Wang, S. C. & Norell, M. A. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Curr. Biol. 24, 2386–2392 (2014).CAS 
PubMed 

Google Scholar 
Prum, R. O. et al. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526, 569–573 (2015).ADS 
CAS 
PubMed 

Google Scholar 
Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Methods Ecol. Evol. 3, 803–807 (2012).
Google Scholar 
Schmitz, L. & Motani, R. Nocturnality in dinosaurs inferred from scleral ring and orbit morphology. Science 332, 705–708 (2011).ADS 
CAS 
PubMed 

Google Scholar 
Motani, R. & Schmitz, L. Phylogenetic versus functional signals in the evolution of form–function relationships in terrestrial vision. Evolution 65, 2245–2257 (2011).PubMed 

Google Scholar  More

Barton, N. Evolutionary biology. The geometry of adaptation. Nature 395, 751–752. https://doi.org/10.1038/27338 (1998).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Otto, S. P. & Lenormand, T. Resolving the paradox of sex and recombination. Nat. Rev. Genet. 3, 252–261. https://doi.org/10.1038/nrg761 (2002).CAS 
Article 
PubMed 

Google Scholar 
Becks, L. & Agrawal, A. F. Higher rates of sex evolve in spatially heterogeneous environments. Nature 468, 89–92. https://doi.org/10.1038/nature09449 (2010).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Hughes, T. P. et al. Spatial and temporal patterns of mass bleaching of corals in the Anthropocene. Science 359, 80–83. https://doi.org/10.1126/science.aan8048 (2018).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Thompson, D. M. & van Woesik, R. Corals escape bleaching in regions that recently and historically experienced frequent thermal stress. Proc. Biol. Sci. 276, 2893–2901. https://doi.org/10.1098/rspb.2009.0591 (2009).CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Pandolfi, J. M., Connolly, S. R., Marshall, D. J. & Cohen, A. L. Projecting coral reef futures under global warming and ocean acidification. Science 333, 418–422. https://doi.org/10.1126/science.1204794 (2011).ADS 
CAS 
Article 
PubMed 

Google Scholar 
Sully, S., Burkepile, D. E., Donovan, M. K., Hodgson, G. & van Woesik, R. A global analysis of coral bleaching over the past two decades. Nat. Commun. 10, 1264. https://doi.org/10.1038/s41467-019-09238-2 (2019).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Yund, P. O. How severe is sperm limitation in natural populations of marine free-spawners?. Trends Ecol. Evol. 15, 10–13 (2000).CAS 
Article 

Google Scholar 
Levitan, D. R. & Petersen, C. Sperm limitation in the sea. Trend Ecol. Evol. 10, 228–231 (1995).CAS 
Article 

Google Scholar 
Baird, A., Guest, J. & Willis, B. Systematic and biogeographical patterns in the reproductive biology of scleractinian corals. Annu. Rev. Ecol. Evol. Syst. 40, 551–571. https://doi.org/10.1146/Annurev.Ecolsys.110308.120220 (2009).Article 

Google Scholar 
Wei, N. V. et al. Reproductive isolation among Acropora species (Scleractinia: Acroporidae) in a marginal coral assemblage. Zool. Stud. 51, 85–92 (2012).
Google Scholar 
Kitanobo, S., Isomura, N., Fukami, H., Iwao, K. & Morita, M. The reef-building coral Acropora conditionally hybridize under sperm limitation. Biol. Lett. 12, 20160511. https://doi.org/10.1098/rsbl.2016.0511 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Mercier, A. & Hamel, J.-F. Synchronized breeding events in sympatric marine invertebrates: Role of behavior and fine temporal windows in maintaining reproductive isolation. Behav. Ecol. Sociobiol. 64, 1749–1765 (2010).Article 

Google Scholar 
Levitan, D. R. et al. Mechanisms of reproductive isolation among sympatric broadcast-spawning corals of the Montastraea annularis species complex. Evolution 58, 308–323 (2004).Article 

Google Scholar 
Willis, B. L., Babcock, R. C., Harrison, P. L. & Wallace, C. C. Experimental hybridization and breeding incompatibilities within the mating systems of mass spawning reef corals. Coral Reefs 16, S53–S65 (1997).Article 

Google Scholar 
Nozawa, Y., Isomura, N. & Fukami, H. Influence of sperm dilution and gamete contact time on the fertilization rate of scleractinian corals. Coral Reefs 34, 1199–1206. https://doi.org/10.1007/s00338-015-1338-3 (2015).ADS 
Article 

Google Scholar 
Oliver, J. & Babcock, R. Aspects of the fertilization ecology of broadcast spawning corals: Sperm dilution effects and in situ measurements of fertilization. Biol. Bull. 183, 409–417. https://doi.org/10.2307/1542017 (1992).CAS 
Article 
PubMed 

Google Scholar 
Coma, R. & Lasker, H. R. Small-scale heterogeneity of fertilization success in a broadcast spawning octocoral. J. Exp. Mar. Biol. Ecol. 214, 107–120. https://doi.org/10.1016/S0022-0981(97)00017-8 (1997).Article 

Google Scholar 
Teo, A. & Todd, P. A. Simulating the effects of colony density and intercolonial distance on fertilisation success in broadcast spawning scleractinian corals. Coral Reefs 37, 891–900. https://doi.org/10.1007/s00338-018-1715-9 (2018).ADS 
Article 

Google Scholar 
Marshall, D. J. In situ measures of spawning synchrony and fertilization success in an intertidal, free-spawning invertebrate. Mar. Ecol. Prog. Ser. 236, 113–119 (2002).ADS 
Article 

Google Scholar 
Babcock, R. C., Mundy, C. N. & Whitehead, D. Sperm diffusion-models and in-situ confirmation of long-distance fertilization in the free-spawning asteroid Acanthaster planci. Biol. Bull. 186, 17–28 (1994).CAS 
Article 

Google Scholar 
Omori, M., Fukami, H., Kobinata, H. & Hatta, M. Significant drop of fertilization of Acropora corals in 1999. An after-effect of heavy coral bleaching?. Limnol. Oceanogr. 46, 704–706. https://doi.org/10.4319/lo.2001.46.3.0704 (2001).ADS 
Article 

Google Scholar 
Levitan, D. R., Fogarty, N. D., Jara, J., Lotterhos, K. E. & Knowlton, N. Genetic, spatial, and temporal components of precise spawning synchrony in reef building corals of the Montastraea annularis species complex. Evolution 65, 1254–1270. https://doi.org/10.1111/j.1558-5646.2011.01235.x (2011).Article 
PubMed 

Google Scholar 
Fukami, H., Omori, M., Shimoike, K., Hayashibara, T. & Hatta, M. Ecological and genetic aspects of reproductive isolation by different spawning times in Acropora corals. Mar. Biol. 142, 679–684. https://doi.org/10.1007/S00227-002-1001-8 (2003).Article 

Google Scholar 
Morita, M. et al. Reproductive strategies in the intercrossing corals Acropora donei and A. tenuis to prevent hybridization. Coral Reefs 38, 1211–1223. https://doi.org/10.1007/s00338-019-01839-z (2019).ADS 
Article 

Google Scholar 
Shinzato, C. et al. Development of novel, cross-species microsatellite markers for Acropora corals using next-generation sequencing technology. Front. Mar. Sci. 1, 11 (2014).Article 

Google Scholar 
R: A Language and Environment for Statistical Computing. (R Foundation for Statistical Computing, 2020).Albright, R. & Mason, B. Projected near-future levels of temperature and pCO2 reduce coral fertilization success. PLoS One 8, e56468. https://doi.org/10.1371/journal.pone.0056468 (2013).ADS 
CAS 
Article 
PubMed 
PubMed Central 

Google Scholar 
Iguchi, A., Morita, M., Nakajima, Y., Nishikawa, A. & Miller, D. In vitro fertilization efficiency in coral Acropora digitifera. Zygote 17, 225–227. https://doi.org/10.1017/S096719940900519X (2009).Article 
PubMed 

Google Scholar 
Morita, M. et al. Eggs regulate sperm flagellar motility initiation, chemotaxis and inhibition in the coral Acropora digitifera, A. gemmifera and A. tenuis. J. Exp. Biol. 209, 4574–4579. https://doi.org/10.1242/jeb.02500 (2006).CAS 
Article 
PubMed 

Google Scholar 
Chan, W. Y., Hoffmann, A. A. & van Oppen, M. J. H. Hybridization as a conservation management tool. Conserv. Lett. https://doi.org/10.1111/conl.12652 (2019).Article 

Google Scholar  More