Influences of long-term changes in land cover on mammal populations: an example from Mexico

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Influences of long-term changes in land cover on
              mammal populations: an example from Mexico
                                                                                            CONSUELO LORENZO, EUGENIA C. SÁNTIZ
                                                          JORGE         B O L A Ñ O S - C I T A L Á N and D A R Í O N A V A R R E T E - G U T I É R R E Z

              Abstract Changes in vegetative cover and land use in the                               growth (Mas et al., ; Nájera et al., ), which increases
              southern part of the Isthmus of Tehuantepec in Oaxaca,                                 demand for space for housing and food production, and
              Mexico, were identified by analysing satellite images from                             poor agricultural management, particularly with respect to
               and . Fluctuations in population density of mam-                              livestock raising and prescribed burning of pastures (uncon-
              mal species during – in response to these changes                              trolled burning of grasses to promote new growth of grazing
              were analysed. During – the types of land use that                             pasture for cattle) for seasonal crops (López-Granados et al.,
              increased in area (per year) were prescribed burning (un-                              ; Peralta-Rivero et al., ). Both of these matters have
              controlled burning caused by humans) by . ha, seasonal                            led to reduction, degradation and fragmentation of habitat
              agricultural plots by . ha, areas without vegetation by                            (Bender et al., ; Rioja et al., ; Lorenzo et al., ),
              . ha, and human settlements by . ha. In the same                                directly affecting the amount and quality of vegetation, as
              period, savannahs decreased by . ha, tropical dry forest                          well as food availability for wild animals.
              by . ha, secondary forests by . ha, and human-                                      In the southern region of the Isthmus of Tehuantepec,
              induced grassland by . ha per year. The loss of these                               habitat consists of scrub and mangrove, with submerged vege-
              habitats resulted in low population densities of mammals,                              tation in coastal lagoons (Superior Lagoon and Inferior
              including the eastern cottontail Sylvilagus floridanus and                             Lagoon; Ortiz et al., ), as well as extensive areas of savan-
              the Tehuantepec jackrabbit Lepus flavigularis, which is cate-                          nah with isolated shrubs and trees (Rzedowski, ; Sántiz
              gorized as Endangered on the IUCN Red List. We recom-                                  et al., ). Large areas of low tropical deciduous and low de-
              mend establishing a communal ecological reserve in the                                 ciduous forest (Torres-Colín, ), as well as savannahs and
              study area to implement appropriate management strategies                              human-induced grasslands (native or exotic species that
              for grassland communities and develop a programme of                                   develop in sites in which the original vegetation has been
              semi-captive breeding to conserve L. flavigularis.                                     eliminated or in abandoned agricultural areas; Aristida,
                                                                                                     Bouteloua, Cathestecum, Cenchrus, Digitaria, Eragrostis,
              Keywords Carnivores, conservation, land management,
                                                                                                     Panicum, Paspalum and Schizachyrium) are found in this
              land use, Lepus flavigularis, Oaxaca, population density,
                                                                                                     region (Rzedowski, ; INEGI, ; Sántiz et al., ).
              Sylvilagus floridanus
                                                                                                         Grasslands of the Isthmus of Tehuantepec have been
                                                                                                     transformed for agriculture and livestock raising, causing
                                                                                                     degradation and reduction of the original vegetation
                                                                                                     (Sántiz et al., ). One mammal species of the region
              Introduction                                                                           that depends on savannahs and human-induced grasslands
                                                                                                     is the endemic Tehuantepec jackrabbit Lepus flavigularis

              I n tropical regions changes in land use have been the main
                cause of biodiversity loss, with the associated loss of
              habitats for wild flora and fauna (Arriaga & Gómez, ;
                                                                                                     (Anderson & Gaunt, ; Lorenzo et al., ), which is ca-
                                                                                                     tegorized as Endangered on the IUCN Red List (Cervantes
                                                                                                     et al., ). This lagomorph shares its habitat with other
              González et al., ; INEGI, ). The principal causes of                           lagomorphs, such as the eastern cottontail Sylvilagus florida-
              changes in vegetative cover and land use on the Isthmus of                             nus, as well as their predators: the coyote Canis latrans and
              Tehuantepec in Oaxaca, Mexico, are human population                                    the grey fox Urocyon cinereoargenteus (Lorenzo et al., ;
                                                                                                     López et al., ).
                                                                                                         Interactions among taxa influence abundance and dens-
              CONSUELO LORENZO (Corresponding author), EUGENIA C. SÁNTIZ and JORGE                   ity (Erlinge et al., ; Daniel et al., ; Bailey, ) and
              BOLAÑOS-CITALÁN Departamento de Conservación de la Biodiversidad, El
              Colegio de La Frontera Sur, Unidad San Cristóbal, Carretera Panamericana y             can cause population fluctuations in response to environ-
              Periférico Sur s/n, Barrio de María Auxiliadora, C.P. 29290, San Cristóbal de          mental changes and the amount and quality of available
              Las Casas, Chiapas, México. E-mail clorenzo@ecosur.mx
                                                                                                     food (Keith & Windberg, ; Keith, ). Therefore,
              DARÍO NAVARRETE-GUTIÉRREZ Laboratorio de Información Geográfica, El Colegio            identifying the impact of changes in vegetative cover and
              de La Frontera Sur, Unidad San Cristóbal, San Cristóbal de Las Casas, Chiapas,
              México                                                                                 land use on distribution and population fluctuations of
              Received  August . Revision requested  February .
                                                                                                     mammals, using basic ecological parameters such as popu-
              Accepted  April . First published online  June .                            lation abundance and density, facilitates decision making

              Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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322         C. Lorenzo et al.

                for the conservation of endemic and threatened mammal                                  turn to subsistence hunting, which has a negative effect on
                species, such as L. flavigularis, as well as adequate manage-                          populations of lagomorphs, mainly L. flavigularis. Grey
                ment of agriculture and other land-based economic                                      foxes and coyotes are also hunted, although not on a regular
                activities.                                                                            basis (J. Antonio, pers. comm.).
                    Densities of – individuals km− have been reported
                for L. flavigularis (Lorenzo et al., ; Vargas, ;
                                                                                                       Methods
                Sántiz, ), and abrupt changes in density have been re-
                corded over  years of monitoring. These changes have been
                                                                                                       Surveys
                partially attributed to management practices such as pre-
                scribed burning and grazing (Lorenzo et al., ).                                    The population density of L. flavigularis, S. floridanus and
                    Given the importance for wildlife management and con-                              the carnivores U. cinereoargenteus and C. latrans was esti-
                servation of identifying long-term fluctuations in the popu-                           mated annually during at least  month in the dry season
                lation density of lagomorphs, the relationship of these                                (November–April) and  month in the rainy season
                fluctuations to those of carnivores, and the responses of                              (May–October) over  years (–). As there were
                lagomorph population densities to changes in vegetative                                few sightings of U. cinereoargenteus and C. latrans, these
                cover and land use (such as grazing and prescribed burn-                               were grouped together as carnivores. We used the line tran-
                ing), the objective of this study was to examine the relation-                         sect method (Buckland et al., ) to monitor mammal spe-
                ships between fluctuations in population densities of                                  cies along an  km transect of variable width between
                mammals (L. flavigularis, S. floridanus, and their natural                             Montecillo Santa Cruz and Huamuchil. This transect was
                predators U. cinereoargenteus and C. latrans) and changes                              established according to accessibility (traversable roads)
                in land use and vegetative cover over a -year period.                                and visibility (flat open areas) (Fig. ). Number of indivi-
                                                                                                       duals was recorded while travelling along the transect in a
                Study area                                                                             pickup truck at – km per hour, at night, using halogen
                                                                                                       lanterns for illumination. Geographical location, altitude,
                Land use and vegetative cover were analysed in four muni-                              habitat type, and perpendicular distance (the shortest dis-
                cipalities in the southern region of the Isthmus of                                    tance from the individual to the transect, in m; Buckland
                Tehuantepec in the Mexican state of Oaxaca:                                            et al., ) were recorded for each individual observed.
                San Francisco del Mar, San Francisco Ixhuatán, San                                     The perpendicular distance was recorded for jackrabbits
                Dionisio del Mar, and Juchitán de Zaragoza. Mammals                                    and cottontail rabbits since , and for carnivores since
                were monitored in Montecillo Santa Cruz in the municipal-                              . For this, an electronic distance measuring device
                ity of San Francisco del Mar, and Huamuchil in the muni-                               (Bushnell Yardage Pro, Bushnell Corporation, Overland
                cipality of San Dionisio del Mar (Fig. ). The study area                              Park, USA) was used. All surveys were conducted during
                comprised . km and the area surveyed for mammals                                   .–., when lagomorphs and carnivores are most ac-
                was . km. Climate is warm sub-humid, with a mean an-                                tive (Carrillo-Reyes et al., ; Rioja et al., ). Each night,
                nual temperature of °C, and mean total annual precipita-                             the same  km transect was travelled in both directions, at
                tion of . mm (García, ).                                                       least  hours apart to avoid duplicating data (in number of
                    Residents of Montecillo Santa Cruz are traditional fish-                           records and kilometers travelled); each mammal observed
                ers, although over the past decade they have begun to                                  was recorded, and density was estimated using these data.
                plant sorghum for sale and use as fodder for livestock,                                   Data were analysed using Distance . (Buckland et al.,
                and maize and watermelon for family consumption                                        ) to estimate density for each mammal species, and
                (Vargas, ; Sántiz et al., ). Cattle are raised extensive-                      we considered each direction travelled along the transect
                ly in all grassland areas, and prescribed burning is carried                           to be independent (i.e. we analysed the data for each direc-
                out to promote regeneration of grasses (Lorenzo et al.,                                tion independently). To evaluate the probability of detecting
                , ; Vargas, ).                                                             animals, the four models used by Distance were tested: half-
                    Within the study area a sand bar connects the Inferior                             normal, uniform, hazard rate, and negative exponential.
                Lagoon with the Gulf of Tehuantepec (Fig. ). The sand                                 Each model varies depending on homogeneity of sightings,
                bar closes and opens cyclically, changing in length as a result                        probability of detecting individuals according to the dis-
                of natural events (e.g. changes in sea level, hurricanes). This                        tance from which they are observed, and number of sight-
                process affects local people’s socio-economic activities, as                           ings (Buckland et al., ). In each case a cosine
                they principally engage in fishing. When the sand bar is                               adjustment was assessed using likelihood ratio tests. The
                closed, movement of fish and nutrients from the Gulf of                                longest % of the perpendicular distances recorded through-
                Tehuantepec to the Inferior Lagoon is inhibited, and fish                              out the analyses were eliminated to avoid bias introduced by
                and shrimp populations decrease, and therefore local people                            atypical outlier distance sightings. The final model was

                                                                                           Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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https://doi.org/10.1017/S0030605317000485
Effects of land cover change in Mexico                    323

                                                                                                                                  FIG. 1 Study area in Montecillo Santa
                                                                                                                                  Cruz, south-eastern Oaxaca, Mexico, in
                                                                                                                                  which vegetative cover and land use were
                                                                                                                                  analysed, with the location of an  km
                                                                                                                                  transect between Montecillo and
                                                                                                                                  Huamuchil along which lagomorphs and
                                                                                                                                  carnivores were monitored during –
                                                                                                                                  .

              chosen based on a combination of the lowest value for
              Akaike’s information criterion (AIC) and the lowest vari-
              ance (Rizo-Aguilar et al., ) for each year of monitoring.
              The population density obtained corresponds to the area
              monitored for mammals (. km; i.e. the length of the tran-
              sect ( km) multiplied by the maximum perpendicular dis-
              tance ( m each side of the transect) at which it is possible
              to identify observed animals). The results obtained for the
              density in the monitoring area (. km) were extrapolated
              to the total study area (. km).
                                                                                                     FIG. 2 Fluctuations in population densities of the Tehuantepec
                                                                                                     jackrabbit Lepus flavigularis, the eastern cottontail Sylvilagus
                                                                                                     floridanus, and carnivores (Canis latrans and Urocyon
              Changes in vegetative cover and land use
                                                                                                     cinereoargenteus) in Montecillo Santa Cruz, Municipality
              Maps of vegetation and land use were created using analysis                            San Francisco del Mar, and Huamuchil, Municipality San
                                                                                                     Dionisio del Mar, Oaxaca, Mexico (Fig. ) during –,
              of a  satellite image, from the Landsat Enhanced
                                                                                                     with years indicated in which the sand bar was closed and in
              Thematic Mapper (ETM+), and a  image from the                                      which there was extensive burning.
              Landsat Operational Land Imager (OLI). Changes in vege-
              tative cover and land use were processed using the inter-
              dependent classification method (FAO, ), stratifying                               correctly or incorrectly from satellite images (Velázquez
              the two scenes with ArcGIS . (ESRI, Redlands, USA).                                 et al., ). For these points, types of vegetation and land
              Using this approach we delimited polygons for each type                                use were described.
              of vegetation and land use, to avoid bias in delimiting                                    Once vegetation and land-use maps were drawn up for
              each thematic class. Estimating changes in surface area                                 and , along with the road layer and the digital ele-
              also involved the road layer, taken from a  base map,                              vation model, rates of change were estimated using the tran-
              using ArcGis ., and the digital elevation model of                                  sition matrices (Bocco et al., ) by superimposing maps
              Instituto Nacional de Estadística y Geografía (for ,                               using the Land Change Modeler module of IDRISI Selva
              with  m resolution).                                                                 v. . (Clark Labs, Worcester, USA). An equation pro-
                  To verify classification of the satellite image and ensure                         posed by the FAO () was used to obtain rates of percent-
              reliability of vegetative cover and land-use maps,  control                         age change in surface area for each type of vegetation and
              points were recorded using a global positioning system in                              land use. This procedure was applied to all types of vegeta-
              sites where the vegetation could potentially be interpreted                            tion and land use: human-induced grassland, savannah,

              Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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https://doi.org/10.1017/S0030605317000485
324         C. Lorenzo et al.

                TABLE 1 Percentage change (δ) in surface area (ha) for each type of vegetation and land use in Montecillo Santa Cruz, Oaxaca, Mexico
                (Fig. ), in  and , with the change in area (ha), the percentage lost or gained, and the change per year (ha).

                Types of vegetative cover and                  Surface area (ha)                                             Change in               % loss              Change per
                land use                                       2001                 2014                 δ (%)               area (ha)               or gain             year (ha)
                Human-induced grassland                         6,806.5              6,804.8             −0.002                 −1.710                −0.03                −0.13
                Savannah                                        8,174.5              6,823.4             −1.4                −1351.2                 −16.5               −103.94
                Seasonal agriculture                            3,087.1              3,842.9              1.8                  755.8                  24.5                 58.14
                Human settlements                                 202.1                255.9              1.9                   53.7                  26.6                  4.13
                Tropical dry forest                             6,474.9              5,961.4             −0.7                 −513.5                  −7.9                −39.50
                Burned area                                       957.2              2,323.5              7.4                 1366.4                 142.8                105.11
                Areas without vegetation                          833.7              1,152.7              7.4                  319.1                  38.3                 24.54
                Secondary forest                                5,721.8              5,533.8              2.6                 −187.9                  −3.3                −14.46
                Bodies of water                                28,895.2             28,531.4             −0.3                 −363.8                  −1.3                −27.98
                Mangrove swamp                                  3,399.9              3,323.0             −0.1                  −76.9                  −2.3                 −5.92

                seasonal agriculture, human settlements, tropical dry forest,                          with an annual loss of .%, and an increase in
                burned areas, areas without vegetation, secondary forest,                              human-induced grassland (. ha) and burned areas
                bodies of water, and mangrove swamps. The results describe                             (. ha; Table ).
                transitions from one type of land cover to another through-                                From  to  seasonal agriculture, human settle-
                out the region (Velázquez et al., ).                                               ments, burned areas and areas without vegetation showed
                                                                                                       annual increases in surface area. Burned areas increased
                                                                                                       by .% annually, with a total increase of ,. ha
                Results                                                                                (.%). During the same time period, areas without vege-
                In Montecillo Santa Cruz the mean estimated density of                                 tation increased by .% annually, with a total increase of
                L. flavigularis during – was . ± SE .                                    . ha (.%; Table ). Burned areas were located mainly
                individuals per km (range .–.; model hazard rate/                              in savannahs (,. ha), human-induced grasslands (
                cosine; AIC = .; CV = .%) and of S. floridanus . ±                         ha), and forests (. ha; Table ). All changes (losses
                SE . individuals per km (range .–.; model hazard                            and increases) in land use and surface area of vegetative
                rate/cosine; AIC = .; CV = .%). The mean estimated                            cover from  to  are shown in Fig. .
                density of the carnivores U. cinereoargenteus and C. latrans
                during – was . ± SE . individuals per km                                Discussion
                (range .–.; model hazard rate/cosine; AIC = .;
                CV = .%). The total sampling effort (for all  years)                              The estimated population density of L. flavigularis (. in-
                was  km travelled for L. flavigularis and S. floridanus,                            dividuals km−) in the study region is similar to estimates
                and  km travelled for the carnivores.                                               from previous studies (.–. individuals km−; Vargas,
                    Fluctuations in mean annual population density for each                            ; Sántiz, ; Lorenzo et al., ). However, mean
                mammal species/group are shown in Fig. . Although sev-                                densities of L. flavigularis are low compared to other lago-
                eral fluctuations occurred during –, as a general                              morph species, such as Lepus californicus (– indivi-
                trend the density of L. flavigularis indicates a general de-                           duals km−; Hayden, ; Davis & Schmidly, ;
                cline; Sylvilagus floridanus has also declined but appears                             Portales-Betancourt et al., ). The estimated population
                to be stable since ; and carnivores tended towards sta-                            density for S. floridanus in this study (. individuals
                bility throughout the study period (Fig. ).                                           km−) is also low compared to other studies of the same spe-
                    Savannah (,. ha in ; ,. ha in ) and                              cies (.–, individuals km−; Chapman et al., ;
                human-induced grassland (,. ha in ; ,. ha                                Silvano et al., ). Alteration and degradation of lago-
                in ) were the dominant cover types in the study area                               morph habitat has probably led to a decrease in available
                (Table ), accounting for . and .% of total surface                              food and space over time, resulting in lower population
                area on average, respectively. Almost all vegetative cover                             densities (Lorenzo et al., ; Sántiz, ).
                types showed losses in surface area from  to , but                                The density of L. flavigularis decreased drastically over
                the most drastic loss occurred in savannah, with a .% de-                            the study period, whereas the density of S. floridanus de-
                crease annually, and a total loss of ,. ha (.%) of the                         creased significantly only during –. Highs and
                original surface area. This was principally a result of fires                          lows in densities of the two species (Fig. ) did not overlap
                (burned area = ,. ha) and an increase in agricultural                             except in – and ; this may be explained by the
                areas ( ha). Tropical dry forest lost . ha (.%),                              differential response of the two species to habitat use and

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Effects of land cover change in Mexico      325

                                                                                                                                                                                                                                                                                                                                     food availability. Lepus flavigularis inhabits savannahs and

                                                                                                                                                                                                                                                                                                                      22.14 (0.08)
                                                                                                                                                                                                                                                                                                                                     human-induced grasslands (Lorenzo et al., ;

                                                                                                                                                                                                           swamp, ha
                                                                                                                                                                                                           Mangrove
                                                                                                                                                                                                                                                                                                                                     Carrillo-Reyes et al., ; Sántiz et al., ) and feeds
                                                                                                                                                                                                                                                                                                                                     mainly on grasses (Lorenzo et al., ), whereas S. florida-

                                                                                                                                                                                             water, ha (%) (%)
                                                                                                                                                                                                                                                                                                                                     nus inhabits human-induced grasslands, tropical forests,
                                                                                                                                                                                                                                                                                                                                     and scrub-brush, feeding on herbaceous and woody plants
                                                                                                                                                                                                                       9.72 (0.1)

                                                                                                                                                                                                                                                                    6.57 (0.1)

                                                                                                                                                                                                                                                                                         70.02 (8.4)
                                                                                                                                                                              Secondary for- Bodies of

                                                                                                                                                                                                                                                                                                                                     (Chapman et al., ; Chapman & Ceballos, ). It is
                                                                                                                                                                                                                                                                                                                                     likely that the two species avoid competing with each other.
                                                                                                                                                                                                                                                                                                                                         The combined population density of the carnivores U. ci-
                                                                                                                                                                                                                                                                                                                                     nereoargenteus and C. latrans was stable over time. These
                                                                                                                                                                                                                       190.53 (2.8)

                                                                                                                                                                                                                                        98.37 (1.2)

                                                                                                                                                                                                                                                                    46.17 (0.7)
                                                                                                                                                                                                                                                                    76.14 (8.0)
                                                                                                                                                                                                                                      384.75 (12.5)

                                                                                                                                                                                                                                                                     2.07 (0.2)
                                                                                                                                                                                                                                                                                                                                     species are adapted to a range of environments and can tra-
                                                                                                                                                                              est, ha (%)

                                                                                                                                                                                                                                                                                                                                     vel long distances in search of food, without excessive en-
                                                                                                                                                                                                                                                                                                                                     ergy expenditure (Villa & Cervantes, ). In our study
                                                                                                                                                                                                                                                                                                                                     area, at least, they do not appear to be greatly affected by
                   TABLE 2 Change in area (ha) from one type of vegetation and land use to another in Montecillo, Oaxaca, Mexico (Fig. ), during –.

                                                                                                                                                                                                                                                                                                                                     changes in land cover.
                                                                                                                                                                                              Areas without
                                                                                                                                                                              Burned area, ha vegetation, ha

                                                                                                                                                                                                                       2.7 (0.04)

                                                                                                                                                                                                                                                                                                        2.97 (0.05)
                                                                                                                                                                                                                                                                                                       381.87 (1.3)
                                                                                                                                                                                                                                                                                                        37.17 (1.1)
                                                                                                                                                                                                                                                                                                                                         In the study area, burned areas, areas without vegetation,
                                                                                                                                                                                                                                                                                                                                     and human settlements increased in surface area from 
                                                                                                                                                                                                                                                                                                                                     to . Undisturbed land has been converted to agriculture,
                                                                                                                                                                                              (%)

                                                                                                                                                                                                                                                                                                                                     with an increase in surface area of . ha during this per-
                                                                                                                                                                                                                                                                                                                                     iod. This land was initially planted with corn; however, as a
                                                                                                                                                                                                                       286.02 (4.2)

                                                                                                                                                                                                                                      1024.11 (12.5)
                                                                                                                                                                                                                                         10.44 (0.3)

                                                                                                                                                                                                                                                                    190.89 (2.9)

                                                                                                                                                                                                                                                                                                       178.74 (3.1)

                                                                                                                                                                                                                                                                                                                                     result of changes in geographical and environmental condi-
                                                                                                                                                                                                                                                                                                                                     tions (sandy soil, drought and intense rainfall), cultivation
                                                                                                                                                                                                                                                                                                                                     of sorghum has increased since  (Vargas, ;
                                                                                                                                                                              (%)

                                                                                                                                                                                                                                                                                                                                     Lorenzo et al., ; Rioja-Paradela et al., ).
                                                                                                                                                                                           forest, ha (%)

                                                                                                                                                                                                                                                                                                                                         Savannahs and forests have increasingly been converted
                                                                                                                                                                                           Tropical dry

                                                                                                                                                                                                                                      8.82 (0.11)
                                                                                                                                                                                                                                      1.08 (0.03)

                                                                                                                                                                                                                                                                                   64.08 (6.69)
                                                                                                                                                                                                                                                                                    0.09 (0.01)

                                                                                                                                                                                                                                                                                                                                     to pastures for livestock raising (Lorenzo et al., ;
                                                                                                                                                                                                                                                                                                                                     Rioja-Paradela et al., ), affecting the coverage and diver-
                                                                                                                                                                                                                                                                                                                                     sity of vegetation (Altesor et al., ). Trampling by cattle
                                                                                                                                                                                                                                                                                                                                     modifies the soil’s physical properties, causing it to lose its
                                                                                                                                                                                                                                                     16.74 (0.54)
                                                                                                                                                                                                                       1.98 (0.03)

                                                                                                                                                                                                                                                                                                       15.93 (0.28)
                                                                                                                                                                                                                                                                                                       19.08 (0.07)
                                                                                                                                                                              Seasonal agri- settlements,

                                                                                                                                                                                                                                                                                                                                     capacity for water retention, thereby facilitating erosion
                                                                                                                                                                                                                                                                                                                                     (Drewry & Paton, ). It has been observed that the per-
                                                                                                                                                                                              Human

                                                                                                                                                                              culture, ha (%) ha (%)

                                                                                                                                                                                                                                                                                                                                     manence of the height and cover of species of bunchgrass
                                                                                                                                                                                                                                                                                                                                     such as Muhlenbergia macroura and Jarava ichu benefits
                                                                                                                                                                                                                                                                                                                                     the presence and abundance of other threatened lagomorph
                                                                                                                                                                                                                       246.24 (3.6)

                                                                                                                                                                                                                                      301.95 (3.7)

                                                                                                                                                                                                                                                                    82.17 (1.3)
                                                                                                                                                                                                                                                                    25.74 (2.7)

                                                                                                                                                                                                                                                                                                       524.43 (9.2)
                                                                                                                                                                                                                                                                                                        3.24 (0.01)

                                                                                                                                                                                                                                                                                                                                     species, such as the volcano rabbit Romerolagus diazi, be-
                                                                                                                                                                                                                                                                                                                                     cause it provides refuge from potential predators and nest-
                                                                                                                                                                                                                                                                                                                                     ing (Hoth et al., ; Rizo-Aguilar et al., ).
                                                                                                                                                                                                                                                                                                                                         To manage grazing areas, residents of Montecillo Santa
                                                                                                                                                                                                                                                                    22.95 (0.4)
                                                                                                                                                                                                                                                                    69.75 (7.3)

                                                                                                                                                                                                                                                                                                       0.63 (0.01)

                                                                                                                                                                                                                                                                                                                                     Cruz carry out prescribed burning, which consists of delib-
                                                                                                                                                                                           Savannah,

                                                                                                                                                                                                                                                                                                                                     erately causing fires to promote regeneration of grasslands
                                                                                                                                                                                           ha (%)

                                                                                                                                                                                                                                                                                                                                     during the dry season. However, these fires frequently be-
                                                                                                                                                                                                                                                                                                                                     come uncontrolled, affecting large areas of land (Lorenzo
                                                                                                                                                                                                                                                                                                                                     et al., , ; Vargas, ).
                                                                                                                                                                                           grassland, ha (%)
                                                                                                                                                                                           Human-induced

                                                                                                                                                                                                                                                                                                                                         Changes in land use to accommodate agriculture, includ-
                                                                                                                                                               Modified to:

                                                                                                                                                                                                                       Seasonal agriculture 14.94 (0.5)
                                                                                                                                                                                                                                             11.25 (0.1)

                                                                                                                                                                                                                       Tropical dry forest 238.77 (3.7)
                                                                                                                                                                                                                                             88.11 (9.2)
                                                                                                                                                                                                                                             33.48 (4.0)

                                                                                                                                                                                                                                            263.25 (4.6)
                                                                                                                                                                                                                                             23.76 (0.1)
                                                                                                                                                                                                                                             61.92 (1.8)

                                                                                                                                                                                                                                                                                                                                     ing prescribed burning, have been the main cause of de-
                                                                                                                                                                                                                                                                                                                                     struction of large areas of savannah (.% of the original
                                                                                                                                                                                                                                                                                                                                     surface area), which is the preferred habitat of L. flavigularis.
                                                                                                                                                                                                                                                                                                                                     This is a significant threat to this Endangered species, as la-
                                                                                                                                                                                                                       Human settlements

                                                                                                                                                                                                                                                                                                                                     gomorphs are unable to quickly recolonize intensively
                                                                                                                                                                                                                       Mangrove swamp
                                                                                                                                                                                                                       Secondary forest
                                                                                                                                                                                                                       Human-induced

                                                                                                                                                                                                                                                                                                                                     burned areas (Meslow & Keith, ; Driessen, ;
                                                                                                                                                                                                                       Bodies of water
                                                                                                                                                                                                                       Areas without

                                                                                                                                                                                                                                                                                                                                     Salvatori et al., ). The time it takes for lagomorphs to
                                                                                                                                                                                                                         vegetation
                                                                                                                                                                                                                       Burned area
                                                                                                                                                                                                                         grassland
                                                                                                                                                                                                                       Savannah

                                                                                                                                                                                                                                                                                                                                     recolonize these areas depends on how long it takes for habi-
                                                                                                                                                                                                                                                                                                                                     tat to recover its capacity to provide them with sufficient
                                                                                                                                                                                                                                                                                                                                     food and protection (Parker, ), and burning results in

              Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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https://doi.org/10.1017/S0030605317000485
326         C. Lorenzo et al.

                                                                                                                                             FIG. 3 Vegetative cover and land
                                                                                                                                             use in the municipalities of San
                                                                                                                                             Dionisio del Mar, San Francisco
                                                                                                                                             del Mar, San Francisco Ixhuatán,
                                                                                                                                             and Juchitán de Zaragoza, Oaxaca,
                                                                                                                                             Mexico, in (a)  and (b) .

                temporary habitat fragmentation, which could affect popu-                              habitat. Such a programme could be certified as a Wildlife
                lation densities of several species (Fahrig & Paloheimo, ;                         Use and Management Unit by the Department of the
                Diffendorfer et al., ; Klok & De Roos, ). Our find-                            Environment and Natural Resources, to obtain legal use per-
                ings support this observation, as the population density of                            mits for a threatened mammal species, and economic sup-
                L. flavigularis decreased in , when burned areas in-                               port for programme development. () As part of the
                creased (Fig. ). However, fires have had a positive effect                            Wildlife Use and Management Unit a Communal
                on the abundance of R. diazi, as they maintain and promote                             Ecological Reserve could be established, to be administrated
                new growth of the dominant zacaton bunchgrass, the main                                by community authorities, with universities playing a tech-
                food source for this rabbit (Hunter & Cresswell, ).                                nical advisory role. Local protected areas have been shown
                Additionally, the closing of the sand bar affects lagomorph                            to promote appropriation of territory by communities, and
                population density. The sand bar closed in  and ,                              the patrimony of natural resources, and their management
                and the following years the mean population density of                                 could be a viable method of conserving habitat and the
                L. flavigularis decreased (from . individuals km− in                               Tehuantepec jackrabbit, as well as improving the quality
                 to . individuals km− in , and from . indivi-                           of life of local residents. This could ensure the persistence
                duals km− in  to . individuals km− in ). This                              of the Tehuantepec jackrabbit, with economic support
                process appears to be affecting survival of the species, al-                           from the government.
                though the sand bar remained open in  and  and
                the population recovered slightly.
                                                                                                       Acknowledgements
                Conservation and management strategies
                                                                                                       We thank Miguel Aquino, Roberto Gutiérrez, Leyberto
                We offer several recommendations to conserve L. flavigularis                           Gutiérrez, Juan Antonio, Julieta Vargas, Felipe Barragán,
                and establish appropriate management strategies for grass-                             and the people of the municipality of San Francisco del
                land and savannah communities. () Grasses should be con-                              Mar for their valuable help during fieldwork, Ann
                trolled annually by fire, taking care that fire does not spread                        Greenberg for editorial support, and the families of
                beyond targeted areas. () Livestock-raising practices should                          Antonio Gutiérrez and Gutiérrez Vázquez for housing our
                be improved to avoid a negative impact on grasses; for ex-                             crew. We thank El Colegio de la Frontera Sur, the Chicago
                ample, by periodically rotating cattle among pastures to                               Zoological Society, the Lincoln Park Zoo Neotropic Fund,
                avoid overgrazing, thereby facilitating recovery of grasses                            and Mexico’s Consejo Nacional de Ciencia y Tecnología,
                to increase their vegetative cover and fodder productivity.                            and Secretaría de Medio Ambiente y Recursos Naturales
                () A programme of reproduction in semi-captivity in situ                              for funding this research. Recommendations and comments
                in protected enclosures should be developed. If successful,                            from Martin Fisher and an anonymous reviewer improved
                jackrabbits could be released and reintroduced into the                                this article.

                                                                                           Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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https://doi.org/10.1017/S0030605317000485
Effects of land cover change in Mexico                           327

              Author contributions                                                                                  D AV I S , B. & S C H M I D LY , D.J. () Black-tailed jackrabbit. In The
                                                                                                                       Mammals of Texas – Online Edition. Texas Tech University. Http://
              CL has provided funding for this study since . She has                                               www.nsrl.ttu.edu/tmot/Default.htm [accessed  April ].
              monitored mammals in the field since , and managed                                                D I F F E N D O R F E R , J.E., G A I N E S , M.S. & H O LT , R.D. () Habitat
                                                                                                                       fragmentation and movements of three small mammals (Sigmodon,
              the mammal database. She supported the analysis of popu-
                                                                                                                       Microtus and Peromyscus). Ecology, , –.
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              monitoring mammals in the field since . She managed                                                  natural amelioration on soil physical properties and pasture under
              the mammal database, reviewed satellite images and ana-                                                  dairy farming in Southland, New Zealand. New Zealand Journal of
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                                                                                                                    D R I E S S E N , M. () Effects of fire on the broad-toothed mouse,
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                P O R T A L E S -B E T A N C O U R T , G., H E R N Á N D E Z , L., L A U N D R É , J. &                   Instituto de Biología, Universidad Nacional Autónoma de Mexico
                    C E R VA N T E S , F.A. () Reproducción y densidad de la liebre                                   and Grupo Editorial Iberoamérica, Mexico.
                    cola-negra (Lepus californicus) en relación a factores ambientales, en
                    la Reserva de la Biosfera Mapimí, Desierto Chihuahuense. Therya, ,
                    –.                                                                                          Biographical sketches
                R I O J A , T., L O R E N Z O , C., N A R A N J O , E., S C OT T , L. & C A R R I L LO -R E Y E S ,
                    A. () Breeding and parental care in the endangered Tehuantepec                                CO N S U E L O LO R E N Z O is interested in the molecular systematics,
                    jackrabbit (Lepus flavigularis). Western North American Naturalist,                               taxonomy and conservation of lagomorphs and tropical rodents
                    , –.                                                                                        and has been monitoring populations and habitat of the
                R I O J A -P A R A D E L A , T., C A R R I L LO -R E Y E S , A. & L O R E N Z O , C. ()           Tehuantepec jackrabbit Lepus flavigularis for  years. EU G E N I A
                    Análisis de población viable para determinar el riesgo de extinción                               SÁ N T I Z studies the ecology and effects of global climate change on
                    de la liebre de Tehuantepec (Lepus flavigularis) en Santa María del                               the distribution of threatened mammals, and the conservation and
                    Mar, Oaxaca. Therya, , –.                                                                  management of the Tehuantepec jackrabbit and its habitat. JO R G E
                R I Z O -A G U I L A R , A., D E L F Í N -A LO N S O , C., G O N Z Á L E Z -R O M E R O , A. &        BO L A Ñ O S -CI T A L Á N ’ s work is focused on the conservation and
                    G U E R R E R O , J.A. () Distribution and density of the zacatuche                           management of tropical mammals, especially bats and rodents.
                    rabbit (Romerolagus diazi) at the Protected Natural Area “Corredor                                DA R Í O NA V A R R E T E -GU T I É R R E Z studies changes in distribution of
                    Biológico Chichinautizin”. Therya, , –.                                                    mammals, as well as consequences of land-use changes.

                                                                                                           Oryx, 2019, 53(2), 321–328 © 2017 Fauna & Flora International doi:10.1017/S0030605317000485
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https://doi.org/10.1017/S0030605317000485
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