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An Updated Compilation of World Soil Ciliates (Protozoa, Ciliophora), with Ecological Notes, New Records, and Descriptions of New Species

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An Updated Compilation of World Soil Ciliates (Protozoa, Ciliophora), with Ecological Notes, New Records, and Descriptions of New Species Europ. J. Protistol, 34, 195-235 (1998) European Journal of June 16, 1998 PROTISTOLOGY Review An Updated Compilation of World Soil Ciliates (Protozoa, Ciliophora), with Ecological Notes, New Records, and Descriptions of new Species Wilhelm Foissner Universitat Salzburg, Institut fur Zaalagie, Salzburg, Austria Summary community is different from freshwater and other cili­ ate assemblages. Since then many new species and This review compiles and analyses the taxonomic and bio­ records of soil ciliates have been described. The present geographical information available on soil ciliates. Key liter­ review summarises and analyses these and older taxo­ ature (257 references) is tabulated for each species. Further­ nomic and biogeographical data. Furthermore, about more, about 500 new records are reported from hundreds of samples collected world-wide, and four new species are de­ 500 new records are listed from hundreds of samples scribed from Kenyan soils, including the new genus Apo­ collected world-wide. During these studies, more than bryophyllum; Metopus ovalis Kahl, 1927 is redescribed. 643 500 new species, a few of which are described here, ciliate species were originally described or reliably recorded were found, confirming a previous estimation that from about 1000 soil samples world-wide, 49 (7.6%) of them there are at least 1330-2000 species of soil ciliates [96]. were later recognized as junior synonyms, and 78 (13.2%) have been poorly described, leaving a total of 516 well­ known species. However, the studied samples contained about 700 new species, most of which (about 500) have not Material and Methods yet been described. Thus, the total number of soil ciliates amounts to at least 1000 species. The overwhelming portion Areas and sampling of soil ciliates belongs to three systematic assemblages, viz. the hypotrichs (39%), gymnostomatids (26%) and colpodids About 817 samples were collected and analysed from (13%) if the undescribed species are taken into account. countries world-wide, including all main biogeographic re­ Most soil ciliates feed on bacteria (39%) or are predatory gions (Tab. 1, Fig. 1). Generally, collections were made from a (34%) or omnivorous (20%). Some are strictly my­ variety of biotopes covering most principal soil and vegeta­ cophagous and highly characteristic for terrestrial habitats; tion types of the respective region. Detailed site descriptions and a few (mainly metopids) are anaerobic. Only about one for the new records will be provided in later publications. fourth of the soil ciliate species has been reliably reported The material collected usually included mineral top soil from freshwater habitats and from more than three out of (0-5 em depth) with fine plant roots, the humic layer, and the five main biogeographical regions, indicating a high speci­ deciduous and!or grass litter from the soil surface. Further­ ficity of the soil ciliate biota and a limited distribution of at more, many samples contained some terrestrial or arbori­ least some species. This is supported by the observation that colous mosses with adhering soil and!or bark from trees. All some very conspicuous species, such as Krassniggia auxiliaris these habitats are referred to as "terrestrial", as opposed to and Bresslauides discoideus, have so far been found only in freshwater, because they contain, although in varying Gondwanan, respectively, Laurasian soils. amounts, true humic and mineral soil [73]. Usually, about 10 small subsamples were taken from an area of about 100 m2 Key words: Biodiversity; Community structure; Geo­ and mixed to a composite sample. All samples were air-dried graphic distribution; New species; Soil ciliates; Taxonomy. for at least one month and then sealed in plastic bags. Such samples can be stored for years without any significant loss of species [73, and unpub!. results]. Introduction Sample processing and community analysis The last comprehensive review on diversity and All collections were analysed with the non-flooded Petri ecology of soil ciliates was published in 1987 [73]. It dish method as described in [73, 83]. Briefly, this simple listed almost 300 species and showed that the soil ciliate method involves placing 10-50 g terrestrial material in a Petri © 1998 by GustavFischer Verlag 196 W. Foissner Table 1. Origin and number of samples investigated. dish (10-15 em in diameter) and saturating but not flooding it with distilled water. Such cultures were analysed for ciliates by Biogeographic region Number of inspecting about 2 ml of the run-off on days 2, 7, 14,21, and 28. samples investigated The non-flooded Petri dish method is selective, that is proba­ bly only a small proportion of the resting cysts present in a HOLARCTIS sample is reactivated, and undescribed species or species with specialized demands are very likely undersampled [96]. Thus, Austria about 300 1,3) the real number of species, described and undescribed, in the Germany about 10 1,3) samples investigated is probably much higher. Unfortunately, a Denmark 12 2) Finland 10 better method for broad analysis of soil ciliates is not known. Iceland 11 2) Portugal 2 Identification of species and species concept Italy 2 Greece 10 2) Identification of species was according to the literature Japan 232) cited in this paper. Most of the species found were either new or described or redescribed by my students and myself. Thus, subtotal 380 identification was mainly of live specimens using a high­ power oil immersion objective and differential interference contrast. However, all "difficult", new or supposedly new, PALAEOTROPIS species were treated with the silver staining techniques de­ Kenya 56 2) scribed in [82]. Usually, these methods yield permanent slides Tanzania 6 which have been or will be deposited in the Oberosterreichis­ Cameroon 8 che Landesmuseum in Linz (LI). Namibia (Etosha Pan) 12 The species concept, of course, influences the number of Republic of South Africa 10 species found and/or recognised as "undescribed". I usually apply the morphospecies concept which is, according to Fin­ subtotal 92 lay et al. [55], as valid as any, and probably more pragmatic than any other. I do not consider myself a splitter, that is I classify new species as such only when populations can be AUSTRALIS separated from their nearest relatives by at least one distinct (non-morphometric) morphological character. For examples, Australia and Tasmania see papers cited and species described in this review. NEOTROPIS Species descriptions Costa Rica 33 2) The species described were found in soils from Kenya, Venezuela 16 equatorial Africa. The soils of this region contain a very rich Brazil 52) ciliate community. 507 species were found in 92 samples, Peru 4 240 of them were undescribed [96]. Chile 29 Protospathidium terricola: Grassland soil (0-3 em) from Mt. Kenya near the lodge "The Ark", about 2300 m above sea subtotal 87 level. Soil reddish, humic, pH 6.4. Collected on 1. 7. 1985, in­ vestigated on 25. 7. 1986. Apobryophyllum terricola and Tachysoma humicola ARCHINOTIS longisetum: Litter, humus, and mineral soil from a small plant 1 Continental and maritime Antarctis about 73 ) island within young (some hundred years) lava masses of the Gough & Marion Islands 27 1) Shetani vulcano, a black and almost bare, fascinating area in the Tsavo National Park. Soil brownish, pH 7.6. Collected on subtotal 100 8.7. 1985, investigated on 1. 4. 1986. Metopus ovalis and Keronopsis dieckmanni: Grassland soil Total number of samples investigated 817 from shore of Lake Baringo, one of the highly saline, out­ standing Flamingo lakes in Kenya. Soil brownish, humic and 1) Taxonomic and faunistic data almost completely published, saline (salt crystals were formed when a drop of the soil water e.g. [2,9, 12-14, 16,62-65,69,70,73-75,77,81,94,95, 108, evaporated on the slide), pH 7.3. Collected on 2. 7.1985, in­ 111,113,161,162,183,184,189]. vestigated in October 1985. 2) Taxonomic and faunistic data partially published, e.g. [8, Methods follow those used in my previous papers [e.g. 82, 11,20,21,49,71, 72, 73, 75, 78, 87, 93, 97]. Note, however, 84, 95]. Two or three type slides (1 holotype and 1 to 2 para­ that almost all colpodids from all samples investigated so types) each of the new species described and two voucher far have been published [84- 86, 89, 90, 93]. slides of the species redescribed have been deposited in the 3) Many sites were investigated several times during a period Obcrosterreichischc Landesmuseum in Linz (LI), Austria. of 1-3 years; all samples from the other countries are from The slides usually contain many protargol-impregnated cells, different sites. Details of these sites will be provided in later with relevant specimens marked by a black ink circle on the papers. cover glass. Soil ciliate diversity 197 Fig. 1. Biogeographicregions according to [174]. Hatched areasare transition zones. Results and Discussion Uniqueness of the soil ciliate biota Only 25% (16% if the undescribed species are in­ The data are summarized in Tables 2-4, as in my pre­ cluded) of the described species have been reliably vious review [73]. The present compilation is more recorded from both soil and freshwater habitats (Tab. complete, considers extensively the older literature, and 4). Thus, there would appear to be a high proportion of includes many new records from soils world-wide species which occur only or mainly in terrestrial envi­ (Tab. 2). Furthermore, the quality of species descrip­ ronments and very likely evolved in such habitats [73]. tions was analysed and detailed taxonomic literature The most characteristic soil ciliates are colpodids, but compiled for each species (Tab. 3). many autochthonous genera and species occur also The new compilation largely confirms and among hypotrichs (e.g., Circinella spp., Hemisincirra strengthens the conclusions from my previous revie.w spp., Erniella filiformis, Terricirra spp.) and gymno­ [73], for instance, that soil ciliates are on average stomatids (e.g., Coriplites terricola, Pleuroplitoides smaller and more slender than freshwater ciliates (Tab.
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