Degradation of agro-pastoral village land in semi-arid southeastern Cuba

Degradation of agro-pastoral village land in semi-arid southeastern Cuba

ARTICLE IN PRESS Journal of Arid Environments Journal of Arid Environments 59 (2004) 299–311 www.elsevier.com/locate/jnlabr/yjare Degradation of agro...

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ARTICLE IN PRESS Journal of Arid Environments Journal of Arid Environments 59 (2004) 299–311 www.elsevier.com/locate/jnlabr/yjare

Degradation of agro-pastoral village land in semi-arid southeastern Cuba A. Wezel*, S. Bender International Nature Conservation, Institute for Zoology, University of Greifswald, Grimmer Str. 88, Greifswald 17487, Germany Received 8 November 2002; received in revised form 8 August 2003; accepted 5 December 2003

Abstract Land degradation was investigated in a mountainous semi-arid area at the coast of southeastern Cuba in 2001. The study area included two villages and part of a nature reserve. Degree of degradation was investigated by walking across the study area to delineate different land use classes. Further information about the area was collected by interviewing 24 farmers and conducting rankings with key persons of the study villages. Six land use classes were distinguished: village areas, highly, moderately, and little degraded areas, areas with high tree density and areas with natural vegetation. Grazing was found to be the main cause of degradation in the study area, followed by wood cutting and forest fires. Impact of grazing was found to be highest near the villages, decreasing with greater distance towards the mountains. Woody vegetation is mostly cut to provide firewood or charcoal for the local population. The vegetation of the Nature Reserve is only affected to smaller degrees by grazing and wood cutting due to geographical factors limiting access. Some management options to combat land degradation, such as better stoves to reduce lavish use of firewood, seem to be feasible, but most options are limited by lack of finances, lack of personal responsibility or the failure to see short-term effects. Although villagers’ perception of degradation factors such as over-grazing is low, local initiatives to conserve their natural environment exist. r 2004 Elsevier Ltd. All rights reserved. Keywords: Caribbean; Coastal Cuba; Grazing; Livestock; Protected area; Semi-desert

*Corresponding author. Tel.: +49-3834-864185; fax: +49-3834-864187. E-mail addresses: [email protected] (A. Wezel), [email protected] (S. Bender). 0140-1963/$ - see front matter r 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.jaridenv.2003.12.010

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1. Introduction Livestock production in rural areas of Cuba takes place in different domains. The most important one, excluding poultry, is cattle production on fenced pastures in humid and sub-humid areas. Between 1998 and 2001, the number of cattle was about 4.38 million (FAO, 2002). Pig production is second with about 2.55 million pigs (1998–2001), followed by sheep (0.31 million) and goats (0.21 million). The latter three species are produced by state farms and co-operatives, but also by individual households which raise a limited number of animals on the farmyard. Often these animals roam freely during daytime around the houses, along streets or on unused land near villages. Most of the cattle pastures are found in the larger humid parts of Cuba, only a smaller part, and less recognized, in the semi-arid area of coastal southeastern Cuba, the Franja Costera Sur. In this region, herds of cattle, sheep and goats are often kept on large tracts of freely accessible state land; only few pastures are fenced. In contrast, pastures in the humid areas are normally fenced or enclosed by live fencing with Gliricidia sepium. In total, pasture land in Cuba covered 3.89 Mio. ha in 1945 (World Bank, 1951 cited in D!ıas-Briquets, 2000), decreased after the Cuban revolution to 1.90 Mio. ha in 1961, increased to 2.97 Mio. ha until 1989 and decreased again gradually after the collapse of the Soviet Union to 2.20 Mio. ha in 1999 (FAO, 2002). Livestock production in the Franja Costera Sur region can be traced back to the 18th century, when the Spanish practiced sheep breeding. From the beginning of the 19th century until the mid 20th century, forests were cleared to extend pasture areas, but also for timber export and wood for the construction of ships. In 150 years forest areas were reduced by 38% (Gonza! lez, 1999). At the end of the 1940s, a large livestock production enterprise was installed at Baitiquir!ı, one of the study villages. After the Cuban Revolution in 1959, this enterprise was nationalized and moved to the neighbouring town Antonio del Sur in 1985. This, together with the ‘special period in times of peace’ in Cuba caused by the collapse of the Soviet Union (Alvarez, 2000), led to a significant decrease of livestock numbers in this region (Gonza! lez, 1999), but degradation of the area still continued. To investigate degradation in semi-arid Franja Costera Sur, the present study analyses the situation of land use, in particular livestock production, and its influence on vegetation and soils in a selected area. In addition, degradation of village land was compared to that of a protected area.

2. Materials and methods The study was conducted from July to October in 2001 in the Franja Costera Sur region in the Bay of Baitiquir!ı (Fig. 1). There are two villages located in the area: Baitiquir!ı with 865 person and El Naranjo with 419 inhabitants in 2001 (personal communication, Statistical Office San Antonio del Sur, 2001). Most people in the villages are pensioners or have a regular job as technicians in road construction, hydraulic engineering or in a state-run agricultural enterprise and cooperatives.

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Fig. 1. Location of the study area in eastern Cuba.

Some of them are also engaged in part-time agricultural activities. A minority of the people in the study villages are full-time private farmers. The two villages are located adjacent to the Nature Reserve Baitiquir!ı, parts of which were included in this study. This protected area was proposed as a Nature Reserve of national significance in 1975 and is administered as an ecological monitoring and research area since 1978 (Bender, 2002). According to the new legislation about protected areas in Cuba, the first 32 protected areas were accepted by the Council of Ministers in February 2001. Additional 232 protected areas, including the Nature Reserve Baitiquir!ı, are enrolled on the proposal list of the Ministry of Science, Technology and Environment (CITMA) and thus receive a restricted, preliminary protection (Niekisch and Wezel, 2003). The Nature Reserve Baitiquir!ı is listed as a protected area of national importance (CNAP, 2002). The climate of the study area is semi-arid with an annual precipitation of 400– 500 mm. There are two rainy seasons: a short one during May and a longer one from August to November. Mean annual temperature is about 27 C. The aridity is caused by its position in the rain shadow of the Sagua de Baracoa mountains which block the dominating northeast and east winds from the Caribbean Sea (Instituto de Meteorolog!ıa de la Academia de Ciencias de Cuba, 1987). Natural vegetation of the study area is characterized by different types of xeromorphic formations, including semi-desert cactus scrub, as well as coastal vegetation such as mangroves and coastal rock pavement vegetation (Borhidi, 1991). In this region, mostly Leptosols, but also Cambisols, Ferralsols and Fluvisols (according to FAO-classification) are found. Dominant land use practice is grazing with cows, sheep and goats. Where irrigation is possible, different crops are cultivated either in homegardens or on fields outside the villages. During field work, land use and degree of land degradation was investigated by walking a few times across the study area, often along paths, roads or mountain ridges, to get a first impression. Throughout these walks, it became obvious that six major land use classes are suitable to bring together the major features of the study area. Then, further walks were carried out to exactly delineate and transfer them into a topographic map. The various grazing areas to be outlined were visually

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distinguished by the parameter shrub density, cover of herb layer and crusted or bare soil areas. The following land use types were distinguished: (i) village areas, (ii) highly degraded grazing areas (large areas of bare soil, soil crusts, shrub density strongly reduced or missing, very reduced cover of herb layer), (iii) moderately degraded grazing areas (reduced cover of herb layer, scattered shrubs, livestock paths between shrubs, almost no soil crusts), (iv) little degraded grazing areas (normally a continuous cover of herb layer existent, medium to high shrub densities, no soil crusts or bare soil areas), (v) areas with high tree densities and undergrowth shrubs, (vi) areas with undisturbed or almost undisturbed natural vegetation. Within the land use units delineated, special areas with specific forms of degradation or land use were included into the map to enrich the map with additional small-scale information (see consecutive numbers in Fig. 2). Further information about the study area was gathered by using participatory methods such as semi-structured interviews and rankings during a survey from July to October 2001. Interviews were held with 12 private farmers in each of the two villages. They were interviewed about their socio-economic situation, agricultural land use and resource use related problems. Farmers’ households were selected by the staff

Fig. 2. Land use map of the Bay of Baitiquir!ı study area, Cuba.

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of the Ecological Station and thus could not be randomly chosen. However, as the number of 24 farmers in the two study villages covered about 85% of the private farmers existent in the two villages, the interviews can be considered as to be representative. Besides the interviews, different rankings were conducted with key persons of the two study villages such as representatives of the Communist Party and the Federation of Cuban Women, the mayor, the leader of the local school, doctors, scientists and staff from the Ecological Station in Baitiquir!ı. Rankings in each village were carried out to the questions: ‘‘Which problems exist in the village related to resource use’’, and ‘‘Which elements are used or exist for environmental education in the village’’. In the current study, only the rankings concerning resource use related problems are presented. Rankings were conducted in groups of 4–12 persons. First, responses or ideas were collected through brainstorming with the group, and were summarized under various items by the group. Then each person could individually distribute a certain amount of points to the different items. Finally the points distributed by the participants to the different items were summed up and the items ranked according to their sum of points. Often, not all items collected for the rankings received points by the participants. Scientific plant names in this study follow Rehm and Espig (1996) for cultivated plants and Borhidi (1991) for natural ones.

3. Results 3.1. Household characteristics On average, a farmer’s household consists of four persons. All farmers interviewed are owners of houses and small gardens. Three out of five farmers own private land besides the house area, which is used to about 25% for cultivation, the other area as private pastures. Nine farmers with private land live in El Naranjo (see Fig. 2), where agriculture is more widely practiced by the villagers compared to Baitiquir!ı (5 farmers with private land). The size of private agricultural land holdings in the two study villages varies enormously between 1.5 and 67 ha. Average size of private land per household in Baitiquir!ı is 9.5 ha (range: 1.5–30.9 ha); in El Naranjo it is 27.5 ha (range: 4.0–67.1 ha). Among the farmers of El Naranjo are three big farmers who own over 53 ha of private land. Four farmers possess land of a more than 100 years old heritage; in most other cases, farmers’ private land was obtained shortly before or after the Cuban Revolution in 1959. The rest of the farmers interviewed use state land. Most private land is located 2–4 km from villages. Due to high temperatures, low precipitation and lack of irrigation, cultivation of crops is limited in the study area and only practiced on very few fields and in homegardens. In the latter, mainly fruit trees such as mango (Mangifera indica), avocado (Persea americana), coconut palms (Cocos nucifera) or sweet bananas and plantains (Musa x paradisiaca) as well as different pulses (Cajanus cajan, Phaseolus

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Table 1 Number of livestock of interviewed households engaged in agriculture in the study villages in southeastern Cuba Cows

Sheep

Goats

Baitiquir!ı (n ¼ 12) Average Minimum–maximum Number of farmers with the respective livestock

7 3–16 5

34 5–90 8

8 4–17 9

El Naranjo (n ¼ 12) Average Minimum–maximum Number of farmers with the respective livestock

32 2–135 8

36 2–140 10

18 2–50 10

Total livestock in both villages

291

251

636

lunatus, P. vulgaris) and yams (Dioscorea spec.) are cultivated for self-sufficiency (for detailed plant inventory see Wezel and Bender, 2003). In a few fields of a state enterprise, located between the study villages (see number 1 in Fig. 2), fruit trees and legumes are grown for food provision of towns and villages in the region. One farmer tries to cultivate tobacco (Nicotiana tabacum) for export (2), which is traditionally a crop in western Cuba. Some farmers have remote small fields (3), where they grow maize, beans or fruit trees. In the study villages many people do not keep livestock, but all farmers interviewed own livestock. In El Naranjo, most farmers keep cows, sheep and goats, whereas in Baitiquir!ı most farmers have only sheep and goats. The number of cows and goats owned by farmers in El Naranjo is significantly higher (average and maximum), whereas the number of sheep is similar in both villages (Table 1). The highest numbers of livestock were found with the three big farmers in El Naranjo. In total, 1178 animals were kept by the farmers interviewed in the two study villages. In addition, 186 cattle and 75 calves (in 2001) were kept by the Cuban state on the communal pastures near Baitiquir!ı. 3.2. Grazing In the north-west of the villages Baitiquir!ı and El Naranjo highly degraded areas were found (Fig. 2). Main cause of degradation is over-grazing by livestock. The degradation characteristics are a generally reduced or non-existent grass and herb cover, the dominance of species such as Jatropha gossypiifolia or Calotropis procera (4), the latter a well-known degradation species (Francis, 2002). Large areas show bare soil or broad paths created by trampling of livestock. In some areas Calotropis, Jatropha or different cacti species are cut down by livestock owners or state workers to increase area for growth of herbs and grasses (5); in other locations the cacti were not touched (6). In one area, small shrubs were not pruned to promote regeneration

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of herbs and grasses under the crown of shrubs (7). This area might be affected by water erosion after heavy rainfalls because two riverbeds of temporary small rivers cross this area. The highly degraded areas are mostly state land with free access. Private livestock is normally kept near or in the village overnight, and the state cattle herd is kept in a corral (8). During daytime, the different herds leave the village area with shepherds to move towards the mountains, as fodder resources are scarce in the vicinity of the villages. On their way back, they are watered at a water hole (9) with water of a water pipe from the mountains. In the west of El Naranjo, some fenced pastures also exist (10, 11). The vegetation of the grazed areas is strongly affected by livestock but also by cutting of woody species by the local population. Here, a gradient from the villages to the higher altitudes of the mountains is clearly visible, with a reduced number of trees, e.g. Guajacum officinale, Guasuma species or different cacti species near Baitiquir!ı. In higher altitudes of the mountains dense ‘green’ woody vegetation is found along small temporary rivers (12). At the foot of the mountains, species of the genera Acacia, Albizia and Cassia are found with a relatively high density in a small valley (13). However, also in smaller areas located further away from Baitiquir!ı, relatively highly degraded areas (14) can be found along preferred livestock tracks towards the mountains. Grazing intensity in the Nature Reserve is low-to-moderate at the foot of the mountain ridges; on top of the ridge, vegetation seems not to be influenced by grazing.

3.3. Cutting of trees and shrubs A second important cause for degradation in the study area is cutting of trees and shrubs by the local population. Often this is done in order to get firewood, but also for the production of charcoal. Firewood is cut near the villages (e.g. 15, 16), in areas with high tree density (13), but also at higher altitudes. Cutting of small trees was found in the Nature Reserve as well (17). The natural vegetation of the Nature Reserve is mainly characterized by a semi-desert vegetation with the dominance of columnar and tree-shaped cacti such as Ritterocereus hystrix and Consolea macracantha (Borhidi, 1991). The sea-grape (Coccoloba uvifera) is often dominant on the first limestone terrace along the coast. Along the Bah!ıa de Baitiquir!ı (Bay of Baitiquir!ı), mangrove stands (18, 19) are found, interrupted by a small harbour for the local fishermen, a small base of the Cuban Marine (20) and a saltern (21). At one location, mangroves have been cut and scrap material deposited (18). Charcoal is mainly produced in areas with higher tree density (13). In both study villages, firewood and charcoal are mainly used for cooking because gas, petroleum, kerosene and electricity are not sufficiently available or too expensive. In a small part of the study area, a secondary thicket of the invasive species Dichrostachys cinerea occurs (22). This Mimosaceae has been introduced from

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Africa as an ornamental plant and has spread spontaneously throughout the country (Bisse, 1988). If it is cut down, it regenerates relatively quickly. 3.4. Fire Human-caused bush or forest fires are relatively frequent in the mountains, near the villages, as well as in the Nature Reserve. They are mainly caused by uncontrolled fires of charcoal production or burning cigarettes. Recently burnt zones are clearly visible in the landscape (23, 24), whereas older areas are characterized by burnt tree stumps or large areas of dense grass layer (25). In the area of Baitiquir!ı, 1.722 ha had burned during the period from 1995 to 2000 (Nicomedes Cobas, 2001, personal communication). 3.5. Ranking of resource use related problems Lack of water and increasing desertification (generally used by the villagers as expression for land degradation) are seen as major problems for resource use in both study villages (Fig. 3). In Baitiquir!ı, the insufficient availability of fodder, which is linked to the semi-arid conditions, is ranked even higher. In contrast, this factor was not mentioned in El Naranjo. Here, lack of discipline concerning firewood cutting or bird-chasing, as well as lack of knowledge regarding the environmental problems, was ranked at second place. In both villages, the lack of reforestation was mentioned as well as the factors forest fires and illegal cutting of firewood in Baitiquir!ı. Overgrazing of pasture areas is not really perceived as a problem in the study area. Other factors, such as lack of protection of local fauna, charcoal production in the mountains and slow regeneration of the vegetation after forest fires, were listed by the key persons during brainstorming, but did not receive points during the rankings.

Fig. 3. Resource use related problems as seen by key persons of the two study villages.

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4. Discussion Presently, the most important factor for land degradation in the study area is grazing. Close to the villages, where more livestock passes, the land is more degraded than in remote areas. Features such as small erosion gullies along preferred livestock paths, pruning of unwanted species and expansion of indicator species for overgrazing such as Calotropis procera or Jatropha gossypiifolia indicate an intensive use of the pastures. This is also obvious when comparing vegetation cover in the Nature Reserve, to vegetation cover in the preferred grazing area from north to west of the study villages. In most parts of the Nature Reserve, a continuous vegetation cover is found. In smaller areas only, vegetation cover is reduced by a few grazing animals. In contrast, vegetation in the main grazing areas near the villages is significantly affected. The most important reason why the Nature Reserve seems to be relatively unaffected by grazing is the difficult access. It is almost blocked by the saltern, the Bah!ıa de Baitiquir!ı, the field area between the two villages and, to a smaller degree, by the steepness of the coastal mountain ridges. It is not especially protected by the local population because almost nobody knows about the Reserve status (Bender, 2002). Degradation of the study area due to grazing is not directly recognized by the local population. Instead, insufficient fodder for livestock is seen as a major problem, which of course is linked to overgrazing, or more generally the lack of water and desertification (see Fig. 3). This might be due to the mentioned lack of knowledge and discipline, but also due to the heterogeneity in livestock endowment. In the study villages most people do not keep livestock, others have only few animals and do not rely on income from livestock production because they have a regular job. Thus, degradation of pastures does not seem to be very important or even relevant to them. In contrast, the herdsmen of the state herd as well as some village people realize the changes on the pastures. They cut down unwanted plant species or try to ease regeneration of herbaceous vegetation at one location. To what extent degradation of pastures is affected by the state herd or by private livestock remains an open question. In general, in the free access area north and west of Baitiquir!ı the degradation by livestock of the state herd and private farmers of Baitquir!ı and El Naranjo seems to be more pronounced than in the area west of El Naranjo, where most pastures below 200 m altitude are fenced. Destruction of forests and woody vegetation by cutting of firewood and by forest fires seems to be perceived more clearly by the local population as compared to degradation caused by grazing. This might be due to the fact that almost all people in the villages continuously need firewood for cooking. Burnt zones are also clearly visible in the landscape. Although woody vegetation is affected by browsing of livestock as well, illegal wood cutting and charcoal production are the main causes for its degradation. In many parts of the study area, cut trees and shrubs and the work of charcoal-burners could be observed. The people in the villages know that the collection of firewood or purchase of charcoal affects their local environment, but they often mention lack of discipline as a reason to continue cutting (Bender, 2002). Reduction of woody vegetation is also seen as a lack of reforestation. At present,

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trees are cut for the expansion of fields as well (1, 2). Like the degradation of pastures, the destruction of woody vegetation follows a gradient from the heavily used vicinity of the villages to nearly untouched natural vegetation in remote areas in the mountains. However, also in the more remote mountains forest fires occur (23). Household economics influence resource degradation because farmers depend on their income from livestock production. Cropping is very limited in the region because only homegardens can be irrigated. The part-time farmers who have a regular job and the pensioners also rely on livestock and homegarden production. They mostly receive very low wages or pensions and need the additional agricultural production to assure food security for their households. The protection of the Nature Reserve Baitiquir!ı is of great importance for Cuba because of its particular semi-desert habitats as well as coastal vegetation and mangrove stands. Therefore, the reception of a full legal status from the Council of Ministers in the near future is essential. Access for livestock to the Reserve area is difficult, thus degradation of vegetation by grazing is small. The largest problems are bush fires which occur every few years in the Reserve (24, 25). Regeneration of the natural woody and succulent vegetation occurring normally is often inhibited because a dense grass layer establishes rapidly in burnt areas. Further problems in the Nature Reserve are illegal cutting of trees (17) and the destruction of mangrove stands along the Bahia de Baitiquir!ı. Efforts to stop degradation of woody vegetation and pastures in the study area are few. Near Baitiquir!ı, people try to promote regeneration of herbaceous vegetation by leaving small unpalatable shrubs and spreading pruned tree branches (7). Some private farmers try to protect their land from free moving livestock by fencing. This can be critical in semi-arid areas because of the highly variable inter-annual carrying capacity of pastures due to rainfall variability (Hiernaux, 2001). In some areas, problems with bush encroachment were found (10). After a few years, these areas have to be re-cleared for pastoral use. To minimize the loss of forest areas, reforestation is carried out by the Forest Department. In 2001 about 57 ha were planted with trees (Servicio Forestal San Antonio del Sur, 2001, personal communication). This area is much too small to compensate for an area of about 344 ha annually lost by forest fires. A project of the municipality and the local school for the reforestation of the mangrove stands along the Bah!ıa de Baitiquir!ı also exists, where mangroves should be planted. If degradation is to be stopped, the management of natural resources in the study area has to be significantly improved. This of course might be a difficult problem as management options are limited by various constraints. A general grazing management does not seem realistic at present, as it requires an institution not only in charge, but implementing the measures. The Ecological Station in Baitiquir!ı which belongs to the Ministry of Science, Technology and Environment of Cuba (CITMA) could be such an institution, but currently has other objectives than combating land degradation. Another possibility could be to limit grazing at least in crucial areas such as on ridges, on steep slopes and along temporary

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Table 2 Recommendations for land management and resource conservation in the study area in southeast Cuba Proposed management Grazing Institution for supervision of grazing management Limited grazing on ridges, steep slopes and along the temporary rivers Reduction of livestock numbers

Wood consumption Better stoves Gas stoves Planting of trees in homegardens Planting of wood lots Nature reserve Sensitising of local population Improvement of the legal status

Limitation Ecological Station seems to have other priorities at present Supervision, responsibility Limited influence on the state enterprise; private persons rely on livestock production, problem of compensation

Costs (although low) Too expensive, incentives or credits needed No measure for the near future No measure for the near future

Costs for information material; number of multipliers Very limited influence on national legislation

rivers during certain periods of the year to reduce erosion risk and to promote regeneration of woody species (Table 2). The impact of grazing could also be reduced by reducing livestock numbers. The number of cattle of the state enterprise’s herd could be discussed with the local state authorities. Reductions of livestock numbers for the private owners seem difficult, because farmers rely on livestock production. Perhaps compensation could solve this problem, but that is unlikely to happen. To combat destruction of forest and woody vegetation, various strategies can be followed. One is the reduction of wood consumption in the households. This could be accomplished by using better stoves which reduce the amount of firewood or by new gas-stoves. Presently most households use simple fireplaces surrounded by stones. For most villagers the purchase of gas-stoves is only affordable if incentives or cheap credits are supplied, because gas and gas-stoves are too expensive. Reduction of firewood consumption in the households would also decrease illegal charcoal production in the study area, an important cause for forest fires. Another measure is to increase fire wood production by planting suitable trees in the homegardens or establishing wood lots through the Forest Department. Although some species, mostly exotic ones such as Azadirachta indica or Albizia lebbeck, are presently being tested (26), this is not an option for the near future. To secure the Nature Reserve, reception of the full protection status has to be fostered and the local population must be informed about the protected area and its importance for the region.

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5. Conclusions Degradation of natural pasture and woody vegetation in the study region has reached a high intensity. Only strong management efforts may be able to stop or reverse this crucial trend. Although degradation of the area already started a long time ago, the current situation is characterized by too high numbers of livestock and an over-use of the woody vegetation. In particular, the pastures with free access near the villages are the most heavily degraded areas. Fodder resources in many areas have already been reduced to an extent that farmers are forced to track greater distances with their herds, mostly up to higher altitudes, which are now increasingly degraded. Even though the local population perceives the degradation of woody vegetation to a certain degree, they do not declare overgrazing as a problem. This seems to be strongly related to the individual perspective of the subject. Firewood and charcoal are needed by almost all households in the study villages, whereas livestock production is only pursued by a part of the households. Although some management options could mitigate degradation in the study area, most of them are heavily constrained by lack of financial sources, lack of personal responsibility or failure to see short-term effects. Nevertheless, various villagers are ready to participate in local environmental initiatives such as planting of cacti or mangroves, an environment theatre group or active pupil groups (Bender, 2002). At present, the most practicable option to combat degradation seems to be the introduction of better stoves to reduce lavish use of firewood because it can be implemented in the short-term and with low costs.

Acknowledgements We gratefully acknowledge the help of Adalberto Matos Leyva during field work. We thank the DAAD (German Academic Exchange Service) for funding and the people from CITMA in the province of Guanta! namo, particularly the Ecological Station in Baitiquir!ı, for their support during the research in Cuba. We also thank Eva Schlecht, Elke Mannigel and Pam Knittel for improvements on the manuscript.

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