Invertebrate

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Invertebrates
Drosophila melanogaster - side (aka).jpg
The common fruit fly, Drosophila melanogaster , has been used extensively for research.
Scientific classification Red Pencil Icon.png
(unranked): Filozoa
Kingdom: Animalia
Groups included
Cladistically included but traditionally excluded taxa

Invertebrates are animals that neither possess nor develop a vertebral column (commonly known as a backbone or spine), derived from the notochord. This includes all animals apart from the subphylum Vertebrata. Familiar examples of invertebrates include arthropods (insects, arachnids, crustaceans, and myriapods), mollusks (chitons, snails, bivalves, squids, and octopuses), annelids (earthworms and leeches), and cnidarians (hydras, jellyfishes, sea anemones, and corals).

Contents

The majority of animal species are invertebrates; one estimate puts the figure at 97%. [1] Many invertebrate taxa have a greater number and variety of species than the entire subphylum of Vertebrata. [2] Invertebrates vary widely in size, from 50  μm (0.002 in) rotifers [3] to the 9–10 m (30–33 ft) colossal squid. [4]

Some so-called invertebrates, such as the Tunicata and Cephalochordata are more closely related to vertebrates than to other invertebrates. This makes the invertebrates paraphyletic, so the term has little meaning in taxonomy.

Etymology

The word "invertebrate" comes from the Latin word vertebra, which means a joint in general, and sometimes specifically a joint from the spinal column of a vertebrate. The jointed aspect of vertebra is derived from the concept of turning, expressed in the root verto or vorto, to turn. [5] The prefix in- means "not" or "without". [6]

Taxonomic significance

The term invertebrates is not always precise among non-biologists since it does not accurately describe a taxon in the same way that Arthropoda, Vertebrata or Manidae do. Each of these terms describes a valid taxon, phylum, subphylum or family. "Invertebrata" is a term of convenience, not a taxon; it has very little circumscriptional significance except within the Chordata. The Vertebrata as a subphylum comprises such a small proportion of the Metazoa that to speak of the kingdom Animalia in terms of "Vertebrata" and "Invertebrata" has limited practicality. In the more formal taxonomy of Animalia other attributes that logically should precede the presence or absence of the vertebral column in constructing a cladogram, for example, the presence of a notochord. That would at least circumscribe the Chordata. However, even the notochord would be a less fundamental criterion than aspects of embryological development and symmetry [7] or perhaps bauplan. [8]

Despite this, the concept of invertebrates as a taxon of animals has persisted for over a century among the laity, [9] and within the zoological community and in its literature it remains in use as a term of convenience for animals that are not members of the Vertebrata. [10] The following text reflects earlier scientific understanding of the term and of those animals which have constituted it. According to this understanding, invertebrates do not possess a skeleton of bone, either internal or external. They include hugely varied body plans. Many have fluid-filled, hydrostatic skeletons, like jellyfish or worms. Others have hard exoskeletons, outer shells like those of insects and crustaceans. The most familiar invertebrates include the Protozoa, Porifera, Coelenterata, Platyhelminthes, Nematoda, Annelida, Echinodermata, Mollusca and Arthropoda. Arthropoda include insects, crustaceans and arachnids.

Number of extant species

By far the largest number of described invertebrate species are insects. The following table lists the number of described extant species for major invertebrate groups as estimated in the IUCN Red List of Threatened Species , 2014.3. [11]

Invertebrate groupLatin nameImageEstimated number of
described species [11]
Insects Insecta European wasp white bg02.jpg 1,000,000
Arachnids Arachnida Hobo-spider.jpg 102,248
Molluscs Mollusca Grapevinesnail 01.jpg 85,000
Crustaceans Crustacea J J Wild Pseudocarcinus cropped.jpg 47,000
Corals Anthozoa FFS Table bottom.jpg 2,175
Velvet worms Onychophora Velvet worm.jpg 165
Horseshoe crabs Xiphosura Carcinoscorpius rotundicauda (mangrove horseshoe crab).jpg 4
Others
jellyfish, echinoderms,
sponges, other worms etc.
68,658
Total:~1,300,000

The IUCN estimates that 66,178 extant vertebrate species have been described, [11] which means that over 95% of the described animal species in the world are invertebrates.

Characteristics

The trait that is common to all invertebrates is the absence of a vertebral column (backbone): this creates a distinction between invertebrates and vertebrates. The distinction is one of convenience only; it is not based on any clear biologically homologous trait, any more than the common trait of having wings functionally unites insects, bats, and birds, or than not having wings unites tortoises, snails and sponges. Being animals, invertebrates are heterotrophs, and require sustenance in the form of the consumption of other organisms. With a few exceptions, such as the Porifera, invertebrates generally have bodies composed of differentiated tissues. There is also typically a digestive chamber with one or two openings to the exterior.

Morphology and symmetry

The body plans of most multicellular organisms exhibit some form of symmetry, whether radial, bilateral, or spherical. A minority, however, exhibit no symmetry. One example of asymmetric invertebrates includes all gastropod species. This is easily seen in snails and sea snails, which have helical shells. Slugs appear externally symmetrical, but their pneumostome (breathing hole) is located on the right side. Other gastropods develop external asymmetry, such as Glaucus atlanticus that develops asymmetrical cerata as they mature. The origin of gastropod asymmetry is a subject of scientific debate. [12]

Other examples of asymmetry are found in fiddler crabs and hermit crabs. They often have one claw much larger than the other. If a male fiddler loses its large claw, it will grow another on the opposite side after moulting. Sessile animals such as sponges are asymmetrical [13] alongside coral colonies (with the exception of the individual polyps that exhibit radial symmetry); alpheidae claws that lack pincers; and some copepods, polyopisthocotyleans, and monogeneans which parasitize by attachment or residency within the gill chamber of their fish hosts).

Nervous system

Neurons differ in invertebrates from mammalian cells. Invertebrates cells fire in response to similar stimuli as mammals, such as tissue trauma, high temperature, or changes in pH. The first invertebrate in which a neuron cell was identified was the medicinal leech, Hirudo medicinalis. [14] [15]

Learning and memory using nociceptors in the sea hare, Aplysia has been described. [16] [17] [18] Mollusk neurons are able to detect increasing pressures and tissue trauma. [19]

Neurons have been identified in a wide range of invertebrate species, including annelids, molluscs, nematodes and arthropods. [20] [21]

Respiratory system

Tracheal system of dissected cockroach. The largest tracheae run across the width of the body of the cockroach and are horizontal in this image. Scale bar, 2 mm. Tracheal system of dissected cockroach.tif
Tracheal system of dissected cockroach. The largest tracheae run across the width of the body of the cockroach and are horizontal in this image. Scale bar, 2 mm.
The tracheal system branches into progressively smaller tubes, here supplying the crop of the cockroach. Scale bar, 2.0 mm. Cockroach tracheae supplying crop.tiff
The tracheal system branches into progressively smaller tubes, here supplying the crop of the cockroach. Scale bar, 2.0 mm.

One type of invertebrate respiratory system is the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues. [22] The distribution of spiracles can vary greatly among the many orders of insects, but in general each segment of the body can have only one pair of spiracles, each of which connects to an atrium and has a relatively large tracheal tube behind it. The tracheae are invaginations of the cuticular exoskeleton that branch (anastomose) throughout the body with diameters from only a few micrometres up to 0.8 mm. The smallest tubes, tracheoles, penetrate cells and serve as sites of diffusion for water, oxygen, and carbon dioxide. Gas may be conducted through the respiratory system by means of active ventilation or passive diffusion. Unlike vertebrates, insects do not generally carry oxygen in their haemolymph. [23]

A tracheal tube may contain ridge-like circumferential rings of taenidia in various geometries such as loops or helices. In the head, thorax, or abdomen, tracheae may also be connected to air sacs. Many insects, such as grasshoppers and bees, which actively pump the air sacs in their abdomen, are able to control the flow of air through their body. In some aquatic insects, the tracheae exchange gas through the body wall directly, in the form of a gill, or function essentially as normal, via a plastron. Note that despite being internal, the tracheae of arthropods are shed during moulting (ecdysis). [24]

Reproduction

Like vertebrates, most invertebrates reproduce at least partly through sexual reproduction. They produce specialized reproductive cells that undergo meiosis to produce smaller, motile spermatozoa or larger, non-motile ova. [25] These fuse to form zygotes, which develop into new individuals. [26] Others are capable of asexual reproduction, or sometimes, both methods of reproduction.

Social interaction

Social behavior is widespread in invertebrates, including cockroaches, termites, aphids, thrips, ants, bees, Passalidae, Acari, spiders, and more. [27] Social interaction is particularly salient in eusocial species but applies to other invertebrates as well.

Insects recognize information transmitted by other insects. [28] [29] [30]

Phyla

The fossil coral Cladocora from the Pliocene of Cyprus Cladocora.jpg
The fossil coral Cladocora from the Pliocene of Cyprus

The term invertebrates covers several phyla. One of these are the sponges (Porifera). They were long thought to have diverged from other animals early. [31] They lack the complex organization found in most other phyla. [32] Their cells are differentiated, but in most cases not organized into distinct tissues. [33] Sponges typically feed by drawing in water through pores. [34] Some speculate that sponges are not so primitive, but may instead be secondarily simplified. [35] The Ctenophora and the Cnidaria, which includes sea anemones, corals, and jellyfish, are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus. [36] Both have distinct tissues, but they are not organized into organs. [37] There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, they are sometimes called diploblastic. [38]

The Echinodermata are radially symmetric and exclusively marine, including starfish (Asteroidea), sea urchins, (Echinoidea), brittle stars (Ophiuroidea), sea cucumbers (Holothuroidea) and feather stars (Crinoidea). [39]

The largest animal phylum is also included within invertebrates: the Arthropoda, including insects, spiders, crabs, and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. In addition, they possess a hardened exoskeleton that is periodically shed during growth. [40] Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits. The Nematoda or roundworms, are perhaps the second largest animal phylum, and are also invertebrates. Roundworms are typically microscopic, and occur in nearly every environment where there is water. [41] A number are important parasites. [42] Smaller phyla related to them are the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom. Other invertebrates include the Nemertea or ribbon worms, and the Sipuncula.

Another phylum is Platyhelminthes, the flatworms. [43] These were originally considered primitive, but it now appears they developed from more complex ancestors. [44] Flatworms are acoelomates, lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha. [45] The Rotifera or rotifers, are common in aqueous environments. Invertebrates also include the Acanthocephala or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and the Cycliophora. [46]

Also included are two of the most successful animal phyla, the Mollusca and Annelida. [47] [48] The former, which is the second-largest animal phylum by number of described species, includes animals such as snails, clams, and squids, and the latter comprises the segmented worms, such as earthworms and leeches. These two groups have long been considered close relatives because of the common presence of trochophore larvae, but the annelids were considered closer to the arthropods because they are both segmented. [49] Now, this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla. [50]

Among lesser phyla of invertebrates are the Hemichordata, or acorn worms, [51] and the Chaetognatha, or arrow worms. Other phyla include Acoelomorpha, Brachiopoda, Bryozoa, Entoprocta, Phoronida, and Xenoturbellida.

Classification of invertebrates

Invertebrates can be classified into several main categories, some of which are taxonomically obsolescent or debatable, but still used as terms of convenience. Each however appears in its own article at the following links. [52]

History

The earliest animal fossils appear to be those of invertebrates. 665-million-year-old fossils in the Trezona Formation at Trezona Bore, West Central Flinders, South Australia have been interpreted as being early sponges. [53] Some paleontologists suggest that animals appeared much earlier, possibly as early as 1 billion years ago. [54] Trace fossils such as tracks and burrows found in the Tonian era indicate the presence of triploblastic worms, like metazoans, roughly as large (about 5 mm wide) and complex as earthworms. [55]

Around 453 MYA, animals began diversifying, and many of the important groups of invertebrates diverged from one another. Fossils of invertebrates are found in various types of sediment from the Phanerozoic. [56] Fossils of invertebrates are commonly used in stratigraphy. [57]

Classification

Carl Linnaeus divided these animals into only two groups, the Insecta and the now-obsolete Vermes (worms). Jean-Baptiste Lamarck, who was appointed to the position of "Curator of Insecta and Vermes" at the Muséum National d'Histoire Naturelle in 1793, both coined the term "invertebrate" to describe such animals and divided the original two groups into ten, by splitting Arachnida and Crustacea from the Linnean Insecta, and Mollusca, Annelida, Cirripedia, Radiata, Coelenterata and Infusoria from the Linnean Vermes. They are now classified into over 30 phyla, from simple organisms such as sea sponges and flatworms to complex animals such as arthropods and molluscs.

Significance of the group

Invertebrates are animals without a vertebral column. This has led to the conclusion that invertebrates are a group that deviates from the normal, vertebrates. This has been said to be because researchers in the past, such as Lamarck, viewed vertebrates as a "standard": in Lamarck's theory of evolution, he believed that characteristics acquired through the evolutionary process involved not only survival, but also progression toward a "higher form", to which humans and vertebrates were closer than invertebrates were. Although goal-directed evolution has been abandoned, the distinction of invertebrates and vertebrates persists to this day, even though the grouping has been noted to be "hardly natural or even very sharp." Another reason cited for this continued distinction is that Lamarck created a precedent through his classifications which is now difficult to escape from. It is also possible that some humans believe that, they themselves being vertebrates, the group deserves more attention than invertebrates. [58] In any event, in the 1968 edition of Invertebrate Zoology, it is noted that "division of the Animal Kingdom into vertebrates and invertebrates is artificial and reflects human bias in favor of man's own relatives." The book also points out that the group lumps a vast number of species together, so that no one characteristic describes all invertebrates. In addition, some species included are only remotely related to one another, with some more related to vertebrates than other invertebrates (see Paraphyly). [59]

In research

For many centuries, invertebrates were neglected by biologists, in favor of big vertebrates and "useful" or charismatic species. [60] Invertebrate biology was not a major field of study until the work of Linnaeus and Lamarck in the 18th century. [60] During the 20th century, invertebrate zoology became one of the major fields of natural sciences, with prominent discoveries in the fields of medicine, genetics, palaeontology, and ecology. [60] The study of invertebrates has also benefited law enforcement, as arthropods, and especially insects, were discovered to be a source of information for forensic investigators. [40]

Two of the most commonly studied model organisms nowadays are invertebrates: the fruit fly Drosophila melanogaster and the nematode Caenorhabditis elegans . They have long been the most intensively studied model organisms, and were among the first life-forms to be genetically sequenced. This was facilitated by the severely reduced state of their genomes, but many genes, introns, and linkages have been lost. Analysis of the starlet sea anemone genome has emphasised the importance of sponges, placozoans, and choanoflagellates, also being sequenced, in explaining the arrival of 1500 ancestral genes unique to animals. [61] Invertebrates are also used by scientists in the field of aquatic biomonitoring to evaluate the effects of water pollution and climate change. [62]

See also

Related Research Articles

Nematomorpha A phylum of parasitoid animals, horsehair worms

Nematomorpha are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size from 50 to 100 millimetres long, reaching 2 metres in extreme cases, and 1 to 3 millimetres in diameter. Horsehair worms can be discovered in damp areas, such as watering troughs, swimming pools, streams, puddles, and cisterns. The adult worms are free-living, but the larvae are parasitic on arthropods, such as beetles, cockroaches, mantids, orthopterans, and crustaceans. About 351 freshwater species are known and a conservative estimate suggests that there may be about 2000 freshwater species worldwide. The name "Gordian" stems from the legendary Gordian knot. This relates to the fact that nematomorphs often tie themselves in knots.

Nemertea Phylum of invertebrates, ribbon worms

Nemertea is a phylum of invertebrate animals also known as ribbon worms or proboscis worms. Alternative names for the phylum have included Nemertini, Nemertinea and Rhynchocoela. Most are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many have patterns of yellow, orange, red and green coloration.

Coelom The main body cavity in most animals

The coelom is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it remains undifferentiated. In the past, and for practical purposes, coelom characteristics have been used to classify bilaterian animal phyla into informal groups.

Arachnid Class of arthropods

Arachnida is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Spiders are the largest order in the class, which also includes scorpions, ticks, mites, harvestmen, and solifuges. In 2019, a molecular phylogenetic study also placed horseshoe crabs in Arachnida.

Sclerite hardened body part

A sclerite is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly to the hardened parts of arthropod exoskeletons and the internal spicules of invertebrates such as certain sponges and soft corals. In paleontology, a scleritome is the complete set of sclerites of an organism, often all that is known from fossil invertebrates.

Ecdysozoa Superphylum of protostomes including arthropods, nematodes and others

Ecdysozoa is a group of protostome animals, including Arthropoda, Nematoda, and several smaller phyla. They were first defined by Aguinaldo et al. in 1997, based mainly on phylogenetic trees constructed using 18S ribosomal RNA genes. A large study in 2008 by Dunn et al. strongly supported the Ecdysozoa as a clade, that is, a group consisting of a common ancestor and all its descendants.

Nerve net simplest form of a nervous system: neurons are found spread apart, allowing the organism to respond to its environment

A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a central nervous system, organisms with radial symmetry are associated with nerve nets. Nerve nets can be found in members of the Cnidaria, Ctenophora, and Echinodermata phyla, all of which are found in marine environments. Nerve nets can provide animals with the ability to sense objects through the use of the sensory neurons within the nerve net.

Invertebrate zoology is the subdiscipline of zoology that consists of the study of invertebrates, animals without a backbone.

In biology, a tagma is a specialized grouping of multiple segments or metameres into a coherently functional morphological unit. Familiar examples are the head, the thorax, and the abdomen of insects. The segments within a tagma may be either fused or so jointed as to be independently moveable.

Body plan A set of morphological features common to members of a phylum of animals

A body plan, Bauplan, or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrate body plan is one of many: invertebrates consist of many phyla.

Terrestrial animal animals living on land

Terrestrial animals are animals that live predominantly or entirely on land, as compared with aquatic animals, which live predominantly or entirely in the water, or amphibians, which rely on a combination of aquatic and terrestrial habitats. Terrestrial invertebrates include ants, flies, crickets, grasshoppers and spiders.

The following outline is provided as an overview of and topical guide to zoology:

Animal Kingdom of motile multicellular eukaryotic heterotrophic organisms

Animals are multicellular eukaryotic organisms that form the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Over 1.5 million living animal species have been described—of which around 1 million are insects—but it has been estimated there are over 7 million animal species in total. Animals range in length from 8.5 millionths of a metre to 33.6 metres (110 ft). They have complex interactions with each other and their environments, forming intricate food webs. The kingdom Animalia includes humans, but in colloquial use the term animal often refers only to non-human animals. The scientific study of animals is known as zoology.

Taxonomy of commonly fossilised invertebrates Classification of ancient, commonly preserved, spine-lacking animals

Although the phylogenetic classification of non-vertebrate animals remains a work-in-progress, the following taxonomy attempts to be useful by combining both traditional (old) and new (21st-century) paleozoological terminology.

Marine invertebrates

Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.

Worm Animal that typically has a long tube-like body, no limbs, and no eyes

Worms are many different distantly related animals that typically have a long cylindrical tube-like body, no limbs, and no eyes. Worms vary in size from microscopic to over 1 metre (3.3 ft) in length for marine polychaete worms, 6.7 metres (22 ft) for the African giant earthworm, Microchaetus rappi, and 58 metres (190 ft) for the marine nemertean worm, Lineus longissimus. Various types of worm occupy a small variety of parasitic niches, living inside the bodies of other animals. Free-living worm species do not live on land, but instead, live in marine or freshwater environments, or underground by burrowing. In biology, "worm" refers to an obsolete taxon, vermes, used by Carolus Linnaeus and Jean-Baptiste Lamarck for all non-arthropod invertebrate animals, now seen to be paraphyletic. The name stems from the Old English word wyrm. Most animals called "worms" are invertebrates, but the term is also used for the amphibian caecilians and the slowworm Anguis, a legless burrowing lizard. Invertebrate animals commonly called "worms" include annelids, nematodes (roundworms), platyhelminthes (flatworms), marine nemertean worms, marine Chaetognatha, priapulid worms, and insect larvae such as grubs and maggots.

Arthropod Phylum of invertebrates with jointed exoskeletons

An arthropod is an invertebrate animal having an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Euarthropoda, which includes insects, arachnids, myriapods, and crustaceans. The term Arthropoda as originally proposed refers to a proposed grouping of Euarthropods and the phylum Onychophora.

Pain in invertebrates

Pain in invertebrates is a contentious issue. Although there are numerous definitions of pain, almost all involve two key components. First, nociception is required. This is the ability to detect noxious stimuli which evokes a reflex response that moves the entire animal, or the affected part of its body, away from the source of the stimulus. The concept of nociception does not imply any adverse, subjective feeling; it is a reflex action. The second component is the experience of "pain" itself, or suffering—i.e., the internal, emotional interpretation of the nociceptive experience. Pain is therefore a private, emotional experience. Pain cannot be directly measured in other animals, including other humans; responses to putatively painful stimuli can be measured, but not the experience itself. To address this problem when assessing the capacity of other species to experience pain, argument-by-analogy is used. This is based on the principle that if a non-human animal's responses to stimuli are similar to those of humans, it is likely to have had an analogous experience. It has been argued that if a pin is stuck in a chimpanzee's finger and they rapidly withdraw their hand, then argument-by-analogy implies that like humans, they felt pain. It has been questioned why the inference does not then follow that a cockroach experiences pain when it writhes after being stuck with a pin. This argument-by-analogy approach to the concept of pain in invertebrates has been followed by others.

The evolution of nervous systems dates back to the first development of nervous systems in animals. Neurons developed as specialized electrical signaling cells in multicellular animals, adapting the mechanism of action potentials present in motile single-celled and colonial eukaryotes. Simple nerve nets seen in animals like Cnidaria (jellyfish) evolved first, consisted of polymodal neurons which serve a dual purpose in motor and sensory functions. Cnidarians can be compared to Ctenophores, which although are both jellyfish, have very different nervous systems. Unlike Cnidarians, Ctenophores have neurons that use electrochemical signaling. This was perplexing because the phylum Ctenophora was considered to be more ancient than that of Porifera (sponges), which have no nervous system at all. This led to the rise of two theories which described how the early nervous system came about. One theory stated that the nervous system came about in an ancestor basal to all of these phylum, however was lost in Porifera. The other theory states that the nervous system arose independently twice, one basal to Cnidarians and one basal to Ctenophores. Bilateral animals – ventral nerve cords in invertebrates and dorsal nerve cords supported by a notochord in chordates-- evolved with a central nervous system that was around a central region, a process known as cephalization.

Annelid Phylum of segmented worms

The annelids, also known as the ringed worms or segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.

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Further reading