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Survey of a Coastal Tiger Beetle Species, Cicindela marginata Fabricius, in Maine
Mark A.Ward and Jonathan D. Mays

Northeastern Naturalist, Volume 21, Issue 4 (2014): 574–586

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Northeastern Naturalist 574 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 Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 575 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 Northeastern Naturalist 576 M.A.Ward and J.D. Mays 2014 Vol. 21, No. 4 (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. Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 577 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. Northeastern Naturalist 578 M.A.Ward and J.D. Mays 2014 Vol. 21, No. 4 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 Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 579 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 Northeastern Naturalist 580 M.A.Ward and J.D. Mays 2014 Vol. 21, No. 4 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. Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 581 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. Northeastern Naturalist 582 M.A.Ward and J.D. Mays 2014 Vol. 21, No. 4 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 Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 583 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. Northeastern Naturalist 584 M.A.Ward and J.D. Mays 2014 Vol. 21, No. 4 Literature Cited Brust, M.L., W.W. Hoback, K.F. Skinner, and C.B. Knisley. 2005. Differential immersion survival by populations of Cicindela hirticollis Say (Coleoptera: Cicindelidae). Annals of the Entomological Society of America 98(6):973–979. Brust, M.L., W.W. Hoback, K.F. Skinner, and C.B. Knisley. 2006. Movement of Cicindela hirticollis Say larvae in response to moisture and flooding. Journal of Insect Behavior 19(2):251–263. Choate, P.M., 2003. A Field Guide and Identification Manual for Florida and Eastern US Tiger Beetles. University Press of Florida, Gainesville, FL. 197 pp. Drummond, M.R. 2002. The effects of geophysical factors on the distribution of the Northeastern Beach Tiger Beetle, Cicindela dorsalis dorsalis Say. M.Sc. Thesis. Christopher Newport University, Newport News, VA. Dunn, G. 1981. The tiger beetles of New Hampshire. Cicindela 13:1–28. Gawler, S., and A. Cutko. 2010. Natural Landscapes of Maine: A Guide to Natural Communities and Ecosystems. Maine Natural Areas Program, Department of Conservation, Augusta, ME. 347 pp. Gedan, K.B., A.H. Altieri, and M.D. Bertness. 2011. Uncertain future of New England salt marshes. Marine Ecology Progress Series 434:229–337. Holdredge, C., M.D. Bertness, and A. Altieri. 2009. Role of crab herbivory in die-off of New England salt marshes. Conservation Biology 23:672–679. Kelley, J.T., A.R. Kelley, and O.H. Pilkey, Sr. 1989. Living with the Coast of Maine. Duke University Press, Durham, NC. 174 pp. Kirwan, M.L., G.R. Guntenspergen, A. D’Alpaos, J.T. Morris, S.M. Mudd, and S. Temmerman. 2010. Limits on the adaptability of coastal marshes to rising sea level. Geophysical Research Letters 37:L23401–L23405 doi:1029/2010GL0454489. Knisley, C.B. 1997. Microhabitat preferences for Cicindela dorsalis dorsalis, the Northeastern Tiger Beetle. Report to the VA Department of Agriculture and Consumer Service. Richmond, VA. Knisley, C.B., and T.D. Schultz. 1997. The Biology of Tiger Beetles and a Guide to the Species of the South Atlantic States. Special Publication #5. Virginia Museum of Natural History, Martinsville, VA. 210 pp. Leonard, J.G., and R.T. Bell. 1999. Northeastern Tiger Beetles: A Field Guide to Tiger Beetles of New England and Eastern Canada. CRC Press, Boca Raton, FL. 176 pp. Maine Geological Survey. 2002. Mile and Half Mile Beaches at Reid State Park. Available online at http://maine.gov/doc/nrimc/mgs/explore/marine/sites/apr02.htm. Accessed 7 7 February 2013. NatureServe. 2009. NatureServe Explorer: An online encyclopedia of life. Version 7.1. NatureServe, Arlington, VA. Available online at http://www.natureserve.org/explorer. Accessed 8 December 2009. Neil, K., and C.J. Majka. 2008. New records of tiger beetles (Coleoptera: Carabidae: Cicindelinae) in Nova Scotia. Journal of the Acadian Entomological Society 4:3–6. Pearson, D.L., and A.P. Vogler. 2001. Tiger Beetles: The Evolution, Ecology, and Diversity of the Cicindelids. Cornell University Press, Ithaca, NY. 333pp. Pearson, D.L., C.B. Knisley, and C.J. Kazilek. 2006. A Field Guide to the Tiger Beetles of the United States and Canada: Identification, Natural History, and Distribution of the Cicindelidae. Oxford University Press, Inc., New York, NY. 211 pp. Schlesinger, M.D., and P.G. Novak. 2011. Status and Conservation of an Imperiled Tiger Beetle Fauna in New York State, USA. Journal of Insect Conservation 15:839–852. Northeastern Naturalist Vol. 21, No. 4 M.A.Ward and J.D. Mays 2014 585 Sikes, D.S. 1998. Connecticut tiger beetle status survey (Carabidae: Cicindelinae) 1996– 1998: Conservation status, taxonomy and ecology. Research final report to The Connecticut Department of Environmental Protection. 72 pp. Available Online at http://www. biodiversity.uconn.edu/Collections/insects/CTBnew/projectreport.htm#Databasing. Accessed 26 September 2012. United Nations Environmental Program (UNEP). 2006. Marine and coastal ecosystems and human well-being: A synthesis report based on the findings of the millenium ecosystem assessment. 76 pp. Available Online at: http://www.unep.org/pdf/Completev6_LR.pdf. US Fish and Wildlife Service (USFWS). 1994. Northeastern Beach Tiger Beetle (Cicindela dorsalis dorsalis Say) Recovery Plan. Hadley, MA. 60 pp. USFWS. 2009. Northeastern Beach Tiger Beetle (Cicindela dorsalis dorsalis), five year review: Summary and evaluation. Gloucester, VA. 21 pp. Whitman, A., B. Vickery, P. deMaynadier, S. Stockwell, S. Walker, A.Cutko, and R. Houston. 2010. Climate change and biodiversity in Maine: A climate-change exposure summary for participants of the Maine climate-change species-vulnerability assessment. Manomet Center for Conservation Sciences (in collaboration with Maine Beginning With Habitat Climate Change Adaptation Working Group) Report NCI-2010-2. 22 pp. 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 586 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