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Kevin Anderson1,7, Mark Fate2,7, Ching-Han (Eric) Hsieh3,7, Lorenz Kim4,7 Katy Lazarus3,7, J. Patrick Kociolek*5,7 and Rex L. Lowe6,7 1 University of Northern Carolina at Chapel Hill, USA 2 Program in the Environment, University of Michigan, Ann Arbor, MI, USA 3 Dept. of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, USA, 4 College of Literature, Science and the Arts, University of Michigan, Ann Arbor, MI, USA, 5 Museum of Natural History and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO, USA 6 Department of Biology, Bowling Green State University, Bowling Green, OH, USA 7 University of Michigan Biological Station, Pellston, MI, USA


The Lake of the Clouds, located inside the Porcupine Mountains and two miles away from Lake Superior, is a famous vista site within the Porcupine Mountains Wilderness State Park, Michigan. Even though several fishery surveys of the lake were released by the Michigan Department of Nat- ural Resources in the 1940s, no known algal studies have been conducted. The Porcupine Mountains Wilderness State Park has experienced a relatively low level of anthropogenic disturbance compared to other regions of the Great Lakes, which makes it an appealing site for algal studies. This paper re- ports the exploratory research done by professors and students at the University of Michigan Bio- logical Station on the algal diversity of the Lake of the Clouds. Water chemistry shows this lake to be slightly acidic and with low nutrients and low conductivity. In this study, fresh samples of algae and cleaned diatom samples were investigated. From the living algal samples, 7 algal divisions were observed. The inventory showed that, based on the number of genera recorded, the Chlorophyta is more diverse than other algal divisions. A total of 78 genera (excluding diatoms) and 29 species of algae were identified from the living samples. Nearly 150 species in 58 genera of diatoms (Bacillar- iophyta) were also recorded from the cleaned samples. Among the diatoms, three species new to sci- ence were identified and are described here. These are Brachysira ontonageniana Kociolek & Lowe, B. gatesii Kociolek & Lowe, and Gomphonema porcupiniana Kociolek & Lowe. This rich list of algal taxa is only the first step in exploring the algal diversity, microhabitats, and ecosystem of the Lake of the Clouds. The diversity of algal taxa found in this study suggests that further algal re- search on the Lake of the Clouds is warranted.

K E Y W OR D S : A l g a l d iv e r s i t y, L a k e o f t h e C l o u d s , n e w d i a t o m s p e c i e s , B r a c h s i r a , Gomphonema


Although there are large-scale surveys of the algal flora of Michigan (e.g., Prescott 1962), and individual groups of algae have been documented from spe- cific sites or habitats (Ackley 1929, 1932), there are few surveys of the entire algal flora from individual sites. In particular, we know of no previous work on

*Author for correspondence (


the algal flora of lotic (flowing water) or lentic (still water) ecosystems in the Porcupine Mountains, which is located in the northwestern part of the Michi- gan’s Upper Peninsula.

The Porcupine Mountains contain one of North America’s largest stands of northern hardwoods, stretching over 31,000 acres of old growth forest (Davis 1993). The Porcupine Mountain range is on the southern edge of the Canadian Shield, a vast region of common geologic origin (Larson and Schaetzl 2001; Der- ouin et al. 2007). The bedrock in and around the Porcupine Mountains is primar- ily Precambrian, making it over 3.5 billion years old (Ojakangas and Matsch 1982).

Porcupine Mountains Wilderness State Park was established in 1945 and has experienced relatively little human disturbance. The first European contact re- sulted from the fur trade in the 1700-1800s, although this attracted settlers in only small numbers compared to subsequent waves of miners. Between 1845 and 1910, 45 copper mines operated on present day parkland. Most mines went bank- rupt or closed before the mid 1900s. The area is still extremely rich in copper ore, in part because the copper particles are particularly fine and difficult to mine. White pines along the shore of Lake Superior were targeted by loggers for their ease of harvest and drew contact to the area in the early 1900s (Davis 1993).

The Lake of the Clouds, one of the four lakes in the Porcupine Mountains Wilderness State Park, is located in Section 22 of Township 51 North, Range 43 West in Ontonagon County of Michigan’s Upper Peninsula (Taube 1949) and is a part of the Carp River Valley. The lake, which is approximately 330 meters above sea level and is overlooked by an escarpment rising an additional 120 me- ters overhead (Taube 1949), has a surface area of 133 acres and a maximum depth of 4 meters (Moffett 1940). Lake of the Clouds drains into Lake Superior to the north. The lake has sufficient levels of dissolved oxygen to support fish life at all depths, a circumneutral acidity, and a soft water composition at ap- proximately 25 ppm dissolved salts (Moffett 1940). Around two dozen species of vascular aquatic plants occupy the littoral zone, of which three-way sedge (Dulichium arundinaceum), spike rush (Eleocharis sp.), yellow water lily (Nuphar sp.), and a pond weed (Potamogeton angustifolius) are the most preva- lent macroflora species. Perch, smallmouth bass, and brook trout are the most common species of fish in the lake (Moffett 1940).

No previous effort to catalog the algal diversity of Lake of the Clouds is known. Due to the lake’s unknown algal flora and its relatively short history of human interaction, Lake of the Clouds presents an ideal environment for con- ducting a survey of algal diversity.


On July 9, 2012, a survey of the algal diversity in the Lake of the Clouds was conducted. Five different microhabitats were sampled to include epiphytic, epipelic, epilithic, deep benthic, and planktonic algae.

Collection Sites

Five collection sites were chosen for this study. Four of them were located nearly equidistantly across an approximately1290 meter transect along the southern shore of Lake of the Clouds. Sites 1-


4 were located at ca. 300 m, 600 m, 900 m and 1290 m from the outflow. Site 1 was a sheltered cove with emergent aquatic vegetation; Site 2 was an exposed site with obvious benthic algal production; Site 3 had a gravel bottom with epilithic algae evident; and Site 4 was at a small spit at which evi- dent wave action was present. The fifth site was located at the bridge over the outflow of the lake (Carp River Outflow), and thus had flowing water. The five sites were chosen to include a diversity in vegetative density, amount of current flow, sunlight intensity, substrate availability, and similar factors. Spacing out the collection sites was necessary for gaining a fair representation of the entire lake.

Sampling Processes Algae

Epiphytic algae were collected by squeezing liquid and associated algae out of aquatic plants into 100 mL Whirlpaks®. A small sample of each plant was also collected. Epipelic algae were sampled from the upper few millimeters of the sediment with a turkey baster. Planktonic algae were collected with a size 25 plankton net with 64-.m mesh openings.

Each sample of epilithic algae (algae attached to rock), epidendric algae (algae attached to wood) and deep benthic algae (algae living in the sediment surface of the lake) was collected from sites 2 - 3 meters below lake surface. For sampling the epilithic algae and epidendric algae, a pocketknife was used for scraping algal specimens from the substrate. A snorkel, diving goggles, and a turkey baster were used for diving into the lake and collecting all samples from these three algal habitats. Each sample was placed into a 100 mL resealable Whirlpaks. A total of 25 samples were brought back to the lab on ice for analysis and stored in a refrigerator to slow down the biological processes of the organisms and prevent their degradation.

Sampling Process for Water Temperature, pH and Water Chemistry

At each site, temperature and pH were measured and recorded. A Fisher Scientific Accumet® portable meter was used to measure both pH and the temperature. A water sample was taken in a 1 L plastic Nalgene bottle, kept on ice, and returned to the University of Michigan Biological Station for analysis by standard methods with an autoanalyzer. The following environmental variables were measured: Total phosphorus, total nitrogen, silica, chloride, conductivity, alkalinity, dissolved or- ganic carbon and turbidity.

Samples Analysis—Living Algae

Samples were examined microscopically using microscope slides, pipettes, cover glasses, Amer- ican Optical Spencer light microscopes and an Olympus BX-51 light microscope. Genera and species were documented by site number and habitat, and photomicrographs were taken to document taxa. Soft algae (non-diatom) were identified at 400X magnification. Identification of the algal taxa was based on Canter-Lund and Lund 1995; Croasdale et al. 1983; Dillard 1989a, 1989b, 1990, 1991a, 1991b, 1993, 1999; Komarek and Anagnostidis 1999, 2005 Prescott, 1962, 1978; Prescott et al. 1972, 1975, 1977, 1981, 1982; Wehr & Sheath 2003; West and West 1904, 1905, 1908, 1912; and West et al. 1912.

Identified genera and species were recorded on a master list. Photomicrographs of specimens were taken with the cellSens® entry program and the Olympus microscope. Approximately 5 mL of several rich samples were preserved with glutaraldehyde so that the soft algae could be further ana- lyzed over time.

Additional subsamples were allocated for cleaning of diatoms.

Sample Analysis—Cleaned Diatoms

Subsamples of the living algae were cleaned by the method of van der Werff (1955). After clean- ing, the diatom valves were mounted in Naphrax®, and specimens were observed using an Olympus BX51 light microscope. Diatoms were viewed at 1000x, and light micrographs of the specimens were captured with an Olympus DP71 camera. Cleaned specimens were mounted onto coverslips, coated with gold-palladium, and prepared for observations with scanning electron microscopy (SEM). Electron micrographs were obtained by using a JEOL 7401F field emission scanning elec- tron microscope and the microscope JEOL software. Stria densities were measured at the valve mar-


gin at the valve center across a full length of 10 microns. Cleaned diatoms and permanent slides have been deposited in the Kociolek Collection at the University of Colorado Herbarium (COLO).


Living Algae Samples

Samples containing living algae from Lake of the Clouds contained a wide variety of taxa. A total of 7 divisions and 88 genera from the 5 collection sites were identified. The division with the most genera was Chlorophyta, with 48 genera. Eighteen genera of Cyanophyta and 11 genera of Bacillariophyta were identified prior to cleaning. In addition, 4 genera of Euglenophyta, 4 genera of Pyrrophyta, 3 genera of Chrysophyta and 2 genera of Xanthophyta were identi- fied. The complete list of taxa from these samples is presented in Table 1.

Four genera, Microcystis, Staurastrum, Mougeotia, and Desmidium, were found in all five collection sites. Of these, Microcystis and Mougeotia were found in several different microhabitats among the sites. Eleven genera were found in 4 of the 5 collection sites; these included the diatoms Fragilaria and Tabellaria, the chlorophytes Bulbochaete, Closterium, Cosmarium, Hyalotheca, Pediastrum, Pleurotaeniu and, Zygnema, and the cyanophytes Gloeotrichia, and Oscillatoria. Of those found in four sites, Fragilaria, Tabellaria, Closterium, Cosmarium, Hyalotheca, Pediastrum, and Zygnema were found in several dif- ferent microhabitats among the sites.


Community richness was also analyzed as a function of the various algal mi- crohabitats—epiphytic, epipelic, epilithic, epidendric, deep benthic, metaphytic and planktonic. The largest number of genera, 68, was found in epiphytic micro- habitats. In addition, 38 genera were found in metaphytic microhabitats, 22 gen- era in deep benthic microhabitats, 21 genera in epipelic microhabitats, 18 genera in planktonic microhabitats, and 15 genera in epidendric habitats. The genera found in epilithic microhabitats were mainly diatoms.

Cleaned Diatoms

A total of 147 diatom taxa from 58 different genera were identified from the cleaned material. Three species new to science are included in this list. Formal descriptions are presented later in this article. Genera with the most taxa repre- sented in this survey included Navicula (10), Gomphonema (9), Eunotia (9), Neidium (8), and Stauroneis (8). Several monoraphid genera were also well rep- resented (e.g., Psammothidium, Planothidium). Although there were some planktonic species present in our samples (e.g., Fragilaria crotonensis, Stephan- odiscus superiorensis), the vast majority of the species were from benthic mi- crohabitats. The cumulative list of diatoms encountered from the 5 sites is pre- sented in Table 2.


TABLE 1. List of algal taxa from live samples collected from Lake of the Clouds. For each taxon, the habitat or habitats in which it was collected at each site is indicated by the following letters: E=epiphyton, M-metaphyton, PL=plankton, B=benthic, PE=epipelon and D-epidendron.

Taxa Sites 1 2 3 4

CHLOROPHYTA Chaetophorales Aphanochaete sp. Chaetophora sp. Chlamydomonales Eudorina sp. Chlorellales Zoochlorella sp. Chlorococcales Characium sp. Coelastrum sp. Dictyosphaerium sp. Dictyosphaerium pulchellum Wood Nephrocytium sp. Radiofilum flavescens G.S. West Coleochaetales Chaetosphaeridium sp. Coleochaete sp. Desmidiales Bambusina sp. Closterium sp. Closterium kutzingii Brébisson Cosmarium sp. Cosmocladium sp. Desmidium sp. Desmidium grevillii (Kützing) De Bary Euastrum sp. Euastrum dubium Nägeli Gonatozygon sp. Hyalotheca sp. Hyalotheca mucosa (Dillwyn) Ehrenberg Micrasterias sp. Micrasterias laticeps Nordstedt Micrasterias radiata Hassall Onychonema sp. Pleurotaenium sp. Pleurotaenium ehrenbergii (Brébisson) De Bary Pleurotaenium trabecula (Ehrenberg) Nägeli Sphaerozosma sp. Sphaerozosma vertebratum (Brébisson) Ralfs Spondylosium sp. Spondylosium aubertianum var. archerii (Gutwinski) W. & G.S. West Staurastrum sp. Triploceras sp. Xanthidium sp. Xanthidium brebissonii Ralfs M M E E, M E E E E E, M E E E, M M E, PE E, M E E E E E, M E E E E E E, M E, M E, M, PL PE E, M M E B PL B PL PL E B B E E B B B B E, PE E, B E E, PL E, D E, D E E, D E, M, PL E, M, PL E E, D E, D E, M, PL E, M, PL PE PE E, PE E, PE PE E E E E E, PE E E PE E, PE E


TABLE 1. (Continued).

Taxa Sites 1 2 3 4

CHLOROPHYTA (Continued) Glaucocystales Glaucocystis nostochinearum (Itzigsohn) Rabenhorst Microsporales Microspora sp. Oedogoniales Bulbochaete sp. Oedogonium sp. Sphaeropleales Ankistrodesmus sp. Gloeocystis sp. Kirchneriella lunaris (Kirchner) Möbius Pediastrum sp. Pediastrum boryanum (Turpin) Meneghini Pediastrum duplex Meyen Quadrigula sp. Scenedesmus sp. Scenedesmus quadricauda (Turpin) Brébisson in Brébisson et Godey Sorastrum americanum (Bohlin) Schmidle Tetraedron sp. Tetraedron minimumm (A. Braun) Hansgirg Ulotrichales Ulothrix sp. Ulothrix variabilis Kützing Volvocales Pandorina sp. Volvox sp. Zygnematales Cylindrocystis sp. Mougeotia sp. Netrium sp. Spirogyra sp. Zygnema sp. CYANOPHYTA Chroococales Aphanothece sp. Chroococcus sp. Dactylococcopsis fascicularis Lemmermann Microcystis sp. Nostocales Amphithrix sp. Anabaena sp. Dolichospermum spiroides (Kleb.) Wacklin, L.Hoffm. & Komárek Calothrix sp. E, M M E E, M E E, PL E, M E E, M, PE E E E E E, M E E, M, PL E E, M E,M E, M E, M E E, M, PL E, M E E E M B B PL PL E, B B B B, PL B, PL B E E, PL E, B PL E, D E, D B, PL E, D D, PE, PL E, D E, D PL M, PL E, M, PL E E E E PE M, PL E PE E, PE E, PE E E PE PE PE E


TABLE 1. (Continued).

Taxa Sites 1 2 3 4

CYANOPHYTA (Continued) Hapalosiphon sp. Nostoc sp. Stigonema sp. Tolypothrix sp. Oscillatoriales Lyngbya sp. Oscillatoria sp. Pseudanabaenales Spirulina sp. Synechococcales Aphanocapsa Coelosphaerium sp. Coelosphaerium kutzingianum Nägeli Merismopedia sp. E E E E E M E, M E E E, M E B E B E E E, B E E CHRYSOPHYTA Chromulinales Chrysosphaerella sp. Dinobryon sp. Hibberdiales Derepyxis sp. M M E PL EUGLENOPHYTA Euglenales Euglena sp. Euglena acus Ehrenberg Lepocinclis acuta Prescott in Prescott et al. Phacus sp. Trachelomonas sp. PYRROPHYTA Gonyaulacales Ceratium sp. Ceratium hirudinella (O.F. Müller) Dujardin Peridiniales Peridinium sp. Phytodinales Cystodinium sp. Stylodinium sp. XANTHOPHYTA Mischococcales Ophiocytium sp. Ophiocytium bicuspidatum (Borge) Lemmermann Trebouxiales Botryococcus braunii Kützing E, PE, PL E E E E E E, M E E PE PL B, PL B, PL E PL E E E PE


TABLE 2. Inventory of the Diatom Taxa Identified from Lake of the Clouds, All Sites Combined.

Achnanthidium exiguum (Grunow) Czarnecki Achnanthidium exiguum var. heterovalvum (Krasske) Czarnecki Achnanthidium minutissimum (Kützing) Czarnecki Amphipleura pellucida (Kützing) Kützing Amphora pediculus (Kützing) Grunow Asterionella formosa Hassall Aulacoseira italica (Ehrenberg) Simonsen Brachysira gatesii Kociolek et Lowe, sp. nov. Brachysira ontonageniana Kociolek et Lowe, sp. nov. Brachysira serians (Brébisson) Round et Mann Brachysira vitrea (Grunow) Ross Brachysira zellensis (Grunow) Round & Mann Caloneis bacillum (Grunow) Cleve Caloneis lewisii Patrick Caloneis silicula (Ehrenberg) Cleve Caloneis tenuis (Gregory) Krammer Cavinula cocconeiformis (Gregory ex Greville) Mann et Stickle Cavinula jaernefeltii (Hustedt) Mann et Stickle Cavinula pseudoscutiformis (Hustedt) Mann et Stickle Chamaepinnularia mediocris (Krasske) Lange-Bertalot Cocconeis placentula var. euglypta (Ehrenberg) Grunow Cocconeis pseudothumensis Reichardt Craticula cuspidata (Kützing) Mann Cyclotella comensis Grunow Cymbella aspera (Ehrenberg) Cleve Cymbella ehrenbergii Kützing Cymbella lanceolata (Agardh) Agardh Cymbopleura hybrida (Grunow) Krammer Decussata placenta (Ehrenberg) Lange-Bertalot Diploneis elliptica (Kützing) Cleve Diploneis marginestriata Hustedt Discostella stelligera (Cleve et Grunow) Houk et Klee Encyonema elginense (Krammer) Mann Encyonema hebridicum Grunow ex Cleve Encyonema lunatum (Smith) Van Heurck Encyonema minutum (Hilse) Mann Encyonopsis microcephala (Grunow) Krammer Entomoneis alata (Ehrenberg) Ehrenberg Eolimna minima (Grunow) Lange-Bertalot Eucocconeis alpestris (Brun) Lange-Bertalot Eunotia didyma Grunow in Moller Eunotia formica Ehrenberg Eunotia implicata Nörpel, Alles et Lange-Bertalot Eunotia incisa Smith ex Gregory Eunotia monodon Ehrenberg Eunotia naegelii Migula Eunotia pectinalis var. undulata (Ralfs) Rabenhorst Eunotia praerupta Ehrenberg Eunotia tetraodon Ehrenberg Fragilaria crotonensis Kitton Fragilaria vaucheriae (Kützing) Petersen Fragilariforma constricta (Ehrenberg) Williams et Round Frustulia krammeri Lange-Bertalot et Metzeltin Frustulia saxonica Rabenhorst Frustulia vulgaris (Thwaites) De Toni Frustulia weinholdii Hustedt


TABLE 2. Continued.

Geissleria paludosa (Hustedt) Lange-Bertalot et Metzeltin Gomphonema acuminatum Ehrenberg Gomphonema affine Kützing Gomphonema augur Ehrenberg Gomphonema brebissonii Kützing Gomphonema intricatum Kützing Gomphonema parvulum (Kützing) Kützing Gomphonema patricki Kociolek et Stoermer Gomphonema porcupiniana Kociolek et Lowe, sp. nov. Gomphonema truncatum Ehrenberg Gomphosphenia grovei (M. Schmidt) Lange-Bertalot Gyrosigma sp. Handmannia radiosa (Grunow) Kociolek & Khursevich Hantzschia amphioxys (Ehrenberg) Grunow Hippodonta capitata (Ehrenberg) Lange-Bertalot, Metzeltin et Witkowski Karayevia clevei (Grunow) Bukhtiyarova Karayevia laterostrata (Hustedt) Bukhtiyarova Karayevia suchlandtii (Hustedt) Bukhtiyarova Meridion circulare (Greville) Agardh Navicula aboensis (Cleve) Cleve-Euler Navicula angusta Grunow Navicula constans Hustedt Navicula laterostrata Hustedt Navicula pseudoventralis Hustedt Navicula radiosa Kützing Navicula rhynchocephala Kützing Navicula schadei Krasske Navicula tridentula Krasske Navicula viridula (Kützing) Kützing Neidiopsis levanderi (Hustedt) Lange-Bertalot et Metzeltin Neidium affine (Ehrenberg) Pfitzer Neidium affine var. humerus Reimer Neidium amphigomphus (Ehrenberg) Pfitzer Neidium apiculatum Reimer Neidium bisulcatum (Lagerstedt) Cleve Neidium firma (Kützing) Pfitzer Neidium hitchcockii (Ehrenberg) Cleve Neidium productum (Smith) Cleve Nitzschia dissipata (Kützing) Grunow Nitzschia recta Hantzsch ex Rabenhorst Nupela impexiformis (Lange-Bertalot) Lange-Bertalot Nupela sp. Opephora olsenii Møller Pinnularia biceps Gregory Pinnularia microstauron (Ehrenberg) Cleve Pinnularia nodosa (Ehrenberg) Smith Pinnularia viridis (Nitzsch) Ehrenberg Placoneis clementis (Grunow) Cox Placoneis gastrum (Ehrenberg) Mereschkowsky Planothidium apiculatum (Patrick) Lange-Bertalot Planothidium dubium (Grunow) Round et Bukhtiyarova Planothidium joursacense (Héribaud) Lange-Bertalot Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot Planothidium peragalli (Brun et Héribaud) Round et Bukhtiyarova Planothidium pseudotanense (Cleve-Euler) Lange-Bertalot Psammothidium bioretii (Germain) Bukhtiyarova et Round


TABLE 2. Continued.

Psammothidium chlidanos (Hohn et Hellerman) Lange-Bertalot Psammothidium didymum (Hustedt) Bukhtiyarova et Round Psammothidium rossii (Hustedt) Bukhtiyarova et Round Psammothidium subatomoides (Hustedt) Bukhtiyarova et Round Psammothidium ventralis (Krasske) Bukhtiyarova et Round Pseudostaurosira brevistriata (Grunow) Williams et Round Pseudostaurosira elliptica (Schumann) Edlund, Morales et Spaulding Reimeria sinuata (Gregory) Kociolek et Stoermer Rhopalodia gibba (Ehrenberg) Müller Rossithidium pusillum (Grunow) Round et Bukhtiyarova Sellaphora americana (Ehrenberg) Mann Sellaphora laevissima (Kützing) Mann Sellaphora pupula (Kützing) Meresckowsky Stauroneis anceps fo. gracilis Rabenhorst Stauroneis anceps var. americana Reimer Stauroneis kriegeri Patrick Stauroneis livingstonii Reimer Stauroneis phoenicenteron (Nitzsch) Ehrenberg Stauroneis smithii Grunow Stauroneis stodderi Greenleaf Staurosira construens Ehrenberg Staurosirella pinnata (Ehrenberg) Williams et Round Stenopterobia anceps (Lewis) Brébisson ex Van Heurck Stenopterobia curvula (Smith) Krammer Stenopterobia delicatissima (Lewis) Van Heurck Stephanodiscus superiorensis Stoermer & Theriot in Theriot & Stoermer Surirella angusta Kützing Surirella brebissonii Krammer et Lange-Bertalot Surirella minuta Brébisson Surirella tenera var. nervosa Schmidt Tabellaria fenestrata (Lyngbye) Kützing Tabellaria flocculosa (Roth) Kützing Tryblionella victoriae Grunow Ulnaria delicatissima (W. Smith) Aboal et Silva

Water Chemistry

At the time of collection, the water temperature was 23°C, and the pH ranged from 6.92 to 7.70. Nutrients were relatively low, with total P measured at 8.7 .g/L and total N at 0.157 mg/L. Soluble SiO2 was 2 mg/L and chloride was 0.3 mg/L. Alkalinity (measured as mg CaCO3/L) was 24.2 and conductivity was

42.6 .S at 25°C), indicating rather dilute water. Dissolved organic carbon was 10.4 mg/L, and turbidity, as measured in Jackson units, was 20, which together indicate the water was relatively clear. DISCUSSION

Lake of the Clouds Water Profile

Circumneutral pH often coincides with high algal diversity. According to the research results, Lake of the Clouds satisfied this correlation. Its moderate tem-


perature is also a probable cause of the wide range of genera of Chlorophyta that were observed (Denicola 1996).

Lake of the Clouds Algal Flora

Based on the number of genera observed, diatoms were by far the dominant algal group in Lake of the Clouds. This is typical of alkaline to circumneutral bodies of water in North America (Stevenson et al. 1996). The discovery of three species new to science leads us to believe that there may be more undescribed di- atoms that our initial survey did not detect.

Desmidiales was the most abundant order among the Chlorophyta found in Lake of the Clouds. Desmids can indicate good water quality, because they do not prefer anthropogenically disturbed or extremely eutrophic bodies of water (Ngearnpat and Peerapornpisal 2007). Most of the samples collected were from the eulittoral zone, the relatively shallow portions of the lake. This study sug- gests that Lake of the Clouds has a soft water quality because an abundance of desmids in the eulittoral zone can indicate a calcium poor water body (Steven- son et al. 1996). The lake’s geology is a potential cause of its lack of calcium. The lake’s bedrock is part of the Canadian Shield, which contributes monovalent cations instead of divalent cations to water bodies. A higher divalent cation con- centration would lead to harder water, reduce the occurrence of desmids, and in- crease the prevalence of cyanobacteria (Molot and Dillon 2008).

Based on the number of genera, Chlorophyta was the most diverse non-di- atom algal division in the samples. The abundance of Chlorophyta is typical for the soft water chemistry of the lake, since soft water lakes often have algal dis- tributions characterized by Mougeotia and other filamentous genera as well as Cosmarium, and Staurastrum (Stevenson et al. 1996).

Based on the number of genera, Cyanophyta was the second most diverse non-diatom algal division. A few genera of Cyanophyta, such as Microcystis and Anabaena, produce toxic metabolites that can be hazardous to the ecosystem (Burgess 2001). These two genera were observed only in relatively small num- bers. Other indicator taxa, such as desmids (which prefer soft water and a lower calcium level) and members of the Zygnemataceae (which also dominate the lit- toral zone of soft water lakes), were more common than Cyanophyta (Stevenson et al. 1996).

Comparison of Algal Taxa Between Lake of the Clouds and Other Lakes in the Midwest

Comparison of our results with other published records for the state of Michi- gan or the Midwest region is difficult, since there are few studies of all algal groups from specific ecosystems, especially where taxa are treated to the level of species. Unlike in Lake of the Clouds, Bambusina and Desmidium were not ob- served in great numbers in other Michigan lakes such as Wycamp Lake, Lark’s Lake, Paradise Lake, Monro Lake, Lancaster Lake, Walloon Lake, Burt Lake, Long Lake, and Douglas Lake (unpublished, Warren 1995). Scenedesmus, Oscil- latoria, and Anabaena, which were reported to be abundant in Douglas Lake


(Gulley and Kennedy 1987), were not found in Lake of the Clouds in large num- bers. Similarly, Anabaena, Oscillatoria, and Schizothrix, which were abundant in Northern Lower Michigan groundwater springs and seeps (Smith 1995) were not common in Lake of the Clouds.

The description of three new species of diatoms from this lake in the mid- western United States, one of the more intensively studied regions of the coun- try, suggests that this oligotrophic lake may harbor other new species of algae. Unlike other algal groups (e.g., Prescott 1962), the diatoms of Michigan, espe- cially the Upper Peninsula, are not well known. Further study of the algal flora of the region is warranted and may result in the description of other new taxa.

Sources of Error and Possible Future Study

A few possible error sources should be addressed and considered for future investigations. A more comprehensive floral survey could have incorporated more diversity in collection sites. For instance, the northern shoreline of the lake and the inflow have not yet been investigated for algal flora. Considering that the inflow is a transition point from a river to lake ecosystem, algae may be present due to changes in both biotic and abiotic conditions. Additionally, it may be in- teresting to compare inflow taxa data with the outflow data. Collecting near the landslide on the northern shoreline could show what impact this particular dis- turbance has on the flora, and the area could be a unique habitat for algae.

The abundance and distribution of algal species will vary seasonally in the lake. This is partly due to the availability of various nutrients, such as nitrogen and phosphorus, which are necessary in different quantities for different taxa. The abundance of a nutrient might benefit one alga more than another, confer- ring a competitive advantage. For example, an aquatic system low in nitrogen greatly limits the ability of green algae (Chlorophyta) to photosynthesize and thus to grow and reproduce, but does not affect some blue-green algae (Cyanophyta), since they have the ability to fix their own nitrogen through use of a heterocyte (Canter-Lund and Lund 1995). Sampling at various times throughout the season would increase the likelihood of finding more taxa of algae, leading to a more complete floral record and understanding of the algal ecology in the lake.


Since no previous algal floristic studies have been conducted in Lake of the Clouds, the location was suitable for a biodiversity survey. In this research, the diatoms were the most abundant and diverse in genera. However, the presence of several taxa of Chlorophyta indicated that Lake of the Clouds has a soft water body and relatively low levels of pollutants. The results showed that the lake is a clean water body with a relatively rich algal community. Our records of the rich algal taxa is consistent with the history of relatively few anthropogenic ac- tivities in the Porcupine Mountains Wilderness State Park. Because this study is


not exhaustive, we recommend further algal diversity research on Lake of the Clouds.


Brachysira ontonageniana Kociolek et Lowe, sp. nov.

Description: Valves lanceolate-clavate, apices not protracted, rounded at the headpole and footpole. Length 16–40 .m, breadth 4.5–5.5 .m. Axial area nar- row, straight, forming a small, indistinct, narrowly lanceolate central area. An elongated areola may be positioned on one side of the central area. Raphe fili- form, straight, with external proximal raphe ends dilated, round. Striae distinctly punctate, areolae within a stria appear slightly offset from neighboring striae. Striae radiate the entire length of the valve. Striae number 28-31/10 .m.

Holotype: JPK Collection, slide number 273055, University of Colorado (COLO)

Etymology: This species is named for the Michigan county in which it is found.

Figures 1–10, 15, 16; Figure 1 is of the holotype.

Brachysira gatesii Kociolek et Lowe, sp. nov.

Description: Valves lanceolate, barely clavate, with headpole rounded and footpole protracted, broadly rounded. Length 16–32 .m, breadth 4-5 .m. Axial area narrow, straight, forming a small but distinct, rounded central area, bordered on both sides by shortened striae the shortest being in the middle. Raphe fili-

FIGURES 1–10. LM, Brachysira ontonageniana, sp. nov. Valve view, size diminution series. Scale bar = 10 .m. Figure 1 is of the holotype.


FIGURES 11–14. LM. Brachysira gatesii, sp. nov. Valve view, size diminution series. Scale bar = 10 .m. Figure 11 is of the holoytpe.

form, external proximal raphe ends not rounded. Striae distinctly punctate, areo- lae within a stria slightly offset from those in neighboring striae giving the im- pression of longitudinal undulations. Striae radiate throughout the length of the valve, numbering 29-32/10 .m.

Holotype: JPK Collection, slide number 273056, University of Colorado (COLO)

Etymology: The species is named for former UMBS director, Dr. David Gates.

In the SEM, both Brachysira ontonageniana (Figures 15, 16) and B. gatesii (Figures 17, 18) have valve exteriors with a thick marginal ridge along each side of the valve with the raphe bordered on both sides by longitudinal ribs. The valve face has elongated to ellipsoidal areolae that are covered internally by hy- mens (Figure 19). Siliceous nodules (“warts” of Round et al. 1990, p. 540) are found scattered along the interstriae. Elongated areolae occur on the mantle of the valve. The elongated areolae are evident internally. The central nodule is small and barely elevated internally. At the poles are indistinct helictoglossae.

Brachysira gatesii differs from B. ontonageniana in the shape of the valve, striae of the central area, and undifferentiated external proximal raphe ends.

These two species possess the features typical for the genus Brachysira (Round et al. 1990). In addition, they are slightly asymmetrical to the transapical axis, and this feature has been documented for other Brachysira species. Lange- Bertalot and Moser (1994) illustrate several species of the genus with this asym- metry, including species from New Caledonia (e.g. B. microclava Lange- Bertalot & Moser, B. archibaldi Coste & Ricard among others), the Philippines

(e.g. B. irawanae (Podzorski & Håkansson) Lange-Bertalot & Moser), and Aus- tralia (B. archibaldii var. crassistriata Lange-Bertalot). A similar asymmetry is not well known in the northern hemisphere. Lange-Bertalot and Moser (1994) il- lustrate unnamed species from the Rocky Mountains (see plate 31, figs. 20, 21 Page  17 2013 THE MICHIGAN BOTANIST 17

FIGURES 15–19. SEM. Figs 15, 16. Brachysira ontonageniana, sp. nov. Figure 15, External valve view. Note thickened siliceous rib bordering the raphe and scattered nodules oriented along the striae. Scale bar = 5 .m. Figure 16. Internal valve view. Scale bar = 5 .m. Figures 17–19, B. gatesii. Figure 17. External valve view. Note thickened siliceous rib bordering the raphe and scattered nod- ules oriented along the striae Striae on the mantle are elongated areolae. Scale bar = 5 .m. Figure 18. Internal valve view. Note that the areolae are occluded by hymenate coverings. Scale bar = 5 .m. Fig. 19. Close up of areolae showing the fine hymenate occlusions. Scale bar = 0.5 .m


FIGURES 20–23. LM. Gom- phonema porcupiniana, sp. nov. Valve view, size diminu- tion series. Scale bar = 10 .m. Figure 20 is of the holotype.

and plate 46, fig. 26 therein) that resemble these species from Lake of the Clouds. Figures 11–14, 17–19; Figure 11 is of the holotype.

Gomphonema porcupiniana Kociolek et Lowe, sp. nov.

Description: Valves large, length 77–110 .m, breadth 9–12 .m. Valves linear- clavate with headpole protracted, sometimes bent to one side and rounded, foot- pole. Axial area straight, forming a unilaterally expanded central area bordered by 2–4 shortened striae. On the other side of the central area is a small, rounded stigma positioned at the end of the central stria. The central nodule is large and prominent. Raphe distinctly lateral, slightly undulate, with external distal raphe ends deflected towards the same side of the valve, opposite the stigma. Striae are distinctly punctate, radiate about the center becoming strongly radiate near the poles. Striae continue around the headpole. Striae number 7–8/10 .m in the mid-


FIGURES 24–27. SEM. Gomphonema porcupiniana. External views. Figure 24. Valve view show- ing outline of the valve. Scale bar =10 .m. Figure 25. Headpole, with deflected distal raphe end. Are- olae are narrow slits. Scale bar = 1 .m. Figure 26. Footpole with porelli of the apical pore field phys- ically separate and morphological distinct from the areolae. Distal raphe end bisects the pore field. Scale bar = 1 .m. Figure 27. Central area with dilated external proximal raphe ends. Stigmal open- ing is elliptical. Striae are narrow slits. Scale bar = 1 .m.


FIGURES 28–31. SEM. Gomphonema porcupiniana. Internal views. Figure 28. Valve view showing outline of the valve. Scale bar = 10 .m. Figure 29. Large septum present at the headpole. Scale bar = 1 .m. Figure 30. Footpole, with large pseudoseptum and below it a helictoglossa offset from the raphe branch. Scale bar = 1 .m. Figure 31. Central nodule, with recurved proximal raphe ends and slit-like stigma opening. Scale bar = 1 .m.


dle of the valve, 11–13/10 .m at the apices. A distinct, bilobed apical pore field is positioned at the footpole on the valve face. Septa and pseudosepta are present at both poles.

Holotype: JPK Collection, slide number 273054, University of Colorado (COLO)

Etymology: This species is named for the Porcupine Mountains of Michigan, where it is found.

In the SEM, the valve exterior is covered with narrow, slit-like areolae that may be bifurcated (Figures 24–27). The headpole shows the distal raphe ends to be deflected on the mantle (Figure 25). At the footpole (Figure 26), the apical pore fields are comprised of round porelli, that are physically separate and dif- ferentiated from the slit-like areolae. The pore field extends from the valve face to the mantle, and appears to be asymmetrical, with one side being more exten- sive than the other. Small depressions are evident in the central area of the valve (Figure 27). The proximal raphe ends are dilated and tear-drop shaped, and the stigma opening is elongated, quite different as compared to the slit-like areolae. The slit-like appearance of the areolae is unlike most “typical” Gomphonema species that typically have “c”-shaped openings (Kociolek and Kingston 1999; Round et al. 1990; Reichardt 1999, 2001). The slit-like areolae in G. porcu- pinensis appear to differ from the lineolate striae seen in G. reimeri Kociolek & Kingston (Kociolek & Kingston 1999).

Internally, there is a large septum associated with the girdle band (Figures 28- 30), especially at the headpole. The central nodule is elongate and bears recurved proximal raphe ends and at the base of the central nodule an elongate stigma opening (Figure 31). Striae have fine siliceous struts oriented perpendicular to the striae (Figure 31).

This species resembles G. sagitta Schumann in overall shape, but is much larger. Patrick & Reimer (1975) report the length of G. sagitta (as G. subtile Ehrenberg var. sagitta (Schumann) Cleve) to be 40-51 .m; a similar size range is given by Krammer and Lange-Bertalot (1986), though they lump G. sagitta with G. subtile. Krammer and Lange-Bertalot (1986) suggest tropical forms may be up to 70 .m in length, but these are still smaller than this Lake of the Clouds endemic.

Figures 20–31; Figure 20 is of the holotype.


We would like to acknowledge the assistance of Tyler Elias and Michael Grant, who conducted water chemistry tests for this study. We would also like to acknowledge Katie Kavulic, graduate stu- dent from University of Colorado at Boulder, for sampling the water chemistry of Lake of the Clouds, collecting pH and GPS measurements, and accompanying us on the trip. We would also like to acknowledge Dr. Bob Anderson, whom we met at the study site, for his assistance. Finally, we would like to thank the University of Michigan Biological Station staff who have supported us with the report. This includes, but is not limited to, Bob Vande Kopple, Sherry Webster and Karie Slavik. The comments of two anonymous reviewers are greatly appreciated.



Ackley, A. B. (1929). New species and varieties of Michigan algae. Transactions of the American Microscopical Society 48: 302–309.

Ackley, A. B. (1932). Preliminary report on Michigan algae, exclusive of desmids and diatoms. Pa- pers of the Michigan Academy Science, Arts and Letters 15: 1–49.

Burgess, C. (2001). A wave of momentum for toxic algae study. Environmental Health Perspectives

109: 160–161. Canter-Lund, H. and J. W. G. Lund. (1995). Freshwater algae: Their microscopic world explored. Biopress Ltd., Bristol. Croasdale, H. T., C. E. M. Bicudo, and G. W. Prescott. (1983). A synopsis of North American desmids. Part II. Desmidiaceae: Placodermae Section 5. The filamentous genera. University of Nebraska Press, Lincoln.

Davis, M.B. (1993). Old growth in the East: A survey. Wild Earth: 18–145.

Denicola, D. M. (1996). Periphyton responses to temperature at different ecological levels. In R. J. Stevenson, R. Lowe and M. Bothwell (eds). Algal ecology: Freshwater benthic ecosystems. Aca- demic Press, San Diego.

Derouin, S. A., T. V. Lowell, and I. Hajdas. (2007). Landscape evolution and deglaciation of the upper peninsula, Michigan: An examination of chronology and stratigraphy in kettle lake cores. Journal of Great Lakes Research 33: 875–886.

Dillard, G. E. (1989a). Freshwater algae of the southeastern United States. Part 1. Chlorophyceae: Volvocales, Tetrasporales and Chlorococcales. Bibliotheca Phycologica. Band 81. J. Cramer, Stuttgart.

Dillard, G. E. (1989b). Freshwater algae of the southeastern United States. Part 2. Chlorophyceae: Ulotrichales, Microsporales, Cylindrocapsales, Sphaeropleales, Chaetophorales, Cladophorales, Schizogoniales, Siphonales and Oedogoniales. Bibliotheca Phycologica. Band 83. J. Cramer, Stuttgart.

Dillard, G. E. (1990). Freshwater algae of the southeastern United States. Part 3. Chlorophyceae: Zynematales: Zygnemataceae, Mesotaeniaceae and Desmidiaceae (Section 1). Bibliotheca Phyco- logica. Band 85. J. Cramer, Stuttgart.

Dillard, G. E. (1991a). Freshwater algae of the southeastern United States. Part 4. Chlorophyceae: Zygnematales: Desmidiaceae (Section 2). Bibliotheca Phycologica. Band 89. J. Cramer, Stuttgart.

Dillard, G. E. (1991b). Freshwater algae of the southeastern United States. Part 5. Chlorophyceae: Zygnematales: Desmidiaceae (Section 3). Bibliotheca Phycologica. Band 90. J. Cramer, Stuttgart.

Dillard, G. E. (1993). Freshwater algae of the southeastern United States. Part 6. Chlorophyceae: Zygnematales: Desmidiaceae (Section 4). Bibliotheca Phycologica. Band 93. J. Cramer, Stuttgart.

Gulley, A. R. and C. A. Kennedy. (1987). Comparison of phytoplankton, periphyton and benthic algae in two different beach pools off of Douglas Lake. Unpublished class report, University of Michigan Biological Station.

Kociolek, J. P. and J. C. Kingston. 1999. Taxonomy, ultrastructure and distribution of gom- phonemoid diatoms (Bacillariophyceae: Gomphonemataceae) from rivers of the United States. Canadian Journal of Botany 77: 686–705.

Komárek, J. and K. Anagnostidis. (1999). Cyanoprokaryota. 1. Chroococcales. In Ettl, H., G. Gärt- ner, H. Heynig, and D. Mollenhauer (eds). Süßwasserflora von Mitteleuropa. 19/1. Spektrum, Hei- delberg, Berlin.

Komárek J. and K. Anagnostidis. (2005). Cyanoprokaryota 2. Oscillatoriales. In: Büdel B., L. Krienitz, G. Gärtner, and M. Schagerl (eds). Süßwasserflora von Mitteleuropa. 19/2. Else- vier/Spektrum, Heidelberg.

Krammer, K. and H. Lange-Bertalot. (1986). Bacillariophyceae. 1. Teil: Naviculaceae.—In Ettl, H.,

J. Gerloff, H. Heynig & D. Mollenhauer (editors): Süßwasserflora von Mitteleuropa 2/1. G. Fis- cher, Stuttgart. Lange-Bertalot, H. and G. Moser. (1994). Brachysira. Monographie der Gattung. Bibliotheca Diato- mologica 29: 1–212.


Larson, G. and R. Schaetzl. (2001). Origin and evolution of the Great Lakes. Journal of Great Lakes Research 27: 518–546.

Moffett, J. W. (1940). A fisheries survey and management suggestions for some lakes of the Ottawa National Forest, Michigan. (Fisheries research report: 630). Michigan Department of Natural Re- sources, Fisheries Division. Available at (Ac- cessed 18 April, 2014).

Molot, L. A. and P. J. Dillon. (2008). Long-term trends in catchment export and lake concentrations of major cations in the Dorset study area, central Ontario. Canadian Journal of Fisheries and Aquatic Sciences 65: 809–820.

Ngearnpat, N. and Y. Peerapornpisal. (2007). Application of desmid diversity in assessing the water quality of 12 freshwater resources in Thailand. Journal of Applied Phycology 19: 667–674.

Ojakangas, R. W. and C. L. Matsch. (1982). Minnesota’s Geology. University of Minnesota Press, Minneapolis.

Patrick, R. M. and C. W. Reimer. (1975). The diatoms of the United States. Vol. II, Part 1. Mono- graph 13. Academy of Natural Sciences of Philadelphia.

Prescott, G. W. (1962). Algae of the western Great Lakes area. W.C. Brown Company Publishers, Dubuque, Iowa.

Prescott, G. W. (1978). How to know the freshwater algae. W. C. Brown Company Publishers, Dubuque, Iowa.

Prescott, G. W., H. T. Croasdale, and W. C. Vinyard. (1972). A synopsis of North American desmids. Part 1. Saccodermae, Mesotaeniaceae. North American Flora, Series 2, Part 6. The New York Botanical Garden, Bronx, New York.

Prescott, G. W., H. T. Croasdale, and W. C. Vinyard. (1975). A synopsis of North American desmids. Part 2. Placodermae. Section 1. University of Nebraska Press, Lincoln.

Prescott, G. W., H. T. Croasdale, and W. C. Vinyard. (1977). A synopsis of North American desmids. Part 2. Placodermae. Section 2. University of Nebraska Press, Lincoln.

Prescott, G. W.,H. T. Croasdale, W. C. Vinyard, and C. E. M. Bicudo. (1981). A synopsis of North American desmids. Part 2. Placodermae. Section 3. University of Nebraska Press, Lincoln.

Prescott, G. W., C. E. M. Bicudo, and W. C. Vinyard. (1982). A synopsis of North American desmids. Part 2. Placodermae. Section 4. University of Nebraska Press, Lincoln.

Reichardt, E. (1999): Zur Revision der Gattung Gomphonema. Die Arten um G. affine/insigne, G. angustatum/micropus, G. acuminatum sowie gomphonemoide Diatomeen aus dem Oberoligozän in Böhmen. Iconographia Diatomologica. 8: 1–203.

Reichardt, E. (2001). Revision der Arten um Gomphonema truncatum und G. capitatum. Pp. 187–224 In: R. Jahn, J. P. Kociolek, A. Witkowski, and P. Compère, (editors): Festschrift: Studies on diatoms (dedicated to Prof. Dr. Dr. H. C. Horst Lange-Bertalot on the occasion of his 65th birthday). Gantner Verlag, Ruggell, Liechtenstein

Rominger, C. L. (1895). Geological report on the upper peninsula of Michigan: Exhibiting the progress of work from 1881 to 1884. Iron and copper regions. Volume 5. R. Smith & Co, Lansing, Mich.

Round, F. E., R. M. Crawford, and D. G. Mann. (1990). The diatoms. biology & morphology of the genera. Cambridge University Press. Cambridge.

Smith, B. (1995). A survey of the algal flora in ground water spring and seeps in Northern Lower Michigan. Unpublished class report, University of Michigan Biological Station.

Stevenson, R. J., M. L. Bothwell. and R. L. Lowe, eds. (1996). Algal ecology: Freshwater benthic ecosystems. Academic Press, San Diego.

Taube, C. M. (1949). The fishes and the physical aspects of Lake of the Clouds, Mirror Lake, Big Carp River, and Little Carp River, of the Porcupine Mountains State Park. (Fisheries Research Re- port 1211) Michigan Department of Natural Resources, Fisheries Division. Lansing.

van der Werff, A. (1955). A new method of concentrating and cleaning diatoms and other organisms. Verhandlungen Internationalen Vereingung für Theoretische und Angewandte Limnologie 13: 276–277.

Warren, M. (1995). A survey of phytoplankton communities in ten northern Michigan lakes. Unpub- lished class report, University of Michigan Biological Station.

Wehr, J. D. and R. G. Sheath, editors. (2003). Freshwater algae of North America: Ecology and clas- sification. Academic Press, San Diego.


West, W. and G. S. West. (1904). A monograph of the British Desmidiaceae. Volume I. The Ray So- ciety, London.

West, W. and G. S. West. (1905). A monograph of the British Desmidiaceae. Volume II. The Ray So- ciety, London.

West, W. and G. S. West. (1908). A monograph of the British Desmidiaceae. Volume III. The Ray So- ciety, London.

West, W. and G. S. West. (1912). A monograph of the British Desmidiaceae. Volume IV. The Ray So- ciety, London.

West, W., G. S. West, and N. Carter, N. (1912). A monograph of the British Desmidiaceae. Volume

V. The Ray Society, London.