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A Simple Key to the Bdelloid Rotifers

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A Simple Key to the Bdelloid Rotifers Quekett]ournal ofMicroscopy, 1999,38,351- 356 351 A simple generic key to the Bdelloid rotifers PAUL N. TURNER Summary comparisons or genetic tests, since cross-breeding or inbreeding does not occur and cannot be Rotifers of the Order Bdelloidea occupy a vast number of induced. natural habitatS, from ice to soil, lichens to mosses. fresh· water interstitial to salrwater sands, from the tropics to the The proper identification of bdelloids requires Antarctic. Because of their apparently cosmopolitan that they be viewed in-vivo, which in turn requires presence, an understanding of their taxonomy is critical, special care and handling of the animals and the especially since more and more ecologists are confronted medium. Bdelloids are particularly sensitive to with the need tO identify them in sampling regimes. changes in temperature, light, oxygen content, or 'bdelloid sp.' Since descriptions such as 'rotifer sp.' and vibrations, so these factors must be controlled are becoming more and more unacceptable in a modern to be successful. species list, this paper presents a relatively simple numeri· cal key that will allow non-expens tO identify most The customary use of a high-intensity light­ common bdelloid rotifers to genus. Although the Order source for microscopic examination may need to is in much need of revision at species level, the current be altered while viewing bdelloids, since the heat generic names and criteria will probably persist through generated by the light source may provoke any such revision, and thus generic-level identifications animal contraction. Likewise, physical vibrations accurate in the produced with this key will likely be in the water, on slide or watch-glass, created by future as well. focusing or by moving the mechanical stage, may induce contraction. If the animal stays con­ Introduction tracted for more than a few minutes, it is not likely to recover for some time. If this occurs, AccuRATE identification of Bdelloid rotifers can the viewer may be forced to wait for the rotifer re on rotifer taxonomy be very difficult. Literatu tO recover from the agitation, or seek another seldom discusses details of the Bdelloids in example for identification. common terms. Works that deal with the taxo· nomy of the Order in any detail are usually large, technical, and aimed at specialists. As a Background result, there are very few references of note A short history of the research, and a partial occupying the 'middle' ground between the listing of references available, begins with the amateur and specialist. many works of Murray (1902- 1913) and Bryce When presented with the identification, non­ (1892- 1929). They combine to assume the most specialists t urn to the existing taxonomic refer­ significant baseline information used for the ences, but this difficult chore is compounded in later works on taxonomy and ecology of three ways. (1) the subtleties of acceptable specific Remane (1 932/33) and Voigt (1957). Subsequent features, (2) concurrent questions about what is a works by Bartos (1951 ) in English, and Donner true species in a completely parthenogenetic (1965) in German, are the most recent and most group of animals, and (3) the difficulties of detailed compilations of significance. Additional bdelloid rotifer handling. Suitable identifications efforts include Koste & Shiel (1986) discussing are also hindered by the extreme amount of Australian bdelloids as a part of a series on time usually required of the investigator in the Australian rotifers, and the six works of H aigh attempt to wade through the subtle specific (1963- 1971) that discuss New Zealand bdelloids. elements for good identifications. As a result, These later works are among the last compre­ "bdelloid sp." too often shows up on microfauna hensively to examine the taxonomy, ecology and lists. systematics of the Order. Non-taxonomic & The entire Order Bdelloidea (bdelloids) con­ research in the o rder is described by Bateman Zullini & Ricci (1980), sists only of female rotifers. There are no Davis (1980), Ricci (1987), known males. In one sense this simplifies the and others. chore of identification, since there is only one The current state of understanding is au_gmen­ gender for which to accumulate identification ted by virtual works such as the Aydin Orstan elements. Without males, however, acceptable bdelloid web page a~ http://members.aol.com/ species identifications must rest on phenotypic bdelloidl/deloid.htm. Orstan discusses the entire 352 PAUL N. TURNER group with very useful diagnostic keys, photos, treatment techniques. Donner (1965) is the drawings and ecological data. preferred, and possibly the most authoritative work on bdelloid rotifers, but published only in Discussion German. Use of other keys such as that by Bdelloid rotifers can be found in almost any TABLE 1 environment. temporary or permanent wet Generic Identification Elements Many are considered aquatic, flourishing in the benthos or littoral of ponds and streams, while 1. Rostrum imperfectly developed others occupy terrestrial habitats such as moist 2. Rostrum non-retractile soil, mosses, leaf litter and lichens. These rotifers 3. Wheel organ (ciliated field) prone and flat are notable for their ability to resist severe 4. Wheel organ ring, with dorsal and ventral breaks environmental changes to their habitat. The pro­ 5. Upper (dorsal) lip undivided cess and morphology of their ability to resist 6. Upper (dorsal) lip bilobed, or divided dehydration (anhydrobiosis) is documented in 7. Pedicels rudimentary, or absent well developed Jacobs (1909), Rahm (1926), Ricci (1987) and 8. Pedicels 9. T rochal disc inset, between 2 horn-like processe.s (1958) to mention a few. Only a rela­ Umezawa 10. Trochal disc between hoodlike expansion of upper d r otifers tively small number of specific bdelloi lip have been actually documented performing 11. Throat distinct anhydrobiosis, (Philodina roseola, Adineta vaga, 12. Gullet absent Macrotrachela musculosa, Mniobia magna, Macro­ 13. Rami protrusile trachela quadricornifora, Habrotrocha constricta, 14. Lumen wide Philodina acuticornis odiosa, Mniobia russeola) 14a. Lumen absent, where food pellets 611 body cavity under special, slow drying conditions (Gilbert, 15. Lumen narrow 1974), yet the process is logically extrapolated 16. Food in pellets (do not mistake oil droplets for food) for the rest of t he Order by their reappearance 17. Food NOT in pellets lichen and soil habitats in which in the moss, 18. Abdominal transverse cuticle folds correspond to they often occur. body segments Non-bdelloid species (those of the other major 19. Abdominal transverse cuticle folds numerous, class of Rotifera, Eurotatoria (Monogononta)) NOT corresponding with body segments also sometimes suddenly reappear in open perma­ 20. Foot long, slender nent or temporary waters after a long stretch of 21. Foot short, stout ending in sucker-like plate, disc or discs dryness. The mechanism responsible for their 22. Foot 23. Foot glands in longitudinal series can be attributed to an equally significant presence 24. Foot glands in transverse series or 'resis­ survival phenomenon of laying resting 25. Foot shorter than pre-intestinal part tant' eggs, prior to, or during environmental 26. Toes 2 stressful conditions such as the evaporation of 27. Toes 3 their water habitat. This phenomenon is thought 28. Toes 4 to be an extension of their ability to reproduce 29. Spurs 1 parthenogenetically, which includes episodes of 30. Spurs 2 males appearing and joining with females to 31. Spurs long produce 'resting' eggs and a new generation of 32. Spurs short 33. Spurs modified or absent parthenogenetic females. 34. Cuticle coarse, leathery, bumps, spines, plates or There are too few rotifer taxonomic references knobs properly to support bdelloid rotifer identifi­ 35. Viviparous (able to see young forming inside cations for the non-specialist. Ongoing research mother) by interested ecologists, environmentalists and 36. Oviparous other biological workers has suffered from a 37. Commensal lack of suitable references to assist even the 38. Limnetic (free-swimming) (found in moss) simplest of bdelloid rotifer identifications. 39. Anhydrobiotic 40. Eyes generally absent Works such as Pennak (1978) offer a modicum 41. Eyes cervical or in neck, rarely absent of key elements of help, but the arrangement 42. Eyes in rostrum or rarely absent can sometimes be confusing, and some primary 43. T rophi aberrant elements of identification are not visible without 44. Sometimes in 'bottle' secretion shell the application of suitable and often difficult A simple generic key to the bdelloid rotifers 353 Koste & Shiel (1986) (for Australian fauna), Dorsey of Philadelphia, Pennsylvania, during although adequate for genera, can be misleading their cooperation in rotifer research. Modifica­ for species if used for other parts of the world, tions to the Myers key were made by Alben E. since rotifers in general are now being acknowl­ Turner Ill, in his unpublished aborted attempt edged to be endemic to a larger extent than to revise the Order. Noticeable elements of previously thought (Ricci, 1987). Relying on important characters are taken from Bryce experts for bdelloid rotifer identifications can (1910), Bartos (1951) and Donner (1965). also be difficult, since such people are all too few. The Tables are arranged into Generic Identifi­ cation Elements (Table 1), and the Numeric Key The Key of genera based on Table 1. A simplistic This paper is designed to be a non-specialist diagram
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