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Halacaroidea (Acari): a Guide to Marine Genera

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Halacaroidea (Acari): a Guide to Marine Genera Org. Divers. Evol. 6, Electr. Suppl. 6: 1 - 104 (2006) © Gesellschaft für Biologische Systematik URL: http://www.senckenberg.de/odes/06-06.htm URN: urn:nbn:de:0028-odes0606-0 Halacaroidea (Acari): a guide to marine genera Ilse Bartsch Forschungsinstitut Senckenberg, Deutsches Zentrum für Marine Biodiversitätsforschung, Notkestr. 85, 22607 Hamburg, Germany Corresponding author, e-mail: [email protected] Received 13 December 2004 • Accepted 8 July 2005 Abstract Halacarid mites (Halacaroidea: Halacaridae) are meiobenthic organisms. The majority of species and genera are marine, only few are restricted to freshwater. Halacarid mites are present from the tidal area to the deep sea. It is the only mite family completely adapted to per- manent life in the sea. The first record was published more than 200 years ago. At present, 51 marine and brackish water genera of halacarid mites are known, including more than 1000 species. The genera are Acantho- halacarus, Acanthopalpus, Acarochelopodia, Acaromantis, Acarothrix, Actacarus, Agaue, Agauides, Agauopsis, Anomalohalacarus, Areni- halacarus, Arhodeoporus, Atelopsalis, Australacarus, Bathyhalacarus, Bradyagaue, Camactognathus, Caspihalacarus, Coloboceras, Co- lobocerasides, Copidognathides, Copidognathus, Corallihalacarus, Enterohalacarus, Halacarus, Halacarellus, Halacaroides, Halacaropsis, Halixodes, Isobactrus, Lohmannella, Metarhombognathus, Mictognathus, Parhalixodes, Pelacarus, Peregrinacarus, Phacacarus, Rhombo- gnathides, Rhombognathus, Scaptognathides, Scaptognathus, Simognathus, Spongihalacarus, Thalassacarus, Thalassarachna, Thalass- ophthirius, Tropihalacarus, Werthella, Werthelloides, Winlundia, and Xenohalacarus. The guide, which includes marine and brackish water genera, starts with an introduction to methods of collection, extraction and examination of halacarid mites, an outline of the external morphology and life history, and an overview of the commonly used terminology. Both a dichoto- mous key and tabular keys to the genera are presented. The keys have been prepared on the basis of adults. In general, in adults and nymphs the outline of idiosoma, gnathosoma and legs is similar, whereas the outline of plates, the sculpturing and number of setae on idiosoma and legs differ. In the tabular keys idiosoma, gnathosoma, palps, legs, tarsi and shape of claws are treated separately. The major part of the guide deals with descriptions of the 51 genera. Each genus is diagnosed and illustrated, namely a dorsal and ventral aspect of the idiosoma, the gnathosoma, leg I and tarsi I, II and IV. The diagnoses mention both, characters expected to be relevant in a phylogenetic sense and those thought to be mainly correlated with environment and mode of life. Rare character variants are included in the diagnoses; more variants are expected to be found in the future. In addition to the diagnoses, short notes are given on biology and geographi- cal distribution, on similar-looking genera, and distinguishing characters. At the end of the presentation of a genus, relevant and most recent descriptive or phylogenetic references are listed. Keywords: Halacaroidea; Acari, General introduction; Marine genera; Keys; Diagnoses Introduction The first record of a marine mite (‘insecta marina’) have been described, and the family Halacaridae is from coralline algae of a seashore was published given the rank of a superfamily Halacaroidea. more than 200 years ago (Baster 1758: pl. 10, fig. 7). Halacarid mites are meiobenthic organisms within a A century later, Gosse (1855) introduced the generic length range from 150 µm to about 2000 µm. Halacarids name Halacarus for marine mites, and Murray (1877) can be found at altitudes of 5000 m (freshwater established the family Halacaridae for mites ‘living halacarids in lakes and springs) as well as at depths habitually under the sea’. Since those days, more than of 7000 m (marine mites in deep-sea trenches). They 1000 species in 51 marine and 13 freshwater genera generally live in submerged habitats, but a few species Org. Divers. Evol. 6, Electr. Suppl. 6 (2006) Bartsch: Halacaroidea (Acari): a guide to marine genera 2 are adapted to survive in the splash zone. Halacarids are stirring with hot water is recommended. A minority present at all latitudes, from polar to tropical regions. of species has a hydrophobic cuticle; these mites are Halacarid mites live in a variety of substrata: within trapped at the water surface film when the sample is tufts of algae, on large fronds, in and on colonies of treated with air bubbles. Techniques using density sponges, hydrozoans, bryozoans, barnacles, mussels, gradient separation can also be used when extracting polychaetes, in interstices of coarse sediment or mites, with a flotation medium prepared from cooking flocculent ooze, amongst surface structures and gill salt, sugar or colloidal silica polymers. filaments of crustaceans and molluscs, and between The material held back in the fine-mesh sieve is spines and in the gut of echinoderms. Some few species placed in a petri dish or narrow-channel sorting dish and are known or suspected to be parasites. Halacarids scanned for halacarids under a dissecting microscope are rare or absent in silty sediments, oxygen-free (magnification: 12–25x). If possible, sorting should be habitats, and biotopes regularly defaunated due to done whilst the mites are alive, crawling around in the heavy pollution, destruction of the substratum, extreme dish and thus more conspicuous. Halacarids may be temperatures or desiccation. Several species of marine badly fouled with debris attached to idiosoma and legs. halacarids inhabit brackish waters, with rare records The mites can be cleaned somewhat by allowing them even from freshwater habitats. to run around in sand. In preserved sediment samples, The keys below deal with genera described from staining with rose bengal will facilitate sorting. marine and brackish water habitats. Genera widespread Ethanol (70%) should be used for fixation and stor- in fresh water (salinity less than 0.5 ‰) are not included. ing. Formalin hardens the mites and the necessary The drawings of the genera are generalized; rare clearing will be difficult. exceptions in external characters are excluded. Both For detailed examination, the mites have to be cleared a tabular and dichotomous key are given, as each has and mounted on slides. The halacarids are placed into a advantages and disadvantages. The references given at clearing medium, pepsin or lactic acid; gentle warming the end of each chapter or genus are only recent and (40–50 °C) may facilitate the clearing. The gnathosoma major ones. is torn off using a sharp needle, and body contents are squeezed out by applying gentle pressure with a blunt Collection, extraction and examination needle. Material stored in formalin for several years In intertidal and shallow subtidal zones, collecting by should be transferred into glycerine-acetic acid (with hand at low tide is recommended. Colonies of mussels, 2% acetic acid); after 1–3 months, the contents of the polychaetes, barnacles, bryozoans and hydrozoans are idiosoma can be dissolved with lactic acid. Animals scraped off; algal tufts, holdfasts and fronds, shells or with very delicate plates should not be left in acetic pebbles covered with unicellular algae, fine and coarse acid or lactic acid for more than a few hours. deposits from a beach surface as well as from deeper Permanent mounting media are glycerine jelly, glyc- layers, or rooted saltmarsh sediments are gathered; erine or Hyrax. Careful sealing of the coverslips with 50–500 ccm of the substratum may yield hundreds of a standard microscopical sealing agent, marine epoxy halacarid mites. From the sublittoral, collections can paint or nail varnish is necessary. Berlese, modified be made with trawl, dredge and grab. Halacarids inhab- Hoyer's fluid or polyvinyl lactophenol can be used iting the flocculent ooze are collected with epibenthic for temporary mounts but are not recommended for sleds. permanent slides. Specimens cleared in lactic acid The mites are extracted from the substratum by wash- have to be rinsed (in glycerine) to prevent subsequent ing. Large algal fronds, stones, decapods, or echinoids crystallization in the permanent mount. Over-cleared are washed in a bucket with a strong jet of water; specimen can be stained with chlor-azol black. To allow subsequently the washing water is poured through a a microscopic examination of both sides, mounting be- 60–100 µm sieve. Algal tufts and colonial organisms tween coverslips is recommended. The coverslips can are placed in a series of sieves, a 1–2 mm mesh sieve be stored in a metal slide frame or attached to a glass over a 60–100 µm sieve, and washed with a jet of water. slide with a drop of glycerine. Sediment samples are either stirred vigorously in water, References. Newell (1947, 1984), Green and and the supernatant water decanted through a 60–100 MacQuitty (1987), Bartsch (1988a, 1993a). µm mesh sieve, or placed in a separatory funnel where lighter particles (such as halacarids) are elutriated by External morphology (Figs 1A, B, 2A–F, 5E) a water stream and retained in a sieve at the outflow. The body of an adult halacarid is divided into idiosoma, As intertidally living species are resistent to extreme with four pairs of legs, and gnathosoma, with palps and conditions, shock techniques with fresh water, ice water chelicerae. or anaesthetizing media may prove to be inefficient. The idiosoma is typically covered with sclerotized For sediment samples
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