thumb|upright=1.35|[[Killer whales (orcas) are highly visible marine apex predators that hunt many large species. However, most marine activity takes place among microscopic organisms that cannot be seen individually with the naked eye, such as marine bacteria and phytoplankton. The study of marine life spans into multiple fields, which is primarily marine biology, as well as biological oceanography.

By volume, oceans provide about 90% of the living space on Earth, Marine microorganisms have been variously estimated as constituting about 70% or about 90% of the total marine biomass. Marine primary producers, mainly cyanobacteria and chloroplastic algae, produce oxygen and sequester carbon via photosynthesis, which generate enormous biomass and significantly influence the atmospheric chemistry. Migratory species, such as oceanodromous and anadromous fish, also create biomass and biological energy transfer between different regions of Earth, with many serving as keystone species of various ecosystems. At a fundamental level, marine life affects the nature of the planet, and in part, shape and protect shorelines, and some marine organisms (e.g. corals) even help create new land via accumulated reef-building.

Marine life can be roughly grouped into autotrophs and heterotrophs according to their roles within the food web: the former include photosynthetic and the much rarer chemosynthetic organisms (chemoautotrophs) that can convert inorganic molecules into organic compounds using energy from sunlight or exothermic oxidation, such as cyanobacteria, iron-oxidizing bacteria, algae (seaweeds and various microalgae) and seagrass; the latter include all the rest that must feed on other organisms to acquire nutrients and energy, which include animals, fungi, protists and non-photosynthetic microorganisms. Marine animals are further informally divided into marine vertebrates and marine invertebrates, both of which are polyphyletic groupings with the former including all saltwater fish, marine mammals, marine reptiles and seabirds, and the latter include all that are not considered vertebrates. Generally, marine vertebrates are much more nektonic and metabolically demanding of oxygen and nutrients, often suffering distress or even mass deaths (a.k.a. "fish kills") during anoxic events, while marine invertebrates are a lot more hypoxia-tolerant and exhibit a wide range of morphological and physiological modifications to survive in poorly oxygenated waters.

Water

thumb|upright=1.3|Elevation histogram showing the percentage of the Earth's surface above and below sea level

There is no life without water. It has been described as the universal solvent for its ability to dissolve many substances, and as the solvent of life. Water is the only common substance to exist as a solid, liquid, and gas under conditions normal to life on Earth. The Nobel Prize winner Albert Szent-Györgyi referred to water as the mater und matrix: the mother and womb of life.

thumb|upright=1.3|Composition of seawater. Quantities in relation to 1 kg or 1 litre of sea water.

The abundance of surface water on Earth is a unique feature in the Solar System. Earth's hydrosphere consists chiefly of the oceans but technically includes all water surfaces in the world, including inland seas, lakes, rivers, and underground waters down to a depth of . The deepest underwater location is Challenger Deep of the Mariana Trench in the Pacific Ocean, having a depth of . The mass of this world ocean is 1.35 metric tons or about 1/4400 of Earth's total mass. The world ocean covers an area of with a mean depth of , resulting in an estimated volume of .

thumb|left|The Earth's [[water cycle]]

About 97.5% of the water on Earth is saline; the remaining 2.5% is fresh water. Most fresh water – about 69% – is present as ice in ice caps and glaciers. The average salinity of Earth's oceans is about of salt per kilogram of seawater (3.5% salt). Some salts are released from volcanic activity or extracted from cool igneous rocks. averaging nearly in depth. By volume, the ocean provides about 90 percent of the living space on the planet.

However, water is found elsewhere in the Solar System. Europa, one of the moons orbiting Jupiter, is slightly smaller than the Earth's Moon. There is a strong possibility a large saltwater ocean exists beneath its ice surface. It has been estimated the outer crust of solid ice is about thick and the liquid ocean underneath is about deep. This would make Europa's ocean over twice the volume of the Earth's ocean. There has been speculation Europa's ocean could support life, and could be capable of supporting multicellular microorganisms if hydrothermal vents are active on the ocean floor. Enceladus, a small icy moon of Saturn, also has what appears to be an underground ocean which actively vents warm water from the moon's surface.

Evolution

Historical development

The Earth is about 4.54 billion years old. The earliest undisputed evidence of life on Earth dates from at least 3.5 billion years ago, during the Eoarchean era after a geological crust started to solidify following the earlier molten Hadean Eon. Microbial mat fossils have been found in 3.48 billion-year-old sandstone in Western Australia. Other early physical evidence of a biogenic substance is graphite in 3.7 billion-year-old metasedimentary rocks discovered in Western Greenland as well as "remains of biotic life" found in 4.1 billion-year-old rocks in Western Australia. According to one of the researchers, "If life arose relatively quickly on Earth ... then it could be common in the universe." as early as 4.25 billion years ago according to one study, and 4.4 billion years ago according to another study.--->

All organisms on Earth are descended from a common ancestor or ancestral gene pool.

Highly energetic chemistry is thought to have produced a self-replicating molecule around 4 billion years ago, and half a billion years later the last common ancestor of all life existed. The current scientific consensus is that the complex biochemistry that makes up life came from simpler chemical reactions. The beginning of life may have included self-replicating molecules such as RNA and the assembly of simple cells. In 2016 scientists reported a set of 355 genes from the last universal common ancestor (LUCA) of all life, including microorganisms, living on Earth.

Current species are a stage in the process of evolution, with their diversity the product of a long series of speciation and extinction events. The common descent of organisms was first deduced from four simple facts about organisms: First, they have geographic distributions that cannot be explained by local adaptation. Second, the diversity of life is not a set of unique organisms, but organisms that share morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits and finally, that organisms can be classified using these similarities into a hierarchy of nested groups—similar to a family tree. However, modern research has suggested that, due to horizontal gene transfer, this "tree of life" may be more complicated than a simple branching tree since some genes have spread independently between distantly related species.

Past species have also left records of their evolutionary history. Fossils, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record. By comparing the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species. However, this approach is most successful for organisms that had hard body parts, such as shells, bones or teeth. Further, as prokaryotes such as bacteria and archaea share a limited set of common morphologies, their fossils do not provide information on their ancestry.

More recently, evidence for common descent has come from the study of biochemical similarities between organisms. For example, all living cells use the same basic set of nucleotides and amino acids. The development of molecular genetics has revealed the record of evolution left in organisms' genomes: dating when species diverged through the molecular clock produced by mutations. For example, these DNA sequence comparisons have revealed that humans and chimpanzees share 98% of their genomes and analyzing the few areas where they differ helps shed light on when the common ancestor of these species existed.

Prokaryotes inhabited the Earth from approximately 3–4 billion years ago. No obvious changes in morphology or cellular organization occurred in these organisms over the next few billion years. The eukaryotic cells emerged between 1.6 and 2.7 billion years ago. The next major change in cell structure came when bacteria were engulfed by eukaryotic cells, in a cooperative association called endosymbiosis. The engulfed bacteria and the host cell then underwent coevolution, with the bacteria evolving into either mitochondria or hydrogenosomes. Another engulfment of cyanobacterial-like organisms led to the formation of chloroplasts in algae and plants.

thumb|upright=1.3 |right|Phylogenetic and symbiogenetic tree of living organisms, showing a view of the origins of eukaryotes and prokaryotes

The history of life was that of the unicellular eukaryotes, prokaryotes and archaea until about 610 million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period. The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae, cyanobacteria, slime moulds and myxobacteria. In 2016 scientists reported that, about 800 million years ago, a minor genetic change in a single molecule called GK-PID may have allowed organisms to go from a single cell organism to one of many cells.

Soon after the emergence of these first multicellular organisms, a remarkable amount of biological diversity appeared over a span of about 10 million years, in an event called the Cambrian explosion. Here, the majority of types of modern animals appeared in the fossil record, as well as unique lineages that subsequently became extinct. Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen in the atmosphere from photosynthesis.

About 500 million years ago, plants and fungi started colonizing the land. Evidence for the appearance of the first land plants occurs in the Ordovician, around , in the form of fossil spores. Land plants began to diversify in the Late Silurian, from around . The colonization of the land by plants was soon followed by arthropods and other animals. Insects were particularly successful and even today make up the majority of animal species. Amphibians first appeared around 364 million years ago, followed by early amniotes and birds around 155 million years ago (both from "reptile"-like lineages), mammals around 129 million years ago, homininae around 10 million years ago and modern humans around 250,000 years ago. However, despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this process continue to be highly successful and dominate the Earth, with the majority of both biomass and species being prokaryotes.

Estimates on the number of Earth's current species range from 10 million to 14 million, of which about 1.2 million have been documented and over 86 percent have not yet been described.

Microorganisms

Microorganisms make up about 70% of the marine biomass. or multicellular. Microorganisms are diverse and include all bacteria and archaea, most protozoa such as algae, fungi, and certain microscopic animals such as rotifers.

Many macroscopic animals and plants have microscopic juvenile stages. Some microbiologists also classify viruses (and viroids) as microorganisms, but others consider these as nonliving.

Microorganisms are crucial to nutrient recycling in ecosystems as they act as decomposers. Some microorganisms are pathogenic, causing disease and even death in plants and animals. As inhabitants of the largest environment on Earth, microbial marine systems drive changes in every global system. Microbes are responsible for virtually all the photosynthesis that occurs in the ocean, as well as the cycling of carbon, nitrogen, phosphorus, other nutrients and trace elements.

thumb|upright 1.2|The range of sizes shown by [[prokaryotes (bacteria and archaea) and viruses relative to those of other organisms and biomolecules]]

Microscopic life undersea is diverse and still poorly understood, such as for the role of viruses in marine ecosystems. Most marine viruses are bacteriophages, which are harmless to plants and animals, but are essential to the regulation of saltwater and freshwater ecosystems. They infect and destroy bacteria in aquatic microbial communities, and are the most important mechanism of recycling carbon in the marine environment. The organic molecules released from the dead bacterial cells stimulate fresh bacterial and algal growth.

thumb|upright=1.3 |right| Sea spray containing marine microorganisms can be swept high into the atmosphere where they become [[aeroplankton, and can travel the globe before falling back to earth.]]A stream of airborne microorganisms circles the planet above weather systems but below commercial air lanes. Some peripatetic microorganisms are swept up from terrestrial dust storms, but most originate from marine microorganisms in sea spray. In 2018, scientists reported that hundreds of millions of viruses and tens of millions of bacteria are deposited daily on every square meter around the planet.

Microscopic organisms live throughout the biosphere. The mass of prokaryote microorganisms — which includes bacteria and archaea, but not the nucleated eukaryote microorganisms — may be as much as 0.8 trillion tons of carbon (of the total biosphere mass, estimated at between 1 and 4 trillion tons). Single-celled barophilic marine microbes have been found at a depth of in the Mariana Trench, the deepest spot in the Earth's oceans. Microorganisms live inside rocks below the sea floor under of ocean off the coast of the northwestern United States, as well as beneath the seabed off Japan. The greatest known temperature at which microbial life can exist is (Methanopyrus kandleri). In 2014, scientists confirmed the existence of microorganisms living below the ice of Antarctica. According to one researcher, "You can find microbes everywhere — they're extremely adaptable to conditions, and survive wherever they are." Viruses can infect all types of life forms, from animals and plants to microorganisms, including bacteria and archaea. The linear size of the average virus is about one one-hundredth that of the average bacterium. Most viruses cannot be seen with an optical microscope so electron microscopes are used instead.

Viruses are found wherever there is life and have probably existed since living cells first evolved. The origin of viruses is unclear because they do not form fossils, so molecular techniques have been used to compare the DNA or RNA of viruses and are a useful means of investigating how they arise.

Viruses are now recognized as ancient and as having origins that pre-date the divergence of life into the three domains. But the origins of viruses in the evolutionary history of life are unclear: some may have evolved from plasmids—pieces of DNA that can move between cells—while others may have evolved from bacteria. In evolution, viruses are an important means of horizontal gene transfer, which increases genetic diversity. They are considered by some to be a life form, because they carry genetic material, reproduce by creating multiple copies of themselves through self-assembly, and evolve through natural selection. However they lack key characteristics such as a cellular structure generally considered necessary to count as life. Because they possess some but not all such qualities, viruses have been described as replicators

Bacteriophages, often just called phages, are viruses that parasite bacteria and archaea. Marine phages parasite marine bacteria and archaea, such as cyanobacteria. They are a common and diverse group of viruses and are the most abundant biological entity in marine environments, because their hosts, bacteria, are typically the numerically dominant cellular life in the sea. Generally there are about 1 million to 10 million viruses in each mL of seawater, or about ten times more double-stranded DNA viruses than there are cellular organisms, although estimates of viral abundance in seawater can vary over a wide range. Tailed bacteriophages appear to dominate marine ecosystems in number and diversity of organisms. Inoviridae and Microviridae are also known to infect diverse marine bacteria.

Microorganisms make up about 70% of the marine biomass.

The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms. These viruses have been studied in most detail in the thermophilic archaea, particularly the orders Sulfolobales and Thermoproteales.

Viruses are an important natural means of transferring genes between different species, which increases genetic diversity and drives evolution. It is thought that viruses played a central role in the early evolution, before the diversification of bacteria, archaea and eukaryotes, at the time of the last universal common ancestor of life on Earth. Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth. and the deep portions of Earth's crust. Bacteria also live in symbiotic and parasitic relationships with plants and animals.

Once regarded as plants constituting the class Schizomycetes, bacteria are now classified as prokaryotes. Unlike cells of animals and other eukaryotes, bacterial cells do not contain a nucleus and rarely harbor membrane-bound organelles. Although the term bacteria traditionally included all prokaryotes, the scientific classification changed after the discovery in the 1990s that prokaryotes consist of two very different groups of organisms that evolved from an ancient common ancestor. These evolutionary domains are called Bacteria and Archaea.

The ancestors of modern bacteria were unicellular microorganisms that were the first forms of life to appear on Earth, about 4 billion years ago. For about 3 billion years, most organisms were microscopic, and bacteria and archaea were the dominant forms of life. Although bacterial fossils exist, such as stromatolites, their lack of distinctive morphology prevents them from being used to examine the history of bacterial evolution, or to date the time of origin of a particular bacterial species. However, gene sequences can be used to reconstruct the bacterial phylogeny, and these studies indicate that bacteria diverged first from the archaeal/eukaryotic lineage. <!-- The most recent common ancestor of bacteria and archaea was probably a hyperthermophile that lived about 2.5 billion–3.2 billion years ago. -->

Bacteria were also involved in the second great evolutionary divergence, that of the archaea and eukaryotes. Here, eukaryotes resulted from the entering of ancient bacteria into endosymbiotic associations with the ancestors of eukaryotic cells, which were themselves possibly related to the Archaea. This is known as secondary endosymbiosis.

<gallery mode="packed" heights="140" style="float:left;">

File:Sulphide bacteria crop2.jpg|The marine Thiomargarita namibiensis, the largest known bacterium

File:Potomac river eutro.jpg|Cyanobacteria blooms can contain lethal cyanotoxins.

File:Glaucocystis sp.jpg|The chloroplasts of glaucophytes have a peptidoglycan layer, evidence suggesting their endosymbiotic origin from cyanobacteria.

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The largest known bacterium, the marine Thiomargarita namibiensis, can be visible to the naked eye and sometimes attains .

Marine archaea

The archaea (Greek for ancient) constitute a domain and kingdom of single-celled microorganisms. These microbes are prokaryotes, meaning they have no cell nucleus or any other membrane-bound organelles in their cells.

Archaea were initially classified as bacteria, but this classification is outdated. Archaeal cells have unique properties separating them from the other two domains of life, Bacteria and Eukaryota. The Archaea are further divided into multiple recognized phyla. Classification is difficult because the majority have not been isolated in the laboratory and have only been detected by analysis of their nucleic acids in samples from their environment.

Archaea and bacteria are generally similar in size and shape, although a few archaea have very strange shapes, such as the flat and square-shaped cells of Haloquadratum walsbyi. Despite this morphological similarity to bacteria, archaea possess genes and several metabolic pathways that are more closely related to those of eukaryotes, notably the enzymes involved in transcription and translation. Other aspects of archaeal biochemistry are unique, such as their reliance on ether lipids in their cell membranes, such as archaeols. Archaea use more energy sources than eukaryotes: these range from organic compounds, such as sugars, to ammonia, metal ions or even hydrogen gas. Salt-tolerant archaea (the Haloarchaea) use sunlight as an energy source, and other species of archaea fix carbon; however, unlike plants and cyanobacteria, no known species of archaea does both. Archaea reproduce asexually by binary fission, fragmentation, or budding; unlike bacteria and eukaryotes, no known species forms spores.

Archaea are particularly numerous in the oceans, and the archaea in plankton may be one of the most abundant groups of organisms on the planet. Archaea are a major part of Earth's life and may play roles in both the carbon cycle and the nitrogen cycle.

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File:Halobacteria with scale.jpg|Halobacteria, found in water near saturated with salt, are now recognized as archaea.

File:Methanosarcina barkeri fusaro.gif|Methanosarcina barkeri, a marine archaea that produces methane

File:Thermophile bacteria2.jpg|Thermophiles, such as Pyrolobus fumarii, survive well over 100&nbsp;°C.

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Marine protists

<!-- thumb|100px| [[Dinoflagellate]]

thumb| Marine [[slime nets form labyrinthine networks of tubes in which amoeba without pseudopods can travel]] -->

Protists are eukaryotes that cannot be classified as plants, fungi or animals. They are usually single-celled and microscopic. Life originated as single-celled prokaryotes (bacteria and archaea) and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics, because they are paraphyletic (lacking a common ancestor). Protists can be broadly divided into four groups depending on whether their nutrition is plant-like, animal-like, fungus-like, or a mixture of these.

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! Description

! colspan=2 | Example

! Other examples

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! width=90px | Plant-like

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| Autotrophic protists that make their own food without needing to consume other organisms, usually by using photosynthesis

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| Red algae, Cyanidium sp.

| Green algae, brown algae, diatoms and some dinoflagellates. Plant-like protists are important components of phytoplankton discussed below.

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! Animal-like

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| Heterotrophic protists that get their food consuming other organisms

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| Radiolarian protist as drawn by Haeckel

| Foraminiferans, and some marine amoebae, ciliates and flagellates.

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! Fungus-like

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| Saprotrophic protists that get their food from the remains of organisms that have broken down and decayed

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| Marine slime nets form labyrinthine networks of tubes in which amoeba without pseudopods can travel

| Marine lichen

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! Mixotropes

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| Mixotrophic and osmotrophic protists that get their food from a combination of the above

| 100px

| Euglena mutabilis, a photosynthetic flagellate

| Many marine mixotrophs are found among protists, including among ciliates, Rhizaria and dinoflagellates

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16px Getting to know our single-celled ancestors - MicroCosmos

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Protists are highly diverse organisms currently organized into 18 phyla, but are not easy to classify. Studies have shown high protist diversity exists in oceans, deep sea-vents and river sediments, suggesting a large number of eukaryotic microbial communities have yet to be discovered. Recent studies in marine environments found mixotrophic protests contribute a significant part of the protist biomass.

<gallery mode="packed" heights="130" style="float:left;" caption="Single-celled and microscopic protists">

File:Diatoms through the microscope.jpg|Diatoms are a major algae group generating about 20% of world oxygen production.

File:Diatom algae Amphora sp.jpg|Diatoms have glass like cell walls made of silica and called frustules.

File:Podocyrtis papalis Ehrenberg - Radiolarian (30448963206).jpg|Radiolarian

File:Gephyrocapsa oceanica color (lightened).jpg|Single-celled alga, Gephyrocapsa oceanica

File:CSIRO ScienceImage 7609 SEM dinoflagellate.jpg|Two dinoflagellates

File:Zooxanthellae.jpg|Zooxanthellae is a photosynthetic algae that lives inside hosts like coral.

File:Paramecium bursaria.jpg|A single-celled ciliate with green zoochlorellae living inside endosymbiotically.

File:Euglenoid movement.jpg|Euglenoid

File:The ciliate Frontonia sp.jpg|This ciliate is digesting cyanobacteria. The cytostome or mouth is at the bottom right.

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In contrast to the cells of prokaryotes, the cells of eukaryotes are highly organized. Plants, animals and fungi are usually multi-celled and are typically macroscopic. Most protists are single-celled and microscopic. But there are exceptions. Some single-celled marine protists are macroscopic. Some marine slime molds have unique life cycles that involve switching between unicellular, colonial, and multicellular forms. Other marine protist are neither single-celled nor microscopic, such as seaweed.

<gallery mode="packed" heights="130" style="float:left;" caption="Macroscopic protists (see also unicellular macroalgae →)">

File:Chaos carolinensis Wilson 1900.jpg|The single-celled giant amoeba has up to 1000 nuclei and reaches lengths of .

File:Xenophyophore.jpg|The xenophyophore, another single-celled foraminiferan, lives in abyssal zones. It has a giant shell up to across.

File:Giant Kelp.jpg|Giant kelp, a brown algae, is not a true plant, yet it is multicellular and can grow to 50m.

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Protists have been described as a taxonomic grab bag where anything that doesn't fit into one of the main biological kingdoms can be placed. Some modern authors prefer to exclude multicellular organisms from the traditional definition of a protist, restricting protists to unicellular organisms. This more constrained definition excludes seaweeds and slime molds.

Marine microanimals

As juveniles, animals develop from microscopic stages, which can include spores, eggs and larvae. At least one microscopic animal group, the parasitic cnidarian Myxozoa, is unicellular in its adult form, and includes marine species. Other adult marine microanimals are multicellular. Microscopic adult arthropods are more commonly found inland in freshwater, but there are marine species as well. Microscopic adult marine crustaceans include some copepods, cladocera and tardigrades (water bears). Some marine nematodes and rotifers are also too small to be recognized with the naked eye, as are many loricifera, including the recently discovered anaerobic species that spend their lives in an anoxic environment. Copepods contribute more to the secondary productivity and carbon sink of the world oceans than any other group of organisms. While mites are not normally thought of as marine organisms, most species of the family Halacaridae live in the sea.

<gallery mode="packed" heights="150" style="float:left;" caption="Marine microanimals">

File:Copepod 2.jpg|Over 10,000 marine species are copepods, small, often microscopic crustaceans

File:Gastrotrich.jpg|Darkfield photo of a gastrotrich, a worm-like animal living between sediment particles

File:Echiniscus testudo Doyere 1840 Pl 12 Fig 1.png|Drawing of a tardigrade (water bear) on a grain of sand

File:Squatinella sp. (Rädertierchen - Rotifera) - 160x (13402418244).jpg|Rotifers, usually 0.1–0.5&nbsp;mm long, may look like protists but have many cells and belongs to the Animalia.

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Fungi

thumb|right|[[Lichen on a rock in a marine splash zone. Lichens are mutualistic associations between a fungus and an alga or cyanobacterium.]]

Over 1500 species of fungi are known from marine environments. These are parasitic on marine algae or animals, or are saprobes feeding on dead organic matter from algae, corals, protozoan cysts, sea grasses, wood and other substrata. Spores of many species have special appendages which facilitate attachment to the substratum. Marine fungi can also be found in sea foam and around hydrothermal areas of the ocean.

Mycoplankton are saprotropic members of the plankton communities of marine and freshwater ecosystems. They are composed of filamentous free-living fungi and yeasts associated with planktonic particles or phytoplankton. Similar to bacterioplankton, these aquatic fungi play a significant role in heterotrophic mineralization and nutrient cycling. Mycoplankton can be up to 20&nbsp;mm in diameter and over 50&nbsp;mm in length.

A typical milliliter of seawater contains about 10<sup>3</sup> to 10<sup>4</sup> fungal cells. This number is greater in coastal ecosystems and estuaries due to nutritional runoff from terrestrial communities. A higher diversity of mycoplankton is found around coasts and in surface waters down to 1000 meters, with a vertical profile that depends on how abundant phytoplankton is. This profile changes between seasons due to changes in nutrient availability. Marine fungi survive in a constant oxygen deficient environment, and therefore depend on oxygen diffusion by turbulence and oxygen generated by photosynthetic organisms.

Marine fungi can be classified as: Many more occur in the splash zone, where they occupy different vertical zones depending on how tolerant they are to submersion. Some lichens live a long time; one species has been dated at 8,600 years. However their lifespan is difficult to measure because what defines the same lichen is not precise. Lichens grow by vegetatively breaking off a piece, which may or may not be defined as the same lichen, and two lichens of different ages can merge, raising the issue of whether it is the same lichen.

According to fossil records, fungi date back to the late Proterozoic era 900–570 million years ago. Fossil marine lichens 600 million years old have been discovered in China. It has been hypothesized that mycoplankton evolved from terrestrial fungi, likely in the Paleozoic era (390 million years ago).

Origin of animals

thumb| [[Dickinsonia may be the earliest animal. They appear in the fossil record 571 million to 541 million years ago.]]

The earliest animals were marine invertebrates, that is, vertebrates came later. Animals are multicellular eukaryotes, Marine invertebrates are animals that inhabit a marine environment apart from the vertebrate members of the chordate phylum; invertebrates lack a vertebral column. Some have evolved a shell or a hard exoskeleton.

The earliest animal fossils may belong to the genus Dickinsonia, 571 million to 541 million years ago. Individual Dickinsonia typically resemble a bilaterally symmetrical ribbed oval. They kept growing until they were covered with sediment or otherwise killed, and spent most of their lives with their bodies firmly anchored to the sediment. Their taxonomic affinities are presently unknown, but their mode of growth is consistent with a bilaterian affinity.

Apart from Dickinsonia, the earliest widely accepted animal fossils are the rather modern-looking cnidarians (the group that includes coral, jellyfish, sea anemones and Hydra), possibly from around The Ediacara biota, which flourished for the last 40 million years before the start of the Cambrian, were the first animals more than a very few centimeters long. Like Dickinsonia, many were flat with a "quilted" appearance, and seemed so strange that there was a proposal to classify them as a separate kingdom, Vendobionta. Others, however, have been interpreted as early molluscs (Kimberella), echinoderms (Arkarua), and arthropods (Spriggina, Parvancorina). There is still debate about the classification of these specimens, mainly because the diagnostic features which allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the Ediacarans. However, there seems little doubt that Kimberella was at least a triploblastic bilaterian animal, in other words, an animal significantly more complex than the cnidarians.

Small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and Middle Cambrian periods. The earliest, Cloudina, shows signs of successful defense against predation and may indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to molluscs, while the owners of some "armor plates," Halkieria and Microdictyon, were eventually identified when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals.

Body plans and phyla

thumb|right|[[Kimberella, an early mollusc important for understanding the Cambrian explosion. Invertebrates are grouped into different phyla (body plans).]]

Invertebrates are grouped into different phyla. Informally phyla can be thought of as a way of grouping organisms according to their body plan. A body plan refers to a blueprint which describes the shape or morphology of an organism, such as its symmetry, segmentation and the disposition of its appendages. The idea of body plans originated with vertebrates, which were grouped into one phylum. But the vertebrate body plan is only one of many, and invertebrates consist of many phyla or body plans. The history of the discovery of body plans can be seen as a movement from a worldview centered on vertebrates, to seeing the vertebrates as one body plan among many. Among the pioneering zoologists, Linnaeus identified two body plans outside the vertebrates; Cuvier identified three; and Haeckel had four, as well as the Protista with eight more, for a total of twelve. For comparison, the number of phyla recognized by modern zoologists has risen to 35.]]Historically body plans were thought of as having evolved rapidly during the Cambrian explosion, but a more nuanced understanding of animal evolution suggests a gradual development of body plans throughout the early Palaeozoic and beyond.

In the 1970s there was already a debate about whether the emergence of the modern phyla was "explosive" or gradual but hidden by the shortage of Precambrian animal fossils. Later discoveries of similar animals and the development of new theoretical approaches led to the conclusion that many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups—for example that Opabinia was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it may have been closely related to the modern tardigrades. Nevertheless, there is still much debate about whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it appears unique in the history of animals.

Earliest animals

The deepest-branching animals — the earliest animals that appeared during evolution — are marine non-vertebrate organisms. The earliest animal phyla are the Porifera, Ctenophora, Placozoa and Cnidaria. No member of these clades exhibit body plans with bilateral symmetry.

Marine sponges

thumb|right|Sponges are perhaps the most basal animals. They have no nervous, digestive or circulatory system.

Sponges are animals of the phylum Porifera (from Modern Latin for bearing pores). They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells. They have non-specialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes.

Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues and organs, and have no body symmetry. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where it deposits nutrients, and leaves through a hole called the osculum. Many sponges have internal skeletons of spongin and/or spicules of calcium carbonate or silicon dioxide. All sponges are sessile aquatic animals. Although there are freshwater species, the great majority are marine (salt water) species, ranging from tidal zones to depths exceeding . Some sponges live to great ages; there is evidence of the deep-sea glass sponge Monorhaphis chuni living about 11,000 years.

While most of the approximately 5,000–10,000&nbsp;known species feed on bacteria and other food particles in the water, some host photosynthesizing micro-organisms as endosymbionts and these alliances often produce more food and oxygen than they consume. A few species of sponge that live in food-poor environments have become carnivores that prey mainly on small crustaceans.

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File:Sponges in Caribbean Sea, Cayman Islands.jpg|Sponge biodiversity. There are four sponge species in this photo.

File:Euplectella aspergillum (cropped).jpg|Venus' flower basket at a depth of 2572 meters

File:Barrel sponge (Xestospongia testudinaria).jpg|Barrel sponge

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Linnaeus mistakenly identified sponges as plants in the order Algae. For a long time thereafter sponges were assigned to a separate subkingdom, Parazoa (meaning beside the animals). They are now classified as a paraphyletic phylum from which the higher animals have evolved.

Ctenophores

Ctenophores (from Greek for carrying a comb), commonly known as comb jellies, are a phylum that live worldwide in marine waters. They are the largest non-colonial animals to swim with the help of cilia (hairs or combs). Coastal species need to be tough enough to withstand waves and swirling sediment, but some oceanic species are so fragile and transparent that it is very difficult to capture them intact for study. In the past ctenophores were thought to have only a modest presence in the ocean, but it is now known they are often significant and even dominant parts of the planktonic biomass. The position of the ctenophores in the evolutionary family tree of animals has long been debated, and the majority view at present, based on molecular phylogenetics, is that cnidarians and bilaterians are more closely related to each other than either is to ctenophores.

Placozoa

Placozoa (from Greek for flat animals) have the simplest structure of all animals. They are a basal form of free-living (non-parasitic) multicellular organism that do not yet have a common name. They live in marine environments and form a phylum containing so far only three described species, of which the first, the classical Trichoplax adhaerens, was discovered in 1883. Two more species have been discovered since 2017, and genetic methods indicate this phylum has a further 100 to 200 undescribed species.

thumb|upright=1.5|right| Crawling motility and food uptake by [[Trichoplax adhaerens|T.&nbsp;adhaerens]]

Trichoplax is a small, flattened, animal about one mm across and usually about 25&nbsp;μm thick. Like the amoebae they superficially resemble, they continually change their external shape. In addition, spherical phases occasionally form which may facilitate movement. Trichoplax lacks tissues and organs. There is no manifest body symmetry, so it is not possible to distinguish anterior from posterior or left from right. It is made up of a few thousand cells of six types in three distinct layers. The outer layer of simple epithelial cells bear cilia which the animal uses to help it creep along the seafloor. Trichoplax feed by engulfing and absorbing food particles – mainly microbes and organic detritus – with their underside.

Marine cnidarians

thumb|right|Cnidarians, like this [[starlet sea anemone, are the simplest animals to organise cells into tissue. Yet they have the same genes that form the vertebrate (including human) head.]]

Cnidarians (from Greek for nettle) are distinguished by the presence of stinging cells, specialized cells that they use mainly for capturing prey. Cnidarians include corals, sea anemones, jellyfish and hydrozoans. They form a phylum containing over 10,000 species of animals found exclusively in aquatic (mainly marine) environments. Their bodies consist of mesoglea, a non-living jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. They have two basic body forms: swimming medusae and sessile polyps, both of which are radially symmetrical with mouths surrounded by tentacles that bear cnidocytes. Both forms have a single orifice and body cavity that are used for digestion and respiration.

Fossil cnidarians have been found in rocks formed about . Fossils of cnidarians that do not build mineralized structures are rare. Scientists currently think cnidarians, ctenophores and bilaterians are more closely related to calcareous sponges than these are to other sponges, and that anthozoans are the evolutionary "aunts" or "sisters" of other cnidarians, and the most closely related to bilaterians.

Cnidarians are the simplest animals in which the cells are organized into tissues. The starlet sea anemone is used as a model organism in research. It is easy to care for in the laboratory and a protocol has been developed which can yield large numbers of embryos on a daily basis. There is a remarkable degree of similarity in the gene sequence conservation and complexity between the sea anemone and vertebrates.

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File:Sea anemone in tidepools.jpg|Sea anemones are common in tidepools.

File:Coral detail.jpg|Close up of polyps on the surface of a coral, waving their tentacles.

File:Maldives small island.jpg|If an island sinks below the sea, coral growth can keep up with rising water and form an atoll.

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File:Portuguese Man-O-War (Physalia physalis).jpg|The Portuguese man o' war is a colonial siphonophore

File:Porpita porpita.jpg|Porpita porpita consists of a colony of hydroids

File:Largelionsmanejellyfish.jpg|Lion's mane jellyfish, largest known jellyfish

File:Turritopsis dohrnii (cropped).jpg|Turritopsis dohrnii achieves biological immortality by transferring its cells back to childhood.

File:Chironex fleckeri (sea wasp).jpg|The sea wasp is the most lethal jellyfish in the world.

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Bilateral invertebrate animals

thumb|upright=1.7|right|Idealised wormlike bilaterian body plan. With a cylindrical body and a direction of movement the animal has head and tail ends. Sense organs and mouth form the basis of the head. Opposed circular and longitudinal muscles enable [[peristalsis|peristaltic motion.]]

Some of the earliest bilaterians were wormlike, and the original bilaterian may have been a bottom dwelling worm with a single body opening. Animals with this bilaterally symmetric body plan have a head (anterior) end<!--necessary! not implying there's an actual head--> and a tail (posterior) end as well as a back (dorsal) and a belly (ventral); therefore they also have a left side and a right side. The body stretches back from the head, and many bilaterians have a combination of circular muscles that constrict the body, making it longer, and an opposing set of longitudinal muscles, that shorten the body; They also have a gut that extends through the basically cylindrical body from mouth to anus. Many bilaterian phyla have primary larvae which swim with cilia and have an apical organ containing sensory cells. However, there are exceptions to each of these characteristics; for example, adult echinoderms are radially symmetric (unlike their larvae), and certain parasitic worms have extremely simplified body structures.

thumb|upright=0.6| [[Ikaria wariootia, an early bilaterian]]