Identification Key to the Genera of Marine Rotifers Worldwide
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75 - J - + J Meiofauna Marina, Vol. 16, pp. '75-99,109 figs., March 2008 © 2008 by Verlag Dr. Friedrich PfellpMühírterVVlJbrmany - ISSN 1611-755 R I T ,NST,TUUT V °°R DEZE' ^A N D ER S MARINE INSTITUTO Oostende - Belgium Identification key to the genera of marine rotifers worldwide Diego Fontaneto*'**, Willem H. De Smet*** and Giulio Melone** A bstract A dichotomous key to rotifers is presented for the 28 Families and 66 Genera that have been reported from saline systems of both marine and inland waters. Information is provided on general identification and papers dealing more particularly with certain Families and Genera. A succinct overview of the species found in saline habitats is given for each genus. Keywords: Rotifera, saltwater, brackish water, dichotomous key Introduction ments are a single m edian fulcrum, and 3 paired elements: rami, unci, and manubria (Fig. 3). The Rotifers (phylum Rotifera) are transparent micro fulcrum (absent in bdelloids) is mostly rectangular scopic eutelic m etazoans (50-2000 pm), with three in lateral view, but may be variable in shape. The main body regions: head, trunk, and foot (Figs. rami are hollow, roughly triangular structures. 1-2). The head is characterized by the presence Their inner margin is provided with numerous of a typical anterior ciliated field named corona, e lo n g a te e le m e n ts, th e ra m i sclero p ili, w h ic h m ay with different ciliated areas located anteriorly be fused in a ridge and/or a series of tooth-like and around the mouth. The body wall may be projections. Commonly in monogononts, the rami thickened in a lorica, and may bear variable are provided with lateral projections, the alulae. appendages. The foot usually ends with mov The unci are plates formed by connection of a vari able toe-like extensions named spurs and toes; able num ber of teeth into one rigid structure. The some species lack toes and possess one or more tee th are m o stly u n e q u a l. M in u te su b u n c in al teeth adhesive disks. While many rotifers may lack are mostly present, and are situated under the unci the above features, all of them possess a special plate. The manubria are more-or-less triangular ized masticatory organ, the muscular pharynx or to crescent-shaped, or rod-shaped supports of mastax, containing a set of hard sclerotised jaws the unci. A variable num ber of diverse accessory or trophi, the shape of 'which is very im portant in sclerites o c c u rs in m o n o g o n o n ts . N in e m a in ty p es ro tife r ta x o n o m y (W alla ce et al. 2006). T ro p h i e le of trophi are recognized (malleate, malleoramate, * Imperial College London, Division of Biology, Silwood Park Campus, Ascot Berkshire, SL5 7PY, United Kingdom ** Université degii Studi di Milano, Dipartimento di Biologia, via Celoria 26,1-20113 Milano *** University of Antwerp, Department of Biology, Campus Drie Eiken, Universiteitsplein I, B-2610 Wilrijk, Belgium Corresponding author: Diego Fontaneto; e-mail:[email protected] Meiofauna Marina, Vol. 16 7 6 CO 3) Virgate (Fig. 7-9): fulcrum long, rami recurved dorsally, unci with few teeth or reduced, manubria mostly with elongate shafts; often asymmetrical (e.g., Trichocerca). T h e v irg a te trophi type is the most variable of all. Used for piercing and sucking, or swallowing food by pumping without crushing. 4) Uncinate (Fig. 10): similar to the malleoramate type, but all trophi elements except the unci strongly reduced. Unci teeth few (2-5 teeth ex per uncus), elongate and curved, forming s u p p o rtin g ro d s fo r th e m astax ; n o t im p o rta n t, the food is already predigested. Only in Col lothecidae. 5) Incudate (Fig. 11): fulcrum short, rami elon gate, pincer-shaped. Manubria and unci strongi}' reduced. Specialized for seizing. Only in Asplanchnidae. 6) Cardate (Fig. 12): fulcrum short, rami lvriform, manubria with a shaft and a characteristic, additional ventral projection. Species-specific accessory trophi elements are present, and often numerous. The cardate type is adapted for pum ping. Restricted to Lindiidae. Figs. 1, 2. Scheme of the anatomy of a generic rotifer 7) Forcipate (Figs. 13-19): fulcrum mostly short; in dorsal (1) and lateral (2) view. H, head; T, trunk; rami elongate long pincers, usually with sharp F, foot; b, brain; bl, bladder; cl, cloaca; co, corona; tips and often armed with teeth along medial da, dorsal antenna; ex, excretory apparatus; gg, gastric margin. Unci are strong, but with a single or glands; gv, germo-vitellarium; i, intestine; m, mastax few teeth only. Manubria are long and thin; with trophi; n, nephridium; pg, pedal glands; ra, retro- cerebral apparatus; sg, salivary glands; t, toe. Modified often with intramalleus between uncus and from W allace & Ricci (2002). manubrium. The trophi work by gripping (rami and unci can be extruded in Dicrano phoridae). In Dicranophoridae and Ituridae. uncinate, incudate, virgate, cardate, forcipate, 8) Ramate (Figs. 20, 21): fulcrum absent, rami ramate, and fulcrate), characterized by the shape semicircular, flat; unci teeth numerous, oc a n d size of th e e le m en ts, th e p re sen c e of accesso ry casionally resembling striations; manubria as parts, and the way they operate (Figs. 3-22). lateral bands. Grinding. In Subclass Bdelloi 1) M alleate (Fig. 3-5): fulcrum short, rami more d ea. or less triangular and flat. Unci with several 9) Fulcrate (Fig. 22): fulcrum long, manubria (4-12) firmly connected teeth. The manubria absent; the pum ping action is performed by are provided with a shaft, which is typically the hypopharynx muscle. In Seison. fairly short, but this structure is more elongate For a recent, comprehensive introduction to the in the submalleate trophi of Lecanidae and m o rp h o lo g y , b io lo g y a n d eco lo g y of th e tax o n see some Proalidae. The malleate trophi are W allace & Ricei (2002) and W allace et al. (2006). adapted for gripping, grinding and pump- ing. 2) Malleoramate (Fig. 6): fulcrum short, rami Classification and systematics more-or-less triangular, flat; similar to mal le a te ty p e, b u t u n c i tee th m o re n u m e ro u s , -with Classically, three groups are recognized within weaker connections, occasionally resembling Phylum Rotifera: Seisonacea, Bdelloidea and striated plate; manubria crescent-shaped, M onogononta, while molecular studies indicate without shaft. Only grinding. that the exclusively endoparasitic Acanthocephala actually are rotifers (Mark Welch 2000, Sorensen Fontaneto et al.: Identification key to marine rotifers 7 7 Figs. 3-22. SEM pictures of the nine trophi types. 3-5. malleate: 3, Brachionus plicatilis; 4, Proales similis; 5, Colurel la colurus; 6, m alleoram ate, Testudinella clypeata. 7-9. virgate: 7, Trichocerca pediculus; 8, Cephalodella sp.; 9, Pleuro trocha atlantica. 10, uncinate,Cupelopagis vorax (freshwater species); 11, incudate,Asplanchna priodonta; 12, cardate, Lindia tecusa; 13-19, forcipate: 13,Aspelta europaea; 14, Dicranophoroides caudatus; 15, Encentrum algente; 16, Dicrano phorus forcipatus; 17, Erignatha clastopis; 18, Myersinella uncodonta; 19,Itura myersi. 20-21. ramate: 20, Zelinkiella synaptae; 21, Philodinavus paradoxus. 22, fulcrate, Seison annulatus. Scale bars = 10 pm; a, alula; b, basal chamber; f, fulcrum; i, intramalleus; m, manubrium; r, ramus; s, subbasal chamber; u, uncus. Meiofauna Marina, Vol. 16 78 & Giribet 2006). Nevertheless, only the three different scenario, with areas of high numbers classical groups will be considered in this key. of endemic taxa. The rotifer record is highest in The most recent and comprehensive checklist of the Northern hemisphere, but this may be due all rotifer species, excluding acanthocephalans, to the higher num ber of studies in those regions. is by Segers (2007). Diversity hotspots are northeast North America, Seisonacea, w ith oni}’ three species, are ex tropical South America, Southeast Asia, Australia, clusively m arine and live epizootically on lepto- and Lake Baikal. Endemicity is low in Africa stracan crustacea. Males and females are usually (including Madagascar), Europe, the Indian sub present at a 1:1 sex ratio, and reproduction is by continent and Antarctica (Segers 2008). Data for obligate amphim ixis (Ricci et al. 1993). m arine rotifers are still too scarce to allow sound M onogononta contains 1570 species-level taxa, conclusions about their biogeography (Fontaneto and the vast majority of them (1488) are free-living et al. 2006a). fresh or inland water taxa (Segers 2007, 2008). M onogononts occur in all types of water bodies worldwide; they are particularly abundant and di Collection and specimen handling verse in lentic freshwater habitats, although many and preparation species also occur in haline habitats (Fontaneto et al. 2006a). This group is characterized by a single Rotifers may be found in the plankton and peri gonad and cyclic parthenogenetic reproduction:phyton, and as interstitial fauna or psammon. parthenogenesis dominates, but sexual reproduc Planktonic and littoral rotifers are collected by tion also occurs occasionally. Eggs produced by dragging a plankton net (mesh size: 25-50 pm) sexual reproduction are known as resting eggs through the open water, submerged vegetation, undergoing obligatory diapause. They are able littoral macrophytes or algae. Rotifers inhabit to survive desiccation and other adverse envi ing periphyton may be collected using a flexible ronmental conditions for long periods, and oftencollecting tube attached to a large syringe.