Animalia

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Introduction

Animalia is the name of the animal kingdom in standard taxonomy, coming directly from the Latin animalis, which in turn comes from anima. Also the title of a book by Graeme Base full of illustration and aliteration: see Graeme Base/Animalia.


Development and evolution

Animals are eukaryotes, and diverged from the same group of flagellate protozoa that gave rise to the fungi and choanoflagellates. The last are especially close relatives, with collared cells appearing only among them, the sponges, and rarely in certain other animal forms. In all these groups motile cells have a single posterior flagellum with similar ultrastructure.

Adult animals are typically diploids, producing small motile sperm and large non-motile eggs. In all forms the fertilized zygote intiallvy divides to form a hollow sphere called a blastula. This then undergoes rearrangement and differentiation. Blastulae are probably representative of the sort of colonies animals evolved from; similar forms occur among other flagellates, e.g. Volvox. However none of these other groups really ever progressed further, and large multicellular forms tend to develop by progressive growth instead.

What makes animals different? The answer lies in the way the cells are held together. Instead of being simply stuck together or held in place by thick walls, animal cells are linked by septate junctions, composed mainly of elastic proteins - collagen is characteristic - that make up the extracellular matrix. Sometimes this is calcified to form shells, bones, or spicules, but otherwise it's fairly flexible and can serve as a framework, upon which cells can move about and be reorganized.

The first forms that might represent animals appear in the fossil record around the end of the Precambrian. These are called Vendian Biota and are exceedingly difficult to relate to later forms. Other than them, virtually every phylum makes a more or less simultaneous appearance during the Cambrian. This massive adaptive radiation may have come about because of climate change or a simple genetic innovation, and is so sudden that it is usually called the cambrian explosion.

Classification

In Linnaeus' original scheme, the animals were one of three kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves, and Mammalia. Since then the last four have all been subsumed into a single phylum, the Chordata, whereas the various other forms have been separated out. When they were first discovered, the Protozoa were included as an animal phylum or subkingdom, but as they are generally unrelated and often as similar to plants as animals, a new kingdom, the Protista, was devised to hold them.

Subkingdom Parazoa
        Phylum Porifera
Subkingdom "Agnotozoa" ~ "Mesozoa"
        Phylum Placozoa
        Phylum Orthonectida
        Phylum Rhombozoa         
Subkingdom Metazoa
  "Radiata"
        Phylum Cnidaria
        Phylum Ctenophora
  Bilateria
    Protostomia
        Phylum Platyhelminthes
        Phylum Nemertina
        Phylum Gnathostomulida
        Phylum Gastrotricha
        Phylum Rotifera
        Phylum Kinorhyncha
        Phylum Loricifera
        Phylum Acanthocephala
        Phylum Entoprocta
        Phylum Nematoda
        Phylum Nematomorpha
        Phylum Cycliophora
        Phylum Mollusca
        Phylum Sipuncula
        Phylum Echiura
        Phylum Annelida
        Phylum Pogonophora
        Phylum Vestimentifera
        Phylum Tardigrada
        Phylum Onychophora
        Phylum Arthropoda
        Phylum Phoronida
        Phylum Ectoprocta
        Phylum Brachiopoda
    Deuterostomia
        Phylum Echinodermata
        Phylum Chaetognatha
        Phylum Hemichordata
        Phylum Chordata

The sponges (Porifera) were separated from the other animals early on, and are very different. Sponges are sessile and usually feed by drawing in water through pores all over the body, which is supported by a skeleton typically divided into spicules - the cells are differentiated, but not organized into distinct groups.

There are three problematic phyla - the Rhombozoa, Orthonectida, and Placozoa - that have an unclear position with respect to other animals. After these, all animals belong to a monophyletic group called the Metazoa, characterized by a digestive chamber and separate cell layers that differentiate into various tissues, including nerves and muscles.

The simplest Metazoa are radially symmetric and diploblastic, that is, they have two germ layers. The outer layer (ectoderm) corresponds to the surface of the blastula and the inner layer (endoderm) is formed by cells that migrate into the interior. It then invaginates to form a digestive cavity with a single opening. This form is called a gastrula or planula when it is free-swimming. The Cnidaria (jellyfish, anenomes, corals, etc) are the main diploblastic phylum; the Ctenophora (comb jellies) may also belong here.

The remaining forms comprise a group called the Bilateria, since they are usually bilaterally symmetric, and are triploblastic. The blastula invaginates without filling in first, so the endoderm is simply its inner lining, and the interior then fills in to become a third layer (mesoderm) between the others. Like tissues are grouped into organs. The simplest of such animals are the Platyhelminthes (flatworms) which may be paraphyletic to the higher phyla.

The vast majority of the triploblastic phyla form a group called the Protostomia. These forms have a one-way digestive cavity, with a second opening developing to pass waste through. The mesoderm arises as in the flatworms, from a single cell, and then divides to form a mass on each side of the body. Usually there is a hollow space around the gut, called the coelom, arising from a split within the mesoderm, or at least some reduced version thereof (eg a pseudocoelom, where the split occurs between the mesoderm and endoderm, common in microscopic forms).

Some of the main protostome phyla are united by the presence of trocophore larva, which are distinguished by a special pattern of flagella, into a group called the Trochozoa. These include:

Traditionally the Arthropoda - the largest animal phylum including insects, spiders, crabs, and kin - and two small phyla related thereto, the Onychophora and Tardigrada, have been held to be close relatives of the annelids on account of their common segmented body plan. This relationship is now in doubt, and it appears that the arthropods may instead belong with various pseudocoelomate worms - the Nematoda (roundworms), Nematomorpha (horsehair worms), Kinorhyncha, Loricifera, and Priapulida - that share with them the characteristic of moulting. More work will be needed to resolve between these possibilities.

There are various pseudocoelomate protostomes that are hard to classify because of their small size and reduced structure. The Rotifera and Acanthocephala are closely related to each other and probably belong near the Trochozoa. Other groups include the Gastrotricha, Gnathostomulida, Entoprocta, and Cycliophora. The last was discovered only recently, and as little investigation has been done into the marine world more will probably turn up.

The Brachiopoda (lamp shells), Ectoprocta (=Bryozoa, moss animals), and Phoronida form a group called the Lophophorata, thanks to the shared presence of a fan of cilia around the mouth called the lophophore. The evolutionary relationships of these forms are very unclear, and the group may belong among the Deuterostomes and is possibly paraphyletic.

The Deuterostomes differ from the Protostomes in various ways. They also have a one-way digestive tract, but in this case the second opening is the mouth, so it goes the other way, and the mesoderm and coelom do not form in the same way, but rather through evagination of the endoderm called enterocoelic pouching. And, finally, the embryonic cleavage is different. All this suggests that the two lines are separate and monophyletic. The Deuterostomes include:

There are also some extinct animal phyla that, not having knowledge of their embryology or structure, are very difficult to place. These are mostly from the cambrian period, and include Archaeocyantha (possibly sponges), Conulariida (possibly cnidarians), and Conodonta (possibly chordates).