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Metazoa: Information
Metazoans
Vielzellige Tiere



Protozoa - Mesozoa - Metazoa

Some time ago most zoologists would divide the animals into Protozoa, Mesozoa and Metazoa. Protozoans are primitive, unicellular animals, and metazoans are the more derived multicellular animals. Mesozoans were believed to be at the junction between these two large groups- living remnants linking unicellular and multicellular animals.

This subdivision of the animals, however, does not reflect natural groups and today it is no longer in use. Especially the group Protozoa has been recognized as a polyphyletic assemblage (i.e. containing species that are not directly related). The term "protozoan" is now used only as a general name to denote unicellular eukaryotic organisms that do not primarily feed by photosynthesis.

The earliest organisms on Earth were simple cells. Their offspring are the bacteria and archaebacteria. The origin of eukaryotic organisms from these bacteria and archaebacteria is still unclear and debated. Most biologists, however, agree that a single event gave rise to eukaryotic cells, and that Eukaryota, therefore, are a monophyletic group (i.e. all derive from a single common ancestor).

The Eukaryota are subdivided into three large monophyletic groups: (1) Tetramastigota. (2) Discicristata, Chromalveolata, Rhizaria, Planta. (3) Amoebozoa, Opisthokonta. This subdivision already shows that old systematic entities like "plants" and "animals" are para- or even polyphyletic groups.


Tetramastigota
These are primitive eukaryotes. They do not possess mitochondria, which has previously been regarded as a primitive character. However, genomic studies revealed that mitochondrial genes are present; now tetramastigotes are believed to have lost their mitochondria at an early stage of their evolution. The hydrogenosomes and mitosomes found in some Tetramastigota might be remnants of mitochondria.
The Tetramastigota comprise the following taxa:

(1) Retortamonadea. Tiny, mostly commensalic organisms, some species also are parasitic and pathogenic (e.g. Chilomastix mesnili as the cause of diarrhea in humans). No mitochondria.

(2) Diplomonadea. Two subtaxa. The primitive species of the subtaxon Enteromonadida are  commensals in the gut of vertebrates including humans (e.g. Enteromonas hominis). The derived subtaxon Diplomonadina comprises species with a so-called diplozoic organisation that is basically a not fully separated 2-cell stage. The Diplomonadina are mostly free living or commensalic. Some species, however, are pathogenic in animals and humans (e.g. Giardia intestinalis). No mitochondria, but the so-called mitosomes might represent reduced mitochondria.

(3) Oxymonadea. Commensalic organisms and symbionts, especially in the gut of termites (e.g. Oxymonas grandis). No mitochondria.

(4) Parabasalea. Similar to the Oxymonadea. No mitochondria, but similar organelles called hydrogenosomes are present. Most species are symbionts in the gut of insects. Some species are pathogenic in humans. Four subtaxa: Trichomonadida (e.g. Trichomonas vaginalis, pathogenic in humans), Cristamonadida, Trichonymphida, and Spirotrichonymphida (the latter three are symbionst in the gut of termites and other insects).


Discicristata, Chromalveolata, Rhizaria, Planta
This large monophyletic group comprises many species formerly classified as "algae", "protozoa" or "plants". The four large subgroups each comprise several subtaxa.

(1) Discicristata
Named for the disc-like shape of the cristae in their mitochondria. Five subtaxa: (a) The Heterolobosa are amoebae and slime-molds living in the soil or in limnic and marine sediments. Some species can be pathogenic and cause severe illnesses in humans (e.g. Naegleria fowleri). The Heterolobosa comprise two subtaxa: Schizopyrenida and Acrasea. The latter form cell aggregates that function as quasi multicellular fruiting bodies. (b) The Euglenozoa (=Euglenophyta) comprise heterotrophic forms and forms that have acquired the ability for photosynthesis by incorporating other photosynthetic organisms. Two subtaxa: Kinetoplastida (e.g. Trypanosoma brucei, the agent causing sleeping illness) and Euglenida (=Euglenata) (e.g. Euglena viridis, the popular study object in university courses). (c) The Diplonemida are a small group of flagellate organisms. Mainly marine, in sediments or planctonic. (d) The Hemimastigophora are very similar to the Euglenida and might be early offshoots of the Euglenozoan lineage. The few known species live in the soil. (e) The Pseudociliata superficially recall ciliates (and previously have been classified with them as "Protociliata"). Ultrastructural data suggest that the Pseudociliata are closelely related to the Euglenozoa. The few species are marine and feed on other microorganisms.

(2) Chromalveolata
A very large taxon that comprises species formerly classified as "algae", "protozoa" or "fungi". Two large subtaxa, Chromista and Alveolata:

(a) Chromista. Most species incorporate a photosynthetic symbiont. Three subtaxa: (I) Prymnesiomonada (=Haptomonada, =Haptophyta, =Prymnesiophyta), mainly marine (e.g. Chrysochromulina polylepis, the "killer algae" producing newspaper headlines regularly in the summer). (II) Cryptomonada (=Cryptophyta). Mainly in fresh water. (III) Heterokonta. Largest subtaxon. Comprising the following:

        Proteromonadea
        Bicosoecidea (=Bicocoecida)
        Opalinea
        Actinophryida
        Chrysophyceae (=Chrysomonadea)
        including the following taxa that are sometimes regarded as independent:
               Synurophyceae (=Mallomonadales)
               Pedinellophyceae (=Pedinellidida)
               Dictyochophyceae (=Silicoflagellida)
               Parmophyceae
               Sarcinochrysidophyceae
        Bacillariophyceae (=Diatomophyceae, =Diatomeae)
        Xanthophyceae (=Heteromonadea)
        Eustigmatophyceae
        Raphidophyceae (=Raphidomonadea, =Chloromonadophyceae)
        Phaeophyceae (=Fucophyceae)
        Labyrinthulomycetes (=Labyrinthulea)
        Hyphochytriomycetes (=Hyphochitridiomycetes)
        Oomycetes (=Oomycota)

The first 4 taxa were formerly regarded as "protozoans", whereas the taxa 5 - 10 were regarded as "algae" and taxa 11 - 13 as "fungi".
Proteromonadea are endosymbionts in vertebrates. Bicosoecida are reminiscent of choanoflagellates and can occur in colonies; mainly marine. Opalinea are commensalic in the gut of vertebrates. Actinophryida were formerly grouped with the Heliozoa, but are not related to them. Most species are limnic or marine, some species are terrestrial.
Chrysophyceae (golden brown algae) live mostly in fresh water. Bacillariophyceae (diatomeans) feed mainly phototrophic; marine or limnic. Xanthophyceae (yellow algae) live in the soil, fresh- and saltwater; many species form cell threads. Eustigmatophyceae and Raphidophyceae are unicellular organisms that live in freshwater or saltwater and comprise less than 20 species each. Phaeophyceae (brown algae) are multicellular; most species are attached to the ground in marine habitats; only few species live in freshwater.
Labyrinthulomycetes (web slime molds) and Hyphochytriomycetes are closely related to the Oomycetes. Most species of these three groups are multicellular organisms and many are parasites.

(b) Alveolata. This taxon comprises 7 subtaxa (3 of which are uncertain) that were previously classified either as "algae" or as "protozoans". (I) Dinoflagellata (=Dinophyta). Most species are armored with cellulose plates; most species are phototrophic; limnic or marine. (II) Perkinsozoa. Parasites of molluscs. (III) Colpodellidae. Classification of this family is uncertain; Only 5 known species; parasites of other "protozoa". (IV) Apicomplexa. Formerly a component of the "protozoan" taxon "Sporozoa". Many are extracellular parasites and commensals in the gut of a large variety of metazoans, mainly arthropods. Others are intra- and extracellular parasites in different organs; many human pathogens e.g.: Cryptosporidium parvum (agent of cryptosporidiosis), Toxoplasma gondii (agent of toxoplasmosis) and Plasmodium vivax (agent of malaria). (V) Ciliophora. Limnic and marine "protozoans" (ciliates) with many well-known forms, e.g. Paramecium caudatum, popular study object in university zoology courses. Few species are parasites or commensals. (VI) Haplosporea. Classification with the Alveolata uncertain. Formerly classified in the "protozoan" taxon "Ascetospora". Parasites of various metazoans, mainly molluscs. (VII) Paramyxea. Parasites of polychaetes, crustaceans and molluscs. The classification with the Alveolata is uncertain and is probably incorrect. Currently the true relationships of the Paramyxea are unknown.

(3) Rhizaria
This taxon comprises mainly amoebae that have been formerly classified with the "Rhizopoda", "Heliozoa" and "Radiolaria". These old taxa were found to be polyphyletic. Many subgroups have been transferred to other groups (e.g. to the Heterokonta), but some remaining amoeba-like forms have been found to constitute a monophyletic group now called Rhizaria. 8 Subtaxa. (I) Cercozoa. Some parasitic species, others are limnic, marine or terrestrial; some species have previously been classified as "slime molds" (Phytomyxa (=Plasmodiophorea)); some phototrophic species have previously been considered as "algae" (e.g. Paulinella chromatophora, Chlorarachnion spec.). (II) Foraminifera (=Granuloreticulosa) (foraminiferans). Most species have a chambered shell made from proteins, hardened organic matrix (sometimes with iron or silicium), or calcium carbonate. This shell preserves well and thus these organisms have a large fossil record. (III - VIII) Phaeodarea, Acantharea, Polycystinea, Centrohelida, Desmothoracida and Taxopodida. These 6 taxa comprise species with spicules made from silica and other minerals, that give them a "star-" or "sun-like" appearance.

(4) Planta (=Plantae)
This taxon comprises the "green plants" and many "algae". Two large subtaxa:
(a) Biliphyta. Two subtaxa: (I) Rhodophyta (red algae). Mostly multicellular. Some species are intracellular symbionts of Chromista or parasites in other Rhodophyta; (II) Glaucocystophyta (=Glaucophyta) (blue green algae); unicellular, limnic.
(b) Viridiplanta (=Chlorobionta, =Viridiplantae). Two subtaxa: (I) Chlorophyta (green algae and green plants); large group with unicellular and multicellular species; limnic, marine, or terrestrial. A well-known example is the colonial Volvox spec. The higher plants (Embryophyta) also belong to this taxon. (II) Streptophyta. These "algae" comprise species that formerly have been classified with the Chlorophyta, but are now known to represent an early offshoot of the Viridiplanta lineage.

Amoebozoa, Opisthokonta
This is the largest eukaryotic taxon that comprises species that have been classified as "protozoa", "fungi" and "animals".

(1) Amoebozoa. This taxon comprises a large portion of the "Rhizopoda". Two subtaxa: (a) Lobosa. Naked and shelled amoebae, including Amoeba proteus, the popular study object of university zoology courses. (b) Conosa. This taxon contains the Archamoebae that have previously been regarded as primitive Eukaryota, because of the lack of mitochondria; however, these have been lost secondarily. The second subtaxon of the Conosa are the Mycetozoa comprising the majority of the slime molds (Eumyxa and Dictyostela) and Aconchulina (formerly members of the "Rhizopoda").

(2) Opisthokonta. Four subtaxa:
(a) Fungi. Comprises all true fungi (Chytridiomycota, Zygomycota, Ascomycota, and Basidiomycota) and the unicellular parasitic Microspora (=Microsporidia), that have previously been regarded as the most primitive Eukaryota. Many true fungi live in symbiosis with bacteria or "algae" and together they form a new grade of organisation, the lichens ("Lichenes").
(b) Ichthyosporea (=Mesomycetozoa). Parasitic species that are closely related to the fungi. Some are pathogenic in humans.
(c) Choanoflagellata. The choanoflagellates are the sister group of the Metazoa. They comprise unicellular and colonial species. Choanoflagellate colonies are similar to sponges and could represent a model for the first step towards multicellularity.
(e) Metazoa. Multicellular animals; all species that occur in Germany will be covered in this Web repository.




This page has been updated on February 15, 2008
This site is online since May 31, 2005
Copyright © 2008 by Nikola-Michael Prpic. All rights reserved.



Related information:

Protozoa, Mesozoa, Metazoa

Tetramastigota

Discicristata to Planta

Amoebozoa, Opisthokonta

References