Invertebrate

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Invertebrates
Temporal range: Cryogenian to Present, 665–0 Ma
Invertebrate montage (2022).jpg
Left to right: Chrysaora fuscescens (Cnidaria), Fromia indica (Echinodermata), Caribbean reef squid (Mollusca), Drosophila melanogaster (Arthropoda), Aplysina lacunosa (Porifera), Pseudobiceros hancockanus (Platyhelminthes), Hirudo medicinalis (Annelida), Polycarpa aurata (Tunicata), Milnesium tardigradum (Tardigrada).
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Choanozoa
Kingdom: Animalia
Groups included

Invertebrates is an umbrella term describing animals that neither develop nor retain a vertebral column (commonly known as a spine or backbone), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum Vertebrata, i.e. vertebrates. Well-known phyla of invertebrates include arthropods, mollusks, annelids, echinoderms, flatworms, cnidarians, and sponges.

Contents

The majority of animal species are invertebrates; one estimate puts the figure at 97%. [1] Many invertebrate taxa have a greater number and diversity of species than the entire subphylum of Vertebrata. [2] Invertebrates vary widely in size, from 10  μm (0.0004 in) [3] Myxozoans to the 9–10 m (30–33 ft) colossal squid. [4]

Some so-called invertebrates, such as the Tunicata and Cephalochordata, are actually sister chordate subphyla to Vertebrata, being more closely related to vertebrates than to other invertebrates. This makes the term "invertebrates" rather polyphyletic, 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 groupPhylumImageEstimated number of
described species [11]
Insects Arthropoda European wasp white bg02.jpg 1,000,000
Arachnids Arthropoda PlatycryptusUndatusFemale.jpg 102,248
Gastropods Mollusca Grapevinesnail 01.jpg 85,000
Crustaceans Arthropoda J J Wild Pseudocarcinus cropped.jpg 47,000
Bivalves Mollusca Argopecten irradians.jpg 20,000
Anthozoa Cnidaria Coral Outcrop Flynn Reef.jpg 2,175
Cephalopods Mollusca Sepia officinalis1.jpg 900
Velvet worms Onychophora Velvet worm.jpg 165
Horseshoe crabs Arthropoda 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. Despite being internal, the tracheae of arthropods are shed during moulting (ecdysis). [24]

Hearing

Only vertebrate animals have ears, though many invertebrates detect sound using other kinds of sense organs. In insects, tympanal organs are used to hear distant sounds. They are located either on the head or elsewhere, depending on the insect family. [25] The tympanal organs of some insects are extremely sensitive, offering acute hearing beyond that of most other animals. The female cricket fly Ormia ochracea has tympanal organs on each side of her abdomen. They are connected by a thin bridge of exoskeleton and they function like a tiny pair of eardrums, but, because they are linked, they provide acute directional information. The fly uses her "ears" to detect the call of her host, a male cricket. Depending on where the song of the cricket is coming from, the fly's hearing organs will reverberate at slightly different frequencies. This difference may be as little as 50 billionths of a second, but it is enough to allow the fly to home in directly on a singing male cricket and parasitise it. [26]

Simpler structures allow other arthropods to detect near-field sounds. Spiders and cockroaches, for example, have hairs on their legs, which are used for detecting sound. Caterpillars may also have hairs on their body that perceive vibrations [27] and allow them to respond to sound.

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. [28] These fuse to form zygotes, which develop into new individuals. [29] Others are capable of asexual reproduction, or sometimes, both methods of reproduction.

Extensive research with model invertebrate species such as Drosophila melanogaster and Caenorhabditis elegans has contributed much to our understanding of meiosis and reproduction. However, beyond the few model systems, the modes of reproduction found in invertebrates show incredible diversity. [30] In one extreme example it is estimated that 10% of orbatid mite species have persisted without sexual reproduction and have reproduced asexually for more than 400 million years. [30]

Reproductive systems

Invertebrates have an extremely diverse array of reproductive systems, the only commonality may be that they all lay eggs. Also, aside from cephalopods and arthropods, nearly all other invertebrates are hermaphroditic and exhibit external fertilization.

Social interaction

Social behavior is widespread in invertebrates, including cockroaches, termites, aphids, thrips, ants, bees, Passalidae, Acari, spiders, and more. [31] Social interaction is particularly salient in eusocial species but applies to other invertebrates as well.

Insects recognize information transmitted by other insects. [32] [33] [34]

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. [35] They lack the complex organization found in most other phyla. [36] Their cells are differentiated, but in most cases not organized into distinct tissues. [37] Sponges typically feed by drawing in water through pores. [38] Some speculate that sponges are not so primitive, but may instead be secondarily simplified. [39] 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. [40] Both have distinct tissues, but they are not organized into organs. [41] There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, they are sometimes called diploblastic. [42]

The Echinodermata are radially symmetric and exclusively marine, including starfish (Asteroidea), sea urchins, (Echinoidea), brittle stars (Ophiuroidea), sea cucumbers (Holothuroidea) and feather stars (Crinoidea). [43]

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. [44] Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share some traits with them, excluding the hardened exoskeleton. 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. [45] A number are important parasites. [46] 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. [47] These were originally considered primitive, but it now appears they developed from more complex ancestors. [48] Flatworms are acoelomates, lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha. [49] The Rotifera, or rotifers, are common in aqueous environments. Invertebrates also include the Acanthocephala, or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and the Cycliophora. [50]

Also included are two of the most successful animal phyla, the Mollusca and Annelida. [51] [52] 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. [53] Now, this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla. [54]

Among lesser phyla of invertebrates are the Hemichordata, or acorn worms, [55] 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. [56]

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. [57] Some paleontologists suggest that animals appeared much earlier, possibly as early as 1 billion years ago [58] though they probably became multicellular in the Tonian. Trace fossils such as tracks and burrows found in the late Neoproterozoic era indicate the presence of triploblastic worms, roughly as large (about 5 mm wide) and complex as earthworms. [59]

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. [60] Fossils of invertebrates are commonly used in stratigraphy. [61]

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. [62] 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). [63]

In research

For many centuries, invertebrates were neglected by biologists, in favor of big vertebrates and "useful" or charismatic species. [64] Invertebrate biology was not a major field of study until the work of Linnaeus and Lamarck in the 18th century. [64] 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. [64] 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. [44]

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. [65] Invertebrates are also used by scientists in the field of aquatic biomonitoring to evaluate the effects of water pollution and climate change. [66]

See also

Related Research Articles

<span class="mw-page-title-main">Nematomorpha</span> 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, reaching 2 metres (79 in) 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, mantises, 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 coil themselves in tight balls that resemble knots.

<span class="mw-page-title-main">Nemertea</span> Phylum of invertebrates, ribbon worms

Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of about 1300 known species. Most ribbon worms 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. The foregut, stomach and intestine run a little below the midline of the body, the anus is at the tip of the tail, and the mouth is under the front. A little above the gut is the rhynchocoel, a cavity which mostly runs above the midline and ends a little short of the rear of the body. All species have a proboscis which lies in the rhynchocoel when inactive but everts to emerge just above the mouth to capture the animal's prey with venom. A highly extensible muscle in the back of the rhynchocoel pulls the proboscis in when an attack ends. A few species with stubby bodies filter feed and have suckers at the front and back ends, with which they attach to a host.

<span class="mw-page-title-main">Bilateria</span> Animals with embryonic bilateral symmetry

Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which extend to pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.

<span class="mw-page-title-main">Coelom</span> The main body cavity in many animals

The coelom is the main body cavity in many 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.

<span class="mw-page-title-main">Sclerite</span> 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.

<span class="mw-page-title-main">Ecdysozoa</span> Superphylum of protostomes including arthropods, nematodes and others

Ecdysozoa is a group of protostome animals, including Arthropoda, Nematoda, and several smaller phyla. The grouping of these animal phyla into a single clade was first proposed by Eernisse et al. (1992) based on a phylogenetic analysis of 141 morphological characters of ultrastructural and embryological phenotypes. This clade, that is, a group consisting of a common ancestor and all its descendants, was formally named by Aguinaldo et al. in 1997, based mainly on phylogenetic trees constructed using 18S ribosomal RNA genes.

<span class="mw-page-title-main">Nerve net</span> Nervous systems lacking a brain

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 condensation of neurons or, in more advanced forms, a central nervous system, organisms with radial symmetry are associated with nerve nets, and are found in members of the Ctenophora, Cnidaria, and Echinodermata phyla, all of which are found in marine environments. In the Xenacoelomorpha, a phylum of bilaterally symmetrical animals, members of the subphylum Xenoturbellida also possess a nerve net. Nerve nets can provide animals with the ability to sense objects through the use of the sensory neurons within the nerve net.

<span class="mw-page-title-main">Cephalization</span> Evolutionary trend of a head region developing

Cephalization is an evolutionary trend in animals that, over many generations, the special sense organs and nerve ganglia become concentrated towards the rostral end of the body where the mouth is located, often producing an enlarged head. This is associated with the animal's movement direction and bilateral symmetry, and cephalization of the nervous system led to the formation of a functional centralized brain in three groups of bilaterian animals, namely the arthropods, cephalopod molluscs, and vertebrates (craniates).

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

<span class="mw-page-title-main">Body plan</span> 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 vertebrates share one body plan, while invertebrates have many.

<span class="mw-page-title-main">Ventral nerve cord</span> Structure of the invertebrate central nervous system

The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, integrating sensory input and locomotor output. Because arthropods have an open circulatory system, decapitated insects can still walk, groom, and mate — illustrating that the circuitry of the ventral nerve cord is sufficient to perform complex motor programs without brain input.

<span class="mw-page-title-main">Terrestrial animal</span> 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, and semiaquatic animals, which rely on both aquatic and terrestrial habitats. Some groups of insects are terrestrial, such as ants, butterflies, earwigs, cockroaches, grasshoppers and many others, while other groups are partially aquatic, such as mosquitoes and dragonflies, which pass their larval stages in water.

<span class="mw-page-title-main">Animal</span> Kingdom of living things

Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Animals form a clade, meaning that they arose from a single common ancestor. Over 1.5 million living animal species have been described, of which around 1.05 million are insects, over 85,000 are molluscs, and around 65,000 are vertebrates. It has been estimated there are as many as 7.77 million animal species on Earth. Animal body lengths range from 8.5 μm (0.00033 in) to 33.6 m (110 ft). They have complex ecologies and interactions with each other and their environments, forming intricate food webs. The scientific study of animals is known as zoology, and the study of animal behaviour is known as ethology.

<span class="mw-page-title-main">Marine invertebrates</span> Marine animals without a vertebral column

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.

<span class="mw-page-title-main">Worm</span> Limbless invertebrate animal

Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and usually no eyes.

<span class="mw-page-title-main">Arthropod</span> Phylum of invertebrates with jointed exoskeletons

Arthropods are invertebrates in the phylum Arthropoda. They possess an exoskeleton with a cuticle made of chitin, often mineralised with calcium carbonate, a body with differentiated (metameric) segments, and paired jointed appendages. In order to keep growing, they must go through stages of moulting, a process by which they shed their exoskeleton to reveal a new one. They form an extremely diverse group of up to ten million species.

<span class="mw-page-title-main">Cestoda</span> Class of flatworms

Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of many similar units known as proglottids—essentially packages of eggs which are regularly shed into the environment to infect other organisms. Species of the other subclass, Cestodaria, are mainly fish infecting parasites.

<span class="mw-page-title-main">Pain in invertebrates</span> Contentious issue

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 necessarily 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. Primitive systems, like those found in protists, use chemical signalling for movement and sensitivity; data suggests these were precursors to modern neural cell types and their synapses. When some animals started living a mobile lifestyle and eating larger food particles externally, they developed ciliated epithelia, contractile muscles and coordinating & sensitive neurons for it in their outer layer.

<span class="mw-page-title-main">Annelid</span> Phylum of segmented worms

The annelids, also known as the segmented worms, comprise a large phylum called Annelida. The phylum contains 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.

References

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Further reading