Short description: Animals that live both on land and in the water
Restoration of Ichthyostega, an early tetrapod from the Devonian
Marine otter of the west coast of South America
Hawaiian monk seal, off Kaʻula Island
This is a list of tetrapods that are semiaquatic; that is, while being at least partly terrestrial, they spend part of their life cycle or a significant fraction of their time in water as part of their normal behavior, and/or obtain a significant fraction of their food from an aquatic habitat. The very earliest tetrapods, such as Ichthyostega, were semiaquatic, having evolved from amphibious lobe-finned fish.
Some marine mammals, such as the marine otter, the polar bear and pinnipeds, are semiaquatic, while others, such as the sea otter, cetaceans and sirenians, are fully aquatic. The only fully aquatic nonmarine mammals are several manatees (the Amazonian manatee and some populations of African manatee) and certain small cetaceans (river dolphins, the tucuxi, and some populations of Irrawaddy dolphin and finless porpoise). No bird species is fully aquatic, as all must lay and incubate their amniotic eggs, as well as begin raising their young, on land or ice. Similarly among marine reptiles, sea turtles are almost fully aquatic, but must come ashore to lay eggs. Marine iguanas and partly marine crocodiles (such as the saltwater crocodile and the American crocodile) are all semiaquatic. Most sea snakes are ovoviviparous (live-bearing) and fully aquatic (the exceptions being the oviparous, semiaquatic sea kraits). A few freshwater snakes are also ovoviviparous and fully aquatic (e.g., Erpeton tentaculatum and Acrochordidae), but the majority are semiaquatic. Most amphibians have an aquatic larval stage and are at least semiaquatic for that reason, but there are many exceptions to this generalization.
The aquatic component of a semiaquatic species' lifestyle may be either obligatory or facultative to varying degrees (examples of the latter are the Arctic fox, jaguar and green iguana).
Note: dagger symbols, "†", have been used to indicate a listed taxon is extinct.
Contents
1Mammals
2Birds
3Non-avian dinosaurs
4Pterosaurs
5Other reptiles
6Amphibians
7See also
8Notes
9References
Mammals
All extant fully aquatic mammals except the sea otter are found in two clades of exclusively aquatic species, Cetacea and Sirenia; the extinct desmostylians are also thought to have been fully aquatic (these groups are thought to have become fully aquatic about 45, 40 and 30 Ma ago, respectively). In contrast, semiaquatic mammals are widely distributed throughout the class. However, extant semiaquatic swimming marine mammals are restricted to Carnivora (among which, pinnipeds apparently appeared about 20 Ma ago). Semiaquatic (carnivorous) rodents have been noted as having larger than normal brains for their size, possibly as a consequence of using their vibrissae for acoustic detection of prey.[1][2]
Platypus, a semiaquatic monotreme, Tasmania
Asian elephant using its trunk as a snorkel,[3] India
Monotremes
Platypus
Marsupials
Lutrine opossum
Water opossum or yapok
Tenrecs
Otter shrews
Web-footed tenrec
†Ancestral sirenians - fossils are known of transitional semiaquatic sirenians, such as the 50 Ma old Pezosiren, which was a hippo-like quadruped[4]
Proboscideans
†Moeritherium
Asian elephant - primarily terrestrial [note 1][3][6][7]
Sloths
†Thalassocnus - this marine genus of Pacific coastal South America arose about 7 Ma ago, and its most recent species may have been fully aquatic
Primates
Allen's swamp monkey
Crab-eating macaque - feeds opportunistically, but some populations dive for crabs in mangrove swamps
Proboscis monkey - inhabits mangrove, riparian and swamp forest and has webbed digits[8]
Humans - some cultures have practiced marine freediving for subsistence for over a thousand years; in the case of the Sama-Bajau of Maritime Southeast Asia, evidence suggests that genetic adaptations facilitate this lifestyle[9][10][11]
North American beaver, Virginia
Invasive coypu, Europe
Muskrat, Ontario
Rodents
Castorimorpha - beavers, pocket gophers, and kangaroo rats
Beavers
Caviomorpha - New World hystricognaths
Capybaras
Coypu
Paca
Myomorpha - mouse-like rodents
'Crab-eating rats'
European water vole
'Fish-eating rats'
Link rat or Congo forest mouse (Deomys ferrugineus), an African murid
'Marsh rats'
Muskrat
'Swamp rats'
'Water mice'
'Water rats' or 'wading rats'
Lagomorphs
Marsh rabbit
Swamp rabbit
Eulipotyphlans
Desmans
Star-nosed mole
'Water shrews'[12]
Bats
Noctilionidae - bulldog bats, a family with only two species, both semiaquatic, one eating aquatic insects, one mainly fish
Vespertilionidae - vesper bats
Mouse-eared bats - several species of this mostly insectivorous genus are piscivorous to varying degrees, such as Myotis vivesi (an exclusively marine species), M. pilosus, M. macropus, M. macrotarsus, M. stalkeri, and M. daubentonii[13]
Jaguar, in the Pantanal
Indian rhinoceros, India
Male waterbuck, Kenya
Female moose, Wyoming
Hippopotamus underwater
Carnivorans - while semiaquatic mustelids are widespread, semiaquatic felids are known primarily from South and Southeast Asia, while semiaquatic canids are best known from the Neotropics
Canidae - foxes, jackals and wolves
Arctic fox - coastal populations exploit marine food sources, such as fish, invertebrates and seaweed, and may hunt ringed seal pups or scavenge polar bear kills on pack ice[14][15][16][17]
British Columbia wolf
Bush dog[18]
Dhole - often drive their prey into water bodies[19]
Short-eared dog
Mustelidae - weasels and otters
Minks
Otters (except the sea otter)
Pinnipedia - seals, sea lions and the walrus
Procyonidae - raccoons
Crab-eating raccoon
Ursidae - bears
Kodiak bear and other salmonid-specializing brown bear populations
Polar bear
Felidae - cats
Fishing cat
Flat-headed cat
Jaguar - facultatively semiaquatic, i.e., have an affinity for water and exploit aquatic prey in a rainforest or floodplain environment, but may also inhabit more arid habitat[20][21]
Tiger
Herpestidae - mongooses
Crab-eating mongoose
Viverridae - civets and genets
Aquatic genet
Otter civet
Perissodactyls
Rhinocerotidae - rhinos
Indian rhinoceros - primarily terrestrial
Javan rhinoceros - primarily terrestrial
Sumatran rhinoceros - primarily terrestrial
Tapiridae - tapirs
Artiodactyls
Bovidae - ruminants
Anoa
Sitatunga
Nile lechwe
North Ronaldsay sheep - Feral populations in Orkney almost exclusively eat seaweed.
Waterbuck
Wild water buffalo
Cervidae - deer
Chinese water deer
Marsh deer
Moose - obtain a substantial fraction of their food from aquatic sources[22]
Pere David's deer - adept swimmers and stand for long periods in water up to their shoulders[23][24]
Hippopotamidae - hippos are the closest extant relatives of the fully aquatic cetaceans, which are considered a subgroup of artiodactyls
Tragulidae - mouse-deer[12][25]
Greater mouse-deer
Water chevrotain
Yellow-striped chevrotain
†Ancestral cetaceans - descended from terrestrial artiodactyls; fossils are known of transitional semiaquatic cetaceans, such as the 43 Ma old Peregocetus, which had both hooves and webbed feet[26]
White-tailed eagle, Norway
Emperor penguins, Antarctica
Brown pelican, Florida
James's flamingos, Bolivia
Male wood duck, Quebec
Birds
The great majority of semiaquatic birds are found within three clades whose members are mostly semiaquatic: Aequorlitornithes, Anseriformes and Gruiformes, thought to be about 64, 47 and 41 Ma old, respectively.[27][note 2]
Accipitriformes - diurnal birds of prey; semiaquatic members are the sea eagles, osprey, and to a lesser degree, marsh harriers and the palm nut vulture
Aequorlitornithes - a waterbird clade recognized in 2015,[27] which includes most marine birds
Ardeae
Aequornithes - wading and diving birds
Boobies and cormorants
Austrodyptornithes
Albatrosses
Penguins
Petrels
Loons
Pelecaniformes
Hammerkops
Herons, egrets and bitterns
Ibises and spoonbills
Pelicans
Shoebills
Storks
Eurypygimorphae
Sunbittern
Tropicbirds
Charadriiformes - shorebirds
Auks - including the flightless †Pinguinus
Gulls, skuas and allies
Waders - sandpipers, painted-snipes, jacanas, ibisbills, avocets and stilts, oystercatchers and plovers
Mirandornithes
Flamingos
Grebes
Anseriformes - waterfowl; mostly freshwater, but includes some marine species (e.g. the kelp goose and sea ducks, including the flightless †Chendytes)
Ducks, geese, and swans
Magpie goose
Screamers
Coraciiformes - many kingfishers are semiaquatic but not the others
Gruiformes - most but not all of the order is semiaquatic, mainly freshwater but with some (e.g. the clapper and mangrove rails) in brackish or marine habitats
Cranes
Finfoots
Limpkin
Rails, including crakes, coots and gallinules
Passeriformes - dippers are the only semiaquatic passerines. Some others like swallows, icterids, the seaside sparrow and the bearded reedling are often found close to water but don't normally enter the water.
Strigiformes - nocturnal birds of prey; fishing owls and fish owls specialize to varying degrees on fish and other aquatic prey
Non-avian dinosaurs
Hesperornis restoration
Spinosaurus restoration
Only a few non-avian dinosaurs are thought to have been semiaquatic. While the Mesozoic had many types of marine reptiles, the combination of being oviparous and endothermic seems to have prevented the evolution of fully aquatic dinosaurs, as in birds.
Theropoda
Paraves
Avialae
†Ichthyornis - a toothed ornithuran analog of modern seabirds such as gulls and petrels
†Hesperornithes - a mostly flightless group of diving bird-like ornithurans
†Dromaeosauridae
†Halszkaraptor - a mallard-sized basal dromaeosaurid from Mongolia with flipper-like forelimbs
†Megalosauria
†Spinosauridae[28] - thought to be partially piscivorous; the group had crocodile-like skulls, paddle-like tails[29] and includes some of the largest known carnivorous dinosaurs. The preference for a semiaquatic lifestyle probably varied by species.
†Ornithischia
†Ornithopoda
†Hadrosauriformes
†Lurdusaurus - an unusually heavy-bodied and short-limbed iguanodont conjectured to have been similar in lifestyle to the modern hippopotamus
†Ceratopsia
†Neoceratopsia
†Koreaceratops
Pterosaurs
Eudimorphodon restoration
Pterodaustro restoration, showing its bristle-like modified teeth, likely used for filter feeding as in flamingos
A number of types of pterosaurs are thought to have been piscivores, and a few are suspected of being molluscivores.
†Eudimorphodontidae
†Eudimorphodon - has been found with fish remains in its stomach
†Rhamphorhynchidae
†Rhamphorhynchus - commonly found with fish and cephalopod remains in its stomach
†Ctenochasmatoidea - most had webbed hindfeet and long torsos, adaptations for swimming and floating, and are inferred to have lived in coastal or lake environments
†Pteranodontia - many in this clade were piscivores
†Boreopteridae - apparently freshwater surface swimmers or divers
†Nyctosauridae - later members were similar to ornithocheirans
†Ornithocheirae - soaring marine forms that were aerial dip-feeders like frigatebirds
†Pteranodontidae - some may have been pelagic plunge-divers like gannets
Semiaquatic forms are widely distributed among extant and extinct reptiles, and extinct semiaquatic or fully aquatic marine forms were once ecologically prominent.
†Choristoderes resembled crocodilians in many respects but were not closely related and were generally more fully aquatic; they survived until the Miocene.
†Champsosaurus - a Late Cretaceous and early Paleogene sexually dimorphic choristodere in which the females appear better adapted for terrestrial life, probably to facilitate egg-laying
Crocodilians[note 3] - a great diversity of extinct relatives dating back to the Late Triassic include terrestrial, semiaquatic as well as fully aquatic marine forms
Lizards
†Aigialosauridae - semiaquatic Late Cretaceous lizards that evolved into fully aquatic marine mosasaurs, who appeared and rose to dominance after an anoxic event eliminated ichthyosaurs and pliosaurs
Basiliscus (basilisks) - corytophanids able to run across water
Chinese crocodile lizard - a monitor lizard relative
Chinese water dragon - an agamid analog of the green iguana
Crocodilurus (the crocodile tegu) and Dracaena (caiman lizards) - Neotropical teiid monitor lizard analogs
Galápagos marine iguana - the only extant marine lizard
Green iguana - facultatively semiaquatic, i.e., dives into bodies of water when available to escape predators,[30] but may also live in xeric habitats[31]
Earless monitor lizard - another monitor lizard relative
Monitor lizards
Asian water monitor
Mertens' water monitor
Mitchell's water monitor
Nile monitor
Sailfin lizards - agamids also able to run across water
Waterside skinks - often live along streams and use the water to escape predators
Snakes
Anacondas
Aquatic coral snake
Cottonmouth
Crayfish snakes
Farancia - mud and rainbow snakes
Garter snakes
Sea kraits
Water cobras
'Water snakes'
Turtles are mostly semiaquatic; fully terrestrial examples include box turtles, tortoises, and some Asian box turtles (there are no fully aquatic examples, as all lay their eggs on land)
Rhynchocephalians - represented today only by the terrestrial tuatara, but several Mesozoic lineages became adapted to aquatic lifestyles
†Palaeopleurosaurus - probably semiaquatic
North American eastern newt as a gilled aquatic larva, aposematic terrestrial juvenile ("red eft") and aquatic adult
Amphibians
Amphibians differ from other semiaquatic tetrapods in that their semiaquatic lifestyle is ancestral, rather than being the result of a secondary evolutionary trend from a terrestrial state back towards an aquatic environment. Thus, they are the only tetrapods to possess gills. All extant amphibians that are semiaquatic or fully aquatic inhabit freshwater habitats, with the exception of the crab-eating frog, which also exploits brackish habitats.
Most amphibians have an aquatic larval stage and thus are at least semiaquatic by virtue of this fact. Many adult amphibians are also semiaquatic (while others are fully aquatic or terrestrial). However, some amphibians lack an aquatic larval stage. Some frogs, such as most leiopelmatids, most ranixalids, some leptodactylids, some myobatrachids, Darwin's frog and the Seychelles frog, have nonaquatic tadpoles. Some caecilians, many frogs such as saddleback toads, most sooglossids and the greenhouse frog,[32] and most plethodontid salamanders lay eggs on land in which the larvae develop into adult form before they hatch. The alpine salamander[33] and African live-bearing toads (Nectophrynoides and Nimbaphrynoides)[34] are ovoviviparous and give birth on land. Additionally, about 75% of caecilians are viviparous.
†Temnospondyls - an early group of amphibians, reaching sizes up to those of crocodiles, whose adult stage was variously fully aquatic, semiaquatic or almost entirely terrestrial. Among the aquatic forms, the Triassic trematosaurids adapted to a marine lifestyle.
Most anurans (frogs and toads), but not the fully aquatic pipids, or fully aquatic members of other families such as Telmatobiidae
Some caecilians, such as ichthyophiids, rhinatrematids, Chthonerpeton and Nectocaecilia, but not including other fully aquatic typhlonectids[35]
Most non-plethodontid salamanders, but not including the fully aquatic amphiumids, cryptobranchids, proteids, sirenids and various neotenic species in other families, such as Ambystoma mexicanum
See also
Amphibious fish
Aquatic animal
Aquatic ecology
Aquatic mammal
Diving bird
Freshwater biology
Marine biology
Marine mammal
Marine reptile
Seabird
Secondarily aquatic tetrapods
Wading bird
Water bird
Notes
↑Elephants have a system of cracks in their skin which retains water and mud for purposes of thermoregulation and protection from insect parasites and UV radiation. This system is less developed in Asian elephants than in African bush elephants; the former generally live in more mesic habitats.[5]
↑These dates are without calibration based on the putative late Cretaceous fossil crown avian Vegavis; its inclusion would push back the date for Anseriformes to ~69 Ma.
↑Although all extant crocodilians are semiaquatic, some recently extinct mekosuchine genera, Mekosuchus and Quinkana, were mostly or entirely terrestrial.
References
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