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Vertebrate Diversity in Stormwater Sewer Systems of Alachua County, Florida
Alan A. Ivory II1*, Matthew T. Hallett1,2, Brett Scheffers1, and Steve A. Johnson1
1Department of Wildlife Ecology and Conservation, University of Florida, 110 Newins-Ziegler Hall, Gainesville, FL 32611 USA. 2Conservation Department, Jacksonville Zoo and Gardens 370 Zoo Parkway, Jacksonville, FL 32218 USA. *Corresponding author.
Urban Naturalist, No. 75 (2024)
Abstract As urbanization continues, animals are increasingly compelled to navigate human-altered environments. Here we investigate wildlife use of stormwater sewer systems (SSS), a widespread, subterranean environment resulting from urbanization. We used camera traps to reveal how wildlife exploit subterranean pathways, shedding light on their presence within this anthropogenic context in Alachua County, Florida. From February to May 2023, we documented a total of 35 species of vertebrates within SSS, including amphibians, reptiles, and birds, although mammals dominated our sample. Raccoons and Southeastern Myotis accounted for more than half of all observations, signifying their prevalence and widespread presence within SSS. Our research offers a comprehensive exploration of vertebrate diversity within an unconventional urban habitat and provides valuable insights into the relationship between SSS and species utilization patterns. Ultimately, our research lays the groundwork for future studies and informs the development of ecologically conscientious urban planning strategies.
Urban Naturalist
Urban Naturalist
A.A. Ivory II, M.T. Hallett, B. Scheffers, and S.A. Johnson
2024 No. 75
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2024 Urban Naturalist 75:1–18
Vertebrate Diversity in Stormwater Sewer Systems of
Alachua County, Florida
Alan A. Ivory II1*, Matthew T. Hallett1,2, Brett Scheffers1, and Steve A. Johnson1
Abstract –As urbanization continues, animals are increasingly compelled to navigate
human-altered environments. Here we investigate wildlife use of stormwater sewer systems
(SSS), a widespread, subterranean environment resulting from urbanization. We used camera
traps to reveal how wildlife exploit subterranean pathways, shedding light on their presence
within this anthropogenic context in Alachua County, Florida. From February to May
2023, we documented a total of 35 species of vertebrates within SSS, including amphibians,
reptiles, and birds, although mammals dominated our sample. Raccoons and Southeastern
Myotis accounted for more than half of all observations, signifying their prevalence and
widespread presence within SSS. Our research offers a comprehensive exploration of vertebrate
diversity within an unconventional urban habitat and provides valuable insights into
the relationship between SSS and species utilization patterns. Ultimately, our research lays
the groundwork for future studies and informs the development of ecologically conscientious
urban planning strategies.
Introduction
As urbanization accelerates, animals are increasingly forced to interact with humanmodified
environments (Jokimaki et al. 2011, Messmer 2009). Development of urban centers
necessarily focuses on human needs and usually neglects the requirements of wildlife.
In addition to direct loss of habitat, urbanization fragments habitats and may create barriers
to animal movements, contributing to population declines (Czech et al. 2000, Lesbarreres
and Fahrig 2012, Markovchick-Nicholls et al. 2008). Impacts are particularly severe when
development and supporting infrastructure bisects or isolates wildlife habitat. For example,
roads that isolate wetland breeding sites from important upland habitats can be especially
detrimental to small vertebrates such as reptiles and amphibians (Schmidt and Zumbach
2008, van Gelder 1973). Roads may also be a significant source of mortality for mammals
and birds (Bond and Jones 2008, Dodd et al. 2004, Husby 2016, O rlowski 2008).
To mitigate the impact of roads on vertebrates, wildlife crossings are often built to
facilitate the movement of animals under or over roadways (Alexander and Waters 2000,
Bond and Jones 2008, Dodd et al. 2004). Historically, wildlife crossings have been constructed
primarily to promote the movement of large mammals (Askins 2012, Andis et al.
2017). However, numerous studies have demonstrated the conservation value of facilitating
the movement of smaller-bodied vertebrates via the installation of simple culvert-type
underpasses (Aresco 2003, 2005; Chen et al. 2021; Dodd et al. 2004). Nonetheless, due to
species-specific life history traits, no single underpass design is ideal across species. Factors
such as traffic noise, cover, light availability, substrate, water availability, and the dimen-
1Department of Wildlife Ecology and Conservation, University of Florida, 110 Newins-Ziegler Hall,
Gainesville, FL 32611 USA. 2Conservation Department, Jacksonville Zoo and Gardens
370 Zoo Parkway, Jacksonville, FL 32218 USA. *Corresponding author: alanivory34428@ufl.edu
Associate Editor: Travis Ryan, Butler Univesity
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sions of the culvert influence how likely wildlife are to use an underpass (Glista et al. 2009,
Jackson and Griffin 2000).
Typically, constructed wildlife crossings demand a large footprint to connect habitats
that are divided by a road (Jackson and Griffin 2000). Due to the space and resources required
for such structures, they are rarely built in urban environments, even though urban
areas are highly fragmented by roads, buildings, and residential areas (Andreu et al. 2017).
This forces urban wildlife to regularly cross roads to meet their basic needs, leading to
road mortality having a significant impact on urban animal populations (Riley et al. 2014).
Despite urban areas not typically being targeted as sites for installing wildlife crossings,
conservation actions in Europe have included constructing small-scale, urban passages for
mesomammals, such as Lutra lutra Linnaeus (Eurasian Otters) and Meles meles Linnaeus
(European Badgers; Philcox et al. 1999, Van der Zee et al. 1992). In the U.S., there has been
success with salamander underpass tunnels, which facilitate animal movements between
breeding and foraging locations (Aresco 2005, Bain et al. 2017, Hedrick et al. 2019). In
these studies, underpasses consisted of straight culverts of varying lengths, ranging in pipe
diameter from 25–100 cm. Pipes of this style and size are typical of stormwater sewer systems
(SSS) across the U.S. (Tran 2016, UF/IFAS Extension 2012).
To avoid flooding, most U.S. cities have SSS that move water from areas with impervious
surfaces (e.g., roads, sidewalks, parking lots) to retention ponds, lakes, and streams.
This is accomplished by diverting water from rainfall via open ditch systems or in a closed
stormwater sewer pipe below ground (Che et al. 2014). Throughout our study area in North
Florida, SSS and sanitary sewers are separate and serve distinct purposes. Sanitary sewers
transport waste from various sources to wastewater treatment plants, where it undergoes
treatment before being discharged into local water bodies. In contrast, stormwater sewers
channel rainwater away from roadways and other impervious surfaces via curb inlets and
grates and release it untreated into natural and manmade water bodies.
Although the wildlife value of stormwater systems has been studied, there is a lack of
research on how animals use the subterranean pipes of SSS. Existing studies have predominantly
examined animal use of retention ponds typically associated with SSS outflows. For
example, amphibians use stormwater retention ponds as breeding sites (Hale et al. 2015,
Scheffers and Paszkowski 2013), whereas birds use stormwater retention ponds for foraging
and breeding (Frederick and McGehee 1994, Sparling et al. 2007). Additionally, while
it is established that many species of wildlife will traverse straight, relatively short culverts
as road underpasses (Ascensao and Mira 2007, Bain et al. 2017, Dodd et al. 2004), the
propensity of animals to use more complex SSS (with multiple points of entry, nodes, and
branches) has received little attention, with the recent exception of Rattus sp. (Guo et al.
2023).
Many species that use SSS in urban habitats are generalists and respond positively to
urbanization and human influences (Graser et al. 2012, Parsons et al. 2018, Prange et al.
2003). Likewise, population density and abundance of synanthropic species may increase
near sources of water and adjacent to roads where resources may be aggregated (Bernasconi
et al. 2022, Bissonette and Rosa 2009, Fidino et al. 2016, Jeffress et al. 2011). The use of
an underground corridor system has the potential to increase survivorship, dispersal, and
population viability of urban wildlife. Additionally, a better understanding of the spatial
movement of wildlife through urban areas can help mitigate future human-wildlife conflicts
(Markovchick-Nicholls et al. 2008, Villalobos-Hoffman et al. 2022).
To better understand the importance of SSS as potential habitat and movement corridors
for vertebrates, we deployed wildlife cameras within pipes of SSS at numerous locations in
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Alachua, Co. Florida. Our objectives were to document which species were present (e.g.,
diversity) in the SSS of our study area as well as their frequency of occurrence (i.e., relative
abundance). We also sought to determine how various species were using these subterranean
corridors. Based on a literature search, our study is one of few to document wildlife use
of complex SSS beyond the use of simple culverts. Since SSS share similarities in materials
and pipe diameters with culverts but are more complex, interconnected, and typically found
in urban settings; they offer a unique research focus. Exploring SSS is important because
they offer enhanced connectivity opportunities to wildlife beyond what simple, straight
culverts can provide.
Materials and Methods
Study area
We conducted our study from February 2023 to May 2023 in Alachua County, Florida,
specifically targeting the SSS of the City of Gainesville and the campus of the University
of Florida. The study area is primarily urban, with intermixed swamp and hardwood upland
habitat (Andreu et al. 2017). The University of Florida’s Gainesville campus spans more
than 800 hectares and includes approximately 400 stormwater drains. Nearly 60% of the
University of Florida campus is covered by the Lake Alice sub-watershed, with about 40%
of this area covered by impervious surfaces such as roads and buildings (UF/IFAS Extension
2012). All stormwater from impervious surfaces within our study area drains into an
associated stormwater sewer system.
Study design
We used the Create Random Points tool in ArcGIS Pro (Environmental Systems Research
Institute, ESRI 2021) to select random manholes along sewer pipes within the SSS
of Alachua County. The SSS shapefiles were sourced from the City of Gainesville and the
University of Florida Facilities Services (UFFS). We selected 100 random points, then removed
sites if the pipe diameter was <30 cm diameter, an opening too small for a camera
trap to function properly (McCleery et al. 2014). If the site was on private property or if the
site was not an access point pre-approved by the City of Gainesville or the UFFS, the site
was excluded from the study. We also excluded locations with damaged or broken sewer
pipes.
We deployed 39 camera traps across 33 unique “sewersheds” in Alachua County, FL
(Appendix 1, Fig. 1) that were each left in the field for 60 trap nights per camera to ensure
sufficient sampling effort to confidently access wildlife diversity within each sewershed
(Hallett et al. 2019, Silver 2004). We define a sewershed as independent sewer pipes which
may or may not be connected in a network. Sewersheds with only two access points are
defined as simple culverts and those with more than two access points are classified as a
system (Fig. 2). Access points are where animals might enter or exit a sewershed including
pipe openings at the ends of the systems (typically at a body of water), curb inlets often associated
with a manhole cover (where runoff water from roads enters SSS), and grated inlets
(Fig. 3). These access points may double as light sources and are important as potential
ways for wildlife to enter and exit a sewershed.
For the systems we studied, pipe openings ranged from 30 to182 cm in diameter (mean
= 72 cm). Sewersheds in our study ranged from 11 m to 6912 m (mean = 942 m) in pipe
length and consisted of 13 simple culverts and 20 more complex systems.
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Figure 1. Locations of camera sites across Alachua County, Florida.
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Figure 2. Examples of 4 stormwater sewersheds in Alachua County, FL. Sewershed systems with
multiple access points denoted as nodes on the line features (A and B) and sewershed culverts with
a maximum of two access points (C and D). Linear, white areas indicate roads and darker shapes are
buildings and stormwater ponds.
Figure 3. Example access points into the SSS of Alachua County. Stormwater grate in a road (A). Pipe
openings at a body of water (B and C). Curb inlet on a roadside with a manhole (D). Photo credit: Alan
Ivory, UF/IFAS
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Camera-trap surveys
We deployed camera traps (Bushnell Care S-4K No Glow #119949C; Prime Combo Low
Glow #119932CB Bushnell®, KS, USA; Moultrie M-880 #MCG12594 Moultrie®, AL,
USA) at each of the 39 sites (Fig. 1). At sites with metal attachment points (i.e., manhole
cover), we used magnetic mounts to secure cameras facing down toward the entrance of
Figure 4. Example of camera-trap mount for magnetic attachment points such as manhole covers.
Photo credit: Alan Ivory, UF/IFAS
Figure 5. Diagram of camera mounting set-up in magnetic attach ment points.
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the pipe (Figs. 4, 5). At sites with pipes constructed of concrete or other materials that were
not magnetic, we attached cameras to a tree at the entrance of the pipe. Due to the nature
of stormwater sewer construction, the distance from the entrance to the pipe could not be
standardized, but, when possible, we set cameras mounted within or outside of SSS within 1
m of a pipe opening (Baker 2015, McCleary et al. 2014, Zeitler et al. 2023). Cameras were
active 24 hours a day, recording date and time, capturing a 10-second video upon detecting
motion, followed by a 30 second delay before the next recording. We used videos, rather
than still images to increase detection probability, due to the proximity of target species to
the cameras. When reviewing camera trap videos, we watched the full 10-second video clip
to confidently identify vertebrates to species even when an individual’s entire body was not
discernable. Videos of the same species that occurred within a period of 30 minutes were
excluded to ensure that camera visits were independent (Silver 2004). We used data from
the cameras to determine vertebrate diversity within the SSS, and to calculate the number of
detections by species, species relative abundance (RA), and naïve occupancy. We calculated
relative abundance by dividing the number of observations of a species by the number of
trap nights and multiplying by 100. Naïve occupancy was calculated by dividing the number
of sites where a species occurred by the total number of sites included in the study.
We deployed camera traps from February 2023 to May 2023, which encompasses late
winter and spring in north Florida. Camera traps were active for 14–92 trap nights/camera
(mean = 58.3 trap nights) for a total of 2273 trap-nights. The variation in trap nights per
camera is attributed to heavy rainfall causing flooding, as well as instances of camera theft.
Results
Of the 39 camera sites, 19 (49%) had more than 50 wildlife observations and 4 sites (10%)
did not record any observations. In total, we documented 3798 animal detections and recorded
35 unique vertebrate species, including 12 species of birds, four species of amphibians, seven
species of reptiles, and 12 species of mammals (Table 1). The majority of species documented
were detected at fewer than five camera sites, with 16 species being detected at just
a single site each. Nine species, mostly mammals, were detected at five or more sites (Table
1). Among the 35 species documented, 21 were observed within a stormwater sewer system
Table 1. Number of observations (Obs), relative abundance (RA), and naïve occupancy (NO) for each
species observed in Alachua County SSS. Relative abundance was calculated by dividing the number
of observations of a species by the number of trap nights and multiplying by 100. Naïve occupancy
was calculated by dividing the number of sites where a species occurred by the total number of sites
in the study.
Scientific Name Common Name Authority Obs RA NO
Mammal
Dasypus novemcinctus Nine-banded Armadillo Linnaeus, 1758 82 3.61 0.18
Didelphis virginiana Virginia Opossum Kerr, 1792 321 14.12 0.23
Felis catus Domestic Cat Linnaeus, 1758 50 2.20 0.28
Lontra canadensis River Otter von Schreber, 1776 29 1.28 0.05
Lynx rufus* Bobcat von Schreber, 1777 1 0.04 0.03
Myotis austroriparius Southeastern Myotis Rhoads, 1897 694 30.53 0.44
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Table 1. Number of observations (Obs), relative abundance (RA), and naïve occupancy (NO) for each
species observed in Alachua County SSS. Relative abundance was calculated by dividing the number
of observations of a species by the number of trap nights and multiplying by 100. Naïve occupancy
was calculated by dividing the number of sites where a species occurred by the total number of sites
in the study.
Scientific Name Common Name Authority Obs RA NO
Odocoileus virginianus* White-tailed Deer Zimmermann, 1780 3 0.13 0.03
Procyon lotor Raccoon Linnaeus, 1758 1810 79.63 0.54
Rattus rattus Black Rat Linnaeus, 1758 324 14.25 0.10
Scalopus aquaticus Eastern Mole Linnaeus, 1758 5 0.22 0.05
Sciurus carolinensis Eastern Gray Squirrel J. F. Gmelin, 1788 58 2.55 0.18
Sigmodon hispidus Hispid Cotton Rat Say and Ord, 1825 124 5.46 0.13
Reptile
Alligator mississippiensis American Alligator Daudin, 1801 50 2.20 0.13
Anolis sagrei Brown Anole Dumeril and Bibron, 1837 19 0.84 0.05
Coluber constrictor Eastern Racer Linnaeus, 1758 4 0.18 0.08
Nerodia fasciata* Banded Watersnake Linnaeus, 1766 1 0.04 0.03
Plestiodon laticeps* Broad-headed Skink Schneider, 1801 1 0.04 0.03
Storeria dekayi DeKay’s Brownsnake Holbrook, 1839 2 0.09 0.03
Trachemys scripta scripta Yellow-bellied Slider Schoepff, 1792 27 1.19 0.08
Amphibian
Anaxyrus terrestris Southern Toad Bonnaterre, 1789 1 0.04 0.03
Osteopilus septentrionalis Cuban Treefrog Duméril and Bibron, 1841 3 0.13 0.03
Rana grylio* Pig Frog Stejneger, 1901 1 0.04 0.03
Scaphiopus holbrooki Eastern Spadefoot Harlan, 1835 127 5.59 0.10
Bird
Anhinga anhinga Anhinga Linnaeus, 1766 13 0.57 0.03
Ardea alba* Great Egret Linnaeus, 1758 24 1.06 0.08
Ardea herodias* Great Blue Heron Linnaeus, 1758 2 0.09 0.03
Buteo jamaicensis* Red-tailed Hawk Gmelin, 1788 1 0.04 0.03
Cardinalis cardinalis* Northern Cardinal Linnaeus, 1758 1 0.04 0.03
Corvus brachyrhynchos* American Crow Brehm, 1822 2 0.09 0.05
Dumetella carolinensis* Gray Catbird Linnaeus, 1766 1 0.04 0.03
Egretta thula* Snowy Egret Molina, 1782 1 0.04 0.03
Parkesia noveboracensis Northern Waterthrush Gmelin, 1789 3 0.13 0.03
Thryothorus ludovacianus Carolina Wren Latham, 1790 10 0.44 0.15
Strix varia* Barred Owl Barton, 1799 1 0.04 0.03
Zenaida macroura* Mourning Dove Linnaeus, 1758 2 0.09 0.03
* signifies species was not observed entering the stormwater sewer system, but was observed on the camera trap.
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beyond a pipe access opening. Nine species of birds, one amphibian species, two species of
reptiles, and two species of mammals were only detected at the mouth of a stormwater sewer
pipe and never within a simple culvert or complex sewershed (Table 1).
Bird diversity
Of the 12 bird species documented (Table 1), only Thryothorus ludovicianus Latham
(Carolina Wren), Anhinga anhinga Linnaeus (Anhinga), and Parkesia noveboracensis
Gmelin (Northern Waterthrush) were observed within a sewershed away from a pipe opening.
Carolina Wrens were observed at six sites, and occasionally with nesting material,
while the other two species were observed at a single location.
Amphibian diversity
Three of the four species of anuran amphibians we documented were only detected at a
single camera site, for a combined five observations (Table 1). One of these, a Rana grylio
Stejneger (Pig Frog), was recorded at the entrance to a pipe. Scaphiopus holbrooki Harlan
(Eastern Spadefoot Toad) was the most frequently detected amphibian and was observed at
four camera sites, though 120 of the total 127 observations cam e from a single site.
Reptile diversity
Of the 33 sewersheds monitored in our study, only three were inundated with water for
the duration of the project. These three sites accounted for all the Trachemys scripta scripta
Schoepff (Yellow-bellied Slider) observations. An Alligator mississippiensis Daudin
(American Alligator) was also detected at two of the three inundated sites. We also observed
American alligators at three more culvert sites linked to either natural or artificial ponds,
each retaining water for over half of the study duration.
Mammal diversity
In addition to being the most species-rich taxon of vertebrates observed among all sites,
mammals were also observed most frequently, accounting for 3501 (92.2%) of the 3798
total camera-trap observations. We documented 12 species of mammals, ten of which were
observed within a sewershed beyond the entrance of a pipe opening (Table 1).
Procyon lotor Linnaeus (Northern Raccoon) was the most commonly observed (1810
occasions) and widespread (detected at 21 of the 39 camera sites) species in our study.
While raccoon detections were similar in simple culverts (865 observations) as compared
to the complex sewersheds (945 observations), raccoon relative abundance index (RAI) was
higher in culverts (RAI = 107.9) than in complex sewersheds (RAI = 64.2).
Among the 33 sewersheds, the second most commonly detected species was Myotis austroriparius
Rhoads (Southeastern Myotis Bat), with 694 bat observations across 17 camera
sites. Eleven of these sites were complex sewersheds and 6 sites were simple culverts, with
bat abundance being higher in complex sewersheds (RAI = 46.1) compared to simple culverts
(RAI = 2.0).
Both Rattus rattus Linnaeus (Black Rat) and Sigmodon hispidus Say and Ord (Hispid
Cotton Rat) were commonly observed, with 324 and 124 observations, respectively. Of the
four sites where we found Black Rats, 223 of the 324 observations came from one camera.
While we observed Lontra canadensis von Schreber (North American River Otter) on 29
occasions, otters were only detected at two of our sites, both of which were simple culverts.
The only mammals observed at pipe openings, and not within SSS, were Odocoileus virginianus
Zimmermann (White-tailed Deer) and Lynx rufus von Schreber (Bobcat).
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Discussion
Our study is one of the first to quantify the use of urban SSS by wildlife. We detected at
least one species at 90% of our camera trap sites, and nearly half of the camera traps recorded
at least 50 observations. While each of the classes of terrestrial vertebrates was found in the
SSS of Alachua County, Florida, mammals dominated our camera trap detections, accounting
for 92% of all observations. Birds, amphibians, and reptiles may not regularly use the SSS,
they nonetheless are able to take advantage of the resources the systems provide as well as
their associated wetlands.
We expected to detect rodents and mesomammals, such as raccoons, Didelphis virginiana
Kerr (Virginia Possums), and Dasypus novemcinctus Linnaeus (Nine-banded Armadillo),
based on previous corridor studies (Bond and Jones 2008, Chen et al. 2021). As there is little
evidence to suggest that SSS are frequented by birds, we did not expect to observe many birds
in our sites. Predators such as bobcats are known to use culvert crossings, and we expected
to document more than one observation (Ascensao and Mira 2007). Additionally, foxes are
common in suburban settings in Florida (Brown 1997), and we were surprised that we did not
detect them during our study.
Birds
Although we documented 12 species of birds across our study area, most of the observations
were dominated by just three species: Anhinga, Ardea alba Linnaeus (Great Egret), and
Carolina Wren. Additionally, only three species of birds were detected within SSS pipes, away
from a pipe opening (Anhinga, Northern Waterthrush, and Carolina Wren). The others were
detected at a pipe entrance, but apparently did not enter the sewer system. Although retention
ponds associated with SSS can provide foraging opportunities for birds (Fidorra et al. 2016),
the subterranean sewer pipes of the culverts and complex sewersheds that we studied were
apparently of little value to birds, with the possible exception of Carolina Wrens.
Anhingas were observed numerous times within a single culvert that was inundated with
water throughout our study period. Anhingas appeared to be using this culvert as a thoroughfare
to move between two permanent water bodies connected by this culvert. Our cameras
also documented wading birds at pipe openings of several sewersheds. The birds appeared
to be foraging, and it’s likely they were attracted to pipe openings because such areas may
concentrate prey (Frederick and McGehee 1994).
Carolina Wrens were detected at more unique camera locations (6) than any other bird in
our study. Additionally, they were the only species observed carrying nest materials within a
sewershed, suggesting they use SSS as nesting sites. Carolina Wrens are well known for their
habit of constructing nests in man-made structures like carports and mailboxes (Laskey 1948,
Nice and Thomas 1948). While their motivations for constructing nests in such locations are
poorly understood, similar to carports and mailboxes, SSS likely provide sheltered areas with
protection from inclement weather and nest predators, as well as a favorable microclimate for
thermoregulation (Labisky and Arnett 2006).
Amphibians
It seems unlikely that the 120 detections of Eastern Spadefoot Toads at a single cameratrap
location are indicative of a breeding aggregation within the sewershed, as the camera
was located at a curb inlet near a retention pond within a suburban neighborhood. We suspect
the toads were making terrestrial movements (possibly for breeding) and were directed to the
inlet by the curb and simply fell into the storm drain and subsequently detected by the camera.
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Although the sewer pipes themselves may not be important habitat for amphibians, it is well
documented that stormwater ponds and wetlands directly associated with SSS are important
breeding habitat (Scheffers and Paszkowski 2013, Hale et al. 2015).
Reptiles
We detected seven species of reptiles overall (3 species were only observed at a single
site), and documented reptiles more often at sites that held water for more than half of the
study period. The most frequently observed reptile (Alligator mississippiensis) was also documented
across the greatest number of sites (5). The most commonly observed reptiles were
aquatic and able to take advantage of the water bodies associated with the SSS (Aresco 2009).
Two of the three sites that were inundated with water for the duration of our study are
complex sewershed systems, suggesting the Yellow-bellied Slider that was observed there
had the opportunity to navigate the pipes further than a single road crossing. In all 3 sites that
documented sliders, the SSS connected 2 or more bodies of water, so the sliders are likely
using the pipes as corridors between ponds. Of the 5 sites, 4 of the sites where alligators were
observed were simple culverts, and for this reason most of the alligator observations were of
animals swimming from one pond to another (35 of 50 observations), thereby avoiding crossing
busy roads.
Mammals
Mammals were the most frequently and widely observed taxonomic group. Eleven of the
12 mammal species were recorded more than once, and 10 of 12 species were documented
at more than 1 site. Species such as raccoons, Virginia opossums, Nine-banded Armadillos,
rats, and Felis catus Linnaeus (Domestic Cat), which dominated our observations, do well in
urban environments (DeGregorio et al. 2021, Fidino et al. 2016, Graser et al. 2012, Parsons
et al. 2018, Prange et al. 2003, 2004). This association with urban areas is due in part to their
use of human resources, such as trash and artificial structures urban areas often provide (Feng
and Himsworth 2014, Webb et al. 2021).
We mainly observed Black Rats at sites nearest to downtown Gainesville and the University
of Florida campus, suggesting a positive relationship with human presence. Black
Rats are excellent climbers that can easily access SSS via curb inlets and open pipes, and are
known for their opportunistic foraging behavior and tolerance of human presence (Feng and
Himsworth 2014). Given their climbing ability and affinity for manmade structures, it was
not surprising that this introduced rodent was regularly detected by our cameras within several
sewersheds. We also documented 124 observations of native Cotton Rats across several
sewersheds. However, unlike Black rats, the locations of these observations were primarily at
the periphery of our study area, away from heavier human presence. Although Cotton Rats are
known pests of agricultural crops in rural settings and are one of the most common mammals
in Florida (Brown 1997), our data indicate that they apparently do not occur in abundance in
suburban and urban centers within our study area.
Many species of bats are known to inhabit urban areas (Lehrer et al. 2021, Webb et al. 2021),
and it is well established that bats commonly roost in culverts and under bridges (Keeley and
Tuttle 1999, Leivers et al. 2019). Culverts can provide a similar thermal regime as natural caves
and are therefore attractive to bats as roosting sites (Leivers et al. 2019). However, few studies
have evaluated bats in SSS. For example, Goehring (1957) documented Eptesicus fuscus
Beauvois (Big Brown Bat) using a storm sewer in Minnesota, U.S. as a winter roost site, and
Wojtaszyn et al. (2013) documented 6 species of bats (including 4 species of Myotis) inhabiting
storm drains in Poland as winter hibernacula. Additionally, Myotis grisescens A. H. Howell
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(Gray Bats) have been documented using storm sewers as maternity sites in Kansas (Hays and
Bingham 1964) and Arkansas, U.S. (Timmerman and McDaniel 1992).
We detected Southeastern Myotis at almost half of our camera sites, including simple
culverts (6 sites) and complex sewersheds (11 sites). However, based on the number of bats
documented per trap-night, they appear to prefer complex sewersheds over simple culverts
consisting of a single pipe open at each end. The conditions in more complex sewersheds
likely provide a microclimate (e.g., light, temperature, humidity) similar to natural caves,
which bats find attractive. Ten of the sites where we detected bats were in suburban neighborhoods,
and an additional 2 sites were in natural areas on the University of Florida campus.
Both types of sites (79.6% of total bat-occupied sites) have small roads with limited traffic.
This finding reinforces previous findings (Kerth and Melber 2009), which indicated that bat
activity is more common near roads with limited traffic as compared to wider, more heavilytrafficked
roads.
We documented bats throughout the duration of our study, but the frequency of observations
declined over time, suggesting the bats were using the SSS as winter roosts, like other
species of Myotis (Wojtaszyn et al. 2013). Southeastern Myotis in north Florida give birth in
mid-May (Elizabeth Braun de Torrez, Florida Fish and Wildlife Research Institute, Gainesville,
FL; Pers. Comm.), which is when our study ended, thus precluding our ability to make
any definitive statement about the utility of the sewer systems to serve as breeding sites. In
addition to roosting, the bats may have also been opportunistically foraging on insects within
the SSS. While their foraging behavior is typically associated with capturing flying insects,
we also observed instances of Southeastern Myotis landing on the floor of the sewer pipe to
capture insects. This behavior could potentially be interpreted as a substrate-gleaning foraging
strategy suitable for confined environments like SSS (Razak 2018). Clearly, additional monitoring
is required to better understand the importance of SSS to bats, many species of which
are declining across their range.
Raccoons were observed at most sites in our study. On several occasions our cameras documented
raccoons within SSS feeding on invertebrates (i.e., crayfish) and accompanied by their
young. This suggests that raccoons regularly use SSS as denning sites, to forage, and possibly
to avoid crossing roads. We also documented raccoons entering and exiting SSS by climbing
manhole ladder rungs as well as using curb inlets as access points for some sewersheds.
Domestic cats and Sciurus carolinensis J.F. Gmelin (Gray Squirrels) also used curb inlets
as access points. These species, like raccoons, are excellent climbers, so with the aid of ladder
rungs and/or sewer wall texture they are able to move freely between SSS and the urban
habitat above ground (Haigh et al. 2017, McClearn 1992, Moseby and Read 2006). Considering
these behaviors as well as the number and geographic breadth of raccoon observations, it
appears raccoon use of SSS of Alachua County is not by chance. Rather, our data suggest that
SSS provide habitat for raccoons in suburban areas. That said, the frequency of movement
between subterranean SSS and above ground habitats by raccoons is unknown and needs to
be studied to assess the relative importance of SSS for this common mesocarnivore.
The SSS we studied likely mimic natural habitats, such as caves and dens, used for roosting
and reproduction. For some species, the sewer systems likely provide protection from
predators as well as foraging opportunities. Raccoons and opossums had a naïve occupancy of
0.54 and 0.23, respectfully; rates similar to those of other urban camera trap studies (Soultan
et al. 2021). Similarly, Black Rats, opossums, raccoons, and Southeastern Myotis all have
a relative abundance greater than 10 (Table 1), suggesting frequent use of SSS. Our data
strongly suggest that the SSS in our study are regularly used by some mammals, namely raccoons
and bats, and to a lesser extent by opossums and invasive Black Rats. However, further
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study is needed to determine how often these species enter stormwater sewers and the extent
of their movements within the systems.
SSS management for wildlife
While it is well established that some species, including raccoons, Southeastern Myotis,
alligators, and aquatic turtles, intentionally enter SSS, smaller species of herpetofauna may
enter a stormwater sewer system inadvertently. For example, we documented a Storeria
dekayi Holbrook (DeKay’s Brownsnake) after a storm carried large amounts of water into the
system, possibly washing this small snake into the system. Additionally, curbs and their inlets
may act as a drift fence with pitfall traps for small reptiles and amphibians. Small individuals
may follow curbs until they fall into SSS via curb inlets. This would explain the numerous
observations of Eastern Spadefoot Toads at one of our sites. A better understanding of
how amphibians and small reptiles enter SSS could help in preventing them from becoming
trapped. If amphibians are falling in from the curb, and are unable to exit, exclusion devices
and climbing aids could be implemented to prevent such wildlife from being trapped within
SSS. In shallow curb inlets, ramps could be used to bridge the elevation gap, while in deeper
locations, ropes might act as climbing vines.
Amphibians and, potentially, small reptiles are likely underrepresented due to our sampling
methods and the camera traps reliance on an animal being warmer than its surroundings
to trigger image capture (Meek et al. 2012, Rovero et al. 2013). Aside from the American
Alligator and Yellow-bellied Slider, the remaining amphibian and reptile species likely did
not trigger the camera trap themselves, but instead their presence were only documented as a
byproduct of a mammal also being in the field of view. For this reason, the detection probabilities
of amphibian and reptile species are likely lower than the detection of the endothermic
species (mainly mammals, Hobbs and Brehme 2017)), which comprised the majority of our
observations. For example, Osteopilus septentrionalis Duméril and Bibron (Cuban Treefrogs)
are often heard calling from storm drains, and there is anecdotal evidence they will breed in
SSS under certain situations (Steve A. Johnson, personal observation). This invasive frog in
Florida is common in Gainesville, but we only documented three observations during our
study. We suspect this underrepresents their abundance in SSS as a result of camera trap bias.
To address the issue of underrepresentation of these ectothermic species, future studies could
include camera traps using time lapse image capture, where regardless of animal temperature
or motion, the camera records a video on a repeating interval for the duration of the study.
Our study focused on a small region in northern Florida, and future studies in other regions
should be conducted to determine if our findings are an anomaly or if SSS are in fact important
habitat for some species of urban wildlife. Alachua County, the UF campus, and the city of
Gainesville have numerous natural areas and an extensive tree canopy, so our findings may
differ from other cities where wildlife habitat is sparser. Moreover, our results are based on
four months of sampling, so seasonality is an important factor to explore further in determining
when and how wildlife use SSS. Nonetheless, our research appears to be among the first
of its kind to evaluate the uses of SSS by vertebrates, and therefore serves as a baseline for
similar studies elsewhere.
Additional research is needed to better understand the role of SSS in regions experiencing
variable levels of urbanization. Wildlife may be less likely to use these subterranean corridors
in less developed areas because preferred resources are available aboveground. However, in
more urbanized areas, artificial roosts are beneficial in promoting urban bat populations. Bats
seem to make use of SSS for roosting, but through the use of radio tracking, we could better
understand bat movement through SSS.
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Species from various taxonomic groups appear to use SSS as corridors to avoid roads,
potentially navigating through a network of pipes that link separated patches of habitat. While
we did not collect metrics related to road mortality or track movements of individuals, SSS
have the potential to increase connectivity of highly fragmented environments. Alternatively,
there is also the possibility that these confined corridors may lead to increased predation of
some species, such as raccoons preying on bats (Wojtaszyn et al. 2013). Regardless, a diversity
of vertebrates was found in the SSS of our study area, underscoring the need to consider
these subterranean systems in land-use planning and their potential to help mitigate potential
human-wildlife conflicts.
Acknowledgments
We thank the City of Gainesville, Florida Department of Transportation, City of Alachua, University
of Florida Facilities Services, Celebration Pointe, and Turkey Creek for access to sampling sites.
Thanks to our numerous volunteers who aided in our field samplin g efforts.
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Appendix 1. Camera-trap site identification and locations.
SiteID Lat Long Sewershed Type Deployment Type
13 29.6711 −82.3363 System Manhole
16 29.7136 −82.3589 System Open Pipe
17 29.7118 −82.3591 Culvert Grate
21 29.6612 −82.3433 System Manhole
23 29.6540 −82.3384 System Manhole
24 29.6881 −82.3822 System Manhole
25 29.6533 −82.3293 System Manhole
48 29.6486 −82.3412 System Grate
51 29.6462 −82.3514 System Open Pipe
52 29.7595 −82.4124 System Grate
53 29.7504 −82.4099 Culvert Grate
54 29.7580 −82.4167 System Grate
57 29.6349 −82.3688 System Grate
59 29.7108 −82.3586 Culvert Manhole
60 29.6845 −82.3806 System Manhole
64 29.5954 −82.3957 System Manhole
66 29.5957 −82.3946 System Manhole
67 29.5957 −82.3946 System Manhole
68 29.6386 −82.3525 System Grate
71 29.6629 −82.3581 System Manhole
72 29.6283 −82.3969 System Grate
75 29.6451 −82.3475 System Grate
76 29.6440 −82.3453 System Manhole
77 29.6880 −82.3822 System Manhole
78 29.6629 −82.3582 System Manhole
79 29.6236 −82.3977 Culvert Grate
81 29.6613 −82.3433 System Manhole
82 29.6351 −82.3616 Culvert Grate
83 29.6463 −82.3514 System Open Pipe
84 29.6414 −82.3461 Culvert Grate
85 29.6443 −82.3509 Culvert Open Pipe
86 29.6427 −82.3529 Culvert Open Pipe
89 29.6215 −82.3398 Culvert Grate
90 29.5278 −82.3077 Culvert Open Pipe
91 29.6247 −82.2584 Culvert Open Pipe
92 29.6688 −82.3353 System Manhole
93 29.6688 −82.3352 System Manhole
94 29.7627 −82.4187 Culvert Open Pipe
95 29.6307 −82.3662 Culvert Open Pipe