Protozoan Taxonomy and Secondary article Systematics John O Corliss, University of Maryland, College Park, Maryland, USA Taxonomy

and systematics of major groups of the Protozoa, an assemblage of so-ca lled lower eukaryotes, refer basically to the classification of such groups, that is, their arrangement into a hierarchy of evolutionary interrelated groups (taxa) of scien tifically named phyla, classes, orders, etc. Article Contents .Introduction .Historical Considerations .Major Groups of Protozoa .The Old Phylum Protozoa .Modern Options Concerning the Place of Protozoa in the Biotic World .The New Kingdom Protozoa Introduction The classification of protozoa and other microorganisms above the organizational level of the bacteria has always been dependent on microscopy because the body sizes involved generally range from only one micrometre to one or

two millimetres in length. Any structures these species possess, useful in comparative studies of their morphology and thus their taxonomy and systematics, are at the cellular and subcellular levels, and invisible to the naked eye. The physiological properties of protozoa (and the neighbouring algae)have also played a role in the classification of these ubiquitous eukaryotic microorganisms; and of growing significance

are the findings made possible by research using molecular biological approaches. Because we are continuing to learn more and more about such minute organisms, protozoan systematics that is, the taxonomy (classification)and the evolutionary interrelationships of major groups of protozoa remains a topic of debate and change, still today. Some of the rather large and unwieldy taxonomic

groupings of past years are particularly subject to revision with expansion of and refinement in our knowledge about the members of those and related assemblages. Paradoxically, the protozoa themselves are becoming more difficult to define with precision as our information about them and other microbial assemblages increases. Thus, presenting a single satisfactory circumscribed definition for them is not

an easy task. It is attempted towards the end of this article; but, for sake of clarity, further background information is first supplied. Historical Considerations Until well beyond the middle of the twentieth century, protozoa were widely treated taxonomically as a (mere) subset of the kingdom Animalia, retaining their firstanimal definition dating from 150 years earlier. As a

phylum of unicellular animals, they were thought to exhibit major characteristics typical of that kingdom: colourless (i.e. no photosynthetic pigments present; yet, incongruently, a goodly number of chlorophyll-possessing algal groups were included in the phylum), phagotrophic, and capable of independent locomotion. It is now abundantly clear that this classical definition of protozoa is at best misleading and incomplete,

and that it requires considerable refinement. In former times, zoologists were the principal investigators and namers and claimers of such microorganisms; and they often worked with taxonomic disregard for studies of what might actually be the same group, sometimes even the same species, by the botanists (phycologists and mycologists). Thus, unfortunately, territoriality and authoritarianism also played

major roles in determining the early systematic status of the protozoa, algae and lower fungi (Corliss, 1986). Advances in our knowledge of the protozoa in general have followed progress in microscopy, as mentioned above. As early as a century ago, novel improvements in methods of light microscopy and related techniques of fixing and staining were already beginning to

make possible the revelation of morphological cellular characteristics that would have remained totally unrecognized before the appearance of such technological advances. Then, at mid-twentieth century, the use of electron microscopical approaches in cell biology opened up a whole new epoch of exploration in protozoology and related fields. During the Age of Ultrastructure (as it has been called), a myriad of

previously unknown subcellular structures were revealed that became of immense value in the comparative taxonomy of the lower eukaryotes (especially protozoa and algae). Even more recently, the exciting development of molecular biological approaches, particularly in study of genealogical and phylogenetic relationships within all groups of organisms, large and small, plant or animal, has offered a refinement in taxonomic investigations that

is without parallel in past decades. Here an attempt is made to review and understand the systematics of protozoa both from the more traditional point of view, keeping in mind that many biologists are still familiar mostly with older, conventional classifications of these minute organisms, and from more modern apENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics proaches. The latter are complicated by the relatively recent advent of the protistological perspective lumping protozoa, algae and some other eukaryotic microorganisms together as Protista and by their usage of more sophisticated data. For more detailed information concerning protozoan protistan overlapping relationships, the reader is referred particularly to comprehensive papers by Cavalier-Smith (1993, 1998)and Corliss (1994, 1998)

and to the insightful textbooks by Sleigh (1989)and Hausmann and Hu lsmann (1996). Major Groups of Protozoa For many decades (commencing with the perceptive schemes of Otto Bu tschli, published in the 1880s), it has been convenient to assign species known as protozoa to just four major categories or assemblages, as has been done in numerous textbooks of biology and

zoology as well as protozoology. Despite advances and improvements in our knowledge of eukaryotic microorganisms in general, it is still often rather helpful, for sake of discussion under diverse circumstances, to arrange protozoa taxonomically in such a way. The categories generally recognized are: (1)the amoeboid forms (the Sarcodina, in a broad sense); (2) the flagellated forms (the Mastigophora, including groups

of autotrophic or photosynthetic as well as heterotrophic species); (3) the ciliated forms (the Ciliophora, the most stable and perhaps most circumscribed of all protozoan assemblages); and (4) the various totally symbiotic or parasitic forms (primarily spore-forming species that are typically endoparasites, some highly pathogenic to their hosts, once assigned to a very broad group called the Sporozoa,

a high-level taxon that subsequently became divided into the Sporozoa and the Cnidosporidia). One of the pedagogically oft-regretted but inevitable changes, especially during the busy second half of the twentieth century (even before the protist revolution : see Corliss, 1986), was the tremendous expansion in the total numbers of high-level taxonomic groups (subphyla, classes, orders)of protozoa, while generally still

recognizing the four major top divisions mentioned above. The discovery of new and unique differentiating characteristics useful in classification and in evolutionary and phylogenetic studies as well required such a multiplication and fragmentation of taxa even though it placed/places greater demands on (the memories of)teachers, students and researchers alike. The Old Phylum Protozoa

Basically, the classification scheme of

the widely accepted Honigberg Report of 1964 may be used to illustrate the conventional situation concerning the systematics of the protozoa (Table 1). Although this popular arrangement in comparison with the long-followed classical one of Bu tschli of 80 years earlier contained nearly four times the number of taxonomic units above the level of family, it showed

few novelties of great significance. For an outstanding example of its conservatism, the classification still retained several major groups of algae, all treated as comprising a single class, the Phytomastigophorea, of flagellated protozoa (the Mastigophora). A subsequent revision supported by the international Society of Protozoologists, the Levine Report of 1980, recognized seven separate phyla (in a subkingdom Protozoa)and increased the

number of taxa above the familial level to 229 (Table 2). But, fundamentally, it followed the arrangement of its predecessor (Table 1). Five years later, the well-known and widely used Illustrated Guide to the Protozoa (Lee et al., 1985)appeared, with its classification scheme mainly that of the Levine Report. By then, the protist revolution had already been going strong for a full

decade (see accounts in Corliss, 1986, 1998)and many findings were indicating the pressing need for a fresh look at the old persisting problems of how to treat the systematics of the conventional/ traditional protozoan, algal and fungal assemblages of microorganisms. One reason for including the above discussion here, besides the fact that those classifications of 20 40 years ago

are still accepted by many biologists, is to help bridge the gap between such neoclassical systems and the suggested recent arrangements of the 1990s (see below). The organisms involved, and often their common group names as well, have not changed over the decades, but our ideas concerning their most likely interrelationships at the higher taxonomic levels have done so, primarily

by fresh analyses of the continuing accumulation of data of high phylogenetic and evolutionary significance from precise ultrastructural and molecular biological investigations (Coombs et al., 1998). Modern Options Concerning the Place of Protozoa in the Biotic World There are a number of ways in which modern biologists are viewing the overall placement of the protozoa with respect to other major groups of organisms. Several of

these are not truly taxonomic in their approach: for example, arrangement of species of protozoa based solely on their nutritional preferences or requirements, or considering protozoan groups as representing simply an evolutionary grade or level of cellular organization between the anucleate prokaryotes (essentially the bacteria)and the well-known higher (nucleated)eukaryotic forms of life (particularly the multicellular and multitissued plants and animals).

And popular today, especially among workers employing cladistic approaches in their investigation of ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 1 Taxonomic classification of the single phylum Protozoa according to the Report published by Honigberg et al. (1964), but omitting ordinal and subordinal names. The total number of suprafamilial categories reached 140, nearly four times the number in vogue a mong earlier protozoologists well into the twentieth century. (Modified and abbreviated from Corliss, 1998.) Subphylum I. Sarcomastigophora Subphylum II. Sporozoa Superclass 1. Mastigophora Class 1. Teleosporea Class 1. Phytomastigophorea Subclass (1) Gregarinia Class 2. Zoomastigophorea (2) Coccidia Superclass 2. Opalinata Class 2. Toxoplasmea Superclass 3. Sarcodina Class 3. Haplosporea Class 1. Rhizopodea Subclass (1) Lobosia Subphylum III. Cnidospora (2) Filosia Class 1. Myxosporidea (3) Granuloreticulosia Class 2. Microsporidea (4) Mycetozoia (5) Labyrinthulia Subphylum IV. Ciliophora Class 2. Piroplasmea Class 1. Ciliatea Class 3. Actinopodea Subclass (1) Holotrichia Subclass (1) Radiolaria (2) Peritrichia (2) Acantharia (3) Suctoria (3) Heliozoia (4) Spirotrichia (4) Proteomyxidia phylogenetic interrelationships of life forms, is the idea of clades, distinct evolutionary lines or lineages but ones often with unknown or nonpostulated possible taxonomic relationships with other lines of organisms

(microorganisms or macroorganisms)(see Patterson, 1999). The only true taxonomic options today are the inclusion of the protozoa with other eukaryotic microorganisms in a single kingdom called Protista or the recognition of a number of kingdoms (one called the Protozoa)within the overall assemblage of the Eukaryota. These two approaches require further discussion. Recognition of a single kingdom

to contain all eukaryotic microorganisms (and some of their larger but yet basically unicellular or single-tissued relatives)is a choice, first voiced more than 120 years ago by E. Haeckel and others, still popular today among a number of biologists. In this option, a neoHaeckelian kingdom Protista (or Protoctista )contains commingled phyla, divisions and classes of organisms formerly representing the protozoa, the

algae and taxa of lower fungi. It is given equal standing with the neighbouring kingdoms of the prokaryotic Bacteria, on the one hand, and of the (other) eukaryotic moieties known as the Fungi, Plantae and Animalia, on the other hand (Margulis et al., 1990; Margulis and Schwartz, 1998). This five-kingdom arrangement represents a pragmatic one that is quite

attractive pedagogically (easy to teach at all levels in the educational hierarchy)and is convenient for general nonspecialist usage and for information retrieval systems. But it is becoming evident that, as our store of relevant data continues to expand, the single kingdom Protista hypothesis lumps together lines of microscopic organisms that are proving to be not very closely related evolutionarily and

includes still other lines that are being shown to have closer affinities with forms assigned to one of the other three eukaryotic kingdoms. And the protozoa are not sufficiently given their own identity as an independent high-level taxonomic assemblage. The second major alternative available to us today is to consider the protists, overall, as best assignable to a

number of separate kingdoms, one of which has now been labelled with the familiar name Protozoa (Cavalier-Smith, 1993; Corliss, 1994). The composition of this protozoan assemblage distinctly separated from the four other eukaryotic assemblages or kingdoms recognized, the Chromista (taxonomic home for most lines of algae), Plantae, Fungi and Animalia (see Cavalier-Smith, 1998; Corliss, 1998, 2000) is considered below

in further detail. Two points need emphasis here. (1)A formal high-level taxonomic category called Protista is no longer needed or ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 2 Taxonomic classification of the animal subkingdom Protozoa according to the Report published by Levine et al. (1980), but here omitting all categories below the ra nk of class. The total number of suprafamilial categories reached 229, nearly double the number endorse d in the preceding Honigberg Report of 1964 (see Table 1). (Modified and abbreviated from Corliss, 1998.) Phylum I. Sarcomastigophora Phylum II. Labyrinthomorpha Subphylum 1. Mastigophora Class (1) Labyrinthulea Class (1) Phytomastigophorea (2) Zoomastigophorea Phylum III. Apicomplexa Subphylum 2. Opalinata Class (1) Perkinsea Class (1) Opalinatea (2) Sporozoea Subphylum 3. Sarcodina Superclass 1. Rhizopoda Phylum IV. Microspora Class (1) Lobosea Class (1) Rudimicrosporea (2) Acarpomyxea (2) Microsporea (3) Acrasea (4) Eumycetozoea Phylum V. Ascetospora (5) Plasmodiophorea Class (1) Stellatosporea (6) Filosea (2) Paramyxea (7) Granuloreticulosea (8) Xenophyophorea Phylum VI. Myxozoa Superclass 2. Actinopoda Class (1) Myxosporea Class (1) Acantharea (2) Actinosporea (2) Polycystinea (3) Phaeodarea Phylum VII. Ciliophora (4) Heliozoea Class (1) Kinetofragminophorea (2) Oligohymenophorea (3) Polyhymenophorea recognized; all former protists (spelled with a lower case p )can be reassigned to various of the

five kingdoms mentioned immediately above. (2)The (new)kingdom Protozoa, while containing many conventional groups of protozoa, is purged of others and thus is not identical in overall taxonomic composition to the old phylum Protozoa embraced in very popular classifications of both distant and more recent past years (for example, see preceding sections of this article; see also Honigberg et al., 1964,

and Levine et al., 1980; and contrast the contents of Table 1 and Table 2 with that of Table 3). The New Kingdom Protozoa

Of the two options briefly described above, many biologists now favour the second one, the concept of dividing up major taxa of former protozoa (and of algae and lower fungi, too), with new assignments to several different kingdoms, one of which

is identified as the kingdom Protozoa. The first person to do this systematically and in some detail was T. Cavalier-Smith, starting in the 1980s (see full discussions in Cavalier-Smith, 1993, 1998). The kingdom Protozoa may be briefly described (after Corliss, 1994, 2000)as follows. The kingdom comprises predominantly unicellular, plasmodial or colonial protists (eukaryotic microorganisms) that are

mostly phagotrophic or osmotrophic, colourless, lacking cellulosic cell walls, and microscopic in body size. Included species that are capable of photosynthesis (but some are nutritionally mixotrophic) typically possess unique cytosolic plastids with stacked thylakoids, no stored starch, and surrounded by three membranes. Nearly universally present are tubular mitochondrial cristae; when mitochondria are absent, they are typically replaced by hydrogenosomes. Golgi

bodies and peroxisomes are widely present. Flagellar mastigonemes, if present, are never rigid or tubular. Free-living (typically independently motile via pseudopodia, flagella, or cilia) and symbiotic species are numerous from a great variety of habitats. As one of the two (of five)eukaryotic kingdoms ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Table 3 The 14 phyla, including authorships and dates of their names, comprising the kingdom Protozoa Goldfuss, 1818, with an indication of the kinds and numbers of protists included in each phyletic taxon (based on abbreviated characterization data from Corliss, 1994, 1998, 2000) Archamoebae Cavalier-Smith, 1983 Large, benthic, microaerobic amoebae, amitochondriate, allegedly primitive forms, with endosymbiotic

bacteria; species few in number, all free-living in fresh water Neomonada Cavalier-Smith, 1997 Often small, free-living, marine heterotrophic flagellates and amoeboflagellates; small group, still ill-defined Rhizopoda von Siebold, 1845 Typically amoeboid, with differing kinds of pseudopodia, some flagellated forms; naked or with tests or thecae; 4 5000 species found in soil, fresh-or saltwater habitats

Mycetozoa de Bary, 1859 Plasmodial slime moulds (cellular and acellular), some very large; aerial (stalked) fruiting bodies produce spores; c. 850 species, mostly in decaying vegetation; a few symbiotic forms Foraminifera d Orbigny, 1826 Amoeboid forms in tests (usually calcareous), with alternation of haploid sexual and diploid asexual generations; shells of extinct species reach 15 cm in diameter; reticulate

pseudopodia for feeding and locomotion; mostly marine, with some 45 000 species (largest phylum in the kingdom)of which c. 90% are fossil forms Heliozoa Haeckel, 1866 Mostly freshwater group of the classical actinopod sarcodinids , with slender radiating axopodial type of pseudopodia used in food capture; c. 100 species, many stalked Radiozoa Cavalier-Smith, 1987 Marine, spherical forms,

typically planktonic, often with elaborate symmetrical shell pierced by stiff axopodia; three major subgroups, with total of nearly 12 000 species (c. 65% fossil forms), second largest phylum in the kingdom Percolozoa Cavalier-Smith, 1991 Small heterotrophic flagellates or amoeboflagellates, c. 100 species, some poorly known Euglenozoa Cavalier-Smith, 1981 The old Euglenophyta , mainly free-living, freshwater phytoflagellates , 4

1000 species, plus Kinetoplastidea (parasitic trypanosomes plus free-living bodonids), 4 600 species; commonly with discoidal mitochondrial cristae and a paraxial rod in their main flagellum Dinozoa Cavalier-Smith, 1981 Dinoflagellates, unique biflagellated protists, mostly marine planktonic, one-half pigmented forms; some thecate; a few colonial; cortical alveoli; about half found as fossils; total species c. 4500, with some 100 described

as parasites; many orders Metamonada Grasse , 1952 Biflagellated to multiflagellated forms, typically digestive tract parasites (insects to humans); c. 300 species; hydrogenosomes in place of mitochondria Parabasala Honigberg, 1973 Parasitic multiflagellated forms, amitochondriate and with prominent parabasal (Golgi)apparatus; 4 400 species, in intestines of woodroaches to humans Apicomplexa Levine, 1970 Essentially the Sporozoa

of old; unique complex of apical organelles; all symbiotic, with many minute species as harmful endoparasites (e.g. in birds, livestock, humans: outstandingly, malaria); cortical alveoli; 4 5000 species in three major classes Ciliophora Doflein, 1901 All heterokaryotic (micro-and macronuclei); usually multiciliate, phagotrophic, relatively large protists found mostly free-living in diverse fresh-/saltwater/soil habitats; others symbiotic or epibiotic, mostly in

or on invertebrate hosts; often complex oral ciliature; cortical alveoli; many exhibit sexual phenomenon of conjugation; asexual reproduction by transverse fission; third largest protozoan phylum: c. 8000 species in 8 10 classes, many orders composed purely of protistan forms, the Protozoa As characterized above, the refined kingdom Protozoa, counting fossil as well as extant species embrace nearly

large though it is, is considerably more discriminating and 40% of all protists (numbering some 213 000 species) more restricted in its boundaries than was the old phylum described to date. Protozoa. The latter, traditionally, contained a number of ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net

Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics major taxa (e.g. at class and ordinal levels)that have now been eliminated. For example, purged from the new kingdom have been half a dozen algal groups, the curious labyrinthomorphids, the choanozoa, the opalinids, the myxozoa, and most recently, the microsporidia. All species of these taxa fail to possess major distinguishing features of the kingdom Protozoa listed above and/or

have unique morphological, physiological or molecular characteristics of their own beyond discussion here. Supported by recent molecular (e.g. rRNA sequencing information)as well as morphological and biochemical findings, these other protists have been placed in one or another of the neighbouring kingdoms Chromista, Fungi, Plantae or Animalia (Corliss, 1998, 2000). The 14 phyla (containing nearly 83 000 species, about 60% of

which are fossil and 12% symbiotic forms)that then remain in the refined kingdom Protozoa are listed, with abbreviated descriptions, in Table 3. Representative genera are not mentioned, for sake of brevity within the table, but names of many such genera are included in the long papers by Cavalier-Smith (1993)and Corliss (1994). Several additional observations might be helpful concerning some

of the major differences between this new classification of the Protozoa and the older, more conventional ones discussed on preceding pages. Two distinctive eukaryotic algal lines the euglenids and the dinoflagellates are retained as Protozoa, but they have strikingly different characteristics (including presence of three, not two, plastidic membranes and frequent exhibition of nonphotosynthetic modes of

nutrition, among others)from other major assemblages of algae now appearing in the separate kingdoms Chromista (e.g. the chrysophytes, phaeophytes, haptomonads, cryptomonads and lesser heterokontic groups)and Plantae (e.g. the prasinophytes, chlorophytes (with such long-claimed protozoa genera as Volvox and Chlamydomonas), ulvophytes, charophytes, rhodophytes and glaucophytes). In fact, the euglenophytes of old are phylogenetically closely related to the colourless kinetoplastideans (bodonids and

parasitic trypanosomatids), the combining single phyletic name now being the Euglenozoa. The dinoflagellates (Dinozoa), furthermore, are linked with the ciliates (Ciliophora)and the sporozoa (Apicomplexa) in a supraphyletic assemblage often designated as the Alveolata because of their common possession of cortical alveoli in their pellicles, a character not particularly known in other phyla of protists. Mostly on

the basis of very recent rRNA data, but also supported by other differentiating characteristics, the longlinked protozoan-cnidosporidian groups, the enigmatic microsporidia and myxosporidia, turn out to be not only not closely related to each other but also not members of the kingdom Protozoa. The first, the Microspora, along with chytrids (but not with oomycetes and hyphochytriomycetes, which are now

in the Chromista), are probably primitive or degenerate protistan members of the kingdom Fungi, while the complex Myxozoa are better classified in the Animalia (perhaps close to the cnidarians). Until rather recently, the Archamoebae (with genus Pelomyxa and possibly others such as the important parasite of humans, Entamoeba)were considered as very primitive protists in a kingdom

of their own. Latest studies (see references in Cavalier-Smith, 1998; Corliss, 2000) suggest that, while probably composed of very ancient species (now without mitochondria or hydrogenosomes), the phylum belongs more appropriately in the revised kingdom Protozoa, as seen in Table 3. The unique Choanozoa, collar-celled, loricate, colourless flagellates with flattened mitochondrial cristae, presumably at the crossroads of origins of

both kingdoms Animalia and Fungi, have been removed from the Protozoa and tentatively placed in the Animalia (Corliss, 2000), where another protistan group and former protozoan phylum, the Myxozoa, is also now located (see above). References Cavalier-Smith T (1993)Kingdom Protozoa and its 18 phyla. Microbiological Reviews 57: 953 994. Cavalier-Smith T (1998)A revised six-kingdom system of life. Biological

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JJ, Hutner SH and Bovee EC (eds)(1985)An Illustrated Guide to the Protozoa. Lawrence, KS: Society of Protozoologists. Levine ND, Corliss JO, Cox FEG et al. (1980)A newly revised classification of the Protozoa. Journal of Protozoology 27: 37 58. Margulis L and Schwartz KV (1998) Five Kingdoms: an Illustrated Guide to the Phyla of Life on Earth, 3rd edn. New York: WH

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Protozoan Taxonomy and Systematics Protozoan Taxonomy and Systematics Further Reading Andersen RA (1992)Diversity of eukaryotic algae. Biodiversity and Conservation 1: 267 292. Cavalier-Smith T (1989)The kingdom Chromista. In: Green JC, Leadbeater BSC and Diver WL (eds) The Chromophyte Algae: Problems and Perspectives, pp. 381 407. Oxford: Clarendon Press. Cavalier-Smith T (1995)Evolutionary protistology comes of age: biodiversity and molecular cell biology. Archiv fur Protistenkunde

145: 145 154. Corliss JO (1998)Haeckel s kingdom Protista and current concepts in systematic protistology. Stapfia 56: 85 104. Grell KG (1973) Protozoology. Berlin: Springer-Verlag. Kudo RR (1966) Protozoology, 5th edn. Springfield, IL: Charles C Thomas. Margulis L (1996)Archaeal eubacterial mergers in the origin of Eukarya: phylogenetic classification of life. Proceedings of the National Academy of Sciences of the

USA 93: 1071 1076. Patterson DJ (1994)Protozoa: evolution and systematics. In: Hausmann K and Hu lsmann N (eds) Progress in Protistology (Proceedings of the IX International Congress of Protozoology, Berlin, July August 1993), pp. 1 14. Stuttgart: G Fischer. Vickerman K (1998)Revolution among the Protozoa. In: Coombs GH, Vickerman K, Sleigh MA and Warren A (eds) Evolutionary Relationships Among

Protozoa, pp. 1 24. Dordrecht: Kluwer Academic Publishers. ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www.els.net