Vertebrates (Latin for
joints of the spine) are a
subphylum of
chordates. They are chordates that have a
vertebral column (backbone). The vertebral column provides the central support structure for an
internal skeleton which gives shape, support, and protection to the body and can provide a means of anchoring fins or limbs to the body. The vertebral column also serves to house and protect the
spinal cord that lies within the vertebral column. Marine vertebrates can be divided into marine
fish and marine
tetrapods.
Marine fish Fish typically breathe by extracting oxygen from water through
gills and have a skin protected by
scales and
mucous. They use
fins to propel and stabilise themselves in the water, and usually have a
two-chambered heart and
eyes well adapted to seeing underwater, as well as other
sensory systems. Over 33,000 species of fish have been described as of 2017, of which about 20,000 are marine fish. }}
Jawless fish protruding from its back, may be an early jawless fish. Early fish had no
jaws. Most went extinct when they were outcompeted by jawed fish (below), but two groups survived:
hagfish and
lampreys. Hagfish form a class of about 20 species of
eel-shaped,
slime-producing marine fish. They are the only known living animals that have a
skull but no
vertebral column.
Lampreys form a superclass containing 38 known extant species of
jawless fish. The adult lamprey is characterized by a toothed, funnel-like sucking mouth. Although they are well known for boring into the flesh of other fish to
suck their blood, only 18 species of lampreys are actually parasitic. Together hagfish and lampreys are the sister group to vertebrates. Living hagfish remain similar to hagfish from around 300 million years ago. The lampreys are a very ancient lineage of vertebrates, though their exact relationship to
hagfishes and
jawed vertebrates is still a matter of dispute. Molecular analysis since 1992 has suggested that hagfish are most closely related to lampreys, and so also are vertebrates in a
monophyletic sense. Others consider them a sister group of vertebrates in the common taxon of craniata. The
Tully monster is an extinct genus of soft-bodied bilaterians that lived in tropical estuaries about 300 million years ago. Since 2016 there has been controversy over whether this animal was a vertebrate or an invertebrate. In 2020 researchers found "strong evidence" that the Tully monster was a vertebrate, and was a
jawless fish in the lineage of the
lamprey, while in 2023 other researchers found 3D fossils scans did not support those conclusions. File:Eptatretus polytrema.jpg|
Hagfish are the only known living animals with a
skull but no
vertebral column. File:Eudontomyzon mariae Dunai ingola.jpg|
Lampreys are often parasitic and have a toothed, funnel-like sucking mouth. File:Pteraspidomorphi.gif|The extinct
Pteraspidomorphi, ancestral to
jawed vertebrates
Pteraspidomorphi is an extinct
class of early jawless fish ancestral to jawed vertebrates. The few characteristics they share with the latter are now considered as primitive for all
vertebrates. Around the start of the
Devonian, fish started appearing with a deep remodelling of the vertebrate skull that resulted in a
jaw. All vertebrate jaws, including the human jaw, have evolved from these early fish jaws. The appearance of the early vertebrate jaw has been described as "perhaps the most profound and radical evolutionary step in vertebrate history". Jaws make it possible to capture, hold, and chew prey.
Fish without jaws had more difficulty surviving than fish with jaws, and most jawless fish became extinct during the Triassic period.
Cartilaginous fish Jawed fish fall into two main groups:
fish with bony internal skeletons and
fish with cartilaginous internal skeletons. Cartilaginous fish, such as
sharks and
rays, have jaws and skeletons made of
cartilage rather than
bone.
Megalodon is an extinct species of shark that lived about 28 to 1.5 Ma. It may looked much like a stocky version of the
great white shark, but was much larger with estimated lengths reaching . it was one of the largest and most powerful predators in vertebrate history, and probably had a profound impact on marine life. The
Greenland shark has the longest known lifespan of all vertebrates, about 400 years. Some sharks such as the great white are partially warm blooded and give live birth. The
manta ray, largest ray in the world, has been targeted by fisheries and is now
vulnerable. File:Acanthodes BW spaced.jpg|Cartilaginous fishes may have evolved from
spiny sharks. File:Myliobatis aquila sasrája.jpg|
Stingray File:MantaAlfrediLCouterier.jpg|
Manta ray, the largest ray File:Pristis clavata 2.jpg|
Sawfish, rays with long
rostrums resembling a saw. All species are now
endangered. File:Megalodon_size_chart.png|The extinct
megalodon resembled a giant
great white shark. File:Somniosus microcephalus1.jpg|The
Greenland shark lives longer than any other vertebrate. File:Rhincodon typus (recropped).jpg| The largest
extant fish, the
whale shark, is now a
vulnerable species.
Bony fish '', the earliest-known bony fish lived during the Late
Silurian 419 million years ago. Bony fish have jaws and skeletons made of
bone rather than
cartilage. Bony fish also have hard, bony plates called
operculum which help them respire and protect their gills, and they often possess a
swim bladder which they use for better control of their buoyancy. Bony fish can be further divided into those with
lobe fins and those with
ray fins. The approximate dates in the phylogenetic tree are from Near et al., 2012 and Zhu et al., 2009.
Guiyu oneiros, the earliest-known bony fish, lived during the Late
Silurian 419 million years ago. It has the combination of both
ray-finned and lobe-finned features, although analysis of the totality of its features place it closer to lobe-finned fish. Lobe fins evolved into the legs of the first tetrapod land vertebrates, so by extension an early ancestor of humans was a lobe-finned fish. Apart from the coelacanths and the lungfishes, lobe-finned fishes are now extinct. The remaining bony fish have ray fins. These are made of webs of skin supported by bony or horny spines (rays) which can be erected to control the fin stiffness. • The main distinguishing feature of the
chondrosteans (
sturgeon,
paddlefish,
bichir and
reedfish) is the cartilaginous nature of their skeletons. The ancestors of the chondrosteans are thought to be bony fish, but the characteristic of an ossified skeleton was lost in later evolutionary development, resulting in a lightening of the frame. •
Neopterygians (from Greek for
new fins) appeared sometime in the Late Permian, before dinosaurs. They were a very successful group of fish, because they could move more rapidly than their ancestors. Their scales and skeletons began to lighten during their evolution, and their jaws became more powerful and efficient.
Teleosts , where the upper half mirrors the lower half. About 96% of all modern fish species are teleosts, of which about 14,000 are marine species. Teleosts can be distinguished from other bony fish by their possession of a
homocercal tail, a tail where the upper half mirrors the lower half. In general, teleosts tend to be quicker and more flexible than more basal bony fishes. Their skeletal structure has evolved towards greater lightness. While teleost bones are well
calcified, they are constructed from a scaffolding of struts, rather than the dense
cancellous bones of
holostean fish. Teleosts are found in almost all
marine habitats. They have enormous
diversity, and range in size from adult
gobies 8mm long to
ocean sunfish weighing over 2,000 kg. The following images show something of the diversity in the shape and colour of modern marine teleosts... File:Istiophorus platypterus.jpg|
Sailfish File:Anguilla japonica 1856.jpg|
Eel File:Seahorse MKL Bd. 14 1890 (128643088).jpg|
Seahorse File:Sunfish.jpg|
Ocean sunfish File:Humpback anglerfish.png|
Anglerfish File:Tetraodon-hispidus.jpg|
Pufferfish File:Synchiropus splendidus 2 Luc Viatour cropped.png|
Mandarin dragonet Nearly half of all extant vertebrate species are teleosts.
Marine tetrapods '', an extinct lobe-finned fish, developed limb-like fins that could take it onto land. A
tetrapod (Greek for
four feet) is a vertebrate with
limbs (feet). Tetrapods evolved from ancient
lobe-finned fishes about 400 million years ago during the
Devonian Period when their earliest ancestors emerged from the sea and adapted to living on land. This change from a body plan for breathing and navigating in gravity-neutral water to a body plan with mechanisms enabling the animal to breathe in air without dehydrating and move on land is one of the most profound evolutionary changes known. Tetrapods can be divided into four classes:
amphibians,
reptiles,
birds and
mammals. }} Marine tetrapods are tetrapods that returned from land back to the sea again. The first returns to the ocean may have occurred as early as the
Carboniferous Period whereas other returns occurred as recently as the
Cenozoic, as in cetaceans,
pinnipeds, and several
modern amphibians.
Amphibians (from Greek for
both kinds of life) live part of their life in water and part on land. They mostly require fresh water to reproduce. A few inhabit brackish water, but there are no true marine amphibians. There have been reports, however, of amphibians invading marine waters, such as a Black Sea invasion by the natural hybrid
Pelophylax esculentus reported in 2010.
Reptiles Reptiles (Late Latin for
creeping or
crawling) do not have an aquatic larval stage, and in this way are unlike amphibians. Most reptiles are oviparous, although several species of squamates are
viviparous, as were some extinct aquatic clades — the fetus develops within the mother, contained in a
placenta rather than an
eggshell. As
amniotes, reptile eggs are surrounded by membranes for protection and transport, which adapt them to reproduction on dry land. Many of the viviparous species feed their
fetuses through various forms of placenta analogous to those of
mammals, with some providing initial care for their hatchlings. Some reptiles are more closely related to
birds than other reptiles, and many scientists prefer to make Reptilia a monophyletic group which includes the birds.
Extant non-avian reptiles which inhabit or frequent the sea include
sea turtles,
sea snakes,
terrapins, the
marine iguana, and the
saltwater crocodile. Currently, of the approximately 12,000 extant
reptile species and sub-species, only about 100 of are classed as marine reptiles. Except for some sea snakes, most extant marine reptiles are
oviparous and need to return to land to lay their eggs. Apart from sea turtles, the species usually spend most of their lives on or near land rather than in the ocean. Sea snakes generally prefer shallow waters nearby land, around islands, especially waters that are somewhat sheltered, as well as near estuaries. Unlike land snakes, sea snakes have evolved flattened tails which help them swim. File:Marine-Iguana-Espanola.jpg|
Marine iguana File:Leatherback sea turtle Tinglar, USVI (5839996547).jpg|
Leatherback sea turtle File:SaltwaterCrocodile('Maximo').jpg|
Saltwater crocodile File:Micrurus fulviusHolbrookV3P10AA.jpg|Marine snakes have flattened tails. File:Ichthyosaurus BW.jpg|The ancient
Ichthyosaurus communis independently evolved flippers similar to dolphins. Some
extinct marine reptiles, such as
ichthyosaurs, evolved to be
viviparous and had no requirement to return to land.
Ichthyosaurs resembled dolphins. They first appeared about 245 million years ago and disappeared about 90 million years ago. The terrestrial ancestor of the ichthyosaur had no features already on its back or tail that might have helped along the evolutionary process. Yet the ichthyosaur developed a
dorsal and
tail fin which improved its ability to swim. The biologist
Stephen Jay Gould said the ichthyosaur was his favourite example of
convergent evolution. The earliest marine reptiles arose in the
Permian. During the
Mesozoic many groups of reptiles became adapted to life in the seas, including
ichthyosaurs,
plesiosaurs,
mosasaurs,
nothosaurs,
placodonts,
sea turtles,
thalattosaurs and
thalattosuchians. Marine reptiles were less numerous after
mass extinction at the end of the
Cretaceous.
Birds Marine birds are
adapted to life within the
marine environment. They are often called
seabirds. While marine birds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking
convergent evolution, as the same environmental problems and feeding
niches have resulted in similar adaptations. Examples include
albatross,
penguins,
gannets, and
auks. In general, marine birds live longer,
breed later and have fewer young than terrestrial birds do, but they invest a great deal of time in their young. Most
species nest in
colonies, which can vary in size from a few dozen birds to millions. Many species are famous for undertaking long annual
migrations, crossing the
equator or circumnavigating the Earth in some cases. They feed both at the ocean's surface and below it, and even feed on each other. Marine birds can be highly
pelagic, coastal, or in some cases spend a part of the year away from the sea entirely. Some marine birds plummet from heights, plunging through the water leaving vapour-like trails, similar to that of fighter planes.
Gannets plunge into the water at up to 100 kilometres per hour (60 mph). They have air sacs under their skin in their face and chest which act like
bubble-wrap, cushioning the impact with the water. File:Goéland argenté - Julien Salmon.jpg|
European herring gull attack herring schools from above. File:Pygoscelis papua -Nagasaki Penguin Aquarium -swimming underwater-8a.jpg|
Gentoo penguin swimming underwater File:Royal Albatross - east of the Tasman Peninsula, Tasmania.jpg|Albatrosses range over huge areas of ocean and some even circle the globe. The first marine birds evolved in the
Cretaceous period, and modern marine bird families emerged in the
Paleogene.
Mammals , a classic
keystone species which controls
sea urchin numbers
Mammals (from Latin for
breast) are characterised by the presence of
mammary glands which in
females produce
milk for feeding (nursing) their young. There are about 130 living and recently extinct marine
mammal species such as
seals,
dolphins,
whales,
manatees,
sea otters and
polar bears. They do not represent a distinct taxon or systematic grouping, but are instead unified by their reliance on the marine environment for feeding. Both cetaceans and sirenians are fully aquatic and therefore are obligate water dwellers. Seals and sea-lions are semiaquatic; they spend the majority of their time in the water, but need to return to land for important activities such as
mating,
breeding and
molting. In contrast, both otters and the polar bear are much less adapted to aquatic living. Their diet varies considerably as well: some may eat
zooplankton; others may eat fish, squid, shellfish, and sea-grass; and a few may eat other mammals. In a process of
convergent evolution, marine mammals, especially cetaceans such as dolphins and whales, redeveloped their
body plan to parallel the streamlined
fusiform body plan of
pelagic fish. Front legs became
flippers and back legs disappeared, a
dorsal fin reappeared and the tail morphed into a powerful horizontal
fluke. This body plan is an adaptation to being an active predator in a high
drag environment. A parallel convergence occurred with the now extinct marine reptile
ichthyosaur. File:Bluewhale2 noaa.jpg|Endangered
blue whale, the largest living animal File:Tursiops truncatus 01.jpg|The
bottlenose dolphin has the highest
encephalization of any animal after humans File:Beluga03.jpg|
Beluga whale File:Noaa-walrus22.jpg|
Walrus File:Polar Bear - Alaska (cropped).jpg|
Polar bear ==Primary producers==