Road Mortality of Reptiles and Other Wildlife at the Ojibway Complex and Greater Park Ecosystem in Southern

Jonathan D. C hoquette 1, 3 and LinDsey VaLLiant 2

1sCC ecological, P.o. Box 221 station a, Windsor, ontario n9a 6K1 2Wildlife Preservation Canada, Guelph, ontario n1h 6J2 Canada 3Corresponding author: [email protected]

Choquette, Jonathan D., and Lindsey Valliant. 2016. Road mortality of reptiles and other wildlife at the ojibway Prairie Complex and Greater Park ecosystem in southern ontario. Canadian Field-naturalist 130(1): 64–75. the ojibway Prairie Complex in Windsor contains the largest protected tallgrass prairie ecosystem in ontario and supports numerous species at risk. Despite its ecological significance, it is crossed by multiple high-traffic roads. Road mortality is a major threat to endangered species in Canada, particularly reptiles. the main goal of this study was to describe the nature and extent of vertebrate road mortality, with a focus on reptiles, on roads bisecting the ojibway Prairie Complex, and the Greater Park ecosystem, in Windsor and Lasalle, ontario. a systematic road mortality survey was conducted by bicycle along seven roads (12.5 km) in 2010, 2012, and 2013. also, opportunistic observations ( n = 103) spanning over 30 years were assembled from a variety of sources. in total, 2083 vertebrates (49 species), including 446 reptiles (11 species), were recorded “dead on road” during systematic surveys. the highest diversity of reptiles was recorded on Matchette Road, whereas the highest rate of reptile mortality was recorded on Malden Road. Reptile species at risk were killed on all roads surveyed. Combining sys - tematic and opportunistic data, we found seven reptile species at risk: Butler’s Gartersnake ( Thamnophis butleri ), eastern Foxsnake ( Pantherophis vulpinus ), eastern Massasauga ( Sistrurus catenatus catenatus ), Blanding’s turtle ( Emydoidea blandingii ), eastern Musk turtle ( Sternotherus odoratus ), northern Map turtle ( Graptemys geographica ), and snapping turtle ( Chelydra serpentina ). Reptile road mortality “hotspots” occurred where each road is intersected by a naturalized utility right-of-way. our results can be used to focus mitigation efforts in space and time to reduce mortality rates and enhance connectivity in the ojibway Prairie Complex and Greater Park ecosystem. Key Words: Reptiles; vertebrates; species at risk; road mortality; ojibway Prairie Complex; Windsor; Lasalle; utility right- of-way Introduction by residential, industrial, commercial, and agricultural our understanding of the negative impacts of roads lands as well as an extensive network of local, collector on wildlife is increasing. in Canada, road mortality is and arterial roads, in addition to a newly built provin - now considered a major threat to the persistence of en - cial highway. Many species of wildlife, including at-risk dangered species, particularly reptiles (e.g., Row et al. reptiles, have been observed killed on these roads over 2007). Road mortality surveys have been used in areas the previous three decades (P. Pratt, unpublished data). of ecological importance to identify the nature and ex - these data were collected opportunistically, and no at - tent of wildlife road mortality (ashley and Robinson tempt has been made to document road mortality in this 1996; Vijayakumar et al. 2001 ; smith and Dodd 2003; region systematically. Langen et al. 2007; Coelho et al. 2008; shepard et al. the main goal of this study was to describe the nature 2008), and to measure the effectiveness of mitigation and extent of vertebrate road mortality, with a focus on measures (Dodd et al. 2004; aresco 2005; Baxter- reptiles and species at risk, on roads bisecting the oPC Gilbert et al. 2015). and Greater Park ecosystem in the city of Windsor and the ojibway Prairie Complex (oPC), in extreme the town of Lasalle. our objectives were to identify south western ontario, is recognized as a Carolinian which vertebrate species are killed on roads, estimate Canada signature site (Johnson 2005); it contains the road mortality rates for each group, and identify spatial largest protected tallgrass prairie remnant in ontario, and temporal patterns of vertebrate road mortality. (Rodger 1998). this “complex” of tallgrass , savannahs, forests, and provincially significant wetlands Methods supports a multitude of regionally, provincially, and Systematic Road Mortality Surveys globally significant species of flora and fauna, some of seven collector and arterial roads in the study land - which are found nowhere else in Canada (City of Wind - scape were surveyed (Figure 1; table 1). they were sor 2013). Furthermore, as many as 10 reptile species divid ed into two groups, reflecting different survey ef - listed federally as at risk have been recently document - fort: in section B, all amphibians, birds, mammals, ed in the oPC and the surrounding greater park ecosys - snakes, and turtles found dead on a road were recorded tem (City of Windsor 2013; CoseWiC 2015). systematically, whereas, in section a, only dead snakes situated within an urban landscape, the oPC and and turtles were recorded systematically. also, more Greater Park ecosytem is surrounded and fragmented surveys were conducted in section B ( n = 157) than in A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalist’s Club.

64 ©The Ottawa Field-Naturalists’ Club (2016) 20 16 Choquette anD VaLLiant : R oaD MoRtaLity at oJiBWay PRaiRie CoMPLex 65

FiGuRe 1. Map of the ojibway Prairie Complex and Greater Park ecosystem study area showing approximate boundaries of protected areas and roads surveyed for dead vertebrates during 2010–2013. BohP = Black oak heritage Park, Cnhs = Candidate natural heritage site, LW esa = Lasalle Woods environmentally significant area, oP = ojibway Park, oPPnR = ojibway Prairie Provincial nature Reserve, sGna= spring Garden natural area, tC PsW = turkey Creek Provincially significant Wetland, and tPhP = tallgrass Prairie heritage Park.

taBLe 1. Length, traffic intensity, and adjacent land use for roads surveyed during a systematic road mortality study in the ojibway Prairie Complex and Greater Park ecosystem in Windsor and Lasalle, ontario, from 2010 to 2013. Length of Road estimated average adjacent survey route (km) (length surveyed, km) annual daily traffic* land use† section a (7.85) armanda street (1.17) 2200 (2006 data) 91% res, 9% row spring Garden Road (1.09) 2500 (2005 data) 81% res, 8% ins, 8% row, 3% com Malden Road (1.82) 8654 (2006 data) 67% res, 19% row, 6% com, 6% nat, 2% ins 8000 (2013 data) todd Lane (1.74) 9580–12 027 (2006 data) 85% res, 11% row, 4% rec 15 236 (2008 data) normandy Road (2.03) 6619–8744 (2006 data) 70% res, 18% nat, 6% rec, 4% row, 2% ins section B (4.75) Matchette Road (3.00) 6836–9300 (2006 data) 54% res, 38% nat, 6% row, 2% rec sprucewood avenue (1.75) 4619–6235 (2006 data) 58% res, 25% rec, 12% nat, 5% row 5700–9402 (2008 data) *sources: Dillon Consulting (2007, 2009); P. Bouliane, personal communication, 2014; County of essex (2014). †sources: City of Windsor (2007); town of Lasalle (2014). “ adjacent land use” is estimated using a Gis by dividing the length of road frontage for a given land use or zoning designation (both sides of the road) by the total length of road frontage (i.e., double the road length). note: com = commercial, ins = institutional, nat = natural heritage/environment, rec = recreational, res = residential, row = opened and unopened road right-of-way. 66 the CanaDian FieLD -n atuRaList Vol. 130

section a ( n = 135). Results for snakes and turtles from years (1984–2014) and included records of snakes and both sections were pooled. turtles found dead ( n = 106) and alive ( n = 17) on roads Roads were surveyed by bicycle at speeds of 12–17 in the study landscape. Many observations were pro - km/h on 3 days a week (on average every other day, ex - vided by staff at the ojibway nature Centre. in most cept section a was surveyed on average every 4 days in cases, records were verified by qualified personnel. all 2010) for 52 non-consecutive weeks. three technicians results reported here are based on our systematic data, conducted road surveys: surveyor 1 in 2010–2013, sur - unless otherwise specified. veyor 2 in 2012, and surveyor 3 in 2013. surveys took Data Analysis and Mapping place from May to mid-august in 2010 and 2013 and Mortality rates are reported either as number of dead from late-august to october in 2012 and 2013. (Data on road (DoR) individuals per 100 km surveyed (e.g., from a single late-april survey are combined with May [446 DoR /1798 km surveyed] × 100 = 24.8 DoR/ data.) the posted speed limit on all roads surveyed was 100 km surveyed) or as number of DoR individuals 50 km/h. the survey route was traveled in a loop such per km per survey (e.g., [446 DoR] / [11.5 km per sur - that both lanes of each road were surveyed and it took vey, on average] / [157 surveys] = 0.25 DoR/km/sur - about 3 h to complete a full survey. surveys were al - ways conducted with the flow of traffic; however, for vey). Rates per month or per survey road were calculat - each survey we alternated between completing section ed in the same way, but using only the relevant subset a or section B first. Most surveys (> 70%) were com - of the data. Frequency estimates (DoR/km/survey) are pleted between 1100 and 1700. When all effort is com - assumed to be representative of daily mortality rates bined, we surveyed the equivalent of almost 1800 km (i.e., DoR/ km/day) for birds, mammals, snakes, and of roads, an average of 300 km/month (range 244–382 amphibians, as other research has shown that most car - km/ month). casses of these animals remain on the road for only one For each specimen found dead on a road, the follow - day (enge and Wood 2002; DeGregorio et al. 2011; ing data were recorded: utM coordinates (accuracy of santos et al. 2011). For turtles, however, per survey 5–10 m), road name, and location on road (yellow line, rates may not be synonymous with daily rates as some centre of lane, white line, or shoulder). During periods investigators have found most specimens remain on the of high amphibian mortality, utM coordinates were road for two or more days (Langen et al. 2007; santos not recorded for these species; rather, these were tab - et al. 2011). ulated by pre-defined road segment. species, age class, to determine whether there were significant differ - and sex were also recorded when possible; however, ences in the numbers of dead animals recorded per tax - on across calendar months (data pooled by month, many amphibians (54%), birds (46%), and mammals 2 (31%) could not be identified to species. Photographs regardless of year), we made comparisons using a χ were taken of all species listed as at risk by the Com - goodness-of-fit test using sPss 22.0 (iBM, armonk, mittee on the status of endangered Wildlife in Canada new york, usa). Departures from expected equal fre - (CoseWiC 2015). quencies across all months were determined by resid - to avoid duplication of records, all dead specimens uals that were greater or less than the critical value of were removed from the road and discarded in adjacent ± 1.96. When a significant difference from an expect - vegetation or roadside swale. When physical removal ed frequency was found, pairwise goodness-of-fit tests was not possible (because of carcass condition or safe - were performed to determine which months were sig - ty concerns), specimens were left in situ and the white nificantly different from the others. sample sizes were line adjacent to the carcass was marked with a water- too low to compare monthly mortality rates for at-risk resistant cattle marker or lumber crayon. these marks turtles. continued to be visible for at least a week after mark - scientific names of all reptiles and amphibians fol - ing, which was generally sufficient to allow for the low Crother (2012). Maps were produced in arcGis speci men to be removed from the road by other means. version 9.1 (esRi, Redlands, California, usa). all dis - Carcass persistence rates were not estimated. all ani - tances were determined using the “Measure” tool in mals encountered alive on roads were noted and either arcGis. left in situ (if on the road shoulder) or helped across Spatial Analysis the road in the direction in which they appeared to be the distribution of reptile road mortality events was traveling. analyzed using siriema version 1.1 (Coelho et al. Opportunistic Data Collection 2006). Roadkill data were weighted for species at observations of snakes and turtles on roads, dead or risk (saR) using the following scheme based on the alive, were also solicited from various local naturalists CoseWiC (2015) list: non-saR/not at risk = 1, special (including the authors) and organizations to assemble concern = 2, threatened = 3, endangered = 4 (table 2); a dataset of opportunistic records. opportunistic data thus, “hotspots” (and any future mitigation efforts) were kept separate from those collected during our would be biased toward such species. (a sensitivity surveys. observations in this dataset spanned over 30 analysis on Malden Road demonstrated that the location 20 16 Choquette anD VaLLiant : R oaD MoRtaLity at oJiBWay PRaiRie CoMPLex 67

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of mortality hotspots was sensitive to the saR weight - Results ing.) Species Composition of Road Mortality the siriema analysis consisted of two steps (Coelho overall, 2083 vertebrates of 49 species were found et al. 2008). First, Ripley’s K function is used to test for dead during systematic surveys. this includes four significant spatial aggregations of road mortality events species of amphibians, 21 species of birds, 13 species (L(r) values) on each study road, and, if found, to deter - of mammals, five species of snakes, and six species of mine at which spatial scales (i.e., radius length) such turtles (table 2). eastern Gartersnakes ( Thamnophis aggregations occur. second, using a relevant radius sirtalis sirtalis ) and northern Brownsnakes ( Storeria length from step one as an input, hotspot analysis is dekayi dekayi ) made up the majority (70%) of snakes used to identify the relative locations along each road recorded, while Midland Painted turtles ( Chrysemys where significant spatial aggregations occur ( Nevents - picta marginata ) and snapping turtles ( Chelydra ser - Nsimulated ). We used a linear, as opposed to a two- pentina ) accounted for the majority (82%) of turtles. dimensional, K-function and hotspot analysis (Coelho on average, dead snakes were observed seven times et al. 2006) because all roads are linear (except for a more frequently than dead turtles (table 3). minor curve on two of the roads), and no major differ - Just over a fifth (21%) of all dead snakes and turtles ences were detected during an initial analysis of a sam - were saR. six such species were found in this study: ple of roads using both methods. Blanding’s turtle ( Emydoidea blandingii ), eastern For the Ripley’s K-function analysis, we used a 95% Musk turtle ( Sternotherus odoratus ), northern Map confidence interval (Ci), an initial radius of 0.025 km, turtle ( Graptemys geographica ), snapping turtle, But - a radius step of 0.025 km, and 100 simulations. For the ler’s Gartersnake ( Thamnophis butleri ), and eastern hotspot identification, we used a 95% Ci, 100 simula - Foxsnake ( Pantherophis vulpinus ). Butler’s Garter - tions, 500 road divisions, and two radii for each road: snakes, eastern Foxsnakes, and snapping turtles made a radius for which the greatest intensity of spatial clus - up the vast majority (92%) of saR records. on aver - tering was reported (from Ripley’s K) and a radius of age, dead saR snakes were observed twice as often as 0.050 km. two radii identified as having significant saR turtles (table 3). an additional saR, the eastern aggregations were used (one relatively longer than the Massasauga ( Sistrurus catenatus catenatus ) was found other), so that results could better inform placement of dead opportunistically during the study period (table both fine-scale (e.g., ecopassages) and broad-scale (e.g., 2). seven provincially listed snakes and turtles were barrier fencing or traffic calming) mitigation strategies observed dead at the oPC and Greater Park ecosystem, (Coelho et al. 2006) . only roads with significant spatial and, of these, three species appeared to be dispropor - aggregations of road mortality (based on Ripley’s K tionately represented. analysis) were subjected to hotspot identification. For Temporal and Spatial Patterns of Road Mortality north–south roads, kilometre 0.00 was set at the south - Mortality rates differed significantly between months ern end and for east–west roads, kilometre 0.00 was set 2 for amphibians ( χ = 1483.12, df = 5, P < 0.001; peak in at the western end. 2 July–august), birds ( χ = 73.87, df = 5, P < 0.001; peak

taBLe 3. Vertebrate mortality rates recorded during a systematic road mortality survey in the ojibway Prairie Complex and Greater Park ecosystem in Windsor and Lasalle, ontario, from 2010 to 2013. Rates for amphibians, birds, and mammals are based on data from section B only; rates for all other groups are based on combined data from both sections. above average mortality rates by month, above average mortality rates by average no./100 km road, no./100 km*† Vertebrate mortality rate, group no./100 km May Jun Jul aug sep oct nor Mal Mat spr tod amphibians 176.2 — — 607.2 192.3 — — Birds 24.7 32.3 48.0 40.4 — — — n/a Mammals 20.1 — 26.9 24.4 23.0 — — snakes 21.4 — — — 36.2 26.3 28.7 — 52.3 — — — turtles 3.4 6.0 5.9 — — 4.1 — — — 7.2 4.7 — species at risk all† 5.3 — 6.3 — — 9.1 7.1 — 9.4 7.4 — — snakes 3.8 — — — — 6.6 6.3 4.0 7.0 5.3 — 3.9 turtles† 1.4 — 3.2 — — 2.5 — — 2.5 2.1 1.8 — *nor = normandy Road, Mal = Malden Road, Mat = Matchette Road, spr = sprucewood avenue, tod = todd Lane. above average mortality rates for snakes, turtles, and species at risk were not observed on spring Garden or armanda roads. †χ2 tests were not conducted. 70 the CanaDian FieLD -n atuRaList Vol. 130

in May–July), snakes ( χ2 = 98.5, df = 5, P < 0.001; peak combined, were observed on Malden and Matchette in august–october), saR snakes ( χ2 = 32.8, df = 5, P < roads (table 3). the highest diversity of dead snakes 0.001; peak in september–october), and turtles ( χ2 = and turtles (11 species) and saR (six species) were 24.71, df = 5 , P < 0.001; peaks in May–June and observed on Matchette Road. three saR (Butler’s 2 september), but not for mammals ( χ = 6.08, df = 5, P = Gartersnake, eastern Foxsnake, and snapping turtle) 0.299). temporal patterns remained after controlling were each observed at least once during all months for the number of kilometres surveyed per month (Fig - and on most roads. ure 2, table 3). turtle mortality was hatchling-biased in May and Road Mortality Hotspots september (67% and 62%, respectively), but not in significant aggregations of snake and turtle road June (13%). over half (62%) of all dead saR turtles mortality events were detected at multiple scales on were found in June and september, whereas most dead five of seven roads: Matchette, Malden, spring Garden, non-hatchlings (70%) were found in June. snake mor - todd, and normandy roads (table 4, Figure 3). a ggre - tality was adult-biased (88%) in august, whereas, in gation intensity was highest at the scale of 0.300 km to september and october, mortality was more evenly 1.050 km for each of these five roads (table 4). signif - represented by younger age classes (55% and 67%, icant aggregations were also detected at the scale of respectively). over half (65%) of saR snake mortality 0.050 km for all five roads except spring Garden Road. was observed in september and october. above aver - significant dispersion was detected only on Matchette age mortality rates for saR turtles and saR snakes, Road ( table 4, Figure 3 ).

FiGuRe 2. Mortality rates recorded during 52 non-consecutive weeks from 2010 to 2013 on seven collector and arterial roads in the ojibway Prairie Complex and Greater Park ecosystem in Windsor and Lasalle, ontario.

taBLe 4. Radii with significant aggregations and dispersions (to the nearest 0.025 km) and locations of hotspots (to the nearest 0.050 km) determined during a reptile road mortality study in the ojibway Prairie Complex and Greater Park ecosystem, Windsor and Lasalle, ontario, from 2010 to 2013. Length (and relative location) of road mortality hotspots* with greatest intensity, km Radii with Radii with Radii with significant* highest significant* Finer scale Broader aggregations, aggregation dispersions, (0.050 km scale Road km intensity, km km radii) (radii varies)

Malden Road 0.025–1.375 1.050 n/a 0.150 (km 0.400 (km 0.000 – km 0.400); 1.475–1.550 1.125 0.200 – km 0.350) 0.250 km radius Matchette Road 0.025–1.400 0.825 1.675–2.725 0.200 (km 2.750 1.000 (km 2.000 – km – km 2.950) 3.000); 0.825 km radius normandy Road 0.025–1.225 0.850 n/a 0.300 (km 0.550 0.600 (km 0.650 – km 1.250); – km 0.850) 0.450 km radius todd Lane 0.025–0.600 0.300 n/a 0.200 (km 0.250 0.650 (km 0.000 – km 0.650); – km 0.450) 0.300 km radius spring Garden Road 0.225–0.325 0.475 n/a n/a n/a 0.375–0.400 0.450–0.500 *Based on a 95% confidence interval. 20 16 Choquette anD VaLLiant : R oaD MoRtaLity at oJiBWay PRaiRie CoMPLex 71 ] ) r ( L [

y t i s n e t n i

n o i t a g e r g g A

Radius length (km) FiGuRe 3. Ripley’s K analysis for significant spatial aggregations of reptile road mortality events recorded on seven roads during a systematic road mortality study in the ojibway Prairie Complex and Greater Park ecosystem in Windsor and Lasalle, ontario. in each graph, aggregation intensity [ L(r)] is a function of radius length [r (km)], and 95% confidence limits are represented by the two light black lines. significant aggregations of road mortality events occur where the bold black line exceeds the upper confidence limit. Results of hotspot analysis were aberrant for Malden, that bisects the study landscape (parallel natural gas and normandy, and spring Garden roads when using the high-voltage hydro transmission lines) and crosses four radii with the highest aggregation intensity for each of study roads (Matchette, Malden, todd, and normandy; these roads (1.050 km, 0.850 km, and 0.475 km, respec - Figures 1, 4). For each of these four roads, and at two tively). subsequent analyses were conducted for Mal - scales of analysis, the highest intensity road mortality den and normandy roads using the radii of the next hotspot occurred in close proximity to where each road smallest “peaks” in L(r) from the Ripley’s K analyses intersects the right-of-way (Figure 4). Depending on the (0.250 km and 0.450 km, respectively; Figure 3). spring scale of analysis used, approximately a third (33/82 or Garden Road was dropped from further analysis after 40%) to half (45/82 or 55%) of all saR records from trials with four radii continued to produce aberrant the four roads were observed within hotspots (a com - results. bined length of 0.850–2.650 km; table 4, Figure 3). snake and turtle road mortality hotspots appeared to be associated with the presence of a utility right-of-way 72 the CanaDian FieLD -n atuRaList Vol. 130 ) d e t a l u m i s N

– s t n e v e N ( y t i s n e t n i n o i t a g e r g g A

Distance (km) along road from survey start FiGuRe 4. hotspot analysis for significant spatial aggregations of reptile road mortality events recorded on four roads in the ojibway Prairie Complex and Greater Park ecosystem in Windsor and Lasalle, ontario. in each graph, aggregation inten - sity ( Nevents – Nsimulated ) is a function of distance (km) along road from survey start (i.e., km 0.0), and 95% confidence limits are represented by light gray horizontal lines. significant aggregations of road mortality events (i.e., hotspots) occur where the bold dashed line exceeds the upper confidence limit. aggregations of greatest intensity are highlighted in light gray, and their approximate extent (to the nearest 0.05 km) is indicated by the light gray vertical bars. the proportion of species at risk (saR) observations that occur within the hotspots (i.e., within the gray bars) is indicated. above each graph, roads are depicted as a solid black line and segments with significant aggregations of road mortality events are represented in light grey. the scale of analysis (i.e., radii) is indicated next to each road name. Black arrows indicate the approximate location of the utility right-of-way intersection with each road. (note: for normandy Road, 50% of saR records occured between km 0.550 and km 1.250) 20 16 Choquette anD VaLLiant : R oaD MoRtaLity at oJiBWay PRaiRie CoMPLex 73

Discussion enging pressure is relatively high in those areas. after Limitations of this Study placing fresh snake carcasses, DeGregorio et al. (2011) We recognize three major limitations to our study: found they were removed more often in certain habitat underestimation of the number of species found dead as types (forested versus dune habitats). also, santos et al. well as mortality rates; overgeneralization of temporal (2011) found that lower traffic rates facilitate scavenger patterns; and under-representation of spatial aggrega - access to carcasses. although scavengers are present in tions of roadkill. the study landscape (see table 2), we did not conduct ours is not a comprehensive list of all vertebrates tests to determine how scavenging pressure is dis - killed on roads in and surrounding the oPC. Consid - trib uted along our study roads. hotspots were not iden - ering the short time frame of our study, the fact that a tified on three roads — sprucewood, armanda, and large proportion (41%) of specimens we collected were spring Garden roads — likely because of low sample not identified to species, and that we only recorded 17% sizes. Fur thermore, our analysis was conducted using of the approximately 301 vertebrate species known data from only two full May–october periods, which locally (City of Windsor 2013), it is likely that addi - might not be sufficient to identify all hotspots. addi - tional spe cies are being killed on roads in the study tional spatial aggregations may become apparent with landscape. a larger dataset that spans a longer time. there is no doubt that the rates of road mortality that Impacts of Roads on Ojibway Prairie Vertebrates we observed are underestimates of the true mortality Roads are a conspicuous cause of anthropogenic rates in the study landscape during our study. there are mortality of vertebrates, especially snakes and species a number of reasons for this. First, most animals killed at risk, in the oPC and Greater Park ecosystem. the on roads are removed or obliterated within a day by average snake road mortality rate of 0.21/km/day (as - scavengers, high traffic volumes, or other forces (Kline suming [384 DoR snakes] / [11.5 km per survey, on and swann 1998; Clevenger et al. 2001; hels and average] / [157 surveys]) is higher than those reported Buchwald 2001; enge and Wood 2002; smith and at other ontario sites: 0.06/km/day at the Long Point Dodd 2003; DeGregorio et al. 2011; santos et al. 2011; Causeway, Port Rowan (ashley and Robinson 1996), Farmer and Brooks 2012), which could result in road 0.15/km/day at Rondeau and Point Pe - mortality being underestimated by a factor of 12–16 lee national Park combined (Farmer and Brooks 2012), (scavenging: slater 2002, as cited by DeGregorio et al. and 0.19/km/day at Dyer’s Bay, northern Bruce Penin - 2011). second, surveying by bicycle led to slight under - sula (Reed and Mackenzie 2010). estimates of mortality rates of small snakes compared We confirmed that seven reptile saR are being sub - with surveying on foot (s. Boyle, personal communi - jected to road mortality at the oPC and Greater Park cation, 2013). third, we did not survey for carcasses in ecosystem, and that mortality occurs on all major, and roadside swales, where some animals killed on roads some local, roads in the focal area. During our study, we likely ended up (e.g., Dodd et al. 2004). Fourth, we did estimate that saR were being killed on roads across not estimate or correct for differences in detection rates the oPC at a minimum average rate of 19 individuals between observers (i.e., observer bias). Finally, we sur - a month (assuming: [5.3 DoR/100 km surveyed] × veyed only the arterial and collector roads in the study [11.5 km surveyed/survey day, on average] = [0.61 area, not the multiple “local” roads, on which eastern DoR/day] × [30.7 days/month]). the population-level Foxsnakes have been confirmed killed in 2010, 2012, impacts of the rates of road mortality experienced by and 2013 (P. Pratt, unpublished data). saR in general, and each species in particular, re - the temporal patterns in road mortality that we ob - main un known. Regardless, road mortality undoubted - served after pooling data from all three survey years ly places additional pressure on small populations of may not be representative of within-year patterns. how - reptiles already experiencing a wide range of threats ever, more in-depth analyses of the data are hindered and stressors as a result of inhabiting an urban land - by unequal survey effort within years and, in 2013, a scape (Mitchell et al. 2008). Furthermore, management series of temporary road closures on our study roads. documents for at least two of these species highlight the Regardless, the patterns we observed for snakes and tur - need to address road mortality and habitat fragmenta - tles are consistent with the biology of these two faunal tion across their range ( agency 2013; groups. For example, high mortality of turtles in May oMnR 2011). and June can be explained by adult dispersal during a precedent has already been set for mitigating road nesting and emergence of hatchlings that overwintered. mortality at protected areas across ontario. For exam - also, high mortality of turtles and snakes from august ple, using today’s saR designations for reptiles (Cos - to october can be explained by emergence of neonates eWiC 2015), the diversity of saR affected by roads as well as snake dispersal to hibernation sites. at the oPC (seven) is equal to or greater than that ob - additional aggregations of road mortality may re - served at Long Point Provincial Park (seven saR: ash - main undetected in the study landscape because of ley and Robinson 1996), Rondeau Provincial Park scavenging pressure, low sample size, or short length (seven saR: Farmer and Brooks 2012), Point Pelee of study. some hotspots could be “masked” if scav - national Park (four saR: McKay and Brown 2007; 74 the CanaDian FieLD -n atuRaList Vol. 130

Farmer and Brooks 2012), and Bruce Peninsula nation - potential benefits to all vertebrate groups of permanent al Park (four saR: eco-Kare international 2010; Reed road closures or traffic speed reductions should not be and McKenzie 2010). efforts to mitigate road mor tal - overlooked (see Martinson 2009; Farmer and Brooks ity are complete or underway in all four of these parks. 2012). our results provide insight into where and when sim - ilar mitigation efforts would have the greatest impact Acknowledgements in terms of reducing road morality at the oPC and We thank R. Brooks, K. Gunsen, and C. McLaren for Greater Park ecosystem. their comments on earlier versions of this manuscript. Management Considerations Funding for the 2012–2013 portions of this study was Mitigation measures would produce the greatest ben - provided mainly through the Ministry of natural Re - efit for saR and other reptiles if they are prioritized at sources’ species at Risk stewardship Fund. Partial locations where saR road mortality is highest or dur - funding in 2012–2013 was also provided by ojibway ing periods of peak mortality rates, or both. our results Defence (made possible by a Climate Grant and a Pro - suggest that the following four roads should be target - ject Grant by Freshwater Future). Partial funding for ed: Malden, Matchette, normandy, and todd. at a min - a field technician in 2013 was provided by Carolini - imum, the installation of physical mitigation structures an Canada Coalition through their Conservation intern - (e.g., barrier walls or fencing), 150–300 m in length, at ship Program. Funding for previous years was provid - the intersection of each of these roads and the utility ed by ojibway Defence (through a grant from right-of-way, for a total of 850 m, would target road Freshwater Future) and the save ojibway Group. sections where, collectively, over a third of saR mor - B.-a. Jaksic and e. Primeau conducted systematic tality was recorded. if these structures were extended road mortality surveys in 2012 and 2013, respectively. to 400–1000 m, depending on the road, for a total of the ojibway nature Centre (onC) graciously provid - 2650 m, mitigation measures would target road sections ed a base of operations for the field technician in 2012 where just over half of all saR mortality was recorded and allowed us to gather incidental observation data on these roads. Furthermore, there is an opportunity for from their wildlife sightings book. Wildlife Preservation mitigation measures along the four roads mentioned Canada provided important administrative assistance above to contribute not only to reducing road mortality, and support to the lead author and field technicians but also to increasing landscape connectivity for snakes from 2013 to 2015. and turtles. other opportunistic records from the study land - the utility right-of-way that bisects the study land - scape were obtained from naturalists and citizen sci - scape is managed to prevent development of a forest entists (K. Fawdry, R. Jones, s. Marks, a. L. Meloche, canopy, thus providing a continuous corridor of suit - M. Montsch, n. Pancheshan, t. Preney, and J. Row). able open habitat for snakes, particularly in areas dom - thanks to the following organizations for providing op - inated by residential development or dense forest. portunistic records or allowing us to search their Results from this road mortality study, in combination databases: Ministry of natural Resources and with previous habitat suitability modeling for eastern Forestry (aylmer), natural heritage information Cen - Massasauga (Choquette 2011) and radiotelemetry data tre/ontario herpetofaunal summary, Windsor/essex on eastern Foxsnakes (s. Marks, personal communica - County hu mane society, and Wings Wildlife Rehabil - tion, 2013), suggest that the right-of-way is either used itation Centre. onC employees, s. Marks and R. as a movement corridor or has potential to function as Jones, provided assistance identifying unknown verte - a corridor for saR snakes in the study area. Consid - brates from digital photos. the essex Region Conser - ering the importance of protecting and restoring con - vation authority provided boundaries for Candidate nectivity in this fragmented landscape, mitigation work natural heritage sites in the study landscape and the done on roads in the vicinity of the utility right-of-way City of Windsor Planning Department provided should combine efforts to reduce road mortality and boundaries of the ojib way Prairie Complex. thank increase connectivity (e.g., diverting animals to newly you to P. Bouliane (transportation planner, City of installed, or existing culverts, with the use of barrier Windsor), s. Boyle (graduate student, Lakehead uni - fencing). versity) and s. Marks (species at risk reptile specialist, in addition to permanent solutions, and to address amec Foster Wheeler) for sharing relevant informa - the widespread nature of saR road mortality, tempo - tion. rary mitigation measures (e.g., seasonal road closures, seasonal speed limit reductions, etc.) could be used dur - Literature Cited ing peak mortality periods for certain taxa and age Aresco, M. J. 2005. Mitigation measures to reduce highway mortality of turtles and other herpetofauna at a north Flori - classes, at a minimum, in June, september, and octo - da lake. Journal of Wildlife Management 69: 549–560. ber, when relatively high mortality rates of snake and Ashley, E. P., and J. T. Robinson. 1996. Road mortality of turtle saR were observed. efforts could be further ex - amphibians, reptiles and other wildlife on the Long Point tended to include May and august to cover additional Causeway, , ontario. Canadian Field-naturalist periods of high turtle and snake mortality. 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