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Invertebrate

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Template:Short description Template:Pp-vandalism Template:Paraphyletic group Invertebrates are 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, molluscs, annelids, echinoderms, flatworms, cnidarians, and sponges.

The majority of animal species are invertebrates; one estimate puts the figure at 97%.<ref>Template:Cite journal</ref> Many invertebrate taxa have a greater number and diversity of species than the entire subphylum of Vertebrata.<ref name="isbn0-412-61390-5">Template:Cite book</ref> Invertebrates vary widely in size, from 10 μm (0.0004 in)<ref>Template:Cite book</ref> myxozoans to the 9–10 m (30–33 ft) colossal squid.<ref>Roper, C.F.E. & P. Jereb (2010). Family Cranchiidae. In: P. Jereb & C.F.E. Roper (eds.) Cephalopods of the world. An annotated and illustrated catalogue of species known to date. Volume 2. Myopsid and Oegopsid Squids Template:Webarchive. FAO Species Catalogue for Fishery Purposes No. 4, Vol. 2. FAO, Rome. pp. 148–178.</ref>

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 "invertebrates" paraphyletic, so the term has no significance in taxonomy.

EtymologyEdit

The word "invertebrate" comes from the Latin word Template:Wikt-lang, 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.<ref name="Latin Tucker">Template:Cite dictionary</ref> The prefix in- means "not" or "without".<ref>Template:Cite book</ref>

Taxonomic significanceEdit

The term invertebrates does not describe a taxon in the same way that Arthropoda, Vertebrata or Manidae do. Each of those 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<ref name=" Pech">Template:Cite book</ref> or perhaps Bauplan.<ref>Template:Cite book</ref>

Despite this, the concept of invertebrates as a taxon of animals has persisted for over a century among the laity,<ref name="isbn0-19-861271-0">Template:Cite book</ref> 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.<ref name="Agassiz2013">Template:Cite book</ref> 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 speciesEdit

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.<ref name=IUCN1014>The World Conservation Union. 2014. IUCN Red List of Threatened Species, 2014.3. Summary Statistics for Globally Threatened Species. Table 1: Numbers of threatened species by major groups of organisms (1996–2014) Template:Webarchive.</ref>

Invertebrate group Phylum Image Estimated number of
described species<ref name=IUCN1014 />
Insects Arthropoda File:European wasp white bg02.jpg 1,000,000
Arachnids Arthropoda File:PlatycryptusUndatusFemale.jpg 102,248
Gastropods Mollusca File:Grapevinesnail 01.jpg 85,000
Crustaceans Arthropods File:J J Wild Pseudocarcinus cropped.jpg 47,000
Bivalves Mollusca File:Argopecten irradians.jpg 20,000
Sea anemones,
corals, sea pens 		Cnidaria File:Coral Outcrop Flynn Reef.jpg 2,175
Cephalopods Mollusca File:Sepia officinalis1.jpg 900
Velvet worms Onychophora File:Velvet worm.jpg 165
Horseshoe crabs Arthropods File:Carcinoscorpius rotundicauda (mangrove horseshoe crab).jpg 4
Others
jellyfish, echinoderms,
sponges, etc. 		— — 68,658
Total: ~1,300,000

The IUCN estimates that 66,178 extant vertebrate species have been described,<ref name=IUCN1014 /> which means that over 95% of the described animal species in the world are invertebrates.

CharacteristicsEdit

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 symmetryEdit

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.<ref name="gastropod development">Template:Cite journal</ref>

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<ref name=Columbia>Symmetry, biological Template:Webarchive, cited at FactMonster.com from The Columbia Electronic Encyclopedia (2007).</ref> 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 systemEdit

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.<ref name="Nicholls and Baylor, 1968">Template:Cite journal</ref><ref name="Pastor et al., 1996">Template:Cite journal</ref> Learning and memory using nociceptors have been described in the sea hare, Aplysia.<ref name="Byrne et al., 1978">Template:Cite journal</ref><ref name="Castellucci et al., 1970">Template:Cite journal</ref><ref name="Fischer et al., 2011">Template:Cite journal</ref> Mollusk neurons are able to detect increasing pressures and tissue trauma.<ref name="Illich and Walters, 1997">Template:Cite journalMechanosensory neurons innervating Aplysia siphon encode noxious stimuli and display nociceptive sensitization. The Journal of Neuroscience, 17: 459–469</ref>

Neurons have been identified in a wide range of invertebrate species, including annelids, molluscs, nematodes and arthropods.<ref name="Eismann et al., 1984">Eisemann, C.H., Jorgensen, W.K., Merritt, D.J., Rice, M.J., Cribb, B.W., Webb, P.D. and Zalucki, M.P., (1984). "Do insects feel pain? — A biological view". Cellular and Molecular Life Sciences, 40: 1420–1423</ref><ref name="St John Smith and Lewis, 2009">Template:Cite journal</ref>

Respiratory systemEdit

File: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.
File: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.<ref>Wasserthal, Lutz T. (1998). Chapter 25: The Open Hemolymph System of Holometabola and Its Relation to the Tracheal Space. In "Microscopic Anatomy of Invertebrates". Wiley-Liss, Inc. Template:ISBN.</ref> 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.<ref>Template:Cite journalTemplate:Dead link</ref>

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).<ref>Template:Cite journal</ref>

HearingEdit

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ReproductionEdit

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.<ref>Template:Cite book</ref> These fuse to form zygotes, which develop into new individuals.<ref>Template:Cite book</ref> 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.<ref name="Picard2021">Template:Cite journal</ref> 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.<ref name = Picard2021/>

Reproductive systemsEdit

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Social interactionEdit

Social behavior is widespread in invertebrates, including cockroaches, termites, aphids, thrips, ants, bees, Passalidae, Acari, spiders, and more.<ref name=ChoeCrespi1997>Template:Cite book</ref> Social interaction is particularly salient in eusocial species but applies to other invertebrates as well.

Insects recognize information transmitted by other insects.<ref name="Riley">Template:Cite journal</ref><ref>Template:Cite journal</ref><ref name="Frisch1967">Frisch, Karl von. (1967) The Dance Language and Orientation of Bees. Cambridge, Massachusetts: The Belknap Press of Harvard University Press.</ref>

PhylaEdit

File: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.<ref>Template:Cite book</ref> They lack the complex organization found in most other phyla.<ref>Template:Cite book</ref> Their cells are differentiated, but in most cases not organized into distinct tissues.<ref>Template:Cite book</ref> Sponges typically feed by drawing in water through pores.<ref>Template:Cite book</ref> Some speculate that sponges are not so primitive, but may instead be secondarily simplified.<ref>Template:Cite journal</ref> 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.<ref>Template:Cite book</ref> Both have distinct tissues, but they are not organized into organs.<ref>Template:Cite book</ref> There are only two main germ layers, the ectoderm and endoderm, with only scattered cells between them. As such, they are sometimes called diploblastic.<ref>Template:Cite book</ref>

The Echinodermata are radially symmetric and exclusively marine, including starfish (Asteroidea), sea urchins, (Echinoidea), brittle stars (Ophiuroidea), sea cucumbers (Holothuroidea) and feather stars (Crinoidea).<ref>Template:Cite book</ref>

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.<ref name=forensic>Template:Cite book</ref> 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.<ref>Template:Cite book</ref> A number are important parasites.<ref>Template:Cite book</ref> 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.<ref name=Gilson>Template:Cite book</ref> These were originally considered primitive, but it now appears they developed from more complex ancestors.<ref>Template:Cite journalTemplate:Dead link</ref> Flatworms are acoelomates, lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha.<ref name="umodena">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The Rotifera, or rotifers, are common in aqueous environments. Invertebrates also include the Acanthocephala, or spiny-headed worms, the Gnathostomulida, Micrognathozoa, and the Cycliophora.<ref name="IntroCyclio">Template:Cite journal</ref>

Also included are two of the most successful animal phyla, the Mollusca and Annelida.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite video</ref> 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.<ref>Template:Cite journal</ref> Now, this is generally considered convergent evolution, owing to many morphological and genetic differences between the two phyla.<ref>Template:Cite journalTemplate:Dead linkTemplate:Cbignore</ref>

Among lesser phyla of invertebrates are the Hemichordata, or acorn worms,<ref>Template:Cite book</ref> and the Chaetognatha, or arrow worms. Other phyla include Acoelomorpha, Brachiopoda, Bryozoa, Entoprocta, Phoronida, and Xenoturbellida.

ClassificationEdit

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.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

HistoryEdit

The earliest animal fossils are 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.<ref name="roseMaloof">Template:Cite journal</ref> Some paleontologists suggest that animals appeared much earlier, possibly as early as 1 billion years ago<ref>Template:Cite book</ref> 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.<ref name="Seilacher1998">Template:Cite journal</ref>

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.<ref name=clarkson>Template:Cite book</ref> Fossils of invertebrates are commonly used in stratigraphy.<ref name=Kummel>Template:Cite book</ref>

ClassificationEdit

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.

SignificanceEdit

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.<ref name=rskbarnes>Template:Cite book</ref> 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).<ref>Template:Cite book</ref>

In researchEdit

For many centuries, invertebrates were neglected by biologists, in favor of big vertebrates and "useful" or charismatic species.<ref name="Aristotle"/> Invertebrate biology was not a major field of study until the work of Linnaeus and Lamarck in the 18th century.<ref name="Aristotle">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> 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.<ref name="Aristotle"/> 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.<ref name=forensic />

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 1,500 ancestral genes unique to animals.<ref>Template:Cite journal</ref> Invertebrates are also used by scientists in the field of aquatic biomonitoring to evaluate the effects of water pollution and climate change.<ref>Template:Cite journal</ref>

See alsoEdit

ReferencesEdit

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Further readingEdit

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External linksEdit

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