Freshwater Silica-Scaled Heterotrophic Protista: Heliozoa, Thaumatomonad Flagellates, Amoebae, and Bicosoecids, from the Lake Itasca Region, Minnesota
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JOURNAL OF THE MINNESOTA ACADEMY OF SCIENCE VOL. 78 NO. 2 (2015) FRESHWATER SILICA-SCALED HETEROTROPHIC PROTISTA: HELIOZOA, THAUMATOMONAD FLAGELLATES, AMOEBAE, AND BICOSOECIDS, FROM THE LAKE ITASCA REGION, MINNESOTA Daniel E. Wujek Central Michigan University, Mt. Pleasant, MI Forty-nine plankton samples were collected from the Lake Itasca Region, Minnesota over a period sporadically covering the summers of 1980, 1981 and 1987. A total of 22 freshwater heterotrophic siliceous-scaled species were observed: 18 heliozoa, two thaumatomonad flagellates, one bicosoecid, and one testate amoeba. Scale identifications were based on transmission electron microscopy. New records for North America include two heliozoans and one thaumatomonad flagellate. Five heliozoa taxa and one thaumatomonad flagellate are new records for the U.S. Wujek DE. Freshwater silica-scaled heterotrophic protista: heliozoa, thaumatomonad flagellates, amoebae, and bicosoecids, from the Lake Itasca Region, Minnesota. Minnesota Academy of Science Journal. 2015; 78(2):1-14. Keywords: bicosoecids, heliozoa, protista, testate amoeba, thaumatomonad flagellates Daniel E Wujek, Department of Biology, Central microscopy (EM) usually is necessary to distinguish Michigan University, Mt. Pleasant, MI 48859, e- sufficient morphology for species identification in the mail: [email protected]. scaled chrysophyte groups3 and now have become the This study was in part funded by a grant from the tool for other scaled protists. CMU FRCE committee. North American heterotrophic protist studies using INTRODUCTION electron microscopy in general have lagged behind The protist groups of golden brown algae those for Europe and Asia. The U.S. Great Lakes and (Chrysophyceae and Synurophyceae), heliozoa inland water’s heterotrophic freshwater protistan (Heliozoa, Filosea), thaumatomonads (Cercozoa, taxa, in particular the silica-scaled heliozoa and Thaumatomonadida), bicosoecids (Protozoa, scaled flagellates, are poorly documented using Bicosoecidida), and testate amoebae (Rhizopoda, electron microscopy (EM) with only a single species Himatismenida) are characterized by an exogenous being reported 4. On the contrary, more extensive siliceous or calcified envelope composed of scales Canadian studies by Nicholls and coworkers5-8 have and/or bristles or loricae. These organisms have long been reported. In contrast, the North American been recognized as important indicators of silica-scaled chrysophyte protist group environmental conditions. The silica-scaled (Chrysophyceae and Synurophyceae) have been chrysophytes are second in importance only after widely published3,9-15, including the Lake Itasca diatoms as indicators of the past ecology of lacustrine Region11-15. environments1,2. For example, the effects of acid rain were demonstrated by examining present and past Heliozoans are cosmopolitan, free-living aquatic sediments for scaled chrysophytes to demonstrate protozoans. Species are found in marine, brackish changes in the environment over time. and fresh waters. Some species have endosymbiotic algae, but most obtain their nutrition by phagocytosis, Identification of scale-bearing protists is based upon usually using characteristically long axopodia and the morphology of scales, which are often preserved short pseudopodia to capture their food. For recent in sediments1. Light microscopy-based reviews of the heliozoa see Febvre-Chevalier16, Lee identifications are of limited value, as electron et al.17, Page and Siemensma18 and Patterson and 1 JOURNAL OF THE MINNESOTA ACADEMY OF SCIENCE VOL. 78 NO. 2 (2015) Hedley19. An assessment of the centrohelid group Table 1. Lake Itasca Region sampling sites and and their worldwide distribution has been compiled dates containing heterotrophic silica-scaled by Mikrjukov20. protistans. Date In this study, the silica-scaled heterotrophic protists Sampling Site 1980 1981 1987 from samples collected during the summer months of Arco Lake x Beaver Pond, Hwy 4 x x 1977, 1980 and 1987 from 32 locations in the Lake Bohall Lake, north end x x Itasca Region, Minnesota, were observed using Bohall Pond, south end x transmission electron microscopy. All the sampling Chamber Creek x sites have a substantial and published silica-scaled Dahlberg Lake x phytoplankton record11-15. Darling Pond x x Deer Park Lake x MATERIALS AND METHODS Deming Pond, north x x x Forty-nine phytoplankton samples were collected Deming Pond, south x x x from 32 sampling sites with 20 µm mesh plankton Elk Creek x x x Hay Creek x x net from the Lake Itasca Region, Minnesota during Lasalle Creek x x the summer months of 1980, 1981 and 1987 (Table Lake Itasca, north arm x 1). Samples were preserved with acid Lugol’s in Lake Itasca, east arm x x plastic screw-cap vials. Samples for transmission Lake Itasca, west arm x x electron microscopy (TEM) were sub-sampled onto Long Lake x x Formvar-coated, carbon stabilized, 3 mm copper Mary Lake x Mississippi River culvert x x grids. After air drying, they were examined with Nicollet Lake x either a Philips EM300 or JEOL CM-10 TEM. All Pickerel Lake x identifications were based on TEM. The heliozoan Squaw Lake x 20 classification used is according to Mikrjukov . Twin Lake, east x Twin Lake, west x RESULTS Two Spot Trail, north x Twenty two heterotrophic freshwater silica-scaled pond #1 protists were observed during this investigation Two Spot Trail, north x (Table 2): 18 heliozoa taxa, two thaumatomonad pond #2 Two Spot Trail, south x flagellates, and one taxon each of a bicosoecid and pond #1 rhizopod amoeba (Plates 1-23). Of the eighteen Two Spot Trail, south x heliozoa observed, two are newly reported for North pond #2 America (Plates. 11, 19) and six for the U.S. (Plates. Upper Rice River x x 5, 7, 8, 15, 16, 17). The heliozoans observed all have Wild Rice River x x a cosmopolitan distribution20,21. Raphidocystis Wilderness Drive, south x tubifera was the most widely observed heliozoan in pond #1 Wilderness Drive, pond #2 x this study with the genus Acanthocystis having the greatest number of species (Table 2). Included in my using electron microscopy, Raineriophrys observations was the first heliozoan characterized erinaceoides (Plates 12-13), a species whose original description was as an Acanthocystis taxon from Denmark22. Thaumatomonad flagellates observed for the first time in North America was Gyromitus limax, and for the U.S., Thaumatomastix triangulata (Plate 21). All of taxa in the study are new records for Minnesota. 2 JOURNAL OF THE MINNESOTA ACADEMY OF SCIENCE VOL. 78 NO. 2 (2015) Table 2. Checklist of silica-scaled heliozoa, (Thomsen) Roijackers & Pickerel Lake thaumatomonad flagellates, amoeba, and Siemensma** bicosoecids from the Lake Itasca Region, Thaumatomonadida Minnesota, 1977, 1980, 1987. Thaumatomastix Dahlberg Lake Taxon Location triangulata (Balanov) Centrohelida Beech & Moestrup* Acanthocystis bicornis Deming Lake Gyromitus limax Belcher S. Deming Pond Dürrschmidt & Swale** Acanthocystis cornuta Nicolett Lake Himatismenida Dürrschmidt* Cochliopodium Hay Creek, Acanthocystis nichollsi Pickerel Lake bilimbosum Auerbach Pickerel Lake Siemensma & Roijackers Bicosoecida Acanthocystis penardi Hay Creek, Wild Cyathobodo crucifera Deming ponds Wailes* Rice River Swale & Belcher Acanthocystis polymorpha Deming Lake *New record for the United States Dürrschmidt* ponds, Wild **New record for North America (see Table 1 for Rice River dates) Acanthocystis turfacea Arco Lake Carter DISCUSSION Choanocystis aculeata Squaw Lake Heliozoa (Hertwig & Lesser) Siemensma & Roijackers Heliozoa are defines as predatory organisms, Pterocystis tropica Long Lake, Wild distinguished by their spherical body that is (Dürrschmidt) Rice River characterized with fine radiating cytoplasmic Siemensma** projections - the axopodia. These cytoplasmic Raineriophrys Hay Creek, E. projections participate in the capture of prey, erinaceoides (Petersen & Twin Lake, Two movement of cells, and adhesion to various Hansen) Mikrjukov Spot ponds, substrates. Long considered a natural taxonomic Wild Rice Lake group, recent molecular data indicate they are rather a Raphidiocystis tubifera Fish Hook Lake, polyphyletic assemblage of protists23,24. Penard Pickerel Lake, S. Deming Pond, Order Centroheliozoa (Centrohelids) Squaw Lake, Pickerel Lake Family Raphidiophryidae Raphidiophrys elegans E. Twin Lakes, Hertwig & Lesser Upper Rice Lake All taxa have external scales surrounded by a hollow Polyplacocystis marginata N. Deming Pond marginal rim. Spine scales, if present, are always (Siemensma) Mikrjukov symmetrical and either tubular, trumpet or funnel- Pompholyxophrys Dahlberg Lake, shaped. ovuligera Penard* Long Lake Pompholyxophrys punicea Wild Rice River Polyplacocystis marginata (Siemensma) Mikrjukov Archer* Figure 1, Plate 1. Pompholyxophrys stellata Deming Lake, Polyplacocystis marginata, originally described as Dürrschmidt and Hay Creek, 21 Nicholls* Pickerel Lake Raphidiophrys marginata Siemensma , it was later 25 Pinaciophora fluviatilis Lake Itasca, all transferred by Mikrjukov to his newly described Greeff arms genus Polyplacocystis. The genus contains five Rabdiophrys anulifera E. Twin Lake, species. Rainer emend. Siemensma Upper Rice Lake Rabdiophrys monopora Hay Creek, 3 JOURNAL OF THE MINNESOTA ACADEMY OF SCIENCE VOL. 78 NO. 2 (2015) Figure 1. Plates 1-12. Isolated surface scales. Plate 1. Polyplacocystis marginata, body scale. Plate 2. Raphidiophrys elegans, body scale. Plate 3. Raphidiocystis tubifera, plate scale. Plates 4-9. Acanthocystis. 4. A. bicornis,