Effect of the Edge on Eastern Cottontail Density: Urban Edges are Harder than Agricultural
Sofia Abisag Montes-Rodríguez1, Yury Glebskiy1,2*, and Zenón Cano-Santana1
1Laboratorio de Interacciones y Procesos Ecológicos, Departamento de Ecología y Recursos Naturales, Facultad de Ciencias, UNAM, Mexico City, Mexico. P.O. 04510. 2Posgrado en Ciencias Biológicas UNAM, Mexico City, Mexico. P.O. 04510. *Corresponding author.
Urban Naturalist, No. 72 (2024)
Abstract
Fragmentation is a very common process in today’s ecosystems, and understanding its effect on species is key to their management. In this particular case, we concentrate on the effect of a natural-urban edge on Eastern Cottontail population density. To estimate this, we measured Eastern Cottontail density using fecal pellet count on 21 transects drawn inwards from the edge of an urban reserve. A GLIM analysis was applied to the data and showed that Eastern Cottontail density was smaller at the edges of the reserve. Our result differs from previous studies and the common assumption that Eastern Cottontails benefit from the edges. This may be because previous studies concentrated on the natural-agricultural edge, which is generally softer and divides two areas that can provide certain resources for the Eastern Cottontail, whereas we studied a natural-urban border, which is more abrupt. Therefore, we propose incorporating buffer areas into the design of urban reserves.
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Urban Naturalist
S.A. Montes-Rodríguez, Y. Glebskiy, and Z. Cano-Santana
2024 No. 72
1
2024 Urban Naturalist 72:1–5
Effect of the Edge on Eastern Cottontail Density; Urban
Edges are Harder than Agricultural
Sofia Abisag Montes-Rodríguez1, Yury Glebskiy1,2*, and Zenón Cano-Santana1
Abstract: Fragmentation is a very common process in today’s ecosystems, and understanding its
effect on species is key to their management. In this particular case, we concentrate on the effect
of a natural-urban edge on Eastern Cottontail population density. To estimate this, we measured
Eastern Cottontail density using fecal pellet count on 21 transects drawn inwards from the edge
of an urban reserve. A GLIM analysis was applied to the data and showed that Eastern Cottontail
density was smaller at the edges of the reserve. Our result differs from previous studies and the
common assumption that Eastern Cottontails benefit from the edges. This may be because previous
studies concentrated on the natural-agricultural edge, which is generally softer and divides
two areas that can provide certain resources for the Eastern Cottontail, whereas we studied a
natural-urban border, which is more abrupt. Therefore, we propose incorporating buffer areas
into the design of urban reserves.
Introduction
Habitat fragmentation is one of the major ecological threats to animals since it can
isolate populations or divide natural areas into fragments too small to host certain species,
although this rule is not universal (Betts et al. 2019). At the same time, fragmentation
brings another challenge for conservation: the edge effect (Betts et al. 2019). The conditions
at the edges could be beneficial for certain species or detrimental for others, thus
reducing the effective area of conservation. However, it has to be noticed that the same
species could respond differently to edges between different vegetation types (Ries and
Sisk 2010), and understanding the effect of these edges is key to conservation efforts since
these responses determine how much fragmentation a species can endure and the optimal
design of a protected area.
In this case, we aim to understand the effect of the edge on Sylvilagus floridanus (Allen,
1890) (Eastern Cottontail). There is a fair amount of information on how edges affect
this rabbit (Bertolino et al. 2011a, b, Morgan & Gates 1983, Pierse et al. 2011, Roseberry
1998), and from it, we can draw a general conclusion that this species benefits from the
edges. However, the aforementioned studies concentrate on the edge between natural vegetation
and agricultural fields, which is a softer transition between two vegetation types,
both of which can provide cottontails with some resources. In our study, we concentrate
on a much sharper edge between natural vegetation and an urban area. In this case, the
disturbance by humans and vehicles presence is much higher and the resources provided
by the urban area are lower. At the same time, this type of edges is less studied, at least for
this species (although there are reports for the genus suggesting that the urban edge has
little effect on S. aquaticus; Stevens et al. 2023). Therefore, the objective of this study is
to understand the effect of an urban-natural edge on the density of Eastern Cottont ails.
1Laboratorio de Interacciones y Procesos Ecológicos, Departamento de Ecología y Recursos Naturales,
Facultad de Ciencias, UNAM, Mexico City, Mexico. P.O. 04510. 2Posgrado en Ciencias Biológicas
UNAM, Mexico City, Mexico. P.O. 04510. *Corresponding Author: agloti@ciencias.unam.mx
Associate Editor: José R. Ramírez-Garofalo, Rutgers University
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Materials and Methods
Study location
For this study, we chose El Pedregal de San Angel natural reserve (henceforth El Pedregal),
Mexico City, Mexico. This reserve has a surface area of 237 ha and is divided
into 18 fragments surrounded and separated by the urban areas of Mexico City (SEREPSA
2022). The vegetation of El Pedregal consists of well-preserved xerophytic shrubs (Lot
and Camarena 2009) that are separated from the urban areas by fences (Zambrano et al.
2016). This makes a perfect location for studying the edges between natural and urban
areas. The Eastern Cottontail is the main herbivore in the area (Glebskiy 2016) that increases
vegetation diversity (Fortis-Fernandez 2023), and is distributed in 4 El Pedregal
fragments (Dorantes-Villalobos 2017). For this study, we excluded the smallest one (the
buffer zone called “A8-Biológicas”, since it is too small, consists entirely of edge area,
and it is not clear if the population there is resident or Eastern Cottontails are visitors
from a bigger area). The areas we chose for this study are 23.8, 53.7, and 114.4 ha, and
are dominated by well-preserved natural vegetation. The three areas are surrounded by
fences that separate them from the urban areas, and in most cases, there is no vegetation
buffer between the reserve and urban infrastructure (roads and buildings). Fences prevent
Eastern Cottontails from exiting the reserve and protect them from vehicular collisions
and direct contact with humans, but there is no infrastructure that prevents them from approaching
the fences, and they offer limited protection against sound and light from the
urban area. For the purposes of this study, those fences were considered to be the border
between natural and urban areas.
Method
We used fecal pellets count as an indirect measure of Eastern Cottontail density; previous
studies have proven that this is an efficient method to estimate rabbit populations (Palomares
2001), and since the rugged terrain and vegetation of our study area prevent the use
of direct observation technics, this method has been commonly used in the area (Dorantes-
Villalobos 2017, Fortis-Fernandez 2023, Glebskiy et al. 2018). Previous studies show that
Eastern Cottontail density is greatly influenced by terrain ruggedness (Glebskiy et al. 2018);
therefore, this factor was also included in the analysis. We measured fecal pellet density
between February and March 2018 using 21 lines 51 m long; each line consisted of 34 quadrants
of 1.5 by 1.5 m (Fig. 1). All lines run from the edge fence inwards into the natural area;
the distance between lines was at least 150 m. Pellet density was measured by meticulously
inspecting the ground in 0.5 by 0.5 m quadrants. Terrain ruggedness was estimated using
by dropping a measuring tape on the terrain and dividing the resulting length by the linear
distance, similar to the method presented in Risk (1972) and Aronson and Swanson (1997);
however, in our case, we measured the ruggedness of each quadrant inde pendently.
Statistical analysis
Lines that contained no fecal pellets were excluded from the analysis, since those lines
provide no information on Eastern Cottontails. We used a GLIM (Generalized Linear
Model) with Poisson distribution to analyze the data. The response variable is the number
of pellets per 0.5 by 0.5 quadrant, and the distance from the edge and terrain ruggedness are
independent variables. To evaluate the contribution of each of the predictors, we evalu ated
the GLIM model using a step Akaike information criteria analysis.
Urban Naturalist
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Results
A total of 13 lines and 442 quadrants were used for this analysis (8 lines were excluded
since they did not contain pellets). The average pellet density was 22.8 per m2 (range:
0–628). The GLIM analysis showed an intercept of 1.723 and estimates of 0.0297 for distance
from edge and -2.8419 for terrain heterogeneity index (p < 0.001 in all cases). The
best model, according to the AIC, contained both predictors and scored 8535.6; the model
using only distance to the border scored 9875.4; and only terrain ruggedness scored 9008.9.
Discussion
Our results show that both terrain heterogeneity and distance from edge affected Eastern
Cottontail density. As previously reported (Glebskiy et al. 2018), Eastern Cottontails preferred
plain terrains. The interesting result here is that Eastern Cottontail density is higher
away from the edge, suggesting that the natural-urban edge is n ot suitable for Eastern Cottontails.
This conclusion differs from most previous studies that show that Eastern Cottontails
are tolerant to edges and are common in the natural-agricultural edges (Bertolino et al.
2011a, b; Morgan and Gates 1983; Pierse et al. 2011; Roseberry 1998). Therefore, we can
conclude that urban areas surpass the ability of Eastern Cottontails to withstand disturbance.
However, it is not clear what stops Eastern Cottontails from using edge zones in El Pedregal
and two main hypotheses can be proposed: 1) the direct human disturbance such as light,
sounds, smells, and trash thrown into the reserve scare Eastern Cottontails (although these
factors had no impact on Swamp rabbit; Stevens et al. 2023), or 2) the vegetation associated
with the edge is less suitable for their activities. In the last case, it is known that in the edges
Figure 1. Schematic of the data collection lines. Data from each 1.5 by 1.5 quadrant was used for the
GLIM analysis.
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of El Pedregal exotics plants dominate the landscape (Noguez-Ledesma and Cano-Santana
2020) and some exotic vegetation has been shown to decrease the Eastern Cottontail population
(Glebskiy et al. 2018). We consider that it is unlikely that the fences by themselves
affect Eastern Cottontails since in previous studies we placed fences to exclude Eastern Cottontails
from fragments of El Pedregal (Fortiz-Fernandez 2023, Glebskiy 2019) and those
fences (between natural vegetation) had no ef fect on Eastern Cottontails.
This suggests that for Eastern Cottontails, the effective conservation area in urban reserves
is smaller than the geographical area, this is important since many lagomorphs are sensible to
fragmentation (Barbour and Litvaitis 1993) and cannot inhabit most urban reserves (Mora et
al. 2020, Stevens et al. 2023, Verde-Aregoitia et al. 2015). Therefore, the ecological functions
of lagomorphs (vegetation consumption, seed dispersal, prey for many carnivores, among others;
Chapman et al. 1980, Glebskiy 2016, Glebskiy 2019) could be lost. And because of that,
conservation of this species (including S. floridanus) becomes essential to the ecosystems’
functions. Currently, the Eastern Cottontail is not an endangered species (Nielsen and Lanier
2019), however we consider that its conservation is important since losing this species even
on a local level could affect other trophic levels for example, in our location Fortiz-Fernandez
(2023) proved that in absence of Eastern Cottontails, plant diversity is diminished.
At the same time, given this species’ endurance to edges in several other habitats (Bertolino
et al. 2011a, b, Morgan and Gates 1983, Pierse et al. 2011, Roseberry 1998) it can be
hypothesized that some other species could be even more affected by edges inside urban
reserves, although more research is needed in this area. Therefore, we propose that the implementation
of buffer areas around urban reserves must be considered, probably in a form of
parks or gardens that soften the transition between the urbanized and strict protection areas.
Acknowledgements
We are thankful to M. J. Samano-Silva, I. Castellanos-Vargas for technical support, D. A. Parra-
Campos for helping with the illustration, and the working team of SEREPSA for permits for this project.
This article was financially supported by the PAPIIT project IN222816 “Ecología de Sylvilagus
floridanus (Lagomopha: Leporidae) en la Reserva Ecológica de Ciudad Univers itaria, D.F.”
References
Aronson, R.B., and D.W. Swanson. 1997. Video surveys of coral reefs: Uni-and multivariate applications.
Proceedings of the 8th International Coral Reef Symposiu m 2:1441–1446.
Barbour, M.S., and J.A. Litvaitis. 1993. Niche dimensions of New England cottontails in relation to
habitat patch size. Oecologia 95:321–327.
Bertolino, S., B. Ingegno, and M. Girardello. 2011a. Modelling the habitat requirements of invasive
Eastern Cottontail (Sylvilagus floridanus) introduced to Italy. European Journal of Wildlife Research
57(2):267–274.
Bertolino, S., A. Perrone, L. Gola, and R. Viterbi. 2011b. Population density and habitat use of the introduced
Eastern Cottontail (Sylvilagus floridanus) compared to the native European Hare (Lepus
europaeus). Zoological Studies 50(3):315–326.
Betts, M.G., C. Wolf, M. Pfeifer, C. Banks-Leite, V. Arroyo-Rodríguez, D.B. Ribeiro, J. Barlow, F.
Eigenbrod, D. Faria, R.J. Fletcher, A.S. Hadley, J.E. Hawes, R.D. Holt, B. Klingbeil, U. Kormann,
L. Lens, T. Levi, G.F. Medina-Rangel, S.L. Melles, D. Mezger, J.C. Morante-Filho, C. D.L. Orme,
C.A. Peres, B.T. Phalan, A. Pidgeon, H. Possingham, W.J. Ripple, E.M. Slade, E. Somarriba, J.A.
Tobias, J.M. Tylianakis, J.N. Urbina-Cardona, J.J. Valente, J.I. Watling, K. Wells, O.R. Wearn,
E. Wood, R. Young and R.M. Ewers. 2019. Extinction filters mediate the global effects of habitat
fragmentation on animals. Science 366(6470):1236–1239.
Urban Naturalist
S.A. Montes-Rodríguez, Y. Glebskiy, and Z. Cano-Santana
2024 No. 72
5
Chapman, J.A., J.G. Hockman, and C. Ojeda. 1980. Sylvilagus floridanus. Mammalian Species 136:1–8.
Dorantes, D. 2017. Distribución y abundancia del conejo castellano, Sylvilagus floridanus (Lagomorpha),
en la Reserva del Pedregal de San Ángel, Cd. Mx., México. Bs. Sc. Dissertation. Facultad de
Ciencias, Universidad Nacional Autónoma de México, Mexico City, Mexico. 56 pp.
Fortis-Fernandez, G. 2023. Efecto de 3 y 4 años de exclusión del conejo castellano (Sylvilagus floridanus)
sobre la estructura de la comunidad vegetal de la Reserva Ecológica del Pedregal de San
Ángel, Ciudad de México, México. Bs. Sc. Dissertation. Facultad de Ciencias, Universidad Nacional
Autónoma de México, Mexico City, Mexico. 78 pp.
Glebskiy, Y. 2016. Factores que afectan la distribución y abundancia del conejo castellano (Sylvilagus
floridanus) en la Reserva del Pedregal de San Ángel, D.F. (México). Bs Sc. Dissertation. Facultad
de Ciencias, Universidad Nacional Autónoma de México, Mexico City, Mexico. 36 pp.
Glebskiy Y. 2019. Efecto del conejo castellano (Sylvilagus floridanus) sobre la comunidad vegetal
del Pedregal de San Ángel. Masters Dissertation. Facultad de Ciencias, Universidad Nacional
Autónoma de México, Mexico City, Mexico. 50 pp.
Glebskiy, Y., I. Castellanos-Vargas, D. Dorantes-Villalobos, and Z. Cano-Santana. 2018. Role of
predators, terrain ruggedness, and plant composition for the abundance of the Eastern Cottontail
rabbit (Sylvilagus floridanus). The Southwestern Naturalist 63(1):59–63.
Lot, A., and P. Camarena. 2009. El Pedregal de San Ángel de la ciudad de México: Reserva ecológica
urbana de la Universidad Nacional. Pp. 19–25, In A. Lot, Z. Cano-Santana (Eds.). Biodiversidad
del Ecosistema del Pedregal de San Ángel. Universidad Nacional Autónoma de México, Mexico
City, Mexico.
Mora, J.M., S.M. Silva, L.I. López, M.K. Burnham–curtis, D.J. Wostenberg, J.H. French, and L.A.
Ruedas. 2020. Systematics, distribution, and conservation status of Dice’s cottontail, Sylvilagus
dicei Harris, 1932 (Mammalia, Lagomorpha, Leporidae), in Central America. Systematics and
Biodiversity 19(1):74–88.
Morgan, K.A., and J.E. Gates. 1983. Use of forest edge and strip vegetation by Eastern Cottontails.
The Journal of Wildlife Management 47(1):259–264.
Nielsen, C., and H.C. Lanier. 2019. Sylvilagus floridanus. The IUCN Red List of Threatened
Species 2019. Avaliable online at https://dx.doi.org/10.2305/IUCN.UK.2019-1.RLTS.
T41299A45191626.en. Accessed 17 August 2023.
Noguez-Ledesma, L.P., and Z. Cano-Santana. 2020. Efecto de borde sobre la estructura de la comunidad
de vegetal en el matorral xerófilo del Pedregal de San Ángel, Ciudad de México. Pp. 32, In J.
C. Herrera (Ed.). Jornadas de Diversidad Biológica y Recursos Naturales, Volume 1. Facultad de
Ciencias Biológicas, Universidad Juárez Autónoma de Durango, Gómez Palacio, Mexico. 162 pp.
Palomares, F. 2001. Comparison of 3 methods to estimate rabbit abundance in a Mediterranean environment.
Wildlife society bulletin 29:578–585.
Pierce, J.E., R.T. Larsen, J.T. Flinders, and J.C. Whiting. 2011. Fragmentation of sagebrush communities:
Does an increase in habitat edge impact pygmy rabbits? Animal Conservation 14(3):314–321.
Roseberry, J.L. 1998. Landscape characteristics and spatial patterns of Eastern Cottontail abundance
in Illinois. Transactions of the Illinois State Academy of Science 91:167–178.
Ries, L., and T.D. Sisk. 2010. What is an edge species? The implications of sensitivity to habitat edges.
Oikos 119(10):1636–1642.
Risk, M.J. 1972. Fish diversity on a coral reef in the Virgin Islands. Atoll Research Bulletin 153:6.
SEREPSA. 2022. Ficha Descriptiva de la REPSA. Available online at http://www.repsa.unam.mx/
index.php/objetivos/caracteristicas/ficha-descriptiva. Accessed 16 August 2022.
Stevens, T.K., A.M. Hale, and D.A. Williams. 2023. Environmental and anthropogenic variables influence
the distribution of a habitat specialist (Sylvilagus aquaticus) in a large urban forest. Conservation
Science and Practice 5:1–12.
Verde Arregoitia, L.D., K. Leach, N. Reid, and D.O. Fisher. 2015. Diversity, extinction, and threat
status in Lagomorphs. Ecography 38(11):1155–1165.
Zambrano, L., S. Rodríguez-Palacios, M. Pérez-Escobedo, G. Gil-Alarcón, P. Camarena, and A. Lot.
2016. La Reserva Ecológica del Pedregal de San Ángel: Atlas de Riesgos. Universidad Nacional
Autónoma de México, Mexico City, Mexico. 53 pp.