Is It Kingdoms or Domains

Confusion & Solutions

W I L L H. B L A C K W E L L

onfusion has arisen, at higher levels of classification of organisms, over two points: 1. How many kingdoms are there, i.e., which kingdoms should we recognize? 2. How do potentially larger, supra-kingdom groupings such as domains fit into the picture of a tradition of kingdom-based classification?

more than five kingdoms (Blackwell & Powell, 1995, 2001). The American Biology Teacher has thus been a significant forum for discussion of kingdom and suprakingdom concepts. Much additional debate is, of course, found in other sources as well (cf. Blackwell & Powell, 1999). The purposes of the present paper do not include repetition of a voluminous literature on the details of putative kingdom categories, but are: 1. to review pertinent historical points concerning the development of kingdom and domain concepts 2. to address questions arising from previous suggestions of kingdom and domain composition 3. to discuss recent investigations into such questions, particularly the taxonomic breakup of the heterogeneous kingdom Protista, and to suggest additional sorts of evidence that may contribute to our understanding of kingdom-level groupings 4. to acknowledge that future modification of classification systems will surely occur, and to consider which kingdoms should be recognized based on present knowledge.

Even if no other source is consulted, one may ascertain from the pages of The American Biology Teacher (ABT) that there is no universal agreement on these questions. Consider the appearance in ABT of the presentation (Margulis, 1981) of the popular five-kingdom concept, and a recent discussion favoring a system of three domains over the five-kingdom concept (Offner, 2001). Such papers have fostered additional discussion in ABT (e.g., Biermann, 2001). Also, papers in ABT have been thrown into the mix suggesting the efficacy of

WILL H. BLACKWELL is Professor Emeritus, Miami University (Ohio), and Adjunct Professor at The University of Alabama, Tuscaloosa, AL 35402; e-mail W60BUBBA@aol.com.

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Brief History of Kingdom (& Domain) Concepts

Understanding of the distinction between the animal and plant kingdoms is traceable to the science of ancient Greece (Margulis & Schwartz, 1998). Formal nomenclatural recognition of these two kingdoms, however, is dated from works of Linnaeus in the mid-eighteenth century (cf. Raven & Johnson, 2002). By the mid-nineteenth century, it was recognized that not all organisms are neatly pigeon-holed as either animals or plants, e.g., Euglena, and a third kingdom, containing mainly one-celled organisms was recognized. This unicellular kingdom was named Protista by Haeckel (1866). Earlier, Hogg (1860) had conceived of a similar kingdom Protoctista for organisms, unicellular or multicellular, which simply werent either plants or animals (cf. Margulis & Sagan, 1985). Though roughly equivalent, Hoggs Protoctista, interpreted to include large multicellular algae, is viewed as somewhat broader conceptually than Haeckels Protista. Various authors, including Haeckel, have recognized four kingdoms; however, a four-kingdom concept is often associated with Copeland (1947, 1956) who recognized: Mychota (for bacteria, not fungi as the name would seem to imply), Protoctista, Plantae, and Animalia. Regardless of name choices, Copelands careful exposition distinguished two basic kinds of unicellular organisms: bacteria (prokaryotes) and protists (eukaryotes)a distinction initially made by Chatton (1937). Some authors (e.g., Dodson, 1971) felt that Protista (or Protoctista) were not definable, and recommended a return to a system of three kingdoms (plants, animals, and prokaryotes). Authors with a yet greater reductionist bent (Dillon, 1963) favored recognition of only a single kingdom for all of life. The general trend, however, has been toward acceptance of an increased number of kingdoms. The five-kingdom approach represented a major milestone in kingdom concepts. The inception of the five-kingdom systemMonera (bacteria or prokaryotes), a name traceable to Haeckel, and a better name choice than Mychota; Protista (Haeckels kingdom); Fungi (added by Whittaker as a kingdom); Plantae; and Animaliais attributed to Whittaker (1969). This system was further elaborated by Whittaker and Margulis (1978). Margulis (1981) adopted the five-kingdom system, utilizing the name Protoctista, rather than Protista. Margulis use of Protoctista, consistent with Hogg (1860), was broader than Whittakers use of Protista, and included red and brown algae in addition to more usual protists. Since nomenclatural priority, i.e., priority of publication or authorship, does not often apply, in a formal sense, at the rank of kingdom (more stringently applied at family rank, and ranks below family), either the name

Protista or Protoctista may be selected. A little known fact (cf. Blackwell & Powell, 2001) is that Jahn & Jahn (1949) had recognized these five kingdoms some years earlier, as: Monera, Protista, Fungi, Metaphyta or Embryophyta (for Plantae) and Metazoa (for Animalia); they also recognized a sixth kingdom, Archetista, for viruses. Regardless of author credit, the five-kingdom approach dominated biology texts through the second half of the twentieth century. The five-kingdom approach is still used, but often with modification (e.g., Solomon et al., 2002); two bacterial kingdoms, Archaebacteria and Eubacteria, are often recognized, bringing the kingdom total to six. Additional kingdoms have been proposed, with varying acceptance. The purpose of increasing the number of kingdoms has not been, as some might suppose, to confound the readership; rather, the goal has been to establish more phylogenetically cohesive kingdom units (cf. Campbell & Reece, 2002). A major problem was presented by the heterogeneous nature of kingdom Protista, composed of both related and unrelated kinds of organisms (cf. Corliss, 1984; Blackwell & Powell, 1999, 2001). Most of the increase in kingdoms (pursuant to the five-kingdom concept) has resulted from dividing up the Protista into more logical groupings (cf. Campbell & Reece, 2002). Jeffrey (1971) proposed a seven-kingdom scenario including the kingdoms Rhodobiota (red algae) and Chromobiota (brown, golden, yellow-green algae, diatoms and Oomycetes), carved out of the Protista. Edwards (1976) had a similar scenario, using the name Erythrobionta instead of Rhodobiota, and Ochrobionta instead of Chromobiota. There is no shortage of neologisms, as Margulis & Sagan (1985) pointed out. Cavalier-Smith (1981) used the name Biliphyta for Rhodobiota, and (in 1986) Chromista for Chromobiota. Patterson (1989) introduced the informal term Stramenopiles for a natural grouping (possessing tripartite, tubular flagellar hairs) within the Chromista; the long-range effect of Pattersons proposal has been to question the inclusion of unrelated forms, such as Haptophytes and Cryptomonads, in the Chromista (cf. Blackwell & Powell, 2000, 2001). The category Stramenopiles has been set forth as a candidate kingdom, Stramenopila, by Campbell et al. (1999) and Campbell & Reece (2002); it was spelled Straminipila by Dick (2001). Not only the spelling of this name, but its status as a formally proposed kingdom could bear further scrutiny. Both Protozoa and more primitive Protozoa, the Archezoa, have been suggested as kingdoms (see discussion or listing of kingdoms in Cavalier-Smith, 1993; Corliss, 1994). The number of kingdoms recognized has usually ranged from seven to nine (cf. Cavalier-Smith, 1981; Blackwell & Powell, 1999). In an extreme view, Leedale (1974) suggested as many as 19 kingdoms.
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There is no over-riding consensus on which kingdoms should be recognized; and, there is also the matter of domains to consider. The introduction of domains could be said to begin with Woeses (1981) presentation on Archaebacteria. Woese designated three primary kingdoms: Archaebacteria, Eubacteria, and Eukaryotes. Woese contended that, in terms of ribosomes, transfer RNAs, and membrane lipids, the two major groupings of prokaryotes or bacteria (the Archaebacteria and the Eubacteria) are as different from each other as each is from the eukaryotes. Nine years later Woese et al. (1990) proposed that these most major groupings be elevated to the status of domains, with the names: Archaea, Bacteria, and Eucarya. The basis of recognition was that these three groupings represent, biochemically, the most fundamental subdivisions of cellular life. Woese apparently felt that taxonomic systems in use were inadequate in regard to recognition of these most basic categorical distinctionsthe primary tripartite division of the living world, as Woese stated it. Woese et al. (1990) thus formally proposed a new (higher) taxon or systematic category, Domain. Woeses three domains received acceptance by biologists, with some notable exceptions (e.g., Margulis & Schwartz, 1998). Consequently, in addition to the perpetuation of the five (or six)-kingdom concept (borne of Jahn & Jahn, 1949; Whittaker, 1959, 1969; Margulis, 1981), the three-domain concept of Woese has also found support in textbooks (e.g., Campbell & Reece, 2002; Raven & Johnson, 2002; Solomon et al., 2002). However, it is often unclear how kingdoms and domains are envisioned to interface. To hearken back to Offner (2001) and Biermann (2001), to some this is an either/or option. But, as I discuss, choosing between kingdoms and domains may be unnecessary.

priority when dealing with higher taxonomic ranks such as Kingdom, Phylum (Division), or Class. This truly becomes a moot point when dealing with alleged categories above kingdom level, since such categories are not recognized by the Codes. A more salient concern is, which proposal (three domains, Woese; or two superkingdoms, Jeffrey) is more biologically accurate? This is a difficult question, since cogent arguments could be made for either. Regardless, preference has generally gone to Woeses system in recent years, especially in terms of adoption by textbooks. Solomon et al. (2002), who surveyed research on this question, concluded that Archaea, which look like Bacteria, may be even more distinct from Bacteria (Eubacteria), biochemically, than from Eucarya. However, not all authors reached the same conclusion. Cavalier-Smith (1987) indicated that sequence homology between Eubacteria and Archaebacteria is greater than first thought; this closer relationship has been supported by certain recent evidence, cf. Brinkmann & Philippe (1999). In the alternative kingdom classification offered in Solomon et al. (2002), attributed to Cavalier-Smith (cf. Cavalier-Smith 1986, 2001), Prokaryota is recognized at the kingdom level, and any systematic distinction between Archaebacteria and Eubacteria would thus be below kingdom level, i.e. subkingdoms. As we shall see, however, Cavalier-Smith was not averse to changes of mind on such matters. In any case, a pragmatic problem associated with the recognition of Archaea and Bacteria as separate domains, is that the criteria of distinction are mainly biochemical, e.g., whether peptidoglycan is present in the cell wall or not. As indicated by Solomon et al. (2002), when viewed under the microscope, Eubacteria and Archaebacteria are very similar. It may be difficult for students to conceive that organisms, which appear to be virtually identical, are not in the same kingdom, perhaps not even the same domain. The above is not to suggest that the proposed, fundamental biochemical distinctions between Archaea and Bacteria are incorrect; there is, in fact, plenty of evidence that Woese was correct in his domain assessments. Yet, it is unquestionably easier to perceive, initially, the prokaryote/eukaryote distinction, than that of Archaea/Bacteria. Correspondingly, several authors, e.g., Barnes (1998), have adopted the essence of the superkingdom system of Jeffrey (Prokaryota and Eukaryota). Similarly, Cavalier-Smith (1993), in a change of mind from his 1986 publication, recognized as supra-kingdom categories, an Empire Bacteria (prokaryotes) and an Empire Eukaryota. Margulis & Schwartz (1988) had endorsed a similar distinction, but at a lower rank, employing the kingdom Prokaryotae (listing Monera as a synonym). Unexpectedly, though, Margulis (1992) raised the level

Problems Encountered with Present Domain & Kingdom Concepts

Domain is considered to be a supra(above or greater than)-kingdom category. The domains outlined by Woese (1990): Archaea, Bacteria and Eucarya, seem straightforward; so what could be the problem? For one thing, Woese was not the only author to propose suprakingdom categories. Jeffrey (1982) proposed two Superkingdoms: Prokaryota and Eukaryota, related to Chattons (1937) recognition of two basic structural types of cells, prokaryotic and eukaryotic. Since Jeffreys proposal of supra-kingdom categories preceded Woeses, then shouldnt Jeffrey receive priority recognition? This is not a significant problem, since all three kingdom-based Codes of NomenclatureBotanical (ICBN 2000); Zoological (ICZN 2000); and Bacteriological (ICNB 1992)place little emphasis on
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of Prokaryota (and Eukaryota, for that matter) to Superkingdom (or Domain, as she indicated in parenthesis). Margulis & Schwartz (1998) demonstrated that supra-kingdom, kingdom, and subkingdom categories may be intercalated without difficulty. Thus, it is frankly not clear at present whether we should favor a system of two supra-kingdom categories (sensu Jeffrey) or three supra-kingdom categories (sensu Woese). Campbell & Reece (2002) held the position that the three-domain system of Woese (1990), particularly recognition of Archaea and Bacteria, had rendered kingdom (or superkingdom) Monera (Prokaryota) obsolete. But, of course, this is so only if one accepts Woeses system of three domains. If the distinctions between the domains Archaea and Bacteria turn out to be not as great as first thought, then the broader category, Prokaryota, would seem to be reestablished. Bearing on the above, however, is a different, quasi-legal sort of question, one of a fundamental nomenclatural nature. Can we, in fact, accept (in a strict, formal taxonomic sense) any domains, empires, or super-kingdoms? I raise this issue because there is no provision for any sort of suprakingdom category in any of the three kingdom-based codes of nomenclature (ICBN, ICZN & ICNB). No taxonomic rank above kingdom is officially allowed, or even mentioned, regardless of Woeses (1990) formal domain proposal. Thus, any putative category above kingdom is spurious, in a nomenclatural sense (cf. Blackwell & Powell, 1999). One may ask, though, shouldnt such ranks be permitted? My answer would be, probably yes, with so many authors thinking that such are needed. But, as of the present, we dont legally have these categories. It may well be part of the solution that we need to bring biological nomenclature up to speed so that it will reflect improved biological knowledge (cf. Blackwell & Powell, 1999). This will require nomenclature proposals to this effect, and approval by the respective international congresses of botanical, zoological, and bacteriological nomenclature. But, on the other hand, should the lack of a formal, supra-kingdom nomenclature prevent us from utilizing, at least informally, such ranks as domains? I should think, clearly not! As far as formal naming methodology of the future is concerned, we may eventually have a unified

code of biological nomenclature (not three separate kingdom codes), in which such issues as supra-kingdom taxonony may be resolved (cf. Blackwell & Powell, 1999; Blackwell, 2002). It should be mentioned here in passing, that there are those (e.g., Hibbett & Donoghue, 1998) who would do away with all ranks (even the concept of rank), in the promotion of rankless classification. However, a rank-free system could cause chaos, providing little taxonomic structure. It is improbable as a reality. In any event, such possible changes are in the future; so the question now is, how do we solve present taxonomic problems concerning higher ranks? What

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recent evidence do we have? What are the most reasonable taxonomic categories to recognize based on present evidence?

Recent Evidence for Kingdom (or Domain) Decisions & Evidence Still Needed
Considering prokaryotic domains, there is, as has been mentioned, some evidence of a closer relationship of Archaea and Bacteria than initially suspected; additional work along the lines of that of Aravind et al. (1998), who demonstrated gene exchange between the two groups, might be a fruitful area of investigation. Among eukaryotic kingdoms, the recognition of animal, plant, and fungal kingdoms is relatively uncontroversial; although it should be mentioned that there is a greater relationship, based on molecular data, between animals and fungi than might have been expected a priori (cf. Borchiellini et al., 1998). However, it is really the recognition of new (ex-protistan) eukaryotic kingdoms pursuant to the breakup of the presumably simplest eukaryotic kingdom, Protista, or more broadly, Protoctistathat has been associated with the most uncertainty; see discussion in Blackwell & Powell (1995, 1999, 2001). Formerly within the encompassing Protoctista, the Rhodophyta (red algae) and the related glaucophytes (algae containing cyanelles derived Cyanobacteria) constitute a part of the consideration (cf. Margulis et al., 1990). Together, rhodophytes and glaucophytes constitute, according to some authors, the Biliphyta, in reference to possession by their plastids of bile pigments, phycobiliproteins, contained in small structural units, the phycobilisomes. The often multicellular red algae and related forms are a good case in point as to whether or not a given major group should be recognized as a kingdom. One need look no further than the work of Cavalier-Smith to sense there is a debate here; CavalierSmith (1981) recognized the Biliphyta as one of possibly nine kingdoms of superkingdom Eukaryota. Later, Cavalier-Smith (1987) relegated Biliphyta to a subkingdom of kingdom Plantae. And, the debate is still on. Solomon et al. (2002) considered red algae closely related to the plant kingdom; whereas, Campbell & Reece (2002) positioned them as related to plants, but somewhat more distantly. Molecular data give comparable results. Baldauf et al. (2000), based on protein data, connected red algae and glaucophytes to the plant kingdom. Kumar & Rzhetsky (1996), using small-subunit ribosomal RNA, also connected red algae to plants, but much more distantly; a distant relationship was suggested as well by large-subunit rRNA sequences (Perasso et al., 1989). Broadwater et al. (1992) had

noted that different ribosomal molecular-genetic sequences gave conflicting results. Based on rbcL data, some (e.g., Delaney et al., 1995) have even postulated a plastid connection between red algae and chromophytous algae (browns, goldens, etc.). Although ultrastructural studies have taken a back seat to molecular approaches, investigations of red algae could benefit by a return to the examination of ultrastructure. Distinct structures seen during mitosis, nuclear-associated organelles (Patrone et al., 1991) known as polar rings support the supposition of the unique nature of the group (Broadwater et al., 1992). Also, the absence of any flagellated structure or remnant of such (Gabrielson et al., 1990; Broadwater et al., 1992) in rhodophytes, speaks against a connection to green algae/green plants, in which flagellated cells figured heavily in their early evolution (Mattox & Stewart, 1984; Van den Hoek et al., 1995, Nakayama et al., 1998). Further work on the cytology of red algae could be potentially enlightening; Broadwater et al. (1992) decried the small number of workers in this specialty field. In any event, the paper of Kumar & Rzhetsky (1996) is particularly interesting in that a time line is estimated, on the basis of nucleotide sequencing. Red algae are suggested to be an ancient lineage, arising before the separation of green algae and land plants. This matches well with the fossil record (ancient for putative red algae), and with the speculation (Blackwell & Powell, 1995) that, whereas the plastids of both Viridiplantae (green algae/green plants) and Biliphyta (including red algae) were derived from Cyanobacteria, these were separate events. These respective symbioses were lineage-defining events, and a separation of green and red algal lines (and, for that matter, the brown algal line) has been maintained. This supposition is supported by a difference in structure of plastids, e.g., singlethylakoid lamellae, bearing phycobilisomes, in red algae vs. granal thylakoids, lacking phycobilisomes, in green algae and green plants (cf. Lee, 1999); chromophytes (brown algal line) have complex plastids (surrounded by four membranes), distinct from those of red or green algae (cf. Blackwell & Powell, 1995). I tentatively favor recognition of red algae (and relatives) as a separate kingdom, Biliphyta, based on present evidence, but additional molecular and ultrastructural studies would be helpful for any final decision. Other former protistan kingdoms (cf. Cambell & Reece, 2002) are equally important, but are accorded less discussion here because they have been thoroughly discussed elsewhere. The kingdom Chromista has been generally accepted since its intense exposition by Cavalier-Smith (1986, 1989). The remaining question with kingdom Chromista has been whether to recognize this grouping sensu lato, or in the more restricted sense of Stramenopilesa fundamentally heterokont

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(unequally flagellate) assemblage within the original Chromista (cf. Patterson, 1989; Blackwell & Powell, 2000). Stramenopiles are held together taxonomically, among a number of other characters, by the synapomorphy of composite (usually three-parted) tubular, reverse-thrusting, mastigonemes (flagellar hairs); these retronemes (Cavalier-Smith, 1989) are considered highly significant in evolution of the group. Blackwell & Powell (2000) provided an account of features suggesting that the Stramenopile lineage (consisting of chromophyte algae; pseudofungi; and certain primitive, heterotrophic flagellates) is fundamentally monophyletic. The problem with the kingdom Chromista is that it includes not only Stramenopiles, but relatively incongruous groups such as Haptophytes and Cryptomonads. A number of authors, e.g., Leipe et al. (1994) and Daugbjerg and Andersen (1997), have shown that Haptophytes (Prymnesiomonads) are not closely related to other chromophyte algae. Haptophytes, and the equally unrelated Cryptomonads, resemble types of flagellated Protozoa (cf. Blackwell & Powell, 2001). The preferred kingdom, for phylogenetic coherency, is thus Stramenopila (cf. Campbell & Reece, 2002), not Chromista. The remaining ex-protistan kingdom, Protozoa, has rather recently reemerged from the obscurity of submersion within kingdom Protista (cf. Cavalier-Smith, 1993; Blackwell & Powell, 2001; Corliss, 2002). The former kingdom Protista was the ultimate catch-all category, for placement of unicellular eukaryotes of uncertain relationship. The smaller, but still heterogeneous, kingdom Protozoa has replaced the Protista in this taxonomic holding-bin function. For example, Haptophytes and Cryptomonads are, at present, only loosely placed in kingdom Protozoa. All is not phylogenetically bleak, however, because the Protozoa also contain a number of related forms, such as the Alveolates: ciliates, dinoflagellates, and apicomplexans (sporozoans), cf. Corliss (1991). Alveolates are perhaps related to Stramenopiles (Van de Peer & De Wachter, 1997; Sogin & Silberman, 1998), but it is probably unwise to merge these groups at present. The relationship of Stramenopiles (Chromista) and Alveolates, however, holds significant potential for ultrastructural research into evolution. Recently, a residual plastid-like structure found in the alveolate (in this case, malarial) parasite Plamodium (Hopkins et al., 1999) has been linked to dinoflagellate plastids and ultimately to plastids of Chromista (Cavalier-Smith 1999); more work is needed to elucidate these relationships. Progress has been made in understanding relationships of a number of groups within the Protozoa, e.g. a relationship of Euglenoids and Trypanosomes (cf. Margulis & Sagan, 1985; Blackwell & Powell, 2001). Knowledge of relationship of Trichomonads and Hypermastigids,

as Parabasalids, has also been gained (cf. Levine et al., 1980; Blackwell & Powell, 2001). Protozoa have reemerged as a now somewhat more phylogenetically cohesive group than the unwieldy assemblage recognized early in the twentieth century (cf. Calkins, 1933). In any event, new phylogenetic insights, and the new kingdoms pared from the old Protista (Protoctista), render it unlikely that this large, hodge-podge kingdom (Protista, or Protoctista) will continue to be recognized, except informally.

Which Kingdoms & Subkingdoms, or Domains, Should Be Recognized? (Table 1)

As we have seen, the number of kingdoms recognized has varied widely although some consensus of seven or eight kingdoms (considering both prokaryotes and eukaryotes) has emerged in recent years (Corliss 1994, 2002; Blackwell & Powell, 1995, 1999, 2001; see also D. Maddison, The Tree of Life, http://tolweb.org/ tree?group=Life&contgroup=). The number of suprakingdom categories, such as domains, superkingdoms or empires, has usually been either two (Jeffrey, 1982; Cavalier-Smith, 1993) or three (Woese, 1990). Domain has been the most common supra-kingdom designation, and, for the sake of argument, I will settle for use of this term. But, the question remains (e.g., Biermann, 2001), is it domains or kingdoms? Discussed earlier in the paper (Problems Encountered with Present Domain & Kingdom Concepts), there is a question as to the nomenclatural legitimacy of the category domain, or of any suprakingdom category (empire, superkingdom, etc.). This, however, has not prevented authors of articles (e.g., Woese, 1990) and textbooks (e.g., Campbell & Reece, 2002) from using the rank, domain, and, in my opinion, should not. Nonetheless, the safest course of action, if one wishes to abide by present rules of biological nomenclature, is to use the largest category formally allowed by any of the nomenclatural codes, i.e., kingdom, and work down in inclusiveness from there (next, to subkingdom, for example). This approach was adopted by Margulis and Schwartz (1988). If we thus decide to have kingdom as our pinnacle of ranks, the question becomes, which kingdoms do we recognize? Better stated, perhaps, which system of kingdoms best represents the major phylogenetic lineages as we now understand them? As indicated at various points throughout this paper, particularly in Recent Evidence for Kingdom (or Domain) Decision & Evidence Still Needed, biological information recently available has taken us beyond the initially useful five-kingdom concept, which featured the large and admittedly heterogeneous

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Table 1. Four Plausible Kingdom, Kingdom/Subkingdom & Domain/Kingdom Scenarios, Based on Current Understanding of Major Phylogenetic Lineages.
Except for category emphasis, each presentation (A-D) below conveys essentially the same information. To render Woeses domains (C) compatible with codes of nomenclature, C may be converted to kingdom/subkingdom levels (D). The concern over domains versus kingdoms may be unnecessary; kingdoms and domains are readily interfaced, regardless of precise spelling or suffix selected. The category of domain (or empire, or superkingdom) is, however, not recognized by nomenclatural codes. But, utilization of subcategories, as allowed by codes, permits ample groupings (ranks) with which to represent the depth of variation in nature.

A. Kingdom Archaebacteria Kingdom Eubacteria Kingdom Protozoa Kingdom Stramenopila Kingdom Biliphyta Kingdom Plantae Kingdom Fungi Kingdom Animalia

B. Kingdom Monera (Prokaryota) Subkingdom Archaebacteria Subkingdom Eubacteria Kingdom Protozoa Kingdom Stramenopila Kingdom Biliphyta Kingdom Plantae Kingdom Fungi Kingdom Animalia

C. Domain Archaea Domain Bacteria Domain Eucarya Kingdom Protozoa Kingdom Stramenopila Kindgom Biliphyta Kingdom Plantae Kingdom Fungi Kingdom Animalia

D. Kingdom Archaebacteria Kingdom Eubacteria Kingdom Eukaryota Subkingdom Protozoa Subkingdom Stramenopila Subkingdom Biliphyta Subkingdom Plantae Subkingdom Fungi Subkingdom Animalia

kingdom Protista (or Protoctista), to a situation in which several kingdoms (Protozoa, Stramenopila or Chromista, and Biliphyta) have been split out of this former mega-kingdom. If we wished to recognize a straight system of kingdoms, eight in total, without subkingdoms, then the listing in Table IA, might be appropriate. Table 1B represents basically the same system, but one in which seven kingdoms are recognized, and Archaebacteria and Eubacteria are considered subkingdoms of prokaryotes (compare Margulis & Schwartz, 1988, with Blackwell & Powell, 1995). If we wish, however, to establish (regardless of rules of nomenclature) a system utilizing domains, i.e. following a Woesian paradigm, then Table 1C conveys basically the same information as IA, but employs Archaea, Bacteria and Eucarya as domains; under eukaryotes, in this system, there would thus be six kingdoms. Margulis & Schwartz (1998) unnecessarily concluded that use of domains mandated a substantial increase in kingdom number. To the contrary, one may extrapolate a relatively simple system of domains and kingdoms, such as we outline (Table IC), from the presentation in Campbell and Reece (2002). At a minimum, it is obvious that domains and kingdoms may be readily interfaced, if this is deemed desirable. In Table ID, we observe the same system as IC, converted to the kingdom/subkingdom level. I do not know of the existence of ID per se in other publications, but it is a system that

A. Eight kingdom approach, after Blackwell and Powell (1999, 2001). B. Seven kingdom, two subkingdom scenario, after Blackwell and Powell (1995), and Margulis and Schwartz (1988) as regards Monera and its subkingdoms. C. Three domains, six kingdoms; modified from Woese (1990), Blackwell and Powell (1999), and Campbell and Reece (2002). D. Presentation C, converted to kingdom/subkingdom levels: three kingdoms, six subkingdoms.

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could be defended based on our present knowledge, and it is in compliance with the rules of nomenclature. Perhaps of the systems discussed herein, ID is presently the best compromise.

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