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The small, the forgotten and the dead: Highway

impact on vertebrates and its implications for
mitigation strategies
Article in Biodiversity and Conservation February 2013
DOI: 10.1007/s10531-012-0396-x

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Biodivers Conserv (2013) 22:325342

DOI 10.1007/s10531-012-0396-x
ORIGINAL PAPER

The small, the forgotten and the dead: highway impact

on vertebrates and its implications for mitigation
strategies
Alberto Gonzalez-Gallina Griselda Bentez-Badillo
Octavio R. Rojas-Soto Mircea G. Hidalgo-Mihart

Received: 4 May 2012 / Accepted: 30 October 2012 / Published online: 19 December 2012
Springer Science+Business Media Dordrecht 2012

Abstract Roads affect wildlife in many ways, and probably the most conspicuous of
these is by producing roadkills. In Mexico few studies have addressed this issue. In the
present study, we look at the richness and abundance of vertebrate roadkills along a 14 km
stretch of the Amozoc-Cantona-Perote highway, at the border of the states of Veracruz and
Puebla and across two environments: shrubland and grassland. The highway was sampled
on 34 days between June 2010 and March 2011. A total of 946 animals were recorded as
killed by traffic, and belonged to 58 species. Ten of these are under some protection status
by Mexican law. Diversity values for shrubland were D = 0.52 and for grassland D = 1.2,
both with BrayCurtis similarity values of Ibc = 0.12 and complementarity (CAB) was
98.5. The most common roadkilled animals were rodents (genus Peromyscus). We estimate
that 11,899 vertebrates are killed annually in this section of the highway. This study
highlights the need for designing and applying policies and actions for road construction
and maintenance that, together with drivers education, will allow us to increase landscape
connectivity and decrease wildlife roadkills.
Keywords

Impact  Highway  Mexico  Richness  Roadkills  Vertebrates

A. Gonzalez-Gallina (&)  G. Bentez-Badillo  O. R. Rojas-Soto

Departamento de Ambiente y Sustentabilidad, Instituto de Ecologa A.C., Carretera Antigua a
Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico
e-mail: fodopo@hotmail.com
G. Bentez-Badillo
e-mail: griselda.benitez@inecol.edu.mx
O. R. Rojas-Soto
e-mail: octavio.rojas@inecol.edu.mx
M. G. Hidalgo-Mihart
Division Academica de Ciencias Biologicas, Universidad Juarez Autonoma de Tabasco, km 0.5
Carretera Villahermosa-Cardenas, 86039 Villahermosa, Tabasco, Mexico
e-mail: mhidalgo@yahoo.com

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Introduction
The effects of roads on environment have been widely documented According to Trombulak and Frisell (2000) road affects the environment by: (1) altering the natural habitat,
(2) dispersing exotic species, (3) inducing behavioral modifications, (4) causing direct
mortality from road construction and also (5) from wildlife-vehicle collisions (roadkills).
The latter is a permanent effect and hence potentially has a greater overall effect. Road
construction tends to reduce landscape permeability and impacts population movement
dynamics with the result that the populations that are more isolated and smaller become
more prone to local extinction (Bissonette 2002).
The factors that affect highway-related wildlife mortality in different ways can be
divided into: (a) those of the road itself, i.e., the relationship between structure and
landscape; (b) those related to the driver, such as the speed of the vehicle and the drivers
agility for maneuvering to avoid colliding with the animals; (c) the weather and its effect
on driving conditions, such as rain or fog; (d) those characteristics of the species themselves. Regarding the latter, species with low densities and reproductive rates, together
with long-lived generations are more vulnerable, the same way that species with high
intrinsic mobility (such as habitat generalists) are typically more prone to roadkills mortality (Case 1978; Lye 1987; With and King 1999; Foufopolous and Ives 1999; Forman
et al. 2003).
Some authors suggest that roadkill rates can exceed those of natural mortality (Ferreras
et al. 1992) and this will affect population density if there is no natural compensation
mechanism of increased breeding (Fahrig et al. 1995; Huijser and Bergers 2000). Similarly,
imbalances can occur in the sex structure of populations (Gibbs and Steen 2005) and for
many species, highway related mortality can serve as a limiting factor for foraging and
dispersion. Although the impact of road related mortality is not as notable in populations
with high abundances, it may have a considerable effect on threatened or endangered
species, which are characterized by their low abundances (Forman and Alexander 1998;
Glista et al. 2007). Roads and their traffic could, in fact, be compared to a new predator for
which the animals have not developed any defense mechanisms or behavioral adaptations
that might increase their survival. As a result, some populations become unstable due to the
additional mortality of adult individuals (Andrews et al. 2006).
Roadkills have been documented for over 50 years (Stoner 1925, 1936; Haugen 1944;
Finnis 1960) and in several countries (Bashore et al. 1985; Reijnen et al. 1995; Mumme
et al. 2000). However, to date, the information available about the impact of roads on
wildlife in biodiverse regions such as the Neotropics is still mostly in an early descriptive
phase and there is little information compared to that available for the Holarctic regions.
There, the majority of recent studies focus on practical aspects. The contrast is noteworthy
given the rapid growth in infrastructure and traffic of the tropics. In Latin America, road
ecology is a relatively new field of research, with only a few studies addressing the effects
of transportation infrastructure on wildlife. As this line of research grows, it will be of
particular importance to address whether there are differences in how individuals and
species are affected.
Studies have been done in Brazil (Pfeifer et al. 2008; Ferreira Da Cunha et al. 2010;
Caceres et al. 2010), Colombia (Arroyave et al. 2006; Delgado-V 2007); Costa Rica
(Monge-Najera 1996; Gomez-Figueroa and Monge-Najera 2000), Panama (Oliveira da
Silva et al. 2007) and Argentina (Comita 1984; Barri 2010). However, in Mexico, there
have been few studies of road ecology (Polaco and Guzman 1991, 1993; Morales-Mavil
et al. 1997; Becerril-Morales 2001; Grosselet et al. 2008; Lara-Gomez 2008; Pozo-Montuy

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327

et al. 2008); unfortunately most of them are unpublished or constrained to thesis, or

Environmental Impact Assessment (EIA) requested by the Department of the Environment
and Natural Resources (SEMARNAT) that are available until authorized and published in
the official gazette of SEMARNAT (Gaceta Ecologica). Under Mexican law, all those
involved in infrastructure development projects, such as highway expansion, must present
the authorities with an evaluation of the environmental impact that their project will
produce (Mexican law: Ley General del Equilibrio Ecologico y la Proteccion al Ambiente,
Artculo 28, Reglamento de la LGEEPA en materia de Impacto Ambiental). The absence of
specific data on the impact of highways has important consequences when mitigation
measures are proposed in the EISs. Thus, roadkill assessment for Mexican highways is
important because it will allow us to document the importance of roadkills to wildlife
populations, and improve the way new roads are built and existing roads are upgraded.
Mexico offers a good place to study this phenomenon in Latin America considering that
over the next 10 years the road system across the country is going to be significantly
expanded (SCT 2007). The state of Veracruz, where this study was done, is one of the most
biodiverse in Mexico, but it is also one of those most fragmented by anthropogenic
activities. Currently in Veracruz, the construction of five new highways is underway or
being planned (Gobierno del Estado de Veracruz 2011).
The main objective of this study was to analyze the richness and abundance of vertebrate
roadkills along a segment of highway in central Veracruz in order to estimate the potential
impact of the highway on its associated wildlife. The information obtained should allow us to
make recommendations for mitigating any impact detected, particularly for small species,
which are usually underestimated in EIAs and several of which are protected by law.

Materials and methods

Study area
The study area is located in the semi-arid region of the states of Puebla and Veracruz, at the
juxtaposition of two mountain ranges: the Cofre de Perote Mountain and the Sierra Norte
of Puebla mountain range, which form part of the Tehuacan-Cuicatlan Valley Province. It
is characterized by a degree of high endemicity in its flora (Smith 1965) and by the
presence of species with vicariant distributions from the dry zones in the north of the
country (Rzedowski 1978), making this an area of particular importance to the biological
diversity of the state of Veracruz.
Elevation is 2,300 and 2,500 m a.s.l. and forms the semi-arid endorheic Perote-Libres
Basin. Climate is at the humid extreme of the semi-arid classification, similar to temperate
with a warm, rainy summer. Mean annual temperature is 14 C, mean annual precipitation
is 500 mm, and mean annual evaporation is 1,700 mm. The Cofre de Perote Mountain has
a shadow effect on the region and there are dry winds from the north and southeast. The
vegetation of the area is highly fragmented with several vegetation types: pine, cypress and
oak forests interspersed with grassland, shrubland, and temporal crops (Gerez-Fernandez
1985).
The highway segment we studied runs NE-SW and connects Amozoc-Cantona-Perote
(ACP) and is classified as an A2 highway (SCT 2011). It has a crown width of 12 m
surfaced with asphalt, and has two lanes that are each 3.5 m wide and two shoulders that
are 2.5 m wide. This type of road is apt for speeds of 90110 km h-1. The main body of
the road was built to one side of the right of way to leave space for the second stage of the

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project during which two more lanes will be added. It is almost 112 km long in total.
105 km corresponds to the Amozoc-Perote axis and there is a 7 km-long branch extending
from the Cantona turnoff to the archeological site of the same name. Of the 104.96 kmlong segment, 15.0 km lie within the state of Veracruz, and of the remaining 91.52 ha,
59.43 ha belongs to the town of Totalco and 28.38 % to the town of Perote (Fig. 1). In
2010, 22,988 vehicles crossed this highway and 24,082 in 2011, giving us a traffic flow
average of 62.98 vehicles per day in 2010 and 65.97 for 2011. Fitch Ratings informed that
the highway traffic flow incremented 4.8 % from 2010 to 2011, and since its construction
(2004) until 2011 it has incremented at a 33.2 % every year (Imagen Poblana 2012).
The highway segment we studied runs through two different environments. The first is
associated with shrubland that grows on a lava spill from the Xalapaxquillo volcano
(badlands, malpas in Spanish) (from km 90 to 98 ? 360) and the second is flat grassland
known as a llano in Spanish (from km 98 ? 360 to 103 ? 820) approximately 14 km in
length (Fig. 1). The malpas is characterized by a shrubland formed mainly by the following species: Hechtia roseana, Agave obscura, Nolina parviflora, Yuca periculosa,
Opuntia sp., Juniperus deppeana, Dasylirion acrotriche, Muhlemebergia sp., Croton
dioicus, Quercus microphylla, Mimosa biuncifera and Bouteloua curtipendula (Ingenieros
Civiles de Mexico 2002). It seems to be a secondary community maintained by fire where
Nolina parviflora appears to be favored by the local burning (Rzedowski 1978). The area,
especially the grassland had notable signs of disturbance and defaunation, having been rid
of big wildlife since a long time ago (deer, cougars, etc.) only medium-size (coyotes, hares,
etc.) to small fauna remains. The ACP highway was opened on July 2, 2004, and divided
the malpas, which to that time had been relatively well preserved, from the llano which,
being appropriate for crops, has a long history of human activity.

Fig. 1 Amozoc-Cantona-Perote highway (double black line), the study area falls along the section that lies
in the state of Veracruz (thicker double line) (ca. 14 km) and crosses two environments: a shrubland on a
relatively well preserved lava spill (badlands or malpas) and a flat grassland that highly fragmented by
agriculture (llano)

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Vertebrate roadkill search

This study took place from June 2010 to May 2011, during which there were a total of 34
sampling days (11 fieldtrips with a mean duration of 3 days). We searched for vertebrate
roadkills on the asphalted surface together with culverts while driving a vehicle along the
roads shoulder at a speed of 30 km h-1. Surveys were done between 10:00 and 14:00 h so
we could collect nocturnal and diurnal animals. When encountered, roadkill was identified
(whenever possible to the species level) and photographed. Each animal was removed from
the roads surface to avoid duplicate records. On each fieldtrip, the first day dedicated to
removing all of the animals that had accumulated to that point so we could be sure that the
following days we would be recording daily roadkills rather than the sum from the previous days, and starting from zero for estimating the annual death toll. Data obtained
during the cleaning passes were used for overall richness and abundance counts.
Persistence of roadkills on the road have been considered as an important factor to be
considered on roadkill counts (CITA). To deal with this issue, we considered the decay values of
road kill estimated by Santos et al. (2011) on road kill decay. Their estimation for all taxa
persistence time, as a mean, was one day and corresponded to a 0.5 probability of persistence.
This decreased to 0.241 on the second day with a substantial drop afterwards. Maximum
persistence time varied among different taxonomic groups, ranging from 4 (lizards) to 158 days
(carnivores). Due to these, following Santos et al. (2011) recommendation in which the
advisable periodicity of road monitoring is daily monitoring specially for all small species.
We calculated species richness (number of vertebrate species found as roadkills) and
abundance (number of individual animals found as roadkill during the study) for each
group observed (amphibians, reptiles, birds and mammals). Owing to the low number of
roadkill observations for amphibians, amphibians and reptiles were combined into the
group we refer to as herpetofauna.
Data analysis
We calculated the diversity for each environment using Simpsons index (Simpson 1949).
To estimate the number of species expected for each environment we used species accumulation curves for each vertebrate group based on the cumulative number of species for
each sampling day. For this analysis we used EstimateS 7.5 software (Colwell 2005) with
the jackknife nonparametric first order estimator (Jack1). Similarity amongst sites (Ibc) was
calculated using the BrayCurtis formula (Bray and Curtis 1957):
Ibc

2Nab
Na Nb

where Na = total number of individuals at site A (malpas), Nb = total number of individuals at site B (llano), and Nab = sum of the minimum abundance of each of the species
shared between sites.
Complementarity amongst sites (CAB) was calculated according to Colwell and Coddington (1994):


Sj Sk  2Vjk
100
CAB
Sj Sk  Vjk
where Sj = number of species at site A, Sk = number of species at site B and
Vjk = number of species shared by both sites.

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We calculated the annual death toll (ADT) by calculating the daily roadkill average
(dra = summary of the number of roadkills each surveyed day, divided by the number of
surveyed days after cleaning the highway) for all animal groups, times the number of days
in a year.
ADT dra  365

Results
Total species richness was 58 vertebrate species (Tables 1, 2, and 3), and this was dominated by mammals (82.87 %), followed by birds (9.64 %) and reptiles (7.8 %). For the
regional diversity estimate we used the Perote Valley fauna inventory by Gonzalez-Romero (2010; Table 1) which reports a total of 162 potential species in the area. Our roadkill
results include 35 % of the total expected vertebrates in the area. By taxonomical group
there were 58.3 % of the expected mammals, 20.4 % of the expected birds and 52.3 % of
the expected reptiles and amphibians.
Based on the comparison of richness and abundance between both environments
(Table 2) we calculated a complementarity value of CAB = 98.49 and a BrayCurtis
similarity value of Ibc = 0.12. Of the total roadkill richness recorded for the highway (58
species), 10 (18.87 %) were observed during cleaning sessions and 43 (81.13 %) on the
other surveys.

Table 1 Number of species with number of individuals by taxonomic group found as roadkill during the
20102011 period on the Amozoc-Cantona-Perote highway (km 90103 ? 860)
Taxonomic
group

Percent
of total
species
found as
roadkill

Potential
number
of species
in the Perote
Valley*

Species
richness

Number of
individuals
found as
roadkill

Mammals

28

827

48

58.33

Birds

19

65

93

20.43

Herpetofauna

11

54

21

52.38

Total

58

946

162

35.80

The third column is the total number of species expected to be found in the Perote Valley for the different
taxonomic groups and the last column gives the percentage of the total expected number of Perote Valley
species found as roadkill
* Gonzalez-Romero (2010)

Table 2 Diversity (richness and abundance) of the vertebrate roadkill on the Amozoc-Cantona-Perote
highway (km 90103 ? 860) in two types of environment
Habitat

Richness

Number of
individuals

Simpsons
index (D)

BrayCurtis
similarity

Complementarity

0.12

98.50

Shrubland

38

475

0.52

Grassland

34

471

1.20

Total

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Table 3 Vertebrate species found road-killed along the Amozoc-Cantona-Perote highway (km
90103 ? 860) with their records, displayed by taxonomic group, and habitat
Species

Common name

Records

Shrubland
(malpas)

Grassland
(llano)

Mammals
Bassariscus astutus

Ringtail

Canis familiaris

Domestic Dog

Canis latrans

Coyote

1
0

Chiroptera sp.

Bat

Cratogeomys fulvescens

Oriental Basin Pocket Gopher

Didelphis virginiana

11

Dipodomys phillipsii

Opposum
Phillips Kangaroo Rat

21

17

Felis catus

Domestic Cat

Lasiurus borealis

Eastern Red Bat

Lepus callotis

White-sided Jackrabbit

Liomys irroratus

Mexican Spiny Pocket Mouse

Microtus mexicanus

Mexican Vole

Mustela frenata

Weasel

Neotoma mexicana

Mexican Woodrat

52

50

Otospermophilus variegatus

Rock Squirrel

Perognathus flavus

Silky Pocket Mouse

Peromyscus difficilis

Southern Rock Deermouse

291

288

Peromyscus maniculatus

Deer Mouse
Pinon Mouse

363

21

342

Peromyscus truei(?)

Procyon lotor
Raton

Raccoon
Mouse

Rattus norvegicus

Common Rat

Reithrodontomys megalotis

Southern Marsh Harvest Mouse

Sigmodon hipsidus

Hispid Cotton Rat

Sylvilagus audubonii

Desert Cottontail

Sylvilagus cunicularius

Mexican Cottontail

Sylvilagus sp.

Rabbit

Xerospermophilus perotensis

Perote Ground Squirrel

Total

33

33

827

401

426

Birds
Passeriform

Bird

34

21

13

Ammodramus savannarum

Grasshopper Sparrow

Carpodacus mexicanus

Mexican Sparrow

Colaptes auratus

Northern Flicker

Cyrtonyx montezumae

Montezuma Quail

Eremophila alpestris

Horned Lark

Falco sparverius

Kestrel

Gallus gallus

Domestic Hen

Geococcyx californianus

Roadrunner

Guiraca caerulea

Blue Grosbeak

Hirundo rustica

Barn Swallow

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Table 3 continued
Species

Common name

Records

Shrubland
(malpas)

Grassland
(llano)

Pipilo fuscus

Canyon Towhee

Salpinctes obsoletus

Rock Wren

Sturnella sp.

Meadowlark

Toxostoma curvirostre

Gray Thrasher

Trochilidae sp.

Hummingbird

Turdus sp.

Thrush

Tyto alba

Barn Owl

Zenaida macroura

Mourning Dove

65

35

30

Total
Reptiles and amphibians
Barisia imbricata

Plated alligator Lizard

Bufonidae sp.

Toad

Conopsis lineata

Lined Tolucan Ground Snake

Crotalus molossus

Black-tailed Rattlesnake

11

11

Crotalus scutulatus

Mohave Rattlesnake

18

15

Crotalus sp.

Rattlesnake

Pituophis deppei

Salvadora bairdi

Mexican Bullsnake
Bairds Patchnose Snake

Sceloporus mucronatus

Eastern Crevice Spiny Lizard

Sceloporus spinosus

Spiny Lizard

Tamnophis cyrtopsis

Black-necked Garter Snake

Total
Total

54

38

16

946

474

472

Of the 58 roadkilled species, 10 are under some degree of legal protection according to
the Official Mexican Standard NOM-059-SEMARNAT-2010 (2010) (DOF 2010;
Th = threatened, Pr = under special protection): Xerospermophilus perotensis (Th),
Dipodomys phillipsii (Th), Crotalus scutulatus (Pr), Crotalus molossus (Pr), Crotalus
intermedius (Th), Pituophis deppei (Th), Cyrtonyx montezumae (Pr), Tamnophis cyrtopsis
(Th), Barisia imbricata (Pr), and Salvadora bairdi (Pr). Of these, only the Perote ground
squirrel (Xerospermophilus perotensis) is considered endangered by the IUCN (2011).
We calculated the annual toll by traffic related death as 11,899 vertebrates using the
daily average obtained during this study (32.6 roadkills day-1).
Species accumulation curves by group (Fig. 2) estimate that along this stretch of
highway 33 mammal species, 24 bird species and 20 reptile and amphibian species; though
none of the curves reached its asymptote. The species accumulation curves using the Jack1
estimator show similar values between habitats, with 41 species for the malpas and 40 for
the llano.
The number of roadkills varied over time for the three groups with higher values mainly
in August and October, though mammals were more active during April. The pattern of
roadkill variation by environment was similar (Fig. 3).

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Discussion
Vertebrate roadkill diversity analysis
The impact of highways on wildlife has been underestimated and even ignored; however,
this could represent a major factor in the conservation of biota, especially that of small
vertebrates. The number of vertebrates we observed as roadkill was unexpectedly high
(946 animals) over a relatively short distance (*15 km). Despite the fact we did not
surveyed the right of way or the adjacent area, we noticed a similar situation to Adams and
Geis (1983), we found that generally grassland species were common in the right of way
habitat than adjacent habitat, whereas many less habitat specific species occurred in both
right of way and adjacent habitats. More small mammals were captured in right of way
than in adjacent habitat.
There was no difference in either richness or abundance between environments but in
species composition both habitats are highly complementary, important when it comes for
conservation planning.
Differences in species composition between environments can likely be attributed to the
distinct landscape conditions, i.e., the geomorphology of each site, given that the malpas is
rough and rocky and the grassland is flat and free of rocks. This is the main reason why, as
observed in the analysis (CAB = 0.98) there is a high degree of complementarity between
the two environments and together increase the biodiversity of the highway segment.
The long history of modifications to the landscape in the study area include fragmentation by agriculture, along with changes induced by herding livestock and reforestation in
some parts, and make it difficult to identify the original conditions of the natural vegetation, particularly in the grassland. The presence of species that are specific to coniferous
forests, such as the Plated Alligator Lizard (Barisia imbricata) and the Mexican Vole
(Microtus mexicanus), suggests that the highway might be acting as a corridor between
different habitats.
Local diversity values would probably have been higher if it had been possible to
identify all of the animals properly, particularly small species. Taxonomic identification of
roadkills was difficult due to the condition of the corpses, an expected result of traffic
intensity and impact with a vehicle, the effects of weather and the activities of scavengers
(Ashley and Robinson 1996; Antworth et al. 2005; Langen et al. 2007; Glista et al. 2007;
Hobday and Minstrell 2008). We, along with other researchers, have been going to the area
since 2005, observation of scavengers moving along the highway either birds (Cathartes
aura) or mammals (Canis latrans), are a rare sight. We have never seen scavengers feeding
on carrion along the highway. Dogs are the most abundant scavengers, but they are usually
moving along herders and we have never seen them eating carrion (Pers. obs.). Because of
their low densities in the study area, scavengers did not likely increase our difficulty in
recording individuals, but it is possible that they, along with necrophagous insects, did
remove some corpses. In this study, weather and traffic were more relevant. Rainy days
during the cooler months of the wet season should promote faster carcass dismemberment
by vehicles passing over them. The location of carcasses on road lanes makes them more
susceptible to being repeatedly damaged by vehicles. Small-sized animals, such as most in
this study have smaller persistence probabilities, because they are crushed and torn apart
faster by continuous impact of car wheels, and are more easily removed (Santos et al.
2011). Also, small species are difficult to spot and some may have been missed. This likely
explains why some of the species that are common in the area and frequently associated
with roadsides, such as the lizards Sceloporus megalepidurus and S. scalaris, were not

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Fig. 2 Species accumulation
curves of species found as
roadkill for the different
vertebrate groups on the
Amozoc-Cantona-Perote
highway (km 90103 ? 860).
The curves were generated based
on the upper limit of observed
species (Sobs Mao Tao) compared
to the nonparametric first order
jackknife estimator (Jack1).
Sampling effort was the total
number of survey days on the
highway searching for vertebrate
roadkill (34). Data are shown by
taxonomic group a reptiles and
amphibians, b birds, and
c mammals

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335

Fig. 3 Time fluctuations in the roadkill values for vertebrates on the Amozoc-Cantona-Perote highway (km
90103 ? 860) for a taxonomic group and b habitat type, both standardized on the Y-axis (log ? 1)

recorded. On the other hand, amphibians do not last long after being hit by cars (Antworth
et al. 2005), and thisalong with the low amphibian density of the areaexplains why we
found only one individual.
In this study, of the total number of roadkilled animals (946), mammals are in the first
place, followed by birds, reptiles and amphibians. As occurs in natural communities,
within each group the abundance of the roadkilled species is dominated by a few taxa
(Taylor and Goldingay 2004; Smith-Patten and Patten 2008). As found in other studies
(Polaco and Guzman 1993, Orlowski and Nowak 2006; Tenes et al. 2007) we found that
rodents were most abundant (95 % of all individuals), followed by lagomorphs and carnivores (\2 % each). The two species for which roadkill was highest were mice belonging
to the genus Peromyscus (P. difficilis and P. maniculatus), one of which inhabits the
malpas and the other, the llano, although both species can be found in both of the habitats
along the highway. In contrast, the Mexican Vole (Microtus mexicanus) and the Hispid Rat
(Sigmodon hispidus), for each of which there was only one roadkill, appear to avoid roads
(Swihart and Slade 1984). It is remarkable that both species were new records for the area,
suggesting that they may be using the highway as corridor (Sigmodon was detected on the
shrubland rather than in the grassland).
Species vulnerability to being killed on the road depends on their behavior (Clevenger
et al. 2003). For instance, in birds it might relate to their flight altitude because the majority

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of species found as roadkill had low flight patterns or spent time on the ground, e.g., the
Montezuma Quail (Cyrtonyx montezumae) or the Roadrunner (Geococcyx californianus).
For all fauna groups, however, regardless of body size, all species are susceptible to being
killed on or over roads (Ferreira Da Cunha et al. 2010).
Although a faunistic inventory of the area adjacent to the highway (Gonzalez-Romero
2010) has been constructed along the several studies (unpubl.) in the area mainly on
squirrels (Xerospermophilus perotensis), but also with, hares (Lepus callotis), coyotes
(Canis latrans) and other taxonomic groups such as rodents (Gonzalez-Christen et al.
2009). The present study makes a contribution by reporting several species that have not
been previously observed; particularly cryptic species including several snakes (Pituophis
deppei, Conopsis lineatus, Tamnophis cyrtopsis) and two rodents (Sigmodon hispidus and
Microtus mexicanus). While this suggests that data from highway surveys can complement
biological inventories and document the richness of associated environments, we must
keep in mind that the proportion of wildlife represented will depend on both road characteristics and the species.
The species accumulation curves did not reach the asymptote because the survey was
not designed as a taxonomic inventory. The record of each species depended on the
probability of species dying as roadkill. Another factor that may explain the lack of
completeness of the inventory is the distance between the road we surveyed and the
vegetation types present in the Perote Valley, such as the case of coniferous forests which
are located a fair distance away from the road.
The variation in the number of roadkills over time was probably associated with
changes in the seasonal activity of the animals (e.g., breeding periods, dispersal, etc.
Polaco-Guzman 1993; Morales-Mavil et al. 1997). This is exemplified by the low numbers
of reptiles observed during the drier, colder months; a time of the year associated with a
natural dip in their activity patterns. Temporal variations could also result from changes in
traffic flow, with increases in roadkill numbers associated with vacation periods when there
is more traffic. However, this needs to be analyzed to determine the relative weight of the
factors in explaining the incidence of roadkills.
Annual roadkill death toll
As suggested by Case (1978), our results are underestimates as a result of our not noticing
some small animals, and because others might have been thrown clear of the highway
when hit, or were injured and died elsewhere. The road ecologist only sees a portion of the
whole phenomenon. Collisions with harmless wildlife are generally not reported because
they do not represent a direct threat to humans or automobiles (Andrews et al. 2006).
This study estimated that there are approximately 11,899 roadkilled animals per year
and this is alarming even though most of the species involved are common rodents with
abundant populations. The magnitude of this result stands out when we consider other
studies (Polaco and Guzman 1993). To appreciate what this roadkill number represents, we
can compare with known populations on the area. Knowing how many individuals per
species are being roadkilled and how their natural populations are, gives us clear idea on
which we should concentrate conservation efforts, and which we can leave as they are.
In this area, studies have focused on the Perote endemic ground squirrels (X. perotensis)
which is considered endangered by the IUCN (2011). In this study, we recorded along the
highway 33 individuals or 3.92 squirrels ha-1 along the 8.4 ha of road surface. CastilloCastillo (2009) recorded densities of 3217 squirrels ha-1 in her study area and 2 years
later Montero-Bagatella (2012), who did her study simultaneously to ours, obtained in

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general terms lower densities of this squirrel, the highest being 24 individuals ha-1
(August in natural grassland) and lowest 2 individuals ha-1 (June). Most of the densities
reported in these studies were lower than 20 individuals ha-1, which, according to
Hayssen (2008) means X. perotensis is a species with reduced and declining populations.
We think this is due to the change of resource availability and human induced changes
(including the highway). Looking at the number of roadkills together with the quick habitat
loss, together with the lack of conservation actions and finally that the highway is planned
to be upgraded up to four lines, the need for specific experimental studies is evident.
Some other species despite the fact they are susceptible to roadkill do not require further
compensation, this is the case of other rodents. Gomez-Vasquez (2005) found rodent (not
counting squirrels or gophers) densities from 0.3 in croplands to 11 individuals ha-1 on the
less disturbed neighboring Pizarro Hill. Even though apparently both densities seem lower
than the 45 roadkilled rodents ha-1 we found on the road surface, their biology and drastic
population fluctuations compensate for the lost in the highway. Similar to the study of
Adams and Geis (1983) where small mammal mortality on the highways was higher for
those species with highest densities in the right of way and did not appear to be detrimental
to populations of these species.
Recommendations for mitigating the impact of highways
The results of Bojorquez-Tapia and Garcas (1998) evaluation of EIAs carried out for
highways in Mexico reveal that these impact assessments contain inadequate descriptions
and biased evaluations. The impact studies often lack any focus on the environmental
conflicts that these projects generate. This is reflected in the limited proposals for impact
mitigation that generally only focus on replanting neighboring zones and making some
effortrarely effectiveto rescue fauna during the building phase. Little is done to
maintain the connectivity of the natural elements of the landscape or diminish roadkill,
despite the fact that vehicle traffic is one of the most serious causes of mortality for many
endangered species of mammals around the world (Harris and Gallagher 1989; Rodrguez
and Delibes 1992; Rodts et al. 1998; Van Gelder 1973; Blaustein and Wake 1990; Fahrig
et al. 1995; Seiler 2001). The mitigation actions mentioned above were undertaken by the
ACP highway for the segment that crosses the protected area of the Parque Nacional de La
Malinche (bordering the states of Puebla and Tlaxcala) where, according to the EIAs, the
bridges that are there provide sufficient crossings to maintain connectivity. The impact of
this highway on wildlife (especially small species) was clearly underestimated from the
time of writing up the highway proposal (Ingenieros Civiles de Mexico 2002) to proposing
mitigation measures. On the rest of the highway and particularly the area we studied (km
90103 ? 860), there are no mitigation measures designed to assist or protect fauna while
the highway is being used. The only action done was to relocated plant species rescued
from the shrubland to the grassland (where they do not belong), and to reforest with plant
pines along the right of way.
We believe as mentioned by McGregor et al. (2008), that measures aimed at reducing
traffic amount or traffic speed or altering the timing of traffic seasonally or daily would not
mitigate effectively the effects of roads on small mammal movement. More effective
measures might be construction of suitable wildlife passages (Clevenger et al. 2001;
McDonald and Clair 2004), reduction of the width of the road and/or the road right-of-way,
and in extreme cases (e.g., mitigation for endangered species), removal of the road itself.
Our results also support the suggestion that conservation of large roadless areas should be a
priority (Crist et al. 2005).

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In this case, there are a series of drainage culverts (of different dimensions and designs)
and overpasses that could be remodeled to be more attractive to wildlife. The creation of
multipurpose bridges for people to cross with their livestock and vehicles, and that also
serve as wildlife crossings could be successful in Mexico because it offers a socially viable
and cost-effective solution. Ideally, these crossings would be designed with surface that
resemble a natural surface and should have a continuous line of vegetation, perhaps herbaceous plants, that connects both sides of the highway. Structures could be refitted and
combined with fences to steer organisms to safe passage (Clevenger and Waltho 2000;
Jackson and Griffin 2000; Dodd et al. 2004; Glista 2006), and this would achieve two
objectives. It would reduce vehicle-animal collisions and would enhance connectivity
between populations (Clevenger et al. 2003). If something like this is not done, the results
could be more damaging to the animal populations in the long term. If the highway we
studied is expanded to four lanes, measures to facilitate wildlife crossing would be crucially important, as would conserving the vegetation of the right of way. The latter serves
as refuge from the cropland and a corridor for the remaining wildlife that could be in
danger if this land use is lost. Finally, as observed in this study, wildlife roadkills could be
reduced if proper drivers education is provided so that drivers understand that animals
could cross their path. This is especially important in areas where larger animals are
vulnerable in terms of conservation.
Future studies and considerations
Future studies must include the systematic year round sampling of the highways, as well as
weather information and daily traffic flow fluctuations. The highway studies should also be
linked to wildlife population studies to provide a better picture of the impact of the road on
wildlife mortality. The use of genetic techniques could throw more light on the true long
term impact of the road, by assessing the maintenance of the wildlife populations in the
long run.
Last but not least, it should be a legal requirement to carry out before and after studies
when roads are built, in order to facilitate the evaluation of their impact on wildlife.
However, the most important action is to design and apply policies and constructive actions
that allow for more responsible and environmentally friendly development. Together with
improved drivers education and raising the awareness of society about this serious
problem, these strategies can reduce the impact of roads on wildlife.

Conclusions
The impact highways have on wildlife through vehicle collisions spans all taxonomic
groups, although a few taxa dominate the roadkills scene. As expected, the most roadkilled
species are usually the most abundant on the neighboring area as well. To completely
understand how profound the roadkill impact is, we should have an idea of how wild
populations are behaving, particularly when it comes to threatened species. The roadkill
phenomena is underestimated in almost every EIA especially on small fauna, and therefore, no specific mitigation measures to reduce roadkills and promote connectivity are
made along highways. In Mexico, drainage culverts and overpasses have not been assessed
and both EIA together with highway developers suppose they are an effective measure for
addressing connectivity but no evidence has been gathered to substantiate this hypothesis.
More road ecology studies are required within Mexican landscapes and cultural context if

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we want to have cost-effective mitigation measures that lead to more environmentally

friendly road development, and also if we wish to have conservation measures that insure
sensitive species permanence in the long term.
Acknowledgments Our sincere thanks to A. Gonzalez-Romero and S. Bagatella for help with the field
work. This study was supported by the Instituto de Ecologa A.C. and CONACYT. We are grateful to M.
Equihua and B. Delfosse for reviewing this document in Spanish and English. The Operadora Concesionaria
Mexiquense gave us permission to work along the Amozoc-Cantona-Perote highway, segment km
90103 ? 860.

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