1066

Historical versus contemporary measures of

seaweed biodiversity in the Bay of Fundy
Colin R. Bates, Gary W. Saunders, and Thierry Chopin

Abstract: Anthropogenic changes in many components of the Bay of Fundy biota have recently been reported, however a
detailed assessment of benthic seaweed biodiversity has not been performed since the late 1970s. Here, we summarize 7
years (20002006) of temporal and spatial variation in species richness, composition, and percent cover of intertidal sea-
weeds in the outer Bay of Fundy, New Brunswick, Canada. In doing so, we provide up-to-date baseline records using re-
peatable and well-documented protocols. We document the presence of 72 species of seaweeds across four sites, and
demonstrate that the biota at one of our sites is indicative of a degraded assemblage: large patches of unutilized substrate,
low algal richness and abundance, with assemblages composed of hardy algal species and grazing and filter feeding inver-
tebrates. Additionally, we compare these contemporary data with historical presenceabsence records compiled from herba-
rium collections (19541978) and site surveys from 1963 to 1964 and 1978. Comparisons reveal 16 species not relocated
in contemporary surveys. However, conclusions of anthropogenic changes in seaweed biodiversity are difficult to assess in
the absence of high quality baseline information, and seaweeds historically found but not relocated in current sampling are
easily misidentified, subtidal, or rare. We discuss the nature and value of baseline data, and make suggestions for future
research.
Key words: biodiversity, change, historical data, intertidal, monitoring, seaweed.
Resume : On a recemment rapporte des modifications anthropogenes du biote de la Baie de Fundy, cependant aucune eva-
luation de la biodiversite des algues marines benthiques na ete realisee depuis la fin des annees 1970. Les auteurs resu-
ment sept annees (20002006) de variation spatio-temporelle dans la richesse en especes, la composition et le pourcentage
de couverture des algues marines de la zone intertidale, dans la partie externe de la Baie de Fundy du Nouveau-Brunswick
au Canada. Ceci a permis aux auteurs detablir une ligne de base de donnees en utilisant des protocoles reproductibles et
bien documentes. Ils font etat de 72 especes dalgues marines reparties sur quatre sites, et demontrent que le biote dun
des quatre sites manifeste un assemblage en degradation, soient de grandes surfaces de substrat non utilisees avec faibles
abondance et richesse en algues, constituees despeces dalgues rustiques habitees par des invertebres brouteurs et filtreurs.
De plus, les auteurs comparent ces donnees contemporaines avec les mentions historiques de presence/absence obtenues a
partir de collections dherbiers (19541978) et des suivis de sites de 19631964 et 1978. Ces comparaisons revelent que
16 especes nont pas ete retrouvees dans les suivis contemporains. Tout de meme, il reste difficile de tirer des conclusions
sur les modifications anthropogenes dans la diversite des algues marines en absence dinformations de base de qualite, et
il se peut que les algues historiquement rapportees, mais non retrouvees dans les echantillonnages actuels, puissent avoir
ete mal identifiees, rares, ou encore appartenir a la zone subtidale. Les auteurs discutent la nature et la valeur des donnees
de bases et presentent des suggestions pour les recherches futures.
Mots-cles : biodiversite, change, donnees historiques, zone intertidale, suivi, algues marines.
[Traduit par la Redaction]

Introduction the world. With a tidal amplitude of up to 19 m, the largest

The Bay of Fundy is one of the marine natural wonders of
Received 7 November 2008. Published on the NRC Research
Press Web site at botany.nrc.ca on 3 November 2009.
C.R. Bates1,2 and G.W. Saunders. Centre for Environmental
and Molecular Algal Research, Department of Biology,
University of New Brunswick, Fredericton, NB E3B 6E1,
Canada.
T. Chopin. Centre for Coastal Studies and Aquaculture,
Department of Biology, University of New Brunswick, P.O. Box
5050, Saint John, NB E2L 4L5, Canada.
1Corresponding author (e-mail: colinba@interchange.ubc.ca).
2Presentaddress: Department of Botany, University of British
Columbia & Bamfield Marine Sciences Centre, 3529-6270
University Boulevard, Vancouver, BC V6T 1Z4, Canada.
in the world, this ecosystem offers many opportunities for
use by humans. Indeed, over the past 200 years the array of
anthropogenic pressures on the Bay of Fundy has greatly di-
versified and intensified (Lotze and Milewski 2004), partic-
ularly in the Passamaquoddy Bay region (Fig. 1). Examples
of anthropogenic pressures include increased physical dis-
turbance from coastal development, addition of chemical
contaminants, addition of nutrients and silt through finfish
aquaculture and agricultural runoff, and an increase in both
the amount and variety of biological resources harvested
(Percy and Wells 1996; Lotze and Milewski 2004).
Long-term changes in the abundance and diversity of fish,
mammals, birds, invertebrates, and plankton in the Bay of
Fundy are relatively well documented (Lotze and Milewski
2004). However, gauging of human impacts on seaweeds in
the Bay of Fundy is precluded by a lack of detailed baseline

Botany 87: 10661076 (2009) doi:10.1139/B09-067 Published by NRC Research Press

Bates et al.
Fig. 1. Map of contemporary and historical survey sites along the New Brunswick coast of the outer Bay of Fundy. Lepreau, Grand Manan,
Letite, and Meadow Cove are contemporary monitoring sites, and we make comparisons with herbarium collections (19541978) and sur-
veys taken at South Beach Head (1978), Point Lepreau (19631964), and Letite (1978). Inset box shows eastern Canada.
information (Lotze and Milewski 2004). Seaweeds, which
are the benthic marine macroalgae, are integral components
of coastal ecosystems because as primary producers they
support higher trophic levels by contributing energy to food
webs (Gollety et al. 2008). Additionally, seaweeds provide
three-dimensional structure to nearshore environments and
thus provide habitats for fish (Levin and Hay 1996) and in-
vertebrates (Bates and DeWreede 2007). However, despite
their importance, seaweeds are rarely considered in coastal
management and conservation plans (Norton et al. 1996).
These oversights, likely due to a lack of current information,
could lead to erroneous conclusions about energetic budgets,
productivity, nutrient cycling, and the efficacy of conserva-
tion measures in the Bay of Fundy.
Historical records of seaweed diversity in the Bay of
Fundy tend to be vague, generally occurring in the form of
checklists (MacFarlane and Milligan 1965; Stone et al.
1970; South 1984) or species lists from a poorly defined
geographic area (Hay 1887; Fowler 1902; Klugh 1917; Col-
invaux 1966, 1970; Wilson et al. 1979). Most studies are of
short temporal scope (Colinvaux 1970) or concentrate only
on the economically important species (Neish 1973; McKay
1975). These studies lack details about exact sampling loca-
tions and the specific methods used for data collection, pre-
venting use of the same methods in a contemporary study.
Available historical information lacks an abundance compo-
nent, yet many observed changes in intertidal assemblages
involve shifts in abundance of constituent species (Barry et
al. 1995).
There is one available study with potential value as sea-
1067
weed biodiversity baseline data. Smith et al. (1978) summa-
rize macroalgal herbarium collections spanning 19541978,
(stored at the University of New Brunswick), as well as sev-
eral seaweed surveys at sites along the New Brunswick
coast of the Bay of Fundy. Although suffering from many
of the above-described shortcomings, this study is to our
knowledge the most detailed historical information available
about Fundy seaweeds.
In the absence of comprehensive and current records of
seaweed communities in the Bay of Fundy, there is little
context to directly assess the flora for changes (e.g., recov-
ery) induced by chronic stressors (e.g. pollution, eutrophica-
tion); introduced species such as Codium fragile (Sur.)
Hariot subsp. tomentosoides (van Goor) Silva, which is cur-
rently invasive along the Atlantic coast of North America
(Levin et al. 2002); or pulse stressors such as oil spills
(Paine et al. 1996). Several short term field observations in
the Bay of Fundy (Worm 2000; Bates et al. 2001, 2005)
suggest that seaweed assemblages adjacent to eutrophied
sites are characterized by reduced richness (number of sea-
weed species) and abundance, and compositional shifts
from longer-lived perennial algal species to assemblages
composed of ephemeral annual algae and herbivorous or
filter-feeding invertebrates. While this type of assemblage is
consistent with impacted sites seen in the northeast Atlantic
(Kautsky et al. 1986; Schramm and Nienhuis 1996; Scott
and Tittley 1998; Ducrotoy 1999), these observations are
from short-term sampling events, and may not reflect the
prevailing status of marine algae in the Bay of Fundy.
In response to these concerns, in May 2000 we initiated a
Published by NRC Research Press
1068
Table 1. Descriptive data for four New Brunswick long-term seaweed biodiversity monitoring sites.
Site Upper right GPS Upper left GPS
Grand Manan 44.62433 N, 44.62449 N,
66.86006 W 66.85977 W
Lepreau 45.07312 N, 45.07286 N,
66.47020 W 66.47024 W
Letite Point 45.03811 N, 45.03818 N,
66.89126 W 66.89091 W
Meadow Cove 45.10397 N, 45.10413 N,
66.38308 W 66.38300 W
Note: Global positioning system (GPS) coordinates designate the upper corners of our 30 m sampling sites. Upper boundary of sampling zone
(as dictated by the lower boundary of the Ascophyllum zone) is listed in metres above chart datum. Fetch calculations and wave exposure classifi-
cations according to Boizard and DeWreede (2006).
seaweed-biodiversity-monitoring program at several interti-
dal sites along the New Brunswick coast of the outer Bay
of Fundy (hereinafter referred to as the outer Bay of
Fundy). By applying well-defined protocols to routine
monitoring of species richness, composition, and percent
cover within distinct geographic locations, our primary goal
was to establish a detailed long-term baseline data set for
outer Bay of Fundy seaweeds. The focus of this current re-
port is to document the temporal and spatial dynamics of
seaweeds in the outer Bay of Fundy since that time. Also,
we compare these contemporary data with available histori-
cal records (Smith et al. 1978) to look for evidence of floris-
tic change. Specifically, we ask the following questions: (i)
How does seaweed biodiversity vary across sites in southern
coastal New Brunswick? (ii) Is there evidence of within-site
changes in richness or composition over the past seven
years? (iii) Have long-term changes in seaweed species rich-
ness and composition taken place in the outer Bay of Fundy
since the late 1970s?
Materials and methods
Sampling was initiated in the summer of 2000, after
which we annually sampled seaweeds at three sites along
the New Brunswick side of the outer Bay of Fundy (Fig. 1):
Letite Point, Lepreau, and Grand Manan Island. In 2003 we
added a fourth site at Meadow Cove. In 2006, we were not
able to access Meadow Cove for abundance sampling owing
to stormy conditions.
Each site is delimited by a 30 m horizontal stretch of
shore, and bound at the top and bottom by the extent of the
algae and the waterline at chart datum, respectively. Global
positioning system coordinates for the upper corners of each
site are listed in Table 1. Using modified effective fetch and
maximum fetch measurements (Boizard and DeWreede
2006), these sites are classified as either semi-wave-exposed
or wave exposed (Table 1). The sites have similar topogra-
phy and were selected along a gradient of perceived level
of human influence including proximity to eutrophic out-
flow, finfish aquaculture sites, nuclear power stations, hu-
man traffic, and resource harvesting (for detailed
descriptions, see Bates et al. 2005). Once per summer, we
sampled one site per day over a 4 d low-tide series
(30 July 1 August 2000; 2224 July 2001; 1114 August
2002; 1417 June 2003; 36 June 2004; 2326 June 2005;
2729 May 2006).
Botany Vol. 87, 2009
Lower boundary Modified
of Ascophyllum effective Maximum Wave-exposure
zone (m) fetch (km) fetch (km) classification
1.3 958. >1000. Exposed
2.9 37. >1000. Semi-exposed
2.1 27. 92. Semi-exposed
2.8 157. >1000. Exposed
Seaweed sampling
We used two separate sampling approaches to summarize
richness, composition, and abundance of seaweeds. Sam-
pling for each type of data was performed according to a
set of predetermined rules in an attempt to maximize stand-
ardization for future collections and to minimize variation
due to sampler bias.
Site inventories were achieved by performing a ca. 1 h
roaming site search to identify all species that could be
found within the defined site. Collections made during these
searches also formed the basis for examination for micro-
scopic epiphytic species. This haphazard sampling targeted
the littoral zone as well as tide-pools. Here, these data are
used to calculate species richness and composition of the
flora for each site.
Seaweed abundance sampling was restricted to the lower
intertidal zone, the top of which was delineated by the lower
extent of Ascophyllum nodosum (Linnaeus) Le Jolis (this
boundary height for each site is listed in Table 1). This limit
was chosen because of the sharp lower boundary of the As-
cophyllum zone that dominates the mid-intertidal. Seaweed
abundance (percent cover) was estimated along four line-
transects that were randomly selected perpendicular to the
water line. Along each transect, two quadrats were sampled,
resulting in a total of eight quadrats per site, per sampling
event. Additionally, eight quadrats were sampled in each
of the mid (within the Ascophyllum) and upper (supra-
Ascophyllum) zones at each site, but only the results for
the lower zone sampling are reported here. Quadrat loca-
tions were randomly selected, and were rejected if they
fell within a tide pool or if the sampling area was covered
with unattached drift algae.
Bates et al. (2007) demonstrated that sampling at the ge-
nus level saves substantial time with a negligible decrease in
sampling resolution; compared with species-level identifica-
tions, genus-level seaweed abundance measurements re-
sulted in a 97% similar rank order of similarity
relationships between samples. Within each quadrat, percent
cover of algal genera, substrate, and two categories of mac-
roinvertebrates (mussels, other) were determined using
0.5 m  0.5 m random-point encounter quadrats (Jones et
al. 1980). Quadrats were strung with a 10  10 fishing-line
grid, resulting in 100 possible sampling points. Prior to sam-
pling, a random selection of 50 of these points was made,
each identified by a bright orange string tied to the intersec-
Published by NRC Research Press
Bates et al.
tion of two lines. During sampling, the quadrat was placed
on top of the seaweed without disturbing the canopy. Abun-
dance measurements were gathered by identifying the entity
found under each of the 50 points. Each point count repre-
sented 2% cover. After recording the percent cover values
for the 50 points, the seaweed canopy was then lifted to de-
termine which other species were present within the quadrat
area, yet did not fall under the random points (referred to as
non-point encounters). The identity of these species was
recorded, and each was assigned a cover value of one per-
cent. These data are used here to assess changes over time
in the assemblage structure of each site.
In studies that employ multiple observers, sampling bias
is always a concern. In an effort to minimize bias, three
safeguards were implemented: (i) the Northeast Algal Soci-
ety taxonomic key (Sears 1998) was adopted as a taxonomic
standard; (ii) samplers were instructed to collect any and all
samples of uncertain taxonomic identity for collaborative
identification in the lab; and (iii) a voucher collection was
compiled to allow comparisons with previously identified
samples (personal collections of G.W.S., herb. U.N.B.).
Historical records of seaweed diversity
No historical abundance data were available for our sites.
However, presenceabsence data were available (Smith et
al. 1978) for the Letite site, and for several locations in
close proximity to our sampling sites (Fig. 1), including
Point Lepreau (near our Lepreau study site), and South
Head Beach (near our Grand Manan study site). These his-
torical data were compiled by Smith et al. (1978) from three
sources: (i) a summary of collections housed in the Univer-
sity of New Brunswick Herbarium, which spanned from
1954 to 1978; (ii) data from one-time transect surveys, per-
formed in June 1978; and (iii) for Point Lepreau, summaries
from 2 years of informal monthly site surveys. To maximize
data matching between historical and contemporary data, we
updated the taxonomy of these records using Sears (1998)
and AlgaeBase (www.algaebase.org). As well, we truncated
the historical records to include only littoral collections that
took place between the months of June and August. For fur-
ther discussion about the utility and caveats of using herba-
rium collections as historical data, see Ponder et al. (2001).
Abiotic parameter sampling
Concurrent with the biotic sampling during 20002001,
seawater temperature and salinity, as well as nitrogen and
phosphorus levels (through seawater and algal tissue) were
also determined. Seawater temperature was measured at low
water using a thermometer (0.5 8C). Salinity was measured
at low water using a refractometer (1 ppt). Algal-tissue-
based nitrogen and phosphorus levels were determined using
Ulva lactuca L. Tissue collections were obtained from both
the lower and upper intertidal zones. Samples were cleaned
of invertebrates and decaying material, paper-towel dried,
and placed in a drying oven at 60 8C until a constant dry
3 Abioticdata are listed in online supplemental material (Table S1) to serve as a baseline for future comparisons.
4 Supplementary data for this article are available on the journal Web site (http://botany.nrc.ca) or may be purchased from the Depository of
Unpublished Data, Document Delivery, CISTI, National Research Council Canada, Building M-55, 1200 Montreal Road, Ottawa, ON K1A
0R6, Canada. DUD 5282. For more information on obtaining material refer to http://cisti-icist.nrc-cnrc.gc.ca/eng/ibp/cisti/collection/unpub-
lished-data.html.
1069
weight was achieved (approximately 72 h). The tissue was
then ground to fine powder and assessed for levels of nitro-
gen and phosphorus as in Chopin et al. (1999). Nitrogen and
phosphorus levels were also estimated using seawater sam-
ples at each site. Triplicate samples were taken at maximal
ebb tide. Water was filtered in the field using syringes
equipped with GF/C Whatman filters (pore size = 45 mm).
Samples were then frozen and stored until processing
(Chopin et al. 1999). Temperature and salinity did not vary
across sites, and the nutrient measurements were highly var-
iable, both within and across sites3,4. Abiotic parameter val-
ues for the area are also discussed in Worm and Lotze
(2006).
Data analysis
Seaweed monitoring data were analysed using univariate
(Underwood 1997) and multivariate (Clarke 1993) ap-
proaches. Seaweed species richness data met the assump-
tions of normality (ShapiroWilk test, W ranged from 0.837
(p = 0.10) to 0.921 (p = 0.53)) and homogeneity of variance
(Levenes test, F = 0.223 (p = 0.876)), so site differences in
seaweed species richness were assessed using one-way AN-
OVA followed by Tukey post-hoc tests. Compositional dif-
ferences of seaweed assemblages were assessed by
calculating BrayCurtis similarity values on species lists
and square-root transformed genus-level abundance data.
Similarity matrices were visualized using nonmetric multidi-
mensional scaling (nMDS), and differences between sample
groups were assessed using analysis of similarities
(ANOSIM) tests. When testing for temporal variation within
sites, ANOSIM test alpha values were Bonferroni adjusted
(pcrit = 0.002381) to account for 21 multiple comparisons
per site. Dominant taxa (i.e., taxa responsible for 90% of
within-group similarity) were identified using the similarity
percentages (SIMPER) routine. Univariate data analyses
were performed using JMP (version 4.0, SAS Institute Inc.
Cary, North Carolina, USA), and multivariate data analyses
were performed using PRIMER software (version 6.1.6,
Primer-E Ltd., Lutton, Ivybridge, UK). Comparisons be-
tween historical and contemporary collections are limited to
qualitative assessments.
Results
General patterns
Over 7 years of sampling, we encountered 72 seaweed
species (see supplementary data,4 Table S2). This is likely a
slight underestimate because we lumped together some of
the fleshy and coralline red crusts. Of these 72 encountered
seaweed species, 31 were red (division Rhodophyta), 25
were brown (class Phaeophyceae), and 16 were green (divi-
sion Chlorophyta). Seventeen species were ubiquitous
throughout our sampling program, whereas only 8 species
were found at only one site (see supplementary data,4 Table
S2).
Published by NRC Research Press
1070 Botany Vol. 87, 2009

Table 2. Comparison of seaweed species richness between con- cords, 16 species were collected between 19541978 that
temporary (between 20002006) surveys, historical surveys (Le- were not encountered in contemporary sampling (Table 4).
preau, 1963 and 1964; South Head Beach and Letite, 1978), and Of these 16 species, 5 are reds, 6 are browns, and 5 are
herbarium collections (19541978) from the University of New greens.
Brunswick, Fredericton.
Assemblage structure: spatial variation
Site Species richness
Generic percent cover of lower zone seaweeds was differ-
20002006 surveys ent among all of the four sites (ANOSIM Global R = 0.486
Grand Manan 49.1 p < 0.001). Average percent cover values of dominant taxa
Letite 40.3 for each site are listed in Table 5.
Lepreau 51.3 The Grand Manan samples were characterized by 12 algal
Meadow Cove 52.3 genera, but dominated by Laminaria, Ulva, and Fucus. Red
Historical surveys algae were most diverse, with seven of the remaining nine
South Beach Head 20 dominant genera being rhodophytes. On average, generic
Letite 29 composition of Grand Manan samples was 29.8% similar
Point Lepreau 11 (1963), 28 (1964) across samples.
Letite samples lacked many of the seaweed genera found
Herbarium
South Beach Head 10
at other sites. Instead, the dominant cover was rock (i.e., no
Letite 37
macrobiota), followed by invertebrates that were typically
Point Lepreau 74
blue mussels, Mytilus edulis, or green sea urchins, Strongy-
locentrotus droebachiensis. Algae typically made up a small
percentage of the lower-zone cover at Letite, with only two
Table 3. Results of a one-way ANOVA for differences in species dominant algal genera: Fucus and Palmaria. On average,
richness at 4 different sites in the outer Bay of Fundy, New generic composition of Letite samples was 28.5% similar
Brunswick. across samples.
Eight genera were dominant at Lepreau, slightly less than
df SS MS F p found at Grand Manan. Typical samples from Lepreau were
Site 3 572.1 190.7 6.60 0.003 dominated by beds of Chondrus, with patches of Laminaria
Error 21 606.5 28.9 and Mastocarpus, as well as Fucus, Ulva, Cystoclonium,
Total 24 1178.56 Corallina, and to a lesser extent, Chaetomorpha. Like Grand
Manan, the generic composition of Lepreau samples was
29.8% similar across samples.
Seaweed species presenceabsence Meadow Cove had a very similar flora to Lepreau, with
Across sites, seaweed species richness differences were the two dominant genera being Chondrus and Laminaria.
observed (Tables 2 and 3), with Letite (40.3 2.0 SE) hav- Mastocarpus was slightly less abundant at Meadow Cove,
ing a lower average species richness compared with Grand but levels of Fucus were similar between the two sites. Al-
Manan (49.1 2.0), Lepreau (51.3 2.0), and Meadow together, 10 genera (Table 5) characterized Meadow Cove
Cove (52.3 2.6). There was no evidence that seaweed spe- samples, and on average, composition of samples was more
cies richness has changed over the duration of our sampling similar compared with other sites, at 42.6%.
program, except for perhaps a slight increase in richness
after the onset of our sampling (Fig. 2). Assemblage structure: temporal variation
Composition of seaweed species was also similar across The assemblage structure at Letite and Meadow Cove did
sites, except for Letite, which had a different composition not change significantly (ANOSIM p = 0.084 and p = 0.33,
of seaweed species (ANOSIM p < 0.05, Fig. 3) compared respectively) over the course of our sampling (Fig. 4). Some
with the other sites5. Within sites, seaweed species composi- temporal differences were observed at Grand Manan (2000
tion was minimally variable from year to year, with Letite vs. 2004, 2001 vs. 2004; p < 0.002). Specifically, in 2004,
inventories being the least similar (80.2%), and Meadow there was a lower percent cover of Fucus, Laminaria, Cysto-
Cove being the most similar (89.4%) in species composition. clonium, and Ulva/Ulvaria and a higher percent cover of
Alaria and Acrosiphonia compared with 2000. Also, in
Comparison with historical data 2004, there was a lower percent cover of Laminaria, Alaria,
Depending on the data source, the historical data sets re- Ulva/Ulvaria, and Porphyra and higher percent cover of
vealed either higher or lower seaweed species richness than Palmaria and Acrosiphonia compared with 2001. Long-
the contemporary collections (Table 2). The 1978 and 1963 term changes were not observed at our Grand Manan site,
1964 collections were lower in seaweed richness than con- as the structure of assemblages in 2005 and 2006 was not
temporary sampling revealed. The herbarium summaries different than that in 2000 and 2001. We also observed tem-
were variable, with the South Beach Head and Letite collec- poral differences at Lepreau (2001 vs. 2004, 2001 vs. 2006,
tions having low species richness (10 and 37 species, respec- and 2003 vs. 2004). These differences were due to variation
tively), and the Point Lepreau herbarium collections having in the dominant cover genera: Laminaria, Chondrus, and
much higher species richness (74). Across the historical re- Mastocarpus, and some of the less dominant components:
5 Percent encounters per species are listed in online supplemental material, Table S2.

Published by NRC Research Press

Bates et al.
Fig. 2. Annual patterns of seaweed species richness for four sites in
the Bay of Fundy. *, significant difference from other sites
(ANOVA, Tukey, P < 0.05).
Cystoclonium, Corallina, Palmaria, Fucus, and Acrosipho-
nia. However, these differences appear to reflect natural var-
iation in abundance of these genera, and not long-term
reductions.
Discussion
Changes in marine benthic vegetation have been well
documented in some parts of the world, particularly in Eu-
rope (Schramm and Nienhuis 1996). These changes can be
mediated through reductions in seawater quality, light pene-
tration, substrate suitability, herbivore dynamics, competi-
tive interactions, and reduced recruitment potential (Walker
and Kendrick 1998). Although these stressors often act in
conjunction with one another, making it difficult to link
changes in macrophyte biodiversity to a particular anthropo-
genic pressure, there is evidence of declines in water quality
and increases in nutrient levels and silt in the Bay of Fundy
(Lotze and Milewski 2004), and earlier surveys indicate im-
pacted macroalgal assemblages (Worm 2000). Here, we
demonstrate that, although conclusions of human impact are
not possible, the flora at one of our four monitoring sites is
consistent with published accounts of environmental stress.
Further, 16 seaweed species previously encountered in the
outer Bay of Fundy have not been relocated during 7 years
of contemporary sampling.
1071
Fig. 3. Non-metric multidimensional scaling plot showing similarity
in species composition of seaweeds at four sites along the New
Brunswick coast of the Bay of Fundy. Presenceabsence records
were collected in summer, in 20002006 (except Meadow Cove,
20032005). Composition is the same at all sites except Letite,
which is different from all other sites (ANOSIM, P < 0.05).
Spatial differences in seaweed biodiversity
Spatial differences in seaweed biodiversity were observed
among our monitoring sites, but the extent of the differences
depended on the type of data examined. Richness and com-
position analyses showed only the Letite site to be different
from the others, whereas all four sites were different based
on generic abundance measurements. Abundance data have
a greater multidimensionality than presenceabsence data,
so with quantitative measurements it is possible to detect
differences that do not involve the complete extirpation of
species from the flora.
Of the four sites we examined, Letite was the most likely
to be classified as impacted. It is possible that the flora ob-
served at Letite is not the result of floristic change, but in-
stead due to differences in abiotic site conditions or natural
variability. However, two lines of evidence suggest that the
flora observed at Letite is a result of temporal changes.
First, we have been informally collecting seaweeds at Letite
on an annual basis for the past 17 years, and our anecdotal
observations suggest that the flora at Letite was previously
similar to the other sites in our sampling program; we have
perceived major changes since our initial collecting trips.
Second, the type of assemblage currently seen at Letite is
consistent with previously reported impacted floras
(Schramm and Nienhuis 1996; Middelboe and Sand-Jensen
2000). Letite had a lower species richness than the other
sites, and the dominant cover was bare rock, which could
be explained by the presence of herbivorous mobile inverte-
brates, in this case sea urchins and periwinkle snails. We ac-
knowledge that it is not possible to attribute the shift of the
Letite flora to a particular anthropogenic stressor, and that
our sampling was not initiated in time to demonstrate the
changes. However, we suggest that it would be worthwhile,
in future studies, to formally investigate the possibility of
links between anthropogenic stressors and biotic impact at
Letite.
One macroalgal phenomenon often linked with eutrophi-
cation is green tide (Raffaelli et al. 1998; Middelboe and
Sand-Jensen 2000), where nutrient scavenging annuals (typi-
cally ulvoid green algae) proliferate to substantial biomass.
Published by NRC Research Press
1072 Botany Vol. 87, 2009

We did not witness green tides at any of our current moni-

toring sites, but this could reflect the rocky substrate and
herbivorous invertebrates observed at our sampling sites.
Point Lepreau 1964

Herbivores can effectively prevent the bloom-forming algae

from establishing (Worm and Lotze 2006) because the
newly recruited algae are consumed before they can estab-
lish. Green tides have been observed elsewhere in the Bay
of Fundy, and are typically found on sheltered mudflat areas
(Auffrey et al. 2004), or at more sheltered rocky intertidal
0
0
0
0
0
0

0
0
0
0
0

0
0
0
0
0
sites (Bates et al. 2005).
One response that we did not measure here is a change in
the depth distributions of macroalgae in response to changes
Point Lepreau 1963

in light regimes mediated through excessive silt or phyto-

Table 4. Seaweed species present in the flora from the period of 1954 to 1978 but not encountered during contemporary (20002006) sampling.

Monthly survey

plankton in the water, or through altered competitive inter-

actions with fast-growing epiphytic species that respond
quickly to elevated nutrient levels (Sand-Jensen and Borum
1991). The Bay of Fundy has a naturally high silt load ow-
0
0
0
0
0
0

0
0
0
0
0

0
1
0
0
0
ing to substantial tidal flux and the dominance of mud flats
in the upper part of the Bay, so it would be interesting to see
if shifts in vertical distribution of subtidal seaweeds have oc-
South Head Beach

curred in response to more frequent phytoplankton blooms

(Martin et al. 1999).

Temporal changes in seaweed biodiversity

One time surveys 1978

No major changes in richness, composition, or assemblage

0
0
0
0
0
0

0
0
0
0
0

0
1
0
0
0

structure of the floras at our monitoring sites were seen over

the duration of contemporary sampling, but there were small
variations in species composition seen from year to year.
The slight increase in richness at some sites after the onset
Letite

Note: Data from Smith et al. (1978). Taxonomic authorities for these species can be found in Sears (1998).

of sampling likely reflects an improvement in the ability of

0
0
0
0
0
0

0
0
0
0
0

0
0
0
0
0

our team to locate and identify constituent species. At Le-

tite, the relative consistency of assemblage structure over
South Head Beach

time is consistent with the findings of Bertness et al. (2002)

who found that, in the Gulf of Maine intertidal zones, recov-
ery from natural disturbance is largely deterministic and can
be influenced by the water flow regime and presence of con-
sumers. Specifically, they report that mussel beds are favored
0
0
0
0
0
0

0
0
1
0
0

0
0
0
0
0

in higher flow areas (such as our monitoring sites), and where

consumers are present, recovery to either mussel-dominated
Herbarium collections 19541978

or seaweed-dominated assemblages is halted. This may

Point Lepreau

explain why bare rock was dominant in a part of the intertidal

at Letite, where space is typically the limiting resource. At the
other monitoring sites, the consistency in the flora may
indicate the lack of major disturbance.
1
1
1
1
1
1

1
1
0
1
0

0
1
1
1
1

Comparisons with historical data

Comparisons with historical collections such as herbaria
Letite

can often prove problematic because species present at a

0
0
0
1
0
0

0
0
0
0
1

1
0
0
0
0

site can get overlooked if the goal of the original survey or

herbarium collection was not to document the presence of
all species (Ponder et al. 2001). This appears to be the case
for many of historical data sets used in this study. Several
common and abundant species that were ubiquitous during
Chaetomorpha linum/aerea

our collections (e.g. Ascophyllum nodosum, Chondrus

Leptonematella fasciculata

Audouinella membranacea

Capsosiphon groenlandica
Rhodomela lycopodioides

crispus Stackhouse) are absent from most historical data

Urospora wormskjoldii
Division Chlorophyta
Punctaria plantaginea

Division Rhodophyta

Erythrodermis traillii
Class Phaeophyceae

sets. Recognizing that absence of proof is not proof of ab-

Hincksia granulosa

Ulothrix laetivirens
Cladophora albida
Ceramium elegans
Punctaria latifolia

Ralfsia verrucosa

sence, we restrict our assessments of floristic change to

Tsengia bairdii
Chorda filum

species that were previously found, but have not been en-
countered during our recent sampling.
Sixteen species recorded between 19541978 were not
re-encountered during contemporary surveys (Table 3).

Published by NRC Research Press

Bates et al. 1073

Table 5. Average percent cover of dominant seaweed genera, invertebrates, and substrate that are typical of the
lower intertidal zone at four monitoring sites along the New Brunswick Coast of the Bay of Fundy, Summer
20002006 (except Meadow Cove, 20032005).

Grand Manan Letite Lepreau Meadow Cove

Acrosiphonia 3.13
Alaria 5.59 1.92
Ceramium 3.09
Chaetomorpha 0.71
Chondrus 16.32 18.58
Coralline Crust 0.96
Corallina 4.04 1.63
Cystoclonium 4.89 6.14 1.38
Devaleraea 4.05
Fucus 7.43 5.82 5.66 6.08
Laminaria 16.23 8.8 12.04
Mastocarpus 2.55 7.77 2
Mytilus 9.8
Palmaria 3 3.3 2.71
Polysiphonia 2
Porphyra 2.5
Rhodomela 1.27
Ulva/Ulvaria 9.11 3.32
Rock 22.8
Urchins, snails 7.61
Note: Values shown in the table are those retained by the SIMPER routine of PRIMER to differentiate the sites.

Fig. 4. Non-metric multidimensional scaling plots showing temporal variation in BrayCurtis similarity of intertidal assemblage structure at
four sites along the New Brunswick coast of the outer Bay of Fundy. Similarity was calculated using genus-level percent cover measure-
ments of lower intertidal seaweeds and invertebrates, as well as substratum. Records were collected in summers, 20002006 (except Mea-
dow Cove, 20032005). The stress values for nMDS plots are as follows: Grand Manan, 0.18; Letite, 0.14; Lepreau, 0.18; Meadow Cove,
0.09. Despite some year to year variation at Grand Manan and Lepreau, no major shifts in composition were observed.

There are several potential explanations for why we have
not relocated these species. Many of the species listed in
Table 4 may have been misidentified, either by earlier in-
vestigators, or by our team. For example, Capsosiphon
groenlandica (J. Agardh) K.L. Vinogradova can easily be
mistaken for an Ulva species in the field, Ceramium can
be challenging to identify to the correct species, and to the
untrained eye, Ralfsia verrucosa (Areschoug) Areschoug
may have been mistaken for the Petrocelis phase of the
Mastocarpus life history. For several genera (e.g., Clado-
phora), the taxonomic characters available to differentiate
species are ambiguous, and other genera such as Rhodo-
mela require taxonomic reworking (Sears 1998). Although
we excluded subtidal collections from our assessment of

Published by NRC Research Press

1074
historical data, several subtidal species such as Erythroder-
mis traillii (Holmes ex Batters) Guiry & Garbary, Hincksia
granulosa (J.E. Smith) P.C. Silva, and Leptonematella fas-
ciculata Reinke were historically reported from lower tide-
pools. These species, which are often found as subtidal
epiphytes, may in fact be present in the subtidal area of
our study sites. Many of the species listed in Table 3 dwell
in crevasses (e.g., Chaetomorpha linum/aerea), are small in
stature, such as Audouinella membranacea (Magnus) Pa-
penfuss, or rare (Tsengia bairdii (Farlow) K.C. Fan &
Y.P. Fan), and may simply have been overlooked in the
field. While sampling elsewhere in the Bay of Fundy, we
have encountered some species from Table 4, such as
Chorda filum (Linnaeus) Stackhouse and Erythrodermis
traillii, indicating that we may be witnessing shifts in
metapopulation structure, and not loss of species. It is also
possible that some of these species have actually been ex-
tirpated from the flora or from the local area by the ele-
vated anthropogenic stressors, for several larger species
have not been seen in the Bay of Fundy by us (e.g., Punc-
taria latifolia Greville and P. plantaginea (Roth) Greville).
On the surface, it appears as though the marine algae of
the Bay of Fundy are not subject to widespread anthropo-
genic influence, unlike many other biotic components of the
Bay of Fundy (Lotze and Milewski 2004). This may be par-
tially true, reflecting the resilience of many marine algae to
disturbance. Although numerous exceptions exist, seaweeds
are typically very fast growing and tend to have high repro-
ductive output (Graham and Wilcox 2000). However, the
major difficulty in trying to assess change in Bay of Fundy
seaweed biodiversity is the lack of quantitative measures of
abundance for comparison. Where species have not been lo-
cated in contemporary sampling, we rely on taxonomic keys
and general descriptions of species to determine estimates of
abundance or rarity. Further, we have not been able to as-
sess the likelihood of shifts in abundance of constituent spe-
cies. These difficulties highlight the importance of collecting
baseline data that are appropriate to the task of assessing
change in assemblage structure.
Several previously published floristic studies (Edelstein et
al. 1970; South et al. 1988) reported a significantly higher
number of species in the Bay of Fundy (187 and 171, re-
spectively). In our study we identified only 72 species, so a
discussion of why we found substantially fewer species is
warranted. First, we largely confined ourselves to the macro-
algae and didnt identify most of the microscopic seaweeds
(unless, as for A. secundatum, we routinely uncovered them
in identifying the larger taxa). Second, in light of advances
in the general understanding of species with heteromorphic
life histories we did not allow for alternate stages to be re-
corded as separate species (e.g., both Edelstein et al. (1970)
and South et al. (1988) list Conchocelis, the sporophyte
stage of the Porphyra life history, as a separate species
from the gametophyte). Third, ca. 100 algal records from
South et al. (1988) are indicated as rare (or only found once
or twice), ca. 45 algal records are reported as only being
from Nova Scotia (and thus not part of the study area), and
ca. 10 algal records are only from Campobello Island, which
was not included in our study. For the previous, many of
these uncertain records were taken by South from published
Botany Vol. 87, 2009
accounts (ca. 80; which the authors indicate in their list) and
thus were not confirmed as being in the flora. Fourth, we
put taxonomically complex genera (e.g., taxonomy is known
to be uncertain and identification to species based on current
keys suspect at best, or species that were too difficult to
identify past the genus level in the field) into bins to facili-
tate ecological sampling (i.e., ours was not strictly a floristic
account). Based on the previous, it is not unfair to infer that
we actually have no idea as to the number of species in the
Bay of Fundy, not even of our relatively confined study area
in Southwest New Brunswick, which ones are representative
of the flora (except for the obvious common species), which
are occasional invaders (e.g., during particularly warm
years), and which records are simply based on misidentifica-
tions (seaweeds are notoriously difficult to identify; see Sa-
unders (2005, 2008)).
The utility of our collections as a seaweed biodiversity
baseline
The grounds for assessing changes in biodiversity are de-
pendant on high quality baseline information (e.g., Barry et
al. 1995; Beuchel et al. 2006), so we feel that the true value
of this study will be for future assessments of floristic
change. The protocols that we established to estimate sea-
weed biodiversity have been through several rounds of re-
finement (Bates 2002; Bates et al. 2007) to facilitate
efficient collection of effective measures of seaweed biodi-
versity. The protocols are described in detail here, providing
opportunities for other investigators to collect comparable
data with a similar level of sampling effort. The multiple de-
scribed types of data (species richness, composition, and as-
semblage structure) comprise a multifaceted and
complementary approach to assess temporal and spatial dif-
ferences in seaweed biodiversity. The richness and composi-
tion data are high resolution; species level identifications
were sought in most cases, and the reference collection that
we established is available to verify against future collec-
tions. Assemblage structure data, although documented at
the genus level, have proven sufficient to determine site-to-
site differences, as well as within-site differences over time.
When has sufficient baseline data been collected? This
depends on the questions being asked, and hence, the type
of data being collected. In our study, asymptotes in species
richness and composition were comparatively quickly
reached, however, the variation in assemblage structure
took longer to capture, and the range of natural variation in
seaweed assemblage structure is likely still not fully de-
scribed. The data we have documented here do not take
into account seasonal variation in assemblage structure.
However, it appears as though our sampling has encom-
passed the major extent of the natural variation taking place
in summer months. Further, it may be argued that our study
sites are already impacted by human activities, and that no
pristine reference sites exist in the Bay of Fundy. However,
we suggest that there are undoubtedly more changes yet to
come. Interestingly, the project rationale outlined by Smith
et al. (1978) is very similar to our own. Long-term monitor-
ing programs such as the one described here are a valuable
resource for the future, because information collected today
will become tomorrows historical data.
Published by NRC Research Press
Bates et al.
Acknowledgements
We are grateful for the enthusiastic early morning field
support of countless volunteer undergraduate and graduate
sampling technicians. Discussions with B. Clarke,
S. Murray, and G. Pohle helped to solidify sampling and
analysis protocols. Natural Sciences and Engineering Re-
search Council of Canada (NSERC) and Canada Research
Chair grants to G.W.S., NSERC and AquaNet grants to
T.C., Huntsman Marine Science Centre grants to
G.W.S. and C.R.B, and a Vaughan Fellowship to
C.R.B. supported this research. This paper is contribution
no. 81 from the Centre for Coastal Studies and Aquaculture.
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