Invertebrate

Invertebrates are a paraphyletic group of animals that

neither possess nor develop a vertebral column (commonly Invertebrates
known as a backbone or spine), derived from the notochord. Temporal range: Cryogenian to Present,
This is a grouping including all animals apart from the
chordate subphylum Vertebrata. Familiar examples of
invertebrates include arthropods, mollusks, annelids,
echinoderms and cnidarians.

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,
Left to right: Chrysaora fuscescens
from 50 μm (0.002 in) rotifers[3] to the 9–10 m (30–33 ft)
(Cnidaria), Fromia indica
colossal squid.[4]
(Echinodermata), Caribbean reef squid
Some so-called invertebrates, such as the Tunicata and (Mollusca), Drosophila melanogaster
Cephalochordata, are more closely related to vertebrates than
(Arthropoda), Aplysina lacunosa
to other invertebrates. This makes the invertebrates
paraphyletic, so the term has little meaning in taxonomy. (Porifera), Pseudobiceros hancockanus
(Platyhelminthes), Hirudo medicinalis
(Annelida), Polycarpa aurata (Tunicata),
Contents Milnesium tardigradum (Tardigrada).

Etymology Scientific classification

Taxonomic significance (unranked): Filozoa
Number of extant species Kingdom: Animalia
Characteristics Groups included
Morphology and symmetry
Nervous system All animal groups not in
Respiratory system subphylum Vertebrata
Reproduction
Social interaction
Phyla
Classification of invertebrates
History
Classification
Significance of the group
In research
See also
References
Further reading
External links

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]
 Estimated number of

Invertebrate group Phylum Image

described species[11]

Insects Arthropoda 1,000,000

Arachnids Arthropoda 102,248

Snails Mollusca 85,000

Crustaceans Arthropoda 47,000

Clams Mollusca 20,000

Corals Cnidaria 2,175

Octopi/Squid Mollusca 900

Velvet worms Onychophora 165

 Estimated number of

Invertebrate group Phylum Image

described species[11]

Nautilus Mollusca 6

Horseshoe crabs Arthropoda 4

Others
jellyfish, echinoderms,
— — 68,658
sponges, other worms etc.

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

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 Tracheal system of dissected
diameters from only a few micrometres up to 0.8  mm. The cockroach. The largest tracheae run
smallest tubes, tracheoles, penetrate cells and serve as sites of across the width of the body of the
diffusion for water, oxygen, and carbon dioxide. Gas may be cockroach and are horizontal in this
conducted through the respiratory system by means of active image. Scale bar, 2 mm.
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. The tracheal system branches into
progressively smaller tubes, here
Insects recognize information transmitted by other
[28][29][30] supplying the crop of the cockroach.
insects. Scale bar, 2.0 mm.

Phyla

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 The fossil coral Cladocora from the
they are not organized into organs.[37] There are only two main Pliocene of Cyprus
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 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.[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]

Sponges (Porifera)
Comb jellies (Ctenophora)
Medusozoans and corals (Cnidaria)
Acoels (Xenacoelomorpha)
Flatworms (Platyhelminthes)
Bristleworms, earthworms and leeches (Annelida)
Insects, springtails, crustaceans, myriapods, chelicerates (Arthropoda)
Chitons, snails, slugs, bivalves, tusk shells, cephalopods (Mollusca)
Roundworms or threadworms (Nematoda)
Rotifers (Rotifera)
Tardigrades (Tardigrada)
Scalidophores (Scalidophora)
Lophophorates (Lophophorata)
Velvet worms (Onychophora)
Arrow worms (Chaetognatha)
Gordian worms or horsehair worms (Nematomorpha)
Ribbon worms (Nemertea)
Placozoa
Loricifera
Starfishes, sea urchins, sea cucumbers, sea lilies and brittle stars (Echinodermata)
Acorn worms, cephalodiscids and graptolites (Hemichordata)
Lancelets (Amphioxiformes)
Salps, pyrosomes, doliolids, larvaceans and sea squirts (Tunicata)

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] 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.[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
Biology portal

Invertebrate zoology
Invertebrate paleontology
Marine invertebrates
Pain in invertebrates

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Further reading
Hyman, L. H. 1940. The Invertebrates (6 volumes) New York : McGraw-Hill. A classic work.
Anderson, D. T. (Ed.). (2001). Invertebrate zoology (2nd ed.). Oxford: Oxford University Press.
Brusca, R. C., & Brusca, G. J. (2003). Invertebrates (2nd ed.). Sunderland, Mass. : Sinauer
Associates.
Miller, S.A., & Harley, J.P. (1996). Zoology (4th ed.). Boston: WCB/McGraw-Hill.
Pechenik, Jan A. (2005). Biology of the invertebrates. Boston: McGraw-Hill, Higher Education.
pp. 590 pp. ISBN 978-0-07-234899-6.
Ruppert, E. E., Fox, R. S., & Barnes, R. D. (2004). Invertebrate zoology: a functional evolutionary
approach. Belmont, CA: Thomas-Brooks/Cole.
Adiyodi, K.G. & Adyiodi, R.G. (Eds) 1983- . Reproductive Biology of Invertebrates. Wiley, New
York. (Many volumes.)
Giese, A.G. & Pearse, J.S. (Eds) 1974- . Reproduction of Marine Invertebrates. Academic Press,
New York. (Many volumes.)
Advances in Invertebrate Reproduction. Elsevier Science, Amsterdam. (Five volumes.)

External links
 A. R. Maggenti; S. Gardner (2005). Online Dictionary of Invertebrate Zoology (http://digitalcommon
s.unl.edu/onlinedictinvertzoology/).
Buglife (UK) (https://web.archive.org/web/20160305154711/https://www.buglife.org.uk/News/Enda
ngeredspeciessculpture.htm)
African Invertebrates (https://web.archive.org/web/20080820143549/http://www.africaninvertebrate
s.org.za/)

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