2011 NORTHEASTERN NATURALIST 18(4):457–474
The Algal Flora of Acadia National Park, Maine
Melissa A. Vaccarino1, Jana Veselá2, and Jeffrey R. Johansen1,*
Abstract - The inland algal flora of Acadia National Park was studied based on over 200
samples from 119 localities. A total of 160 algal genera were found, representing 12 algal
phyla. Among these were 140 new generic records for the Park. The most frequently
encountered taxa were desmids and diatoms, and the genera and species in these groups
were acidophilous, reflecting the low pH of the ponds, lakes, and streams of the Park.
Ponds and streams were the most heavily sampled habitats in the Park, and coincidentally
had the highest diversity (103 and 91 genera, respectively). Based on other similar
studies, we estimate that the Park likely contains as many as 1000 algal species within its
boundaries, indicating that the diversity is exceptionally high for an area of this size.
Introduction
Acadia National Park is located on the Atlantic coast of Maine and was established
90 years ago as the first national park east of the Mississippi (Tree and
Oxnard 2003). It is composed of Mount Desert Island (MDI), Schoodic Peninsula,
Isle au Haut, and several small islands nearby. Although it is the sixth smallest
national park, it is one of the most popular in the USA, with about 3 million visitors
per year (Eugene et al. 1994). Mountains as well as deep valleys, seashores
with steep cliffs or sand beaches, numerous lakes and wetlands, barren mountain
tops, meadows, and deep forests all can be seen and enjoyed by the visitor.
The land where Acadia National Park is now situated was not inhabited by
European-Americans until the middle of the 19th century. At that time, the area
attracted the attention of several wealthy industrialists, such as the Rockefellers,
Vanderbilts, Carnegies, and Astors (Karr 2005, Mitchell 2005). They built more
than 200 massive summer “cottages” in and around Bar Harbor on MDI (Tree
and Oxnard 2003). George Dorr and Harvard president Charles Eliot, along with
others (e.g., John D. Rockefeller, Jr.), started to acquire the public land beginning
in 1901 (Kandell 2008). Eventually, this land was donated to the federal
government to preserve it. John D. Rockefeller, Jr. alone donated 4450 ha, which
is about one third of the present Park area (Tree and Oxnard 2003). In 1919, the
donated land was named Lafayette National Park (Fahey 2005) and was renamed
Acadia National Park in 1929 (Karr 2005).
The Park’s position on the coast, isolation from the mainland, great diversity
of habitats, and protection as a national park have all contributed to a rich fauna
and flora. There are a reported 219 bird species (Killion and Foulds 2007). The
Park also has 37 species of terrestrial mammals, 7 species of reptiles, 11 species
1Department of Biology, John Carroll University, 20700 North Park Boulevard, University
Heights, OH 44118. 2Botany Department, Faculty of Science, University of South
Bohemia, Na Zlaté stoce 1, České Budějovice, Czech Republic. *Corresponding author
- johansen@jcu.edu.
458 Northeastern Naturalist Vol. 18, No. 4
of amphibians (Bank et al. 2006, Crocker et al. 2007), 28 species of fish, 42
species of ants (Ouellette et al. 2010), and thousands of species of other invertebrates
(Killion and Foulds 2007, Stone et al. 2001). Migrating whales, dolphins,
and seabirds are also common in the area. There are 1135 vascular plant records
for the Park (Greene et al. 2005), and a rich marine macroalgal community is
known on the coast. An early work on the flora of Mount Desert Island reported
87 genera of algae, with 142 species and 6 varieties (Rand and Redfield 1894).
However, there is virtually no more recent knowledge of the diversity of algae
from freshwater and brackish habitats within the Park.
In summer of 2008, 119 localities within the Park (110 freshwater, 9 brackish
water) were sampled for algae. All algal genera in these samples were identified.
The purpose of this study is to give an initial report of the algal diversity in inland
aquatic habitats in Acadia National Park.
Field-site Description
Algal samples were collected from freshwater and brackish habitats within
Acadia National Park, ME, during 4–14 June 2008. Most localities were on
Mount Desert Island (109 sites). Eight sites on Schoodic Peninsula were sampled.
Two additional samples were collected from Isle au Haut in October 2008 by
Park naturalists. Altogether, 119 sites were sampled within the Park or from large
lakes bordering the Park (Fig. 1): 7 lakes, 41 ponds, 41 streams, 16 wetlands,
9 brackish sites, and 6 wet walls (samples from site 81 were divided into two
habitat types, brackish and wetland). In this study, a lake was defined as a large
pond with an area of >0.4 km2 (100 acres; usually much larger). A pond was a
water body with an area smaller than 0.4 km2 and usually with developed aquatic
vegetation along the shores. Wetland was a habitat with developed aquatic vegetation
and no sizeable open water. This category included marshes, bogs, forested
wetlands, etc. Brackish habitats were all sites with tidal influence and elevated
conductivity (i.e., ponds, wetlands, and streams with brackish water). A wet wall
was a rock face wetted by seasonal or perennial spring water with developed algal
assemblages. The sites were chosen using a topographic map with a 1:35000
scale (Acadia National Park Trail Map, National Geographic). Every water body
visible on the map was included in the study.
Methods
Samples of phytoplankton—periphyton, epilithon, and epipsammon—were
taken from the 119 sample localities, providing a total of over 200 samples
from the Park because many localities had multiple types of samples. Phytoplankton
was collected using a plankton net (35-μm mesh). Other algal
assemblages were collected using a toothbrush (from rocks, stones) or a spoon.
Pieces of wood or stones were also included in the samples. Algae were refrigerated
after collection and examined fresh for up to one week after sampling.
All samples were preserved in 2% glutaraldehyde within one week of sampling
and returned to the laboratory for further examination.
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 459
UTM coordinates (using Garmin GPSII plus) and several water characteristics
were recorded for each site (Veselá 2010). The temperature (ºC), conductivity
(μS/cm), and salinity (ppt) were measured using a YSI meter model 63. The pH
Figure 1. Sample sites within Acadia National Park, ME. All sites except site 51 were
confirmed to be inside of Park boundaries (delineated by shading). This map is a modifi-
cation of a tourist map prepared for the National Park Service and posted online at www.
nps.gov/acad/planyourvisit/upload/ACADmap2005.pdf.
460 Northeastern Naturalist Vol. 18, No. 4
measurements were taken using ColorpHast® pH-indicator strips with a resolution
of 0.1–0.2 pH units. The temperature, conductivity, and salinity were not
measured on wet walls due to measurement limitation of the YSI meter.
Algae were examined and photographed using Zeiss Axioskop and Olympus
BX60 photomicroscopes with the highest resolution planapochromatic Nomarski
DIC optics available from those manufacturers for 40X, 50X, 60X, 63X, and
100X objectives. Taxa were identified following the most modern taxonomic
texts available for each group, including but not limited to: Bourrelly (1988,
1990), Croasdale et al. (1983), Ettl and Gärtner (1995), Irenee-Marie (1939),
Komárek and Anagnostidis (1999, 2005), Komárek and Fott (1983), Prescott
(1951), Prescott et al. (1972, 1975, 1977, 1981, 1982), Round et al. (1990), Starmach
(1972), Wehr and Sheath (2003), and Wołowski and Hindák (2005). Higher
level classification follows Johansen et al. (2007).
Results
A total of 160 algal genera were recovered from Acadia National Park (Appendix
1), representing a diversity of algal phyla: Cyanobacteria (29 genera;
Figs. 2–4), Charophyta (28 genera; Figs. 5–8), Chlorophyta (39 genera; Figs.
9–10, 15–16, 20), Rhodophyta (2 genera; Fig. 14), Cryptophyta (1 genus), Dinophyta
(5 genera, Fig. 21), Euglenophyta (4 genera; Fig. 12), Bacillariophyta (39
genera; Fig. 17), Chrysophyta (4 genera; Fig. 18), Raphidophyta (1 genus), Synurophyta
(3 genera; Fig. 13), and Tribophyta (5 genera; Fig. 19). The dominant
phyla were Charophyta and Bacillariophyta, particularly acidophilous taxa such
as the desmids and Eunotiophycidae. This dominance is not surprising given the
acidic nature of nearly all waters in the Park (Table 1). The ten most frequently
encountered genera were (in order of decreasing frequency): Tabellaria, Mougeotia,
Staurastrum, Eunotia, Closterium, Dinobryon, Cosmarium, Peridinium,
Euastrum, and Pinnularia.
The most diverse habitats were the ponds, with 103 different genera reported.
Streams were a close second, with 91 taxa. These results were expected, as these
habitats were the most intensively sampled (Appendix 1). Indeed, the number of
Table 1. Ranges and means (with standard errors) of four measured water characteristics for individual
freshwater categories, all freshwater sites, and all brackish sites. Number of replicates
within each site category is indicated in parentheses.
Site category pH T (°C) Conductivity [μS/cm] Salinity [ppt]
Lakes (7) 4.9 ± 0.0 (7) 18.2 ± 0.7 (7) 39.0 ± 4.2 (7) 0.0 ± 0.0 (7)
Ponds (41) 4.9 ± 0.1 (41) 20.4 ± 0.5 (41) 73.0 ± 19.9 (40) 0.0 ± 0.0 (40)
Streams (41) 5.0 ± 0.1 (41) 14.4 ± 0.7 (40) 57.0 ± 8.8 (38) 0.0 ± 0.0 (38)
Wetlands (16) 4.9 ± 0.2 (16) 18.6 ± 1.2 (16) 65.4 ± 15.2 (15) 0.0 ± 0.0 (15)
Wet walls (6) 4.8 ± 0.2 (5) NA NA NA
All freshwater sites (111) 4.9 ± 0.0 (110) 17.7 ± 0.5 (104) 63.4 ± 8.9 (100) 0.0 ± 0.0 (100)
Range measured FS 4.2–6.8 9.0–26.2 9.0–824 0.0–0.5
Brackish (9) 6.9 ± 0.3 (9) 18.3 ± 0.9 (9) 30,000 ± 4700 (5) 21.0 ± 3.2 (5)
Range measured BS 5.4–8.1 14.9–20.9 13,630–41,730 8.5–26.8
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 461
genera from the different habitats is directly proportional to the number of samples
taken: ponds (103 genera/41 samples), streams (91/41), wetlands (61/16),
lakes (48/7), brackish waters (47/9), and wet walls (27/6).
A number of species were identified in species-poor genera. When a
genus was represented by only a single species, we reported that species
(Appendix 1). However, more species than those listed were identified. Some
of the more unusual and rare species included: Chaetophora incrassata
(Fig. 15), Draparnaldia platyzonata (Fig. 16), Binuclearia trana (Fig. 20),
Ducellieria chodatii (Fig. 19), Peridinium limbatum (Fig. 21), Scytonema coactile
(Fig. 4), and Stigonema multipartitum.
Discussion
Prior to this study, an examination of freshwater algae restricted to Acadia
National Park boundaries had not been completed. However, an early work,
The Flora of Mount Desert Island (Rand and Redfield 1894) did list algae in
addition to lichens, mosses, liverworts, and vascular plants. Rand and Redfield
reported 87 algal genera, with 148 infrageneric taxa. Their list included
mostly marine species of algae, with only 20 clearly freshwater or brackish
species in 17 genera. Some overlap between their list and our list exists, notably
Nitella, Batrachospermum, Spirogyra, Vaucheria, Cladophora, Ulothrix,
Rhizoclonium, Enteromorpha, Calothrix, Stigonema, Lyngbya, Phormidium,
Oscillatoria, Chroococcus, and Gloeocapsa. We lacked the freshwater reds
Lemanea and Tuomeya which they saw, although we have seen these genera
in the region outside of the Park. Later additions to the flora by Taylor (1921)
included identifications of eleven freshwater green algae whose genera was not
previously reported: Mougeotia genuflexa (Dillwyn) Agardh, Botryococcus
braunii Kützing, Tetraspora lubrica (Roth) Agardh, Chlorococcum petrocelidis
(Nägeli) Rabenbhorst, Zoochlorella parasitica Brandt, Ankistrodesmus falcatus
(Corda) Ralfs, Kirchneriella contorta (Schmidle) Bohlin, Scenedesmus (3 spp.),
Chaetosphaeridium pringsheimii Klebahn, Draparnaldia glomerata (Vaucher)
Agardh, and Herposteiron vermiculoides (Wolle) Collins. Of these eleven, we
observed six. Thus, only 7 genera additional to what we found have been reported
from the Park (actually, Mount Desert Island), and this report has at least
140 new generic records for the Park.
The algal flora of Maine was mostly studied by Harvey and West in the late
1800s. The majority of samples in Harvey (1888) came from around the Penobscot
River, mostly near Orono, and included sampling sites of small ponds,
streams, and wells. Harvey (1889) included algae from Augusta and Moosehead
Lake, and the final work in the series (Harvey 1892) included samples mostly
from Pushaw Stream. In total, he described 190 species from 56 genera. He
also mentioned several other genera not able to be described to species either
due to seasonality or lack of study (e.g., Oscillatoria, Ulothrix). Many species
were identified for those desmids displaying great diversity (Staurastrum: 30
spp., Cosmarium: 28 sp., Closterium: 16 spp., Euastrum: 13 spp., Micrasterias:
7 spp.), and the desmidps collectively account for more than half of species
462 Northeastern Naturalist Vol. 18, No. 4
Figures 2–13. Representative taxa from Acadia National Park. 2. Eucapsis minor (scale
= 10 μm). 3. Symplocastrum friesii (scale = 10 μm). 4. Scytonema coactile (scale = 20
μm). 5. Bambusina borreri (scale = 10 μm). 6. Closterium kuetzingii (scale = 50 μm).
7. Micrasterias muricata (scale = 10 μm). 8. Pleurotaenium verrucosum (scale = 20 μm).
9. Zygnema sp. (scale = 50 μm). 10. Oophila ambylostomalis (scale = 10 μm). 11. Microspora
sp. (scale = 10 μm). 12. Phacus longicauda (scale = 10 μm). 13. Chrysosphaerella
longispina (scale = 10 μm).
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 463
Figures 14–21. Representative taxa from Acadia National Park. 14. Batrachospermum sp.
(scale = 50 μm). 15. Chaetophora incrassata (scale = 50 μm). 16. Draparnaldia platyzonata
(scale = 50 μm). 17. Neidium sp. (scale = 20 μm). 18. Hydrurus foetidus (scale = 50
μm). 19. Ducellieria chodatii (scale = 10 μm). 20. Binuclearia tatrana (scale = 10 μm).
21. Peridinium limbatum (scale = 10 μm).
464 Northeastern Naturalist Vol. 18, No. 4
described. West continued Harvey’s first flora with a more extensive listing of the
desmids from Orono, identifying 73 additional species and 5 varieties and forms
(West 1888). He later produced more findings (West 1891) in which he described
additional desmid species, some new to the USA and some new to science, and
also included a small listing of diatoms (in 15 genera). Other floristic reports of
the soft algae in Maine since this time have been sporadic (Colt 1995, Wheldon
1943), although ecological studies have recorded the appearance of some taxa
(Bailey et al. 1979, Norton et al. 1981). In an unpublished summary of algal
records for New England, Colt (1999a, b) summarized the genus and species records
for Maine. According to this record, and the records in Prescott et al. (1972,
1975, 1977, 1981, 1982) and Croasdale et al. (1983), two of our reports represent
new desmid genus records for Maine, Mesotaenium and Phymatodocis.
The diatoms of Maine have been more recently reported from the Casco Bay
watershed (Wang et al. 2006) and Maine as part of larger studies in the northeastern
United States (Charles et al. 2006, Davis and Anderson 1985). Davis and Anderson
(1985) reported finding a total of 455 taxa from New England samples (25 sites in
Maine, 6 sites in Vermont and New Hampshire). However, only 152 taxa were actually
listed in their paper. Common species reported include Cyclotella glomerata
Bachmann, C. stelligera (Cleve et Grunow) Van Heurck, Eunotia spp., Frustrulia
rhomboides Ehrenberg (De Toni), Navicula spp., Neidium spp., and Pinnularia
spp. Less abundant (less than 5%) but commonly encountered species seen were in Synedra,
Achnanthes, Eunotia, Navicula, Neidium, and Pinnularia. Wang et al. (2006)
recorded appearances of Nitzschia, Synedra, Surirella, Neidium, Eunotia, and
Pinnularia, where the percentage of Synedra, Nitzschia, Neidium, and Eunotia
were highly correlated to human disturbance. Charles et al. (2006) reported a
high appearance of the following species in the northeast (mostly Pennsylvania,
New Jersey, Maryland, and Maine): Achnanthes lanceolata (Brébisson) Grunow,
A. minutissima Kützing, Cocconeis placentula var. euglypta Ehrenberg, Cyclotella
meneghiniana Kützing, Cymbella minuta Hilse, Fragilaria vaucheriae
(Kützing) Petersen, Gomphonema parvulum (Kützing) Kützing, Melosira varians
Agardh, Navicula cryptocephala Kützing, N. minima Grunow, N. pupula Kützing,
N. rhynchocephala Kützing, N. salinarum var. intermedia (Grunow) Cleve,
Nitzschia amphibia Grunow, N. frustulum (Kützing) Grunow, N. kuetzingiana
Hilse, N. palea (Kützing) W. Smith, Surirella angusta Kützing, and Rhoicosphenia
curvata (Kützing) Grunow. These three studies were focused on diatoms as metrics,
and although they provided biogeographic information about some taxa, the
rare and unusual taxa were generally not reported or discussed. All of the genera
reported in these studies are reported in this study.
The algal flora of Acadia National Park appears to be very diverse given its
relatively small area and the narrow limits of pH and conductivity (Table 1).
Since species were not determined, it is difficult to make comparisons to other
floras of comparable geographic size from which species were identified. Still,
such comparisons do show that the flora of the Park is likely very diverse.
The algal flora of the Great Smoky Mountains National Park has been well
studied and is a worthy comparison. The Great Smoky Mountains has fewer
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 465
lentic habitats, but more lotic habitats and wet walls than Acadia. In the list
of taxa from aquatic habitats in that park, we found representatives from all
phyla in Acadia, with Prymnesiophyta, Eustigmatophyta, Phaeophyceae, and
Cyanidiophyta being additionally present in the Great Smoky Mountains (Johansen
et al. 2005, 2007). There were 1020 species in 265 genera (Johansen
et al. 2007, Khaybullina et al. 2010). The waters of Great Smoky Mountains
National Park were also quite acidic (Furey et al. 2009, Gomez et al. 2003), and
were similarly rich in Eunotia, Frustulia, and diverse desmids (Fučíková et al.
2008). One difference in the flora of the two parks is that at least some soils in
the Great Smoky Mountains have been studied, and they contribute 14 genera
and 23 species to the totals listed above (Khaybullina et al. 2010). The Great
Smoky Mountains flora differs in level of effort as well, with diatoms being
examined in cleaned and prepared slides in that park. We did not include diatoms
in prepared samples from Acadia National Park in this report. However, in
another study of these same samples, a total of 87 diatom genera were observed
(Veselá 2010), with 47 genera being found in that study that were not present in
this one. If those numbers are included, there is a total of 207 algal genera in the
Park. We conclude that if the effort of study in Acadia National Park were equal
to that invested in the Great Smoky Mountains National Park, there would likely
be a similar number of algal genera, even though the Park is much smaller
than Great Smoky Mountains National Park. Acadia National Park does have a
brackish water component not present in the Great Smoky Mountains, and there
were a total of 15 genera restricted to this habitat.
Navajo National Monument was studied with respect to its algal flora, and was
much less diverse, with 137 taxa in 30 genera (Johansen et al. 1983). This park is
much smaller than either Acadia or Great Smoky Mountains, and is additionally
situated in semi-arid shrub steppe habitat.
In the coastal region of Orissa State, in India, a survey of the algal flora yielded
307 species in 87 genera (Adhikary et al. 2009). This region was very different
from either park studied in the United States in that well-inhabited regions with
anthropogenically impacted waters were studied. Adhikary et al. (2009) did not
include diatoms in their flora, so it may be that diversity of this region would be
similar if that component were added. Acadia National Park had 121 non-diatom
genera, which is still high by comparison to this tropical site.
It is interesting to note that in those studies discussed above for which both
species and genera were determined, there are typically 3–4 times as many
species as genera. If this rough estimate held true in Acadia, we could expect
600–800 species to be present (if we base this on the count of 207 genera).
Preliminary results indicate that the diatom flora alone will consist of more
than 500 species, so it seems possible that there may be around 1000 species in
Acadia National Park.
The number of species found in any given geographic area is highly dependent
upon effort. The more samples that are examined, the more rare taxa
are more likely to be found. Even areas with more limited variation in water
chemistry and climate can have high numbers of taxa if sufficient samples are
466 Northeastern Naturalist Vol. 18, No. 4
studied. We hope further work will be conducted on the algal flora of Acadia
National Park. This study has shown that this protected area is home to a diverse
and interesting algal flora.
Acknowledgments
Funding for this project was funded in part by the Grant Agency of University of
South Bohemia GAJU 138/2010/P (Czech Republic). Collection efforts were funded by
an L.L. Bean Acadia Research Fellowship to Jana Veselá and Jeff Johansen. We thank
Acadia National Park for use of researcher housing during the collection effort. The
Humboldt Research Institute at Eagle Hill also provided some travel support to the authors
in conjunction with a seminar taught at the institute concurrently with collection
efforts. Markéta Krautová and Jan Pilný assisted in field collections.
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470 Northeastern Naturalist Vol. 18, No. 4
Appendix 1. Freshwater and brackish algal genera (and species) observed in Acadia National
Park. Frequencies of occurrence (in %) are divided into 6 categories: L = Lakes,
P = Ponds, S = Streams, W = Wetlands, WW = Wet walls, and B = Brackish; numbers in
parentheses indicate the number of replicates within each habitat type.
Taxon L(7) P(41) S(41) W(16) WW(6) B(9)
CYANOBACTERIA
Synechococcophycidae
Aphanocapsa Nägeli 29 0 0 0 0 0
Eucapsis minor (Skuja) Elenkin 0 2 0 0 0 0
Geitlerinema (Anagnostidis et Komárek) 0 15 5 0 0 0
Anagnostidis
Heteroleibleinia pusilla (Hansgirg) 0 2 0 0 0 11
Anagnostidis et Komárek
Leptolyngbya Anagnostidis et Komárek 0 2 2 0 0 22
Merismopedia Meyen 0 24 2 19 0 0
Pseudanabaena Lauterborn 0 2 0 0 0 0
Rhabdoderma Schmidle et Lauterborn 0 5 0 0 0 0
Rhabdogloea Schröder 0 2 0 0 0 0
Snowella Elenkin 0 2 0 0 0 0
Oscillatoriophycidae
Chroococcus Nägeli 0 24 0 13 0 33
Gloeocapsa Kützing 0 0 0 0 17 0
Gloeocapsopsis magma (Brébisson) 0 2 7 0 33 0
Komárek et Anagnostidis
Gomphosphaeria Kützing 0 0 0 0 0 11
Lyngbya Agardh ex Gomont 0 0 2 0 17 11
Microcystis Kützing 0 2 0 0 0 0
Oscillatoria Vaucher ex Gomont 14 32 10 6 33 11
Phormidium Kützing ex Gomont 0 15 10 0 0 11
Symplocastrum (Gomont) Kirchner ex 0 0 0 6 0 0
Engler et Prantl
Trichodesmium lacustre Klebahn 0 5 0 0 0 0
Nostocophycidae
Anabaena Bory 14 5 2 0 0 11
Calothrix Agardh 0 5 5 0 17 11
Cylindrospermum Kützing 0 29 7 6 0 11
Fischerella Gomont 0 17 2 13 0 0
Hapalosiphon Nägeli in Kützing 0 5 0 0 0 0
Nostoc Vaucher 0 0 0 0 0 11
Scytonema Agardh 14 2 10 0 17 0
Stigonema Agardh 43 10 22 0 33 0
Tolypothrix Kützing 0 10 2 0 0 11
RHODOPHYTA
Batrachospermum Roth 14 10 10 0 0 0
Polysiphonia Greville 0 0 0 0 0 11
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 471
Taxon L(7) P(41) S(41) W(16) WW(6) B(9)
CHAROPHYTA
Charophyceae
Chara L. 0 5 0 0 0 0
Nitella (Agardh) Leonhardi 0 2 2 6 0 0
Klebsormidiophyceae
Klebsormidium Silva, Mattox et Blackwell 0 0 5 6 33 0
Zygnemophyceae
Actinotaenium (Nägeli) Teiling 0 0 0 0 17 0
Bambusina Kützing 14 37 5 0 0 0
Closterium Nitzsch 14 76 24 31 0 11
Cosmarium Corda ex Ralfs 29 63 24 19 0 11
Cylindrocystis Meneghini 29 22 12 19 17 0
Desmidium Agardh 14 27 2 0 0 0
Docidium Brébisson 0 5 0 6 0 0
Euastrum Ehrenberg ex Ralfs 14 63 10 38 17 0
Gonatozygon De Bary 0 2 0 0 0 0
Hyalotheca Ehrenberg 14 22 7 13 0 0
Mesotaenium Nägeli 0 0 2 0 0 0
Micrasterias Agardh ex Ralfs 14 22 0 13 0 0
Mougeotia (Agardh) Wittrock 86 78 32 31 0 11
Netrium (Nägeli) Itzigson et Rothe 0 7 0 0 17 0
Penium Brébisson 14 12 12 0 0 0
Phymatodocis Nordstedt 0 5 0 0 0 0
Pleurotaenium Nägeli 14 37 5 25 0 0
Spirogyra Link 43 29 7 19 0 11
Spondylosium Brébisson 29 46 2 13 0 0
Staurastrum Meyen 43 78 15 44 17 0
Staurodesmus Teiling 29 46 2 19 0 0
Tetmemorus Ralfs 0 10 0 0 0 0
Triploceras J.W. Bailey 0 15 0 0 0 0
Xanthidium Ehrenberg 14 34 2 6 0 0
Zygnema Agardh 71 27 10 6 0 0
CHLOROPHYTA
Chlorophyceae
Ankistrodesmus Corda 0 24 5 6 0 0
Bulbochaete Agardh 43 49 2 31 0 0
Carteria Diesing 0 0 2 0 0 0
Chaetophora incrassata Hazen 0 0 5 0 0 0
Characium A. Braun in Kützing 0 0 2 0 0 0
Chlamydomonas Ehrenberg 29 2 0 6 0 0
Coccomyxa subglobosa Pascher 0 0 0 0 17 0
Coenochloris sphagnicola Hindák 0 0 0 0 0 11
Dictyosphaerium Nägeli 0 24 0 0 0 0
Dimorphococcus A. Braun 0 7 0 0 0 0
Draparnaldia Bory 14 0 2 6 0 0
472 Northeastern Naturalist Vol. 18, No. 4
Taxon L(7) P(41) S(41) W(16) WW(6) B(9)
Eudorina Ehrenberg 0 2 0 0 0 0
Gloeocystis Nägeli 0 0 0 6 33 0
Microspora Thuret 0 10 10 13 0 0
Oedogonium Link 14 41 7 19 0 11
Palmodictyon viride Kützing 0 0 2 0 0 0
Pediastrum Meyen 0 17 2 0 0 0
Planktosphaeria G.M. Smith 14 0 0 0 0 0
Quadrigula Printz 0 2 0 0 0 0
Scenedesmus Meyen 0 37 7 0 17 11
Schizochlamys A. Braun 0 0 2 0 0 0
Stigeoclonium Kützing 14 5 2 0 0 0
Tetraedon minimum (A. Braun) Hansgirg 0 0 0 6 17 0
Ulothrix Kützing 0 7 32 6 0 22
Westella linearis G.M. Smith 0 0 2 0 0 0
Trebouxiophyceae
Botryococcus Kützing 14 2 0 0 0 0
Kirchneriella Schmidle 0 5 0 0 0 0
Microthamnion Nägeli 0 2 0 0 0 0
Monoraphidium Komárková 0 0 2 0 0 0
Oocystis Nägeli 0 2 2 6 0 11
Oophila ambylostomalis Lambert 0 2 0 0 0 0
Selenastrum Reinsch 0 2 0 0 0 0
Ulvophyceae
Binuclearia tatrana Wittrock 0 5 2 6 17 0
Cladophora Kützing 0 0 2 0 0 11
Enteromorpha intestinalis (L.) Ness 0 0 0 0 0 44
Geminella Turpin 14 0 0 0 0 0
Gloeotila Kützing 0 0 2 0 0 0
Rhizoclonium Kützing 0 0 0 0 0 11
CRYPTOPHYTA
Cryptomonas Ehrenberg 29 32 20 25 0 11
DINOPHYTA
Ceratium Schrank 14 5 0 0 0 0
Gymnodinium Baumeister 0 0 0 0 0 11
Katodinium Fott 0 2 2 6 0 0
Peridinium Ehrenberg 57 51 2 19 17 0
Rufusiella insignis (Hassall) Loeblich 0 0 0 0 17 0
EUGLENOPHYTA
Euglena Ehrenberg 0 15 10 19 17 0
Ophiocytium Nägeli 0 7 0 0 0 0
Phacus Dujardin 0 7 0 6 0 0
Trachelomonas Ehrenberg 0 24 5 31 0 0
2011 M.A. Vaccarino, J. Veselá, and J.R. Johansen 473
Taxon L(7) P(41) S(41) W(16) WW(6) B(9)
BACILLARIOPHYTA
Achnanthidium Kützing 0 2 2 0 0 0
Actinella punctata Lewis 0 10 5 0 0 0
Amphora Ehrenberg ex Kützing 0 0 0 0 0 11
Asterionella formosa Hassall 14 5 2 0 0 0
Aulacoseira Thwaites 0 5 0 0 0 0
Berkeleya Greville 0 0 0 0 0 11
Cavinula Mann et Stickle 0 0 2 0 0 0
Chaetoceros Ehrenberg 0 0 0 0 0 11
Cosmioneis Mann et Stickle 0 0 0 0 17 0
Craticula Grunow 0 0 0 6 0 0
Ctenophora (Grunow) Williams et Round 0 0 0 0 0 11
Cylindrotheca Rabenhorst 0 27 0 0 0 22
Cymbopleura (Krammer) Krammer 0 0 2 13 0 0
Diploneis Ehrenberg ex Cleve 0 0 0 0 0 11
Encyonema Kützing 0 0 2 0 0 0
Entomoneis Ehrenberg 0 0 0 0 0 11
Eunotia Ehrenberg 43 41 34 44 33 0
Fragilaria Lyngbye 0 0 2 6 0 11
Fragilariaforma (Ralfs) Williams et Round 0 0 2 0 0 0
Frustulia Rabenhorst 14 46 15 44 0 0
Gomphonema Ehrenberg 14 15 15 13 0 11
Gyrosigma Hassall 0 2 0 13 0 11
Hantzschia Grunow 0 2 0 0 0 0
Melosira Agardh 14 2 5 0 0 22
Meridion Agardh 0 0 2 0 0 0
Navicula Bory 0 10 12 6 0 22
Neidium Pfitzer 0 12 5 13 0 11
Nitzschia Hassall 14 17 10 25 0 33
Pinnularia Ehrenberg 0 44 20 44 17 11
Rossithidium Round et Buhtkiyarova 0 0 2 0 0 0
Sellaphora Mereschkowsky 0 0 5 0 0 0
Stauroneis Ehrenberg 0 2 2 6 0 0
Staurosirella leptostauron (Ehrenberg) 0 0 2 0 0 0
Williams et Round
Stenopterobia Brébisson ex Van Heurck 43 39 7 6 0 11
Surirella Turpin 29 17 2 13 0 11
Synedra Ehrenberg 14 10 10 19 0 33
Tabellaria Ehrenberg 100 88 68 50 0 0
Tryblionella W. Smith 0 0 0 0 0 11
Urosolenia Round et Crawford 14 0 0 0 0 0
CHRYSOPHYTA
Chrysosphaera Pascher 0 0 5 0 0 0
Dinobryon Ehrenberg 71 49 7 19 0 0
Hydrurus foetidus (Villars) Trevisan 0 0 2 0 0 0
Phaeosphaera W. et G.S. West 0 0 5 0 0 0
474 Northeastern Naturalist Vol. 18, No. 4
Taxon L(7) P(41) S(41) W(16) WW(6) B(9)
RAPHIDIOPHYTA
Gonyostomum semen (Ehrenberg) Diesing 0 0 2 0 0 0
SYNUROPHYTA
Chrysosphaerella longispina Lauterborn 0 2 0 0 0 0
Mallomonas Perty 14 7 0 6 0 0
Synura Ehrenberg 0 5 0 6 0 0
TRIBOPHYTA
Bumilleria Borzi 0 0 0 0 17 0
Ducellieria Teiling 0 10 5 6 0 0
Tribonema Derbés et Solier 14 2 10 25 50 0
Vaucheria De Candolle 0 0 0 0 0 11
Xanthonema P.C. Silva 0 0 0 0 33 0