The Protozoa, a Kingdom by Default?
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A Kingdom By Default? WILL H. BLACKWELL MARTHAJ. POWELL Those Many Kingdoms A Thumbnail on Protozoology Downloaded from http://online.ucpress.edu/abt/article-pdf/63/7/483/50291/4451168.pdf by guest on 02 October 2021 A good start to a revaluationof the Protozoa is a hanges in concepts of kingdoms of brief look at some main events in the history of study organisms over the latter 20th century of the group, especially in North America. We can have been substantial (Blackwell & Powell here venture the bottom line first. If we span the 1999), sometimes tedious, yet with the worthy goal course of protozoology during the 20th century, we of establishing groupings of organisms reflective of eventually arrive at two apparently contradictory major evolutionarylineages. An increasing number of conclusions: (1) the classification of no group we kingdoms is recognized, typically including two have recognized as a phylum, subkingdom or king- prokaryotic kingdoms, Archaea(Archaebacteria) and dom has changed more, (2) nor has the classification Bacteria(Woese et al. 1990), and six or more eukary- of any group ultimately come more full-circle. otic kingdoms: Protozoa (the amitochondrial mem- Perhaps these seemingly disparate points will be bers sometimes separated as kingdom Archezoa), clear by the end of our discussion. Chromista (including the major group, Stramenopiles), Biliphyta, Plantae, Fungi, and The first major textbook on protozoology in the Animalia (cf. Cavalier-Smith 1987; Corliss 1994; United States, The Protozoa, by Gary N. Calkins Blackwell& Powell 1995). Still more kingdoms have (1901) of Columbia University, probably had more been suggested; Leedale (1974), for example, out- impact in this field of study than any other single lined as many as 19 kingdoms among scenarios he work. Seemingly heedless of the three-kingdom discussed. approach of Haeckel (1866), who proposed a unicel- lular kingdom, Protista, Calkins (1901) followed the Difficulty is encountered in seeking an exclusive Linnaeanview of life as divisible along plant and ani- definition of several of these kingdoms. In particular, mal lines. He recognized the phylum Protozoa as an the Protozoa is loosely characterized;its membership apparently definable group, i.e. unicellular organ- is decided partially by the fact of not belonging to isms, within the otherwise multicellular kingdom another kingdom (Cox 1991; Cavalier-Smith1993). Animalia. But since Protozoa are, in a selection of recent litera- ture, referred to as a kingdom (e.g. Cavalier-Smith Calkins'system of four fundamentalclasses with- 1993; Blackwell& Powell 1995; Beakes 1998; Corliss in the phylum Protozoa-Mastigophora (flagellates), 1994, 1998; Vickerman 1998), it is of possible bene- Sarcodina (amoebae), Infusoria (ciliates), and fit to teachers of biology to reassess the characteriza- Sporozoa (gregarines,coccidea)-remains influential tion, confines and membership of this potential king- (a basis for our proposition of the "full-circle"arrival dom of organisms. of knowledge in protozoology). With minor changes, Calkins' system was echoed in works on Protozoa (and in the "protozoan section" in general works of zoology) through the first half of the 20th century. WILLH. BLACKWELLis an AdjunctProfessor and MARTHAAJ.POWELL For example, R. P. Hall (1953), of New York is Professorand Chair of the Departmentof BiologicalSciences at University,produced an excellent survey of Protozoa, the Universityof Alabama, Tuscaloosa,AL 35487-0344. titled "Protozoology";Hall employed the traditional, THEPROTOZOA, AKINGDOM BYDEFAULT? 483 locomotion-based,light microscopic charactersof some authors (e.g. Levine et al. 1980), but not others Protozoaestablished by Calkins,adding additional (e.g. Jahn et al. 1979). In more recent classifications informationgained during the 50 years following (e.g. Cox 1991; Sleigh 1991), formal headings for Calkins'original text (1901). For reference,we pres- major protozoan groups were often dropped. ent the outlineof Hall'ssystem (Table 1), representa- tiveof mid-century,traditional protozoological classi- Recognition of the Protista; fication.But, changes were in the offing.Questioning the ultimate divisibilityof amoeboid and flagellate Obscuring of the Protozoa protozoans,Honigberg et al. (1964) combinedthese as "Sarcomastigophora'-amodification followed by Having seemingly forgotten, for a hundred years, Haeckel's (1866) proposal for a king- dom Protista,realization reoccurred that a number of unicellular organisms, e.g. TABLE 1. Euglena, dinoflagellates, chrysamoebas, Groups of Protozoa, Hall 1953._ cryptomonads, are not readily fitted into plant and animal kingdoms. As a conse- Downloaded from http://online.ucpress.edu/abt/article-pdf/63/7/483/50291/4451168.pdf by guest on 02 October 2021 Subphylum1.Mastigophora Subclass3.Haemosporidia quence, the kingdom Protista was again Class1. Phytomastigophorea Class2. Cnidosporidea invoked (e.g. Whittaker 1969). With this "rebirth"of Protista came a similar Order1.Chrysomonadida Order1.Myxosporida realizationthat fungal organisms are nei- Order2.Heterochlorida Order2.Actinomyxida ther plant nor animal, and the kingdom Order3.Cryptomonadida Order3.Microsporida Fungi was also established. Whittaker's Order4.Dinoflagellida Order4.Helicosporida five-kingdomproposal - Monera (bacte- OrderS.Phytomonadida Class3. Acnidosporidea ria, prokaryotes), Protista (unicellular Order6.Euglenida Subclass1.Sarcosporidia eukaryotes), Plantae, Animalia and Fungi - pervaded texts of biology for Order7.Chloromonadida Subclass2.Haplosporidia two decades, and is still occasionally Class2. Zoomastigophorea Subphylum4.Ciliophora advocated. Whittaker (1969), and later Order1.Rhizomastigida Class1. Ciliatea Margulis(1974, 1981), who preferenced Order2.Protomastigida Subclass1.Protociliatia the name Protoctista over Protista, are Order3.Polymastigida Order1.Opalinida associated with the development of the Order4.Trichomonadida Subclass2.Euciliatia five-kingdomconcept; however, the core of this proposal (and a suggested addi- Order5.Hypermastigida Order1.Holotrichida tional kingdom for viruses) appeared in Subphylum2.Sarcodina Suborder1.Astomina Jahn and Jahn's (1949) "How to Know Class1. Actinopodea Suborder2.Gymnostomina the Protozoa." Order1.Helioflagellida Suborder3.Trichostomina What did establishment of the Order2.Heliozoida Suborder4.Hymenostomina five-kingdom system mean to the recog- Order3.Radiolarida Suborder5.Thigmotrichina nition and classification of Protozoa?At Class2. Rhizopodea Suborder6.Apostomina first it simply implied a transfer of the Order1.Proteomyxida Order2.Spirotrichida unicellular phylum (or subkingdom) Protozoa from the multicellular king- Order2.Mycetozoida Suborder1.Heterotrichina dom, Animalia, to the unicellular-based Order3.Amoebida Suborder2.Tintinnina kingdom Protista (e.g. Jahn et al. 1979). Order4.Testacida Suborder3.Oligotrichina However, delimitation of Protozoa soon Order5.Foraminiferida Suborder4. Entodiniomorphina blurred amid the taxonomic melange of Subphylum3.Sporozoa Suborder5.Hypotrichina the inflated kingdom Protista. John Corliss' (1981) paper, "Whatare the tax- Suborder6.Ctenostomina Class1.Telosporidea onomic and evolutionary relationships Order3.Peritrichida Subclass1.Gregarinidia of the Protozoa to the Protista?,"went to Order1.Eugregarinida Order4.Chonotrichida the heart of this issue. This question of Order2.Schizogregarinida Class 2. Suctorea protozoan, and other protist group, Subclass2.Coccidia identities was again voiced by Corliss 484 THEAMERICAN BIOLOGY TEACHER, VOLUME 63, NO.7,SEPTEMBER 2001 (1983) in a paper titled, perhaps tongue-in-cheek,"A nonetheless continue to be recognized in some Puddle of Protists." recent biology texts (e.g. Johnson 2000). Corliss raised a two-partedissue: Purging the Protista; 1. The kingdom Protistawas too large, too inclu- sive, too polyphyletic (a largely unnatural Reemergence of the Protozoa 45 phyla assemblage). Corliss (1984) listed The solution to an identity dilemma (i.e. what under Protista, with more to come. The "uni- does/does not constitute Protista?)lies in acquisition in Protista cellular criterion"for "membership" of knowledge. An advance was Cavalier-Smith's was compromised by inclusion of predomi- (1986, 1989) recognition of the kingdom Chromista. brown nantly multicellular groups, such as The largest assemblage within the Chromista is phy- algae (Phaeophyta) and red algae lum Heterokonta,a name referringto unequal flagel- (Rhodophyta). la of many of its members; the more forwardof these 2. Distinctions of formal, major taxonomic flagella generally bears tubular (2- or, more often, mastigonemes (flagellarhairs). In reference groups within the Protistawere vanishing. 3-parted) Downloaded from http://online.ucpress.edu/abt/article-pdf/63/7/483/50291/4451168.pdf by guest on 02 October 2021 to these mastigonemes, Patterson (1989) gave this Given this breakdown of protistan macrosystem- natural heterokont assemblage the appropriate,but atic infrastructure, Corliss (1981) recognized five informal, name Stramenopiles ("straw hairs"). informalgroupings of protist phyla:(1) the protozoan Stramenopila, more monophyletic than Chromista, group;(2) the protozoalgalgroup; (3) the algal group; has been suggested as a candidate kingdom (e.g. (4) the protozofungalgroup; and (5) the fungalgroup, Campbell et al. 1999). for "fungalprotists," not for the kingdom Fungi. Photosynthetic Stramenopiles typicallypossess a The Protistabecame an unwieldy colossus, hous- "chloroplast endoplasmic