11 th USGS NAWQA Workshop on Harmonization of Algal Taxonomy

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1 11 th USGS NAWQA Workshop on Harmonization of Algal Taxonomy November 21-24, 2003 John Carroll University, Cleveland, OH Report No Phycology Section Patrick Center for Environmental Research Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA February 11, 2009

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3 Table of Contents Introduction 1 The Taxon Hydrocoleum brebissonii in USGS NAWQA Samples. 2 Homoeothrix Species in USGS NAWQA Samples.. 7 Dr. Jeří Komárek s Lecture on Oscillatoriales 19 Literature Cited.. 23 Appendices Appendix A Agenda for the 11 th USGS NAWQA Workshop on Harmonization Of Algal Taxonomy. 25 Appendix B Workshop participants. 27 Appendix C List of Measurements/Characteristics of Tapinothrix-type Homoeothrix Forms from USGS NAWQA samples.. 29 Appendix D Algal images from the 11 th USGS NAWQA Workshop on Harmonization Of Algal Taxonomy.. 31 i

4 Introduction The Eleventh USGS NAWQA Workshop on Harmonization of Algal Taxonomy was held in the Biology Department of John Carroll University November This was the second workshop devoted to soft-bodied, non-diatom forms (the Ninth was the original soft-bodied, non-diatom workshop). The main focus of the workshop was a series of lectures by Dr. Jeří Komárek of the University of South Bohemia in the Czech Republic. Dr. Komárek s lectures included the modern cyanobacteria classification and, in particular, the group Oscillatoriales, filamentous forms without heterocytes and true branching. Dr. Rex L. Lowe (Bowling Green State University) re-examined USGS NAWQA samples with large populations of the Oscillatorialean taxon Hydrocoleum brebissonii, a taxon recently revised with characters not considered previously. Similarily, Dr. Jeffery Johansen (John Carroll University) re-examined samples with taxa now considered members of the genus Homoeothrix. The following sections are a review of the presentations and discussions illustrated with algal images from USGS NAWQA samples that were taken before or during the workshop. Some of the illustrations from Dr. Komárek s lecture were published elsewhere and are so noted. 1

The Taxon Hydrocoleum brebissonii in USGS NAWQA Samples The taxon Hydrocoleum brebissonii Kützing ex Gomont is a major taxon found in a large number of USGS NAWQA samples.

5 The Taxon Hydrocoleum brebissonii in USGS NAWQA Samples The taxon Hydrocoleum brebissonii Kützing ex Gomont is a major taxon found in a large number of USGS NAWQA samples. For the USGS NAWQA program, this taxon has had several different species names, including portions of Drouet s Microcoleus vaginatus Gomont ex Gomont and M. lyngbyaceous Kützing ex Rabenhorst. In previous taxonomies (1997 and 2001-NAWQA start lists) the taxon name for most of these forms was updated to Geitler s Hydrocoleum brebissonii. Important characters included the granulation and ends of the trichome. In the modern cyanobacteria taxonomy (Anagnostidis and Komárek, 1988), H. brebissionii is placed in the genera Blennothrix as B. brebissionii (Kützing ex Gomont) Anagnostidis et Komárek. Since the modern taxonomy uses different characters we decided to reexamine USGS NAWQA samples that had large populations of the taxon that was considered Hydrocoleum brebissionii. In the examination of specimens before the workshop, Dr. Lowe concluded that these were Phormidium (Phormidiaceae) and did not belong in Blennothrix (Oscillatoriaceae). The important characters included size (trichome diameter ( µm), isodiametric cells, presence and absence of a thin sheath and the presence or absence of a thicken (calyptra) or capitate end cell. In the Figure 1, there are Figure 1. Cyanobacteria filament from a sample collected by USGS NAWQA Program in the Mecan River, near Richmond, WI (NAWQA Sample WMIC0593ARE0010B). 2

granulated cell walls and a calyptrate end cell (5.5µm broad). Figure 2 is an example with a thin sheath, broad end cell and no granulation at the cell walls (4.6µm Figure 2.

6 granulated cell walls and a calyptrate end cell (5.5µm broad). Figure 2 is an example with a thin sheath, broad end cell and no granulation at the cell walls (4.6µm Figure 2. Cyanobacteria filament from a sample collected by USGS NAWQA Program from the Jones Loop Road wetlands (Charlotte County, FL; NAWQA Sample SOFL1196ARE1100B). broad). Collections (Fig. 3) containing a mixture of morphologies, including capitate and broad ends, granulated and not granulated were not uncommon (for more examples see Appendix C). 3

Figure 3. Cyanobacteria filament from a sample collected by USGS NAWQA Program on the St. Regis River near St. Regis, MT (NAWQA Sample NROK0800ARE0003B).

7 Figure 3. Cyanobacteria filament from a sample collected by USGS NAWQA Program on the St. Regis River near St. Regis, MT (NAWQA Sample NROK0800ARE0003B). The taxa Phormidium autumnale (Agardh) Trevisan ex Gomont and P. favosum Gomont ex Gomont were suggested as possible species names for taxa we had classified as Hydrocoleum brebissionii. Examination of specimens during the workshop, using Dr. Komárek s updated taxonomy, indicated that many would be considered Phormidium autumnale especially the capitate forms. Absence at times of mature ends makes this taxon difficult to identify. It was cautioned that P. autumnale had a very broad species concept (enough to cover most of the specimens we have seen) and might be revised in the future. Phormidium favosum was not confirmed; the ecology of most USGS NAWQA forms is different (P. favosum has a limited range found in small, fast-flowing, cold, calcareous streams; Dr. Komárek, personal communication). Several other species of Phormidium were identified during the workshop including Phormidium formosum (Bory ex Gomont) Anagnostidis et Komárek from Florida (Fig. 4) and Utah (Fig. 5) and P. granulatum (Gardner) Anagnostidis from North Carolina (Fig. 6). 4

Service Creek (Alamance County, NC; NAWQA sample ALBE0503ARE0029BA). Figure 5.

8 Figure 4. Cyanobacteria filament from a sample collected by USGS NAWQA Program on Service Creek (Alamance County, NC; NAWQA sample ALBE0503ARE0029BA). Figure 5. Cyanobacteria filament from a sample collected by USGS NAWQA Program on Cub River near Richmond, UT (Cache County, UT; NAQWA sample GRSL0701ADE2108B). 5

9 Figure 6. Cyanobacteria filament from a sample collected by USGS NAWQA Program (Tamiami Canal, Collier County, FL; NAWQA sample SOFL0697ARE0024B). 6

10 Homoeothrix Species in USGS NAWQA Samples In USGS NAWQA samples there were many populations of filamentous cyanobacteria colonies (trichomes less than 3 µm diameter) that were identified as Amphithrix janthina Bornet et Flahault. Closer examination (i.e., at a higher magnification than during analysis) indicated that there were differences in the trichome structure (different length to width ratios, constricted or not, size, etc.). It was also realized that A. janthina was actually a complex of a small coccoid cyanobacteria at the base of a colony of tapering filaments. In the last taxonomic revision (2001 NAWQA start list) these forms were transferred to Homoeothrix janthina. The purpose of this workshop is to explore the USGS NAWQA populations and determine if we have distinct species and if they fit descriptions of the modern cyanobacteria taxonomy. As was noted in Dr. Komárek s lecture, the genera Homoeothrix refers to two distinct groups. The true Homoeothrix is larger and has characteristics of the family Oscillatoriaceae; H. juliana is a common example. The other, smaller sized group belonging to the Pseudanabaenaceae, are referred as Tapinothrix-type Homoeothrix; H. janthina belongs to this group along with species of the genera Tapinothrix. Prior to the workshop, Dr. Johansen and his colleagues at John Carroll University, examined over 50 populations of the Tapinothrix-type Homoeothrix from USGS NAWQA samples. From the literature, for these types of samples (freshwater, streams and benthic on rocks) they considered the following taxa: Homeothrix janthina (Bornet et Flahault) Starmach is µm broad, and the crosswalls are mostly unconstricted. Cells are nearly quadratic, about as long as broad. When the filament tapers to a hair, the cells are longer than broad in the hair. Homeothrix varians Geitler is a taxon that in most cases is untapered, but can taper to a hair with elongated cells. Its listed dimensions are filaments µm wide, cells disk-like to almost quadratic (mostly ½ as long as broad). The hair has cells distinctly longer than broad (Fig. 7). Leptochaete stagnalis Hansgirg = Homeothrix stagnalis (Hangsgirg) Komárek et Kováćik is wider than either of the other taxa, µm at the base, is tapered, with cells ½-1 times as long as broad. Very little of the material in the USGS NAQWA samples fit any of these species very well. Either there are many different species, or very few species that must be broadly defined. Whether narrow or broad definitions are used, the USGS NAWQA material does not appear to be conspecific with these taxa. Size measurements seem to overlap and are smaller than described species. Important characters include visibility of crosswalls, tapering, degree of crosswall constrictions and presence of merismatic zones (Figs. 8-11). 7

11 Figure 7. This Homeothrix specimen, except for the tapering hair, fits Homeothrix varians fairly well. The filaments are um at the base. Trichomes taper from um wide at the base to at the tips. Cells are um long (Halfway Brook at Barryville, NY; NAWQA sample ID DELR0901ARE0247B). 8

12 Figure 8. This Homeothrix specimen has nearly quadrate cells, and does not have distinct crosswalls. These trichomes are constricted at the crosswalls. No tapering is evident (Halfway Brook at Barryville, NY; NAWQA sample ID DELR0901ARE0247B). 9

13 a b c d e Figure 9. Various populations with difficult to discern crosswalls, varying degrees of tapering and constrictions (a) Jordan Creek near Schnecksville, PA; DELR 0800ARE0109B; b) Schuylkill River at Philadelphia, PA; DELR0801 ARE0233B; c) Sawkill Creek at Milford, PA; DELR0801ARE0243B; d) Canyon Creek near Glacier, WA; PUGT0897ARE0002B; e) Santa Ana River near Running Springs, CA; SANA0799ARE0001B, respectively). 10

14 Figure 10. This population clearly tapers, has walls that are indistinct, and only slight constrictions at crosswalls (Satus Creek near Toppenish, WA; NAWQA Sample ID YAKI0900ARE0009B). 11

Figure 11. Some of the populations, particularly those with distinct septa, had what appeared to be meristematic zones, as the specimen to the right displays.

Species identifications were only tentative, as the literature, especially for the modern cyanobacteria taxonomy, is rather limited, especially when considering North American forms.

15 Figure 11. Some of the populations, particularly those with distinct septa, had what appeared to be meristematic zones, as the specimen to the right displays. These zones were usually subapical (Halfway Brook near Barryville, NY; NAWQA Sample ID DELR0901ARE0274B). During the workshop, we observed several different forms of the Tapinothrixtype Homoeothrix. Species identifications were only tentative, as the literature, especially for the modern cyanobacteria taxonomy, is rather limited, especially when considering North American forms. Homoeothrix janthina (Fig. 12), as described above, are the smallest forms, with isodiametric cells. Of the other forms we expected, we did not find any form large enough to be considered H. stagnalis and several forms were fairly close to H. varians (see Fig. 7). However, for several populations of a form with short cells and constricted cell walls that was thinner than described for H. varians, we used the taxon H. simplex Voronichin (Figs. 13 and 14). The size (consistently µm) and short cells made this species appear to separate from H. janthina (high end of size range, shorter cells) and H. varians (thinner trichomes). 12

16 Figure 12. A cyanobacteria colony of Homoeothrix janthina (Bornet et Flahault) Starmach collected from rocks in Little Cottonwood Creek at Crestwood Park, Salt Lake City, UT (USGS NAWQA Great Salt Lake Study Unit; NAWQA sample ID GRSL0899ARE0103A). 13

17 Figure 13. A cyanobacteria colony of Homoeothrix simplex Voronichin collected from rocks in Shirtee Creek near Odena, AL (USGS NAWQA Mobile River Basins Study Unit; NAWQA sample ID MOBL0600ARE0193B) There were uncertainties with the other Tapinothrix-type Homoeothrix species observed during the workshop. An abundant form found in the Ozark Plateau Study Unit with barrel-shaped cells (i.e., strongly constricted) with cell diameters tapering from µm was tentatively placed in the taxon H. cf. margalefii Komárek et Kalina (Fig. 15). There was an uncommon form, µm wide with a sheath colored (yellow to brown) at the base that was similar to H. gracilis (Hansgirg) Komárek et Kováčik (Fig. 16). The true Homoeothrix, placed in the family Oscillatoriaceae in the modern cyanobacteria taxonomy was found in several USGS NAWQA samples. Homoeothrix juliana (Bornet et Flahault) Kirchner is larger ( µm diameter), with short cells and tapers to a point (Fig. 17). 14

18 Figure 14. A cyanobacteria colony of Homoeothrix simplex Voronichin collected from rocks in the Middle Fork Holston River at Seven Mile Ford, VA (USGS NAWA Upper Tennessee River Basins Study Unit; NAWQA sample ID UTEN0597ARE0023B). 15

19 Figure 15. A cyanobacteria colony of Homoeothrix cf. margalefii Komárek et Kalina collected from rocks in Yokum Creek near Oak Grove, AR (USGS NAWQA Ozark Plateaus Study Unit; NAWQA sample ID OZRK0895ADE0008E). 16

20 Figure 16. A cyanobacteria colony of Homoeothrix gracilis Komárek et Kováčik collected from rocks in Holes Creek at McEwen Road Montgomery County, OH (USGS NAWQA Great and Little Miami River Basins Study Unit; NAWQA sample ID MIAM0701ARE0085B). 17

21 Figure 17. A cyanobacteria colony of Homoeothrix juliana (Bornet et Flahault) Kirchner collected from rocks in Mikes Creek at Powell, MO (USGS NAWQA Ozark Plateaus Study Unit; NAWQA sample ID OZRK0894ARE0005B). 18

22 Dr. Jeří Komárek s Lecture on Oscillatoriales In addition to a series of taxonomic concept articles (Anagnostidis and Komárek, 1985; Komárek, 1986; Anagnostidis and Komárek, 1988; Komárek and Anagnostidis, 1989; Anagnostidis and Komárek, 1990), Dr. Komárek, along with the late Konstantinos Anagnostidis, used the current Süßwasserflora von Mitteleuropa series to describe the modern cyanobacteria taxonomy. This workshop about species of Oscillatorialean cyanobacteria was held as the editing was being made for the revision of the Oscillatoriales (Komárek and Anagnostidis, 2005). Chroococcales (coccoids) were revised prior to the workshop (Komárek and Anagnostidis, 1998) and a future edition will cover the Nostocales (uniseriate filamentous with heterocytes) and Stigonematales (multiseriate filamentous with heterocytes). The lecture was a portion of a comprehensive series of lectures of all groups of the Oscillatoriales. The summary below follows the emphasis on forms likely to be found in USGS NAWQA samples (i.e., freshwater, benthic forms from temperate regions). Forms from other situations (e.g., planktonic, saline, thermal, terrestrial or from tropical regions) are listed but not fully discussed. Cyanobacteria, prokaryotic phototrophs, were amongst the earliest developing life forms, following only the prokaryotic heterotrophs. They developed in the early Precambrian period, 4000 million years ago and were the only producers of oxygen for hundreds of millions of years. The tendency for symbiotic life with heterotrophic eukaryotes was the origin of plant evolution. The use of a taxonomic system is a convention necessary to study natural diversity. The basic unit of a taxonomic system is the species. The main taxonomic system for cyanobacteria in the early to late 20 th century was developed by Lothar Geitler and used mostly morphological data. There were many disadvantages to this system including: Based only on optical microscopical features; Concept of ubiquitous and cosmopolitan distribution of species; Diversity from tropical regions is unsufficiently registered; Ecological and phytogeographic approaches are omitted; Impossible to apply electron-microscopical and molecular data. During the late 20 th century there were several approaches that changed the taxonomic system of cyanobacteria. These approaches, listed below, incorporated new methods with characteristics used to classify cyanobacteria: Paleobotanical research: Cyanoprokaryotes developed in early Precambrium and survived up to now without substantial change of their genotype, by help of a special diversification strategy. Phenotype (morphological) approach: The very wide induced variation dependent on environmental conditions and wide variability within genotypes was recognized. Ecological approach: The ecological specificity in populations and species was found: -- ubiquitous species do not exist; 19

-- ecological demands are species-specific and must be a part of taxonomic evaluation; -- the geographic distribution of species depends on distribution corresponding to environmental conditions.

23 -- ecological demands are species-specific and must be a part of taxonomic evaluation; -- the geographic distribution of species depends on distribution corresponding to environmental conditions. Electron microscopy explained numerous enigmatic characters: The ultrastructural features, mainly thylakoid patterns and cell walls are in agreement with genotype characters. Molecular approach: Molecular sequencing and DNA/DNA hybridization enable to distinguish genotype differences in cyanoprokaryotic types. Creation of a modern cyanobacteria taxonomy involves using the above mention approaches to identify several defining characters of cyanobacteria. The review of morphological characters includes determining the main differentiating characters (sheath, dimensions, granulation, colonial forms, etc.), the coincidence of morphological and ultrastructural characteristics, cyanobacteria life cycles and ecological specificity. Characters based on molecular sequencing that are in agreement with phenotypic markers can be used for modern taxonomic definition at the genera level. An important character in agreement between these characters is the thylakoidal patterns which, in general, are parietal in the lowest forms (coccoids, Pseudanabaenaceae; Fig. 18), radial in larger Figure 18. Thylakoid patterns in the major families of Oscillatoriales. Parietal (A) in Pseudanabaenaceae, radial (B) in Phomidiaceae and irregular (C) in Oscillatoriaceae. 20

24 filamentous (Phormidiaceae) and irregular in the highest forms (Oscillatoriaceae). Figure 19 shows the relationship between coccoid and filamentous cyanobacteria based on Figure 19. Relations between coccoid and filamentous cyanobacteria according to molecular sequencing and electron microscopy characters. From Komárek and Kaštovský,

25 molecular (sequencing) and ultrastructural (electron microscopy of thylakoids) data. It is interesting to note, a group of coccoid cyanobacteria (Pleurocapsales; revised in Komárek and Anagnostidis, 1998 ), because of thylakoid patterns (irregular) and sequencing, should probably be placed with the higher filamentous forms. For modern cyanobacteria taxonomy, the proposed definition of a species is as follows: Group of populations (strains) which belong to one and the same genotype (genus), they are characterized by stabilized phenotype features (definable and recognizable, with distinct limits of variation), and by the same ecological demands. They should occur repeatedly (in time) in various localities with the same ecological conditions. Note the importance of the ecology. Forms with equal phenotypic characters in different habitats are not necessarily equal, have usually evolved apart and have different molecular characters. The modern cyanobacteria taxonomy places the Oscillatoriales between the coccoids and heterocytic filamentous forms. In addition to molecular data, criteria separating the major families of Oscillatoriales include thylakoid patterns (parietial, radial or irregular), diameter of trichome, cell length to width (longer than wide in the Pseudanabaenaceae to shorter than wide in the Oscillatoriaceae) and division with or without necrid cells. CHROOCOCCALES (coccoid and colonial, binary fission) PLEUROCAPSALES (coccoid, combined reproduction by binary and multipled fission OSCILLATORIALES (combined genetic and phenotype criteria): Pseudanabaenaceae (trichomes< 4µm, parietal thylakoids, solitary trichomes) Schizothrichaceae (trichomes < 5µm, parietal thylakoids, fasciculated trichomes) Borziaceae (trichomes 3-12µm, radial thylakoids, reproduction without necroids) Phormidiaceae (trichomes 3-15µm, radial thylakoids, reproduction with necroids) Gomontielaceae (trichomes 6-20µm, irregular thylakoids, flattened trichomes) Oscillatoriaceae (trichomes 6-60µm, irregular thylakoids, cylindrical trichomes) NOSTOCALES (filamentous, heterocytes and akinetes, false branching) STIGONEMATALES (filamentous, faciltative heterocytes and spores, true branching) The majority of Dr. Komárek s lecture dealt with Oscillatoriales genera and characters used to differentiate species. Since this is the main theme of the soon thereafter updated Süßwasserflora von Mitteleuropa volume, details will not be given here but can obtained from Komárek and Anagnostidis (2005). Note that the volume updates many forms from outside of Europe in addition to specific ecological data that are useful with forms found in the USGS NAWQA samples. 22

26 Literature Cited Anagnostidis, K., Komárek, J Modern approach to the classification system of cyanophytes. 1. Introduction. Archiv für Hydrobiologie/Algological Studies 38/39: Anagnostidis, K., Komárek, J Modern approach to the classification system of cyanophytes. 3. Oscillatoriales. Archiv für Hydrobiologie/Algological Studies 50/53: Anagnostidis, K., Komárek, J Modern approach to the classification system of cyanophytes. 5. Stigonematales. Archiv für Hydrobiologie/Algological Studies 59:1-73. Komárek, J Modern approach to the classification system of cyanophytes. 2. Chroococcales. Archiv für Hydrobiologie/Algological Studies 43: Komárek, J. and Anagnostidis, K Modern approach to the classification system of cyanophytes. 4. Nostocales. Archiv für Hydrobiologie/Algological Studies 56: Komárek, J. and Anagnostidis, K (Cyanoprokaryota) Teil 1 (Chroococcales). In Ettl, H., Gerloff, J., Heynig, H., & Mollenhauer, D. (Eds.). Süsswasserflora von Mitteleuropa. 19/1: Gustav Fisher Verlag, Germany. Komárek, J. and Anagnostidis, K (Cyanoprokaryota) Teil 2 (Oscillatoriales). In Ettl, H., Gerloff, J., Heynig, H., & Mollenhauer, D. (Eds.). Süsswasserflora von Mitteleuropa. 19/2: Elsevier GmbH, Müchen, Germany. Komárek, J. and Kaštovský, J Coincidences of structural and molecular characters in evolutionary lines of cyanobacteria. Archiv für Hydrobiologie/Algological Studies 109 (Cyanobacteria Research 4):

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28 Appendices Appendix A Agenda 11 th USGS NAWQA Workshop on Harmonization of Algal Taxonomy November 2003 John Carroll University, Cleveland, OH Friday November 21 st 7:00-9:00pm Introductory Reception Saturday November 22 nd 9:30-10:00 Introduction to USGS NAWQA and Proposed Agenda 10:00-12:00 Part One of Dr. Jeří Komárek s Oscillatoriales Lecture 12:00-1:00 Laboratory set-up and lunch 1:00-1:30 Discussion of the taxon referred to as Hydrocoleum brebissionii in USGS NAWQA samples 1:30-2:00 Discussion of Homoeothrix species in USGS NAWQA samples 2:00-3:30 Part Two Dr. Jeří Komárek s Oscillatoriales Lecture 3:30-5:30 Laboratory study of specimens identified as Hydrocoleum brebissionii Sunday November 23 rd 9:00-10:30 Continued laboratory study on Hydrocoleum brebissionii 10:30-12:00 Part Three Dr. Jeří Komárek s Oscillatoriales Lecture 12:00-1:00 Laboratory on Homoeothrix and lunch 1:00-2:15 Conclusion of Dr. Jeří Komárek s Oscillatoriales Lecture 2:15-5:30 Laboratory study on Homoeothrix Monday November 24 th 9:00-12:00 Laboratory work on other cyanobacteria taxa from USGS NAWQA samples 12:00-1:00 Lunch break 1:00-4:00 Laboratory work on various forms including non cyanobacteria 4:00-5:30 Image editing, review and exchange 25

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30 Appendix B Workshop participants: Jeffery Johansen, Dale Casamatta and Catie Olsen (John Carroll University) Jeří Komárek (University of South Bohemia) Frank Acker, Marina Potapova and Jackie White-Reimer (Academy of Natural Sciences) R. Jan Stevenson and Kalina Manoylov (Michigan State University) Julie Hambrook (USGS Columbus, OH) William R. Cody (Aquatic Taxonomy Specialists) Rex L. Lowe, Jennifer Resh, Paula Furey and Jessie Knapp (Bowling Green State University) 27

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32 Appendix C Table of characters from Tapinothrix-type Homoeothrix specimens from USGS NAWQA samples from throughout the United States. Sample ID refers to the sample ID used at Academy of Natural Sciences (ANSP). Diameter at Filament Base (µm) Diameter at Trichome Base (µm) Diameter at Trichome End (µm) Cell Length (µm) Trichome Length (µm) Crosswalls Distinct (yes/no) Meristematic Zones (yes/no) Constricted Crosswalls (yes/no) Coccoids Present (yes/no) Tapered Flexuous Sample (yes/no) (yes/no) GSN no no no no no yes GSN yes no no no no no GSN yes <60 no no no no no GSN yes >200 no no no yes yes GSN no 50 no no no no no GSN no maybe no no no no GSN yes no no no yes yes GSN yes <140 no no no no yes GSN no <50 no no no no no GSN no <50 no no no no no GSN yes no no no no no GSN no yes yes no no yes GSN no yes no yes no yes GSN no <50 no no yes no no GSN no >100 no no yes no no GSN no 0 yes no yes no yes GSN no <50 no no yes no no GSN yes yes no yes no yes GSN yes no no yes no yes GSN yes >100 yes no yes yes no GSN no yes no yes no no GSN no <100 yes no yes no no GSN no >50 maybe no yes no no GSN yes no no yes no no GSN yes no no yes no no GSN yes >100 yes no yes yes no GSN no <50 yes yes yes no no GSN no 0 yes yes yes no yes GSN no <40 yes yes yes no yes GSN yes <120 yes yes yes no no GSN no <50 yes yes yes no yes 29

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34 Appendix D Images of algal species observed during the 11 th USGS NAWQA Workshop on Harmonization of Algal Taxonomy. 31

35 Figure D1. The cyanobacteria epiphyte Chaemosiphon incrustans, from the Provo River near Provo, UT (Great Salt Lake Basins Study Unit; USGS NAWQA sample ID GRSL0800ARE0226B). 32

36 Figure D2. The cyanobacteria epiphyte Chaemosiphon polonicus, from Halfway Brook at Barryville, NY (Delaware River Basins Study Unit; USGS NAWQA sample ID DELR0901ARE0247B). 33

37 Figure D3. The coccoid cyanobacterium, possibly Cyanodermatium or Pleurocapsa, from Yokum Creek near Qak Grove, AR (Ozark Plateau Study Unit; USGS NAWQA sample ID OZRK0994ADE0012B). 34

38 Figure D4. The cyanobacteria epiphyte Heteroleibleinia, from the Middle Fork of the Holston River at Seven Mile Ford, VA (Upper Tennessee River Basins Study Unit; USGS NAWQA sample ID UTEN0597ARE0023B). 35

39 Figure D5. The filamentous cyanobacteria Komvophorn, from the Service Creek above Dry Creek at Burlington, NC (Albemarle-Pamlico Drainages Study Unit; USGS NAWQA sample ID ALBE0503ARE0029BA). 36

40 Figure D6. The filamentous cyanobacteria Leptolyngbya, from Cub River near Richmond, UT (Great Salt Lake Basins Study Unit; USGS NAWQA sample ID GRSL0701ADE2108B). 37

41 Figure D7. The filamentous cyanobacteria Oscillatoria, from the Neversink River near Claryville, NY (Delaware River Basins Study Unit; USGS NAWQA sample ID DELR0801ADE0230B). 38

42 Figure D8. The filamentous cyanobacteria Oscillatoria, from the Tamiani Canal near Monroe, FL (Southern Florida Basins Study Unit; USGS NAWQA sample ID SOFL0697ARE0024B). 39

43 Figure D9. The filamentous cyanobacteria Phormidiochaete, from Yokum Creek near Qak Grove, AR (Ozark Plateau Study Unit; USGS NAWQA sample ID OZRK0994ADE0012B). 40

44 Figure D10. The colonial coccoid cyanobacteria Pleurocapsa minor, from Yokum Creek near Qak Grove, AR (Ozark Plateau Study Unit; USGS NAWQA sample ID OZRK0895ADE0008B). 41

45 Figure D11. The filamentous cyanobacteria from the Pseudanabaenaceae family, from from the Neversink River near Claryville, NY (Delaware River Basins Study Unit; USGS NAWQA sample ID DELR0801ADE0230B). 42

46 Figure D12. The colonial coccoid cyanobacteria Radaisia, from Yokum Creek near Qak Grove, AR (Ozark Plateau Study Unit; USGS NAWQA sample ID OZRK0895ARE0008B). 43

47 Figure D13. The cyanobacteria epiphyte Stichosiphon, from Old Mans Creek near Iowa City, IA (Eastern Iowa Basins Study Unit; USGS NAWQA sample ID EIWA0897ARE8820B). 44

48 Figure D14. The heterocytic cyanobacteria Stigonema mammilosum, from the Williams River at Dyer, WV (Kanawha-New River Basins Study Unit; USGS NAWQA sample ID KANA0697ARE0010B). 45

49 Figure D15. The filamentous cyanobacteria Symplocastrum penicillatum, from the Middle Fork of the Holston River at Seven Mile Ford, VA (Upper Tennessee River Basins Study Unit; USGS NAWQA sample ID UTEN0597ARE0023B). 46

50 47

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