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M.A.Ward and J.D. Mays
22001144 NORTHEASTERN NATURALIST 2V1(o4l). :2517,4 N–5o8. 64
Survey of a Coastal Tiger Beetle Species, Cicindela
marginata Fabricius, in Maine
Mark A.Ward1, 2,* and Jonathan D. Mays3, 4
Abstract - We undertook field surveys for Cicindela marginata (Salt Marsh Tiger
Beetle) in Maine and confirmed its presence at previously documented sites and at new
locations in the southern third of the state. We found no evidence that this species has
experienced a recent decline in Maine. We observed adults and larvae in a variety of
sparsely vegetated sandy microhabitats in tidal ecosystems. Our findings suggest broader
ecological amplitude for C. marginata than simply backdune-marsh strands. Variability
within occupied salt marsh habitats suggests the potential for substantial differences in
habitat use and resilience of C. marginata in comparison to other coastal tiger beetle
species. Nevertheless, better baseline documentation and understanding of this species’
biology is needed in light of current and emerging threats to coastal habitats. We recommend
that researchers conduct comprehensive C. marginata inventories throughout its
range that take a far-reaching approach in terms of site selection, microhabitats searched,
and life stages sought.
Introduction
In the northeastern US, many communities within coastal ecosystems are highly
threatened, including tidal marshes, nearshore mudflats, rocky intertidal zones,
beaches, and dunes. Coastal ecosystems receive a disproportionately high concentration
of human impacts (UNEP 2006) and are very susceptible to effects from
climate change such as sea-level rise (Kirwan et al. 2010, Whitman et al. 2010).
Many species that rely on these ecosystems including multiple tiger beetle (Coleoptera:
Cicindelidae) species have also been adversely affected by habitat alteration.
For example, the federally threatened Cicindela dorsalis Say (Northeastern Beach
Tiger Beetle) has been extirpated from almost all of its former range (Pearson et al.
2006, USFWS 2009), and coastal populations of C. hirticollis Say (Hairy-necked
Tiger Beetle) have also experienced decline (Leonard and Bell 1999, Schlesinger
and Novak 2011, Sikes 1998). However, one coastal tiger beetle species that has
received relatively little concerted survey effort is C. marginata Fabricius (Salt
Marsh Tiger Beetle).
C. marginata is restricted to coastal localities in North America, including all 14
Atlantic states from Maine to Florida and several islands in the West Indies (Choate
2003, Pearson et al. 2006). A single individual was recently reported from Nova
Scotia (Neil and Majka 2008). As its common name implies, C. marginata adults
1University of New England, Department of Environmental Studies, Biddeford, ME 04005.
2Current address - 3424 Campus Boulevard NE, Albuquerque, NM 87106. 3Maine Department
of Inland Fisheries and Wildlife, Bangor, ME 04401. 4Current address - 2004 SE 41st
Avenue, Gainesville, FL 32641. Corresponding author - maward@centurylink.net.
Manuscript Editor: Christopher M. Heckscher
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can be found in salt marshes, especially those associated with sandy barrier beaches
(Leonard and Bell 1999). According to Dunn (1981), adults can be found along the
back beach where the dunes and the salt marsh meet, on saline mud flats, and occasionally
on sandy ocean beaches. Adults have also been reported from the mouths
of tidal streams (Pearson et al. 2006) and occasionally on coral outcrops (Choate
2003). In the northeastern US, this is a summer-active species with most adults
observed in July or August (Leonard and Bell 1999, Sikes 1998). Although details
of this species’ life history are not well known (Leonard and Bell 1999), larvae are
reported to occupy relatively shallow (8–15 cm) burrows a few meters back from
high-tide line in open, sandy, sparsely vegetated soil (Pearson et al. 2006). This species
is considered globally secure (ranked G5; NatureServe 2009). However, in the
northeast, C. marginata is considered to be a dwindling species (Leonard and Bell
1999) and has apparently been extirpated in New England except at a few protected
natural areas (Pearson et al. 2006). Published results of surveys for C. marginata in
this region are not available or have not been undertaken.
Prior to 2010, only 4 C. marginata locations had been documented in the state
of Maine (Maine Department of Inland Fish and Wildlife [MDIFW], Bangor, ME,
unpubl. data). All of these occurred along the southern third of the coast, and 2 of
them are considered historical (pre-1970). In response to growing concerns about
the possibility of regional decline, the possible limitations of suitable habitat, and
other factors such as projected rising sea levels (Whitman et al. 2010), we conducted
a survey for this species in Maine in 2010. The aim of our survey was to
gather information regarding the distribution of C. marginata in Maine and assess
its conservation status by (1) reconfirming its presence at previously documented
sites, (2) surveying additional sites with suitable habitat, and (3) documenting adult
abundance and flight times. An additional objective was to record other tiger beetle
species found in similar habitats.
Methods
We developed a list of field sites that included previously documented locations
of C. marginata and additional areas with potential habitat. All 4 previously documented
locations in Maine were in coastal dune–marsh ecosystems (Gawler and
Cutko 2010). These ecosystems are most common along Maine’s southern coast
and are uncommon elsewhere in the state (Gawler and Cutko 2010). We identified
potential habitat by searching orthoimages—geometrically corrected aerial photos—
of coastal Maine for barrier beaches with unvegetated habitat patches adjacent
to salt marsh, and through conversations with knowledgeable individuals. The majority
of identified field sites were located in the southern portion of the state. Of the
4 previously documented sites in Maine, 2 had been surveyed within the previous
10 years and 2 others were surveyed prior to 1970.
During site visits, we surveyed appropriate habitat focusing on the backdune–
marsh ecotone where unvegetated sand lies between the beach backdune and the
salt marsh high-tide line. We refer to this targeted habitat as the backdune–marsh
strand. It normally occurs between stands of Ammophila breviligulata Fernald
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(Dune Beachgrass) and the salt marsh proper, where Spartina spp. (salt marsh
cordgrasses) often emerge (Fig. 1). Typical vegetation on this strand includes scattered
Limonium carolinianum (Walter) Britton (Sea Lavender). At some sites, we
also surveyed the oceanside beach and other potentially suitable habitats, typically
while en route to the primary habitat of interest. We recorded GPS coordinates of all
locations visited. We observed and netted adult tiger beetles and sought to collect
voucher specimens (or photographs) of each species encountered at each site. To
a lesser extent, we surveyed larval burrows and larvae using the fishing technique
(Pearson et al. 2006). We confirmed, labeled, and deposited all vouchered tiger
beetle specimens and photographs with the Maine Department of Inland Fisheries
and Wildlife for addition into the state’s tiger beetle database. We described and/or
photographed occupied habitats at sites where we found C. marginata.
Results
During 29 June–12 August 2010, we made 32 visits to 28 sites in 20 different
tidal river–marsh systems (Fig. 2). We documented C. marginata adults at 12 sites
in 10 river–marsh systems from southernmost Maine (York, York County) to midcoast
Maine (Georgetown, Sagadahoc County; Fig. 2). In 2010, we confirmed the
presence of C. marginata at all 4 of the previously documented sites and at 8 new
sites on 6 new river–marsh systems. We observed adults 29 June–1 August. Observations
of C. marginata at occupied sites ranged from less than 1 to 31 adults per hour. The
Figure 1. A photograph of backdune–marsh strand habitat in Maine, 2010.
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mean number of adults per hour at occupied sites was 11.2 ± 2.9. We recorded ≥20
adults at 6 sites (Appendix 1).
We documented C. marginata adults and/or larvae in 3 tidal ecosystem habitats:
backdune–marsh strand, salt marsh, and oceanside beach (Fig. 3). All adult and
larval microhabitats were characterized by sandy substrates and sparse vegetation.
Larval microhabitats were all located at (or near) the highest high-tide line. The
most common habitat in which we saw adults (9 sites) was the targeted backdune–
marsh strand. At one site, we also observed larvae in this habitat.
We recorded both C. marginata adults and larvae in salt marsh habitat (3 sites)—
the habitat in which larvae were most frequently observed in this study. Salt marsh
microhabitats in which we found larval burrows were all characterized by tightly
packed sand, but differed in some ways. Variations included (1) provenance—natural
vs. artificial, (2) whether associated with a major river or stream system, (3)
proximity to tidal mouth, and (4) degree of human recreational activity (Table 1).
Figure 2. Location of sites surveyed for C. marginata in Maine, 2010.
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We also documented C. marginata adults on oceanside beach habitat where larvae
appeared to be absent (two sites). This habitat was surveyed at only 12 sites, of
which 4 hosted C. marginata.
We observed adults of 5 other tiger beetle species (Table 2). C. hirticollis was
the most commonly recorded species (13 sites)—we saw it at 7 of 12 sites occupied
Figure 3. Example of 3 coastal ecosystem habitats occupied by C. marginata in Maine,
2010 (salt marsh, backdune–marsh strand, oceanside beach).
Table 1. Variation in salt marsh microhabitats for C. marginata in Maine, 2010.
Association Distance
with river/ to tidal
Site Provenance stream system mouth (m) Degree of recreational activity
Georgetown site 2 Natural No 200 Heavy use of lagoon
York site 1 Artificial Yes 1000 None perceived
Wells site 6 Natural Yes 200–250 None perceived
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by C. marginata. Cicindela hirticollis adults were a common co-inhabitant with
C. marginata in backdune–marsh strand habitats (7 of 9) and at both oceanside
beach sites where we observed C. marginata, but we did not document them at any
of the 3 salt marsh microhabitats occupied by C. marginata.
Discussion
Documented sites
Our survey effort increased the number of documented C. marginata sites and
extended the species’ range slightly southward, but not northward, in Maine. The
northernmost sites for C. marginata in the state continue to be located in Sagadahoc
County in Midcoast Maine (43°46'N, 69°43'W; MDIFW, unpubl. data). In 2010,
our searches at the most northern 3 sites were unsuccessful. We identified very few
potential C. marginata sites in Downeast Maine—the region from Penobscot Bay
to the New Brunswick border—from orthoimagery because that part of the state
generally lacks barrier beaches and coastal dune-marsh ecosystems. The Downeast
Maine coastline is characterized instead by a combination of broad intertidal
embayments of mud and mixed mud-gravel flats sheltered by granite islands and
steep, straight bedrock ledges scoured by dramatic tides and floored by extensive
tidal flats (Kelley et al. 1989). Under the survey scheme that we employed, this
part of the state therefore received relatively little survey effort. Although a single
C. marginata adult reported from Nova Scotia more than 450 km northeast of the
nearest documented Maine population was likely a windblown stray (Neil and
Majka 2008), additional surveys in tidal habitats of Downeast Maine and the Bay
of Fundy are warranted to confirm this.
The presence of C. marginata in 2010 at all 4 sites where they had been previously
documented did not suggest to us that this species has declined in Maine.
However, historical data for this species were extremely limited with only a few
specimens from 4 locations and no historical abundance data from any site prior
to 2009.
We report this species’ presence at one site (Georgetown site 2) in a tidal marsh
that was not associated with a significant river or stream system and was in close
proximity to a heavily used recreational area. We found adults and larvae in this
35-ha salt marsh on the fringe of a small tidal lagoon (1 ha). The opposite side of
the small lagoon receives heavy recreational use. The salt marsh and lagoon have
Table 2. Additional tiger beetle species encountered during surveys for C. marginata in Maine, 2010.
# Sites # Sites
at which found with
Species encountered C. marginata
Cicindela hirticollis Say (Hairy-necked Tiger Beetle) 13 7
Cicindela repanda Dejean (Common Shore Tiger Beetle) 2 1
Cicindela formosa generosa Dejean (Big Sand Tiger Beetle) 1 1
Cicindela tranquebarica Herbst (Oblique-lined Tiger Beetle) 1 1
Cicindela punctulata Olivier (Punctured Tiger Beetle) 1 1
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an inlet controlled by a bedrock sill with a narrow opening acting as a kind of lowtide
dam that limits the exchange of salt water during the lower part of the tidal
cycle (Maine Geological Survey 2002). Perhaps more surprising than the presence
of C. marginata at this lagoon was its apparent absence, despite two thorough
searches, at an intact backdune–marsh strand behind a classic river-outlet barrierbeach
spit (less than 1.5 km away).
We also report, for the first time, the presence of C. marginata adults and larvae
at a salt marsh that is both well removed from a tidal river mouth (~1 km upstream)
and artificially constructed. York site 1, a triangular 7-ha artificial marsh (bounded
on the east and west by roads), was originally an intertidal clam flat on the York
River that was created from sandy river-bottom dredge-fill in the 1960s. A restoration
project was undertaken here in 2004 to enhance tidal exchange and improve
the vegetation and wildlife habitat (York Rivers Association 2011). We would
not have targeted this site because it lacked features of documented C. marginata
sites—a barrier beach and backdune-marsh strand—had we not learned during our
site-selection process that adults of the species had been observed here in 2008
(S. DesMeules, Medomak Valley Land Trust, Waldoboro, ME, pers comm.). It was
unclear how long C. marginata have occupied this salt marsh or whether it predated
the 2004 restoration project, but the species appeared to be persisting here
despite significant past and recent human alteration.
Flight period and adult abundance
In 2010, our recorded flight period of 29 June–1 August may have underestimated
the actual flight period for this species, although it likely captured the majority
of the active adult flight period (Leonard and Bell 1999, Sikes 1998). The 29 June
2010 observations preceded our planned site visits by about 1 week. Furthermore,
an unusually warm spring led to the early emergence of other invertebrate species
in 2010 (M. Ward, pers. observ.) and may have accelerated the emergence of
C. marginata adults. Only 2 site surveys took place after 1 August 2010, both at
Midcoast Maine sites with somewhat limited habitat potential. Yet, an adult of this
species was collected in Maine on 4 September 2011 (R. Lambert, Biodiversity
Research Institute, Gorham, ME, pers. comm.); we confirmed its i dentity.
At some sites where adults were present, the number of individuals observed
was quite low: only 1 adult at 2 sites. Adults at these sites may represent stray individuals
from nearby sites, low population sizes, or possibly a failure in our ability
to detect the best habitat at the site. One of the challenges associated with tiger
beetle surveys is ensuring detectability at a site when the species is present. We
attempted to maximize the chances of detection by making visits in good weather
conditions during the adult flight season and at times of the day when adults are
most likely to be active. Nevertheless, it is possible that we may have missed the
species at some sites in 2010, in part because we visited most sites only once. Tiger
beetle subpopulations also may move through space and time (Pearson and Vogler
2001), and some sites at which the species was not detected in 2010 may either have
supported populations in the past or could support viable populations in the future.
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Adult and larval microhabitats
The variety of habitats in which we observed C. marginata adults and larvae
suggests broader ecological amplitude for C. marginata than the backdune–marsh
strands of barrier beaches that were targeted in this study. Our results suggest that
any sparsely vegetated sandy area near the high-tide line within a tidal marsh ecosystem
could offer potential habitat for this species. Although our habitat results
are consistent with those previously reported for C. marginata (Table 3), they
document a heretofore-unreported range of variability within salt marsh habitat
for both adults and larvae. The diversity of salt marsh microhabitats in which we
found C. marginata suggests that this species has a greater degree of flexibility and
resilience than expected. Although this species should still be considered a specialist
of tidal habitats, our findings suggest that it may be somewhat adaptable within
tidal marsh habitats and can be found in a variety of sparsely vegetated sandy areas
near the high-tide line. The ability to utilize several tidal marsh microhabitats may
set C. marginata apart from other coastal tiger beetle species with which it is often
associated (e.g., C. hirticollis).
One of the significant gaps in our knowledge of this species’ biology is an understanding
of its larval habitat preferences or constraints. Habitat descriptions
for C. marginata have tended to focus on adults (Table 3). The locations where we
observed larvae in this study were characterized by tightly packed sand in salt
Table 3. Reported habitat types used by C. marginata in the eastern US. Sources: Maine (this study);
New Hampshire (Dunn 1981); Rhode Island (C. Raithel, Rhode Island Department of Environmental
Management, West Kingston, RI, unpubl. data); Connecticut (Sikes 1998); New York (Schlesinger
and Novak 2011); Northeastern US (Leonard and Bell 1999); South Atlantic States (Knisley and
Schultz 1997); Florida (Choate 2003); and US distribution (Pear son et al. 2006).
Outlet of
Backdune streams
Salt Saline Tidal -marsh emptying Oceanside Coral
Location marsh flats flats strand into ocean beach outcrop
Maine YesA Yes Yes
New Hampshire YesB Yes YesC
Rhode Island YesD Yes E YesF
Connecticut Yes Yes
New York Yes YesB
Northeastern US YesB Yes
South Atlantic States Yes Yes YesB YesG Yes
Florida Yes YesH Yes YesC
US distribution YesI YesB Yes Yes
A Natural and artificial.
B Mudflats.
C Occasionally.
D Edges of fringe marshes.
E Especially with cobble bars or tidal fans.
F Shell and mud beaches.
G Backbeaches.
H Coastal alkali mudflats
IShores.
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marsh or backdune–marsh strand habitats at, or near, the highest high-tide line.
Pearson et al. (2006) suggested that larval burrows may also be found on the sea
beach a few meters back from the high-tide line. It is important to improve our understanding
of larval microhabitat locations because tiger beetle larvae generally
occupy a much narrower range of microhabitats than adults (Pearson and Vogler
2001). The majority of a tiger beetle’s life is spent in the relatively immobile larval
stages, and protection of tiger beetle populations may depend on mitigating
threats posed during these periods when the beetles are at less-mobile and potentially
less-adaptable stages.
Unlike other coastal tiger beetle species, preferences or constraints of C. marginata
larval burrow-site locations are largely unstudied. Factors for other coastal
tiger beetle larvae that have been investigated include soil moisture level (Brust
et al. 2006), flood tolerance (Brust et al. 2005), particle size (Drummond 2002 in
USFWS 2009, Knisley 1997), bulk density (Drummond 2002 in USFWS 2009),
and substrate slope (Drummond 2002 in USFWS 2009, Knisley 1997). Larvae of
some coastal tiger beetle species—C. dorsalis dorsalis Say (Northeastern Beach
Tiger Beetle) and C. hirticollis—can move across the surface to relocate burrows
(Brust et al. 2006; USFWS 1994). In the case of C. hirticollis, larvae relocated
burrows in response to flooding and selected burrow sites based on soil moisture
(Brust et al. 2006). It is unknown whether larvae of C. marginata can relocate
burrows or what, if any, of the above-mentioned factors limit or affect larval
burrow location. These and other aspects of C. marginata biology are in need of
greater study.
Efficacy of site selection and site visits
In selecting sites, we targeted barrier beaches with unvegetated habitat patches
on the salt marsh side of dunes. This proved to be an effective method for finding
previously undocumented sites with C. marginata adults in Maine. However, at
least two sites in Maine that harbored C. marginata would almost certainly have
been missed had we used only this approach to select sites. Once at a site, we found
that searching the backdune–marsh strand was often an effective technique for finding
C. marginata adults. However, we also found adults and larval burrows (mostly
by chance) in other habitats not associated with the backdune of a barrier beach.
At more than one site, we searched the backdune–marsh strand unsuccessfully for
adults, but found adults and larval burrows in a different microhabitat nearby. We
conclude that targeting backdune–marsh strands associated with barrier beaches is
a good first step to locate adults of this species, but that a comprehensive inventory
should take a far-reaching approach in terms of site selection, microhabitats
searched, and life stages sought.
Comprehensive inventory for C. marginata should consider all sites associated
with tidal marsh ecosystems where there are sparsely vegetated, sandy areas at or
near the high-tide line. In regions where tidal fluctuations are great (e.g., the Gulf
of Maine), this may include locations somewhat removed from the immediate
coastline. Site searches should strive to include as many promising microhabitats
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as possible (Table 1). Searches for adults of this species should be accompanied by,
or supplemented with, larval-burrow searches. Documenting the presence of larvae
provides greater evidence of a reproductively viable population at a site, especially
if only low numbers of adults have been observed there. Larval burrow searches
offer the added benefit of being feasible outside (or on the extreme ends) of the
relatively narrow flight-period window, effectively lengthening the search season
for this species. We found that the shiny, blue-headed larvae of C. marginata were
relatively easy to distinguish in the field from C. hirticollis—the only other species
likely to occupy similar larval habitat in Maine.
Other tiger beetle species observed
Cicindela hirticollis and C. marginata adults were often found to co-inhabit
backdune–marsh strands, but C. hirticollis was more commonly found on oceanside
beaches than C. marginata. The number of C. hirticollis adults observed on
oceanside beaches was sometimes extremely high (in the hundreds and even
thousands). Cicindela hirticollis adults seemed less abundant on associated
backdune-marsh strands and absent from salt marsh microhabitats. By contrast,
the number of C. marginata adults recorded was lower than for C. hirticollis (only
approaching 100 at the most abundant site), and tended to be highest along the
backdune-marsh strand.
Conclusions
Our work increased the number of documented C. marginata sites in Maine, and
we did not find evidence of decline. Nevertheless, we recommend that this species
status continue to be special concern in Maine because its known distribution remains
limited to the southern third of the state, where most potential habitat has now
been surveyed and the tidal habitats to which it is restricted are subject to a variety
of current and emerging threats. Although tidal marshes in the northeast have shown
incredible resilience despite centuries of human impact (Gedan et al. 2011), continuing
pressures coupled with emerging threats, such as salt marsh die-off (Holdredge
et al. 2009) and sea-level rise associated with climate change, threaten to overwhelm
the inherent ability of tidal systems to adapt and persist (Gedan et al. 2011, Kirwan
et al. 2010) with potentially devastating consequences for species including C. marginata,
that are restricted to tidal habitats.
Acknowledgments
We are grateful for field observations made by Derek Dawson, Stewart DesMeules, and
Rob Lambert. Jason Czapiga provided valuable GIS-mapping support and assistance with
figures. We would also like to acknowledge assistance with permitting and site logistics
provided by Kate O’Brien of the Rachel Carson National Wildlife Refuge. Thanks to Sue
Bickford and Hannah Wilhelm of the Wells National Estuarine Research Reserve for reporting
that C. marginata had been recorded at York site 1 in July 2008 by Stewart DesMeules.
Financial support for this project came from the Maine Endangered and Nongame Wildlife
Fund and Maine Coastal and Inland Surface Oil Clean-up Fund.
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2014 Vol. 21, No. 4
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Brunswick, ME. Available online at www.manometmaine.org. Accessed 4 May 2011.
York Rivers Association. 2011. Healthy rivers, healthy Gulf of Maine. Available online at
http://www.yorkrivers.org/wheeler.html. Accessed 4 May 2011.
Northeastern Naturalist
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M.A.Ward and J.D. Mays
2014 Vol. 21, No. 4
Appendix 1. Observed numbers of C. marginata adults at survey sites in Maine, 2010.
*Indicates a site where C. marginata had previously been observed. **Indicates a site where
C. marginata had been previously observed, but was unknown to the co-authors prior to
this study.
Search # of # of
Site Name Lat. (N) Long. (W) Date effort (min) adults adults/hr
Kittery site 1 43°05' 70°40' 7/31/2010 42 0 0
Kittery site 2 43°05' 70°39' 7/31/2010 13 0 0
Kittery site 3 43°06' 70°39' 7/31/2010 56 0 0
York site 1** 43°08' 70°39' 7/31/2010 87 20 13.8
Ogunquit site 1* 43°15' 70°35' 7/13/2010 172 89 31
Wells site 1 43°19' 70°34' 7/26/2010 59 2 2
Wells site 2 43°19' 70°34' 7/26/2010 12 0 0
Wells site 3 43°19' 70°33' 7/26/2010 26 0 0
Wells site 3 43°19' 70°33' 8/1/2010 36 0 0
Wells site 4* 43°20' 70°32' 7/13/2010 75 30 24
Wells site 5 43°20' 70°32' 8/1/2010 75 0 0
Wells site 6 43°20' 70°32' 8/1/2010 15 1 4
Kennebunk site 1 43°20' 70°32' 7/16/2010 230 1 0.3
Kennebunk site 1 43°20' 70°32' 8/1/2010 58 0 0
Kennebunk site 2 43°20' 70°28' 7/15/2010 20 0 0
Kennebunkport site 1 43°22' 70°26' 7/26/2010 140 25 10.7
Kennebunkport site 2 43°23' 70°26' 7/15/2010 70 10 8.6
Kennebunkport site 3 43°23' 70°24' 8/1/2010 15 0 0
Saco site 1 43°29' 70°23' 7/15/2010 85 22 15.5
Scarborough site 1 43°32' 70°19' 7/12/2010 160 0 0
Scarborough site 2 43°32' 70°18' 7/12/2010 125 0 0
Scarborough site 3 43°33' 70°16' 7/15/2010 45 0 0
Phippsburg site 1 43°43' 69°49' 7/9/2010 105 1 0.6
Phippsburg site 2* 43°44' 69°48' 7/6/2010 85 40 28.2
Phippsburg site 2* 43°44' 69°48' 7/22/2010 370 39 6.3
Georgetown site 1 43°46' 69°44' 7/6/2010 160 0 0
Georgetown site 1 43°46' 69°44' 7/28/2010 180 0 0
Georgetown site 2* 43°46' 69°43' 7/28/2010 225 1 0.3
Bristol site 1 43°51' 69°31' 8/9/2010 57 0 0
Lincolnville site 1 44°17' 69°00' 8/12/2010 15 0 0
Lubec site 1 45°14' 66°59' 7/27/2010 300 0 0