Grazing Systems and Biodiversity in Latin American Areas



Grazing Systems and Biodiversity in Latin American Areas
Grazing Systems
and Biodiversity in
Latin American Areas:
Colombia, Chile and
Coordinated by
Sergio Guevara Sada and
Javier Laborde
Cover photography credits:
Gerardo Sánchez Vigil
Mariano Guevara Moreno-Casasola
Adi E. Lazos R.
To Lucina Hernández García (1960-2013),
enthusiastic woman and researcher on
environmental history of livestock in Mexico.
PASTOS, 42 (2)
PASTOS 2012. ISSN: 0210-1270
Coordinated by
Sergio Guevara Sada and
Javier Laborde
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
PASTOS, 42 (2), 121 - 122
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S. GUEVARA S.................................................................................................... 125
Acknowledgements.............................................................................................. 131
List of contributors............................................................................................... 133
Flora of the Mediterranean Basin in the Chilean espinales: Evidence of
MIGUEL.............................................................................................................. 137
Historical, biogeographical and ecological factors explain the success of some
native dung beetles after the introduction of cattle in Mexico.
M. E. FAVILA...................................................................................................... 161
From tropical wetlands to pastures on the coast of the Gulf of Mexico.
MEDINA.............................................................................................................. 185
The mesoamerican rain forest environmental history. Livestock and landscape
biodiversity at Los Tuxtlas, Mexico.
S. GUEVARA S. AND J. LABORDE.................................................................. 219
Livestock farming in the Saquencipá Valley, New Kingdom of Granada,
Colombia, in the 16th and 17th centuries.
K. G. MORA PACHECO..................................................................................... 251
The impact of raising cattle in the Totonacapan Region of Mexico: Historical
development and approaches for sustainable cattle ranching.
B. ORTIZ ESPEJEL AND R. JIMÉNEZ MARCE.............................................. 273
PASTOS, 42 (2), 121 - 122
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S. GUEVARA S. .................................................................................................. 301
Gaspar González y González, socio de honor y fundador de la SEEP.
JESÚS TREVIÑO................................................................................................ 307
Instrucciones para los autores de la Revista Pastos.............................................. 317
Introduction, Acknowledgments
and Contributors
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Raising livestock is the most extensive productive system in the tropics and
subtropics. The domestication of animals arose almost at the same time as the
domestication of plants in different parts of the world. Both types of production created
two large guilds, that of the crop farmers and that of the ranchers, who throughout
history have battled over land use. This confrontation has produced different results
over time and in different parts of the world, producing the environmental and cultural
characteristics that are particular to each region.
In the Americas, we see the most recent episode in the crops vs. livestock battle.
Throughout these enormous and diverse continents of mainly crop-oriented heritage,
livestock burst onto the scene in the 16th century when the Europeans introduced
domesticated animals large and small, changing a landscape that had been largely
defined by the crop raising vernacular, and becoming the most powerful tool of European
Owing to its extent and profound effect on biological and cultural diversity, the
ecology of raising livestock is essential to understanding the current landscape if we are
to aspire to balanced and sustainable management of the soil, biodiversity and the natural
resources of the Americas. We know little about the interaction between livestock —
cattle, donkeys, horses, pigs, goats, fowl and bees— and local flora and fauna, or of the
effect that environmental conditions have on the animals. The scarcity and fragmented
nature of our knowledge mythifies the ecological impact of livestock and limits our
possibilities and alternatives for their rational management.
To scientists, raising livestock causes a monumental disturbance of ecosystems and
landscapes that entails the disappearance of native species, facilitates the invasion of
exotic species and causes irreversible changes in the physical structure and fertility of
the soil.
From this perspective, the scientists reaction is extreme, they overlook the
biodiversity associated with the cattle pastures, ignore their role in the structure and
functioning of the landscape and the possibility of understanding certain mechanisms
and ecological processes that facilitate the movement of the flora and fauna that maintain
the diversity of natural systems
When speaking about livestock, we are referring to the diverse species of introduced
herbivores: cattle, pigs, goats, sheep, donkeys, chickens and bees. Each has its own
foraging habits and its impact depends on the ecogeographic region where it occurs,
though it can generally be stated that cattle ranching is the most widespread and common
livestock activity in the Americas and the Caribbean.
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The ecology and behavior of cattle in Europe, Asia and Africa have been under study
for some time now, and the results are useful for understanding cattle ranching in the
Americas. However, the huge diversity of American ecosystems in terms of their flora
and fauna, and the short time that has elapsed since the introduction of cattle, compared
to the long history of domestication on Europe, Asia and Africa, make for quite a
different story.
The European colonists were cattle ranchers, as were their ancestors, who practiced
mixed agriculture and herding from 4500 years before the discovery of America. They
brought cattle to the Americas for the first time in the Antilles in 1512, to Mexico in
1520, to the Andes region in 1530 and to Florida in 1565. By the end of the 16th century
cattle had spread as far west as New Mexico and by 1769, to upper California.
The Iberian cattle (Bos taurus) adapted to the American environment quickly, both
on the arid and semiarid high plains, and in the humid lowlands, as evidenced by the
fact that the herds doubled in size at a much higher rate than they did in Europe. From
the 16th to the 19th centuries, Bos taurus —a fast, lean and average-sized beast— rapidly
occupied each ecosystem, in contrast to the more robust British and French cattle, which
progressed much more slowly. From the time of their introduction in the eastern United
States, they only reached the center of the nation in the 19th century.
The capacity of cattle to transform cellulose into meat, milk, fiber and leather gave
them enormous efficiency and an aptitude for changing the environment, even on a
continental level. On the arrival of the Spanish, the Mesoamerican landscape was molded
by extensive and intensive crop growing activities. Permanent and shifting cultivation
occupied large tracts of flat land, hillsides and ravines. Because of colonization,
the indigenous population decreased and relocated, the best lands were abandoned,
monocultures proliferated and herds of cattle spread, changing the landscapes, leaving
only a few remnants of the natural ecosystems.
Livestock became the main agent of transformation of nature in the Americas and
some authors believe it was the determining factor in European colonization.
When livestock was introduced, some animals were left to roam free in the forests
and rainforests from the last third of the 16th century, and continued to do so until the
end of the 19th century. It was left to move freely on the islands and the mainland, and
in particular pigs and cows adapted to the savannas, scrub, and the sparse and dense
woodlands of warm, moist and dry temperate climates.
It is surprising that, over this period of 500 years, the domesticated European animals
that roamed freely caused no severe damage to the original structure of the vegetation,
nor did they cause any considerable changes in the species composition of ecosystems
and natural formations. It seems that the number of head of cattle and their behavior
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adapted to the prevailing environmental conditions and to the carrying capacity of each
of the natural systems they occupied.
This type of feral cattle still exists in parts of Mexico and the Caribbean, where for
various reasons it continues to inhabit natural systems that are reasonably well preserved.
One explanation of why these ecosystems remained in such good ecological
conditions is that these large herbivores did not have to compete with or displace other
species in the natural American ecosystems. Large herbivores disappeared from the
continent en masse, with the exception of bison, moose, caribou and some deer in North
America, and llamas and vicuñas in South America during the Pleistocene, leaving
a void. Livestock filled this gap in natural grasslands, scrubland and dense forests in
both dry and humid regions, fulfilling the task of dispersing fruit and seeds, preying on
seedlings, reducing plant biomass and recycling soil nutrients. One might suppose that
they contributed to maintaining local biodiversity, favoring those species adapted to the
presence of herbivores for their dissemination and establishment.
The disappearance of large herbivores has been documented for arid and semiarid
regions of the Americas. There, the presence of cattle, horses and donkeys has taken
over the function of the extinct herbivores, rescuing tree species that might otherwise
have disappeared had large livestock not arrived on the scene.
Calculations of the number of native herbivores that were present before the
Pleistocene indicate there was an average of 21 animals per km2, with a variation of 15
to 50 animals per km2, each weighing approximately 450 kg.
This would mean there was an average of five hectares per animal, with a range of
two to seven hectares, suggesting a large carrying capacity, which would explain why
the presence of livestock did not drastically change natural habitats in the Americas.
Unfortunately, we do not have similar calculations for the humid tropics, though
everything seems to indicate that the carrying capacity of the forests was high. Recently,
the rapid decline of the medium-sized to large wildlife species most affected and the
decrease in their populations in the tropical rainforests of southern Veracruz over the
past 25 years has been documented. This defaunation included wild herbivore species,
the disappearance of which could change the physiognomy and species composition of
the rainforest.
From our point of view, the greatest decrease in herbivorous fauna occurred in the
late 19th and early 20th centuries, about 100 years ago, when the feral cattle of Bos taurus,
which had roamed freely in the area was removed and replaced by Bos indicus feedlot
cattle. The disappearance of Bos taurus doubtless had a negative effect on the rainforest
plant populations, which once again were left “widowed”, without their dispersers or
primary consumers and carnivore populations, among others.
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This change in the species of cattle was the result of a trend toward intensive
production in enclosures and planted pastures. Spanish breeds of B. taurus were removed
from natural systems and new breeds of B. indicus introduced. This change induced a
massive opening of pastures at the expense of the natural vegetation, and from that time
onwards, cattle ranching has meant deforestation. The concentration of high densities
of livestock modified the soil and pasture management facilitated the arrival and
establishment of grass species associated with livestock from Africa and Asia. Habitats
were simplified and an extensive and homogeneous system emerged, where vegetation
was cleared to make way for fields and pastures, areas with physical conditions that
led to the proliferation of both exotic and other secondary and weed species, which
eventually colonized these open areas.
In both the dry and humid tropics, clearing for fields has reduced the forest to very
small fragments. Huge areas of the tropics are dominated by livestock landscapes
within which there are fragments of the original forest that vary in size immersed in
the pastures. The forest fragments that remain are on hilltops, very steep slopes, and on
rocky terrain or where flooding occurs. Within the pastures a few trees from the original
forest have been spared, some form strips along rivers to protect the channel, and some
are used as living posts to tend barbed wire, or occur as solitary trees to provide shade
for livestock.
This remnant of the forest is worked into livestock management in an incipient
way. Over the past four decades, the expansion of pastures has been impressive. In
ecological terms deforestation and forest fragmentation is very recent and has happened
very quickly. Given the current degree of fragmentation, we do not know if the sections
of forests that remain on the ranching landscapes can be conserved, or for how long.
Nor do we know for certain which and how many of the forest species, or what kind of
ecological processes, can continue to occur in these landscapes.
Livestock management in fragmented landscapes must not only contemplate grass
and cattle, but also the scattered cover provided by the native trees that are present as
isolated trees, as well as the gallery vegetation and the protection of forest edges to
facilitate the movement of the fauna, seed dispersal and pollen exchange among plants
in dispersed populations.
This can lead to the maintenance of the structure and dynamics of this type of
landscape, protecting a number of the species of the original forest in areas dedicated
to raising cattle, which we could describe as a tropical dehesa, an agroforestry system
that originated in the Mediterranean region where the timber, grazing, livestock and
cultivation activities interact beneficially in economic and ecological terms.
The importance of implementing this optimal landscape design and its new
management rules becomes even more evident when we acknowledge that the entire
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area of dry and rain forest in Mexico, Central and South America and the Caribbean
—which there is still in time to conserve in its current state— is now surrounded by
fragmented livestock landscapes. In large part, the future success of the conservation
of Neotropical forests depends on what we can achieve in the landscapes that surround
these remnants, especially with regards to the maintenance and proliferation of the trees
in pastures.
In this context, cattle as herbivores that are potential fruit and seed dispersers could
contribute to maintaining those fragments and the landscape in general, decreasing the
isolation of species and their populations. Naturally, it would be necessary to reflect on
the most appropriate breed of cattle for this purpose.
Knowledge of the ecology and environmental history of livestock will allow us
to understand the current American landscape and will open new perspectives for the
management of biodiversity in livestock areas. This issue brings together a set original
research articles that represent a substantial contribution to our understanding of the
adaptation of livestock to American landscapes and ecosystems, and of the impact its
management and development has had on biodiversity and on the landscape.
The contributions are organized into three chapters. In the chapter on Biodiversity,
Irene Martín-Forés et al. analyze the Mediterranean region in Chile where more than
25% of the flora is not native. This is especially important in the espinales vegetation,
agroforestry-pastoral systems that are very similar in their functioning to the Spanish
dehesas and are of great ecological and socioeconomic interest. The native/nonnative
characteristic of central Chile is compared with that of areas on the Iberian Peninsula,
highlighting possible mechanisms (filters) that may have been acting on floristic
colonization from the Mediterranean basin toward the Chilean Mediterranean zone.
Evidence is provided by Mario Favila for the success of the introduction of livestock.
The dung beetles of Mexico —and those of the Americas in general— were able to
exploit livestock dung and incorporate it into the nutrient cycles of the tropical and
temperate soils. This ability to make use of an exotic resource is explained in the context
of the evolutionary, biogeographic and ecological history of these beetles.
In the Landscape chapter, Patricia Moreno-Casasola et al. explain how, from the
beginning, wetlands were used for cattle grazing, and they describe the transformations
occurring in grazed wetlands that convert them into flooded pastures. The degree of
impact depends on the number of head of cattle, the time they are in the wetland, and
modifications to hydroperiod and vegetation. The changes in the level of flooding,
soil characteristics (organic matter, water retention, bulk density, pH, micro- and
macronutrients) and floristic composition, are described, along with how all this affects
the environmental services provided by wetlands.
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Sergio Guevara S. and Javier Laborde D. demonstrate that the Mesoamerican
rainforest is home to high biological diversity, in spite of its being intensely fragmented
and the isolation of the remnants of the rainforest. There, biodiversity is rich because
of the good connectivity of the landscape, itself the result of natural events, traditional
agricultural practices and, more recently, by cattle raising activities. This historical
overview allows for landscape management to conserve the biodiversity and develop
sustainable production systems.
In the third chapter on Environmental History, Katherinne Giselle Mora Pacheco
challenges the traditionally accepted explanation that environmental problems in the
Saquencipá Valley of Colombia originated during the colonial period from a combination
of wheat monoculture, deforestation and the introduction and expansion of livestock.
She indicates that low rainfall, the influence of dry winds and the presence of clay
soils were factors that from pre-Hispanic times made most of the inhabitants prefer to
live on the fertile riverbanks. Prior to the Conquest, slash and burn activities in the dry
forests, the demand for firewood and the occupation of land that was less fertile or on
slopes led to the loss of vegetation in specific areas, and this loss was notable, even
before the arrival of the Spanish in the region.
Benjamín Ortiz Espejel and Rogelio Jiménez Marce describe the development
of cattle ranching in the Totonacapan, an indigenous region on the Gulf of Mexico,
examining the pros and cons of three models: indigenous, peasant and agroindustrial.
From their analyses, they extract a proposal to build a model for the sustainable
development of raising livestock in this type of region.
In this volume, a notable group of researchers propose a novel approach to the
relationship between raising livestock and biodiversity in the Americas. This is a
landscape comprised of plant and animal species —both native and those brought
from the Mediterranean— where biodiversity has been managed in the American
and European ways, and supported by the biological precedent resulting from the
disappearance of the great abundance and species richness of large herbivores. This
compilation opens a new vista for the investigation of the introduction of livestock to the
Sergio Guevara Sada
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We wish to first thank Juan Piñeiro Andión because he, as the Director of PASTOS,
the journal of the Spanish Society for the Study of Pastures (Sociedad Española para
el Estudio de los Pastos), enthusiastically embraced the project of this special issue
for the journal and waited, with infinite patience and great friendship, while it was
being prepared. We are grateful to Bianca Delfosse, a member of our team, who did
an impeccable job translating and revising the style of the manuscripts, working hard
and applying her professional abilities to overcome linguistic challenges that seemed
insurmountable. Allison M. Jermain always efficient and capable also translated and
revised the manuscripts. Graciela Sánchez-Ríos, a valued member of our research
team, compiled the contributions in an organized manner, with diligence and care,
and formatted them, playing a key role in the production of the manuscript. Kerenha
Hernández, our smiling and ever enthusiastic collaborator and participant in the
project, organized the manuscripts and controlled communication with the authors with
great tenacity, invariably providing superb design ideas. Our thanks also to Gerardo
Sánchez Vigil, a most passionately involved colleague and accomplice, and Mariano
Guevara Moreno-Casasola enthusiastic and inspired, both who generously provided the
magnificent photographs that illustrate the cover of this issue. The Landscape Ecology
project (Ecología del Paisaje) of the Red de Ecología Funcional of the Instituto de
Ecología, A.C., provided the infrastructure and funding for the preparation of the
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Universidad Complutense de Madrid, Departamento de Ecología, Facultad de Biología,
España, [email protected]
Universidad Complutense de Madrid, Departamento de Ecología, Facultad de Biología,
España, [email protected]
Universidad Autónoma de Madrid, Departamento de Ecología, Facultad de Ciencias,
España, [email protected]
Universidad Complutense de Madrid, Departamento de Ecología, Facultad de Biología,
España, [email protected]
Facultad de Ciencias Agrarias, Universidad de Talca, Chile, [email protected]
Instituto de Ecología, A.C.
Red de Ecoetología
Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, México,
[email protected]
Instituto de Ecología, A.C.
Red de Ecología Funcional
Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, México,
[email protected]
Universidad Iberomericana, Puebla, Departamento de Ciencias Sociales. Boulevard del
Niño Poblano 2901. U. Territorial Atlixcáyotl.CP. 72197 Puebla, Puebla, México,
[email protected]
Instituto de Ecología, A.C.
Red de Ecología Funcional
Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, México,
[email protected]
PASTOS, 42 (2), 133-134
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List of Contributors
Universidad Nacional Autónoma de México, Instituto de Ciencias del Mar y Limnología
Estación El Carmen, Cd. del Carmen, Campeche, México, [email protected]
Universidad Complutense de Madrid, Departamento de Ecología, Facultad de Biología,
España, [email protected]
Universidad Nacional de Colombia, Línea de Historia Ambiental IDEA y Departamento
de Historia, Bogotá, Colombia, [email protected]
Instituto de Ecología, A.C.
Red de Ecología Funcional
Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, México,
[email protected]
Universidad Iberoamericana, Puebla. Departamento de Ciencias Sociales. Boulevard
del Niño Poblano 2901. U. Territorial Atlixcáyotl.CP. 72197 Puebla, Puebla, México,
[email protected]
Instituto de Investigaciones Agropecuarias INIA, Centro Regional de Investigación La
Platina, Santa Cruz, Chile, [email protected]
Instituto de Ecología, A.C.
Red de Ecología Funcional
Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, México,
[email protected]
Universidad Complutense de Madrid, Departamento de Ecología, Facultad de Biología,
España, [email protected]
Chapter 1. Biodiversity
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Departamento de Ecología. Facultad de Biología. Universidad Complutense de Madrid. Madrid (España). 2Departamento
de Ecología. Facultad de Ciencias. Universidad Autónoma de Madrid. Madrid (España). 3Instituto de Investigaciones
Agropecuarias INIA-La Cruz. La Cruz (Chile). 4Facultad de Ciencias Agrarias. Universidad de Talca. Talca (Chile).
*Author for correspondence: M.A. Casado ([email protected]).
In Chile’s Mediterranean region, over 18% of plant species are alien. This is
particularly noteworthy in some agrosilvopastoral systems such as the espinales,
which are functionally very similar to the Spanish dehesas and are of great ecological
and socioeconomic interest. In the present paper we analyse Chile’s non-native flora,
considering three scales of analysis: national, regional (the central region, presenting a
Mediterranean climate) and at community level (the espinales within the central region).
We compare this flora with that recorded in areas of the Iberian Peninsula with similar
lithological and geomorphological characteristics, and land use. We discuss possible
mechanisms that might have been operating in the floristic colonisation from the
Mediterranean Basin to Chile’s Mediterranean region.
Key words: Alien plants, biogeography, Chile, life cycle, Spain.
Historically, the transit of goods, domestic animals and people has favoured the flow
of wild organisms around the planet (Lodge et al., 2006), a fact that is often associated
with the introduction of cultural systems, which have contributed to generating
new environmental and socioeconomic scenarios (Le Houérou, 1981; Hobbs, 1998;
Grenon and Batisse, 1989). The current globalisation process is increasing landscape
changes and ecosystem disruptions by human disturbance and therefore facilitating
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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Martín-Forés et al.
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Plant species colonization in Chilean espinales
the transit of organisms (Paskoff and Manríquez, 1999; Rouget et al., 2003; Dukes
and Monney, 2004; Schwartz et al., 2006). These ‘assisted dispersals’ enable species
to cross biogeographical boundaries that have previously limited their distributions.
Species that have been transported from one region to another are defined as alien or
exotic to that newly occupied region (Richardson et al., 2000). Most of these species
fail to establish self-perpetuating populations, but some of them do succeed and become
naturalized (Sax and Brown, 2000). Regardless of the factors enabling establishment,
the main consequence of this naturalisation is that alien species significantly contribute
to the global floristic (taxonomic and phylogenetic) homogenization of regional floras
(Winter et al., 2009). Despite the fact that introduction of species increases diversity
at short temporal and small spatial scales, in the medium and long term, interactions
with native species can lead to extinctions (Pyšek and Richardson, 2006). The net effect
will depend upon the spatial and temporal scales considered and on the balance between
naturalisations and extinctions (Sax et al., 2002). Nonetheless, the resulting ecological
consequence is the coexistence of native species with exotic ones, quite often in the
same community. The origin and composition of these novel communities are of great
interest to understand their functioning and possible management.
Transcontinental naturalisation in Chile’s flora
Mediterranean-type ecosystems around the world offer a great chance to compare
and understand the mechanisms determining the success of species introduced into a
given region (Kruger et al., 1989; Groves and Di Castri, 1991). The different regions
presenting a Mediterranean climate have had different environmental histories
associated with the density of human populations as well as the time and intensity of the
changes that people have caused in the territory. In the Mediterranean Basin, anthropic
modification of the landscape is millenary; however, rates of species extinction and
naturalisation are low in comparison with other Mediterranean regions (Greuter, 1994).
This fact is explained as a process of co-evolution of plants with people (Di Castri,
1981). Conversely, other Mediterranean areas have undergone a rapid change following
successive cultural colonisations, some of these relatively recent, a fact that accounts for
the current ecological conditions threatening the biodiversity of these areas (Underwood
et al., 2009).
As with other Mediterranean regions, Chile is recognised as a biodiversity hotspot
(Ormazabal, 1993), with high levels of regional and national endemism, possibly related
with its biogeographic isolation (Myers et al., 2000). Its flora comprises 5364 native
taxa, including species and subspecies (Marticorena and Quezada, 1985; Marticorena
and Rodriguez, 1995, 2001) and between 552 and 723 alien species, depending on the
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Martín-Forés et al.
Plant species colonization in Chilean espinales
author considered (Arroyo et al., 2000; Castro and Jaksic, 2008). Paradoxically, despite
the large amount of alien species present and their early colonisation, Chile has been
considered a region that has been less invaded or is in an earlier stage of invasion than
other Mediterranean regions of the world (Arroyo et al., 2000; Figueroa et al., 2004;
Castro and Jaksic, 2008).
The arrival of exotic species to Chile started with the European colonisation in
the XVI century (Arroyo et al., 2000; Figueroa et al., 2004), which marked the first
deliberate introduction of animals and plants (Montenegro et al., 1991; Jaksic, 1998).
The rate of species entry in these early days is unknown, since the initial systematic
botanic descriptions of flora date from the XVIII century and were performed by
botanists who were more interested in describing the native species than the exotic ones
(Gay, 1845-1854; Reiche, 1896-1911). By the end of the XVIII century, numerous exotic
species had become naturalised in Chile (Figueroa et al., 2004), such as Cardamine
hirsuta L., Medicago polymorpha L., Spartium junceum L. and Bromus hordeaceus L.
(Castro et al., 2005). Although this species introduction has not been consistent over
time, a rate of two to three species per year is estimated, which is lower than the four
to six species recorded for other Mediterranean regions (Groves, 1991; Kloot, 1991,
Rejmánek et al., 1991; Wu et al., 2003).
Processes and mechanisms in species introduction
Changes in land use constitute the main factor determining processes of colonisation
and naturalisation of plant species (Le Houérou, 1991; Huston, 1994; Holmgrem et
al., 2000). Among the most influential factors, deforestation, fires and particularly
agricultural practices have been highlighted (Le Houérou, 1991; Cowling et al., 1996;
Williamson, 1996; Hobbs, 1998). With regard to deforestation, although Chile still has
one of the biggest areas of temperate forest in South America (Donoso, 1993), much of
it has been deforested for pastures or croplands. This process started in the XVI century,
although the main boom was during the middle of the XX century, with the expansion
and intensification of wheat crops (Echeverría et al., 2006) and the spread of forestry
plantations. In relation to fire, unlike other Mediterranean climate areas, fire has not
constituted a factor of natural disturbance in Chile (Muñoz and Fuentes, 1989; GómezGonzález and Cavieres, 2009), a fact that accounts of the absence of specific adaptations
in native species. Although there is evidence of fires of human origin in earliest
settlements in the region, around 14 000 years ago, these were not significant until the
arrival of the Spanish (Aronson et al., 1998; Aravena et al., 2003; Villa-Martínez et
al., 2003). Subsequently, fires became more frequent and intense, and currently about
5000 ha of native scrubland are burnt each year, the vast majority of these fires being
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Martín-Forés et al.
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Plant species colonization in Chilean espinales
intentional (Gómez-González and Cavieres, 2009). It has been suggested that plant
communities under novel fire regimes are more susceptible to invasion than those under
a natural (historical) fire regime (Trabaud, 1991; D’Antonio, 2000). However, different
studies that have analysed the effects of fire upon native and non-native Chilean flora
appear to indicate that fire is not a relevant factor with regard to favouring non-native
species (Keeley and Johnson, 1977; Holmgrem et al. 2000; Gómez-González et al.,
2010). Fire does, however, constitute a notable advantage for the establishment of
annual plants, which are poorly represented in Chile’s native flora (Arroyo et al., 1995).
The implementation of European agricultural culture in the XVI century leaded
to big changes in land uses and landscapes in Chile, the extent of which are yet not
well known (Turner et al., 1995). The effects of agriculture have been both direct
(ploughing, cropping and grazing) and indirect (fire and deforestation, employed as
techniques for preparing the land for agriculture and livestock farming). Livestock was
introduced into Chile, perhaps at the same time as the colonisation by Europeans. Other
herbivores, however, such as rabbits and hares, were brought more recently, in the XIX
century (Jaksic and Soriger, 1981). Several studies have associated the naturalisation of
exotic plant species with grazing by both livestock (Holmgren et al., 2000; Pauchard
and Alaback, 2004; Del Pozo et al., 2006; HilleRisLambers et al., 2010) and rabbits
(Sáiz and Ojeda, 1988; Holmgren et al., 2000; Holmgren, 2002). The effect of grazing
on native or non-native species has been characterised using morphofunctional plant
traits, revealing differences in their response. For instance, grazing appears to favour
some exotic creeping species, such as Erodium cicutarium (L.) L’Hér. and some
leguminous species, in detriment to the upright ones, the latter more closely associated
with native species (Holmgren et al., 2000). Grazing also favours substitution of native
hemicryptophytes by both native and non-native annuals, capable of resisting periods of
drought stress as seeds.
Many exotic plants were also introduced associated with crops, and became widely
distributed as a result of the importance of agriculture in the country (Castro et al.,
2005). Crop fields, particularly along their succession stages following abandonment,
constitute the ecosystems presenting the highest values for richness and cover of
non-native plants in Chile (Figueroa et al., 2011). Since colonisation the introduction
process has continued, although with different rates along time. Thus, Aronson et al.
(1998) highlight an initial wave of exotic species from 1880 to 1920 associated with
transformation of the landscape. Fuentes et al. (2008) recognise an initial phase (19101940) associated with intense development of agriculture (Cariola and Sunkel, 1982), as
well as a second phase (1980-2000) related with a sharp increase in the mechanisation of
farms and forest plantations at large scale. Matthei (1995) describes a sustained increase
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in the naturalisation of exotic species from 1894 to 1934 associated with wheat imported
from other countries.
Although many species were accidentally introduced into Chile (Arroyo et al., 2000),
many were taken for agricultural, medicinal, culinary and, more recently, ornamental
purposes. Since the arrival of the Spanish, species were introduced associated with hay
for livestock fodder and with wool and cereals. Endozoochory likely constitutes the most
frequent mechanism of effective seed dispersal. However, during transit from Spain
to Chile, exozoochory was probably the most effective means of dispersal, given the
difference in time between the transoceanic voyage and the time required for seeds to
pass through an animal’s digestive tract (generally less than 1 week; Malo and Suárez,
1997). Europe was not the only centre of introduction of non-native species to Chile.
During the Gold Rush, in the middle of the XIX century, there was intense wheat trading
with California (Davis, 1894), a fact which mobilised other species, together with grain
and straw, in both directions (Le Houérou, 1991; Jiménez et al., 2008), especially from
Chile to California.
Comparative studies of transcontinental naturalisation
Comparison of non-native floras between climatically similar regions constitutes
a very useful tool for understanding aspects associated with the species naturalisation
process (Pauchard et al., 2004, Hierro et al., 2005). It can help us to understand the
effects of changes in the landscape associated with historical or cultural scenarios upon
the naturalisation process (Kruger et al., 1989; Aschman, 1991). Among comparative
studies of different regions with Mediterranean climates, those between Chile and
California have been intensely analysed (e.g. Parsons, 1976; Arroyo et al., 1995;
Holmgren et al., 2000; Pauchard et al., 2004; Jiménez et al., 2008). These researches
highlight the large number of species common to both regions (386, which is 64% of
Chile’s non-native flora; Pauchard et al., 2004), as well as the fact that their communities
are undergoing different process of invasion with similar consequences of floristic
homogenization (Figueroa et al., 2011). However, to date there have been no comparative
studies between Chile and Spain, despite their climatic and geomorphological similarity,
the historical relationships that favoured the entry of species into Chile and the large
number of species common to both countries. Many of these naturalised species are
associated with the espinal, an agroecosystem presenting a management system and
structure similar to those of the Spanish dehesa (Ovalle and Avendaño, 1987; Ovalle et
al., 1990). However, as these agricultural systems are similar in both countries, we are
not fully aware of the mechanisms underlying the arrival of determined species and the
subsequent naturalisation and spread thereof.
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The aim of this paper is to analyse the relative significance of the biogeographic
origin, lifecycle and representativeness of taxonomic families in Chile’s non-native flora.
We considered three scales of analysis: national, regional (the central zone presenting
a Mediterranean climate) and community (the espinales within the central zones). In
the third case, we also conducted a comparative study with Mediterranean grasslands
from the Iberian Peninsula in order to identify the degree of similarity in the floristic
composition of communities with comparable ecological and agronomic characteristics.
Study area
The espinal is an anthropic savannoid formation characterised by dispersed trees of
Acacia caven Mol. (the espino) within an herbaceous matrix comprising mainly annual
plants of Mediterranean origin (Olivares and Gastó, 1971; Ovalle et al., 2005; del Pozo
et al., 2006). It supports a rural population of approximately 350 000 people, as well
as the largest Chilean livestock: 800 000 sheep and 250 000 cows. It covers an area
of two million hectares in Chile’s central zone, ranging along the Central Valley and
the western slopes of the Coastal mountain range (Ovalle et al., 1999). It currently
occupies mainly the dryland sectors (Figure 1), from the river Petorca (32° S), bordering
on the arid Mediterranean region, to the river Laja (37º S), bordering on the perhumid
region (Fuenzalida and Pisano, 1965; Di Castri, 1968; Quintanilla, 1981; Rodríguez et
al., 1983). Further north, in the arid and perarid regions, some espinales can be found,
preferentially located in valley bottoms (Follman and Matte, 1963; Rodríguez et al.,
1983). Their distribution is associated with Mediterranean-climate areas, albeit with
very variable annual rainfall, from 160-200 mm at its northern limit and up to 10001200 mm at the southern one. It presents high species diversity (Gulmon, 1977; Solbrig
et al., 2002; Del Pozo et al., 2002) and to date, 215 species have been identified only in
the Cauquenes region (Ovalle et al., 1987). It originated through changes in land uses
after the Spanish conquest; the native sclerophyllous forest was cleared in order to open
up land for agriculture and livestock farming, which favoured gradual invasion by the
exotic species A. caven (Gulmon, 1977; Armesto and Pickett, 1985; Ovalle et al., 1990),
possibly from South America’s Gran Chaco (Holmgren, 2002).
The espinal is an agrosilvopastoral system presenting much similarity with the
Spanish dehesas and Portuguese montados. It has traditionally been based upon
two models of management: continuous extensive grazing in flatlands, occasionally
inundated during winter, and rotation of grazing and cereal cropping in the better
drained hillsides (Ovalle et al., 2005). In the latter case, the espino is periodically cut
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down for firewood and charcoal. The land is subsequently ploughed for sowing cereal
crops. After one or two years’ harvest, depending upon the fertility of the soil, the land
is abandoned and colonised by herbaceous species, while shoots grow from the stump of
the espino. In this phase, the land is used for extensive grazing with a low stocking rate
of approximately one sheep/ha (Del Pozo et al., 2006). The grazing period prior to the
following cropping cycle is variable, from three to 40 years, depending on the fertility of
the soil.
Location of Chile (in grey) in South America and enlargement of the Central zone. The
shaded area represents the main distribution areas of espinal (modified from Ovalle
et al., 1999).
Localización de Chile (en gris) dentro de Sudamérica y ampliación de la Zona Central. La región
sombreada representa el área de distribución de los espinales (modificado de Ovalle et al., 1999).
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The espinal is currently more degraded than in past times. In the first place, an
increasingly greater area of the espinales in the Central Valley is being replaced by
intensive irrigation agriculture. Furthermore, many owners have abandoned their
traditional farming activities for forest plantations (approximately 80 000 ha/year),
mainly of Pinus radiata D. Don or Eucalyptus globulus Labill. Finally, the territory still
maintaining functional espinales has usually low fertility and soil erosion, leading to low
agricultural production. At present, 40% of the area is occupied by espinales with very
little tree cover (<25%) and 4% is dominated by romerillo (Baccharis linearis (Ruiz and
Pav.) Pers.), which indicates a situation of degradation and abandonment. Under these
circumstances, the cereal-pasture rotation system is in decline and limited to small areas,
almost exclusively for people’s own consumption.
For our research we developed a database containing all non-native species found
in Chile. We used the catalogue of Chilean flora by Marticorena and Quezada (1985,
1987), along with information provided by the Laboratorio de Invasiones Biológicas
de la Universidad de Concepción ( This list was
complemented with an extensive bibliographic revision incorporating some exotic
species recently cited in the country. Each species was characterised by assigning its
taxonomic family, area of origin, life cycle and distribution range within Chile. To assign
the area of origin and life cycle we used different regional floras, fundamentally Flora
Iberica (Castroviejo et al., 1986-2010), Flora Europaea (Tutin et al., 1964-80) and Flora
del Cono Sur (Zuloaga et al., 2008). As regions of origin we considered the four big
continents: Eurasia, Africa, America and Australasia. Apart from these four large regions
we considered independently the Mediterranean Basin (SE, S and SW of Europe, N
Africa and SW Asia), given the non-native typology of species present in Chile. Species
present in more than one continent were classified as Cosmopolitan. As for the life cycle
trait, we classified them into three groups: annual, perennial herbaceous and woody
species. Finally, for the distribution range within Chile we took the 15 administrative
regions as units (from the Tarapacá Region in the far north to the Magallanes Region at
the southern limit), excluding the species present only on the islands belonging to Chile
(Easter Island and the Juan Fernández archipelago). We obtained these data mainly from
Castro et al. (2005), Martincorena ( and Zuloaga et
al. (2008). We considered the set of non-native species present throughout the country,
as well as those distributed only in the central zone (from the Coquimbo region in the
north to the Bío-Bío Region in the south; Figure 1).
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In order to complement our database of non-native species, in 2010 we conducted
field samplings, both in Chilean espinales and in Spanish Mediterranean grasslands. In
each country we selected 15 sites and in each one we chose two 10 x 10 m plots in which
we randomly distributed six 50 x 50 cm quadrants. In each quadrant we recorded the
presence of all plant species. Each species was assigned a value of frequency according
to the number of sites in which it was recorded in each country. The 30 sites selected
presented similar lithological characteristics (acidic materials associated with igneous
or metamorphic rocks), geomorphological ones (undulating topography) and those
relating to use history (management for extensive livestock farming). In the case of
Chile the 15 sites were distributed within the Mediterranean region, over more than 600
km, from 32º 31’ 35’’ to 37º 00’ 10’’ S latitude. In the case of Spain the 15 sites were
distributed within the centre-western areas (Extremadura, N Andalucía and W CastillaLa Mancha, from 40º 14’ 45’’ to 37º 51’ 40’’ N latitude). For all species of European
origin, we standardised the nomenclature in accordance with Flora Iberica (Castroviejo
et al., 1986-2010), and in the case of families as yet unpublished in this study, with Flora
Europaea (Tutin et al., 1964-80).
The list of Chile’s non-native flora comprises 773 species or subspecies, which,
following exclusion of spurious citations, was reduced to 698 taxa (548 in the central
zone and 75 in the espinales), 50 of which are woody species and the rest herbaceous
ones. This large set of species is distributed into 72 families, of which the best
represented ones at country scale are Poaceae (19.8% of species), Asteraceae (13.9%)
and Fabaceae (10.2%). This distribution of species into families is very similar when
the Chile’s central zone was exclusively considered (Figure 2). However, the spectrum
of families present in the flora of the espinales is very different, and is characterised by
a higher proportion of Poaceae, Caryophylaceae, Rubiaceae, Rosaceae and Geraniaceae,
and a decrease in the proportion of Brassicaceae. This distribution into families of the
espinales is relatively similar to that seen in Spanish grasslands, except for the fact that
Spain presents more Fabaceae (18.5% vs 12%) and Brassicaceae (3.5% vs 1.3%), and
the espinales more Caryophylaceae (9.3% vs 7.1%), Rubiaceae (4% vs 1.3%), Rosaceae
(5.3 vs 1.3%) and Geraniaceae (5.3% vs 1.8%).
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Each histogram shows the frequency calculated at country scale (white bars; n = 698 species), of Chile’s central
zone (light grey bars; n = 547 species) and of the espinales (dark grey bars; n = 75 species). In the case of Spanish
grasslands (black bars) frequency was calculated with the total number of species found in the field samplings (n =
229 species)
Frequency histograms of the distribution of non-native species according to taxonomic
families. Only the nine most represented families are shown.
Histogramas de frecuencia de la distribución de las especies no nativas según la familia a la que
pertenecen. Solo se muestran las nueve familias mejor representadas.
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With regard to the area of origin of Chile’s non-native species, at the scale both of
country and of the central zone, species of Eurasian origin (39.7%) dominate and, to a
lesser degree, those of Mediterranean origin (28.4%) (Figure 3). However, when only the
flora of the espinales is considered, the species from the Mediterranean Basin increase to
49.3% and those of Eurasian origin to 48%, thus Eurasia in the broader sense constitutes
the area of origin of 97.3% of non-native species. This tendency is maintained in Spanish
grasslands, with values of 97.8% for species from Eurasia (72.1 and 25.7% for species of
Mediterranean and Eurasian origin, respectively).
Each histogram shows the frequency calculated at country scale (white bars; n = 698 species), of Chile’s central
zone (light grey bars; n = 547 species) and of the espinales (dark grey bars; n = 75 species). In the case of Spanish
grasslands (black bars) frequency was calculated with the total number of species found in the field samplings (n =
229 species).
Frequency histograms of the distribution of non-native species according to region of origin.
Histogramas de frecuencia de la distribución de las especies no nativas según la región
biogeográfica de origen.
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Both in Chile as a whole and in the central zone, the vast majority of non-native
species are herbaceous, both annual (55.8%) and perennial (37%) (Figure 4). This
trend is accentuated in the espinales, where none exotic woody species have been
found and annual plants represent 84%, a value very similar to the 81.2% of annual
species found in Spanish grasslands. Life cycle (annual, herbaceous perennial and
woody) is not independent from the area of origin of the non-native species (table of
contingency between areas of origin x life cycle, c2 = 169.18, p < 0.001): among the
species of African and American origin there are significantly more woody and perennial
herbaceous plants than expected at random; among those of Mediterranean origin there
are significantly more annual species; and among those of Australian origin there are
significantly more woody species.
For Chile, the central zone and the espinal we only considered non-native species. For Spanish grasslands, we
calculated frequency with the total number of species found in the field samplings.
Frequency histograms of the distribution of non-native species according to their life cycle:
annual (black bars), perennial herbaceous (grey bars) and woody (white bars) plant.
Histogramas de frecuencia de la distribución de las especies no nativas según su forma biológica:
anual (barras negras), herbácea perenne (barras grises) y leñosa (barras blancas).
In the field samplings we recorded a total of 229 species in the Spanish grasslands and
152 in the Chilean espinales, 49% of which were non-native. Of the 10 most common
species in Spain, all but one (Agrostis pourreti Willd) can be found in the catalogue of
non-native species of Chile; moreover, they were found in the sampling in the espinales
(Table 1). Likewise, of the 10 most common species in the espinales, all but one (Soliva
sessilis Ruiz and Pav.) are non-native and have been found in sampling conducted in
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List of the 10 most frequent species in Spain (a) and in Chile (b). For each species taxonomic
family, distribution in Spain and Chile and frequency of appearance in each country are
Listado de las 10 especies más frecuentes en España (a) y Chile (b). Para cada especie se indica
la familia taxonómica, su distribución española o chilena y la frecuencia de aparición en cada
a) Spain
Plantago lanceolata
Bromus hordeaceus
Trifolium campestre
Vulpia muralis
Hypochoeris glabra
Plantago coronopus
Agrostis pourretii
Trifolium glomeratum
Tolpis barbata
Leontodon taraxacoides subsp.
b) Chile
Petrorhagia prolifera
Soliva sessilis
Erodium cicutarium
Trifolium dubium
Briza minor
Erodium botrys
Hypochoeris glabra
Bromus hordeaceus
Aira caryophyllea
Leontodon taraxacoides subsp.
Frequency in Frequency in
Spain (%)
Chile (%)
50. 6
58. 9
58. 9
61. 7
50. 6
33. 9
46. 7
61. 7
Arroyo et al. (2000) and Figueroa et al. (2004) consider that in continental Chile there
is a total of 707 naturalised non-native species and subspecies. This value is somewhat
higher than the 698 taxa considered in this paper, approximately 12% of the total flora.
Despite the fact that some recent introductions have been done, a greater amount of
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species have been excluded from the list due to being spurious citations. The amount
of exotic species in Chile, however, is likely higher, considering the little botanical
prospection conducted in certain parts of the country. Thus, during the sampling and
in view of the lack of confirmation, at least 5 species had not been previously cited:
Aphanes microcarpa (Boiss. and Reut.) Rothm., Logfia minima (Sm.) Dumort.,
Moenchia erecta (L.) G. Gaertn., B. Mey. and Scherb., Trifolium cernuum Brot. and
Vulpia ciliata Dumort. Considering only the central Mediterranean zone, Figueroa et
al. (2011) recognised 2395 native species and 507 non-native ones, which account for
18% of the flora in this region (Arroyo and Cavieres, 1997; Arroyo et al., 2000). In this
case, our data contain 548 non-native species in the central zone, a difference possibly
associated with the geographic definition of this zone. This non-native flora contains a
large number of families, none of which show any clear dominance, although the most
common ones are Poaceae, Asteraceae and Fabaceae, in accordance with the three most
invasive families worldwide (Pyšek, 1998).
Most of Chile’s non-native species are annual plants (56%) of EurasianMediterranean origin (68%). The dominance of exotic annual species coincides with
the findings of other authors for other Mediterranean-climate regions (Le Floc’h 1991;
D’Antonio and Vitousek, 1992; Cowling et al., 1996; Figueroa et al., 2004; Norton
et al., 2007). Their rapid growth and high reproduction rates, and capacity to resist
unfavourable periods in the form of seeds makes them more competitive in repeatedly
disturbed open spaces, such as those created by fire, ploughing or grazing (Le Floc’h,
1991; Gómez-González et al., 2010). As for biogeographical origin, our results show
that they are preferentially from the Mediterranean Basin or, in the broader sense,
Eurasia, which would coincide with the findings of several authors for Chile and other
Mediterranean-climate areas (Montenegro et al., 1991; Arroyo et al., 2000; Holmgrem
et al., 2000; Figueroa et al., 2004; Castro et al., 2005).
The annual character and Mediterranean origin predominating in non-native Chilean
plants could be related to the different use history from that of the Mediterranean
Basin. The pastures of the Mediterranean basin have been subjected to an intense
grazing regime involving bovines and other domestic herbivores for over 6000 years
of its evolutionary history (Perevolotsky and Seligman, 1998). This long history of
coexistence with natural and anthropic disturbances has determined processes of
co-evolution between plants and agriculture (Di Castri, 1981; Cowling et al., 1996;
Perevolotsky and Seligman, 1998; Holmgren, 2002; Hayes and Holl, 2003; Kimball
and Schiffman, 2003; Ricotta et al., 2009; HilleRisLambers et al., 2010), which have
selected the plants presenting more competitive traits in a context of continuous grazing
such as, for example, forms of growth, concentration of nutrients in tissues or position of
growth meristems, among others (Adler et al., 2004; Díaz et al., 2007). On the contrary,
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Chile’s Mediterranean region underwent a drastic transformation since the colonisation
only 500 years ago and its native flora might not have co-evolved with large herbivores
in the last 10 000 years. This lack of adaptation to continuous grazing means that, with
the introduction of livestock, native species are negatively affected, which favours the
establishment of alien species (Milchunas and Lauenroth, 1993; Holmgren et al., 2000;
Adler et al., 2004; Díaz et al., 2007).
The flora of Central Chile is a good expression of the country’s as a whole with
regard to percentages of the most represented families, life cycle and origin of nonnative species. This central zone, representing only 20% of Chile’s territory, contains
80% of all the country’s exotic plants. This high concentration of naturalised plants
(Matthei, 1995) can be accounted for by a conjunction of factors relating to history,
environment and land uses. Chile was conquered by land from the north through the
Atacama desert into the central zone. To the south of the Bío-Bío river, the territory was
not colonised until just over a century ago (Aronson et al., 1998). Given the arid and
semiarid conditions of the North, only the central zone, with its Mediterranean climate,
was suitable for agriculture and for 350 years, the species introduced were therefore
relegated to this region (Fuentes et al., 2008). It was only as from the XX century
that non-native species were introduced into the whole country, with mass and intense
deforestation of the austral temperate forest (Donoso and Lara, 1996). In short, it is the
central zone that has undergone the biggest change as a result of intensive agriculture
for over five centuries (Montenegro et al., 1991; Matthei, 1995). Moreover, it is the
most densely populated region (78% of the country’s population, with an average of 75
inhabitants/km2; Pauchard et al., 2006; INE, 2007) and with the densest roads network
(Arroyo et al., 2000).
The representation of the non-native flora tends to be greater at more detailed
spatial scales (Gaertner et al., 2009), a fact that can be seen in the espinales studied.
On one hand, the percentage of non-native species therein is much higher than the
13% for the whole country or the 18% for the central zone, reaching 49%. This value
is higher than the 36.8% reported by Figueroa et al. (2011) for espinales and very
similar to that reported by Montenegro et al. (1991). At family level, the percentages
of Caryophyllaceae, Rosaceae, Geraniaceae and Rubiaceae double and even quadruple
the values characteristic of the country or of the central zone. On the other hand, among
non-native species, annual plants show an increase from 56% for the whole country to
84%; exotic woody species were absent from our study. Finally, compared with the 68%
of species of Eurasian or Mediterranean origin in Chile, the espinales account for 97%.
In this context it is interesting to highlight the relationship between regions of origin and
species’ biological cycle, with a positive association among annual plants for species
from the Mediterranean Basin and among woody plants for those of Australian, African
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or American origin. This relationship is possibly associated with the way in which
the species were introduced into the country and with the climatic characteristics of
the recipient habitat: unintentionally in the case of most of the plants associated with
agriculture and livestock farming (Mediterranean annual plants adapted to grazing;
Holmgren, 2002) or for ornamental or forestry purposes (woody plants generally of
extra-European origin; Pyšek et al., 2011). The extreme values found in the espinales
(compared with the rest of Chile) referring to distribution of families, biogeographic
origin and life cycle are, however, of the same order of magnitude as those found in
Spanish grasslands (although proportionally Rubiaceae, Rosaceae and Geraniaceae
present higher percentages in Chilean espinales and Brassicaceae in Spanish grasslands).
Given that Europe, and specifically the Mediterranean Basin, have constituted the main
source of immigration since America was discovered (Di Castri, 1991; Figueroa et al.,
2004; Jiménez et al., 2008) the floristic similarity between both countries appears to
indicate that the espinales represent a copy of Europe’s agricultural model, in which not
only animal and plant species (livestock, cereals and associated weeds) were introduced
from Europe, but also technology (ploughing, harrowing, animal traction) and the
culture associated with management of the system (fallow, rotation). The final result
is an agroecosystem, the Chilean espinal, which not only presents great physiognomic
and functional similarity with the Spanish dehesa (Ovalle and Avendaño, 1987; Ovalle
et al., 1990), but is also similar with regard to the floristic characteristics evaluated in
this research. There are, however, differences between these two agroecosystems,
particularly with regard to species richness (much higher in Spain). It is known that
the number of alien species becoming established in a given country is lower than the
amount that might potentially arrive (Malo and Suárez, 1997) and the number of exotic
species capable of reaching Chile is estimated at between 7 070 and 35 400 (Castro et
al., 2005). An example is the Santiago Botanic Garden which, in the middle of the XIX
century, cultivated over 2000 exotic species, 43 of which became naturalised (Matthei,
1995; Castro et al., 2005). These data place in doubt whether the flora of the espinales
constitutes an impoverished version of Spanish grasslands because specific filters
existed that limited the number of species arriving or, on the contrary, whether there is a
predominance of the biotic and abiotic filters acting in the assemblage of communities in
the espinales.
The entry of alien species and the extinction of native ones in a region have been
considered a process of global biological homogenisation (McKinney and Lockwood,
2001; Olden, 2006; Olden and Rooney, 2006; Winter et al., 2009). Castro and Jaksic
(2008) conclude that this homogenisation process is not yet significant in Chile, given
that from colonisation times up to the present, only two species have become extinct
(Plazia cheiranthifolia (Remy) Wedd. and Menodora linoides Phil.) and that the floristic
similarity among regions has not significantly changed. Our data, however, and those
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provided by Jiménez et al. (2008) and Figueroa et al. (2011), would seem to suggest that
at least the espinales are being subjected to an intense homogenisation process, as can be
seen in the dominance of certain species (and families) common to Spanish grasslands on
the list of most frequent species in the espinales. The most frequent species is Leontodon
taraxacoides (Vill.) Mérat, first recorded in Chile in 1963 (Castro et al., 2005), which
contradicts the hypothesis that the exotic species appearing subsequent to 1950 present
a narrower range within the country (Arroyo et al., 2000; Pauchard et al., 2004; Castro
et al., 2005). This would appear to indicate that this homogenisation process has become
accentuated in the last few decades.
At present the speed of change of many ecological, economic and social parameters
is reaching heretofore unknown rates, and the predictions are even more drastic in
reference to changes in land uses at global scale (Lugo and González, 2010; USDA,
2006). In this context of global change, there is a pressing need to understand these
processes of species colonisation and naturalisation. The exchange of species, successful
establishment of some of them or extinction of others undoubtedly contributes to the
appearance of new environmental scenarios, with socioeconomic repercussions that are
difficult to evaluate in the short term (Rockstrom et al., 2009). Identifying changes in
ecosystem structure and functioning in the short, medium and long term constitutes a
fundamental objective with regard to evaluating the “health” of ecosystems, as is stated
in the objectives of the Millennium Ecosystem Assessment Programme (Millennium
Ecosystem Assessment, 2005; Carpenter et al., 2009).
This research was supported by Spanish Ministry of Science and Innovation project
CGL2009-08718. We are grateful to the Spanish Ministry of Education, Culture and
Sport for providing a FPU PhD fellowship to I. Martín-Forés, the Government of Aragon
for awarding a research fellowship to the Residencia de Estudiantes; and finally, to the
Residencia de Estudiantes itself for its support.
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En la región mediterránea de Chile más del 18% de las especies de plantas
son exóticas. Este hecho es especialmente importante en algunos sistemas
agrosilvopastorales, como los espinales, muy similares funcionalmente a las dehesas
españolas y de gran interés ecológico y socioeconómico. En este trabajo se analiza la
flora no nativa de Chile considerando tres escalas de análisis: nacional, regional (zona
central de clima mediterráneo) y a escala de comunidad (los espinales dentro de la región
central), comparándola con la flora registrada en áreas equivalentes de la Península
Ibérica. Se discuten los posibles mecanismos que han podido operar en la colonización
florística desde la cuenca mediterránea hacia la región mediterránea chilena.
Palabras clave: Biogeografía, Chile, ciclo biológico, España, especies exóticas.
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Instituto de Ecología, A.C. Carretera Antigua a Coaptepec 351. El Haya 91070. Xalapa (Veracruz; México)
[email protected]
The introduction of different types of livestock following the Spanish conquest of
Mexico generated notable changes in the landscape of the Mexican High Plateau and
tropical regions of the country, replacing the mostly agricultural activities of indigenous
populations with European-style livestock production. While the effects on native
vegetation were significant, the dung produced by the livestock— mainly cattle, horses
and goats—did not create the same degree of environmental problems that later occurred
in Australia and New Zealand. This is explained because the dung beetles of Mexico—
and that of the Americas in general—were capable of exploiting this exotic resource and
incorporate it into the nutrient cycles of the tropical and temperate soils of the American
continent. This ability to utilize an exotic resource can be explained by the evolutionary,
biogeographic and ecological history of the species of beetles native to the American
continent. Biogeographically, Mexican dung beetle species come from phyletic lines
that originated in the Palearctic, Nearctic and Neotropical regions, arriving in Mexico in
different waves during the Miocene and Pleistocene. During these epochs, the megafauna
of Mexico included mammoths, mastodons, gomphotheres, horses, camels, glyptodonts,
bison and antelope, among others. These animals produced dung that was similar to that
of the livestock from Spain, especially that of cattle and horses. This made it possible for
the native dung beetles to easily exploit the resource provided by the livestock introduced
by the Europeans. In an anthropized landscape such as Mexico, we must focus not only
on the conservation of the dung beetles that inhabit our temperate and tropical forests,
but also on the species found in man-made pastures, both of which provide valuable
ecosystem services such as incorporating dung into the soil nutrient cycle.
Key words: Mesoamerica, Scarabaeinae, New Spain , megafauna.
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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Mexico covers approximately 2 million km2 and almost 62.5% of this area is used for
the production of cattle, horses, sheep and pigs. Cattle production alone covers 58% of
the country (INEGI, 2007) and there are more than 30 million heads of cattle in Mexico,
Each cow defecates 15 to 20 times per day and its dung can cover one square meter
per day (De Elias, 2002). The total weight of dung produced daily ranges from 20 to 30
kg per animal, though this varies with the breed of cattle and local climate conditions
(Bavera and Peñafort, 2006). At an average weight of 25 kg of excrement per bovine/day,
the 30 million heads of cattle in Mexico produce a total of 7.5 x 108 kg of dung daily; a
substantial amount, and one that does not even take into account the dung produced by
horses, sheep, goats and pigs. In ecological terms, however, this huge deposition of dung
has not created any substantial problem for the country. This is because a considerable
part of this resource is consumed by dung beetles that are abundant throughout Mexico,
and in the temperate and tropical regions of the Americas in general.
The dung produced by Eurasian livestock represents an exotic resource for the
American dung beetles, and one with which they have been in contact for approximately
500 years, dating back to when the Spanish began to transport animals and plants from
one continent to another. Hernán Cortés brought cattle to Mexico for the first time in
1521, the same year the Spanish conquered Mexico-Tenochtitlan.
There is no doubt that the introduction of livestock in Mesoamerica after the
European conquest was detrimental to the environment, but we must acknowledge
that the indigenous people also had modified their surroundings to a notable degree
(Simonian, 1999). Prior to the conquest, there was considerable exploitation of the
environment in this region, albeit in a manner that differed from that of the Spanish
farmers. For the indigenous cultures, the main productive activity was agriculture. On
the arrival of the Spanish, Mesoamerica may have had a population of around 25 million
people (Simonian, 1999). Efficient agricultural production, particularly in the tropical
regions, implied that large areas of tropical forest had been transformed into agrosystems
that, given cyclic management, could partially recover. Barrera-Bassols (1996) proposed
that prior to the arrival of the Spanish 1.5 million hectares of land were managed for
agricultural purposes, equivalent to 20% of the area of what is now the state of Veracruz.
This is a very large area, and gives an idea of the scale of the alterations to the landscape.
According to Simonian (1999), the anthropologist George A. Collier provides evidence
that the inhabitants of the highlands of Chiapas made extensive use of fire to clear
large areas of forest for agriculture before the Spanish arrived. Various pre-Hispanic
governments, including that of Nezahualcóyotl, the tlatoani (monarch) of the city-state
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of Tezcuco, passed forestry laws, concerned by the excessive clearing and the effects of
the uncontrolled use of fire on the forests and their fauna (see Simonian, 1999).
It is generally thought, however, that there was greater harmony between man and
his environment in pre-Columbian America. Here is an example that clearly illustrates
the lifestyle in Mexico-Tenochtitlan, as described by Escalante-Gonzalbo (2004): What
did the ‘Mexica’ do with human waste? Did they throw it into the water, contaminating
the lake in which their city was built? The answer is no, because people evacuated their
bowels using latrines that were on bridges over canals. Container canoes below the
latrines were used to transport the excrement to specific zones where it was processed
and transformed to an agricultural fertilizer and raw material for tanning leather.
Escalante-Gonzalbo (2004) states that this process, together with other systems of waste
management and in the absence of cattle, resulted in a rather clean city.
Introduction of livestock to Mexico
Livestock was brought to Mexico soon after the conquest. Horses were a key factor
in the conquest, and historians note the fondness of Hernán Cortés for bullfighting. The
fall and virtual destruction of Mexico-Tenochtitlan occurred on the 13th of August 1521,
and the first bullfight was held in Mexico City on the 24th of June 1526 to celebrate the
return of Hernán Cortés from his arduous expedition to Honduras. On the 13th of August
1529, bullfights were held to venerate San Hipólito on the day that Mexico-Tenochtitlan
had been conquered (Martínez, 1990). The first cattle to arrive in Mexico entered by the
port of Villa Rica de la Vera Cruz. According to Barrera-Bassols (1996), this first herd
gave rise to those of the central highlands of New Spain. In 1527, Nuño de Guzmán
introduced cattle from Cuba and Hispaniola (Santo Domingo and Haiti). Cattle from the
Guadalquivir marshes were brought to the tropics; extremely agile animals, they adapted
well to life in the tropical forests and over time became feral cattle, of which there are
still populations in Cuba today. In New Spain, by 1579, some northern ranches had herds
of up to 150,000 head that doubled in population every 15 months (McClung de Tapia
and Sugiyama, 2012). The average grazing density was one head of cattle per hectare,
which is maintained to this day in some regions of the country.
It is estimated that there were 1 300 000 cows in the center of New Spain, but also
8 100 000 sheep and goats (McClung de Tapia and Sugiyama, 2012). Thus, cattle
production was not the most widespread activity during the early stages of the colony;
in fact the production of sheep dominated during this period. This was because sheep
produced wool for exportation. Later, with the development of mining, cattle gained
importance as a source of leather. Their meat was of much less importance, given the
difficulties of preservation and transportation, which made trade in cattle particularly
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difficult in view of the small population of New Spain. The center of the country was
more heavily populated, however the people there mainly consumed the meat of pigs,
sheep and goats (G. Halffter pers. comm.).
Livestock production in Mexico radically transformed the original vegetation. The
expansive prairies of the Mexican High Plateau, coupled with the lack of predators
favored the growth of livestock populations to such an extent that many of these animals
formed feral herds (McClung de Tapia and Sugiyama, 2012). Overall, the introduction
of livestock to Mexico resulted in much deforestation, the appearance of new diseases
and water contamination, although not to such a large extent during the initial stages
(16th and 17th centuries). Indeed, following the conquest, there was actually a recovery
in the forests as a result of the gradual change from a crop-based system to one that
was predominantly livestock-based, leading to the abandonment of crop fields (BarreraBassols, 1996).
While there were severe changes to the environment with the introduction of
livestock, the enormous quantity of exotic dung produced daily by the rapidly
developing populations of Mexican livestock did not create an insurmountable problem
for New Spain, or independent Mexico, or even modern Mexico, with the exception
of certain specific cases. In Australia, the introduction of livestock, mainly cattle, was
catastrophic because the dung of these animals was neither consumed nor incorporated
into the soil nutrient cycle; however in Mexico, and tropical America in general, this was
not a problem, owing to the native dung beetles. Through their feeding and reproductive
behavior, dung beetles—mainly those of the subfamily Scarabaeinae—incorporate the
excrement of mammals and other vertebrates into the soil. Their activities favor the
recycling of nutrients in the ecosystem, which increases soil fertility and bioturbation
(displacement and mixing of soils by animals or plants) and reduces the presence of
parasites. Moreover, the dung beetles act as secondary dispersers of seeds (Andresen and
Feer, 2005; Nichols et al., 2008).
In Australia, the native dung beetle species are adapted to feed and reproduce mainly
upon the dung of marsupials. When livestock were brought from the Old World to the
Australian continent, a new type of dung was introduced that could not be utilized by the
native dung beetles, with enormous economic consequences (Bornemizza, 1979). The
partial solution has been the introduction of dung beetle species, mainly from Africa.
Currently, Australia has 23 introduced beetle species that eat the dung of the livestock,
and 437 known native species (Ridsdill-Smith and Edwards, 2011).
However, it was not necessary to bring in any new species of dung beetle for the
livestock introduced to the Americas because the native fauna easily appropriated this
new resource, leading to the efficient incorporation of dung, fertilization of the soil and
prevention of the development of flies, among other environmental services. In a recent
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introduction of exotic dung beetle species, between 1972 and 1974, which took place
mainly in Texas and California, Digitonthophagus gazella and Euniticellus intermedius
were released. Both of these species are African and were introduced in order to recycle
more efficiently the dung produced on cattle ranches (Anderson and Loomis, 1978;
Fincher, 1986). While the effect of the expansion of these exotic species throughout
Latin America has undergone various analyses, at least for Mexico (Montes de Oca
and Halffter, 1998) there is no clear evidence that they have had a negative effect on
the native populations. Perhaps this is because dung is not a saturated resource in open
tropical regions, but I will return to this point later.
The expansion of livestock production is a process that began recently, between the
1940s and 1950s, as a result of the colonization of the tropics in order to exploit unused
land. Prior to this, livestock production was an extensive activity, but one that was
limited compared to the current situation. Thus, although initial livestock production
caused noteworthy changes to the landscape, these were not as marked as those
occurring under the current rate of livestock expansion.
Certain questions arise in relation to the interaction between the dung of the
livestock introduced to America, and especially Mexico, and the native dung beetles:
Which species utilize this abundant resource? Where did these species originate? How
were they able to adapt to use a new, exotic resource so successfully? In this study, I
explore the evolutionary, biogeographical and ecological reasons behind the successful
appropriation of this exotic resource by the Mexican dung beetles, and identify the
species that have exploited the dung of this livestock, which is mainly from cattle and
THE Orography of Mexico
The complex orography of Mexico has favored the dispersal processes, vicariance
and in situ speciation (mainly in the mountains) of plant and animal species. This
combination of conditions has led Mexico to become one of the most megadiverse
countries in the world. The Mexican High Plateau is extensive, and is highest at its
southern end, close to the Trans-Mexican Volcanic Belt. As one advances northward,
its elevation decreases. In the Valleys of Mexico and Toluca, the Mexican High Plateau
reaches heights greater than 2300 m a.s.l. In the Chihuahuan Desert however, the
elevation is only around 1000 m a.s.l. The northern limit of the High Plateau is located
at the southern end of the Rocky Mountains. The Sierra Madre Occidental mountain
range, delimiting the Mexican High Plateau on its western slope, runs parallel to the
Pacific coast, from the border between the United States and Mexico to the state borders
of Nayarit and Jalisco. Its total length is 1400 km and average width is 200 km. The
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eastern end of the Mexican High Plateau is the Sierra Madre Oriental, which consists
of a series of elongated folds running in a NNW – SSE direction, from the north of
the Texas Plateau southeastward, where it is interrupted by the Trans-Mexican Volcanic
Belt, continuing SE, giving rise to the Sierra de Juárez mountains and terminating in the
Isthmus of Tehuantepec. It is approximately 600 km long and an average of 80 km wide.
The Trans-Mexican Volcanic Belt (or Neovolcanic Axis) is found between the 19th and
20th parallels and is some 950 km long and 50 to 150 km wide. At its eastern extreme,
the Trans-Mexican Volcanic Belt has a discontinuity, but then continues as the Sierra de
Los Tuxtlas mountain range, which is situated on the southern coastal plain of the Gulf
of Mexico with a NW-SE orientation. The tallest peaks in this range are 640 to 1,680
m a.s.l. (Guevara et al., 2004). The Sierra Madre del Sur mountains run parallel to the
Pacific coast, from the state of Jalisco to the eastern part of the Isthmus of Tehuantepec.
Between the Sierra Madre del Sur and the Trans-Mexican Volcanic Belt lies the Balsas
Depression, the lowest part of which is located at 300 to 500 m a.s.l. The Central
Massif and the Sierra Madre de Chiapas mountains are located in southeastern Mexico
and constitute the northern projections of the Central American mountain system. The
coastal plains run down the western and eastern sides of the country. East of Tabasco
is the Yucatan Peninsula, which rose from the sea following the meteorite impact on
Chicxulub at the end of the Mesozoic, ending the era of the dinosaurs. On the other coast
of Mexico, the Baja California Peninsula is crossed from north to south by the Sierra de
Baja California mountains.
This is the geographic landscape inhabited by the dung beetles, but it has not always
been so; physiographic and tectonic changes have been, and still are, very intense in
Mexico. Moreover, the dung beetles that presently inhabit Mexico have diverse
biogeographic origins.
THE biogeographic OriginS of the dung beetles in Mexico
In biogeographical terms, Mexico is a very interesting country. It contains a zone
of transition between the Nearctic and Neotropical regions (Darlington, 1957), which
Halffter (1964) called the Mexican Transition Zone (MTZ). The biota of the Mexican
High Plateau is fundamentally Nearctic and Palearctic in origin, while at the lower
elevations of the southern regions and on the coastal plains, the biota is mainly of recent
or ancient Neotropical origin.
The relationship between dung beetles and the expansion of livestock production in
Mexico must be analyzed in the context of the biogeography of the country. The Mexican
High Plateau is mainly populated by dung beetle species of Nearctic and Palearctic
origins, as well as those that have evolved in situ, as has occurred with other animal
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groups. In the Mexican tropics, we find species of both recent and ancient Neotropical
origins, and of Palearctic genera that have evolved in situ. The mountains host a mixture
of species of diverse biogeographic origins, with a clear dominance of Nearctic species
and those that have evolved in situ.
Halffter (1974, 1976) proposed several distribution patterns in the Trans-Mexican
Volcanic Belt to explain the origin of the insects in this region and they apply to dung
beetles. A distribution pattern refers to the current distribution of a group of organisms
that originated in a given area; organisms which have coexisted over a long period of
time and have a shared biogeographical history (Halffter, 1976).
The Paleoamerican Pattern
With origins in the Old World (Europe and Eurasia), the penetration of the insects
with this pattern into America, and also therefore the MTZ, was from the north. The
greatest diversity of insects at the supraspecific and specific levels in the Old World,
compared to that of North America, suggests that the main center of evolution was the
region comprising Europe and Eurasia. As with other insects exhibiting this pattern,
beetle genera of Paleoamerican origin are ancient and widely distributed at the global
Onthophagus and Sisyphus are beetle genera of Paleoamerican origin. There are
only two species of Sisyphus in the Americas (Halffter, 2003); however, at the species
level, there is a notable diversification in Onthophagus, especially in warm and warmtemperate climates. Its penetration into the MTZ was likely to have occurred prior to the
Miocene, when strong orogenic activity caused the uplift of the Mexican High Plateau
and the Trans-Mexican Volcanic Belt and the definitive formation of the Sierra Madre
mountain ranges of Mexico. The widespread distribution of Onthophagus in Mexico,
from the Mexican High Plateau and the mountains to the tropical forests, suggests
that they penetrated the MTZ before the existence of any large mountainous barriers.
These same mountains influenced speciation, giving rise to endemic species that are
found at different elevations, even in the foothills of the large mountain ranges. One
example is Onthophagus hippopotamus, which inhabits mole tunnels in the high reaches
of the Cofre de Perote Mountain in Veracruz. At the other extreme are the species of
Onthophagus which inhabit the Mexican Desert. Pre-Pliocene fossils of Onthophagus
and Oniticellus have been found in North America; the latter also belongs to the
Paleoamerican Pattern. Copris is another genus of Palearctic origin with 24 species in
the Americas, in addition to a fossil species (Matthews, 1962; Matthews and Halffter,
1968). In the United States of America this genus is found east of the 100th Meridian W,
it is also found in Mexico and Central America, and there is one species in the Antilles.
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Copris has a wide distribution, and more than 180 species with great diversification
in tropical Asia and Africa, as well as in tropical America. Within the genus of Copris,
one group of species (fricator) diversified in the mountains and another (minutus) has
two complexes, one in the eastern United States of America and another (complex
incertus) that is considered tropical. Various species of Paleoamerican genera certainly
originated in the New World from Palearctic lineages.
The Nearctic Pattern
This is formed by Holarctic genera of relatively recent penetration (Plio-Pleistocene)
that are restricted to the high reaches of the mountains of Mexico and of northern Central
America in the MTZ. They correspond to genera that are ecologically associated with
cold climates. Their habitat is mainly coniferous forest and high plains. In this pattern,
there are Nearctic and Holarctic lineages. In the mountains of the MTZ, species with
Nearctic affinities are generally found at a lower elevation than the Holarctic species
are. The typically Holarctic boreo-montane elements of the MTZ have moved toward
the south, exploiting the cold and humid conditions characteristic of the four glaciations
during the Pleistocene and especially the periods immediately following these events.
Within the Nearctic Distribution Pattern, some representatives of the family
Geotrupidae, the Geotrupini, are an example of an ancient group that has a sufficiently
long history in the Americas to be reflected in the differences between species in the
United States and those in the mountains of Mexico (Howden, 1964). Geotrupes and
Ceratotrupes are found in the MTZ where Ceratotrupes is considered endemic, while
Geotrupes is Holarctic and clearly related to the species of Asia and Europe (Howden,
1964). Both genera have numerous species endemic to Mexico.
The Mexican High Plateau Distribution Pattern
Formed by South American genera of ancient penetration into Mexico and Central
America, with species endemic to the Mexican High Plateau of Mexico, Chiapas and
Guatemala. They tend to dominate the Mexican High Plateau, and the high valleys of
Oaxaca, Chiapas and Guatemala. They have penetrated the temperate forests of the
mountains that border these geographic formations to a very limited extent. In terms
of the species and individuals in the arid regions of Mexico, the canthonine species are
generally the most numerous (Halffter, 2003). Species of the genus Canthon are found
throughout Mexico, except in the high reaches of the cordilleras above 2500 m a.s.l
(Halffter, 2003). According to the cladistic analysis done by Kholmann and Halffter
(1990), a group of canthonine species found in arid formations, temperate pasture, pine,
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pine-oak and temperate deciduous forest originated in an expansion during the Miocene.
The species of this expansion belong to the genus Canthon (C. humectus, C. pilularius,
C. imitator, C. vigilans, C. chalclites and C. obliquus), the subgenus Boreocanthon and
to the genus Melanocanthon (Halffter, 2003). With the exception of Canthon obliquus,
a species endemic to the Sierra de la Laguna mountains on the southern tip of the Baja
California Peninsula (Halffter, 2003), each of these species rolls a ball of dung, on which
they feed or reproduce.
Canthon humectus Say is considered one of the most successful species of the
Mexican High Plateau. It is distributed throughout the high valleys and plateaus of
Oaxaca, Chiapas and the northeastern and eastern Mexican High Plateaus, especially
in Jalisco and the warm dry areas of the Balsas River Basin, the valleys of TehuacánCuicatlán and Guatemala (Halffter et al., 2012). The Mexican High Plateau has been
inhabited by this species for a very long time and eight subspecies are recognized
(Halffter and Halffter, 2003). Its presence on the Mexican High Plateau suggests that
this species of Canthon, of Neotropical origin, arrived there during the Miocene prior
to the elevation of the Trans-Mexican Volcanic Belt (Halffter, 1976, 1987; Kohlmann
and Halffter, 1990). Within the genus Canthon, the pilularius lineage is also of old
Neotropical origin and four of its species inhabit the plains of the United States of
America, with one reaching the High Plateau: Boreocanthon puncticollis, which is a
canthonine of ancient origin.
One characteristic of the dung beetles on the High Plateau is the low number of
species and the dominance of some of these species as a result of recent anthropogenic
changes. For example, the extensive current distribution of Canthon humectus in the
High Plateau is considered a direct consequence of the introduction of livestock by
the Spanish, and also of the intense deforestation associated with the expansion of
this productive activity (Verdú et al., 2007; Halffter et al., 2012). Cattle produce an
abundant amount of dung, which is the most important source of food for C. humectus
and deforestation has resulted in the high light intensity that is favorable to most of
its subspecies, though some subspecies mainly live in the relatively shady conditions
of the High Plateau forests (Halffter et al., 2012). While it seems contradictory, the
introduction of livestock has favored some Mexican species. A similar process occurs
in parts of South America; Alfaro et al. (2008) describes how, in the coastal dunes of the
Atacama and Coquimbo regions in Chile, the presence of herds of donkeys ensures the
availability of food for Megathopa villosa. What did Canthon humectus and Megathopa
villosa feed on before the introduction of livestock? Were they as abundant as they are
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The Typical Neotropical Pattern
This pattern includes South American genera and species that have entered the MTZ
in relatively recent times, following the low lying tropical regions and with minimal
taxonomic separation from the South American forms. The northern limit of penetration
of these elements is formed by the Mexican coastal plains covered in tropical forest,
although these taxa can reach the southern and even eastern United States. The
Canthonini tribe is very well represented in the Americas, and comprises a large number
of species.
Canthonine species are roller beetles. These beetles, as described previously, are
named for their ability to make a food ball out of dung, roll and bury it a certain distance
away. The female modifies this food ball into a nesting ball, covering it with earth and
laying an egg in the ball (Halffter and Matthews, 1966). Within the Mexican canthonines,
there are strictly coprophagous, copronecrophagous and strictly necrophagous species
(Halffter, 2003). In the tropical forest, there are a considerable number of species, but the
relative number of individuals in open areas is greater than that found within the forest.
Canthon indigaceus (a species typical of pastures), C. cyanellus and C. morsei, as well
as species of the subgenus Glaphyrocanthon, form the group that expanded from South
America in a second Pleistocene wave, and are associated with tropical ecosystems;
while the viridis group specialized in Mexico to the south of the Trans-Mexican Volcanic
Belt (Halffter, 2003). The ecological equivalent of the Coprini tribe in the Neotropical
fauna is the tribe Dichotomiini, the ecological role of which is very similar to that of the
Coprini, but with an area of evolution that is Gondwanian, specifically American.
The Mesoamerican Montane Pattern
Its origin is in the montane Central American Nucleus, from where expansion could
have taken place to the north and south. The majority of its elements have an ancient
South American affinity.
Native Mexican vertebrates currently equivalent to the
introduced livestock
Animals heavier than 44 kg are classified as megafauna, therefore cows, horses and
donkeys can be considered part of this group, although the term is more commonly
used in the context of extinct animals. Arroyo-Cabrales et al. (2008) state that there
are currently only four large native species in Mexico: the pronghorn (Antilocapra
americana), the mule deer (Odocoileus hemionus), the white-tailed deer (O. virginianus)
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and the bighorn sheep (Ovis canadensis), although in relatively recent times there has
also been the bison, which was reintroduced in northern Mexico. In the tropical regions
of Mexico, we could also include the tapir which is 70 cm to one meter in height and
weighs 180 to 300 kilos. Its feces are quite similar to those of the horse, but contain a
greater quantity of fibrous material (Elizondo, 1999). However, unlike cattle, the tapir
lives mainly in the tropical forest and on the banks of rivers. It is therefore difficult to
imagine that the shift by Mexican dung beetles from using native animal dung to that of
the introduced livestock can be explained by the similarity between their dung and that
of the current native vertebrates. It is also clear that, given the complex orography of the
country and the biogeographic history of the Scarabaeinae, we cannot easily explain the
success of this group of insects in terms of the appropriation of a food resource that is
apparently new to them.
The megafauna
How do we explain the fact that the introduction of livestock to Mexico and to
America in general, has not created a serious ecological problem in terms of the
recycling of dung? I suggest that the explanation lies in the type of resources used by the
beetles in past eras. I refer mainly to the dung of the megafauna that began to disappear
ten thousand years ago. The greatest diversification of dung beetles is associated with
the spread of herbivorous mammals in the Mesozoic (Davis et al., 2002; Arillo and
Ortuño, 2008). Fifty million years ago, in the Miocene, the continents were populated by
more than 150 genera of megafauna, but by the end of the Pleistocene, 10 000 years ago,
97 of these had disappeared (Barnosky et al., 2004). In the case of Mexico, and the MTZ
in particular, the successful penetration of the South American Scarabaeinae species in
the Miocene and Pleistocene must have occurred thanks to the presence of a mammalian
fauna and megafauna that provided abundant food resources for the dung beetles. We
can say the same for the dung beetles of Holarctic and Nearctic origins.
In Mexico, the Miocene and Pleistocene mammal communities were more
diverse than those of today. Some 10 000 years ago, Mexico had a wide diversity of
medium-sized and large mammals with at least 61 large species of mammals during
the Pleistocene, the majority of which were herbivores. These include animals
weighing more than a ton, such as the mammoths (Mammuthus sp.), the American
mastodon (Mammut americanum), the gomphotheres (Cuvieronius, Stegomastodon)
and the ground sloth (Glossotherium, Eremotherium), as well as glyptodonts, horses,
camels, llamas, bison and antelope. These lived alongside similarly large predators
such as wolves, lions, saber-toothed tigers and bears, among others (Arroyo-Cabrales
et al., 2008; Pérez-Crespo et al., 2012). Even Baird’s tapir, Tapirus bairdii, currently
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distributed from Tehuantepec to the coast of Ecuador, also lived in northern Mexico and
the southern United States (from California to Florida) during the Pleistocene, where it
died out 10,000 years ago. In addition to these, rhinoceros (Teleoceras) remains have
been found in Miocene sites in northern, central and southeastern Mexico (CarbotChanona et al., 2009).
Additionally, in the state of Oaxaca, in southeastern Mexico, there are records of
Mammuthus columbi, the Columbian mammoth, and of another two proboscidean
species of the family Gomphoteriidae: Rhynchotherium praecursor and Cuvieronius
tropicus (Pérez Crespo et al., 2008). For Oaxaca, these authors describe finding fossil
evidence of Bison sp. Bison fossils have even been found in Nicaragua. In the river
basins of Oaxaca, the remains of megafauna from the Tertiary have been found, dated
at 2 to 3 million years old. These include the camel Camelops, horse Equus, elephant
Elephas, cow Bos and the pronghorn Antilocapra.
The successful appropriation by dung beetles of the dung of introduced livestock
in Mexico, and in America in general, was possible because they had been in contact
with the dung of prairie mammal herbivores that certainly shared the properties and
characteristics of the dung of current livestock. This is true mainly for cattle, mules
and horses but also for sheep and goats, the dung of which is similar to that of deer.
During the Miocene, pastures similar to savannas appeared in North and South America
as a consequence of global cooling and the progressive increase in the aridity of the
climate. Using biogeochemical markers, Pérez-Crespo et al. (2012), conclude that the
Columbian or prairie mammoth (Mammuthus columbi), should be more appropriately
named the pasture mammoth, after these pastures that resembled the savannas of Africa
and had trees that allowed the animals to enjoy a mixed diet from grazing on the grass
and browsing the trees. Such a diet would certainly have generated dung similar to that
of present day cattle.
What quantity of dung was produced by the extinct megafauna? We cannot say,
but it would have been enormous. Consider the fact that that one elephant produces
approximately 200 kg of dung every day. If the 30 million head of cattle in Mexico were
elephants, they would produce a total of 60 x 108 kg of dung, daily. Something had to
remove such a huge amount of mega-dung in the past or an ecological collapse would
have occurred very quickly. The natural candidates for this task are the dung beetles of
that epoch.
However, this megafauna declined significantly during the Pleistocene due to various
factors including changes in the environment and in human activity (Arroyo-Cabrales et
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al., 2008). These authors state that, of the Mexican pleistocenic paleomastofauna, one
order (Notoungulata), six families (20.9%), 29 genera (19.9%) and 77 species (28.1%)
died out, including two of the three families of the order Proboscidea (Gomphotheriidae
and Mammutidae), and three of the six families of the superorder Xenarthra
(Glyptodontidae, Megatheriidae and Mylodontidae). The Proboscidea disappeared
completely from Mexico and the rest of the Americas. Some families did not suffer
worldwide extinction, but others, such as Antilocapridae, Bovidae and Equidae, lost
most of their species. Of 78 extinct species of large mammals, 62 were herbivorous.
This suggests that the dung beetles associated with those herbivores also suffered a
notable reduction in species. A good indication of this is the low number of dung beetles
species on the Mexican High Plateau, and the success and expansion of C. humectus
with the introduction of livestock by the Europeans. All of this suggests that this species,
or its ancestors, were very well adapted to consuming the dung of the megafauna present
from the Miocene to the Pleistocene. Such a quantity and diversity of types of dung
must have sustained a great number of dung beetle species that are now extinct. Another
element that suggests that there was a more diverse beetle fauna in the past is the fact
that cattle dung is not fully exploited in the grasslands of Mexico and the rest of the
Americas. It has been explained that this is a consequence of an ecological semi-vacuum
(Halffter et al., 2008), but what has not been explained is why this vacuum exists.
I propose that this ecological vacuum is the result of the extinction of many dung
beetles species that consumed the dung of the megafauna that started dying out during
the Miocene, and mostly disappeared during the Pleistocene. The disappearance of a
significant number of megavertebrates must have produced a cascade effect that also
caused the loss of many species that depended on their dung. Dung beetle species number
more than 150 in African savannas (Cambefort, 1991) and are supported by the dung
of the big mammals such as elephants, rhinoceros, giraffes and impalas, among others.
According to Davis et al. (2002), two principal ecological factors influence the present
tribal, generic and species distribution patterns of Scarabaeinae dung beetles around
the world: climate suitability and the number of types of dung. There is a relationship
between dung diversity and the biogeographical and evolutionary history of mammals.
Regions with low dung diversities (mostly comprising pellets and small omnivore/
carnivore droppings) feature low taxonomic diversification and high proportions of
ball rolling taxa (Davis et al., 2002). However, regions with higher dung diversities,
comprising the dung of large herbivores, have greater taxonomic diversification,
and one that is numerically dominated by tunneling taxa (including endocoprids and
kleptocoprids). If that is true for the recent dung beetles, the same rule is likely to have
applied in the past. Thus, a high species diversity of dung beetles (perhaps 50 to 100
species, or more) could have existed in the past on the Mexican High Plateau and in the
tropical regions of the country. If this high dung beetle diversity did not exist, then the
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few species present at that time would have been hyperabundant in terms of the number
of individuals.
Current Scarabaeinae species IN THE cattle production
AREAs and LIGHT-FILLED environments OF Mexico
Table 1 lists the species that consume livestock dung present in the pastures and
open areas of the Mexican High Plateau, as well as in tropical regions and mountains,
according to a bibliographic search and interviews with some Scarabaeinae experts.
Most of the taxa are species, but some are subspecies. I will consider all of them as
species in this analysis. This list should be considered preliminary and will be expanded
and refined with the support of colleagues and as more specific studies of dung beetle
diversity in areas of livestock production in Mexico become available. Many species
found in pastures are not on the list as they were considered to be unrepresentative of
these environments; however, future studies may well modify this point of view. Open,
light-filled environments—suitable for heliophilous organisms—are characterized by
herbaceous plants and grasses, ranging from open prairies to semi-wooded and wooded
savannas that allow sunlight to enter relatively shady sites.
The highest diversity of heliophilous species in Mexico is found in the tropical
regions. I have recorded 24 species, including three subspecies of Canthon indigaceus
and one of Phanaeus tridens. Almost 83% of the species are tunnelers or resident and
about 17% are rollers. Canthon indigaceus chevrolati is very abundant on the coastal
plains of the Gulf of Mexico and in the Yucatan (Halffter et al., 1992; Montes de Oca et
al., 1991; Basto-Estrella, 2012), C. i. indigaceus is found on the tropical Pacific slope,
while C. c. chiapas is in southeastern Mexico. Species of Phanaeus are very abundant
in the light-filled regions of the Mexican tropics, especially Phanaeus damon, which
is widely distributed on both coastal plains (Deloya and Moron, 1994) and which has
benefitted from livestock production. Dichotomius colonicus is a very aggressive species
that quickly invades open territory and is found distributed throughout Mexico, except
for the Baja California peninsula (Kohlmann, 2003). Copri incertus is another highly
abundant species in tropical Mexico, as is Onthophagus incensus in certain areas.
Onthophagus landolti is very abundant in the open regions of Chiapas (Zunino, 2003)
and a recent study showed it to be the most abundant species in terms of numbers of
individuals on the livestock ranches of the Yucatan Peninsula (Basto-Estrella, 2012).
Digitonthophagus gazella and Euoniticellus intermedius are introduced species.
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M.E. Favila
Preliminary list of dung beetle species that consume livestock dung and are present in
pastures and open areas in Mexico.
Lista preliminar de escarabajos del estiércol que se alimentan de estiércol de Ganado y están
presentes en pastizales y áreas abiertas de México.
Tropical regions
Ateuchus rodriguezi DeBorre, 1866
Canthon (Boreocanthon) ateuchiceps Bates, 1887
Canthon (Canthon) indigaceus chevrolati Harold, 1868
Canthon (Canthon) indigaceus chiapas Robinson, 1948
Canthon (Canthon) indigaceus indigaceus LeConte, 1866
Copris incertus Say, 1835
Copris sallei Harold, 1869
Dichotomius centralis Harold, 1869
Dichotomius colonicus Say, 1835
Digitonthophagus gazella Fabricius, 1787
Euoniticellus intermedius Reiche, 1848
Phanaeus (Phanaeus) amithaon Harold, 1875
Phanaeus (Phanaeus) damon Castelnau, 1840
Phanaeus (Phanaeus) daphnis Harold, 1863
Phanaeus (Phanaeus) nimrod Harold, 1863
Phanaeus mexicanus Harold, 1863
Phanaeus (Phanaeus) tridens Laporte & Castelnau, 1840
Phanaeus ( Phanaeus) tridens pseudofurcosus Balthasar, 1939
Phanaeus (Phanaeus) furiosus Bates, 1887
Phanaeus (Phanaeus) scutifer Bates, 1887
Onthophagus batesi Howden & Cartwright, 1963
Onthophagus hoepfneri Harold, 1869
Onthophagus incensus Say, 1835
Onthophagus landolti Harold, 1880
Mexican High Plateau
Canthon(Canthon) humectus hidalgoensis Bates, 1887
Canthon(Canthon) humectus humectus Say, 1832
Canthon (Canthon) humectus incisus Robinson, 1948
Phanaeus ( Phanaeus) adonis Harold, 1863
Phanaeus (Phanaeus) quadridens Say, 1835
Phanaeus (Phanaeus) vindex Macleay, 1819
Onthophagus mexicanus Bates, 1887
Copris armatus Harold, 1869
Copris klugi Harold, 1869
Onthophaugus chevrolati chevrolati Harold, 1869
Onthophagus fuscus parafuscus Zunino & Halffter, 2005
Subfamila Geotrupinae:
Geotrupes (Megatrupes) cavicollis Bates, 1887
Geotrupes (Onthotrupes) herbeus Jekel, 1865
Geotrupes (Halffterius) rufoclavatus Jekel, 1865
Ceratrotupes bolivari Halffter & Martínez, 1962
Ceratrotupes fronticornis Erichson, 1847
Onthotrupes herbeus Jekel, 1865
Onthotrupes sallei Jekel, 1866
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On the Mexican High Plateau, there are approximately seven species associated with
open environments. Four of these are tunnelers and three are rollers, and they can be
found on some mountains within their distribution area. Here, the genus Phanaeus of
the mexicanus and daphni groups is formed by species that are undergoing expansion in
central Mexico as a result of livestock production. Canthon humectus is a very successful
heliophilous roller on the Mexican High Plateau and also owes its success to livestock
production (Verdú et al., 2007; Halffter et al., 2012). In the Barranca de Metztitlán,
Puebla, C. humectus hidalgoensis is the most dominant species. The frequency of this
heliophilous roller is high on the Mexican High Plateau, as is that of the other subspecies
C. humectus humectus, and C. humectus sayi in similar agroecosystems (Halffter et
al., 2012). Verdú et al. (2007) propose that an alternative food source for C. humectus
hidalgoensis could be the decomposing fruit and plant matter provided in Metztitlán,
Puebla by the widely cultivated species of Agave and Opuntia. If this is the case, it
is very likely that their populations would have been considerably smaller before the
conquest than they are now, thanks to the abundance of the dung produced mainly by
In the mountains of Mexico, there are 11 heliophilous species that consume the dung
of cattle and horses. Four of these species belong to the subfamily Scarabaeinae, and
seven to the subfamily Geotrupinae. All of these are tunnelers: there are no roller species
associated with the upper montane regions.
In an environment that is increasingly anthropized, the conservation policies that
limit traditional human activities such as livestock production and agriculture can have
a negative impact on the biota of certain regions by homogenizing the environment
and reducing the rate of species exchange between habitats (Halffter et al., 2012).
The current trend is to favor silvopastoral systems that create a complex, but highly
productive and economically beneficial environments, one that is also considered
to be the most environmentally friendly. For the dung beetle, traditionally managed
landscapes constitute a complex system that favors the interaction of livestock, grazing,
vegetation structure and dung beetle community systems (Verdú et al., 2007; Díaz et al.,
2010; Halffter et al., 2012).
New paradigms in conservation biology propose that connectivity, via corridors
of natural vegetation between patches of natural habitat, should be analyzed in the
context of the agricultural or managed matrix. The objective is to achieve a matrix
that is biodiversity friendly, and one that facilitates the movement of forest species
between forest fragments (Perfecto and Vandermeer, 2008). Living fences appear
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Livestock and dung beetles in Mexico
to act as continuous habitat corridors and allow forest beetles to colonize other forest
fragments (Estrada and Coates-Estrada, 2002; Díaz et al., 2010). With dung beetles,
however, we must also consider facilitating the movement of open area species between
pastures: the high turnover of species between pastures and forest fragments shows that
few forest species colonize the pastures and vice versa (Halffter et al., 1992; Estrada
and Coates-Estrada, 2002; Díaz et al. 2010). Connections between natural areas and
managed landscapes can therefore help sustain biodiversity and the natural processes
of ecosystems, in addition to transforming the unfriendly matrix into a more hospitable
managed matrix (Bennett, 2003; Vandermeer and Perfecto, 2007; Perfecto and
Vandermeer, 2008).
While the introduction of livestock has had a strong impact on the original
ecosystems of the Americas, and Mexico in particular, livestock dung itself has not
caused a problem. Nevertheless, the beetles that consume this dung now face new
challenges that threaten their survival, and the environmental service of dung recycling
where livestock is produced is therefore being affected. Currently, the most serious
challenge is posed by the use of parasiticides and insecticides in livestock management
and these can kill or reduce the populations of dung beetles (Martínez and Lumaret,
2006; Nichols et al., 2008; Cruz, 2011; Cruz et al., 2012). This practice could end up
having a very costly impact on livestock production. The presence of introduced species
may pose another threat to the survival of heliophilous dung beetles, but we do not yet
have any specific studies on this topic. The view of the experts has been that the dung
beetles are the guardians of the temperate and tropical forests; a vision of guardianship
that is now justifiably widening to encompass the areas where livestock is raised.
I am grateful to Dr. Sergio Guevara for inviting me to write this contribution. A
single comment from him prompted the development of this study. The conversations
I had with Dr. Gonzalo Halffter allowed me to clarify many of my ideas regarding the
biogeography of dung beetles, as well as the process of livestock production in Mexico.
Any error in this vision is my sole responsibility. The list of heliophilous dung beetle
species common to light-filled areas, a term he helped me to define, was also discussed
with Dr. Halffter. I also assume responsibility for any omissions or errors on this list.
Bianca Delfosse significantly improved the style of my original version in English, I
am grateful to her for her diligent work. I humbly dedicate this contribution to Violeta
Halffter, who will always be remembered and admired.
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La introducción de diferentes tipos de Ganado después de la conquista de México
por los españoles generó notables cambios en el paisaje del Altiplano Mexicano y de las
regiones tropicales del país, al reemplazarse las actividades fundamentalmente agrícolas
de las poblaciones indígenas por el estilo Europeo de producción ganadera. Aunque el
efecto de esta actividad sobre la vegetación fue notable, el estiércol producido por el
ganado, principalmente el proveniente de vacas, caballos y ovejas, no generó el mismo
nivel de problemas ambientales que si se generaron con la introducción del ganado
en Australia y Nueva Zelandia. Esta diferencia se explica porque los escarabajos del
estiércol de México y de América en general fueron capaces de utilizar este recurso
exótico e incorporarlo al ciclo de nutrientes de los suelos templados y tropicales
del continente americano. La capacidad para utilizar un recurso exótico, como lo es
el estiércol del ganado introducido en América, puede ser explicado por la historia
evolutiva, biogeográfica y ecológica de las especies nativas de los escarabajos del
estiércol del continente americano. Biogeográficamente, los escarabajos del estiércol
de México provienen de líneas filéticas que se originaron en las regiones Paleárticas,
Neárticas y Neotropicales, llegando a México en diferentes oleadas que ocurrieron
durante el Mioceno y el Pleistoceno. Durante esas épocas, la megafauna de México
incluía mamuts, mastodontes, gonfoterios, caballos, camellos, gliptodontes, bisontes y
antílopes, entre otros. Estos animales producían estiércol que fue similar al del Ganado
introducido por los españoles, principalmente el de vaca y caballo. Esta situación hizo
posible que los escarabajos del estiércol nativos explotaran fácilmente el Nuevo recurso
provisto por el Ganado introducido por los Europeos. En un paisaje tan antropizado
como lo es actualmente México, debemos de enfocarnos no solo a la conservación de
los escarabajos del estiércol que habitan nuestros bosques templados y tropicales, sino
también tenemos que proteger a las especies encontradas en los pastizales ganaderos las
cuales proveen valiosos servicios al ecosistema, tales como la incorporación del estiércol
al ciclo de nutrientes, pero que se están viendo amenazadas por otras actividades
humanas como el usos de desparasitantes e insecticidas para el ganado y por el cambio
climático global.
Palabras clave: Mesoamérica, Scarabaeinae, Nueva España, megafauna.
Chapter 2. Landscape
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¹Red de Ecología Funcional, Instituto de Ecología, A.C., Carretera antigua a Coatepec No. 351
El Haya, Xalapa 91070, Veracruz, México. ²Estación El Carmen, Instituto de Ciencias del Mar y Limnología, UNAM, km
9.5 carretera Carmen-Puerto Real, Cd. del Carmen, 24157 Campeche, Mexico.
*Author for correspondence: [email protected]
Animal husbandry in Mexico began with the arrival of the Spaniards and the creation
of New Spain. It changed significantly in the middle of the 20th century with the
introduction of the Zebu breed of cattle and improved pastures. From the beginning,
wetlands were used for cattle grazing, and we describe the transformations that occur
in grazed wetlands that convert them into flooded pastures. The degree of impact
depends on the number of cows, the time they are in the wetland, and modifications
to hydroperiod and vegetation. We describe the changes in the level of flooding, the
soil characteristics (organic matter, water retention, bulk density, pH, micro- and
macronutrients) and floristic composition, and how all this affects the environmental
services produced by wetlands. With the introduction of cattle breeds tolerant of
tropical environments, mainly Zebu cattle, and of exotic forage grasses that can grow
in wetlands, the impact has increased. These grasses drastically alter the environment
(water and soil) and can become invasive. Therefore there is a gradient of transformation
from wetlands with no cattle impact, to those with slight changes that continue to
function as wetlands, and finally to heavily transformed wetlands. Management based
on low livestock intensity maintains the functions and environmental services provided
by wetlands while constituting a sustainable economic activity that permits these
ecosystems to be preserved.
Key words: Cattle ranching, compaction, invasives, sustainability, vertical accretion.
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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The most extensive tropical wetlands in Mexico are located along the coastal plains.
They include mangroves, freshwater marshes and swamps forming gradients, which
differ in their salinity, degree and temporality of flooding. Wetlands are transitional
between terrestrial and aquatic systems where the water table is usually at or near the
surface, or the land is covered by shallow water (Cowardin et al., 1979). These wetlands
are rich in plant and animal biodiversity and provide numerous valuable ecosystem
services. These are functions that contribute to human welfare and help sustain the
biosphere (Costanza et al., 1997). Thus, wetlands are not only places; they should be
considered as entities that benefit society. Their genetic diversity helps to maintain
wetland processes such as water storage, sediment trapping and nutrient cycling. Among
the most valued services of wetlands are disturbance regulation, waste treatment, water
supply, cultural and recreational uses, habitat, food production, and nutrient cycling
functions, such as processing nitrogen and phosphorus (Vörösmarty et al., 2005; Mitsch
and Gosselink, 2007). In particular, tropical coastal wetlands have been recognized
because they increase fisheries (Aburto-Oropeza et al., 2008), are important carbon
sinks (Moreno et al., 2002; Campos et al., 2011; Marín Muñiz et al., 2011), store water
(Campos et al., 2011), and they function as protective shields—bioshields—against
storms and surges (Selvam et al., 2005; Thuy et al., 2012). In a noteworthy study,
Costanza et al. (1997) estimated that tidal marshes, mangroves, swamps and floodplains
produced 4879 trillion dollars in services per year.
Mexico has lost or degraded 62% of its wetlands (Landgrave and Moreno-Casasola,
2012). Wetland degradation is not always obvious as direct physical destruction or
alteration. Among the major causes of loss and degradation are both human actions
and natural threats. Direct human actions include drainage, dredging and stream
channelization, deposition of fill, diking and damming, tilling for crops, levee
construction, logging, mining, construction, runoff, air and water pollutants, changing
nutrient levels (increased nutrient inputs and eutrophication), releasing toxic chemicals,
introducing nonnative species, grazing by domestic animals, and urbanization. Indirect
human actions are colmatation and eutrophication in downstream wetlands derived
from agricultural runoff and from erosion, respectively, due to deforestation and
farming upstream. Natural threats include erosion, subsidence, rising sea level, drought,
hurricanes and other storms (
Some of these natural threats will increase with global climate change (increased air
temperature; shifts in precipitation; increased frequency of storms, drought, and floods;
increased atmospheric carbon dioxide concentration; rising sea level; and increased
salinity in freshwater wetlands). All of these changes could impact species composition
and wetland functions. Moreover, these detrimental changes could favor the invasion of
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exotic species, which are more successful in disturbed, fragmented and/or species-poor
environments (Crawley, 1987; Rejmánek et al., 2005).
Semarnat (2008) indicates that 56% of the Mexican territory (1.09 millions km2) is
used for cattle ranching and in 2007, there were 23.3 million bovines (INEGI, 2007).
In 2002, natural and induced grasslands, as well as livestock areas covered 15% of the
territory; thus the remaining 41% of the land used for grazing is maintained in areas
covered with natural vegetation (Semarnat, 2008) from arid land, mountainous regions
and humid tropical areas, including wetlands. Only 14.7% is used for agriculture. This
indicates the importance of livestock production both as an economic activity and with
respect to the impact that the management practices associated with cattle ranching have
on the environment (Guevara and Moreno-Casasola, 2008). In the state of Veracruz, in
the Gulf of Mexico, where we have been developing research on the impact of cattle
ranching on wetlands, 43.2% of its surface is used for agriculture and 26.8% for
cattle ranching. This region has a broad coastal plain (39% of the territory is under 50
m.a.s.l.). The climate is humid tropical, bordered inland by a mountain range that filters
precipitation, which drains as subterranean water into the coastal plain. For the lowlands
of the coastal municipalities, an estimated 63.4% of this area is used for cattle ranching
and 15.7% for agriculture (Peresbarbosa Rojas, 2005). Wetlands currently occupy 70
476 ha in Veracruz (Figure 1).
Cattle ranching in Mexico began in the state of Veracruz, along the Gulf coast. When
the Spanish conquest began in 1519, Veracruz was occupied by the Totonacapan (sensu
B. Ortiz E. and R. Jiménez M., see chapter in this volume). Several authors (Heimo et
al., 2004; Beach et al., 2009) report that at that time, raised fields and canals in many
of the wetlands of lowland Mesoamerica were manmade and used to produce a variety
of crops. The fields bordered the wetlands and were subject to repeated, but shallow
flooding. The canals provided access, irrigation water, muck for fertilizer, and fish. It
was possible to have several crops per year by managing water levels. During the dry
season the lower, more humid parts were used; during the rainy season, the higher,
nonflooded parts were cultivated (Siemens, 1998). Even now, there are remains of the
elevated terraces that the indigenous people used for agriculture.
In the early 1500s, European settlers, mostly farmers, brought several breeds of Bos
taurus with them; cattle breeds that over four centuries became naturalized in tropical
Mexico (Guevara and Lira, 2006; Guevara and Moreno-Casasola, 2008). Some of these
breeds came from the Guadalquivir marshes in Spain (Velasco Toro and Skerritt, 2004).
In the terraces described above, it was possible to grow grasses year round, in the lower
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areas during the dry season and on the upper terraces during the rainy season. Cattle
could, therefore, be fed year round and could also browse in the forests, eating leaves
from lower branches, seedlings, etc. (personal communication Sergio Guevara). These
terraces are still being used for cattle ranching.
Map of the state of Veracruz in Mexico, and the distribution of its coastal wetlands. The
locations of the sites mentioned in the text appear on the map. The size of the watersheds
upon which the sites depend are shown.
Mapa del estado de Veracruz (México) y la distribución de sus humedales costeros. En el mapa
se indican las localidades de los sitios mencionados en el texto. Se muestra el tamaño de las
cuencas hidrológicas de las que dependen los sitios de muestreo.
Bovine livestock accounts for 40% of Mexico’s domestic meat production and, in
rural areas, is mainly used to obtain milk. Production is primarily extensive, low-tech,
and disease control is poor. Stocking density of grazing cattle varies from 0.8 ha/head
in the warm-humid tropics to 70 ha/head in the driest areas in the north, with a national
average of 3 ha; meat production is very low, ranging from 10 to 55 kg per hectare
(Toledo et al., 1993).
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From wetlands to pastures in the Gulf of Mexico
From 1950 on, particularly in the Gulf region and southeastern Mexico, agricultural
and livestock production expanded into the territories still covered by forests and
wetlands. This was accompanied by a change in livestock breed, with Zebu cattle
(Bos indicus) replacing the cattle that had been brought by the Spanish (Bos taurus)
(Guevara, 2001). Beginning in 1870, but especially in the first half of the twentieth
century, exotic grasses were introduced in Brazil and the Caribbean, and then Mexico,
and these replaced the native grasses. These species still persist and are favored by
government authorities and cattle ranchers when they have the means to purchase them.
With its bigger, heavier body, Zebu cattle needs open, high quality pastures. Tropical dry
and evergreen forests, and tropical oak woodland have been felled and anthropogenic
pastures have taken their place. Wetlands have also been used for raising Zebu cattle
(Guevara and Moreno-Casasola, 2008). Parsons (1972) and William and Baruch (2000)
recount the history of exotic grass introduction to the Americas. African grasses were
introduced to the continent in the 17th century, even before they were widely used
for grazing (Parsons, 1972). Different regions of Africa are the centers of production
of various forage grasses, and the flooded African savannas have numerous species
adapted to wetland conditions. Adaptation to foraging by these species is related to
their simultaneous evolution with ruminants in their areas of origin during the late
Pliocene and Pleistocene (Parsons, 1972; Matthews, 1982; Milchunas et al., 1988). As a
highly specific, frequent and intense disturbance, foraging can quickly alter the species
composition (Huston, 1994). One of the primary adaptations of grasses to foraging is the
ability to reproduce vegetatively by producing viable canes that disperse from the parent
plant. This is common in grasses from Africa and the Mediterranean region of Eurasia,
but is absent in many of the native grasses of America. As a result, the native grasses
of America have been largely displaced by invasive Old World grasses (Parson, 1972;
Huston, 1994).
Wetlands have been extensively used for raising cattle not only in Veracruz, but all
over Mexico and other parts of the Americas, such as Cuba (Caraballoso et al., 2011)
and in the Pantanal in Brazil (Junk and Nunes da Cunha, 2012). In some locations, cattle
have been introduced directly without modifying the plant species composition or the
flooding regime though they are moved to drier land during the months when the water
level increases. In some places the wetlands have been drained and in others, African
grasses, tolerant to flooding, have been introduced. Thus there are several levels of
wetland transformation.
The aim of this paper is to integrate and synthesize the information of numerous
studies that our research group has done on the impact of cattle herding on tropical
freshwater coastal wetlands, focusing on how the soil, the hydrological patterns and the
vegetation structure, composition and diversity are affected, as well as any alterations
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to wetland processes that have been documented. This analysis is based on experience
acquired in the analysis of wetlands used for raising cattle in Veracruz, Mexico, by
our research group. Some of these results have been published in Travieso-Bello et al.
(2005), who analyzed biodiversity and soil changes in the flooded pastures of La Mancha,
Veracruz, and management practices in the same region (Travieso-Bello and MorenoCasasola, 2011). López-Rosas (2007), López-Rosas et al. (2005 and 2006) analyzed the
degree of transformation and the type of impact that the introduction of the African grass
Echinochloa pyramidalis had on a freshwater marsh, and also ran several experimental
trials to eradicate the grass. López-Rosas and Moreno-Casasola (2012) analyzed the
results of a competition experiment using different levels of flooding between the grass
mentioned above and two native wetland hydrophytes: Sagittaria lancifolia and Typha
domingensis. Moreno-Casasola et al. (2010) analyzed the vegetation’s composition and
structure, as well as the water level fluctuations of thirteen freshwater marshes along the
coastal plain of Veracruz, some of them with either native or introduced grass species,
and grazing cattle. Rodríguez Medina (2011) and Rodríguez-Medina and MorenoCasasola (2013) studied the vegetation and soil properties of freshwater marshes in an
extensive wetland complex in southern Veracruz and compared areas where cattle had
been excluded with those with cattle. Moreno-Casasola et al. (submitted) have studied
the vegetation, water level fluctuations and soil properties of several flooded pastures
along the coastal plain of Veracruz.
Mitsch and Gosselink (2007) developed a conceptual model for describing
the fundamental role of hydrology in wetlands, which starts with climate and the
geomorphology of the basin. In this model, hydrology (water level, flow, frequency
of flooding, etc.) strongly interacts with the physical environment (sediments, soil
chemistry, water chemistry, etc.) and biota (vegetation, animals and microbes). Our
analysis of flooded pastures is based on this model, thus we will be discussing cattle
ranching impacts on hydrology, soils and vegetation.
Flooding and hydroperiod
Tropical Mexican freshwater wetlands can be dominated by either trees that form
freshwater swamps, or herbaceous species that form freshwater marshes (Olmsted,
1993; Moreno-Casasola et al., 2010; Infante Mata et al., 2011). They are found on
both mineral and organic soils. Wetlands differ not only in the dominant growth forms
and species composition, but also in their hydroperiod. This term defines water level
fluctuations, i.e. the seasonal pattern in the water level of a wetland, which is like a
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hydrological signature for each wetland type (Mitsch and Gosselink, 2007). Wetlands
are ecosystems whose functioning relies on hydrologic regimes and small variations in
flooding pulses or flooding levels may produce massive changes in the local biota.
Figure 2 shows examples of water fluctuation patterns for three sites, using data from
three types of wetlands (freshwater forested wetlands, marshes and flooded pastures)
present in the same area in Veracruz. The graph shows the flooding behavior for a period
of 18 to 24 months (October 2007 to November 2009). Zero represents the soil surface
and values above it denote flooding. Flooding behavior is site specific as is the type of
wetland, but in general pastures have more pronounced oscillations; that is, during the
dry season the phreatic level is lower. All of the wetlands remain flooded for part of the
year, except for pastures in Ciénaga del Fuerte. Water rises and saturates the soil where
roots are found, but there is no flooding; pastures at the other two sites are flooded for
several months of the year. The general picture is that marshes and pastures at the three
sites become flooded, thus from the hydrological perspective, these pastures behave
similarly to the other wetland types.
- 150
swamp EDulceS3
swamp EDulceS4
marsh EDulceP2
marsh EDulceP7
pasture EDulcep1
pasture EDulcep2
swamp FuerteS3
swamp FuerteS4
marsh FuerteP2
marsh FuerteP3
pasture Fuertep1
pasture Fuertep2
swamp RBlancoS1
swamp RBlancoS2
marsh RBlancoP1
marsh RBlancoP2
pasture Rlimónp1
pasture Rlimónp3
Hydroperiod over a year and a half to two years for three types of wetlands (swamps,
marshes and flooded pastures) in three sites (Ciénaga del Fuerte, Estero Dulce and Río
Blanco-Río Limón) in Veracruz, Mexico. Swamps are indicated with a black continuous
line; marshes with a gray continuous line and flooded pastures with a dashed line. Zero is
ground level. Zero is ground level and vertical axis is in centimeters. Horizontal lines at the
bottom of the figure indicate the rainy season.
Hidroperíodo de un año y medio a dos años para los tres tipos de humedales (arbóreos,
herbáceos y pastizales inundables) en tres sitios (Ciénaga del Fuerte, Estero Dulce y Río BlancoRío Limón) en Veracruz, México. Los humedales arbóreos se indican con una línea negra
contínua, los humedales herbáceos con línea gris contínua y los pastizales inundables con una
línea discontinua. El cero es el nivel del suelo y el eje vertical está en centímetros. Las líneas
horizontales en la parte inferior de la figura indican la época de lluvias.
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Both swamps and marshes are being transformed into pastures. In swamps, trees
are felled and grasses planted. One group of species is replaced by another and in the
new community, tree stumps resprout. In areas that remain flooded for longer periods
of time, small differences in topography allow for the presence of patches dominated by
marsh species. In marshes, even when exotic grass species are introduced, herbaceous
species persist, though with low cover values (López-Rosas et al., 2006). Figure 3 shows
a dendrogram comparing the vegetation from quadrats sampled in marshes and flooded
A matrix with 155 quadrats and 113 species from 23 sites sampled in Veracruz was
used. The data were taken from 12 herbaceous wetlands and 11 flooded pastures. A
cluster analysis was done (program PCOrd -McCune and Grace, 2002) using the flexible
β linkage method and relative Euclidian distance as a distance measure. A dendrogram
with seven floristic groups was formed, with 1.38 percent chaining. Figure 3 shows the
dendrogram, indicating the geographic location of the samples and the dominant species
in each group. Most samples from the same site fell into the same floristic group or were
with samples from only one or two other sites; only in group 6 are there samples from
several sites.
The first major division separates the samples from La Mancha in central
Veracruz from the rest. These samples form two subgroups. The first (1 A) is formed
of hydrophytes dominated by native broadleaf herbaceous plants that comprise a
community known as popal (described in Moreno-Casasola et al., 2010). It is located in
a reserve and has been under restoration so there is no influence by cattle. It is dominated
by Sagittaria lancifolia and Pontederia sagittata, among others. The second subgroup
(1 B), also in La Mancha, is dominated by the African grass Echinochloa pyramidalis
which has become an invasive wetland species, and Typha domingensis, a tall,
herbaceous monocot hydrophyte, locally called tule (a name also used when referring to
cattail), which is widely distributed. This site is separated from the others because of the
presence of an exotic that has become a wetland invader that has taken over most of the
wetland (López Rosas et al., 2005 and 2006). The other major group (indicated by the
number 2) brings together the other floristic groups.
This second big group is subdivided into various subgroups that are characterized
by wetland species, with cover by invasive species still low. Group 3 consists of several
subgroups and group 4 is comprised of samples from five sites from the Papaloapan
River Basin, located along the Río Limón and Río Blanco rivers. In this vast area of
herbaceous wetlands there are popales dominated by P. sagittata and Thalia geniculata
and tulares dominated by narrow leaved species such as T. domingensis, Cyperus
giganteus, Eleocharis cellulosa and Phragmites communis; all of which are being used
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for grazing livestock. Herbaceous wetlands from this area are rich in species and are
dominated by E. cellulosa (Cyperaceae) and Leersia hexandra (Poaceae), both of which
are palatable to livestock, the latter a typical native grass and common in wetlands.
Group 3 was divided into five subgroups. Group 5 was dominated by Cyperus giganteus
and the native wetland grass Hymenachne amplexicaulis, and to a lesser degree by Typha
domingensis, Limnocharis flava, and Thalia geniculata, among others. These species are
distributed in several places both in northern Veracruz (Estero Dulce, Laguna Grande
and Chica) and the Papaloapan (Sombrerete and Río Limón).
Dendrogram comparing the vegetation of marshes and flooded pastures. Data are from
López Rosas et al. (2005), Moreno-Casasola et al. (2010), and Rodríguez-Medina and
Moreno-Casasola (2013).
Dendrograma que compara la vegetación de los humedales herbáceos y los pastizales
inundables. Los datos provienen de López Rosas et al. (2005), Moreno-Casasola et al. (2010), y
Rodríguez-Medina y Moreno-Casasola (2013).
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Group 6 is dominated by Leersia hexandra, which is associated with several
species from the herbaceous wetlands, with few representative quadrats. It is a very
heterogeneous group present in several sites of the Papaloapan. Group 7 is dominated by
native species such as the grass Luziola peruviana. The herb Lippia dulcis (Verbenaceae)
dominated group 8 and only appeared in the north of Veracruz, in Ciénaga del Fuerte
and Estero Dulce. Finally, group 9 is dominated by L. peruviana, H. amplexicaulis and
C. giganteus. The dominant species of most of these groups indicates that wetland plant
cover at these sites is dominated by native species used by livestock. The most frequent
members of the Cyperaceae are Eleocharis cellulosa, Cyperus giganteus, Cyperus
articulatus and Fimbristylis spadicea.
The data were also analyzed with a Principal Component Analysis, and data were
modified with Beals smoothing technique to eliminate zero truncation problems (Beals,
1984), using the same program. Axis 1 and 2 of the ordination account for 38.51% of the
variation found. In Figure 4, samples from the different sites can be seen, using different
symbols to show the herbaceous wetlands in central and northern Veracruz, those in the
Papaloapan Basin, the flooded pastures of the central and northern regions and those
in the Papaloapan Basin. Axis 1 shows a regional gradient with both the herbaceous
wetlands and the flooded pastures of the Papaloapan communities (the largest wetlands
in the state of Veracruz) on the left. The communities located on the northern region
appear on the right of the ordination space. Axis 2 shows a gradient of herbaceous
wetlands along the upper part of the ordination space and flooded pastures at the bottom.
These gradients can also be interpreted as diversity gradients. Both marshes and flooded
pastures along the Papaloapan have higher diversity values (Shannon index: 1.949 and
1.250 respectively) than the herbaceous wetlands and flooded pastures in the central
and northern regions (1.191 and 0.632 respectively). Thus, both site and management
determine some of the properties of these wetlands. The Papaloapan River Basin
harbors the most extensive freshwater marshes in Veracruz allowing for a less impacted
hydroperiod, with fewer fluctuations over time, and this allows for the conservation
of wetland diversity. Cattle stocking rates are also low, with only one or two cows
per hectare, though there are cattle throughout the region. These species richness and
diversity values together with the area of the wetlands and the numerous aquatic bodies
in between the wetlands, probably do not favor the invasion of exotic species. Table
1 lists some of the characteristics of the native and exotic grass species found in the
flooded pastures of Veracruz.
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PCA ordination in which Axis 1 and 2 account for 38.51% of the variation. Wetlands in the
central and northern region of Veracruz are dark circles; wetlands in the Papaloapan Basin,
with very low cattle grazing impact, are open diamonds. Flooded pastures in the central
and northern region are open triangles and those located in the Papaloapan Basin are gray
squares. Along axis 1 there is a geographical gradient, with the samples from the Papaloapan
on the left side and samples from the central and northern region on the right. Axis 2 shows
a grazing gradient, which is clearest for the central and northern sites, where some wetlands
are not used for grazing. These appear toward the upper part of the ordination space. The
Papaloapan samples are mixed because there are sites with intensive grazing and others with
very low impact, but there are no sites which have not been grazed at all.
Ordenación por análisis de componentes principales (ACP) en donde los ejes 1 y 2 explican el
38.51% de la varianza. Los círculos representan a los humedales de la región centro y norte
de Veracruz; los rombos representan a los humedales de la Cuenca del Papaloapan, con muy
bajo impacto de pastoreo de ganado. Los triángulos representan a los pastizales inundables en
la región centro y norte; los cuadros representan a los pastizales inundables de la Cuenca del
Papaloapan. A lo largo de eje 1 hay un gradiente geográfico, en donde los sitios del Papaloapan
se encuentran del lado izquierdo y las de la región centro y norte en el derecho. El eje 2 muestra
un gradiente de pastoreo, el cual es más claro para los sitio del centro y norte, donde algunos
humedales no se utilizan para el pastoreo. Estos aparecen en la parte superior del espacio de
la ordenación. Las localidades del Papaloapan son variadas, ya que hay sitios con pastoreo
intensivo, otros con bajo impacto, sin embargo no hay sitios totalmente libres de pastoreo o que
no hayan sido pastoreados en algún momento.
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Characteristics of the more common native and introduced grass species found in Mexican
Características de las especies de gramíneas nativas e intruducidas más comunes que se
encuentran en los humedales de México.
Common name
English (Spanish)
Arundo donax Giant reed, giant
(carrizo, arundo,
Some ecological traits
Perennial, rhizome; sterile seeds, vegetative C31
growth appears to be well adapted to floods,
which may break up individual clumps,
spreading the pieces, which may sprout and
colonize further downstream. Is poisonous
to cattle and so cannot be used as forage.
Pará grass
Perennial, creeping stolons, stems reclining C46
mutica (Forsk.) (pasto Pará, zacate at base, rooting at the lower nodes, fodder
grass and also one of the worst weeds.
Allelopathic abilities allow it to form dense
monocultural stands. Reproduces and
spreads primarily by stem fragments; can
mutica (Forsk.)
form a stolon mat 1 m or more in depth;
sends up floating stems. Tolerates both
drought and brackish water .
Bermuda grass
Perennial, stolons, rhizomes, reproduces
dactylon (L.) (zacate Bermuda) by seed and vegetatively, widespread, good
animal fodder
Jungle rice
Annual, considered an agricultural weed,
colona (L.)
(arrocillo silvestre) good animal fodder
Barnyard grass
Perennial, rhizomes, reproduces
(pasto alemán,
vegetatively, very productive and “builds
(Lam.) Hitchc. zacate alemán)
soil” through biomass accumulation
& Chase
Annual, caespitose, erect or decumbent,
acuminata (J. cupgrass
sometimes rooting at the lower nodes
Presl) Kunth
Hymenachne Foxtail, West Indian Perennial, stolons, stems floating, creeping, C32
amplexicaulis marsh grass
or ascending to 1 m, rooting at the lower
(Rudge) Ness (azuche, cola de
nodes. Adapted to fluctuating water levels,
America and
zorra, trompetilla) which allow massive regeneration by seed
and ensure persistence after extensive
drought. Grows in water up to 2 m deep in
periodically inundated wetlands, but not in
permanent water
Perennial, forms large clumps, reproduces C41
Hyparrhenia Giant thatching
rufa (Nees)
grass, jaragua grass by seed; medium quality forage grass; fire
(pasto jaragua)
Blady or cogon
Perennial, rhizomes, spreads through seeds C41
cylindrica (L.) grass
and rhizomes, forms compact tufts
(alang alang, sujo)
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Common name
English (Spanish)
Swamp ricegrass,
southern cut grass
(pasto lamedor)
From wetlands to pastures in the Gulf of Mexico
Some ecological traits
Perennial, stolons and small rhizomes,
develops rooted, floating culms during
the rainy season, spreads vegetatively by
rhizomes and stolons and can also reproduce
from seed. Sometimes forms floating islands
and can grow in water up to 1.8 m deep;
animal fodder
Peruvian watergrass Perennial, stolons.
peruviana Juss. (engordador)
Ex J.F. Gmel.
Oryza latifolia Wild rice, broad
Perennial, caespitose, short rhizomes, erect C31
leaved rice
culms up to 2 m.
hexandra Sw.
(Forsk.) Stapf
P.J. Bergius
(syn. P.
south of
Mexico to
there are
grass, Egyptian
Egyptian panic
Water paspalum
Perennial, mat forming, elongated rhizomes, C44
rooting at nodes
Perennial, stolons, stems grow long and
sprawling, spongy and thick, frequently
found submersed or floating
elephant grass
(pasto elefante, p.
Perennial, bamboo-like clumps, spreads
by short rhizomes, rooting from lower
nodes or falling stems rooting at nodes
creating a stolon, reproduces vegetatively
or by seed, recovers from fire
Perennial, thick rhizomes, spread mainly
through vegetative means;
rhizome and stolon fragments, producing
dense mats, tolerates brackish water,
although a native it can become invasive
Perennial, rhizomes, forms pure colonies,
rhizome mass excludes all other
vegetation; reproduces only by seeds,
dispersed by wind, animals
Perennial, caespitose, no rhizomes,
tolerates saline conditions; sprouts are
used as animal fodder
Perennial, rhizomes, grows up to 4 meters
tall in dense bunches from large, creeping
rhizomes. Spreads via functional stolons
and vegetative buds that erupt from the
stems; tolerates small amount of salt in
free soil water
there is
Common reed
(Cav.) Trin. ex
(Trin.) Merr. Zizianopsis
(Michx.) Döll
& Asch. 197
Palm grass,
Buddha grass
Gulf cordgrass
Giant cut grass,
water millet
(1) Waller and Lewis (1979), (2) Medina and Motta (1990), (3) Field Guide to Texas Grasses. Robert B. Shaw, Institute
of Renewable Natural Resource, (4) Giraldo-Cañas (2010), (5) Ainouche et al. (2004), (6) Williams and Baruch (2000).
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Flooding as a plant stressor
Freshwater wetlands are stressful ecosystems for most plants. Flooding or water
saturation of the soil decreases the concentration of oxygen available to plant roots.
Under these reducing conditions there is an increase in soil microbial processes
that produce gases that are potentially toxic to the plant, such as sulfide or methane
(Mendelssohn et al., 1981; Ponnamperuma, 1984). Aquatic plants (hydrophytes) have
features that allow them to endure or avoid reducing (anaerobic) soil conditions. The
flood response varies with plant type, the duration and frequency of flooding, and the
flood water characteristics (Kozlowski, 1984). The tolerance of plants to flooding is
associated with the resistance of air movement through the vascular cambium, the
survival of secondary roots, the development of new secondary roots and adventitious
roots, accelerated anaerobic respiration, and the oxidation of the rhizosphere (Kludze
and DeLaune, 1996). Most hydrophytes develop aerenchyma in their leaves, stems and
roots. This tissue has a dual function: (1) transporting oxygen from the atmosphere to
the rhizosphere, and (2) diluting the toxic gases out of the plant cells (Crawford, 1987).
Plants adapted to flooding capture atmospheric oxygen through their photosynthetic
tissue and this oxygen is directed toward the aerenchyma from where it is spread to the
roots, creating an oxidized microenvironment around them. This process is beneficial to
plants because they oxidize reduced compounds such as iron and manganese ions, which
are abundant in flooded soils and are toxic to the roots (Kozlowski, 1984). As flooding is
a stressor to plants, the number of species in tropical wetlands is low compared to those
in terrestrial environments. Perhaps this is why the exotic hydrophytes are successful
invaders in these environments and produce such a negative impact. In the wetlands
of the Americas, invasive hydrophytes are one of the main causes, whether direct or
indirect, of negative effects. Many of these are reported in the literature, others have not
been reported but they are highly likely to be present. For example, Arundo donax (giant
reed) displaces the trees on the banks of rivers and lakes, reducing riparian diversity
(direct impact) and consequently decreases the shade on the water’s surface with the
result that temperature increases (indirect impact). Another example: invader grasses
such as Echinochloa pyramidalis, Brachiaria mutica, Pennisetum purpureum and
Phragmites australis are aggressive competitors that displace native vegetation (direct
impact). They are highly productive species with high water requirements through
evapotranspiration, which leads to reduced hydroperiods and accelerates the process
of succession (indirect impact) to communities of facultative hydrophytes or terrestrial
environments. Table 2 lists the main wetland invasive hydrophytes of the tropical and
sub-tropical Americas, and the effects that have been reported in the literature.
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From wetlands to pastures in the Gulf of Mexico
Wetland soils are a key element in the ecological services that wetlands provide to
society. Their physico-chemical characteristics are fundamental to water retention during
floods and in the storage of organic carbon.
The effect of livestock on wetland soils
Livestock provides an important source of income in a world that increasingly
requires more space to feed growing populations. In recent decades, wetlands have
been affected as the cattle frontier has expanded into mangroves and freshwater
wetlands where the animals can graze during the dry months when forage supply
decreases in other pastures (Skerrit, 1992; Moreno-Casasola, 2004). The few studies
that have examined the impact of cattle grazing on these ecosystems state that there is
some impact on vegetation, the growth of exotic grasses and small plants with short
life cycles is favored, and there is a reduction in the richness and abundance of species
with large leaves and thick rhizomes, which are characteristic of the native vegetation of
herbaceous wetlands (Travieso-Bello et al., 2005; Jones et al., 2011; Rodríguez Medina,
2011). The impact of livestock on the vegetation is also reflected in the diversity of the
fauna. Jansen and Healey (2003) have shown that if large leaves are not available in
wetlands, frog communities are markedly reduced. Furthermore, Jones et al. (2011)
mentioned that waterfowl breeding is adversely affected, mainly that of ducks, because
they are more likely to use the large leaves of the emergent vegetation as cover for the
nest and when trying to escape.
The type and quality of the vegetation depends partly on the soil, which is one of
the basic components of a wetland (Mitsch and Gosselink, 2007). It is critical because
that is where the stress is produced by oxygen limitation, which affects both the rate of
decomposition and nutrient availability (Úlehlová, 1998).
Currently, there are several studies that report the impact of livestock on the soil
of diverse ecosystems such as grasslands, savannas, and agricultural systems. These
studies report that trampling by livestock over short periods of time affects mainly the
first 15 cm of soil, and significantly increases bulk density and penetration resistance,
thus reducing infiltration and porosity (Lal, 1996), and affecting the development of
plant roots and their productivity (Pinzon and Amezquita, 1991). Studies on this topic in
wetlands are scarce. Travieso-Bello et al. (2005) reported that the values of C and N at
multiple sites under different livestock handling regimes are explained by a combination
of factors including changes in hydrology, the introduction of nonnative species, and
the presence of cattle. Higher values of C and N were found in seminatural wetlands
with little livestock management (with no draining or species introduction, etc.), and
the opposite happened in wetlands where the hydrology had been changed and the
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stocking rate was higher. They also found that soils had higher organic matter content
and retained more moisture when the stocking rate was lower. Rodríguez-Medina and
Moreno-Casasola (2013) evaluated the effect of livestock on the soil of four freshwater
herbaceous wetlands on the central Gulf coast of Mexico, and reported that where the
stocking rate was higher, soil bulk density was also higher and the amount of organic
material was low, reducing its total porosity. They also reported that trampling during
the rainy season affected the soil and made it more prone to compaction.
Some authors have briefly mentioned the impact of livestock on wetlands, indicating
that it affects biodiversity, induces changes in the balance of nutrients because cattle
dung introduces nutrients, and reduces the amount of organic matter and soil moisture,
among other effects (Coffin and Lauenroth, 1988; Archer and Smeins, 1991; Skerritt,
1992; Trettin et al., 1995; Collins et al., 1998; Baron et al., 2002). Each cow defecates
15 to 20 times per day and its dung can cover one square meter per day (De Elias, 2002).
Despite all of the negative impact on wetland soils caused by raising livestock,
Rodríguez-Medina and Moreno-Casasola (2013) mention that if livestock were excluded
from these sites during the flooding period for at least six months, and stocking rate were
maintained between one and two cows per hectare, the impact on the soil would not be
as severe, and important physico-chemical characteristics would be little affected in the
long term. Thus low intensity grazing would favor the preservation and maintenance of
tropical wetlands. Further work is needed to develop a system that ensures sustainable
livestock production with minimal degradation of soil resources (Tian et al., 1999).
Junk and Nunes da Cunha (2012) indicate that cattle ranching in the Pantanal in Brazil
maintains the dominant herbaceous wetlands and hampers shrub and tree growth.
In the following paragraphs we describe the changes that take place in the soils
and the tropical marsh vegetation associated with different management practices:
the introduction of cattle, fire, wetland drainage, and the introduction of exotic forage
species. The information was compiled from Travieso-Bello et al. (2005), López Rosas
et al. (2005), Escutia-Lara et al. (2009), Olsen et al. (2011), Wantzen et al. (2012),
Rodríguez-Medina and Moreno-Casasola (2013) and personal observations.
1. Cattle is introduced into the wetlands, no other management practice is applied
1.1. Hydrology
Flooding is kept similar to the natural regime.
1.2. Soil
Organic material (OM). When livestock is present, the layer with OM decreases due
to grazing and trampling; however, during the flooded months animals are removed and
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From wetlands to pastures in the Gulf of Mexico
grazing and trampling stops; the OM layer increases since the process of mineralization
decreases. Several wetland species tolerate grazing and resprout.
Water retention. Soils that have a layer with OM retain more water because they have
pores of various sizes, resulting from plant residue in various stages of decomposition.
These pores fill with water during flooding. Porosity can decrease with cattle trampling.
Bulk density (BD). BD increases because of animal trampling. The soil can regain
its porosity and part of its structure when livestock is excluded during flooding. This is
possible when the number of cattle is no higher than one or two per hectare.
Micro- and macronutrients. A long period of flooding reduces mineralization and the
amount of micro- and macronutrients (C, N, P, K, Mg, Ca, Na) increases.
pH. The soil remains slightly acidic (characteristic wetland soil) because oxidationreduction processes still occur.
1.3. Vegetation
The species appearing under these conditions are mainly native wetland species:
Sagittaria lancifolia, Pontederia sagittata, Thalia geniculata, Eleocharis cellulosa,
Cyperus articulatus, Hydrocotyle verticillata, Nymphaea ampla, Sporolobus virginicus,
Lippia nodiflora, Fuirena simplex, Typha domingensis, Ipomoea tiliacea, and Bacopa
monnieri. In more flooded areas Salvinia minima and Lemna minor are dominant. There
are few grass species.
2. Cattle is introduced into the wetlands and the hydrological conditions are
2.1. Hydrology
Flooding is reduced.
2.2. Soil
Organic material (OM). The layer of OM is reduced because grazing and trampling
by livestock occurs year round and directly affects the vegetation. Additionally, the small
amount of OM present decomposes very quickly in the absence of anaerobic conditions.
Water retention. In soils where the hydrology has been changed and flooding periods
have been significantly reduced, mineralization processes dominate, so there is little OM
and thus the soil’s water retention capacity is much lower.
Bulk density (BD). In the absence of OM, environmental factors such as wind and
rain can more easily erode the soil’s surface. If we add trampling to this scenario, the
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soil structure is modified as stable aggregates are destroyed and begin to clog the air
spaces. This produces an increase in BD and lower porosity.
Micro- and macronutrients. Micro- and macronutrients decrease because OM is
scarce; there are no long periods of flooding, and therefore mineralization is increased.
pH. The alkalinity at these sites is higher due to the decrease in the flooding periods
and lack of anaerobic conditions, which limits the processes that acidify the soil.
2.3. Vegetation
A few native wetland species are maintained, including indigenous wetland grasses
such as Hymenachne amplexicaulis. Exotic grass species used as forage appear or are
introduced: Echinochloa pyramidalis, Echinochloa colona. Other accompanying species
are Panicum sp., Paspalum sp., Hydrocotyle verticillata, and Typha domingensis. Other
species commonly found in disturbed areas or associated with human activities are
Cucumis anguria, Acacia cornigera, Ipomoea tiliacea, and Solanum campechiense.
3. Cattle is introduced into the wetlands and the vegetation is burned annually
3.1. Hydrology
Flooding is kept similar to the natural regime.
3.2. Soil
Organic matter, micro- and macronutrients. In general, OM, soil nutrients and water
retention decreases with the presence of livestock (see above 1.2 and 2.2) and can further
increase with burning. There are reports stating that if both management activities
are low impact (few cattle, and removing cattle during the period of high floods), the
productivity of a site may increase, and may in fact favor the wetland species and control
the spread of exotic and invasive species (López Rosas, 2007; Escutia-Lara et al., 2009;
Rodríguez Medina, 2011).
Bulk density. It has been mentioned that livestock increases the value of BD, but
when the presence of livestock is combined with a low-intensity burning, this may
favor the growth of wetland species. The latter help decrease BD, because the native
hydrophytes of these ecosystems have many, much longer and thicker roots (due to
aerenchyma), and these increase air space and soil porosity (Davey et al., 2011).
3.3. Vegetation
Native wetland grasses and sedges that tolerate grazing are present, such as
Sporolobus virginicus and Hymenachne amplexicaulis, Eleocharis cellulosa, Cyperus
articulatus, and Fuirena simplex. Other hydrophytes are also maintained: Sagittaria
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lancifolia, Pontederia sagittata, Thalia geniculata, Hydrocotyle verticillata, Nymphaea
ampla, Lippia nodiflora, Typha domingensis, and Bacopa monieri. In more flooded areas
Salvinia minima and Lemna minor are dominant. The exotic grass Echinochloa colona
is present. Species associated with disturbance or human activities are Cucumis anguria,
Acacia cornigera, Ipomoea tiliacea, and Solanum campechiense.
4. Cattle is introduced into the wetlands, vegetation is burned each year, and flooding time is reduced, sometimes with the introduction of exotic forage species
4.1. Hydrology
Flooding is reduced
4.2. Soil
The impact of livestock in the areas where flooding has been reduced by changes to
the hydrology is high, and if the vegetation is burned this management practice may be
more harmful to the soil, mainly because the conditions are no longer suitable (mainly
the hydrology) for the growth of native wetland species. Fires cause increased soil
erosion. The low-impact fires in wetlands that have not been strongly transformed may
benefit the growth of native vegetation because temperature in soil is not enough to kill
seeds and propagules of hydrophytes; then the gaps can be revegetated with natives (Lin
et al., 2005). Intense fires are common when the flooding is reduced; those fires kill
seeds and propagules in soil surface, then only propagules of resistant species, such as
the invader grasses, survive and dominate new gaps (Lin et al., 2005) and the process of
invasion is reinforced.
4.3. Vegetation
Species richness decreases and few native wetlands species remain: Hydrocotyle
bonariensis, Fimbristylis spadicea, Cyperus articulatus. Grass species are favored as
well as those associated with human activities: Echinochloa pyramidalis, Echinochloa
colona, Panicum sp., Paspalum sp., Cucumis anguria, Solanum campechiense, Ipomoea
tiliacea, Acacia cornigera, and Mimosa pigra.
The use of wetlands for cattle grazing brings about important environmental changes.
The number of cows allowed to graze per hectare is closely related to the degree of the
impact, thus forming a gradient from wetlands to flooded pastures, with different species
composition and richness, soil characteristics and hydrology. These flooded pastures
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can be considered wetlands, though when they are invaded by exotic species, or when
their hydrology is altered, these transformations impair their functioning, and there is
a tendency to lose environmental services and for them to function more like terrestrial
systems (Table 2).
Impact of exotic hydrophytes on tropical and subtropical American wetlands.
Impacto de hidrófitas exóticas sobre humedales de América tropical y subtropical.
Type of
Hygrophila polysperma
(Roxb.) T. Anderson
1, 4, 5
Sutton, 1995; Mora-Olivo et al., 2008
1-4, 8
Gordon, 1998
Pistia stratiotes L.
Mimosa pigra L.
1-5, 7, 13- Labrada et al., 1996; Gordon, 1998;
15, 20, 22 Rejmánek et al., 2005
1-3, 5
Pueraria montana var.
lobata (Willd.) Maesen &
S.M. Almeida ex Sanjappa
& Predeep
Gordon, 1998; Rejmánek et al., 2005
Iris pseudacorus L.
1, 6
Pathikonda et al., 2008
Melaleuca quinquenervia
(Cav.) S.T. Blake
1-4, 6, 7,
Arundo donax L.
Gordon, 1998; Mack et al., 2000;
Rejmánek et al., 2005; Zedler and Kercher
Brachiaria mutica
(Forssk.) Stapf
Hydrocharitaceae Hydrilla verticillata (L. f.) 1, 2, 4, 5,
8, 13, 14,
16, 17
Lythrum salicaria L.
1-4, 14, 17 Blossey et al., 2001; Brown et al., 2006;
Lavoie, 2010; Rejmánek et al., 2005;
Zedler and Kercher, 2004
1, 3-5, 7, Guthre, 2007; Flores Maldonado et al.,
11, 12, 15, 2008; Comité Asesor Nacional sobre
18, 21
Especies Invasoras, 2010; Rejmanek et al.,
2005; Yang et al.; 2011
1, 3-5
Echinochloa pyramidalis
(Lam.) Hitchc. & Chase
1, 5, 6
Imperata cylindrica (L.)
7, 18, 20
Pennisetum purpureum
Comité Asesor Nacional sobre Especies
Invasoras, 2010; Gordon, 1998; Sousa,
2011; Langeland, 1996
1, 4, 5, 7,
13, 14
D’Antonio and Vitousek, 1992; Parsons,
López Rosas, 2007; López Rosas et al.,
2005; López Rosas and Moreno-Casasola,
Labrada et al., 1996
Williams and Baruch, 2000; Cronk and
Fuller, 1995; Laegaard and Pozo Garcia,
2004; Schardt and Schmitz, 1991
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Type of
Phragmites australis
(Cav.) Trin. ex Steud.
1, 2, 8, 16
Zedler and Kercher, 2004
Eichhornia crassipes
(Mart.) Solms
1-6, 8-10,
Salvinia molesta D.S.
Barret, 1989; Labrada et al., 1996; Comité
Asesor Nacional sobre especies invasoras,
2010; Gordon, 1998; Mack et al., 2000;
Rejmánek et al., 2005
2, 5, 9, 10, Berret, 1989; Labrada et al., 1996;
13, 14, 19 Rejmánek et al., 2005
Vochysia divergens Pohl
Tamarix ramosissima
1-3, 6
Rejmánek et al., 2005; Zedler and Kercher,
Vourlitis et al., 2011; Sanches et al., 2011
Environmental impact: (1) decrease of biodiversity, (2) changes in the chemical composition of soil or water,
(3) alteration of hydrology (e.g. excessive water loss through evapotranspiration), (4) obstruction of flow water
(stagnation), (5) intercepts light, increasing shade at soil level, (6) vertical soil accretion, (7) altered fire regime, (8)
altered food webs, (9) excessive use of oxygen, (10) reduction of dissolved oxygen, (11) increase in the temperature
of rivers and water bodies (by reduced shade of trees), (12) erosion of river borders. Social impact: (13) obstruction
of navigation channels, (14) obstruction of waterways or hydroelectric plants, (15) accelerated siltation of reservoirs
and irrigation canals, (16) reduction in fisheries productivity, (17) reduction of recreational activities, (18) increase
in habitat for vectors of human or livestock disease (malaria, dengue fever, filariasis, encephalitis, schistosomiasis,
etc.), (19) interferes with the operation of waterworks, (20) aquatic weed in crops (e.g. rice), (21) damage to social
infrastructure (bridges, pipes, etc.), (22) interferes with the movement of people and livestock.
Figure 5 synthesizes in a diagram the information presented in this paper with the
changes in the hydrology, soils and vegetation occurring under two conditions. The first
occurs when flooded pastures originate from swamps (freshwater forested wetlands), and
the second when marshes undergo the transformation. Flooding can either be maintained
or reduced. In the first case flooding only remains aboveground for a few months and
in the second case it can either be maintained (as occurs on floodplains with extensive
wetlands, i.e. Río Blanco in the Papaloapan River basin- Figure 2) or reduced as a result
of draining or the introduction of exotic species. Soil properties change, although this
depends strongly on the original soil type, i.e. organic or mineral. Vegetation also varies
as shown in the classification and ordination figures.
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Diagram showing changes in the hydrology, soil and vegetation in flooded pastures
originating from freshwater forested swamps and from marshes.
Diagrama que muestra los cambios en la hidrología, suelo y vegetación en los pastizales
inundables que se originaron a partir de humedales arbóreos de agua dulce y de humedales
herbáceos de agua dulce.
Some grass species introduced from the African flooded savannas can tolerate
flooding and compete with native wetland plants (López-Rosas and Moreno-Casasola,
2012). Wetlands are particularly vulnerable to invasion processes, where variations in
hydrologic regimes may cause changes in community composition and structure and
are considered one of the causes that make the incorporation of alien species possible
(Kalesnik and Malvárez, 2003). Currently, the most widely distributed grass on the
coastal plain of Veracruz is the African grass Cynodon dactylon, which tolerates both
dry and wet conditions (Travieso-Bello, 2005), but not prolonged periods of flooding. In
wetlands, Echinochloa pyramidalis is preferred by cattle ranchers, because it “dries the
area and builds soil” (Melgarejo-Vivanco, 1980), and because of its high productivity
(Andrade et al., 2008; Braga et al., 2008). When exotic species are introduced into
wetlands, transformations increase. Some of them are able to modify the hydrology of
the particular wetland type, thus initiating a more drastic change in wetland functions.
Figure 6 shows how wetland functions, processes and values are affected by the
transformation of herbaceous wetlands into flooded pastures. The following discussion
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does not apply to swamps that are cut down, because felling trees is, in itself, a major
transformation. The degree of transformation of freshwater marshes varies with cattle
grazing intensity. Arrows indicate permanence or decrease. Impact is grouped according
to the type of environmental and socio-economic impact, based on Table 2 and on the
data presented in this paper. The first type of environmental impact is the alteration
of wetland structure and interactions, which includes the decrease in biodiversity and
the increase in the presence and cover of invasive species (López Rosas, 2007). When
exotic species are introduced to allow for more intensive grazing, and the biodiversity
decreases, the dominance of a few species is promoted, shading reduces habitat for sun
loving wetland species, there are changes to the soil physico-chemical characteristics,
and flooding is reduced. Changes in species composition and community structure affect
the wetland regulatory functions, habitat functions, production functions and information
functions (sensu De Groot et al., 2002), thus important ecosystem services decrease. For
example, species composition and structure are related to carbon sequestration and water
regulation (Campos et al., 2011).
Diagram showing the functions and processes affected by the transformation of wetlands
into pastures as a result of cattle ranching.
Diagrama que muestra las funciones y procesos afectados por la transformación de los
humedales en pastizales para ganadería extensiva.
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The second type of impact involves changes in the physico-chemical composition of
the environment, which in the case of cattle grazing mainly involves alterations to the
chemical composition of either the soil or the water, an increase in the temperature of
rivers and water bodies (because of reduced tree shade), the interception of light which
increases shade at soil level, and soil compaction produced by trampling. Among the
main changes in the soil is an increase in bulk density, reduced pore space, and lower
water retention. The organic layer is reduced, there is less aeration, pH becomes neutral,
nutrient content decreases, and carbon storage potential is reduced, as is the soil’s
capacity to store water. These alterations will affect species composition. Important
regulatory functions are affected as soils lose many characteristics and become similar
to terrestrial soils. Supporting and regulatory ecosystem services decrease. Changes in
the soils, one of the fundamental elements of wetlands, has a profound effect on plant
composition, thus altering provisioning services.
The third is the alteration of the wetland hydrological regime, which includes the
alteration of hydrology, and can result from one or more of the following: excessive
water loss through evapotranspiration, obstruction of water flow, changes in topography
due to soil accretion, or an increase in sedimentation. Soils remain drier for a
longer period of time and organic matter is lost, thus altering the physico-chemical
characteristics of the soils. All types of wetland functions are altered as hydrology is the
main driver of hydric soil processes and wetland vegetation. There is a general reduction
in ecosystem services.
The fourth impact is the alteration of sediment dynamics, which includes vertical
soil accretion and the erosion of the river banks. These changes end up altering the
hydrological regime, which has the strongest negative influence on wetlands. The main
impact is on supporting and regulatory services. Finally, the fifth is the alteration of the
disturbance regime, which impacts species composition.
The main type of social and economic impact includes the alteration of income
generating activities that rely on waterways when navigation channels are blocked,
changes to waterways or the construction of hydroelectric plants, the operation of
waterworks, decreased opportunities for recreational activities, and the obstruction of the
movement of people and livestock. A second type is the damage or functional reduction
of infrastructure, which results in the premature siltation of reservoirs and irrigation
canals and damage to social infrastructure (bridges, pipes, etc.). A third type of impact
includes a decrease in crop production (i.e. rice), invasion by species that reduce the
quality of flooded pastures for cattle grazing (Junk and Nunes da Cunha, 2012), and an
increase in the habitat available for the vectors of human and livestock diseases. Local
and regional economies are affected, and the final result is a reduction of human well
being (sensu Millennium Ecosystem Assessment, 2005).
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Wetlands that are used for grazing produce milk and meat, and therefore make an
important contribution to the family income. Milk in particular, sometimes transformed
into dairy products, is part of the everyday diet. Meat (or the live cow) represents capital,
or a means of having some savings for difficult times. Considering the myriad benefits
provided by wetlands, their conservation and use should be analyzed taking into account
the degree of transformation. When wetland ecosystem services decrease or are lost, the
price paid is very, very high. When cattle grazing does not alter ecosystem functions,
it represents a sustainable use of resources that promotes human welfare. This use is
probably linked to wetland size, and according to Junk and Nunes da Cunha (2012) can
be achieved “…through modern management plans that reconcile the requirements of
environmental protection with the economic needs of the ranchers, who are the owners
of most of the Pantanal. The key to any such plan’s successful implementation is to
consider the Pantanal not as a pristine wetland, but as a valuable cultural landscape”.
Over the last two and a half centuries, the vegetation over large parts of the Pantanal
has been altered due to the presence of cattle ranchers, as it has over most of the tropical
wetlands in the Americas. These authors indicate that cattle ranching has maintained the
environment’s habitat diversity and the multiple services provided to humans and to the
environment, including the enhancement of species diversity. Our results indicate that
cattle ranching in wetlands with a low density of cattle (one head per hectare) maintains
wetland species, hydrological and soil conditions. This was probably the situation when
cattle was first introduced to Mexico. The cattle introduced by the Spanish, Bos taurus,
were small animals, much lighter than Zebu and with smaller hoofs. They occupied
extensive areas of wetlands with low densities (Siemens, 1998), thus their impact was
moderate, maintaining the functions and environmental services provided by wetlands
while constituting a sustainable economic activity that permited these ecosystems to be
High density livestock rearing or modifying the hydrology reduces plant diversity
and results in a loss of wetland functions and services, such as water holding
capacity being reduced by soil compaction. This is aggravated by the presence of C4
invasive species that make more efficient use of water than the native C3 species, can
photosynthesize more efficiently in high temperatures and are very productive, causing
soil accretion and increased shade at the soil level.
Government policies should thus promote the conservation of wetland functions
and services, coupled with sustainable cattle ranching. This implies setting limits
to the number of head of cattle per hectare in wetlands, rotating cattle so that during
the flooding peak the wetland soil and plants can recover, banning wetland drainage
practices and the introduction of African grass species. The sustainable practices
mentioned above should be promoted in areas where the wetlands are not in a good
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state of conservation, and that are dedicated to cattle production, thus promoting better
management practices for these areas, and ensuring that both wetland processes and the
economic activities that are associated with them are maintained.
The funding for the various papers used in this article came from ITTO PD
349/05 Rev.2 (F), ITTO-RED-PD 045/11 Rev.2 (M) and Humedales del Papaloapan
CONACYT-CONAGUA (48247). Roberto Monroy made the drawings. We thank all the
local people who helped us in the field, and the cattle owners and land managers for their
generous assistance, which allowed us to carry out the research and served as the basis
for this paper.
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La cría de ganado en México se inició con la llegada de los españoles y la creación
de la Nueva España. Se modificó de manera significativa en la mitad del siglo XX con
la introducción de la raza de ganado cebú y los pastos mejorados que esta ganadería
requería. Los humedales se usaron para el pastoreo de ganado desde el inicio de esta
actividad. El trabajo describe las transformaciones que se producen en las zonas
inundables usadas para pastoreo y como se van convirtiendo en pastizales inundados.
El grado de impacto depende de la cantidad de cabezas de ganado, el tiempo que
permanecen en el humedal, y las modificaciones al hidroperíodo y a la vegetación.
Se describen los cambios en el nivel de la inundación, las características del suelo
(materia orgánica, retención de agua, densidad aparente, pH, micro y macro nutrientes
) y la composición florística, y cómo todo esto afecta a los servicios ambientales que
proporcionan los humedales. Con la introducción de razas tolerantes a ambientes
tropicales, principalmente el ganado Cebú y las gramíneas forrajeras exóticas que pueden
crecer en zonas inundables, el impacto ha aumentado. Estos pastos alteran drásticamente
el medio ambiente (agua, suelo vegetación nativa) y pueden convertirse en invasoras.
Por lo tanto hay un gradiente de transformación de los humedales sin impacto del
ganado, a aquellos con ligeros cambios que siguen funcionando como humedales, hasta
finalmente transformarse y perder sus funciones. Una gestión basada en un bajo número
de cabezas de ganado mantiene las funciones y servicios ambientales que proporcionan
los humedales al mismo tiempo que constituye una actividad económica sostenible, que
permite que estos ecosistemas se conserven.
Palabras clave: Acreción vertical, compactación, invasión, pastoreo, sostenibilidad.
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Instituto de Ecología, A.C.. Red de Ecología Funcional. Carretera antigua a Coatepec 351. El Haya. Xalapa 91070. Veracruz
(México). *Author for correspondence: [email protected]
The current landscape of the Mesoamerican rain forest has been shaped by the
migration of plant and animal species, natural events, and human intervention. This
current reflection on the effects of livestock ranching in this region focuses on the
impact to both the structure and the functioning of the landscape. Los Tuxtlas, Veracruz,
Mexico, is both a case study and a reflection of the larger Mesoamerican landscape,
where cattle livestock were first introduced in the 16th century and subsequently replaced
with a new variety of cattle in the 19th century.
Thus, Los Tuxtlas is an enclave of tropical rain forest in southeastern Mexico and
illustrative of the environmental history of Mesoamerica as a whole, representing
a landscape of flora and fauna that have resisted climatic changes, intense volcanic
activity, and frequent hurricanes and tropical storms that have continuously occurred
across time. More recently, human activities, such as hunting, gathering, and agriculture,
have also formed the landscape beginning with the Olmecs, considered to be the mother
culture of Mesoamerica.
In the 16th century the European colonizers introduced on the northern tip of the
volcanic range of San Martin in Los Tuxtlas several cattle varieties of Bos taurus,
imported from the region of Guadalquivir, Spain. These cattle rapidly adapted to
free-range grazing, probably due to the absence of large native herbivores that
had disappeared in Mesoamerica by the end of the Holocene, as well as due to the
considerable extensions of secondary vegetation of abandoned agricultural fields,
resulting from the disappearance of the indigenous farmers after the arrival of the
The cattle grazed on “grama” grasses, which proliferated during this time and
emerged as part of the process of secondary succession, in addition to other plants and
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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shrubs found within the rain forest. The livestock were adapted to this form of grazing
for over 500 years, dispersing trees and bushes and enriching the process of secondary
In the 20th century, Bos taurus was substituted for Bos indicus, a variety of cattle
coming from Asia and raised only in pasturelands sown with African grasses, resulting
in a lack of herbivorous activity compatible with the local ecosystems, as in the case
of Bos taurus. A resulting defaunation and deterioration of the secondary vegetation
has since occurred, and many “grama” grasses have since disappeared. In addition,
the management practices associated with Bos indicus have stimulated an increase
in deforestation and fragmentation of the rain forest, signifying a threat to regional
However, numerous individuals and species of rain forest trees still persist in open
fields across the landscape, mitigating, as they have throughout history, the impact of
deforestation. The trees that are left behind are an element of the historical landscape
management practices, implemented by the very first inhabitants of the region. These
isolated trees form in many instances “living fences” that help to maintain the ecological
connectivity between the remaining fragments of vegetation, in addition to sustaining
processes of secondary succession and regeneration of the rain forest.
The main objective of this essay is to demonstrate that by understanding the
environmental history of the rain forest landscape of Mesoamerica, new possibilities
may be presented for its design and management with the explicit goals of conserving
biodiversity, preserving environmental services, and innovating productive and
agricultural systems. Our objective is illuminated by examining historical landscape
practices surrounding the introduction of cattle livestock in the region of Los Tuxtlas in
southeastern Mexico and considering the relevance that they may hold for contemporary
landscape management and the natural regeneration of the rain forest.
Key words: Veracruz, secondary succession, defaunation, landscape connectivity.
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The landscape is continuously shaped by natural events and human activities,
reflected in changes in climate and soil conditions, as well as movements of flora and
fauna. Every landscape has a history that may be interpreted by examining its spatial
array, structure, and the relationship between its elements, including its ecological
processes. Understanding the environmental history of a landscape involves an
awareness of the context and region in which it is situated, as well as a consideration
of the state of surrounding ecosystems and their biodiversity. This essay is concerned
with the landscape of Mesoamerica, an extensive region that includes the southern
half of Mexico and Central America, as well as Guatemala, El Salvador, Belize, and
the western parts of Honduras, Nicaragua, and Costa Rica. The area is delimited by the
shared cultural elements of the Mesoamerican peoples that inhabited the region prior
to the arrival of the Spanish, and whom in many cases continue to carry out traditional
landscape management practices (Kirchoff, 1960).
Mesoamerica is one of the most fascinating regions to study of the environmental
history of the landscape, due to its rich cultural and physical background. Within this
history we may discover details that are applicable and useful for the contemporary
landscape management of the rain forest. The region has been extensively disturbed
by natural events, including the displacement of the continental and insular masses that
gave form to the American continent and the Caribbean and extreme climatic changes
during the Glacial period, in addition to other perturbations caused by hurricanes and
frequent tropical storms. Various human groups have settled across the territory from
a very early date, forming the Mesoamerican civilization, a culture that developed in
distinct ecogeographic regions and that held a relatively high population density over
an extended period of time. Afterwards, Mesoamerica was witness to one of the first
experiments of European colonization, followed by the subsequent industrial and
rural development of Mexico in the 20th century as an independent nation. Thus, the
Mesoamerican landscapes owe in large part their structure and function to the productive
activities carried out by humans during these distinct geo-political scenarios (Guevara et
al., 2006; Guevara, 2011).
Among the abundant ecosystems that thrive in Mesoamerica, the rain forest is one of
the most alluring, due to its attractive aspect and its abundance and richness of species.
The term “tropical forests” integrates several forest formations, including both dense and
wet tropical forest as well as those that are dry and sparse (Figure 1). The rain forest has
always attracted human settlements owing to the large quantity of fauna and flora and
the apparent fertility of the soil. Such was true throughout the Spanish colonial period
and well after the independence of Mexico and remains true presently.
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The variation in the height of the vegetation and the seasonality of the rain forest is related
to the quantity of precipitation and the humidity of the soil.
La variación de la altura de la vegetación y la estacionalidad de la selva está relacionada con la
cantidad de precipitación y humedad del suelo.
The rain forest of this region groups together distinct types of forest that grow in
low altitudes of 700 meters (m) below sea level, with an annual precipitation of greater
than 2000 millimeters (mm) and an average annual temperature above 20° Celsius
(C) (Rzedowski, 1978). Another characteristic of the regional rainforest is its dense,
evergreen canopy of 30 m or more in height (Richards, 1952; Bongers et al., 1988;
Figure 2). The northernmost neotropical rain forest has its boreal limit in Mesoamerica,
precisely at the region of Los Tuxtlas (Dirzo and Miranda, 1991). Originally, the
Mesoamerican landscape was formed by a continuous mass of forested vegetation,
covering in Mexico an area of approximately 22 000 square kilometers (km²) and
stretching across Central America (Rzedowski, 1978).
The rain forest as seen from outside the rain forest. The profile of a rain forest fragment
(Drawing by Manuel Escamilla).
La selva desde fuera de la selva. Aspecto de un fragmento de la selva (Dibujado por Manuel
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However, humans have long existed in this forested landscape, intensively and
extensively managing the land and taking advantage of its natural resources for more
than 5000 years. Many of their traditional management practices are still in use today,
providing a certain kind of continuity between the past and the present. Human activities
incorporated agriculture, domestic animals, and biodiversity within a management
framework compatible with their environment and the surrounding ecosystems, a
similarity found across Mesoamerica in both tropical and temperature regions alike
(Guevara, 2011).
Thus past climatic and geological natural disturbances, pre-Hispanic human
activities, and the introduction of livestock during Spanish colonial times strengthened
natural regenerative processes of the rain forest by facilitating the ecological
connectivity and movement of biodiversity across the landscape. This may explain why
today, in the mountain range of Los Tuxtlas, a high biological diversity is maintained
(González-Soriano et al., 1997) in spite of the rampant deforestation that has reduced
the rain forest to several small fragments dispersed across extensive, open pastures and
agricultural fields (Guevara, 2011; Laborde et al., 2011).
This recent and devastating deforestation of Los Tuxtlas and the entire American
continent is a well-documented fact, and perhaps even more troubling considering that
the magnitude and rate of deforestation in both Mexican and Central American rain
forests in one of the highest in the world (Toledo, 1982). By the end of the 1980s, rain
forest cover in Mexico had been reduced to less than 10% of its original extension,
comprising 1 500 000 hectares (ha). The rest of the original cover had been transformed
in pastures, agricultural fields, and secondary vegetation (Rzedowski and Equihua, 1987;
Laborde et al., 2011).
The main objective of this essay is to demonstrate that by understanding the
environmental history of the rain forest landscape of Mesoamerica new possibilities
may be presented for its design and management with the explicit goals of conserving
biodiversity, preserving environmental services, and innovating productive and
agricultural systems. Our objective is illuminated by examining historical landscape
practices surrounding the introduction of cattle livestock in the region of Los Tuxtlas in
southeastern Mexico and considering the relevance that they may hold for contemporary
landscape management and the natural regeneration of the rain forest.
Los Tuxtlas is complex both environmentally and geologically (Figure 3). The region
stands out due to its long history of human settlement, in addition to being the seat of
the Olmec culture. It has also been isolated by its geography, nestled within the coastal
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plains of the Gulf of Mexico along a volcanic belt 95 km long and 50 km wide, between
18° 05’ and 18° 43’ N and 94° 35’ and 95° 25’ W. The area is characterized by two
large volcanic masses, the volcano of San Martin Tuxtla (1,650 m above sea level) to
the northwest and Santa Marta (1700 m above sea level) to the southeast, along with
several associated cinder cones (Mayer, 1962; Geissert, 2006). The region was formed
by volcanic activity during the Miocene, Pleistocene, and Holocene epochs, and remains
volcanically active today with a temporal sequence of volcanic activity dating back
more than 50 000 years. At least 10 volcanic eruptions have taken place in the past
5300 years, the two most recent occurring in 1793 and 1859. Volcanic ash and sediment
deposits have maintained the fertility of the soil, explaining the desirability of the zone
and its high population density (Santley, 2007). The climate is warm and tropical, with
an average annual temperature of 20° C and a minimum temperature of 18° C.
The rain forest of Los Tuxtlas is located in the southeastern portion of Mexico along the
Gulf Coast, in the once metropolitan area of the Olmec culture.
La Sierra de los Tuxtlas está en el sureste de México sobre la costa del Golfo de México, en el
área metropolitana de la cultura Olmeca.
During the Pleistocene, the mountain range of Los Tuxtlas was a secondary
refuge for the flora and fauna of the rain forest, forming a favorable climate due to
the geomorphology of the zone and the resulting humidity that it trapped due to its
positioning along the Gulf of Mexico. Species flourished in Los Tuxtlas during this
period of scarce precipitation and extreme temperatures (Toledo, 1982).
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The most critical time for the rain forest of Los Tuxtlas was during a period of low
temperatures and humidity that prolonged from 20 000 to 12 000 years ago, during
which the rain forest virtually disappeared from the region. Thus, the rain forest
landscape that is currently found is relatively recent in geological terms (Graham,
1975). The subsequent re-colonization of the rain forest was only interrupted by a dry
and hot period, during which the climate was probably more seasonal in nature than in
actuality, lasting from 9000 years ago up until 2000 years ago. After this period a rapid
re-establishment of evergreen species characteristic of the present rain forest began to
take hold.
Another important climatic event of relevance to Mesoamerica and the geological
landscape is that of the Ice Age, which began over 110 000 years ago and reached
its maximum around 20 000 years ago. Low temperatures caused the extinction of
the megafauna across the continent and led to the disappearance of over 80 species
previously encountered in the region, including the large herbivores, 7 species of sloths,
5 species of armadillos, 5 species or horses, 6 species of tapirs, 27 species of deer, 4
species of mammoths and mastodons, and several more species of camels and buffalo
(Galindo, 2012).
Human intervention in the rain forests of Mesoamerica began over more than 5000
years ago with an intensity unmatched in all of the Americas. Two of the most prominent
cultures of the continent flourished in this region, the Olmecs (VanDerwarker, 2006) and
the Mayans (Stuart, 1993). Nomadic agriculture and slash and burn farming, the oldest
and most extensive forms of land management until recently, were the most frequently
employed agricultural methods in the humid tropics of Mexico and Central America
(Rojas, 1994).
Anthropogenic activities across the landscape may be divided into three phases.
During the first phase (8000 B.C. to 1521 A. C.), the landscape was modeled by hunting
and gathering, as well as intensive and extensive agriculture. This period is marked
by the settling of the territory by the first peoples who arrived to the zone up until the
arrival of the Europeans. Another distinguishing characteristic of this time span is that
hunting and gathering, as well as agriculture, were compatible with the natural processes
of secondary succession. Thus, the management and manipulation of secondary
succession may very well be considered the basis of the Mesoamerican management of
the landscape.
The second phase (1521 to 1900 A. C.) is characterized by the introduction of
livestock and in particular cattle, as different varieties of the species Bos taurus,
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originating in the lowlands of the region of Guadalquivir, Spain, were allowed to
roam free across the American landscape (Jordan, 1993). At the beginning of this time
period in the 16th century, cattle were introduced to Los Tuxtlas atop the northern edge
of the volcanic mass of San Martin (Guevara and Lira-Noriega, 2004). The livestock
were rapidly adapted to free-range grazing, probably due to the absence of large, native
herbivores that had disappeared by the end of the Holocene (Janzen, 1984), in addition
to the large extensions of secondary vegetation that covered the agricultural fields that
were abandoned by the indigenous population. The cattle grazed and trampled in the rain
forest and secondary vegetation, during 500 years, allowing for the dispersion of seeds,
trees, and plants, thereby richening the process of secondary succession. One example of
this is the proliferation of native grasses, or “grama,” upon which the cows grazed.
The third phase (1900 to present) is marked by the substitution of B. taurus for B.
indicus, a livestock variety that in America is only raised in pastures planted with African
grasses. As a consequence, the presence of herbivores, diminished. The dissemination
of tree and bush species decreased, once aided by the herbivorous activity of the cattle
(Hillary et al., 2013). As a result, pasturelands were impoverished, “grama” grasses
diminished, and the processes of deforestation and fragmentation of the rain forest
accelerated, thereby threatening the biodiversity of the landscape (Wilcove et al., 1986).
Historical palynological studies, or those which examine pollen and its spores, have
shown that since 2000 B. C. corn has been cultivated in Los Tuxtlas (Goman, 1992). By
examining archaeological evidence, the first human settlements were confirmed as early
as 1150 B. C. (Santley, 2007; Santley and Arnold, 1996). The patterns of settlement
(Pool, 2007; Stark and Arnold, 1997) were concentrated to the northwestern and western
edge of the Lake of Catemaco, along its coastline, and throughout the watershed of the
Catemaco River, crossing diagonally the mountain range of Los Tuxtlas (Santley and
Arnold, 1996; Stanley, 2007). Along the Atlantic coastline a large quantity of settlements
were distinguished at the base of the mountain range in the flat plains, and many were
linked to the hydrological infrastructure for managing water as well as sites for defense
(Siemens, 2002, 2009).
Although the pre-Hispanic occupation was continous, it is distinguished by two
periods of marked population increase. The first occurred around 1000 B. C., when
population density reached 8.5 inhabitants per km2. This popluation increase coincides
with the apex of two pre-Hispanic cities, located at the base of Los Tuxtlas mountain
range – Tres Zapotes to the west and Laguna de los Cerros to the south. Both are among
the first urban-ceremonial centers of Mesoamerica and were constructed by the Olmec
culture (Coe, 1965; Stuart, 1993), whose peak began in 1000 B. C. and extended for over
800 years until the Preclassic period (1200 B. C. to 400 B. C.). The population density
subsequently declined to 4 inhabitants per km2, remaining at this level for approximatley
1000 years.
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Another notable increase in population followed at around 500 A. C., leading to a
maximum density of 133 inhabitants per km2. The second increment coincides with the
rise and growth of the city of Matacapan at the centre of the mountain range of Los
Tuxtlas, founded as early as 300 A. C. Located to the northeast of the Lake of Catemaco,
the site demonstrated clear Teotihuacan influence (Coe, 1965). Around the year 500 A.
C. this city arrived at a maximum population of between 35 000 to 40 000 inhabitants
in a urban surface area of 7 ha, concentrating approximately 85% of the estimated
population for this period, including across the numerous settlements identified for this
region, which totaled up to 107 sites (Santley and Arnold, 1996). Archeoalogical remains
found in Matacapan and other settlements along the mountain range demonstrate that
Los Tuxtlas was a zone of crucial importance in the commercial route along the Mexican
high plains between zones of Teotihuacan domination and those of Mayan influence,
especially during the Classic period (250 to 900 A. C.) (Coe, 1965; Santley and Arnold,
1996; Siemens, 2002, 2009). This peak in population was followed by another dramatic
decline, which led to a density of less than 30 inhabitants per km2 (Gooman, 1992;
Santley and Arnold, 1996).
Palynolgical studies demonstrated that during both population peaks a corresponding
decrease in tree pollen is found, paralleling an increase in pollen spores found from corn
and other weeds associated with agricultural activities. Both cycles of deforestation,
corresponding with population densities, are followed by a rapid recuperation of the
rain forest vegetation due to decreases in human population. Particularly notable is the
deforestation that occurred from 200 to 700 A. C., corresponding with the pinnacle of
the city of Matacapan.
Most recently and shortly before the arrival of the Spanish to the Mexican coasts,
the Mexicas dominated an area extending from Tenochtitlan towards a large area of
the watershed of the Papaloapan, located in southern Veracruz. The tributary area of
Tuxtepec extended from the eastern part of the state of Oaxaca until the Papaloapan and
Tuxtlas (San Juan) Rivers in southern Veracruz (Scholes and Warren, 1965), of which
Los Tuxtlas formed a part.
Upon their arrival, the Europeans brought with them horses, donkeys, cows, and other
livestock. In addition to offering the brute force needed for construction, agriculture, and
transport, they were also raised for leather, meat and dairy products, some in corrals
and others purposely left to roam free across the rural landscape and agricultural fields.
The majority of the livestock were cattle, belonging to the variety of Bos taurus, which
originated in the southern Iberian peninsula and in the north of Africa. In both regions
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the cow was raised in pasturelands, as well as left to graze freely in open areas, forest
clearings, and wetlands. The number of heads increased remarkably in a short period of
time, some proliferating more than others and often surpassing the growth witnessed in
their zones of origin (Guevara and Lira-Noriega, 2004). Many cattle ranching operations
were consolidated farily quickly, as witnessed at the end of the colonial period when
only seven large cattle ranches existed between Acayucan and Santiago Tuxtla, Veracruz,
with property extensions reaching up to 270 350 ha. The herds in each site ranged in size
from 1000 heads to up to 30 000 (Aguirre-Beltrán, 1992).
The first regions on the American continent that witnessed cattle were the wet and
dry forests of the coastal plains in the central region of the state of Veracruz (BarreraBassols, 1992), the outskirts of the city of Veracruz, and the southeastern region of Los
Tuxtlas. In the rain forest of Los Tuxtlas the first cow was introduced around 1525, and
three types would later become common in the region: 1) “chichihua lechero,” raised in
swamps, 2) “rodeano,” raised in open spaces, and 3) “montaraz,” which roamed freely in
the rain forest (González-Sierra, 1991; Aguirre-Beltrán, 1992).
The cows became an integral part of the colonial landscape that was already finding
itself in a process of change due to the massive abandonment of land once dedicated to
permanent and temporal agricultural cropping. These lands were previously farmed by
indigenous people who emigrated or died due to diseases such as smallpox and typhus,
brought to the Americas by the Spanish. After a massive decrease in the indigenous
population, the deforestation of the rain forest was halted, fields were abandoned, and
secondary succession began to rejuvenate the clearings and open lands with rain forest
vegetation. The result was a landscape mosaic that offered a large quantity of diverse
resources for the livestock to graze.
The herds diminished the populations of some rain forest species, but dispersed
many others along the edges of the rain forest, beside waterways, and in regenerating
pastures, favoring the proliferation of native “grama” grasses. The cattle flourished
in areas of secondary growth, which previously had been integrated into the farming
systems of the indigenous inhabitants. A large quantity of tree and bush species were
disseminated within these areas by the cattle, enriching the secondary vegetation and
possibly shortening the time span required for secondary succession (Purata, 1986). In
many instances, the landscape formed a favorable environment in which the cattle could
thrive. The cows faced little competition, as large herbivores had already disappeared
almost entirely during the end of Pleistocene. The livestock co-existed with other
smaller herbivore species, such as the tapirs and agoutis that moved through the forest
undergrowth. Predators were only a minor threat to the cattle, which faced the coyote in
open areas or jaguars in the rain forest.
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The most dramatic change to the landscape transpired recently, at the end of the 19th
century and beginning of the 20th. The Bos taurus variety of cattle, who roamed free
across the landscape mosaic, was violently substituted for the livestock variety of Bos
indicus, only raised in stabled pastures. The expansion of this mode of cattle ranching
has occurred at the cost of rain forest, transforming a largely forested and dynamic
mosaic into a less varied and more static patchwork of cropfields, pastures, scrubland,
and remnants of rain forest, with a clear dominance in extension by open and permanent
After the decade of the 1920’s, cattle ranching in Los Tuxtlas, as well as in the entire
state of Veracruz and humid tropics of Mexico and Central America, underwent various
transformations. The introduction of humped or Brahman cattle (Bos indicus), improved
and adapted to the humid tropics, were brought from Brazil and quickly favored over
varieties of Bos taurus. New technologies emerged in the management of grasses and
tropical fodder, originating in Australia and Africa (Reveal-Mouroz, 1980). The first
registration of Bos indicus, was in Acayucan in 1923 (Attolini, 1948). By the end of
the 1940’s numerous herds of different varieties of livestock existed in the lowlands
of Veracruz, including guzerat, gir, nelore, and indubrasil (Melgarejo-Vivanco, 1980),
aiding in the eventual disappearance of Bos taurus whom had prospered in the warm
tropics of Veracruz for over 400 years (Guevara and Lira-Noriega, 2004).
The introduction of new humped cattle breeds and fodder techniques, the burgeoning
demand in the large nearby cities for both meat and dairy products, and the economic
support and stimulus offered for cattle ranching by governments and distinct institutions
have formed the basis for a recent and accelerated expansion of livestock in the humid
tropics of Latin America (Melgarejo-Vivanco, 1980; Reveal-Mouroz, 1980). During
the decade of the 1950’s cattle ranching continued its dramatic expansion, principally
on behalf of large private ranches. Over the next three decades the precedence of cattle
ranching continued to take hold in the region until it was eventually transformed into
the primary productive activity of the ejidos, or communal lands, often at the expense of
agricultural production.
Therefore, contemporary livestock ranching that is practiced in the humid tropics of
Veracruz and in the Neotropics is categorically different from that which was practiced
from the time of the Spanish conquest and up until the first two decades of the 20th
century. As a consequence, the scenario has changed drastically. The criollo livestock
of Bos taurus roamed the landscape freely, causing considerable damage to agricultural
fields, while the humped cattle of Bos indicus was limited to pasturelands, having
little interaction with the surrounding rain forest or agricultural fields. Knowledge of
the complexity, structure, and functioning of this young, 100-year old landscape is
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key to maintaining biodiversity and natural regenerative processes of the rain forest,
considering that it is also the most commonly found landscape scenario in Los Tuxtlas
and much of the Neotropical humid tropics (Figure 4).
The current landscape in the region of Los Tuxtlas is a complicated mosaic, where small
fragments of rain forest are dispersed throughout large extensions of pastures used for
cattle ranching and where trees are found along waterways or exist in the form of remnants,
isolated patches, or living fences (Drawing by Manuel Escamilla).
El paisaje actual en la región de Los Tuxtlas es un complicado mosaico, en el que se destacan
pequeños fragmentos de la selva y una gran extensión de pastos para la ganadería salpicados
de árboles remanentes, solitarios o en pequeños grupos a lo largo de los cursos de agua y en las
cercas vivas (Dibujado por Manuel Escamilla).
Today in Los Tuxtlas two kinds of pastures co-exist (Guevara et al., 1992; Laborde
et al., 2011): pastures of native “grama” grasses (Paspalum conjugatum, Axonopus
compressus, Setaria geniculata, Panicum spp., Digitaria spp.), and pastures of cultivated
grasses mainly of African origin, such as “Estrella de África” (Cynodon plectostachyus)
and “Zacate Guinea” (Panicum maximum).
The pastures of “grama” often expand naturally upon open agricultural fields,
following the traditional slash and burn farming technique upon which native vegetation
is cleared through the cutting and burning of woody vegetation. Afterwards, corn is often
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cultivated for two to four seasons of harvest, although the growing cycle is commonly
extended with one or two harvests of beans, rice, chilies, peanuts, or pineapple. Once the
agricultural field is left fallow the pastureland phase begins, where either native “grama”
grasses take over or “estrella” African grasses are sown to convert the field for livestock.
The “grama” grass is induced immediately after harvest, simultaneously accompanied
by the livestock, so that trampling may eliminate the herbs and grasses that may sprout
from the previous harvest, thereby facilitating the establishment of native grasses (LiraNoriega, 2003).
The “estrella” grass is spread by burying sections of stolons in the soil, thus giving
it a branched-out appearance, and may be planted at the moment in which corn plants
reach a height of at least 40 centimeters or when they are beginning to flower. The
grass is sown in furrows between the rows of corn (Martinez, 1980; Barrera-Laez,
1995). Therefore, a pasture of “estrella” grass can be established immediately after
the one seasonal harvest of corn and even after slashing and burning, when it is sown
directly into the ground. In comparison, a pasture with “grama” grass requires several
agricultural cycles, during which seeds from plants of the families of Gramineae and
Leguminosae begin to accumulate in the soil seed bank, eventually dominating the
pasture floristically and structurally.
In the current landscape, biodiversity has been maintained through the natural
regeneration and secondary succession of once modified landscapes, resulting from a
regimen of ancient disturbances and an intense management carried out by indigenous
peoples throughout the history of the region. The management of this landscape has
been based on conserving the connectivity of the system that permits the movement of
biodiversity, as well as the potential for species regeneration in abandoned sites. Thus,
the historical management of the landscape can be largely described as the management
of its connectivity.
One key element in the modified rain forest landscapes of Mesoamerica and Los
Tuxtlas that has contributed to the management of connectivity is the existence of rain
forest trees that are left behind in open fields, in groups or sometimes alone in the middle
of cleared areas (Figure 5). Currently, more trees exist outside the tropical forest than
within. Such trees could form critical points of a conservation strategy at the landscape
scale (Guevara et al., 1986; Guevara and Laborde, 1993) where perhaps the key to
conserving the rain forest lies outside the forest (Guevara et al., 2005).
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Through examining this aerial photograph numerous solitary trees are perceived across the
landscape, amidst pastures used for cattle grazing.
Esta fotografía aérea permite percibir los numerosos árboles solitarios en los campos ganaderos.
The Mesoamerican culture who historically managed the rain forest understood
and controlled the ecological processes involved throughout the life cycles of trees and
also understood and favored the regeneration of the forest. Trees have been found in
permanent and temporary agricultural fields, in gardens, and along infrastructures built
for the capture, transport, and storage of water and in urban designs. Many trees exist as
living fences, along waterways and in small isolated groups throughout the open fields
of Mesoamerica and all of Mexico, Central America, South America, and the Caribbean
(Figure 6).
Diagram of the principal forested elements that form the rain forest landscape of Los
Esquema de los principales elementos arbóreos que forman el paisaje dela selva en Los Tuxtlas.
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The trees in open fields maintain a close relationship with the diverse species in the
rain forest landscape and other open sites. For example, epiphytic species and flying
vertebrates, such as birds and bats, are dependent upon such residual trees. Thus, the
remaining biodiversity and its movement across the landscape depends upon the
existence of isolated trees, where the movements and defecation of seeds by these flying
animals beneath the crowns of these trees promotes the regeneration of the rain forest
(Guevara et al., 2004).
Currently, the notion that the landscape of the humid tropics is principally dominated
by livestock bare pastures is widely accepted. The conceived landscape is one formed
by small and diminishing fragments of rain forest surrounded by deforested areas,
completely stripped of trees and poor in plant and animal species. However, thin
forested strips still exist, often left along river ways and agricultural fields. These,
riparian forested belts, treed living fences, together with isolated patches of trees that
are left behind, are the forgotten elements of these deforested landscapes. Despite the
extensive deforestation and transformation of the original forest into crop-fields and
pastures, there is a remarkable diversity of plants present in current pastures, which is
closely related to the profuse presence of trees and arboreal elements within the pastures
(Table 1). Thus, a full understanding of the scenario in Los Tuxtlas and other fragmented
rain forest regions is impeded, as through the historical management of the landscape its
connectivity and diversity have been maintained by trees in the open, in spite of intense
use and modification (Guevara et al., 1998).
The practice of leaving trees behind in open fields originated in the traditional
management of the landscape by Mesoamerican peoples and diverse ethnic groups
around the world, including in regions of Central and South America, Asia, and Africa,
and appears to be linked to agricultural practices (Guevara, 1986). In the case of nomadic
agriculture, the fallow cycle allows old-fields and vegetation associated with secondary
succession to develop as the field is let to rest and recovers its fertility. Thus upon cutting
down the rain forest or forested secondary vegetation, trees are often left behind with the
express goal of later accelerating the process of regeneration and establishment of forest
species during fallow cycles or when fields are cyclically interchanged. These remaining
trees are important seed sources, allowing for the dispersal and accumulation of rain
forest seeds in their vicinity (Kelly and Palerm, 1952; Gordon 1982; Van Dorp, 1985;
Howe and Smallwood, 1982; Alcorn, 1984), provided that dispersal agents are present.
In Mexico the practice of leaving behind trees in corn fields has been observed in
the Gulf of Mexico, the Huasteca region (Alcorn, 1984), the Totonac territory of central
Veracruz (Kelly and Palerm, 1952), the Mayan region of the peninsula of Yucatan
(Hernández, 1959; Redfield and Villa Rojas, 1962; Sanabria, 1986; Zizumbo and Sima,
1988; Gómez-Pompa and Kauss, 1990), and in Chiapas (Nations and Nigh, 1980). In
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Central America the same practice has been observed in the Mayan rain forest of the
Guatemalan Peten (Wiseman, 1978), zones inhabited by Kekchi indigenous peoples
in Guatemala (Carter, 1969), and regions of Guaymi indigenous peoples in Panama
(Gordon, 1982). In each zone the isolated trees are linked to the use and subsequent
abandonment of the land and as part of the fallow cycles employed in slash and burn
farming. Many indigenous peoples continue to preserve this traditional method of
farming, practicing it to this very day (Dufour, 1990; Guevara and Laborde, 1992; Illsey,
Species richness by growth form for different landscape elements in Los Tuxtlas pastures.
Sampling surface area is indicated (in m²) for each landscape element.
Riqueza de especies agrupada por formas de crecimiento de distintos elementos que están en los
potreros de Los Tuxtlas. La extensión de muestreo para cada uno está en m2.
On / under pasture trees
Landscape Element→ open living riparian Isolated
Under2 Under3
In spp. list
On Tree
pasture fence
+cows -cows
x104 m²
Growth Form ↓
800 m² ~700 m² 2600 m² 173x104 m² ≈15 x104 m² 600 m² 136 m²
640 x104 m²
Tall & medium trees
(>15 m)
Small trees (<15m)
Ferns & palms
Herbs (monocots)
Herbs (dicotiledonous)
Epiphytes and
Every plant found on 110 pasture trees (38 isolated trees and 72 trees of riparian belts).
Vegetation under the canopy of 50 isolated trees in active pastures with cattle (+cows).
Vegetation under the canopy of 5 isolated trees in which cattle was excluded (-cows) with an exclosure during three
* Species list of the Estación de Biología Tropical “Los Tuxtlas” of the National University of Mexico (EBT-UNAM)
reported by Ibarra-Manriquez and Sinaca 1995, 1996a, 1996b.
**Note: 55 species recorded by Ibarra-Manríquez and Sinaca (op. cit.) as small trees are shrubs according to Burger
1971, 1983; as well as reported in Flora de Veracruz, Instituto de Ecología, A.C.
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Tree cover offers an abundance of ecological benefits to both humans and the rain
forest. Trees in the landscape provide forest products, timber-yielding and other uses,
such as commercial, medicinal, ritual, and even edible materials or products. In addition,
they offer other environmental services such as shade, improving the condition of the
soil, attracting frugivorous animals, beautifying the scenery, and serving as symbolic
markers across the landscape (Guevara, 1986; Guevara and Laborde, 1993; Guevara et
al., 2006).
The voluminous crowns of the trees are often covered with flowers and fruits, and
among their branches and trunks exist a variety of bromelias, orchids, and plants from
the Araceae family. Underneath their crowns at the grown level numerous juvenile plants
of woody trees species associated with the rain forest are established. The presence
of these rain forest plants in here is related to bird and bat activity, which aids in the
dispersion of seeds beneath the crown of the isolated trees (Guevara and Laborde, 1992;
Guevara and Laborde, 1993; Laborde, 1996; Galindo-González, 1998, 1999).
Therefore isolated trees are an oasis in an unfavorable habitat for flying animals
that require sites to rest upon crossing the pastures between fragments of rain forest.
According to observations in four isolated Ficus spp. trees in pasturelands with more
than 15 years of use, 47 species of visiting frugivorous birds were registered (Guevara
and Laborde, 1993). This number of species represents a third of the total of frugivorous
birds registered in the neighboring biological reserve of the National Autonomous
University of Mexico (American Ornithologist Union, 1983). The attractive quality that
a tree in the open may have for frugivorous species depends on the type and quantity of
fruits produced by the three throughout the year, as the largest number of visitors are
recorded when the trees are most loaded with fruits (Laborde, 1996). Even when the
trees did not have fruits, more than four visits per hour were registered, indicating that
frugivorous birds also used the trees to perch and rest. In another study, over 652 bats of
more than 20 different species were captured in isolated trees in pastures, totaling 56%
of the species reported for the zone (Galindo-González, 1998, 1999, Galindo-González
et al., 2000). Of the total capture, 81% of the bats were frugivores and the most
abundant species were Sturnira lilum (40%), Artibeus jamaicensis (15%) and Carollia
perspicillata (10%), which are effective dispersers of seeds for numerous species of the
rain forest (Figure 7).
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Solitary trees attract birds and frugivorous bats in flight across open fields, thus forming
connectivity with the surrounding landscape. Seeds deposited from the visiting birds and
bats accumulate beneath the crowns of these trees, thus converting these isolated trees,
amidst abandoned pastures once grazed by cows, into sites or nuclei of potential rain forest
Los árboles solitarios ejercen una gran influencia sobre aves y murciélagos frugívoros que vuelan
a través de los campos abiertos atrayéndolos -efecto de conectividad-, y acumulando las semillas
que dejan caer bajo su sombra convirtiéndose en potenciales núcleos de regeneración de campos
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When cattle are excluded from the vicinity of trees in the open and the effects of
their grazing and trampling are stopped beneath the shade of isolated trees, along with
halting the clearing of vegetation and the application of herbicides, interesting changes
occur in the composition of species and in the structure of the plant communities that
grow beneath the canopy of the trees (Guevara et al., 2004). In a study to assess the
effect of removing cattle, the abandonment of a pasture during three years was simulated
by fencing off five isolated trees of Ficus spp. (Guevara et al., 2006). At the end of
the experiment 96 species of plants were found growing beneath the crowns of the
trees, with an average density of 4.6 plants per m2. During this lapse an arboreal and
closed canopy was formed, with more than 4 m in height, dominated by secondary and
pioneer woody species whose individuals varied in size from small seedlings of 10 cm
in height up to juvenile trees of more than 6 m in height. Underneath this canopy the
ruderal species typical of open grasslands disappeared almost completely, while 46 tree
and shrub species characteristic of the rain forest were successfully established (Guevara
et al., 2005; Table 1).
The main obstacles for the regeneration of the rain forest in grasslands has been
attributed to the absence of seeds from rain forest plants (Guevara, 1986) in addition
to the adverse conditions for seed germination and seedling establishment of rain forest
species (Nepstad et al., 1990; Holl, 1999; Holl et al., 2000). The low or almost null
immigration of seeds from woody species into the open grasslands is explained by the
low availability of propagule sources in and around these sites, as well as the unattractive
nature of these sites for frugivores that disperse the seeds of woody plants. The few
seeds of woody plants that manage to arrive to the grassland face adverse conditions
for their germination and establishment, such as compacted soil due to its trampling by
cattle, as well as extreme oscillations of humidity and temperature of both the air and
soil in open areas. In addition, other unfavorable biotic factors exist for the tree species
of the rain forest, such as competition with grasses and heliophilous or sun-demanding
weeds that grow rapidly, intense depredation of seeds, and herbivory of its plants by
organisms that thrive in crop-fields or pastures.
The findings of Los Tuxtlas shown that trees change favorably in response to certain
conditions, since they attract to pastures frugivorous animals that disperse seeds of rain
forest species (Guevara and Laborde, 1993; Galindo-González et al., 2000) by creating
in their shade favorable microclimatic and edaphic conditions for the establishment of
rain forest plants (Williams-Linera et al., 1998) through abating the competition from
heliophilous species, but only when livestock is excluded and associated management
practices are put to a halt. The isolated trees in pastures of Los Tuxtlas act as
“regeneration nuclei” (sensu Guevara et al., 1986) and are determinant in the successful
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establishment of rain forest plants in the interior of the pastures. After three years of
interrupting livestock practices, the effect of the crown of the isolated trees combined
with that of the secondary canopy that forms below makes a more attractive setting
for birds and frugivore bats and creates microclimatic conditions that are increasingly
favorable for the establishment of rain forest plant species, further excluding ruderal
plants from the pastures.
However, the appeal of isolated trees to frugivores that disperse seeds depends
directly on the presence and abundance of dispersing frugivores and on the existence
of propagule sources in the area (Aguirre, 1976; Arriaga and Lozano, 1980). This is at
the same time a function of the extension, form, and floristic composition of the tree and
forest elements that are left in the landscape (Laborde,1996). Due to this, the effect of
isolated trees, in particular their efficiency as “centers of regeneration” of the rain forest,
obey the spatial pattern of the landscape and its floristic composition. This is to say that
regenerative activity is dependent upon the spatial distribution of forested elements
that have not been eliminated by deforestation, since these represent the habitat of the
frugivores that disperse seeds and are the principal sources of propagules of rain forest
plants in fragmented landscapes.
Landscape Design and Management
Understanding the history of a fragmented landscape is important in order to
recommend measures for the management of its natural resources and biodiversity,
and in this case the goal of ultimately contributing to the natural regeneration of the
rain forest. In the case of Los Tuxtlas and many sites in Mesoamerica, maintaining the
connectivity of the landscape is crucial for natural regeneration of the forest to occur.
Such a vision is only enlightened by understanding the historical management practiced
by indigenous populations, at times corresponding with contemporary management, by
which secondary succession is manipulated by the practice of leaving behind forested
elements in agricultural fields.
Enhancing the connectivity of a landscape may be considered as the opposite
of isolating the elements within the landscape. Thus, connectivity has a structural
component that refers to the quantity of physical contacts, for example, between
remnants of rain forest, as well as the magnitude of the distances by which fragments
are separated. While the functional aspect of connectivity, deals with the frequency
or intensity of the flow of organisms, nutrients, materials, or energy across the
landscape (Forman and Godron, 1986). In the case of forested, fragmented landscapes,
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connectivity determines to what extent organisms remain in remnants separated by the
fragmentation and to what degree they may form a functional demographic unit (Turner,
1989; Laborde, 1996; Figure 8).
The pattern or distribution of trees across open fields allows for the management of
the connectivity of the landscape and the maintenance of biodiversity in spite of the
fragmentation of the rain forest and the large extension of pasturelands for cattle ranching.
El patrón de distribución de los árboles en campos abiertos permite manejar la conectividad del
paisaje y mantener la biodiversidad a pesar de la fragmentación de la selva y la gran extensión
de los campos ganaderos.
In fragmented landscapes the distances that separate the remaining fragments of
rain forest have been utilized to estimate its degree of isolation (Guevara, 1995). The
greater the distance of separation between two fragments, the number of animals
able to move between the fragments would be fewer. However, upon considering the
distance that separates the fragments, one must also consider the existence of complex
and heterogeneous structural and floristic characteristics of the landscape. In particular,
forested elements such as trees in the open are often ignored in such calculations,
despite remaining as an integral part of the landscape and influencing the movement
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of organisms across it. From this perspective, the remaining fragments of rain forest
may only be problematically described as “islands of rain forests” immersed in a “sea
of grasses”. A more accurate assessment would acknowledge the connectivity of the
landscape by recognizing the existence of numerous and varied tree elements that are
still found in pastures and other open and agricultural sites which are used by forest
organisms as stepping stones while crossing open areas. These arboreal elements offer
temporal refuge, rest, or even food to the native fauna of the rain forest that ventures
outside the rain forest fragments (Laborde, 1996; Guevara et al., 1998; GalindoGonzález, 1999; Graham, 2001).
The Future Landscape of Los Tuxtlas
The future landscape of Los Tuxtlas could witness two extreme scenarios. The
first would represent a landscape constituted by a disintegrated group of elements,
dominated by extensive pasturelands and lacking in connectivity. In this scenario, the
native populations of rain forest species that survived the clearing of trees would remain
confined and restricted to the interior of the fragments, totally isolated from the surviving
individuals in other fragments. Given the severe current fragmentation of Los Tuxtlas
forest, the previous situation would provoke in the long run a local extinction of a large
number of species and a decrease in biodiversity. The second scenario is a landscape
of interconnected elements of rain forest fragments, pastures, agricultural fields, and
secondary vegetation, integrated due to elements that maintain connectivity across the
landscape such as isolated trees, riparian corridors, living fences, among others, that
would contribute to the maintenance of rain forest species and their accessibility between
sites. This interconnected landscape could sustain itself long-term and shelter a high and
representative percentage of the original native biodiversity of Los Tuxtlas by allowing
for a natural regeneration of the rain forest without the need to limit agricultural and
livestock activities.
The re-placement or maintenance of pastures with tree cover is easily-carried out
in practice and may be accomplished by controlling the clearing of vegetation and the
spraying of herbicides beneath the crown of the trees, in addition to excluding cattle
from grazing for relatively short periods of time. The rich and diverse forest vegetation
that regenerates under these conditions allows rain forest species to be naturally
selected and to replace trees of the original canopy. For the maintenance of the native
biodiversity of Los Tuxtlas, the conservation of the most extensive and remaining rain
forest fragments is indispensable, for which it is necessary to halt the current clearing of
vegetation and deforestation.
The decree of the Biosphere Reserves of Los Tuxtlas (see Guevara et al., 2006) was a
promising step towards strategic management at the landscape level, where management
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practices that would aid in the connectivity of the landscape may be implemented
(Figure 9). The reserve includes three nucleus zones of strict conservation, totaling
nearly 30 000 ha (approximately 10 000, 18 000 and 2000 hectares, respectively) and
protecting the most extensive remnants of rain forest and cloud forest of the region. The
three nucleus zones are surrounded by a single buffer zone that includes 125 000 ha of
pastures and agricultural fields that contain remnant forested elements and are immersed
between fragments of rain forest of distinct sizes, a large quantity of riparian corridors,
living fences, and numerous isolated trees. The mountain range of Los Tuxtlas is an ideal
scenario to implement different alternatives of natural resource use and management
across both the nucleus and buffer zones, which could later be applicable to the unique
scenario of the humid tropics of Mesoamerica that have high population densities and an
advanced grade of fragmentation.
Map of the Biosphere Reserve of the Los Tuxtlas. 1-2-3 Core zone, 4 -Buffer zone.
Mapa de la Reserve de la Biosfera Los Tuxtlas. 1-2-3 Zona núcleo, 4-Zona de amortiguamiento.
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El paisaje de la selva en el territorio de Mesoamérica se ha construido con las
especies de plantas y animales, que han estado sujetas al impacto de eventos naturales
y a la selección de la intervención humana. Este trabajo se enfoca en el efecto que
pudo haber tenido el ganado vacuno, introducido en el siglo XVI y en el siglo XX, en
la composición y el funcionamiento del paisaje en la región de Los Tuxtlas, Veracruz,
La sierra de Los Tuxtlas es un enclave del paisaje del sureste mexicano, dominado
por la selva húmeda que se formó con especies de flora y fauna provenientes del
neártico y del neotrópico, que se adaptaron a los intensos cambios climáticos, a la
violenta actividad volcánica y a las frecuentes tormentas tropicales que azotan la región
de manera continua. También fue modelado por el manejo del suelo y la selección de
especies vinculadas con la caza, recolección y cultivo, llevados a cabo por el pueblo
Olmeca, la civilización más antigua de Mesoamérica.
Entre los eventos más recientes e influyentes en la construcción del paisaje, se
destaca la llegada del ganado durante la Colonia. En el siglo XVI se introdujeron razas
de Bos taurus, provenientes de la cuenca del Guadalquivir. En Los Tuxtlas, las vacas se
adaptaron a la vida libre, probablemente debido a la ausencia de los grandes herbívoros
nativos que desaparecieron a fines del Holoceno y a la gran extensión de vegetación
secundaria en los campos agrícolas abandonados por los cultivadores indígenas.
Las vacas ramonearon y pacieron en la selva y la vegetación secundaria durante
500 años, dispersando árboles, arbustos y hierbas, con lo cual se enriqueció el proceso
de sucesión secundaria. Un ejemplo de ello es el desarrollo de los pastizales nativos
o de grama, donde pacieron hasta principios del siglo XX. Fue entonces cuando se
substituyeron con razas de B. indicus, originarias de Asia y Brasil, que se criaban en
potreros sembrados con especies de pastos africanos, lo cual provocó un déficit repentino
de la herbivoría, una defaunación muy intensa, que empobreció la vegetación secundaria
(actualmente casi desaparecen los pastizales de grama) y aumentó la deforestación y la
fragmentación de la selva, que ahora son las principales amenazas para la biodiversidad
La sierra tiene todavía una gran diversidad biológica. Probablemente debido a
las constantes e intensas perturbaciones de pequeña y gran escala ocasionadas por
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los eventos naturales y por la intervención humana, cuyos asentamientos ocuparon
profusamente la comarca con altas densidades de población, una influencia que se inició
hace más de 5000 años.
Una prueba de ese manejo son los numerosos individuos y especies de árboles de la
selva que se ven en los campos abiertos. Hoy están incorporados al paisaje por costumbre
o tradición, mitigando como lo hicieron desde que se asentaron los primeros pobladores
en la región, el efecto negativo de la deforestación y atenuando las condiciones
provocadas por la presencia y ausencia de herbívoros nativos y exóticos. Estos árboles
son determinantes para la conectividad ecológica, que contraresta el aislamiento
de los fragmentos de la selva y para sostener el proceso de sucesión secundaria y de
regeneración de la selva.
El reconocimiento que el paisaje está cambiando constantemente como efecto de los
factores naturales y humanos y que algunos de ellos, como la agricultura vernácula y
la llegada del ganado han dejado memoria, como es el caso de los árboles solitarios,
plantea nuevas alternativas para mantener la biodiversidad e impulsar el desarrollo
racional, empleando elementos remanentes en el paisaje.
Palabras clave: Veracruz, sucesión secundaria, defaunación, conectividad del
Chapter 3.
Environmental History
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Universidad Nacional de Colombia, Carrera 45 No 26-85 - Edificio Uriel Gutiérrez. Postal code 11132. Bogotá (Colombia).
E-mail: [email protected]
This document brings together the results of research with the aim of analyzing the
transformation of cattle ranching practices during the Colonial period in the specific
context of the Saquencipá Valley on the Altiplano Cundiboyacense (High Plain) in
Colombia, its relationship with land ownership and the impact that these had on the
ecosystem, especially as regards soil deterioration. In order to reconstruct the farming
practices used during the 16th and 17th centuries and their Iberian origins, colonial
documents from the depths of the Archivo General de la Nación (Nation’s General
Archive) in Bogota, Colombia and the Archivo General de Indias (General Archive of
the Indies) in Seville, Spain and along with the chronicles, geographic relationships and
results from the history and archeology literature were exhaustively reviewed. Taking
into account factors such as the link with agriculture, the mobility and number of head of
cattle, and the natural control exerted by drought and predators, the high environmental
impact attributed to colonial cattle ranching in the region is questioned.
Key words: Environmental history, colony, cattle ranching, soil deterioration.
The lands of the Saquencipá Valley are located in what is currently the Department
of Boyacá, Colombia (Figure 1) but which, during the 16th and 17th centuries, lay within
the jurisdiction of Villa de Leyva in the Viceroyalty of New Granada. These lands were
noteworthy for the quality of their wheat harvests. Grains, flours and biscuits travelled
from there—both legally and illegally—to different regions of New Granada including
productive regions in the surroundings of Santafé, Tunja and Pamplona. However, some
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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K.G. Mora Pacheco
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Colonial livestock farming in Saquencipá Valley
of the visitors of the 18th and 19th centuries left a record of the apparent decline in the
region that followed the solar eclipse of 1691 and highlighted the lack of agricultural
productivity based on different types of land use they came across (Oviedo, 1930;
Ancízar, 1983). Even today, differences in the morphology, climate and plant cover
of the Saquencipá Valley relative to that of the rest of the Altiplano Cundiboyacense
in which it is located disquiet those who visit it and have prompted research to find
historical explanations for this panorama of deterioration, purportedly related to the
adverse effects of agricultural activities during the Colonial period.
Saquencipá Valley, research area.
Valle de Saquencipá, área de investigación.
(By Alfonso Simbaqueba Hurtado)
In the Saquencipá Valley, the geographer Joaquín Molano (1990) was the first to
carry out a historical reconstruction of the erosive processes that were observed in the
1980s. His analysis, based on field work and secondary sources, led him to conclude
that Spanish colonial occupation was a fundamental, causal factor in the aridity and
erosion that occurred in Villa de Leyva and its surroundings. According to Molano, the
intensification of soil use, deforestation and the introduction of cattle ranching and wheat
monoculture were what caused environmental deterioration in the region (Molano,
1990). A systematic review of the colonial documents and chronicles, and deeper inquiry
into aspects such as the transformation of the agricultural activities of the indigenous
people after the Conquest, the role of Villa de Leyva in supplying cereals to other
PASTOS 2012. ISSN: 0210-1270
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Colonial livestock farming in Saquencipá Valley
regions of New Granada, agricultural techniques and the way agricultural production
was managed for each type of land use, however, suggest different conclusions.
In fact, archeological studies (Falchetti, 1975; Boada, 1991) posited that the
severe erosion originated in the pre-Hispanic period rather than during colonial times.
According to Falchetti (1975), during the 16th century the majority of the land was still
fertile. The demand for wood for Spanish buildings and the cultivation of wheat did
indeed contribute to the deterioration of the soil, but the latter was also associated with
both the pre-Hispanic specialization in manufacturing pottery and open air firing that
required large quantities of firewood (Falchetti, 1975). Boada (1991) also observes that
soil deterioration might be rooted in pre-Hispanic practices. She especially points to the
effects of changes in the settlement patterns and the intensification in pottery production
in the 9th and 13th centuries C.E. Unable to do any agricultural work because of the
previous damage done to the soil, people had to specialize in some kind of production
to obtain food supplies from other regions, and chose pottery making. This hypothesis is
open to future archeological and palynological research (Boada, 1991).
In the framework of this discussion, the present study examines the relationship
between the agricultural practices implemented in the Saquencipá Valley during the 16th
and 17th centuries, and their possible connection with soil deterioration. To this end, a
first step was to reconstruct the agroecosystem that the Spaniards employed in the 16th
century, and their introduction of new species of livestock. The cattle ranching practices
of the Iberian Peninsula at the time of the Conquest and colonization of the Americas
are analyzed, as well as the way these practices were implemented in the Tunja Province
of New Granada, especially in the Saquencipá Valley. The information presented here
enriches the debate on the relationship—generally considered a negative one—between
the environment and cattle ranching. The allocation of common indigenous lands
(resguardos1) on poor soil, large scale deforestation to make way for livestock, lax
breeding of all types of livestock and the overpopulation of domesticated animals are
Following comparative examples in the literature (Sluyter, 1997; Aguilar Robledo
and Torres Montero, 2005), the research is based on an exhaustive review of a range of
primary sources from the Archivo General de la Nación (General Archive of the Nation)
in Bogota (AGN) and the Archivo General de Indias (General Archive of the Indies) in
1 Land that was allocated to the indigenous people at the end of the 16th century. In theory, they could not
lease out or sell this land; it was for their joint use.
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Seville, Spain (AGI). Documents such as the visitas2, indigenous common land grants,
land disputes, accounts, and official reports about supplies provided useful information
about topics such as the introduction of domesticated animals and their fiscal importance,
times of abundance and scarcity, the different degrees of economic dependence on cattle
ranching, and the changing regulations governing it. Geographic reports and colonial
chronicles were helpful in reconstructing the prevailing ecosystem conditions in the
study region during the 16th century, the perception of these conditions by the European
colonists, and the structure of their cultural and socioeconomic organization on these
These primary sources were complemented by and discussed in light of the results
of research in archeology, history and geography that made it possible to reconstruct
the approximate location of settlements, land ownership, land use, plant cover, the
introduction and proliferation of foreign species in the study region, and to identify
similarities and differences with other places on the continent.
The analysis was complemented by field reconnaissance and mapping work. This
allowed us to georeference the data mined from the archeological evidence and primary
sources, and include them in a geographic information system (GIS). The main objective
was to establish the approximate location of the indigenous common lands, private
ranches, resguardos, estancias3 and ecclesiastical lands in order to detect any relationship
between the topography, climate and the hydrography along with the introduction and
expansion of cattle ranching, and the speed and intensity of any soil deterioration. With
historical GIS applications one does not expect exact results, especially considering
the changes that have occurred in this vast cultural and natural landscape over the last
400 years. To suggest that there is a straightforward association between currently
arid land and areas that were unproductive during the Colonial period would imply an
underestimation of the recent actions that have contributed to the problem. On the other
hand, the sources often refer to places with indigenous names that have left no trace
on the current landscape, and therefore cannot be included in the map. Additionally,
the sources mention areas that were measured in unstandardized units (cabuyas4) and
defined with reference to neighboring properties which were also poorly delimited in
2 Formal inspections of indigenous villages and resguardos (see next footnote) carried out by an official, the
Visitor, to set tribute taxes and record the size of the indigenous population, the state of people’s health, and
the condition of the land, as well as the relationship with the encomendero, the priest and the neighbors in
the nearby towns and cities.
3 Land assigned to the Spanish. In general, the land was used for raising livestock.
4 A measure of length used in the 17th century that was made from a length of the fique agave (Furcraea
bedinghausii) and kept in each town hall. Its length varied from region to region, but was approximately 80
cm (S.I.).
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spatial terms. In spite of these limitations, it was possible to use the available data to
prepare the figures that accompany the results presented here.
The predominant view of environmental historiography has been that ruminants and
ungulates in were absent from the Americas before the Conquest and that this created
an intrinsic weakness in the continent when domesticated animals and cattle were
introduced (Crosby, 1972; Crosby, 1999; Diamond, 1998). In the absence of previous
adaptation or complementarity between the new fauna and the existing plant cover, soils
and native fauna, the encounter is thought to have resulted in a series of adverse effects,
including deforestation, disease propagation and soil deterioration. This view gives
rise to the following questions: Is this a valid explanation for soil deterioration in the
Saquencipá Valley? What was the ecosystem like when the Spaniards first arrived in the
region in the 16th century?
From the end of the 12th century Spanish customs and, later, colonial rules
(Gutiérrez, 1983) dictated that to found a new settlement (city, villa or pueblo) places
should be chosen with herbaceous areas nearby to allow cattle to graze (Klein, 1979).
Natural herbaceous formations and those created by pre-Hispanic societies offered ideal
conditions for implementing cattle ranching (Aguilar Robledo, 1998), while forested or
wooded areas were avoided because they required more time, labor and tools (Patiño,
1977). According to the literature, the latter were the conditions in the Saquencipá Valley
where the town of Villa de Nuestra Señora de Leyva was founded in 1572.
The chronicle Relación del Nuevo Reino de Granada, written in 1571 by Friar
Gaspar de Puerto Alegre, mentions oaks (Quercus humboldtii) as the most common tree
in the region, though with smaller acorns than those of Spanish oaks. Also abundant
in the surroundings of the Tinjacá Lagoon was alder (Alnus acuminata), which was
good for construction (Tovar, 1988). Another chronicle, the 1610 Relación de Tunja
(Patiño, 1983), confirms that the vegetation was of a type that could be included in
Holdridge’s Lower Montane Dry Forest. It included native plants (local names are
given in parentheses) such as Caesalpinia coriaria (dividivi), Sena viarium (alcaparro),
Furcraea andina (fique or cabuya), Dodonea viscosa (hayuelo), Alnus acumminata
(aliso), Ficus indica (tuna), and Furcraea bedinghausii (fique or cabuya). The chronicler
Zamora recorded these very plants as raw materials used by the indigenous people to
make rope from the fibers of the fique plant, and soap from its flowers for instance. The
miel de muelle, a sweet substance extracted from pepper tree (Schinus molle), was used
to alleviate aches and chills (achaques de frío) or was added to chicha, a traditional
beverage made from fermented maize. The buds of the pepper tree were also used for
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cleaning teeth and to improve gum health. Natural dyes were extracted from the dividivi
plant (black dye) and from the fruit, known as tuna (red dye) (Zamora, 1945).
Having said that, presumably in the early 16th century there were no lush forests in
the region, but rather nuclei of native species with low regeneration capacity (Therrien,
1991), in addition to the herbaceous and shrubby vegetation with xerophytic plants.
Pottery production from pre-Hispanic times with open-air firing required a good deal
of firewood (Falchetti, 1975). Hayuelos (Dodonaea viscosa), oak (Quercus humboldtii),
frailejones (Espeletia spp.) and small shrubs proved to be very useful (Therrien, 1991).
To prepare areas for agriculture and gain ground against the Andean forests, the preHispanic inhabitants used fire (Márquez, 2001; Patiño, 1965; Patiño, 1997). Right before
the rainy season, the understory and largest trees that might crush the surrounding
vegetation were cut, and the fire set upwind of the burning area in order to speed up the
process (Patiño, 1965).
On the Cundiboyacense High Plain, tree felling and burning led to the exuberant
vegetation (which had a slow recovery capacity owing to the high elevation) giving
way to pastures, making burning less and less necessary (Fals Borda, 2006). In the
Saquencipá Valley, aridity became an additional factor that prevented the arboreal
vegetation from recovering and favored the propagation of xerophytic plants. The early
predominance of cacti in the region is supported by references to the use of cochinilla
(Dactylopius coccus), an insect that lives in the tuna fruit of the Opuntia cactus and
from which purple and red pigments were extracted to dye the chiefs’ garments in preHispanic times (Molano, 1990), and a variety of textiles during the Colonial period
(Pérez Arbeláez, 1990). The chronicler, Friar Pedro Simón stated that the lands of Tunja,
which included the Villa and the surrounding indigenous villages, were mountains
covered in grass all year round (Original in Spanish: “las sierras de pasto todo el año”)
(Simón, 1981) and that the Saquencipá Valley was surrounded by steep, bare gullies
(“escarpadas y peladas breñas”) (Simón, 1981). Abundant deer continually interacted
with this herbaceous vegetation that was predominant in the Valley. Rodríguez Freyle
said that in the times when the Spanish arrived, deer were so abundant that they moved
in herds as if they were sheep, eating the inhabitants’ crops and livelihood (“eran tan
abundantes que andaban en manadas como si fueran ovejas, y les comían sus labranzas
y sustentos”), but could not be hunted without the permission of the cacique, the head of
the indigenous people (Rodríguez Freyle, 1979).
On one hand, the description may have been somewhat exaggerated to exonerate the
Spanish and their cattle from the destruction of the crops of the indigenous peoples, or
to emphasize the pre-Hispanic origin of the problem. On the other hand though, it gives
evidence of the abundance of deer which may have contributed to the reduction in plant
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The abundance of deer in Tunja Province is confirmed in the Epítome de la
Conquista, written around 1550. This chronicle mentions that the meat eaten there by
the Indians is venison, with such an abundance of deer as to be sufficient to keep them as
[we keep] cattle here (“las carnes que comen los indios en aquesta tierra son benados de
que ai infinidad en tanta abundancia que los vasta a mantener como acá los ganados”)
(Tovar, 1988). By the middle of the 18th century, the multitude of deer encountered by
the Spanish was such that, in spite of the increased pressure of hunting with shotguns
that began decades before (Zamora, 1945), it is said that of these wild animals, native
to these lands, those most esteemed and of greatest utility are the deer, of which the
mountains and plains of the cold lands are full (“de los animales silvestres montaraces,
propios de estas tierras, los más estimados y de gran utilidad son los venados, de que
están llenos los montes y sabanas de tierra fría”) (Oviedo, 1930).
In short, it seems that the Spanish settled in an area of the Valley where grasses used
to be eaten by wild ruminants and ungulates, and that these animals were often hunted. In
this context, to what extent did the introduced livestock species spread, and what might
the impact of grazing have been on such processes as the decrease in forested areas,
soil deterioration and loss of crops? In Tunja Province, the environmental context was
suitable for breeding domesticated animals, and these quickly multiplied as highlighted
in an anonymous report from 1560:
This city [Tunja] is the largest in this district […]. The Spanish breed
all genera of livestock in great abundance, cows, mares, goats, sheep, and
their multiplicity has been such that they now damage natural crops and
it would be convenient to remedy this. “Esta ciudad [Tunja] es la mayor
deste destrito (…). Crían los españoles todo género de ganados en gran
abundancia, vacas, yeguas, cabras, ovejas, assi que ha sido tanto el
multiplico que ya hacen daño en las labranzas de los naturales y conbiene
poner remedio en ello” (Tovar, 1988)
Only two decades after the city’s foundation (1539), the introduced animals
had successfully reproduced and raising livestock had become an activity of great
importance. According to Zamora, the main reasons for such prosperity were the
nutritiousness and saltiness of the pastures of Tunja Province (“lo pingüe, y salitroso
de sus pastos”) (Zamora, 1945), and the animals’ importance as a source of food, a
means of transportation, raw material for making candles, soaps and leather goods, as
well as their cultural significance in culinary customs and bull fighting (Zamora, 1945;
Oviedo, 1930). The time and money required to import the animals and then maintain
them, together with the pastoral mentality of the Spanish and the economic and cultural
importance of cattle raising, led the colonists to use quality land to breed the animals
(Patiño, 1997; Melville, 1999).
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According to several authors, the delimitation of the ejidos5 and the award of
estancias de ganado (cattle ranches) in the high plains of the New Granada often became
agents of damage for the crops of the indigenous people. Animals were said to invade
their corn fields and displace the indigenous people toward steep areas with lower crop
yields and that were more prone to erosion (Patiño, 1965; Patiño, 1969; Molano, 1990;
Boada, 1991; Orbell, 1995; Langebaek, 2001). In the Saquencipá Valley however, this
is unclear. One piece of evidence comes from the visit of Juan de Valcárcel in 1636.
He records the problems faced by the Suta Indians caused by the livestock (especially
cows and mares) of their encomendero6, Pedro Merchán de Velasco, invading, eating
and trampling their crops. According to this priest, the animals had easy access because
the land did not lend itself to the construction of fences or other types of defense because
it was flat and open everywhere, with no resistance to said livestock (“la tierra no es
dispuesta para hacer talanqueras ni otras defensas por ser rasa y abierta por todas
partes, no tienen resistencia al dicho ganado”) (AGN, VB, T.14, f.757r.). This reference
would allude to the relatively flat relief of the indigenous lands.
Further evidence comes from the indigenous land title documents in the region.
Although there is mention of situations of land on slopes being assigned to indigenous
people, in a number of cases, witnesses consulted during the process and the
topographical description and the vista de ojos (the area that fell in the line of sight of
the topographer) have allowed us to detect indigenous lands on flat areas and, in some
cases, in fertile river valleys and ravines (Figure 2). One case that is notable for having
one of the best locations was that of Turca and Gachantivá, indigenous towns with land
bordering the river that is good for crops because it is irrigated when the Cane River
rises (“tierras que caen en las vegas del río son buenas para labor porque se anegan
con las crecientes del río de Cane”). This favored their keeping more than forty pairs of
oxen (“más de cuarenta yuntas de bueyes”) and producing wheat, corn, anis and squash,
but, paradoxically, left their crops vulnerable to trampling by the livestock that drank in
the salty pastures (AGN, VS, T.2, f.553r., 559v., 553v.).
5 Communal land under the perpetual stewardship of rural inhabitants for agricultural activities.
6 Title given to the Spaniard who was in charge of both receiving tribute taxes from the group of indigenous
people under his purview, and who was responsible for guaranteeing their evangelization.
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Location of settlements and villages, 16th and 17th centuries.
Ubicación de los asentamientos y aldeas, siglos 16 y 17.
(By Alfonso Simbaqueba Hurtado)
In any case, domesticated animals and livestock practices generated a good deal
of conflict among the farmers, regardless of whether they were indigenous, mestizo,
or Spanish. For example, the traditional privilege of the la derrota de las mieses
(defeat of the crops) in which cattle were allowed to graze on the remaining stubble
after the harvest, became a regular occurrence where there were no ravines, regardless
of season and resulted in the livestock becoming invasive and destroying permanent
crops. On the peninsula, the privileges of the livestock farmers, especially those who
raised sheep and herded their animals in la Mesta, were increased under the reign of
Ferdinand and Isabella. With the aim of increasing wool production and export,
agricultural improvement projects were forbidden in Granada and the import of wheat
was authorized to avoid plowing the grasslands in the Kingdom of Castilla and Aragón.
Royal pastures were leased, fences eliminated, the felling of small trees was authorized
to feed the sheep when grass was scarce, and testimonies were required to certify the
use of the land for purposes included in the list of “five forbidden things”, i.e., land
where livestock was not allowed free access: pastures, wheat fields, vineyards, orchards
and hay meadows (“dehesas, trigales, viñedos, huertas o prados de guadaña”) (Klein,
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In this regulatory and historical context, how did the colonial authorities react to the
conflicts between crop and livestock farmers? From Víctor Manuel Patiño (1965), we
learn that permanent fences to protect the wheat and corn fields were banned and fencing
was only allowed around permanent crops or woody crops such as vineyards and olive
groves brought into the region at the end of the 17th century. Nonetheless, in practice,
living fences of cambronera (Lycium spp.), brambles, prickly pear cactus and, in Villa
de Leyva, cabuya (fique plants) were often used, and in a very few cases the fences that
had been built by the indigenous people were kept. This was not just a colonial practice.
Even in Castilla, the rules protected wheat fields, vineyards and orchards, and allowed
for the construction of fences.
In the Americas, once again, rules of this kind sought to guarantee the food supply
and tribute payments. On a visit in 1571, led by Juan López de Cepeda, the Indians of
Monquirá made mention of their being forced to build pens made of adobe (bahareque)
for their encomendero, although this low grade precaution did not prevent damage to
the crops (AGN, VB, T.5, f.374v.). During the 1636 visit the conflicts between the Suta
Indians and their encomendero, Pedro Merchán, caused by his livestock invading their
crops made the need for livestock owners to enclose their animals and build trenches
to protect the crops patently obvious and was recorded in writing; these measures were
set during the presidency of Antonio González, from 1590 to 1597 (AGN, VB, T.14,
f.819r.). Contempt for the preventive measures that had been ordered resulted in the free
movement of livestock, however, the accused encomendero was required to pay a fine
and hire herdsmen. At the same time, the indigenous inhabitants were ordered to build a
trench and a fence (AGN, VB, T.14, f.840).
In Tunja Province, other types of regulations aimed at controlling cattle raising
activities were set when its capital city was founded. The city of Tunja was founded on
August 6, 1539 and the Minutes of its Town Council meeting on the 14th of August in
1539 includes the precise delimitation of the pasture for the neighbors’ horses on the
Paipa road, separate from the land where space for orchards would be granted (Ortega
Ricaurte, 1941). To determine sanctions, resolve conflicts and regulate pasture use, cattle
branding had been required by the City of Tunja since 1541, as recorded in the minutes
of the Town Council for February 4th, and March 26 and 29, in which brands were
assigned to several neighbors (Ortega Ricaurte, 1941), and from then on and for decades
afterwards, a person was appointed to be responsible for the branding irons used to mark
the cattle bred by the indigenous people (AGI, Santafé, 66, Nº95, f.1r.).
Other measures perpetuated the Spanish Mesta custom of holding an annual meeting
to deal with stray animals (Klein, 1979). In the New Kingdom, the Tunja Town Council
minutes of July 1564 included in its determinations that cattle be confined at night so
that it could not damage natural crops or those of the neighbours, otherwise the owners
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would face a fine. Every year after San Laureano Day—July 4th in the Calendar of Saints
(santoral)—any unmarked animals grazing in vacant lots or on the ejidos were to be
collected and appropriated as strays, and then sold to cover city expenses. Owing to the
inconvenience of having livestock in the wetland preserve of the city, only those on land
neighboring the wetland were allowed to take their animals there, and those who were
not on neighboring land could only do so for a period no longer than ten days. The ejido
was set up to be used exclusively for domestic cattle, plow oxen and horses for service
that had been broken, while other livestock (cows, mares, sheep, goats and pigs) were
not allowed to graze there (Rojas, 1962). Specifically, it was ordered:
that, because of the many cattle that wander the ejidos, stripping the
land to the point where no forage is left for the horses nor to sustain them
such that they damage the crops of the natives and the crops of Spanish
that sustain this city and to remedy this we command that no neighbor
may have in the region of the ejidos of this city more than 400 head of
sheep and goats[…] and four head of cattle, or he will face a fine of ten
pesos of good gold […] “que ¿por cuanto los muchos ganados que andan
en los ejidos esquilman tanto la tierra que no se puede hallar yerba para
los caballos ni sustentarse los mismos en tanta manera que dañan a los
naturales sus labranzas y las sementeras de los españoles con que se
sustenta esta ciudad y para remedio de esto mandamos que ningún vecino
pueda tener en comarca de los ejidos de esta ciudad más de 400 cabezas
de ovejas y cabras […] y cuatro reses vacunas y no más, so pena de diez
pesos de buen oro […]” (Rojas, 1962).
Limiting the number of livestock head was thus a measure that guaranteed the
protection of the crops and the availability of grasses and, in consequence, the survival
of the activity of raising livestock itself. Even if these measures were ignored, their
existence reflects the concern of the colonial authorities that neither agriculture nor
future supplies for other activities (construction, raising livestock, mining…) be affected.
Moreover, in the expansion of raising livestock, the indigenous people were not
only passive observers affected by the invasive animals that belonged to their Spanish
neighbors. Patiño (1969) and Villamarín (1975), for example, affirm that well into the
colonial period the Indians did not have any horses, and raised the animals that they did
have on a very small scale. However, colonial documents indicate that, on the contrary,
economic organization and the process of enculturation and mestizaje (i.e., the mixing
of the indigenous and Spanish peoples) contributed to the rapid adoption of raising
livestock by the indigenous people, and the proliferation of the domesticated animals that
they had in their possession. In this regard, during the late 16th and early 17th centuries,
there were several references to the number of oxen, horses, sheep and chickens owned
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by the Indians of the Saquencipá Valley. Stallions and mares were often used to process
the wheat when oxen were scarce (AGI, Santafé, 56A, Nº17 (3), f.11r.) or to take flour to
Honda and Mariquita and, in some cases, were rented to neighbors in Santafé, Tunja and
Vélez (AGN, VB, T.10, f.492r.; T.18, f.530). The visit of the magistrate Luis Henríquez
in 1600 collected testimonies highlighting that, although the Indians of Sáchica planted
wheat, corn, potatoes and figs, and sold flour in Tunja and Santafé, they also specialized
in making halters and girths, many were muleteers, who rented mares and raised sheep
and goats for their livelihood (AGN, VB, T.18, f.528r., 530r., 536r., 570).
Since 1587 the Suta Indians had complained frequently because they had been
stripped of their land and the place they were assigned did not allow them to keep their
livestock (Colmenares, 1997). The visit to Suta in 1636, produced several testimonies
and petitions that emphasize livestock activities of the indigenous people of this
resguardo. The teniente de corregimiento (assistant to the Spanish mayor) stated that
although corn, wheat, potatoes and fruit and unspecified vegetables were grown, the
main activity was that of raising sheep, goats, chickens and, in greater numbers, horses
and oxen that they rented to their neighbors, mainly in Vélez. Both the protector de
naturales (the official responsible for the legal representation of the indigenous people)
and the main Indians interrogated in the secret inquiry agreed with this testimony, though
the number of animals reported was not unanimous, with figures of 300 to 400 oxen and
400 to 500 horses (AGN, VB, T.10, f.487v., 549r. and 622). These numbers demonstrate
the importance of livestock to the indigenous people in the region, though not all the
animals counted were there all the time because of the practice of renting them out7.
Although to a lesser degree than in Sáchica and Suta, it is possible to find noteworthy
data for other resguardos. Also remarkable was that the cacique of Tinjacá had, in 1607,
one estancia de ganado mayor (large-animal ranch) and four ranches with smaller
livestock (Ayape, 1965), and the testament of the Indian Esteban Castro, of Ráquira,
who in 1658 said he had six pairs of oxen, 25 mares and 300 sheep (Orbell, 1995). These
records provide evidence that for many indigenous people raising livestock, even the
large animals, was a privileged and lucrative economic activity that required pastures to
be delimited and generated conflicts with farmers, regardless of ethnic origin.
Along with the cattle and horses, other species did well in Tunja Province. The
spread of chickens resulted from their attractiveness (owing to their feathers or their
clucking), or because of the obligation of the indigenous people to supply the markets of
7 Although in the document there is only a textual reference to renting horses, oxen were also used as beasts
of burden, and in numbers that exceeded the fanegadas (capacity of the surface area) where they were
raised by the Indians themselves. This suggests that these animals also were hired to do agricultural work in
neighboring areas or other resguardos.
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the Spanish towns and ensure that the parish priest was provided with chickens or eggs8
(AGN, VB, T.5, f.408v.; T.7, f.591r.; T.14, f.910v.; T.18, f.834v., tribute fixed during
the visit of 1571 and 1572). This was a transference of the Iberian tradition of paying
tribute in chickens to the landlord or priest depending on the region (Carmona Ruíz,
1998). Poultry were fed whole corn and barley (Tovar, 1988) and raised in the open so
they could also eat insects, thus requiring little investment in their maintenance. This
way, the protein needs of not only the indigenous people were met, but also those of the
mestizos and whites who had fewer resources. Although the proliferation of poultry was
evident from the mid-16th century, the most striking numerical data is recorded in the tax
appraisal imposed by Juan de Valcárcel in 1636, listed in Table 1.
Tribute fixed by Juan de Valcárcel during his visit to the towns of Tunja Province in 1635
and 1636.
Tributo fijado por Juan de Valcárcel en su visita a las ciudades de la provincia de Tunja en 1635
y 1636.
Martín Niño
Pedro Merchán de Velasco
Luis Cárdenas
María de la Peña
Félix de la Serna Mujica
Juan Pérez de Salazar
Pedro Merchán de Velasco
Eugenia Alfonso de los Ángeles
Juan de Avendaño Maldonado
Juan Téllez de Mayorga
Pedro Venegas Torrijos
Number of
Indians taxed
Total number of
barnyard fowl**
1 300
1 846
3 198
1 183
Source: Prepared by the author based on data from AGN, VB, T.11, f. 1-341.
* Spaniard who was in charge of a group of indigenous.
** Approximate number obtained by multiplying the number of Indians taxed by the twelve hens and one rooster that
each was required to have at their home. In the region, this number initially rose to a total of 11.635 barnyard fowl, not
counting their offspring or the birds destined for sale or food.
These approximate data reflect the favorable reception by the Indians of poultry and
its spread, not only as a result of tax requirements, but also because of the low cost of
8 By 1583, when Juan Prieto de Orellana collected data on the tariffs that were charged in practice, the
indigenous people reported that they paid three hens and forty eggs to the doctrinero (priest) each week
(AGI, Santafé, 56A, Nº17 (3), f.15v.).
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maintaining them, their rapid reproduction rate and the incorporation of their meat and
eggs into the diet of the locals.
Although the accounting records of the 17th century indicate that paying tribute with
chickens was only constant in Suta, Sáchica and Chíquiza, interest in raising them did
not disappear. In the mid-18th century, Basilio Vicente de Oviedo observed that there was
no neighbor’s house, Spanish or Indian, where chickens were not raised in abundance,
and commonly traded by the poor, in particular by the Indians (“no hay casa de vecino,
sea español o de indio, en que no se críen con abundancia, y es el común trato y agencia
de los pobres, en particular de los indios”) (Oviedo, 1930).
In the Saquencipá Valley, in addition to barnyard fowl, other domesticated species
multiplied in only a few decades. Goats were important for making cordobán9 (the
leather that was made from their skin), as well as for milk and dried meat. They
reproduced quickly even in arid regions where there was little forage, feeding on waste
and thorns (Tovar, 1988). However, their presence in the region is not emphasized in the
colonial documents examined, which may indicate that raising them there was of less
importance than their abundance would indicate for the “lands of Chita, Chitagolo and
Zativa” (Zamora, 1945).
Sheep did well on the Altiplano Cundiboyacense and, although the situation in
New Granada is not comparable to the importance they attained in Mexico and Quito
(Patiño, 1969), their increase was favored by the climate, the herbaceous vegetation of
the region, and the desire of the indigenous people to pay tribute in blankets. According
to Basilio Vicente de Oviedo,
Countless sheep are bred in the cold lands, particularly in the
jurisdictions of Santafé and Tunja […] All of the fields are populated in
these sheep lands, with herds in the thousands. The Indians, in particular
those of the towns in the jurisdiction of Tunja, have very many, and with
their wool make great quantities of cloth, some of which they call ponchos,
others shirts, and still others blankets, that is their trade and business
for everything and for paying their high taxes “El ganado ovejuno no
tiene número el que se cría en las tierras frías, particularmente en las
jurisdicciones de Santafé y Tunja […] Todos los campos están poblados
en dichas tierras de ovejas, las manadas a miles. Los indios, en particular
de los pueblos de la jurisdicción de Tunja, tienen muchísimas, y con sus
lanas fabrican cantidades de mantas que llaman, unas ruanas, otras
camisetas, otras frazadas, que es su trato y comercio para todo y para
pagar sus crecidos tributos” (Oviedo, 1930)
9 According to the Relación de Tunja de 1610, the city tanned 4.000 hides each year (Patiño, 1983); the
number of animals slaughtered gives an idea of how many goats there were in the province.
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Although the author offers no numerical data, nor does he mention any towns for
their outstanding production, it is possible to establish that wool production was
important in Tunja and its surroundings. In the report or description of 1610, reference
was made to eight mills and five beaters10 (Patiño, 1969) and, owing to transportation
difficulties and the quality of the raw material mentioned by Oviedo, it is logical to think
that to obtain the wool sheep were raised in the areas close to the city such as Villa
de Leyva and its surroundings. However, as noted by Zamora, even in the cold lands,
the location of which he does not specify, haciendas with 2.000 to 3.000 head could be
found and, in the “sheep farms that were called de Suesca there were more than forty
thousand” (“Ovejeras que llaman de Suesca pasaban de quarenta mil”) (Zamora, 1945),
a number that could indicate that sheep were raised in the Saquencipá Valley in much
fewer numbers than in other locations on the Altiplano Cundiboyacense.
In fact, during the visit in 1583 of Juan Prieto de Orellana (AGI, Santafé, 56A,
Nº17 (3), f.10v-16v.), the captain of the Suta Indians complained about the lack of
wool for making mantas (cloth) that the Indians used to pay tribute even though their
encomendero was required to provide it, an affirmation that allows one to suppose that
for his subjects raising sheep was less important than raising horses or oxen, noted
above. In the 1636 visit this fact was expressed in the petition of the Monquirá and
Cucaita Indians to use cash to make the payment that was set in mantas by the tax rate,
given that their work on the estancias and on their own lands, and taking care of their
livestock left them no time to weave. They had no sheep, because there weren’t any, as
the land was not suitable for raising them, and when there were some, most that were
raised, died (por “no haber, como no hay, ovejas por no ser la tierra aparejada para ello
y si algunas hay y se crían se mueren las más”; AGN, VB, T.11, f.83v.). For the region
and the study period, no other references to the scarcity of sheep were found, but rather
their mention in the descriptions of the estancias and resguardos in Tinjacá, Ráquira and
Sáchica, allow for the conclusion that this was not a general problem.
Along with birds, goats and sheep, pigs were also being raised; especially for making
ham in Tunja (López de Velasco, 1971; Zamora, 1945; Patiño, 1969). According to
Fernández de Oviedo, more than three hundred head were taken to the Altiplano, all
females and pregnant (“mas de tresçientas cabezas, todas hembras y preñadas”;
Fernández de Oviedo y Valdés, 1852). Although that number may be an exaggeration,
sources concur in that from the time Tunja was founded in 1539 the pigs that
accompanied the conquerors multiplied quickly (Simón, 1981) and created problems for
10 According to the 22nd edition of the dictionary published by the Real Academia de la Lengua Española
obraje is the place where the cloth is worked (“lugar donde se labran los paños”) and the batán is a
machine, generally hydraulic, made of thick wood mallets, moved by an axis to beat, degrease and felt the
cloth (“máquina generalmente hidráulica, compuesta de gruesos mazos de madera, movidos por un eje,
para golpear, desengrasar y enfurtir los paños”).
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crop farmers and those rearing livestock. There is record of this even in the minutes
of the Town Council meeting held on August 14th, 1539, cited previously: when the
pasture area was delimited for horses, the warning was issued that no person should dare
to allow any of his pigs to walk in the specified place or cross the boundaries (“ninguna
persona sea osado de consentir que ningunos puercos suyos anden en el dicho lugar y
término”) (Ortega Ricaurte, 1941), and as a fine, the payment of two pesos of good gold
per head was set. On December 27, 1540 the Town Council broadened the measure and
forbade pigs from drinking in the fountain of the city and from walking in its plaza and
streets, setting a fine of four silver royals per head (Ortega Ricaurte, 1941). In the mid18th century, Oviedo called attention to the way in which the pigs brought to the New
Kingdom, had multiplied in such a way that there was no place or location that was
not full of pigs (“se ha multiplicado en tanta manera que no hay parte o lugar que no
esté lleno de puercos”) (Oviedo, 1930). Their reproduction was favored by the dietary
demand of the Spanish population and the fact that the animals could be maintained on
waste and abundant corn. When left free to roam they did more damage to crops and
ditches than other species did (Patiño, 1969), owing to their tendency to dig along walls,
around trees and planted areas.
In synthesis, it is possible to state that in the region of the Saquencipá Valley, the
practice of raising large and small species of livestock flourished, with particular
emphasis on bovine, equine, ovine, and porcine livestock along with barnyard fowl,
although on a smaller scale than in other regions of the Altiplano of modern day
Colombia. In spite of the impact that raising domesticated animals had on the ecosystem
and the economy by affecting crops, measures were taken so that their expansion would
not put the production and supply of food or raw materials in danger; however, in
practice, these were ignored or not enforced on land belonging to indigenous people and
by Spaniards who depended on animal husbandry.
The question then becomes, to what degree did the implementation of raising
livestock in the 16th and 17th centuries in the region generate or accelerate the processes
of deforestation and soil deterioration?
Trampling by ungulates and ruminants was not new to the land used as pastures
in the Saquencipá Valley, and the region was not without a millennia-long adaptation
between this type of animal and the herbaceous vegetation, a condition that has been
considered a Eurasian advantage (Crosby, 1972; Diamond, 1998). The abundance of
deer in the region, which continued until well into the 18th century, is an indication of
In any case, raising livestock in the region was of secondary importance compared
to agriculture. Although on different visits the Indians were reported to be raising a
variety of domesticated animals and some estancias were granted to the neighbors of
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Tunja and Villa de Leyva, the main activity of the inhabitants of the region was growing
wheat. On land that belonged to the Sáchica and Suta Indians, raising horses and cattle
was of greater importance and the livestock was rented out to other regions or used
in agricultural work. For the period studied and even currently, the number of head of
livestock in the Saquencipá Valley was far below the number that the land located in the
Bogotá savanna or the Ubaté-Chiquinquirá Valley could support; lands notable for their
deep, fertile soils.
In spite of being invasive species, the domesticated animals that were brought to the
region did not reproduce in an uncontrolled manner. Rather, they were faced with natural
controls that regulated their populations. For the Huasteca Potosina in Mexico, Miguel
Aguilar demonstrated how herd growth was regulated by natural factors such as attacks
by wild predators, the proliferation of insects and meteorological phenomena (Aguilar
Robledo, 1998). In the New Kingdom of Granada and particularly in the Tunja Province,
horses and cattle were reported to fall prey to bears, gatos bermejos (feline predators,
possibly jaguars), birds of prey, and the parasitic niguas (chiggers; Tunga penetrans)
(Patiño, 1983; Oviedo, 1930; Zamora, 1945). Barnyard fowl was devoured by some
type of fox, not specified in the literature, and by the common opossum (Didelphis
In the middle of the 16th century, specific areas of the region of the Saquencipá
Valley close to the steepest slopes but far from the rivers became eroded owing to the
biophysical conditions of the sites, such as geological origin, wind and rain patterns,
and this may have been accelerated by the loss of plant cover that accompanied the
farming and artisan activities of pre-Hispanic times. However, the majority of the
valley was notable for its fertility and deterioration was localized and prevented neither
settlements from being established nor agriculture. Rather, it offered suitable conditions
for obtaining food and raw materials for populating the area and those of other regions
in the New Kingdom of Granada.
The introduction of domesticated animals and livestock species undeniably modified
the ecosystem. Continuous trampling, the spread of disease, the consumption of grass
and the demand for cleared areas indeed occurred. In the study region however, cattle
ranching was less important than in other areas of the High Plain, such as the Bogotá
Savannah and the Ubaté Valley. Even with the biophysical conditions, which could
be considered more fragile in Saquencipá, resilience did not decrease and natural
11 A marsupial that inhabits some parts of the Colombian Andes and often feeds on barnyard fowl.
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mechanisms exerted control on the proliferation of these species, such as attack by
predators and parasites, or the decrease in food and water availability owing to drought.
Colonial society also managed to adapt to this new situation and designed or modified
the regulation of cattle ranching by periodically rounding up ownerless cattle, the design
and registry of branding irons, the delimitation and implementation of restrictions on the
use of ejidos, the construction of fences, ditches and corrals or hiring out the animals to
distant regions.
As a consequence, the adverse effects of cattle ranching on the ecosystem and
particularly on the soil were reduced and most likely were less severe than the effects
that other economic activities would have had in the region during the Colonial
period; especially mining and construction, which have yet to be studied in depth. The
deterioration of the soil that cattle ranching—as it was done then—could cause, did not
become a threat to the recovery capacity of the ecosystem. The descriptions indicate that
agricultural production was constant and there were no substantial changes in the quality
of the land during the study period.
This article is based on the research carried out by the first author for her Master
of Science degree in the Environment and Development program at the Universidad
Nacional de Colombia, Bogota, under the direction of Stefania Gallini. This research
was funded by the 2011 Graduate Thesis Support Program (Apoyo a tesis de posgrado,
Convocatoria 2011) of the Research Division of the Universidad Nacional de Colombia,
Bogota campus (DIB), for a travel grant to Seville (Spain). Bianca Delfosse translated
the text from the original in Spanish. The author is grateful to Stefania Gallini, her
thesis director; to the National General Archive of Bogota (Colombia); to the General
Archive of the Indies in Seville and for their comments on the first results of this study,
to Prof. Manuel González de Molina and the research associates of the Laboratorio de
Historia de los Agroecosistemas at the Universidad Pablo de Olavide in Seville for their
useful advice, and to the geographer Alfonso Simbaqueba for his expertise using ArcGIS
10 software and his generous collaboration. Many thanks to Claudia Medina Uribe,
Director of Biological Collections at the A. Von Humboldt Institute of Research on
Biological Resources, Villa de Leyva (Colombia), for help with the scientific binomials
of the common names of these plant species.
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Colonial livestock farming in Saquencipá Valley
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El presente documento reúne los resultados de una investigación que tuvo como
objetivo analizar la transformación en las prácticas ganaderas durante el periodo
colonial, su relación con la tenencia de la tierra y el impacto que estas tuvieron en el
ecosistema manifestado en la degradación de suelos, en el contexto específico del Valle
de Saquencipá en el altiplano cundiboyacense de la actual Colombia. A través de la
revisión exhaustiva de documentos coloniales en los fondos del Archivo General de la
Nación en Bogotá y al Archivo General de Indias en Sevilla, de las crónicas y relaciones
geográficas y de los resultados arrojados por investigaciones desde la historia y la
arqueología, se reconstruyen las prácticas ganaderas durante los siglos XVI y XVII y
sus antecedentes ibéricos. Teniendo en cuenta factores como la complementariedad con
la agricultura, la movilidad del ganado, el número de cabezas y el control natural de las
sequías o los depredadores, se cuestiona el alto impacto ambiental que se le ha atribuido
a la actividad ganadera colonial en la región.
Palabras clave: Historia ambiental, colonia, ganadería, degradación de suelos.
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Boulevard del Niño Poblano 2901. U. Territorial Atlixcáyotl.CP. 72197 Puebla, Puebla, México.
[email protected], [email protected]
The development of animal husbandry in the northern region of the state of Veracruz,
known as the Totonacapan, produced major environmental changes. The highlights
of this development are the introduction of cattle in the sixteenth century and African
grasses in the second half of the nineteenth century. Environmental changes rapidly
ensued between the years of 1940 and 1970 when livestock operations were intensified.
The analysis of three case studies allowed us to estimate the impact that livestock
intensification initiatives had across the state and to identify elements that would allow
for a sustainable model of cattle ranching in this region.
Key words: Sustainability, deforestation, land use.
Following the appearance of the book, Plague of Sheep, by the American historian
Ellinor Melville who argued that the arrival of ungulates had generated severe
environment degradation in the Valle del Mezquital, Mexico (Melville, 1994), the idea
was generalized among some historians that the arrival of cattle to the Americas had an
overall devastating effect (McClung and Sugiyama, 2012). Others, such as B. Turner
and K. Butzer, argued that the presence of the cattle had few or only moderately negative
consequences (Sluyter, 2001). This divergence of opinion may also be witnessed in
several studies highlighted in the book Historia Ambiental de la Ganadería en México
(The Environmental History of Cattle in Mexico). In the introduction, Sergio Guevara
(2001) discusses the idea, as proposed by many ecologists, that cattle ranching has
resulted in a large-scale disturbance of ecosystems. According to this argument, cattle
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
PASTOS, 42 (2), 273 - 297
B. Ortiz and R. Jiménez
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The Impact of Raising Cattle in the Totonacapan
ranching would have led to the demise of native species, the invasion of exotic species,
and changes in the physical structure of soil and its fertility. However, the author also
mentions that the impact of livestock ranching has been variable and depends on the
type of animal being raised, whether cows, pigs, or goats. Guevara recognizes, as well
as other authors, that the introduction of cattle to the Americas by the Spanish settlers
was part of the colonial inheritance, as many Spaniards came from towns where the
mixed agriculture and grazing were practiced. Even so, the introduction of cattle would
be “violent” and its impact would change the environment drastically, particularly due
to the speed in which herds were reproduced. Every 15 years the herds were doubled,
especially in the arid and semi-arid highlands, which contributed, among other things,
to the modification of the original landscape that harbored only small remnants of native
vegetation. The alterations in the landscape provoked accelerated erosion processes, a
higher incidence of flooding, and a loss of crops due to changes in the flora and fauna.
Thus, the author concludes that the introduction of cattle was a transforming agent of the
American territory, as well as a determining factor in the modification of the agricultural
and natural landscape and the state in which it is found today (Guevara, 2001).
Miguel Aguilar (2001) pointed out that in the Huasteca Potosina region an explosive
growth in the cattle was experienced beginning in the second half of the sixteenth
century and continuing until the early decades of the seventeenth century, to the point
that the documented sources mention that large herds of between 468 000 and 500 000
heads of sheep grazed in Valles and Tanchipa. The rapid growth was explained by the
abundance of pastures and the relative lack of competition and predators, as well as the
demographic decrease of native populations, which diminished the pressure on some
ecosystems and made additional lands available for cattle ranching. Despite the large
herds that grazed the Huasteca, the author proposes that the environmental impact of
the cattle was in fact moderate, explained by the complementary way in which cattle
ranching and agriculture were cyclically interchanged in the same fields. This process
was adapted and adjusted to the specific conditions of the new colony. Thus, in the
region of the Huasteca, agricultural fields were left fallow and grasslands allowed to
recover, where large cattle grazed freely and were seasonally interchanged with smaller
cattle (Aguilar, 2001). Andrew Sluyter came to a similar conclusion, where in his study
on the tropical lowlands in Veracruz he establishes that despite the pressure that livestock
placed on the land’s carrying capacity during the colonial period, the vegetation cover
and soil remained stable, in contrast with evidence of destabilization that occurred in
the Pre-hispanic period as caused by population growth, deforestation by slash and
burn farming, and intensification of land use that provoked landslides and erosion. In
agreement with Miguel Aguilar, Sluyter also considers that the Spanish mode of cattle
ranching was environmentally sustainable, due to the fact that the system of nomadic
herding limited the possibility of overgrazing by allowing the movement of semi-feral
creole cattle across the landscape between the humid and dry savannas (Sluyter, 2001).
PASTOS 2012. ISSN: 0210-1270
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The Impact of Raising Cattle in the Totonacapan
While many hold the opinion that the environmental consequences of cattle ranching
were minimal to moderate, at least during colonial times, the debate on the impact of the
introduction of cattle in New Spain continues. There is some consensus surrounding the
idea that cattle ranching would be modified and categorically distinct in the second half
of the nineteenth century and the first decades of the twentieth century, especially in the
warm-humid regions during what has been named the “Silent Revolution,” a time period
in which the establishment of pasturelands and the introduction of “artificial” African
grasses were increasingly prevalent. The Spanish varieties of cattle were disappeared
to make way for new species, leading to the precedence of grasslands at the expense
of natural vegetation and high rates of deforestation in various regions of the country
(Aguilar, 2001; Sluyter, 2001). The presence of a high density of cattle modified
previous land uses, and management of the pastures allowed for the establishment of
exotic species from Africa and Asia. The simplification of the associated habitats and
the presence of an extensive and uniform system of pastures permitted an easy and
rapid expansion of exotic, secondary, and ruderal species. Sergio Guevara considers
that it is necessary to conduct more studies on the environmental history of cattle
ranching in order to understand and situate ranching with the context of the current
state of the Mexican ecosystems through evaluating current management practices and
performance of ranching systems, as well as their environmental, economic, social, and
cultural impacts. The lack of knowledge of cattle behavior, according to the author, has
contributed to a lack of understanding of the ecological impact of cattle ranching and has
limited the possibilities and alternatives for its rational management (Guevara, 2001).
The present investigation has the objective of bringing to light the impact of cattle
ranching in the Totonacapan region of Veracruz, that in addition to other regions of
Mexico, have suffered environmental changes because of the introduction of cattle
and African pastures during the second half of the nineteenth century. Such changes
were particularly accentuated and accelerated between the years of 1940 and 1979.
In the first section of this investigation, a general historical revision of the region is
elaborated, focusing on relevant land uses employed until the mid-twentieth century. It
is important to highlight that during the colonial times special authorizations for the use
and exploitation of the land were given by the Spanish crown, called “mercedes reales”,
and such land grants often established cattle ranches at the expense of the indigenous
groups that sought to preserve their communal properties. This was the overarching
scenario until the liberal land reforms modified land status in the 20th century with the
goal of modernizing the Mexican territory. In the second section of this investigation,
several case studies are presented for the Totonacapan region and other sites in the state
of Veracruz, detailing how the cattle ranching initiatives and their intensification have
unraveled. The goal is to determine more suitable land uses and management of cattle
ranching regions. The third section of this investigation establishes recommendations
oriented towards concrete actions that promote sustainability, highlighting how cattle
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ranching must be understood within a historical context so that a new model may be
established, not only for the Totonacapan region but for the entire state of Veracruz.
During pre-Hispanic times, the Totonacapan included a vast region delimited to the
north by the Cazones River, to the south by the La Antigua River, to the west by the
mountain range of the Sierra Madre Oriental, and to the east by the Gulf of Mexico.
The largest populations were established in the northwest and southwestern areas,
meanwhile the central region had the lowest population density. Two overarching zones
have been identified: the “Llanura Costera,” or the coastal plains, and the “Sierra of
Papantla,” or the mountain range of Papantla (Ortiz, 1995; Velázquez, 1995; Velázquez,
1996; Chenaut, 1996). After the Spanish conquest, 1,434 mercedes, or land grants,
were given throughout the Totonacapan region. A total of 1383 grants were given to
Spaniards and merely 51 grants to indigenous peoples. Benjamin Ortiz identified that
53% of the granted territory was used for haciendas, or ranches, where the manual labor
of indigenous populations was exploited, 27% of the land was dedicated to raising large
cattle, and 19% for small cattle. Most of the haciendas extended throughout regions that
currently form the municipalities of Jalapa, Misantla and Cempoala. The coastal plains
and the mountain range of Papantla had the fewest cattle concessions (Ortiz, 1995,
Velázquez, 1995). This information can be corroborated in the Relación de Hueytlapa
y su partido (The History of Hueytlapa), published in 1580 and written by Joseph
Velázquez. According to this document, in the Hueytlalpa region of Veracruz cattle
ranching did not take precedence over other activities because grass would not grow
due to the “many stones that can be found there.” The same situation that was seen in
Jujupango, which was not hospitable to grass for grazing either, because of the “many
abysses and landslides.” Meanwhile in localities such as Matatlan and Chila grass was
not grown due to the “thickness of the trees and the interwoven branches” (Velázquez,
A situation to the contrary occurred in Zacatlán, where there was “a lot of grass”
owing to the abundance of water and the presence of lagoons, which formed a propitious
environment for the raising of cows, lambs, goats, and pigs. In the region of Papantla,
the existence of grass was mentioned by Velázquez (1985), and although he noted
that many lands had been granted, only the large cattle ranches of Diego Cepeda
and Diego Larios were mentioned, both of whom lived in Mexico City and whose
properties were located along the rivers of San Pedro and San Pablo (nowadays known
as Tecolutla River) (Velázquez, 1985). Despite the agricultural potential that the region
demonstrated, extensive cattle operations would remain as the main motor of the local
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economy. At the end of the eighteenth century fourteen large cattle ranches existed in
the region of Papantla (Ortiz, 1990; Ortiz, 1995; Velázquez, 1995). The presence of the
cattle generated problems with the indigenous populations, due the incursions of cattle
in their fields. Although their complaints were formalized and the colonial government
notified of their grievances, this conflict eventually provoked an indigenous insurgence
in 1836 lead by Mariano Olarte1. It is necessary to clarify that issues of land invasion
did not motivate the rebellion, because in the decade following 1839 the indigenous
populations located in the coastal plains and the mountain range of Papantla held over
24 000 hectares (ha) where forest predominated, including 200 ha of corn fields, vanilla,
and grassland savannas (Escobar, 1996; Blanco 1996).
During the decade of 1859 to 1869, vanilla plants were largely replaced by citrus
trees and grasses for fattening cattle. The precedence of indigenous communal property,
at least until the decade of 1870, had prevented the establishment of large haciendas in
the region. Emilia Velazquez (1995) mentions that while more than 20 ranches existed
in the area, only three had large extensions: Larios and Malpica in Tecolutla with a
total of 33 017 ha, Palma Sola with 24 270 ha, and San Miguel del Rincon with 21
880 ha. Most of the haciendas were dedicated to growing vanilla, tobacco, sugar cane,
bananas, corn, beans, and chili and the exploitation of rubber and the chicozapote fruit,
in addition to cattle ranching (Velazquez, 1995; Ortiz, 1995). The encumbrance policies
of liberal governments seeking to transform the rural people into small land-holders
would face a stubborn resistance in the region. According to Victoria Cheanut (1996),
the opposition of the Totonacan people to the division of their land found its basis in
the wish to preserve and reproduce their identity as an indigenous group. Facing the
possibility of an insurgence with weapons, the authorities determined, between 1875 and
1878, that the land would be divided into 25 large portions. However, in 1891 another
insurgence with weapons occurred which caused in response, between 1893 and 1898,
that the authorities would create smaller portions of land, divided into individual lots that
would be sold as private property. The division of land did have the expected results and
was not egalitarian, because in localities such as Coxquihui, Chumatlán, and Zozocolco,
a total seven families were bequeathed with 50% of the divided land (Blanco, 1996;
Chenaut, 1996).
In the case of Coxquihui three haciendas were established and dedicated to vanilla
and tobacco cropping and cattle ranching. One of the main consequences of the change
in the land ownership, which shifted from communal ownership to small holders, is
that indigenous groups lost control of natural resources. By 1905, in Papantla Cantón,
34 haciendas were counted with an extension of 154 000 ha, among them particularly
1 Una de las peticiones de Olarte para acabar con el conflicto fue que el gobierno dispusiera lo necesario para
evitar que el ganado invadiera los terrenos de los indígenas.
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eminent were Larios y Malpica, which possessed 33 000 ha, San Miguel del Rincon
with 24 000 ha, San Miguel el Grande with 20 000 ha, and San Lorenzo Palma Sola
with 20 000 ha. The distribution of the communal land constitutes a key element in
understanding the subsequent transformations that followed in the Totonacan landscape
(Ortiz, 1990; Velázquez, 1996). Between 1910 and 1930 the deforestation process
would begin in the region, among other things, due to the discovery of oilfields and the
construction of a railway (Ortiz, 1995). In the decade of 1949 there was a resurgence of
cattle ranching, stimulated by the construction of a road from Tuxpan to Mexico City,
which allowed faster transport of the animals. Before the existence of this road, the cattle
ranchers had to bring their herds from the train stations located in Veracruz, Tampico,
Huauchinango, or Teziutlán, a trip that could last from one to two weeks in which the
animals would incur a significant weight loss (Velazquez, 1996). Between 1940 and
1970 important land use changes were beginning to transpire in the Totonacapan, as the
land was increasingly dedicated to the growing of coffee, citrus, and bananas, and the
clearing of fields also facilitated the introduction of cattle ranching, allowing them to
trample unchecked across the landscape (Ortiz, 1995; Chenaut, 1996).
Benjamín Ortiz (1995) mentions that by 1959 there were 156 005 ha of forest in the
region, but in 1970 only 47 485 ha were registered. In contrast, the area of grassland
incremented from 140 852 ha in 1959 to 216 807 ha in 1969, 309 079 ha in 1970, and
362 108 ha in 1984, which represented a total increment of 157% due to increases in
the extension of cattle ranching (Ortiz, 1990), which converted itself into the most
fundamental activity for rural families, surpassing the supremacy of cultivation of corn
(Velázquez, 1996; Lazos, 2001). Testimony of a rural farmer from “Plan de Palmar”
indicates that while he was able to achieve two regular corn harvests per year, the
uncertainty of the harvests converted him into a rancher, andhe began to grow grass in
pasturelands. Another farmer, further testament to the transition, only cultivated one
hectare of grass but later decided convert his entire property of 5 ha to pastureland. In
this last case, the rural farmer had obtained his land in 1968 and made a full conversion
to cattle ranching within three years, introducing up to four different kinds of grass:
guinea, angola, merqueron, and African star. Another study conducted in Plan del
Palmar showed that another land owner began to substitute interchangeably grass and
corn. When the production cycle was finished and corn was harvested, part of the field
was often sectioned off to sow grass for pasturelands.
Between 1968 and 1992 there was a decrease in the surface area of native vegetation
and other traditional land uses and a corresponding increment of the total area dedicated
to pasture. Thus, traditional means of land management by rural Totonacan communities
were increasingly replaced by cattle ranching, although often interchangeably. Their
agricultural fields, or agro-forest units, were harvested and alternated in a cyclical
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manner with those of pastures, which allowed for more diversification and complex
vegetation mosaics. However, since the most recent introduction and extension of cattle
ranching throughout the zone, new management practices have begun to take hold,
and semi-intensive cattle ranching has risen in prominence. With this method pasture
extensions averaged from three to five ha with five to seven cows per ha, where cows
are sometimes rotated between fields. In cyclical systems a high productivity may be
maintained due to the rotation of land uses, the presence of diversified grasslands, and a
high density of organisms that maintain the fertility of the soil (Ortiz, 1995; Ortiz, 2001).
A characterization of three identified cattle ranching systems, or initiatives, was
carried out by analyzing several indicators, based on the modification of a proposal made
by Astier and Masera to evaluate the sustainability of the primary productive systems
(see Table 1.1) (Astier and Masera, 1997).
The following indicators were taken into consideration:
1. Ecological indicators
a) Richness of species in pasture (flora and fauna).
b) Richness of species in the soil (particularly earthworms).
c) Number of productive activities associated with the fattening of cattle.
2. Technological indicators
a) Capacity of animal load, expressed in units of animals per hectare (U.A./ha),
where 1 U.A. is equivalent to a grown adult cow of up to 450 kg (De Alba, 1980).
b)Energetic productivity expressed in kilocalories (the energetic productivity
corresponds to the relationship between the energy inverted in the system and the
energy obtained). For this calculation it was considered that for every kilogram
of balanced food entering the system, 3300 kilocalories can be generated
(Koeslag, 1982). By this calculation, a day´s production is equivalent to 4200
kilocalories (INN, 1990) and the 56% of the living weight of an animal of 450
kg was considered as useful meat and bone for the human use (Williamson
y Payne, 1975). A liter of fresh milk represents 680 calories (INN, 1990). For
the conversion of electric energy to kilocalorie the conversion factor of 1 watt=
0.2983Kcal/seg (Holliday et al., 1990) was used and for the animal traction,
15,000 Kcal/day (Koeslag, 1982).
c) Work productivity (quantity of a day´s wage/hectare/year).
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Social, environmental, and economic features of three systems of livestock intensification.
Características sociales, ambientales y económicas de los tres sistemas de intensificación de la
Type of Tenure
Total Surface (ha)
Antiquity (years)
Number of Beneficiaries
Division of Useful Species
Number of Acts of Solidarity
Annual Average Precipitation (mm)
Annual Average Temperature (°C)
Dominant Soil Type
Altitude (m above sea level)
Richness Above Ground (species)
Richness Below Ground (worms)
Type of Grass
Type of Production
Rotation Days
Stocking (U.A. ha-1)
Pasture Size (ha)
Annual Production Cows
Annual Production Milk
Productive Divisions
Gross Profit ha-1 year-1
Net Profit ha-1 year-1
Wages (W) ha-1 year-1
Energy Efficiency
Economic Efficiency ($/W)
Private property
Associated ejido
Dual purpose
109500 L
Dual purpose
8640 L
24 hours
$ 3270
$ 2598
$ 1664
$ 1489
$ 9818
$ 3435
3. Economic Indicators
a) Cost of production (day´s wage, food, medicines, vitamins, deparasiting, removal
of ticks, vaccines).
b) Expenses (administration, sales, and financial expenses).
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c) Gross profit (all economic inputs from the sale of the products such as milk,
calves, and cull cows).
d) Net profits (total income minus expenses and production costs).
4. Cultural indicators
a) Number of useful species in the pasture recognized by the ranchers.
b) Number of persons directly benefitted by the productive cattle ranching unit.
c) Number of non-profitable activities that are part of the productive cattle ranching
process, indicating solidarity of the group.
To obtain a sustainability indicator for the cattle ranching operations, 12 indicators
were applied to the three separate initiatives or types of cattle ranching operations:
indigenous, rancher, and technical. A range of categories (high, medium and low) were
assigned and standardized for all of the indicators, so that all of indicators would be
equally weighted. A score of 3 was assigned to the high category, 2 to the intermediate
category, and 1 to the low category, where the highest scores indicate the best fit to the
indicator. A matrix with columns may be obtained for each initiative where values for
the total number of indicators are later summed. Thus the index values of sustainable
cattle ranching oscillate between 1 and 3. These indicators are visually displayed by
“pathogenic amoeba” graphs in order to compare all the indicators that influence the
degree of sustainability of a grazing system (see Figures 1, 2 and 3).
From the point of view of richness of vegetal species, the indigenous and rancher
initiatives presented the highest values with 50 and 34 species, respectively. For the
technical initiative, due to the specialized management and extensive coverage of the
utilized Taiwan grass, only 3 species were recorded. In regard to the species richness
of earthworms, the technical initiative presented the highest value with 8 species
found, followed by the indigenous with 6 species and the rancher with 2. The number
of productive activities integrated with pastures also varied, where the initiative of
indigenous cattle ranching is associated with the rotation of land uses, such as the
cultivation of vanilla and corn plants, sowing of tall grass, and milk production. The
rancher initiative of grazing is complemented with milk production, while the efforts
of the technical initiative system are directed exclusively to the fattening of the cattle.
The capacity of the animal load presented the highest values in the technical system
(5.5 animal units per hectare), while the indigenous initiative as well as the ranchers
presented a value of 1.5 units of animals per hectare. With respect to energy productivity
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the highest values, which represent a better efficiency in the conversion of energy, were
found with the technical and indigenous initiatives, which held a productivity index of
4.66 and 3.84 respectively, while the ranching system presented a negative energetic
balance of 0.70, meaning that more energy is used than obtained.
Upon examining agro-ecosystems from the perspective of environmental fragility
and resilience (Toledo, 1996), they may be characterized based on their capacity to
persist in the face of varied or unavoidable disturbances, both physical, biological,
and socio-economic. The three initiatives of technical, indigenous, and rancher present
different degrees of resilience and distinct strengths and weaknesses. The rancher
initiative presents a scenario of growing fragility, because of its high dependence on
external inputs. In spite of the fact that this system presents a net profitableness per
hectare, there is also a corresponding lack of ecological efficiency (Table 1.2). Ranching
also received the lowest score in species richness. The technical initiative is a scenario
of highly specialized production yet at the time offers high ecological efficiency and
the highest profitableness. However, there is also an associated loss in the biological
richness of the soil. On the other hand, what stands out about the technical system is
its high species richness of earthworms, the highest of the group, despite the strong
disturbance that the field suffered at the time when irrigation systems were introduced.
Energy analysis of the three systems of livestock intensification.
Análisis de la energía en los tres sistemas de intensificación ganadera.
Meal (kcal ha-1 year-1)
Milk (kcal ha-1 year-1)
Total Gain (G) (kcal ha-1 year-1)
Wages (kcal ha-1 year-1)
Balanced Food (kcal ha-1 year-1)
Electricity (kcal ha-1 year-1)
Animal Traction (kcal ha-1 year-1)
Total Expenditure (E) (kcal ha-1 year-1)
Energy Efficiency (G/E)
326 592
744 600
1 071 192
92 400
1 445 400
1 537 800
241 920
24 480
266 400
69 300
69 300
2 969 018
2 969 018
138 600
497 727
636 779
The indigenous farming system presents, curiously, an energetic conversion
that is similar to technical initiative and does not require a large investment in day´s
wages and money. In addition, the indigenous strategy conserves the highest number
of species in the soil. However, it is also the least profitable of the group. While the
results of the present study indicate that the technical initiative is the most profitable,
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from the environmental and cultural point of view the indigenous initiative is the most
resilient, as it implements a diversified production strategy. Such a strategy is important
to maintaining biodiversity, in this case under a regime of communal management,
and the sustainability of this system is reflected in the global index of cattle ranching
sustainability. Although the rancher and technical initiatives managed to obtain the
highest incomes, this occurred at the expense of the total elimination of forest cover
(Table 1.3).
These findings turn the attention toward a search for new models of raising beef cattle
in the Mexican tropics that would manage to incorporate the best of the three initiatives,
such as an intensification in production (technical initiative) and the preservation
and promotion of the above and below ground biodiversity (indigenous initiative).
While perhaps cattle ranching had previously been considered as unsustainable, these
reflections lead the discussion in a new direction that has not yet been addressed in
Mexico: the possibility of sustainable cattle ranching.
Sustainability indexes for the three systems of livestock intensification.
Índices de sostenibilidad en los tres sistemas de intensificación ganadera.
Species Richness Above Ground (Animal and vegetable)
Species Richness Below Ground
Activities Associated with Livestock Production
Stocking (Number of cattle per pasture)
Energy Production
Production Costs
Gross Profit
Net Profit
Wages (kcal ha-1 year-1)
Useful Species Recognized
Solidarity activities
Livestock Sustainability Index
1=Low, 2=Medium, 3= High
Rancher Indigenous
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The history of land use in Mexico and particularly in the Mexican tropics has been
characterized in the second half of the twentieth century by a massive deforestation
that has affected more than 50% of the total surface area of the country (Toledo, 1990;
Masera, 1995) and 91% of the surface in the state of Veracruz (Ordoñez and García,
1992). The grand extensions of cattle ranching may be explained by the fact that most
grazing systems practiced in Mexico are extensive and have made little use of agrotechnology to intensify production, as only a 5% of the total land dedicated to ranching
makes use of intensive cattle ranching techniques (Toledo, 1990). Free grazing requires
large extensions of large yet offers a very low productivity, which varies according to
the region. The carrying capacity of pastures in Mexico range from 0.8 ha per animal
per year in the tropics up to 50 ha per animal per year in arid zones. This section seeks
to evaluate, from the point of view of energy usage and economic profitability, some
initiatives that could potentially intensify cattle ranching over a smaller surface area and
potentially allow for the reforestation of the Veracruz tropics. While the production of
cattle has been one of the most important productive activities in tropical countries, few
technical or economic efforts to increase animal productivity have been successfully
achieved (Preston and Leng, 1987; Preston and Murgueito, 1992). This may be explained
by the absence of frames of reference to understanding the ecological, socio-economic,
and cultural impacts and limitations of the region and, at the same time, a failure to
recognize the enormous potential for a sustainable cattle ranching initiatives.
In this sense, other authors (Toledo, 1992; Serrano and Toledo, 1990) have
suggested that a sustainable system of a cattle ranching is determined by exogenous and
endogenous factors. Exogenous factors refer to the natural quality of the environment
(soil and weather), security in land ownership, the existence of tax incentives, soft loans,
and market demand. Endogenous factors deal with the quality of the germoplasm used,
taking into account the relatively low productivity in the tropics where operations are
unable to finance expensive modifications to the environment. Thus the efficiency of
grazing, the optimization of local fodder resources, and the use of economically viable
technology that would improve the venture’s competitiveness in the market are key.
The management of grazing requires a clear vision and knowledge of the relationship
between over and under-grazing, and the rancher must be aware of the most efficient
management options. Therefore, the principal intensification strategies for cattle
ranching through sustainable grazing may be grouped in the following categories:
1. Agrosilvopastoral systems, which consist in the spatial and temporal association
of forested areas, systems of grazing, and the annual cultivation of crops. This
strategy represents a promising alternative for restoring degraded pastures, as
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well as a means of management in a family and community-oriented economy
(Hecht, 1982).
2. Grazing systems of fast rotation, which is characterized by management
practices where the cattle are moved from pasture to pasture and allowed to graze
in each for a determined length of time. The pastures may be grazed for a few
days or only a few hours, and they are subsequently left to rest according to the
rate of recuperation of the grass (Savory, 1988;Voisin, 1974; Canudas, 1995).
3. Integrated grazing systems, which are grazing systems that are coupled with the
organic cropping of sugar cane, cereals, and plantains, which are also commonly
used as fodder for the cattle (Preston and Murgueito, 1992).
The mentioned grazing systems propose an innovation in management over the
current extensive and specialized models of cattle ranching, representing a shift to a
semi-intensive method of ranching that is based on the efficient use of the available, local
biological resources. Thereby ranching may be integrated as part of a more inclusive
system of rural production. This alternative model is directed toward the creation of
mosaics of productive spaces composed of annual and commercial agricultural crops,
managed forest, and controlled grazing.
Initiatives to intensify the production of cattle ranching in Veracruz have resulted
from a general economic crisis in rural areas due to current national economic policy. The
liberation and opening of the markets to the exterior, including meat and milk products,
has created a decrease in the demand and consumption of local beef (Suárez and López,
1998). This economic model, stemming from the 1990s, has lead both cattle ranchers
and various researchers and technical assistants from academic and governmental
institutions to search for alternative methods that would allow cattle ranching and meat
production to become more efficient and competitive in the broader market. Three kinds
of alternative management approaches for cattle ranching in Veracruz are presented.
The first management alternative, hereafter referred to as technical (Figure 1),
corresponds with proposals on behalf of a the National Institute of Forest, Agricultural,
and Livestock Investigations (Instituto Nacional de Investigaciones Forestales,
Agrícolas y Pecuarias or INIFAP), which holds experimental agricultural fields and
pastures in La Posta and Medellin de Bravo, Veracruz. Several lines of research have
been development, involving the generic improvement of cattle breeds that would have
more than one purpose, the construction of silages to preserve fodder, the creation of
protein banks or the provision of high quality proteins to cattle, and the management
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of grasses through intensive rotational grazing. At the Center for Cattle Research of
the National Autonomous University of Mexico, located in El Clarin, Martinez de la
Torre, Veracruz, training for cattle ranchers in the intensive grazing techniques is
offered, developed based upon previous research (Castillo, 1995). A researcher from the
INIFAP made a production agreement with local communal land holders from Lomas
del Porvenir, Medellin de Bravo, Veracruz, establishing in 1994 a production system to
aid in the fattening of young cattle by quickly rotating them through pastures, as well as
the implementation of a drip water irrigation system to speed the growth of grasses. The
land had a total surface area of 11 ha, where 1.8 ha are dedicated to sugar cane cropping
as a food supplement for the cattle. Waste from sliced bread was also incorporated into
the cows’ diet. However, in order to implement this system it was necessary to remove
all vegetation cover, and heavy machinery was used dig over 30 cm deep, bury the
irrigation lines, and connect the lines to the main water source. The entire surface area
was divided into 46 smaller pastures of 2000 m2, each sown with Taiwan grass (Cynodon
plectostachyus). The entire herd, between 40 and 50 young cows, were grazed on one
section of land during a period of 24 hours before they are rotated to the next section.
Such an example represents a technical alternative that has been implemented through
extensive research.
Technical management of the farm.
Manejo tecnificado del potrero.
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The second management alternative, or the ranching alternative (Figure 2), combines
the personal initiatives and experience of private ranchers with new strategies. One
example of this is a dual production system, located in Vega de Alatorre, Veracruz,
which combines agricultural cropping and cattle ranching. In this example, the ranch
belongs to a family of third-generation ranchers. In 1987 the family adopted a rotation
system for their cattle and electric fencing in order to manage them. Such a system may
be referred to as a slow rotation system, as cattle are rotated every 1.7 days in a space
of 2 to 3 ha. This technique was developed by the previously mentioned Center for
Cattle Research of the National Autonomous University of Mexico, located in El Clarín,
Martínez de la Torre, Veracruz. As in the previous initiative, living fences or borders of
trees that once delimited pastures are replaced in favor of electric fences, where trees are
completed cleared from the pastures. In this system the diet and growth of the cattle is
often balanced or supplemented by commercial crops.
Rancher management of the farm (conventional).
Manejo ranchero del potrero (convencional).
The third management strategy, or indigenous initiative (Figure 3), corresponds
to ranchers with communal land holdings whom have integrated the management of
cattle with that of agro-forest systems, resulting a mosaic of productive spaces (Ortiz
and Toledo, 1998, Ortiz, 1995). An example of this kind of management has been
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implemented by a Totonacan indigenous communal land owner, 50 years in age, and
his 17 year-old son. The area of pasture is located next to areas of rain forest, vanilla,
and corn plants, forming a diversified, agrosilvopastoral mosaic. This configuration
of productive spaces nestled within forested areas is the result of a long-standing
Mesoamerican tradition of natural resource management, which has been conserved by
the communal land owners of Plan de Hidalgo, Veracruz since its founding in 1968.
In this system the pastures are abandoned after a period of approximately 20 years,
after which a regeneration of the tropical rainforest ensues. The regenerating rain forest
has various uses to the rancher family, such as providing sources of fuel, construction
materials, and areas for hunting and collecting useful wild plants. After a maximum
of 10 years, these sections of land are re-used for the annual sowing of crops during
a period of 4 to 5 years, after which the land is converted into pasture. This grazing
system that was developed for slow rotations, where the grazing period lasts for 15 days
in pastures of 4000 m2.
Indigenous management of the farm.
Manejo indígena del potrero
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The economic and political scenario surrounding land use in Latin America is
changing due to the crisis of the traditional mode of cattle ranching, provoked by the
decrease in meat prices, the elimination of subsidies for ranchers, the degradation of
pastures by overgrazing, soil erosion due to inadequate pasture management practices
over inclined terrain, and the exhaustion of the availability of cheap land. On the other
hand, new policies have emerged promoting the reforestation of areas previously
without trees. Such factors will influence decisively in the future management of
grasslands, as the livestock sector must develop new productive strategies. The stimulus
of wood production is a potential opportunity for the conversion of large, uniform areas
of pastures of crops, where the planting of trees may represent future commercial value.
Due to low profits, a spontaneous reforestation of abandoned pastures in beginning to
take hold, where regenerating species of high commercial value may be taken advantage
of. Although for a long time researchers and planners have ignored the phenomenon
surrounding the regeneration of native vegetation in pasturelands, it is well-known to
many ranchers and farmers whom have taken advantage of these processes in the past.
The search for a sustainable cattle ranching includes, from our point of view,
defining a new concept of cattle ranching in the tropics. The management must change
from uniform pasturelands and specialized management to one that takes advantage
of new market opportunities in the preservation of biodiversity (rancher and technical
initiatives). Such models would be more flexible, and with the correct use of natural
resources and biodiversity cattle ranching may become compatible with the natural
environment. The nature of indigenous and peasant farmers, fisherman, and foresters
in these environs has been one of duplicity, in the sense that many have traditionally
dedicated themselves to a variety of productive and cattle ranching activities. Supporting
this mode of farming could uphold a new model of sustainable development, where not
only specialized production projects are supported and financed, but rather projects that
would support multiple economic endeavors of the farmers. Thus, the sustainable future
of cattle is dependent upon an integral management of every aspect of the economy of
rural farmers, under a policy that would promote the rational use of natural resources
(indigenous initiative).
There has been important progress in the management scheme of intensified grazing,
which has demonstrated the possibility of increasing up to 10 times the animal load per
hectare (technical initiative). Even if these values are not achievable in every region,
improvements in grazing that would increase the animal load can be introduced. With
an intensive grazing scheme that, for example, reduces of the surface area of pastures
by between one-third and one-tenth of the ranch’s original area, more land would be
freed for forested, agro-forested, or agrosilvopastoral activities. An agrosilvopastoral
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model allows for a design that spatially and temporally integrates cattle ranching, tree
formations, and agricultural spaces, either interspersed or in rotation with one another
(Ortiz and Toledo, 1998; Nigh, 1995). The key to this kind of diversification of land use
is in the building of “mosaics of productive spaces,” following the indigenous initiative,
which means incorporating the best aspects of the technical initiative and avoiding the
rancher’s model altogether. Such productive mosaics must be supported by a solid base
of social organization that incorporate principles of equality in decision making that
would affect the management of the system. Any effort for productive integration that
does not incorporate the local population will be, as previous attempts have shown, full
of good wishes but ending in failure.
Therefore, with the current data and evidence (ecological, economic, technological,
and cultural) shown, it is possible to design practical, eco-productive models (Ortiz
y Toledo 1998), in which cattle ranching is structured upon “mosaics of productive
spaces,” diversified in land use and including the following design elements:
a) Annual poly cultivation. Interspersed corn fields, livestock pastures, and
intensive agricultural fields based on organic farming (composting, zero tillage,
green fertilizer, etc.).
b) Intensive grazing. Smaller pastures where cattle are rotated, including the
introduction of useful trees species within the pasture.
c) Forest or fruit tree plantations. Plantations implemented in areas of sloping
terrain, interwoven with different crops of ornamental species, forage,
condiments, medicinal crops, and other commercial species.
d) Areas of reforestation and forest management. Areas of potential reforestation
generated by the intensification of agriculture and cattle ranching, where different
groups of characteristic or commercially valuable forest species are grown.
Depending on the state of regeneration, a continuous economical productivity
during the regeneration process is guaranteed.
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Multiple ecosystem management.
Manejo múltiple del ecosistema.
The ecological viability of the proposal is based on the dynamics that would be
established between the productive systems and by the rotation in time and space of
different productive activities so that energetic and biotic synergies may be established
through the recycling of material, providing the foundation for the efficiency of this
model. Deforestation may be decreased by drastically reducing the surface area needed
for agricultural activities. The implication of this eco-productive model are many, and
while legal, planning, social, and other considerations would need to be addressed, this
model represents a possible and real alternative for sustainable development in the future
humid tropics of Mexico.
Based on the experiences of the present investigation, interviews with farmers and
with technicians and medical and zoological veterinarians, as well as an extensive review
of the literature on the region and topic of study, we have listed below some concrete
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actions that would help to achieve a sustainable, tropical cattle ranching, in addition to
providing topics for further research and investigation.
• Establish an intensive grazing program according to the biological capacities of
the grass and the environmental conditions (weather and soil).
• Promote and conserve the biological diversity (fauna and flora) in spaces of
natural vegetation, along vegetation corridors, and alongside rivers (riparian
• Promote the use of living fences with species of semi-deciduous trees (Bursera
spp., Eritrina spp., Gliricida sepium).
• Produce food supplements and fodder for the cattle that are local and adapted to
the socioeconomic environment.
• Implement agrosilvopastoral management practices by integrating diversified
land uses both spatially and temporally.
• Grow plants that serve as protein banks, or nutritional supplements, for the cattle
utilizing species such as (Leucaena spp.; Ichlyomethia spp. Glirisidia sepium).
• Encourage the use of trees within the pastures.
• Promote an ideology of resource conservation by preserving resources such as
water, soil, and vegetation.
• Practice zero tillage to minimize soil erosion and the loss of animal species above
and below ground.
• Avoid the use of chemical products, herbicides, and pesticides that modify and
eliminate the natural processes that promote biological fertility in the soil.
• Eliminate grazing in zones with a slope inclination of more than 20%.
• Protect and renovate the natural vegetation near water sources and rivers.
• Induce and encourage “islands of vegetation” next to pastures that would serve as
carbon sinks and promote biodiversity, in addition to supplying wood and poles
for pasture fences.
• Strengthen the self-reliance of the production system, community, and farmers
through product diversification.
• Reduce the costs from external inputs by using a diversified system that allows
for the production of balanced food, for example.
• Increase the economic viability of the system by taking advantage of the
synergisms between different local and regional economic activities by for
example, using organic waste products generated by corn, sugarcane, and citrus
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• Reduce the fragility of the biological system by implementing rotational grazing
pastures and the reduction of chemical dependency.
• Stimulate the biological control of plagues and diseases.
• Undertake hydraulic works to mitigate the effects of drought.
• Encourage the use of grass silage.
• Avoid mono-cultivation and promote biological diversity within the production
• Promote the development of other economic activities in addition to cattle
ranching to consolidate the family economy of the participants.
• Enhance solidarity in facing market, social, or environmental uncertainties and
• Promote the use of local labor in order to prevent emigration within the
• Impulse the financial autonomy of production projects through wise investment
and avoiding the mediation of institutions that offer credit an rates that put
farmers at a disadvantage.
• Establish equality between participating members of the productive system
regardless of gender, ethic, or religious group.
• Establish mechanisms of equitable and fair distribution of the costs and benefits
• Encourage the participation of all partners in different project activities.
• Promote autonomy and control in the decision making process, whereby farmers
are able to make decisions on the critical functional aspect of the productive
• Consolidate a democratic structure during the decision making process.
• Encourage the use of local knowledge to promote livestock production
While the conversion of native vegetation to pasturelands has been the main factor in
the destruction of the tropical rain forests of Latin America, it is also true and has been
demonstrated by the present investigation that the search for production alternatives
must be part of a team effort between investigators and rural, peasant farmers. The
following themes are suggested as future lines of research:
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• The efficiency of synergistic productive activities, such as using fodder from corn
plants to feed livestock and tree litter to enrich the soil, among others.
• The availability and balance of the flow of nutrients in the system of intensive
grazing under permanent rotation throughout the year.
• The conservation and mobilization of soil nutrients for plants that are grazed
upon through the establishment of fodder trees and protein banks.
• The optimal spatial and temporal arrangements in pastures that would stimulate
the synergisms through the establishment of biological corridors in the form of
living fences and the maintenance of riparian vegetation and forest fragments.
• The adaptability and complementarity of in the use of animals and plants.
• The presence of earthworm communities in other agro-systems, such as in vanilla
plantations, secondary rain forest, tall grass, cornfields, and pastures at rest and
low grazing.
• The synergisms between the communities of saprophagous beetles and
earthworms as regulators of soil fertility in pastures.
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El desarrollo de la ganadería en la región norte del estado de Veracruz conocida como
Totonacapan, produjo cambios ambientales importantes. Los eventos más destacados de
este desarrollo son la introducción del ganado en el siglo XVI y de los pastos africanos
en la segunda mitad del siglo XIX. Los cambios ambientales se precipitaron entre los
años de 1940 y 1970. El análisis de tres casos de estudio permitió estimar el impacto
que tuvieron las iniciativas de intensificación ganadera en el estado, e identificar algunos
elementos para definir un modelo sustentable de ganadería bovina en esta región.
Palabras clave: Sustentabilidad, deforestación, uso de suelo.
Final Remarks
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The contribution of this special issue is the presentation of new information that
supports a novel consideration of the environmental consequences resulting from the
adoption of cattle ranching in the American continent. Several questions for future
projects and lines of work may be noted, such as: 1) the relationship that livestock had
with the local flora and fauna, 2) the adaptation of livestock management practices to
diverse environments with a natural and human history different to that of Europe, 3) the
environmental and ecological history involved in the arrival and reception of livestock,
and 4) the influence that different breeds of livestock had on the structure and function
of Latin American ecosystems and the landscape.
The articles included in the present debate focus on the impact of the introduction
of livestock in the American continent and on the conflict that took place between the
dedication of land to either raise livestock or crop fields. The arrival of livestock to
the American continent was massive and fast, beginning in the first decades of the 16th
century and involving all regions - from wet to dry climates, from herbaceous vegetation
to scrubland to forests, and from the lowlands to the highlands. In a short period of time
extensive areas dedicated to raise old-world livestock became the predominant land use
in all of America, accounting for the prevalence of cows, horses, sheep, goats, and pigs,
as well as “vaqueros,” “gauchos,” or “charros”. The breeds and varieties of livestock
that came from various regions of the Iberian peninsula and from the North of Africa
were interbred, producing new breeds and varieties, as well as new forms of management
stemming from traditional animal husbandry practices of the Mediterranean and Central
Europe that ultimately fused together in a mode of animal husbandry without precedent
in the entire history of livestock ranching.
The possible negative effects that livestock had on the American environment are
generally focused on its impact on biodiversity, which was provoked by the deforestation
and fragmentation of ecosystems, the creation of grasslands, and by the voluntary and
involuntary introduction of exotic species that have eventually created novel ecosystems
(emerging ecosystems) due to the loss of soil quality (fertility, structure, erosion)
and to the contamination produced by livestock manure and the immoderate use of
agrochemicals. These changes occurred rapidly due to the urgency with which livestock
was introduced, and the amazing reproductive success of the herds.
The conflict that ensued over the decision to dedicate land to crop fields or the
raising of livestock had its origin in the colonial redistribution of land, which privileged
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
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Final remarks
the use of livestock over that of agriculture. This rivalry probably had an immediate
precedent in the Mesta in Spain, where a similar dispute in the 15th century resulted
in the “derrota de las mieses” (defeat of the crops), which translated to the right for
livestock to graze freely between vegetation and agricultural fields. In America the
presence of domesticated animals also generated conflicts without discretion between
indigenous, mestizo, and peninsular farmers. In the case of tropical rain forests, livestock
moved freely throughout the landscape, including between the secondary vegetation
and agricultural fields, thereby linking the landscape through the interchange of species
and strengthening the process of secondary succession. These phenomena may be
demonstrated in the proliferation of native grass species commonly called “grama,”
which began to appear spontaneously as they were dispersed by livestock. It is possible
that living fences, or rows of trees left behind in pastures, are an answer on behalf of
the indigenous farmers to impede the entrance of livestock into their crop fields. Such
fences were composed of secondary rapid growth trees, such as Bursera simaruba and
Gliricidia sepium in Los Tuxtlas, cambronera (Lycium spp.), brambles or prickly pear
cactus in Los Espinales, and cabuya (fique plants) in Villa de Leyva.
In some regions the introduced or exotic species had a significant role in the shaping
of the landscape, influencing, for example, the process of secondary succession that takes
place in agricultural or abandoned grazing sites. In the majority of regions and countries
we do not know which species were introduced directly or indirectly with animal
husbandry practices and which were incorporated in the structure of the ecosystems and
landscapes, and in some cases they were invasive species. This introduction of exotic
species has continued up to this very date, increasing particularly after the arrival of Bos
indicus into tropical America and with the modernization of livestock farming. These
exotic species together with the modern or more recent practices are having increasingly
important repercussions in the shaping of grasslands and man-dominated landscapes that
are becoming part of “emerging ecosystems” (novel ecosystems).
The consideration that the introduction of livestock was negative for the American
landscape is shared by many specialists. Particularly in regards to the introduction of
bovine livestock, which were the most abundant and spread more widely. It is necessary
to point out that the arrival of livestock occurred in two qualitatively different fashions,
the initial event stemming from the entrance of different breeds of Bos taurus in the
Americas, originating in the basin of Guadalquivir in the Iberian peninsula. A large
quantity of cows was left to roam free in the rain forests and secondary vegetation,
where they prospered and increased considerably in number. This period began in the
16th century and depending on the region, lasted until the end of the 19th and beginning
of the 20th century. During this time the livestock foraged on native plants, dispersing
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Final remarks
seeds and herbaceous plants, bushes, and trees across the landscape, thus increasing the
availability of species within the rain forest and sites of secondary vegetation.
The second event began at the beginning of the 20th century when cattle ranchers
began to substitute original Mediterranean breeds for those of B. indicus from tropical
Asia, with the goal of increasing meat production in tropical America. These Asian
livestock breeds were raised in man-made pastures and fed with planted grasses, the
majority African in origin, which were accompanied by numerous exotic species
and occurred over a large extent - in the rain forests in southern Mexico and Central
America as well as the Andean region. The proliferation of this kind of livestock led
to an increase in deforestation and intense fragmentation of habitats, provoking also
the loss of mobility of many native species, which further increased the isolation of
remnant habitats into increasingly smaller fragments, presenting an additional threat to
biodiversity the fragments and presented a threat to biodiversity.
The impact of cattle ranching in Latin America should additionally be understood
within the context of several other events such as: 1) the disappearance of large native
herbivores between approximately 10 and 12000 A.D., 2) the large quantity of plant
species that depend on frugivores for seed dispersal 3) the existence of coprophagous
beetles, 4) the large extensions of secondary vegetation resulting from the massive
abandonment of the zones of agricultural cultivation due to the reductions in the
indigenous population, decimated by new European diseases, and 5) the change in the
property regime and schemes of land ownership during the colonial rule.
In each region of the Americas livestock was able to take advantage of variable
quantities of forage, with a distinct proportion of species dispersed by animals, which
established a peculiar relationship, in which predatory species began to suffer an
onslaught of parasites and diseases. In Los Tuxtlas fewer predators were present, along
with a large quantity of resources that could be used as forage additionally, the region
has a dominant presence of zoochorous plant species. In comparison with Los Tuxtlas
more predators and diseases existed in other sites, like Totonacapan.
Perhaps the argument that alleges that soil deterioration was produced by cattle
ranching and resulted in environmental degradation is one of the most specific. This
occurred in sites where livestock were stabled and the land was over-grazed. However,
considering the livestock breed of B. taurus, its weight, hoof size, and the wild habitat
to which it was accustomed to roam, its introduction to Latin America did not result in a
drastic deterioration of the soil. In this scenario, soil deterioration could also have been a
product of an inadequate pre-Hispanic and colonial management.
Another cause of erosion and loss of soil fertility was due to the displacement of the
indigenous population from fertile and productive lands to those that were fragile, on
steep terrain, and susceptible to erosion.
PASTOS, 42 (2), 301 - 304
PASTOS 2012. ISSN: 0210-1270
Final remarks
Considering that the cultivated surface area in Mesoamerica was extensive and that
the agricultural practices were mainly design and implemented around the seasonal use
and subsequent abandonment of the land, it seems clear that productivity depended in
large part on the management of the process of secondary succession or forest regrowth.
This process was rooted in the recuperation of soil fertility, useful native species, and
the regeneration of the rain forest or native vegetation. This kind of management led to
a greater resilience of ecosystems, which explains the biodiversity currently encountered
in Mesoamerica. Such biodiversity is self-maintaining, in spite of the fact that the
fragments are decreasing in size; this scenario is also occurring simultaneously in other
regions, such as the Amazon.
The key to a greater resilience and maintenance of biodiversity most likely rests in
maintaining the connectivity of the landscape. This is demonstrated by the presence of
trees in open fields, a characteristic of forested landscapes - especially of both dry and
wet tropical forests. These trees facilitate the movement of frugivores (mainly birds
and bats), and pollinators throughout the landscape, guaranteeing the dispersion and
pollination of many species. This feature of the landscape seems to have its origin in the
deforestation practiced by many indigenous farmers, in which cycles of land use as crop
fields were alternated with forest re-growth in the old-field.
In this context livestock farming incorporated itself into American nature, much in
the same way that frugivorous mammals and birds dispersed fruits and seeds across the
landscape. Not all fruits or seeds were attainable by birds and bats due to the hardness
and size of the fruit, thus livestock farming has contributed to the connectivity between
fragments and secondary vegetation through dispersion of fruits and seeds.
Livestock also quickly developed a relationship with different species of
coprophagous beetles across the landscape, as livestock manure and its handling by
these beetles increased the fertility of the soil and also contributed to the distribution of
seeds by the beetles.
In the current landscape livestock farming has resulted in numerous isolated trees,
spread out throughout pasturelands – a scenario that has a striking resemblance to the
landscape of the Spanish and Portuguese dehesas. While this may be a coincidence, it
could also represent a new design that has implemented both ancient and traditional
The contributions assembled in this issue and their findings opens up a new range
of explanations and possibilities that should be investigated and integrated quickly in
the debate, as they represent a new possibility of managing the landscape in a way that
would harmonize cattle ranching with biodiversity and development.
Sergio Guevara Sada
In Memoriam
PASTOS, 42 (2), 307 - 314
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Gaspar González y González
4 de enero de 1921 - 27 de octubre de 2012
“Fundador y Socio de Honor de la SEEP”
Gaspar González falleció el día 27 de octubre de 2012, a solo poco más de dos
meses para cumplir la edad de 92 años. Vida larga, intensa y plena durante la que mostró
poseer una personalidad intelectual poliédrica con amplios conocimientos en diferentes
parcelas del saber y muy lejos del hombre de dimensión lineal, hoy tan frecuente, con el
que solo se puede hablar de temas puntuales. No es tarea fácil enmarcar el perfil humano
de cualquier persona. Como decía Alexis Carrel, Premio Nobel de Medicina 1912, en
su libro L’homme, cet inconnu, el hombre es un desconocido incluso para sí mismo.
Solamente al final de la vida, desarrolladas todas las capacidades en el ejercicio de la
vocación, es cuando se puede intentar definir, al menos parcialmente, la personalidad de
Don Gaspar, o Gaspar sin más, generalmente se usaba lo primero, sin necesidad
de apellidos para saber de qué persona se trataba, era un hombre de aspecto elegante,
educado y de trato correcto. No solía dar muchas facilidades para que se le tutease. En
mi caso concreto, esto sucedió al cabo de 25 años de conocernos y trabajar muchas horas
juntos, cuando obtuve la Cátedra que él dejaba por alcanzar la edad de jubilación. No
obstante, estoy totalmente seguro de que me apreciaba sinceramente desde muchos
años antes y así me lo demostró siempre que necesité su apoyo. Se decía también de
él que era un Profesor algo abstraído y que olvidaba fácilmente los nombres, incluso
de personas que trabajaban con él, a pesar de la envidiable memoria que poseía. Su
facilidad de almacenar y evocar información pasada se manifestaba, claramente, cuando
se mantenían conversaciones con él sobre temas científicos o en sus intervenciones
en Reuniones y Congresos científicos. En esos momentos, uno se daba cuenta de que
Gaspar recordaba datos concretos, referencias o frases de revistas o libros que hacía
mucho tiempo que había leído.
El proceso vital de Gaspar tuvo etapas bien definidas encaminadas siempre a su
realización como hombre. En plena juventud descubrió su vocación por la docencia
e investigación, pilares básicos sobre los que posterior y principalmente desarrolló
su vida profesional a través de su paso por la Universidad y el Consejo Superior de
Bases de datos: (España), AGRIS (Italia), CAB Abstracts (Reino
Unido), CABI Full Text (Reino Unido), Catálogo LATINDEX (México), DIALNET (España), ICYT
Ciencia y Tecnología (España)
PASTOS, 42 (2), 307 - 314
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In Memoriam Gaspar Gonzalez y González
Investigaciones Científicas (CSIC). Los muchos años que dedicó a ambas Instituciones
han dejado una huella de buen hacer que, aún hoy día, sigue siendo fácilmente
reconocible. A ello y, además, a la huella dejada por Gaspar a su paso por la Sociedad
Española para el Estudio de los Pastos (SEEP) están dedicadas estas líneas, en un
intento de perfilar algunas facetas representativas de su gran y compleja personalidad.
Comenzando por el principio, como se debe hacer siempre, recordaré que Gaspar
nació el 4 de enero de 1921 en San Adrián del Valle, un pequeño pueblo de la provincia
de León. Primogénito de tres hermanos, su vida se fue desarrollando al lado de unos
padres buenos, tradicionales y de humilde posición, dispuestos a cualquier sacrificio
para asegurar el porvenir de sus hijos en las duras condiciones existentes durante
y después de la Guerra Civil española. Estudió el Bachillerato en el Real Instituto
Jovellanos de Gijón, ciudad a donde se había ido a vivir la familia a resultas del
traslado del padre, motivado por su pertenencia al Cuerpo de la Guardia Civil. Años
más tarde, se matriculó en la Escuela Superior de Veterinaria de León, graduándose
en 1943 a los 22 años de edad. Posteriormente, realizó los cursos de Diplomado en
Estudios Superiores en la Escuela Superior de Veterinaria de Madrid. Transformadas las
Escuelas de Veterinaria en Facultades en el año 1943, Gaspar se licenció y doctoró en
Veterinaria por la Universidad Central, hoy Universidad Complutense de Madrid. Inició
su carrera docente en 1944, como Profesor Ayudante y la continuó con su posterior
nombramiento, en 1947, de Profesor Adjunto Encargado de la Cátedra de Fitotecnia
y Economía Rural. Embarcado ya en la aventura de su propia vocación y espoleado
por su incesante inquietud de adquirir nuevos conocimientos y perfilar con mayor
profundidad su especialización en la temática de la producción animal, Gaspar realizó
estancias de mayor o menor duración en Universidades y Centros de investigación:
Gran Bretaña (Universidad de Birmingham y Grassland Improvement Station), Holanda
(Departamento de Bioquímica de la Facultad de Veterinaria de Utrech) y Dinamarca
(Universidad de Copenhague). Posteriormente, en el transcurrir de su vida hizo nuevas
visitas a Centros de diversos países como Estados Unidos, Francia, Alemania, Suiza,
etc. Esta especialización y formación científica fue acompañada por una formación
humanista conseguida a través de Cursos Ético-Sociales organizados por el Instituto
Social León XIII de la Universidad Pontificia de Salamanca. La cultura humanista
adquirida le permitió entender el valor incondicional de la persona humana y el sentido
de su crecimiento, así como el por qué las ciencias deben encontrar su lugar dentro de la
colaboración al servicio del hombre y favorecer el desarrollo de la conciencia en deberes
morales y sociales.
En 1951, tras brillante oposición, Gaspar obtuvo por unanimidad del tribunal, la
Cátedra de Fitotecnia y Economía Rural, denominación que cambió a la de Agricultura
y Economía Agraria al cabo de algunos años con motivo de la implantación de un nuevo
PASTOS 2012. ISSN: 0210-1270
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In Memoriam Gaspar Gonzalez y González
plan de estudios. Gaspar fue el titular de esta Cátedra de la Facultad de Veterinaria de la
UCM desde aquel año hasta que se jubiló en 1987, siendo nombrado Profesor Emérito
de la UCM. Este largo periodo de tiempo le permitió desarrollar plenamente sus dos
grandes vocaciones: la de maestro universitario y la de investigador científico, esta
última en conjunción con su actividad en el CSIC. El resultado de la labor desarrollada
queda reflejado, entre otras cosas, en las 34 tesis doctorales dirigidas, los 19 profesores
de universidad formados, el más de un centenar de publicaciones en revistas nacionales
y extranjeras, incluyendo trabajos de investigación, revisión o doctrinales, numerosas
ponencias y comunicaciones presentadas a Congresos y Reuniones Científicas,
etc. A esto habría que añadir su actuación como director o participante en cursos
postgrado de especialización, algunos repetidos durante más de 10 años, como es el
caso de los titulados: “El impacto ambiental de la actividad agraria” y el de “Módulo
de introducción a la Agricultura ambiental”, ambos organizados por el Instituto
Universitario de Ciencias Ambientales de la UCM. Además, participó en los Cursos de
Verano organizados por la UCM, por la Universidad de Granada y por la Universidad
Internacional Menéndez Pelayo.
Gaspar no olvidó tampoco la importancia de la denominada Formación Profesional
e intervino en ella contribuyendo a la creación de los Institutos de Enseñanza Laboral
y a la elaboración de los correspondientes programas en la modalidad Agrícolaganadera participando, además, como profesor en los cursos de perfeccionamiento
para el profesorado de los referidos Institutos. Igualmente fue director de los Cursillos
Nacionales de Capacitación Ganadera organizados por la Junta Nacional Sindical de
Labradores y Ganaderos, desde 1955 hasta 1975; es decir, durante 20 años consecutivos.
En resumen, un rendimiento pedagógico que creo que nadie dudaría en calificarlo de
Por otra parte, a su tarea docente unió el desempeño de diversos cargos académicos,
comenzando por dirigir el Departamento de Agricultura y Economía Agraria y,
posteriormente, el de Producción Animal de la Facultad de Veterinaria de la UCM. Fue
Vicedecano (1967-1969) y Decano (1973-1977) de dicha Facultad y, en el cuatrienio
1977-1981, Vicerrector de la UCM formando parte del equipo presidido por el Rector
Prof. Vián Ortuño. Según palabras del propio Rector, fueron años difíciles que Gaspar
sorteó con tan buen talante como talento. Gaspar se ocupó muy especialmente de la
reordenación de los Colegios Mayores, consiguiendo recuperar para la Universidad
los edificios situados en la calle Donoso Cortés, así como los Colegios José Antonio y
Nuestra Señora de la Almudena, pertenecientes a la Secretaría General del Movimiento.
También llevó con habilidad la gestión de liberar a favor de la UCM la herencia
multimillonaria de la Fundación Del Amo, reclamada por otra institución ajena a la
PASTOS, 42 (2), 307 - 314
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In Memoriam Gaspar Gonzalez y González
Paralela y conjuntamente con su actividad en la universidad, Gaspar desarrolló otra
intensa y meritoria labor en el CSIC. Para entender mejor esta situación de pertenencia
a dos diferentes Organismos estatales hay que retroceder en el tiempo hasta la década
de los años 50 del siglo pasado. En aquella época, la Facultad de Veterinaria de Madrid
estaba ubicada en un antiguo inmueble de la calle Embajadores. El edificio había sido
construido en el s. XVIII y parece ser que se trataba de un antiguo palacio donde residió
un asesor político de Godoy. A pesar de la remodelación interior que se había realizado,
fundamentalmente para disponer de aulas suficientes, las instalaciones disponían de muy
pocos laboratorios y, además, sin reunir las mínimas condiciones para realizar trabajos de
investigación. Esta fue la situación que se encontró Gaspar: una Cátedra prácticamente
sin dotación de medios materiales y humanos. El entorno tan poco apropiado para hacer
investigación, unido al hecho de que entonces era relativamente frecuente que Centros
del CSIC estuvieran dirigidos por catedráticos, fue lo que movió a Gaspar a buscar en
este Organismo la solución para llevar a cabo su vocación científica.
La oportunidad le llegó a través de los buenos oficios de su amigo el Prof. Ángel
González Álvarez, Catedrático de Metafísica de la Facultad de Filosofía y Letras, quien
le presentó y dio a conocer al entonces Director del Instituto de Edafología y Fisiología
Vegetal del CSIC, Prof. José María Albareda, Catedrático de Edafología de la Facultad
de Farmacia de la UCM. Entre ambos se establecieron unas buenas relaciones de trabajo
y amistad, interesándose el Prof. Albareda por la propuesta de creación de un grupo de
investigación en producción animal en el propio Instituto de Edafología. Así surgió, a
comienzos de 1952, la Sección Bioquímica de Forrajes, bajo la dirección de Gaspar,
reuniendo a algún personal científico, auxiliar y subalterno procedente de las Facultades
de Farmacia y Veterinaria, así como del propio Instituto de Edafología y Fisiología. En
1965, la Sección, siguiendo adscrita al Instituto de Edafología y Fisiología Vegetal, se
ubicó en los laboratorios de la quinta planta del Centro de Investigaciones Biológicas
(calle Velázquez nº138), con la denominación de Bromatología y Nutrición Animal. Dos
años más tarde esta Sección se transformó en un Departamento con el mismo nombre,
cambiándose éste por el de “Alimentación y Productividad Ganaderas” en 1960. Años
más tarde, en 1966, el Departamento fue elevado al rango de Instituto, siguiendo
dentro del Patronato Alonso de Herrera del CSIC, con el nombre de “Alimentación y
Productividad Animal”. Este Instituto integró no solo al mencionado Departamento,
sino también al Departamento de Productividad y Economía Ganadera escindido
del Departamento de Economía y Productividad Agraria, y a la Sección de Fisiología
Animal de la Universidad de Navarra cuyo Jefe era el Prof. Jesús Larralde, Catedrático
de dicha Universidad. Al cabo de unos años, esta Sección de Fisiología Animal se separó
del Instituto por no resultar operativa la fusión debido a la lejanía existente entre ambos
Centros. Gaspar fue nombrado Director del nuevo Instituto creado.
PASTOS 2012. ISSN: 0210-1270
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In Memoriam Gaspar Gonzalez y González
La falta de espacio de que disponía el Instituto de Alimentación y Productividad
Ganaderas en el Centro de Investigaciones Biológicas, indujo a Gaspar a realizar los
trámites oportunos para proceder a su traslado a la Ciudad Universitaria. Esto pudo
realizarse debido a que se había finalizado la construcción de la Facultad de Veterinaria
y había sido asignado a la Cátedra de Agricultura y Economía Agraria un edificio que
había quedado vacío y un terreno anejo de aproximadamente 4 ha de superficie. Se
firmó un Convenio entre el CSIC y la UCM en el que se estableció la normativa que
regulaba la colaboración entre ambos Organismos. El traslado del Instituto a su nueva
ubicación se realizó en el año 1969. No es fácil resumir la gran labor desarrollada por
Gaspar como responsable del Instituto. Desde el primer momento, con gran empuje y
esfuerzo, se dedicó a conseguir medios económicos del CSIC que permitieron realizar
todas las reformas necesarias para adaptar el edificio a su nuevo uso y transformar su
estructura interna en laboratorios, despachos, etc. Además, también obtuvo financiación
para que se fuera dotando a los distintos laboratorios de material suficiente y moderno.
Fue tan eficaz su labor a este respecto que recuerdo que el Instituto dispuso del primer
autoanalizador de aminoácidos que hubo en aquella época en España. Se reconstruyó
también una nave para adecuarla a su utilización en ensayos con animales: aves,
rumiantes y cerdos. Igualmente, Gaspar puso todo su empeño en transformar el terreno
baldío de que se disponía, en una finca de experimentación para hacer ensayos con
especies pratenses y forrajeras. Se hicieron enmiendas al suelo, se construyó un estanque
y se instaló riego por aspersión. La labor de investigación que se realizó en el Instituto
fue realmente importante y supuso una buena parte de los trabajos publicados sobre
alimentación, nutrición animal y economía ganadera realizados en España.
No hay que olvidar tampoco las numerosas Tesis doctorales realizadas bajo
la dirección de Gaspar y leídas en las Facultades de Ciencias Químicas, Ciencias
Biológicas, Farmacia y Veterinaria. Mención especial se debe dedicar, además, al
personal que se formó o colaboró en el Instituto y, posteriormente, desarrolló su
actividad en empresas zootécnicas que contribuyeron al desarrollo ganadero en España
(Bioter, Biona, Farco, Híbridos Americanos, Industrias Agrícolas de Zaragoza, Lucta,
Gaspar también intervino en la creación de la estación Experimental La Mayora,
integrada en el área de Ciencias Agrarias del CSIC. La idea surgió del Prof. Albareda,
hace ya más de 50 años, y se basaba en aprovechar el potencial que ofrecen las
condiciones climatológicas de las zonas del Levante y Sureste de España para la
producción de hortalizas extratempranas y frutas subtropicales, convirtiendo estas zonas
en una especie de California de Europa. El proyecto contó con el apoyo de Gaspar y
algunos otros investigadores, decidiéndose su ubicación en la provincia de Málaga, lo
cual fue apoyado por las autoridades provinciales e, incluso, por países como Alemania,
PASTOS, 42 (2), 307 - 314
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In Memoriam Gaspar Gonzalez y González
esto último en el marco de un posible programa de cooperación Hispano-Alemán a
través de su Embajada en Madrid. Después de realizar una serie de ensayos previos en
fincas particulares y vistos los resultados tan favorables obtenidos, la Junta del Patronato
Alonso de Herrera acordó la creación de una Estación Experimental en aquella zona.
El Prof. Albareda propuso a Gaspar para que se encargara de preparar la memoria
justificativa para el desarrollo de este Centro. Aprobada la misma, se adquirió en 1969
una finca de una extensión de 51 ha en la zona de Algarrobo-Costa, a unos 40 km de
Málaga. Por sugerencia del propio Gaspar, dos miembros del Instituto de Alimentación y
Productividad Ganaderas se trasladaron a la finca para apoyar el desarrollo del proyecto.
Uno de ellos, el Dr. Rafael Viñarás, Colaborador Científico, con el tiempo fue Secretario
de la Estación Experimental y el otro, D. Antonio Gómez Barcina, doctorando becario,
llegó a ser el Director de la misma. En el año 1968 fue inaugurada oficialmente la
Estación Experimental La Mayora. El éxito a los largo de los años ha sobrepasado todas
las expectativas, constituyendo un referente para muchos de los agricultores de la región
dedicados a la producción de hortalizas y frutos subtropicales, principalmente aguacate,
chirimoya y mango.
El paso de Gaspar por la Universidad y el CSIC, demostrando su preparación y
eficacia, le llevó a ocupar puestos relevantes en ambas Instituciones y en el Ministerio de
Educación y Ciencia: Consejero de Educación Nacional; Vocal de la Comisión Nacional
de Becas; Vocal de la Comisión del Consejo Técnico de Universidades Laborales;
Miembro de la Comisión Asesora para el Fomento de la Investigación en la Universidad;
Director Adjunto de Investigación de la División de Ciencias del CSIC; Consejero de
Número de los Patronatos José María Cuadrado y Alonso de Herrera del CSIC; Vocal
de la Comisión Conjunta de Investigación Agraria (Ministerios de Educación Nacional
y de Agricultura); Vocal de la Comisión Asesora de Investigación Científica y Técnica
de la Presidencia de Gobierno; Vocal del Consejo Regional del CRIDA 06; Vocal de la
Comisión Permanente de la División de Ciencias del CSIC; Vocal de Política Científica
del Patronato Alonso de Herrera. Además, se le concedieron múltiples distinciones
y condecoraciones, entre ellas, el Víctor de Plata al Mérito Escolar, el Botón de Oro
del Colegio Mayor César Carlos, dos Medallas de Plata de la UCM por los relevantes
servicios prestadas a la misma y la Encomienda de Número de la Orden Civil al
Mérito Agrícola. En justicia, a Gaspar también se le abrieron las puertas de las Reales
Academias de Doctores y de Farmacia, ingresando en ambas como Académico de
Las actividades desarrolladas en la Universidad y el CSIC fueron, sin duda, factores
determinantes de la estrecha relación que siempre existió entre Gaspar y la SEEP. No
en vano, esta Sociedad surgió por iniciativa suya y a ella le dedicó mucho esfuerzo y
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In Memoriam Gaspar Gonzalez y González
La idea de crear una Sociedad Científica en España semejante a la British Grassland
Society del Reino Unido, le surgió a Gaspar durante su estancia, en el año 1949, en la
Grassland Improvement Station de Stratford-upon-Avon, Inglaterra. Comentada la idea
con el propio Director del Centro, Dr. Willliam Davies, éste le animó a madurarla y
llevarla a efecto a su regreso a España. Ya de vuelta, Gaspar dejó aparcada la idea durante
algunos años por estar ocupado con la preparación de su oposición a Cátedra y, una vez
ganada la misma, con la creación de una Sección de investigación dentro del Instituto
de Edafología y Fisiología Vegetal del CSIC. Unos años después, concretamente en
1958, retoma la idea de crear la Sociedad y se la expone al Prof. Albareda, Director del
citado Instituto, a quien el proyecto le parece muy bien y promete su apoyo y ayuda.
La buena acogida del proyecto por parte del Prof. Albareda, impulsó a Gaspar a poner
en marcha el proceso de fundación de la SEEP contando para ello con la valiosa y
entusiasta colaboración de los bien conocidos Dr. Pedro Monserrat y Dr. Manuel Ocaña,
este último también fallecido. Como resultado de la labor conjunta realizada, coordinada
por Gaspar, y después de mucho esfuerzo y trabajo, en el año 1959 se crea de facto la
SEEP y se elige al efecto una Comisión Directiva provisional en la que Gaspar figura
como Vocal. Al año siguiente, es decir en 1960, la SEEP queda ya oficialmente inscrita y
legalizada como Sociedad Científica sin ánimo de lucro, eligiéndose mediante votación
directa de los socios a la primera Junta Directiva. A esta Junta pertenece Gaspar en
calidad de Vocal y así continuará hasta que en 1968 es elegido Presidente de la SEEP
y, posteriormente reelegido en 1972 por otro periodo de cuatro años. En el año 1973,
y con motivo de haber sido aprobado que la “6th General Meeting” de la European
Grassland Federation (EGF) se celebraría en Madrid, Gaspar fue nombrado Presidente
de esta Federación por un periodo de dos años (1974-1976). Por este motivo, Gaspar
tuvo que coordinar todo lo relativo al desarrollo de este evento, contando con el apoyo
de Organismos del Ministerio de Agricultura (Instituto Nacional de Investigaciones
Agrarias, Dirección General de la Producción Agraria e Instituto Nacional para la
Conservación de la Naturaleza) y del Ministerio de Educación y Ciencia (Secretaría
General y Patronato Alonso de Herrera del CSIC). En 1994, Gaspar fue nombrado Socio
de Honor de la SEEP en premio a la labor realizada durante tantos años.
Me atrevería a decir, y creo no equivocarme mucho, que excepto en dos o tres
ocasiones como máximo, Gaspar asistió a todas las Reuniones Científicas de la SEEP, en
muchas de ellas presentando ponencias y comunicaciones científicas. El empeoramiento
de su estado físico, motivado por la enfermedad que acabó con él, le impidió asistir a las
dos últimas Reuniones, las celebradas en Toledo, en 2011 y en Pamplona, en 2012.
Quienquiera que haya leído todo lo que antecede, estará de acuerdo conmigo en
que Gaspar fue un hombre de gran personalidad e intelectualidad en su doble vertiente
docente e investigadora, dejando su huella allá por donde pasó. A él se le pueden aplicar
aquellas frases que aparecen en un conocido verso de Antonio Machado:
PASTOS, 42 (2), 307 - 314
J. Treviño
PASTOS 2012. ISSN: 0210-1270
In Memoriam Gaspar Gonzalez y González
Caminante, son tus huellas
el camino y nada más;
caminante, no hay camino,
se hace camino al andar.
Gaspar hizo un camino ancho, muy ancho, en el que dejó huellas profundas debido
a la pesada carga de buen hacer que transportó en la mochila de la vida. Probablemente,
todo eso no habría ocurrido si no hubiera existido en su vida una gran mujer, Ana María
Doncel. Se suele decir que detrás de un gran hombre siempre hay una gran mujer y
esto fue también lo que le ocurrió a Gaspar, tener una esposa de excepción. Ana María,
bella y elegante joven de conocida familia navarra, siempre supo “estar ahí”, en la
retaguardia, aceptando de buen grado todas las renuncias necesarias para facilitar las
múltiples actividades de su marido. De ella y de sus hijos, Gaspar en su discurso de
ingreso en la Real Academia de Farmacia dijo lo siguiente: “tengo una deuda con mi
mujer e hijos, a quienes tal vez, no presté toda la atención que se merecían acuciado por
quehaceres que me alejaban, al menos físicamente, de su lado, en muchas ocasiones. A
ella y a ellos, a su estímulo y comprensión hay que atribuir principalmente los méritos
que se puedan anotar en el haber de mi vida”. Los seis hijos del matrimonio seguro que
habrán recibido muchas de las virtudes de su padre y sabrán llevarle con orgullo en su
memoria: Javier, doctor en Veterinaria, Facultativo Especialista en el Centro Nacional
de Alimentación (Instituto de Salud Carlos III); Ana, licenciada en Periodismo, Jefa
de Relaciones Públicas del Ente Público Radio Televisión de Madrid; Gaspar, doctor
en Ciencias Físicas, Investigador Científico del Departamento de Metalurgia Física
del Centro Nacional de Investigaciones (CSIC); Inés, doctora Ingeniero de Montes,
Catedrática de Dasometría, Inventario, Ordenación de Montes y Aprovechamientos
Forestales en la Escuela Universitaria de Ingeniería Técnica Forestal (UPM); Mamen,
Secretaria administrativa en la UCM; y Miguel, doctor en Biología, Investigador en el
Laboratorio de Ecotoxicología del Departamento de Medio Ambiente (INIA).
No quisiera finalizar este memorial sin hacer referencia a las profundas creencias
religiosas de Gaspar y a su convicción de que el paso por esta vida es el preámbulo de
llegada a la otra, a la VIDA con mayúsculas. Por eso, desde estas páginas de la revista
Pastos, le envío un fuerte abrazo en la seguridad de que le llegará allí donde se encuentre
Jesús Treviño
Profesor Emérito de la Universidad Complutense de Madrid
Diciembre de 2012
Instrucciones Para Autores
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Ámbito de la Revista...................................................................................................... 318
Cesión de derechos de los autores.................................................................................. 318
Idiomas........................................................................................................................... 318
Textos originales............................................................................................................. 318
Formato.................................................................................................................... 318
Extensión.................................................................................................................. 318
Envío de los originales................................................................................................... 318
Proceso de revisión de los originales.............................................................................. 319
Organización del texto.................................................................................................... 319
Título principal............................................................................................................... 319
Título abreviado.............................................................................................................. 319
Nombre del autor o autores............................................................................................ 320
Dirección del autor o autores................................................................................... 320
Resumen (en idioma original)........................................................................................ 320
Palabras clave (en idioma original).......................................................................... 320
Exposición del trabajo.............................................................................................. 320
Tablas.............................................................................................................................. 321
Títulos de las tablas.................................................................................................. 321
Figuras............................................................................................................................ 321
Pies de las figuras..................................................................................................... 321
Fotografías...................................................................................................................... 321
Referencias bibliográficas (dentro del texto).................................................................. 321
Referencias bibliográficas (al final del texto)................................................................. 322
Forma de presentación de las referencias al final del texto...................................... 322
Caso de revistas.................................................................................................. 322
Caso de libros de un solo autor o grupo de autores para toda la obra................ 323
Caso de libros colectivos, con capítulos escritos por distintos autores.............. 323
Título, resumen y palabras clave en idioma alternativo................................................. 323
Titulo en inglés............................................................................................................... 324
Summary......................................................................................................................... 324
Key words................................................................................................................ 324
Unidades de medida........................................................................................................ 324
Expresión algebraica de los símbolos de las unidades SI........................................ 324
Notación numérica.......................................................................................................... 325
Cifras decimales....................................................................................................... 325
Dentro del texto en español................................................................................ 325
Dentro del texto en inglés (summary)................................................................ 325
Siglas, materiales, etc..................................................................................................... 325
Nombres de plantas, cultivares, etc................................................................................ 325
Titulares.......................................................................................................................... 325
Jerarquización de titulares.............................................................................................. 326
Apéndice 1. Síntesis de las instrucciones para autores................................................... 326
Apéndice 2. Resumen del SI (Sistema Internacional de Unidades)............................... 327
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
La revista PASTOS admite artículos originales sobre la producción y utilización de pastos
y forrajes, dentro de las áreas de conocimiento siguientes: recursos naturales (suelo, agua, clima,
etc.) en los que se basa la producción de pastos y forrajes; ecología, nutrición, protección, selección,
mejora, manejo y conservación de especies forrajeras y pratenses; nutrición, alimentación y manejo
de animales; sistemas de producción animal con base en pastos y forrajes; aprovechamiento de
pastos; impacto ambiental de las explotaciones ganaderas; estudios económicos; etc.
Los originales pueden ser artículos científicos, notas de investigación, temas monográficos,
revisiones científicas sobre temas de interés (generalmente por invitación de la Dirección),
conclusiones de reuniones científicas, resúmenes de tesis doctorales, descripción de nuevas
variedades y recensiones de libros.
Dado que la revista es de libre acceso gratuito, la publicación en PASTOS implica la cesión de
los derechos de los autores para que PASTOS pueda difundir sus artículos a través de las bases de
datos que estime oportunas.
La revista PASTOS acepta artículos originales en español, idioma preferente. Se aceptan
también textos originales en francés, inglés, italiano y portugués.
Los textos originales se escribirán con letra Times New Roman de 12 puntos, o equivalente, en
líneas a 1,5 espacios. Los márgenes derecho e izquierdo serán de 2,5 cm a cada lado, y el superior
e inferior de 3 cm, aproximadamente. Cada página tendrá un máximo de 32 líneas (incluidas las
vacías) y unos 86 caracteres por línea (incluidos los espacios vacíos). No se partirán las palabras
con guiones al final de las líneas. El texto se justificará únicamente por la izquierda. Esta norma
afectará a todo el artículo: título, nombre de autores, dirección de autores, resumen, palabras clave,
introducción, material y métodos, resultados, discusión, agradecimientos, referencias bibliográficas,
título en inglés, summary y key words, con excepción de las tablas, que podrán ir a un espacio
entre líneas y con caracteres más pequeños, si fuese necesario, hasta un mínimo de 9 puntos.
La primera línea de cada párrafo se sangrará con tabulador, nunca con la barra espaciadora.
La extensión de los mismos no excederá de 30 páginas para los artículos científicos, 10 para
las notas de investigación, cinco para conclusiones de reuniones científicas y resúmenes de tesis
doctorales, y una para descripción de variedades y recensiones de libros, incluyendo en estos límites
las figuras, las tablas y las referencias bibliográficas. Para las revisiones científicas no hay un límite
prefijado de páginas.
Se enviarán por correo electrónico al Director de la Revista PASTOS, D. Juan Piñeiro Andión, a
la dirección electrónica siguiente: [email protected]
Se pueden enviar también en disco a la dirección postal siguiente:
Centro de Investigacións Agrarias de Mabegondo
Estrada de Betanzos a Mesón do Vento, km 7
15318 Abegondo
A Coruña (España)
PASTOS 2012. ISSN: 0210-1270
PASTOS, 42 (2), 317 - 329
Instrucciones para autores de PASTOS
Los originales recibidos se enviarán a dos miembros del Comité de Redacción para su
evaluación, a los que se adjunta una “Hoja de evaluación”, en la que los evaluadores resumirán
su informe sobre el artículo correspondiente. A su vez, sobre el texto del artículo podrán hacer
los comentarios y sugerencias que estimen oportunas, utilizando los programas de “comentarios”
y de “control de cambios” de windows. El artículo no será publicado en PASTOS si los dos
evaluadores indican que “no es aceptable para publicación”. En el caso de que un evaluador lo
califique como “aceptable” y el otro como “no aceptable”, se pedirá un nuevo informe a otro
miembro del Comité de Redacción, que servirá para resolver el conflicto creado. Los miembros
del Comité de Redacción podrán someter el artículo a la evaluación de los expertos que estimen
oportunos para elaborar su informe.
Los artículos científicos tendrán la siguiente disposición:
-Título principal
-Título abreviado
-Nombre autor/es
-Dirección autor/es
-Resumen en idioma original
-Palabras clave en idioma original
-Material y métodos
-Referencias bibliográficas
Si el inglés no es idioma original:
-Título en inglés
-Resumen en inglés (summary)
-Palabras clave en inglés (key words)
Si el español no es idioma original:
-Ttulo en español
-Resumen en español
-Palabras clave en español
Centrado. Letra normal, MAYÚSCULA y negrita. Debe ser claro, corto y conciso, evitando
términos como “estudios sobre...”, “observaciones...” “contribución al...”. Tendrá un máximo de
25 palabras.
Inmediatamente debajo del título principal se pondrá un título abreviado, para que la imprenta
lo ponga en la cabecera de cada página del artículo. Irá centrado, con letra normal, minúscula y no
negrita. Tendrá un tamaño máximo de 50 caracteres, espacios incluidos.
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
Centrado. Letra normal y MAYÚSCULA. INICIAL/ES del nombre, seguidas de punto,
APELLIDO/S seguido/s de coma. Autores separados por una coma, o la conjunción Y, si son los
dos últimos. Se señalará el autor para la correspondencia con una nota al pie de la primera página.
Dirección del autor o autores
Incluirá la dirección postal completa y la dirección electrónica.
Centrada. Letra normal y minúscula. Debe ser lo más completa posible, con objeto de que los
lectores no tengan dificultades para comunicarse con los autores. Si los distintos autores tienen
direcciones diferentes, debe indicarse con un superíndice numérico.
Area de Ecología. Facultad de Biología. Universidad de Salamanca. Plaza de los caídos s/n.
E-37071 Salamanca (España). Dirección electrónica. 2Area de Ecología. Facultad de Biología.
Universidad de León. E-24071 León (España). Dirección electrónica.
RESUMEN (en idioma original)
Debe ser informativo, no indicativo, para permitir al lector apreciar el contenido e interés del
trabajo. Debe informar sobre objetivos, metodología, resultados y conclusiones. Máximo de 300
palabras para artículos científicos y notas de investigación, y 450 para las revisiones científicas.
Anchura de líneas y tipo de letra igual al resto del artículo. Líneas a 1,5 espacios. Sangrar la primera
línea de cada párrafo utilizando el tabulador, no la barra espaciadora. En su contenido no debe haber
referencias ni al texto, ni a las figuras, ni a las tablas del artículo resumido. La palabra RESUMEN
irá en letra normal MAYÚSCULA, negrita y justificada a la izquierda.
Categoría de TITULAR MAYOR.
Palabras clave (en idioma original)
El resumen irá seguido de un máximo de cinco palabras clave que no estén contenidas en el
título. Se escribirán en letra normal y minúscula, separadas por comas. La última irá seguida de
punto. Las palabras Palabras clave irán en letra normal, minúscula y negrita, justificada a la
izquierda y seguida de dos puntos, después de los cuales se pondrán las palabras clave.
Categoría de titular de 1er orden.
Exposición del trabajo
Líneas a 1,5 espacios. Letra normal y minúscula. La primera línea de cada párrafo se sangrará
con tabulador, nunca con la barra espaciadora. No se partirán las palabras mediante guiones al final
de las líneas. Debe justificarse únicamente por la izquierda. Constará de los apartados siguientes:
Resultados y discusión pueden formar un solo apartado, si se estima oportuno.
PASTOS 2012. ISSN: 0210-1270
PASTOS, 42 (2), 317 - 329
Instrucciones para autores de PASTOS
Las palabras que definen los seis apartados antes citados, irán en letra normal, MAYÚSCULA,
negrita y justificados a la izquierda.
En AGRADECIMIENTOS deben figurar las personas e instituciones que han dado apoyo
técnico y el origen de la financiación para el trabajo.
Podrán utilizarse en el texto los comandos siguientes: negrita, cursiva, subrayado, superíndices,
subíndices, sangrado y tabuladores, si los autores los consideran necesarios para resaltar alguna
parte concreta.
Las tablas deben estar concebidas y estructuradas de tal modo que puedan leerse y entenderse
por sí mismas, con independencia del texto. Se recomienda hacerlas con el “Programa Tablas”
de los procesadores ordinarios de textos. En el caso de no utilizar estos programas, las columnas
deben marcarse con tabuladores, nunca con la barra espaciadora. A ser posible, no llevarán líneas
de separación de columnas, ni irán enmarcadas. Se procurará que las líneas horizontales queden
reducidas a una o dos arriba y a una o dos abajo. Irán numeradas con caracteres arábigos. El
tamaño mínimo de letra será equivalente a 9 puntos del tipo Times New Roman. Se recomienda
no utilizar la palabra cuadro en lugar de tabla.
Títulos de las tablas
El título irá en la parte superior, centrado, con letra normal, minúscula y negrita. Se traducirá
al inglés y al español, si éstos no son los idiomas originales. Las traducciones, en letra cursiva y
minúscula, se situarán inmediatamente debajo del título en idioma original. Líneas a 1,5 espacios.
Las figuras deben estar concebidas y diseñadas de tal modo que puedan leerse y entenderse
por sí mismas, con independencia del texto. Se harán con la hoja de cálculo Excel y se enviarán
siempre acompañadas de un pdf de las mismas, para que la imprenta pueda comprobar que no
falta nada. En el caso de que no se puedan enviar en Excel, se enviarán en formato pdf en alta
calidad, tiff o jpg. Vendrán en original aparte del texto, indicando en el texto del artículo el lugar
donde deben ir situadas. Se numerarán con caracteres arábigos. La versión impresa de la revista
no admite colores, por lo que las figuras han de ser comprensibles en la escala de grises. Se
recomienda no utilizar la palabra gráfico o gráfica en lugar de figura.
Pies de las figuras
El pie (titulo de la figura) no formará parte de la figura. Se escribirá en una página
complementaria, en letra normal, minúscula y negrita, con la correspondiente traducción al inglés
y al español, si éstos no son los idiomas originales. Las traducciones, en letra cursiva y minúscula,
se situarán inmediatamente debajo del pie en idioma original. Líneas a 1,5 espacios.
Las fotografías digitales deben enviarse en archivos TIF, JPG o PSD, con una calidad mínima
de 300 ppp. Se publicarán en blanco y negro.
Referencias bibliográficas (dentro del texto)
Todas las referencias bibliográficas que aparezcan en el texto deben figurar también en el
apartado de REFERENCIAS BIBLIOGRÁFICAS, situado al final del texto, y viceversa.
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
Si el nombre/s del autor/es no forma parte del texto se citarán solamente los apellidos, sin
iniciales, entre paréntesis, en letra minúscula, seguidos del año de la publicación, separado por una
coma, en el lugar que corresponda. Ejemplos: Caso de un autor “...(Treviño, 1975a)...”, caso de
dos autores “...(Amella y Ferrer, 1977)...”, caso de más de dos autores “...(Arevalillo et al., 1979;
Miró-Granada et al., 1975)...”
Si el nombre/nombres del autor/es forma parte del texto se pone el año entre paréntesis.
Ejemplos: “...según los trabajos de Treviño (1975a), Miró-Granada et al. (1975), Lazenby (1969)
y García-Navarro et al. (2009),...”. También en este caso deben ponerse sólo los apellidos, sin las
En el caso de revisiones científicas se admite que las referencias dentro del texto se indiquen
por un número entre paréntesis, si el autor lo considera conveniente, para evitar que el texto sea
demasiado repetitivo y largo.
Las REFERENCIAS BIBLIOGRÁFICAS se ordenarán por orden alfabético de apellidos del
autor o primer autor, si son varios. Para distintos trabajos de un mismo autor, o autores, se seguirá
el orden cronológico del año de publicación. Si en un mismo año hay más de una publicación de
un autor, o autores, se distinguirán añadiendo una letra al año de publicación. Ejemplo: 1994a,
1994b. Líneas a 1,5 espacios. No se sangrarán los párrafos. Será la imprenta quien introduzca
posteriormente una sangría francesa en las distintas referencias. Las palabras REFERENCIAS
BIBLIOGRÁFICAS, que definen el apartado, irán en letra normal, MAYÚSCULA y negrita,
justifcadas a la izquierda.
Categoría de TITULAR MAYOR.
Forma de presentación de las referencias al final del texto:
Caso de revistas:
APELLIDO/S INICIAL/ES [del nombre],...,…,… Y APELLIDO/S INICIAL/ES [del nombre] [de
los autores] (año) Título del artículo. Nombre de la revista [en cursiva], volumen(número) [en
negrita], 1ª página-última página (del artículo).
AMELLA A. Y FERRER C. (1977) Utilización del método fitológico en la determinación del
valor nutritivo de pastos. Pastos, 7(2), 270-279.
AREVALILLO A.M., GONZÁLEZ G. Y GONZÁLEZ V. (1979) Influencia de la frecuencia
de siega sobre el rendimiento, la composición y la digestibilidad “in vitro” de la alfalfa Aragón
(Medicago sativa L.) en regadío. Pastos, 9(2), 147-155.
MIRÓ-GRANADA L., LEÓN A. Y FORTEZA DEL REY V. (1975) Evaluación de recursos y
criterios de actuación en la mejora pratense. Pastos, 5(1), 220-238.
TREVIÑO MUÑOZ J. (1975a) Influencia del momento de siega sobre la productividad de la
alfalfa, medida por los rendimientos en proteína y energía. Pastos, 5(1), 239-246.
PASTOS 2012. ISSN: 0210-1270
PASTOS, 42 (2), 317 - 329
Instrucciones para autores de PASTOS
Caso de libros de un solo autor o grupo de autores para toda la obra:
APELLIDO/S INICIAL/S [del nombre],...,… Y APELLIDO/S INICIAL/S [del nombre] [de los
autores] (año) Título del libro [en cursiva]. Ciudad de la Editorial, País: Nombre de la Editorial.
REMON ERASO J. (1991) Las plantas de nuestros prados. Madrid, España: Ediciones MundiPrensa.
FRAME J., CHARLTON J.F.L. Y LAIDLAW A.S. (1998) Temperate forage legumes. Wallingford,
UK: Commonwealth Agricultural Bureaux.
Caso de libros colectivos, con capítulos escritos por distintos autores:
APELLIDO/S INICIAL/S [del nombre],...,… Y APELLIDO/S INICIAL/S [del nombre] [de
los autores] (año) Título del artículo o capítulo. En: Apellido/s Inicial/s [del nombre],...,… y
Apellido/s Inicial/s [del nombre] [de los editores] (Ed, si es solamente un editor, o Eds, si son dos
o más editores) Título del libro (en cursiva), pp. 1ª página-última página (del artículo o capítulo).
Ciudad de la Editorial, País: Nombre de la Editorial.
En el caso de que haya más de dos editores se pondrá solamente el primero seguido de las
palabras et al.
Ejemplos [con uno o dos editores]:
LAZENBY A. (1969) The pastoral complex. En: James B.J.F. (ed) Intensive utilization of pastures,
pp. 105-124. Sydney, Australia: Angus and Robertson Ltd.
TAINTON N.M. Y MENTIS M.T. (1984) Fire in grassland. En: Booysen P.V. y Tainton N.M.
(eds) Ecological effects of fire in South African ecosystems, pp. 115-147. Berlin and Heidelberg,
Alemania: Springer-Verlag.
Ejemplo [con tres o más editores]:
GARCÍA-NAVARRO R., ALVARENGA J. Y CALLEJA A. (2009) Efecto de la fertilización
fosfórica sobre la presencia de especies en el forraje de prados de montaña. En: Reiné R. et al.
(Eds) La multifuncionalidad de los pastos: producción ganadera sostenible y gestión de los
ecosistemas, pp 197-203. Huesca, España: Sociedad Española para el Estudio de los Pastos.
En el caso de que se hayan utilizado números para las citas dentro de texto, las referencias de
final de texto irán precedidas por el número correspondiente que la identifique.
Título, resumen y palabras clave en idioma alternativo
- Si el idioma original es el español se traducirá el título y resumen al inglés.
- Si el idioma original es el inglés se traducirá el título y resumen al español.
- Si el idioma original es distinto del español o inglés se traducirán título y resumen al inglés y
al español.
- Los resúmenes tendrán una extensión máxima de 300 palabras para artículos científicos y
notas de investigación, y de 450 para revisiones, e irán seguidos de un máximo de cinco
palabras clave, no contenidas en el título, en el idioma correspondiente.
Por ser lo más habitual que el idioma original sea el español, el idioma alternativo es el inglés.
Por ello, a continuación se hace referencia al resumen y palabras clave en inglés:
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
Centrado, con letra normal, MAYÚSCULA y negrita. Líneas a 1,5 espacios.
Debe ser informativo, no indicativo, para permitir al lector apreciar el contenido e interés del
trabajo. Debe informar sobre objetivos, metodología, resultados y conclusiones. Máximo de 300
palabras para artículos científicos y notas de investigación, y 450 para las revisiones científicas.
Anchura de líneas y tipo de letra igual al resto del artículo. Líneas a 1,5 espacios. Sangrar la
primera línea de cada párrafo utilizando el tabulador, no la barra espaciadora. En su contenido no
debe haber referencias ni al texto, ni a las figuras, ni a las tablas del artículo resumido. La palabra
SUMMARY irá en letra normal MAYÚSCULA, negrita y justificada a la izquierda.
Categoría de TITULAR MAYOR.
Key words
El resumen irá seguido de un máximo de cinco key words que no estén contenidas en el título.
Se escribirán en letra normal y minúscula, separadas por comas. La última irá seguida de punto.
Las palabras Key words irán en letra normal, minúscula y negrita, justificada a la izquierda y
seguida de dos puntos, después de los cuales irán las key words.
Categoría de titular de 1er orden.
Para las unidades de medida se seguirá el SI (Sistema Internacional de Unidades), del que
se recoge un resumen en el Apéndice 2 de estas instrucciones.
Los símbolos de las unidades se imprimen en caracteres romanos (rectos). En general, los
símbolos se escriben en minúsculas (antiguamente se escribía Kg, Ha, Km), salvo si se trata de
la primera palabra de una frase o del nombre “grado Celsius”, quedando invariables en plural.
Nunca los símbolos van seguidos de punto, salvo si se encuentran al final de una frase. En este
caso el punto corresponde a la ortografía habitual de la frase pero no forma parte del símbolo (es
incorrecto escribir kg., ha., km.).
El símbolo de litro será L cuando vaya precedido por un número y l cuando lo sea por un
prefijo de fracción (ejemplo, ml). Cuando las unidades no vayan precedidas por un número se
expresarán por su nombre completo, sin utilizar su símbolo. Ejemplos de símbolos comunes:
kilogramo = kg, hectárea = ha, metro = m, kilómetro = km. (en este último caso el punto no forma
parte del símbolo, se pone porque es final de frase).
Expresion algebraica de los símbolos de las unidades SI
1. Multiplicación. Cuando una unidad derivada está formada multiplicando dos o varias
unidades, los símbolos de las unidades se separarán por puntos a media altura o por un
espacio. Ejemplo: N·m o N m. Dada la dificultad de escribir el punto a media altura, se
recomienda utilizar el espacio.
2. División. Cuando una unidad derivada está formada dividiendo una unidad por otra, se
puede utilizar una barra inclinada (/), una barra horizontal o exponentes negativos.
Ejemplo: m/s o m s-1. No debe utilizarze la barra inclinada y los exponentes negativos en un
mismo artículo. Hay que optar por uno de los dos.
3. Nunca, en una misma línea, debe seguir a una barra inclinada un signo de multiplicación
o de división, a no ser que se utilicen paréntesis para evitar toda ambigüedad. Ejemplo 1:
PASTOS 2012. ISSN: 0210-1270
PASTOS, 42 (2), 317 - 329
Instrucciones para autores de PASTOS
m/s2 o m s-2, son expresiones correctas, pero m/s/s, es incorrecta. Ejemplo 2: m kg/(s3 A) o
m kg s-3 A-1, son expresiones correctas, pero m kg/s3/A y m kg/s3 A, son incorrectas.
Notación numérica
1. En el texto se utilizarán palabras para los valores de cero a nueve y cifras para los valores
2. Debe dejarse un espacio entre grupos de tres dígitos, tanto a la izquierda como a la derecha
de la coma (15 739,012 53). En números de cuatro dígitos puede omitirse dicho espacio.
Los números de los años deben escribirse sin separar el primer dígito del segundo (es
correcto escribir año 2011). Ni el punto, ni la coma deben usarse como separadores de
los miles. Ejemplo: el número ciento veintitres millones trescientos veinticinco mil ciento
setenta se escribe 123 325 170 (123.325.170 o 123,325,170 son formas incorrectas).
3. Las operaciones matemáticas solo deben aplicarse a símbolos de unidades (kg/m3) y no a
nombres de unidades (kilogramo/metro cúbico).
4. Debe estar perfectamente claro a qué símbolo de unidad pertenece el valor numérico y qué
operación matemática se aplica al valor de la magnitud. Ejemplo: es correcto escribir 35 cm
x 48 cm o 100 g ± 2 g ( 35 x 48 cm o 100 ± 2g son formas incorrectas).
Cifras decimales
Dentro del texto en español
Se separarán de la parte entera por una coma abajo (,). Ejemplo: 10,17 (10.17 es forma
Dentro del texto en inglés (summary)
Se separarán de la parte entera por un punto a media altura (·). Ejemplo: 10·17 (10,17 o 10.17
son formas incorrectas). Dada la dificultad de escribir el punto a media altura, se acepta en su
lugar el punto abajo (Ejemplo: 10.17 es correcto para PASTOS).
Las siglas, materiales o equipos poco usuales se definirán cuando se utilicen por vez primera,
indicando el nombre y dirección del fabricante en el caso de equipos.
El nombre botánico de las plantas se escribirá en cursiva, en letra minúscula, con excepción de
la primera del género, que será mayúscula. El nombre de las variedades comerciales, o cultivares,
se escribirá con letra normal y entre comillas simples o bien con letra normal precedido de cv
(símbolo de cultivar) cuando sigan al nombre botánico de la especie. Ejemplo: Lolium multiflorum
Lam. ‘Tama’ o Lolium multiflorum Lam. cv Tama.
En el caso de cultivos de microorganismos se indicará la procedencia y denominación cuando
estén depositados en colecciones reconocidas.
Los nombres vulgares de plantas deben ir seguidos del nombre botánico entre paréntesis la
primera vez que aparezcan en el texto.
Son las denominaciones de los distintos apartados del trabajo. Se jerarquizarán del modo
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
TITULARES MAYORES: en mayúsculas, letra normal y negrita.
Titulares de primer orden: en minúsculas, letra normal y negrita.
Titulares de segundo orden: en minúsculas, letra cursiva y negrita.
Titulares de tercer orden: en minúsculas, letra cursiva y no negrita.
TÍTULO PRINCIPAL (centrado, máximo 25 palabras)
Título abrevido (centrado, máximo 50 caracteres, incluídos espacios)
Dirección de los autores (centrado)
RESUMEN en idioma original (máximo 300 palabras para artículos científicos y notas de
investigación, y 450 para revisiones científicas)
Palabras clave en idioma original (máximo cinco)
DISCUSIÓN (puede unirse al anterior)
AGRADECIMIENTOS (puede faltar)
Si el inglés no es idioma original:
SUMMARY (máximo 300 palabras para artículos científicos y notas de investigación, y
450 para revisiones científicas)
Key words (máximo cinco)
Si el español no es idioma original:
RESUMEN en idioma español (máximo 300 palabras para artículos científicos y notas de
investigación, y 450 para revisiones científicas)
Palabras clave en idioma español (máximo 5)
Formato: Los textos originales se escribirán con letra Times New Roman de 12 puntos, o
equivalente, en líneas a 1,5 espacios. El texto se justificará únicamente por la izquierda.
Esta norma afectará a todo el artículo: título, nombre de autores, dirección de autores,
PASTOS 2012. ISSN: 0210-1270
PASTOS, 42 (2), 317 - 329
Instrucciones para autores de PASTOS
resumen, palabras clave, introducción, material y métodos, resultados, discusión, títulos de
tablas, pies de figuras, conclusiones, agradecimientos, referencias bibliográficas, título en
inglés, summary y key words, con excepción de las tablas, que podrán ir a un espacio
entre líneas y con caracteres más pequeños, si fuese necesario, hasta un mínimo de
9 puntos. La primera línea de cada párrafo se sangrará con tabulador, nunca con la barra
Extensión: La extensión de los mismos no excederá de 30 páginas para los artículos científicos,
10 para las notas de investigación, cinco para conclusiones de reuniones científicas
y resúmenes de tesis doctorales, y una para descripción de variedades y recensiones de
libros, incluyendo en estos límites las figuras, las tablas y las referencias bibliográficas.
Para las revisiones científicas no hay un límite prefijado de páginas.
Referencias bibliográficas (en texto): Sólo apellidos y año, en minúsculas. No usar las iniciales
del nombre.
AUTORES en letra normal y mayúscula. Apellidos e iniciales de editores en letra normal
y minúscula. Resto en letra normal y minúscula. Nombre de las revistas y títulos de libros,
en letra cursiva. Volumen y número de la revista en negrita.
Tablas (No utilizar la palabra cuadro/s): Se recomienda hacerlas con el “Programa Tablas” de los
procesadores ordinarios de textos. En el caso de no utilizar estos programas, las columnas
deben marcarse con tabuladores, nunca con la barra espaciadora. A ser posible, no
llevarán líneas de separación de columnas, ni irán enmarcadas. Se procurará que las líneas
horizontales queden reducidas a una o dos arriba y a una o dos abajo. Irán numeradas con
caracteres arábigos. El tamaño mínimo de letra será equivalente a 9 puntos del tipo Times
New Roman
Títulos de tablas: En la parte superior de la tabla, centrado. Letra normal, minúscula y negrita.
Títulos de tablas en idioma/s alternativo/s: Inmediatamente debajo del título en idioma
original. En letra cursiva, no negrita y minúscula.
Figuras (no utilizar la palabra gráfico); Se harán con la hoja de cálculo Excel y se enviarán
siempre acompañadas de un pdf de las mismas, para que la imprenta pueda comprobar
que no falta nada. En el caso de que no se puedan enviar en Excel, se enviarán en formato
pdf en alta calidad, tiff o jpg. Vendrán en original aparte del texto, indicando en el texto
del artículo el lugar donde deben ir situadas. Se numerarán con caracteres arábigos.
Pies de figuras: En página aparte. Letra normal y negrita.
Pies de figuras en idioma/s alternativo/s: Inmediatamente debajo del pie en idioma
original. Letra cursiva, no negrita y minúsculas.
Idiomas: Siempre habrá un resumen y palabras clave en español e inglés, cualquiera que sea
el idioma original. Los títulos de tablas y pies de figuras se escribirán siempre en español
e inglés, cualquiera que sea el idioma original. El número máximo de palabras de resumen
y palabras clave, el tipo de letra y el espacio entre líneas serán los mismos que para el caso
del idioma original.
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
Intensidad de corriente eléctrica
Cantidad de sustancia
Unidad básica
Magnitudes derivadas
Masa en volumen
Volumen másico
metro cuadrado
metro cúbico
metro por segundo
kilogramo por metro cúbico
metro cúbico por kilogramo
Minuto de ángulo
Segundo de ángulo
l, L
Valor en unidad SI
1 min = 60 s
1 h = 60 min = 3600 s
1 d = 24 h = 86 400 s
1º = (π/180) rad
1’ = (1/60)º = (π/10 800) rad
1’’ = (1/60)’ = (π/648 000) rad
1 l = 1 dm3 = 10-3 m3
1 t = 103 kg
1 a = 1 dam2 = 100 m2
1 ha = 1 hm2 = 104 m2
1 bar = 0,1 MPa = 100 kPa = 105 Pa
PASTOS, 42 (2), 317 - 329
PASTOS 2012. ISSN: 0210-1270
Instrucciones para autores de PASTOS
Grazing Systems
and Biodiversity in
Latin American Areas:
Colombia, Chile and
Coordinated by
Sergio Guevara Sada and
Javier Laborde
Cover photography credits:
Gerardo Sánchez Vigil
Mariano Guevara Moreno-Casasola
Adi E. Lazos R.

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