Preventing crop raiding by the Vulnerable common hippopotamus Hippopotamus amphibius in Guinea-Bissau

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Preventing crop raiding by the Vulnerable common hippopotamus Hippopotamus amphibius in Guinea-Bissau
Preventing crop raiding by the Vulnerable
             common hippopotamus Hippopotamus amphibius
             in Guinea-Bissau
                  LUIS M. GONZÁLEZ, FRANCISCO G. D. MONTOTO, TOME MERECK, JUNIOR ALVES
                                 JOSÉ PEREIRA, PABLO FERNÁNDEZ DE LARRINOA, ANA MAROTO
                                                        L U I S B O L O N I O and N U R I A E L - K A D H I R

             Abstract Guinea-Bissau is host to the westernmost subpo-                              protection, such as killing the wild animals involved in
             pulation of the common hippopotamus Hippopotamus                                      crop raiding, are limited, and thus human−wildlife conflict
             amphibius, which is one of only two known populations in-                             jeopardizes the efficacy of protected areas by undermining
             habiting coastal waters. The presence of hippopotamuses                               local support for conservation (Fungo, ). To resolve
             causes conflict with rice farmers as a result of crop damage                          the conflict, wildlife managers usually mitigate the crop raid-
             and the absence of effective measures to protect crops. To                            ing activities of wild animals; for example, by scaring animals
             develop an effective method for protecting rice fields, we                            away from crops, using scare-shooting and barriers, or com-
             studied the patterns of access to flooded and rain-fed rice                           pensating farmers for crop loss. However, the latter measure,
             fields by hippopotamuses and assessed the effect of the in-                           particularly when it involves cash payments, is open to
             stallation of electric fences. Hippopotamuses were detected                           corruption and some farmers may be less likely to improve
             in % of the flooded fields (n = ) and in .% of the                            their management practices or adopt new ones to reduce
             rain-fed fields (n = ). They were detected more frequently                          conflict. Thus, bad managers may be compensated at the ex-
             in fields on offshore islands than on the mainland, in un-                            pense of those who invest in good management techniques
             fenced than in fenced fields, and in fields closer to running                         on their own initiative (Woodroffe et al., ; Fungo, ).
             water. Hippopotamuses entered fenced flooded fields less                                  The common hippopotamus Hippopotamus amphibius,
             frequently than unfenced, and were detected most frequently                           categorized as Vulnerable on the IUCN Red List (Lewison
             at the end of the rainy season and the start of the dry season,                       & Oliver, ), is a major source of human−wildlife con-
             and in the period of vegetative stem growth. Electric fences                          flict in Africa. It causes damage to crops and attacks people
             were an effective deterrent and facilitated increased rice pro-                       (Mkanda, ; Kendall, ). The decline of the species has
             duction. The maintenance and cost of the electric fencing                             been attributed to killing of hippopotamuses by farmers,
             were acceptable to farmers, and therefore the use of such fen-                        and the loss of grazing grounds as a result of agricultural de-
             cing is recommended to resolve the conflict between hippo-                            velopment (Eltringham, ; Lewison & Oliver, ).
             potamuses and farmers in Guinea-Bissau and in other areas                                 The hippopotamus population in Guinea-Bissau is the
             with similar conditions.                                                              westernmost population (Eltringham, ) and is one of
                                                                                                   only two known populations to use seawater and inhabit
             Keywords Common        hippopotamus, crop raiding,
                                                                                                   coastal areas (the other is the subpopulation in Loango
             electric fence, Guinea-Bissau, Hippopotamus amphibius,
                                                                                                   National Park, Gabon; Limoges & Robillard, ; Michez,
             human−wildlife conflict, rice fields
                                                                                                   ). During the th century the hippopotamus became
                                                                                                   rare in Guinea-Bissau, where it was considered to be a game
                                                                                                   species. The decline has been linked to shooting of hippo-
             Introduction                                                                          potamuses in retaliation for crop damage; for example, in
                                                                                                   the region of Cacheu during −  hippopotamus
                                                                                                   deaths resulted from conflict with farmers (Lopes, ).
             C  rop raiding by wild animals results in substantial finan-
                cial losses for farmers living in proximity to wildlife.
             When wildlife is protected, traditional means of crop
                                                                                                   In – the hippopotamus was declared a protected
                                                                                                   species in Guinea-Bissau and two protected areas with
                                                                                                   hippopotamus populations were established: Orango
                                                                                                   National Park and Cacheu Natural Park (IBAP, ).
             LUIS M. GONZÁLEZ (Corresponding author) and FRANCISCO G. D. MONTOTO
             Subdirección General de Medio Natural, Ministerio de Medio Ambiente,                  There are estimated to be c.  individuals in Cacheu,
             Plaza San Juan de la Cruz s/n, Madrid 28071, Spain                                    – in Orango and – in the areas of Bissorao and
             E-mail lmgonzalez@magrama.es
                                                                                                   Carantaba (Lopes, ; Silva & Monteiro, ; IBAP,
             TOME MERECK, JUNIOR ALVES, JOSÉ PEREIRA, PABLO FERNÁNDEZ DE LARRINOA, ANA             ; Silva, ).
             MAROTO, LUIS BOLONIO and NURIA EL-KADHIR Fundación CBD-Hábitat,
             Madrid, Spain                                                                             Hippopotamuses require large areas of savannah grass-
             Received  June . Revision requested  July .                                lands, where they feed on stems of herbaceous species,
             Accepted  August . First published online  January .                      and in times of food stress they will also feed in cultivated

                                                                                        Oryx, 2017, 51(2), 222–229 © 2016 Fauna & Flora International doi:10.1017/S003060531500109X
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. https://doi.org/10.1017/S003060531500109X
Preventing crop raiding by the Vulnerable common hippopotamus Hippopotamus amphibius in Guinea-Bissau
Preventing crop raiding by hippopotamus                      223

                                                                                                                               FIG. 1 Locations of the rice fields (black
                                                                                                                               filled circles) studied in Cacheu Natural
                                                                                                                               Park, Bissorao and Carantaba, on
                                                                                                                               mainland Guinea-Bissau, and in the
                                                                                                                               Orango Islands National Park and Uno
                                                                                                                               Island, with the hippopotamus
                                                                                                                               Hippopotamus amphibius resting places
                                                                                                                               (filled white circles) identified in
                                                                                                                               previous studies (Silva, ).

           rice fields if available (Eltringham, ; Kendall, ). Rice                      is not sustainable, and shooting has not been effective in
           production is one of the main economic activities of the                              minimizing crop damage in other countries with similar
           rural population of Guinea-Bissau and is the most import-                             problems (Mkanda, ; Fungo, ), and may cause hippo-
           ant livelihood activity for people in the protected areas of                          potamuses to become more aggressive (Eltringham, ).
           Orango and Cacheu (Alves & Barros, ; Medina, ).                               The hippopotamus is a charismatic species that attracts tour-
           Past surveys with farmers in Guinea-Bissau showed that                                ists and has the potential to generate significant foreign cur-
           hippopotamuses were responsible for a significant reduction                           rency income. In this context the management plans of
           in rice production and caused significant economic losses                             Orango and Cacheu protected areas, developed in consult-
           (Lopes, ; Medina, ; Silva, ). They also have in-                          ation with local people, prioritize the assessment of mitiga-
           direct effects, such as restricting people’s movements and                            tion measures to resolve the conflict (Campos et al., ;
           causing accidents and deaths. In the th century  people                           IBAP, ).
           were injured and  killed by hippopotamuses in Orango                                    The behaviour of hippopotamuses in response to protec-
           and Cacheu as a result of boat sinking or while guarding                              tion measures has not been well studied (Eltringham, ).
           rice fields, and hippopotamuses are thus widely perceived                             To address human−hippopotamus conflict it is necessary to
           to be one of the major threats to the security and livelihoods                        consider both the patterns and effects of the damage as well
           of rural people, and inspire animosity and fear (Lopes, ;                         as the impacts of mitigating actions on the conservation sta-
           Campos et al., ; IBAP, ). Consequently the rice                               tus of the hippopotamus. With the aim of contributing to
           fields situated near water courses inhabited by hippopot-                             resolving the conflict between hippopotamuses and rice
           amuses have progressively been abandoned in favour of                                 farmers in Guinea-Bissau, we investigated access to rice
           rice fields located within forested areas, and this is contrib-                       fields, and evaluated the effectiveness of a protective meas-
           uting to increases in deforestation and biodiversity loss                             ure (electric fencing), hippopotamus behaviour in response
           (Vasconcelos et al., ; Medina, ).                                             to this measure, and the feasibility of its use by local farmers.
               To prevent hippopotamuses from entering rice fields,                              We propose practical and feasible measures to solve the con-
           farmers in Guinea-Bissau have been using barbed-wire                                  flict between people and hippopotamuses.
           fences, embankments and ditches, and night watchmen
           with fires and weapons. However, because of their high
           cost, complex maintenance or safety hazards these methods                             Study area
           have not been successful (Lopes, ; Campos et al., ;
           Silva & Monteiro, ; Medina, ). Electric fencing,                              The study was conducted in the Orango Islands National
           which is used effectively in other countries (Eltringham,                             Park (, ha; IBAP, ) and Uno Island in the
           ), has not been previously tested in Guinea-Bissau. Al-                           Bijagos archipelago, in Cacheu Natural Park (, ha;
           though farmers in protected areas have requested compen-                              IBAP, ), in Bissorao in the region of Oio, and in
           sation, or elimination of hippopotamuses, the wildlife                                Carantaba in the region of Gabu (Fig. ). Henceforth the
           authorities have not considered either option. Compensation                           term Orango includes Uno Island.

           Oryx, 2017, 51(2), 222–229 © 2016 Fauna & Flora International doi:10.1017/S003060531500109X
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. https://doi.org/10.1017/S003060531500109X
224         L. M. González et al.

                The climate is tropical, with a rainy season during                                fences,  unfenced flooded fields and  unfenced rain-fed
             May−October in the Bijagos and during June−October on                                 fields (Table ). We recorded the location and area of each
             the mainland. The dry season is December−April in the                                 rice field using a global positioning system to an accuracy of
             Bijagos and December−May on the mainland. The mean                                     m. We also measured the distance of each field to the near-
             annual rainfall is , mm in the Bijagos and ,–,                            est running water. All fields were unguarded by night watch-
             mm on the mainland, and the mean annual temperature is                                men during the study period. The number of fenced fields
             .°C; both parameters were stable during the study period,                          increased each year. The electric fence equipment was re-
             with no drought years (Medina, ; CCKP, ).                                     tired each year after the end of the harvest and installed
                The dominant vegetation in the study area consists of a                            again the following year in the same field, until the end of
             mosaic of tropical dry forest and tree and shrub dry savan-                           the study period. During the period of rice cultivation
             nah, with sub-humid isolated and flooded herbaceous                                   each year, the perimeter and interior of fenced fields
             savannah (Catarino et al., ). Two types of rice fields                            was monitored daily by the farmers. In addition, we
             were included in the study: flooded and rain-fed. The                                 surveyed the fenced and unfenced fields every − days dur-
             flooded fields, known locally as bolanha and bas-fond, are                            ing this period to search for hippopotamus footprints.
             flooded by river run-off and are located in the humid savan-                          Information on hippopotamus presence in rain-fed fields
             nah and alluvial depressions. The water is retained and                               was provided by farmers at the end of the growing season,
             regulated by a system of dykes with drainage channels.                                having been verified by protected area wardens.
             Management is communal, with each field divided into                                      We recorded hippopotamus detection without entry if
             owned plots (Biai, ; Medina, ). The long-cycle                                we observed footprints only on the perimeter, and with
             rice seed (-day seed) and occasionally the short-cycle                             entry if the footprints were in the interior of the field. The
             seed (-day seed) are used. Sowing begins after water                                location and date were recorded for each footprint, to avoid
             reaches the maximum retainable level, and the rice growth                             duplication. The hippopotamus populations of Orango and
             depends on the amount of water available in the soil. The                             Cacheu are isolated and, according to annual monitoring by
             duration of the crop in each area varies depending on                                 wardens, hippopotamuses occupied the same areas during
             when the rainy season begins (Medina, ). In Orango,                               the study period as in previous studies (Lopes, ; Silva,
             during the study period, sowing began in July–August and                              ; IBAP, ; Fig. ).
             harvesting in December–January; in Cacheu, sowing started                                 We estimated the rice productivity (kg ha−) in each field
             in June–July and harvesting in December; and in Bissorao                              by interviewing the farmers and recording the number of
             and Carantaba, sowing began in August–September and                                   bags of rice produced each season (each bag weighing
             harvesting in December–January.                                                       c.  kg) and the number of families working the fields. In
                The rain-fed fields, known locally as mpam-mpam, are                               Orango we interviewed , ,  and  farmers during
             located in non-flooded lands, in forested areas where the                             –, respectively; on the mainland we interviewed
             vegetation was previously slashed and burned. Each year a                              and  farmers in  and , respectively. To
             different area of land is prepared, and the same area is cul-                         avoid overestimation of crop losses, we recorded the area af-
             tivated again every – years. The short-cycle seeds are                             fected, and this information was checked by the wardens of
             used most often in these fields. The water comes exclusively                          the protected areas.
             from rainfall, and crop development conforms to the dur-                                  We identified three periods of rice cultivation (sowing,
             ation of the rainy season. Sowing normally begins in                                  vegetative shoot growth and harvesting) and five climatic
             April–May and harvesting in October–November (Biai,                                   periods (end of the dry season, February–April, ,  mm
             ; Medina, ).                                                                  rainfall per month; start of the rainy season, May–June,
                                                                                                   – mm per month; full rainy season, July–September,
                                                                                                   .  mm per month; end of the rainy season, October–
             Methods                                                                               November, – mm per month; beginning of the dry
                                                                                                   season, December–January, ,  mm per month).
             Monitoring of rice fields                                                                 The electric fence system included an aluminium wire of
                                                                                                   . mm diameter, c.  cm above the ground, connected to
             We conducted enquiries with farmers in Orango in                                  an energizer unit, and a rope between wooden stakes, with
             (n = ) and on the mainland in  (n = ) to locate                             strips of red and white striped plastic at intervals of  m
             rice fields and record their state of cultivation.                                    (Plate ). We used two models of portable energizers with
             Subsequently we identified farmers with flooded fields                                integrated solar panels: Speedrite-Viper S (Tru-Test
             who were willing to collaborate with the study and con-                               Group, Auckland, New Zealand), which powered c.  km
             sented to the installation of electric fences, and monitoring                         of wire in Orango, and S, which powered c.  km of
             of their fields. During – in Orango and –                             wire on the mainland. In each field two energizers were
             on the mainland we monitored  flooded fields with electric                          used simultaneously. The charge was activated and switched

                                                                                        Oryx, 2017, 51(2), 222–229 © 2016 Fauna & Flora International doi:10.1017/S003060531500109X
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. https://doi.org/10.1017/S003060531500109X
Preventing crop raiding by hippopotamus     225

                                TABLE 1 Numbers of fenced and unfenced rice fields (flooded and rain-fed) studied in the Orango Islands National Park and Uno Island during −, and on mainland Guinea-Bissau

                                                                                                                                                                                                                                                                            Rain-fed
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       91
                                                                                                                                                                                                                                                                                       91
                                                                                                                                                                                                                                                                            Flooded
                                                                                                                                                                                                                                                                    Total

                                                                                                                                                                                                                                                                                        75
                                                                                                                                                                                                                                                                                        25
                                                                                                                                                                                                                                                                                       100
                                                                                                                                                                                                                                                                            Rain-fed
                                                                                                                                                                                                                       No. of rice fields studied on the mainland

                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                            Flooded
                                                                                                                                                                                                                                                                    2013

                                                                                                                                                                                                                                                                                       14
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       14

                                                                                                                                                                                                                                                                                                  PLATE 1 The electric fencing system in a flooded rice field in
                                                                                                                                                                                                                                                                            Rain-fed

                                                                                                                                                                                                                                                                                                  Orango (Fig. ), with the energizer unit model S with
                                                                                                                                                                                                                                                                                                  integrated solar panel.
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                            Flooded

                                                                                                                                                                                                                                                                                                  on only at night. The cost of a complete fencing system with
                                                                                                                                                                                                                                                                                                  the S and S energizers was c. EUR  and , re-
                                                                                                                                                                                                                                                                    2012

                                                                                                                                                                                                                                                                                        5
                                                                                                                                                                                                                                                                                        9
                                                                                                                                                                                                                                                                                       14

                                                                                                                                                                                                                                                                                                  spectively. The vegetation was cut from within a distance
                                                                                                                                                                                                                                                                                                  of – m around the wires twice per week.
                                                                                                                                                                                                                                                                            Rain-fed
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0
                                                                                                                                                                                                                                                                                       0

                                                                                                                                                                                                                                                                                                  Variables and statistical analyses
                                                                                                                                                                                                                                                                            Flooded

                                                                                                                                                                                                                                                                                                  The rice field was considered to be the experimental unit. To
                                                                                                                                                                                                                                                                    2011

                                                                                                                                                                                                                                                                                                  identify differences in the raiding patterns of hippopot-
                                                                                                                                                                                                                                                                                       25
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       25

                                                                                                                                                                                                                                                                                                  amuses we considered two response variables: hippopot-
                                                                                                                                                                                                                                                                            Rain-fed

                                                                                                                                                                                                                                                                                                  amus detection/no detection, and the number of fields
                                                                                                                                                                                                                                                                                                  where hippopotamuses were detected. We used the follow-
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       30
                                                                                                                                                                                                                                                                                       30

                                                                                                                                                                                                                                                                                                  ing as explanatory variables: the type of field (flooded vs
                                                                                                                                                                                                                                                                                                  rain-fed); the geographical location of the field (island vs
                                                                                                                                                                                                                                                                            Flooded

                                                                                                                                                                                                                                                                                                  mainland); the distance to the nearest water body (in m);
                                                                                                                                                                                                                                                                    2010

                                                                                                                                                                                                                                                                                                  the presence/absence of electric fencing; the cultivation
                                                                                                                                                                                                                                                                                       21
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       21

                                                                                                                                                                                                                                                                                                  phase (sowing, growth or harvesting); and the climatic per-
                                                                                                                                                                                                                                                                                                  iod. To investigate the economic effects of fencing we used
                                                                                                                                                                                                                                                                            Rain-fed

                                                                                                                                                                                                                                                                                                  the following as response variables: the rice productivity of
                                                                                                                                                                                                                       No. of rice fields studied in Orango/Uno

                                                                                                                                                                                                                                                                                                  the field, the number of families working in the field, and the
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       37
                                                                                                                                                                                                                                                                                       37

                                                                                                                                                                                                                                                                                                  number of flooded or rain-fed fields. As independent vari-
                                                                                                                                                                                                                                                                            Flooded

                                                                                                                                                                                                                                                                                                  ables we used the presence/absence of electric fencing, and
                                                                                                                                                                                                                                                                    2009

                                                                                                                                                                                                                                                                                                  the year.
                                                                                                                                                                                                                                                                                        6
                                                                                                                                                                                                                                                                                        7
                                                                                                                                                                                                                                                                                       13

                                                                                                                                                                                                                                                                                                     As quantitative variables could not be fitted to a normal
                                                                                                                                                                                                                                                                                                  distribution we performed non-parametric analysis, using
                                                                                                                                                                                                                                                                            Rain-fed

                                                                                                                                                                                                                                                                                                  as covariates each of the independent variables, with a con-
                                                                                                                                                                                                                                                                                                  fidence interval of %. Based on the characteristics of the
                                                                                                                                                                                                                                                                                        0
                                                                                                                                                                                                                                                                                       24
                                                                                                                                                                                                                                                                                       24

                                                                                                                                                                                                                                                                                                  response and explanatory variables (categorical or continu-
                                during − (Fig. ).

                                                                                                                                                                                                                                                                                                  ous) we performed various tests: frequency analyses (χ)
                                                                                                                                                                                                                                                                            Flooded

                                                                                                                                                                                                                                                                                                  when response and explanatory variables were categorical,
                                                                                                                                                                                                                                                                    2008

                                                                                                                                                                                                                                                                                        4
                                                                                                                                                                                                                                                                                        9
                                                                                                                                                                                                                                                                                       13

                                                                                                                                                                                                                                                                                                  simple regression analyses for continuous variables, and
                                                                                                                                                                                                                                                                                                  Kruskal−Wallis (χ) for comparing mean values of the mul-
                                                                                                                                                                                                                                                                                       Unfenced

                                                                                                                                                                                                                                                                                                  tiple (. ) categories of independent categorical variables.
                                                                                                                                                                                                                                                                                       Fenced

                                                                                                                                                                                                                                                                                                  To compare the mean annual productivity of fields we
                                                                                                                                                                                                                                                                                       Total

                                                                                                                                                                                                                                                                                                  used the same fields before and after the installation of the

           Oryx, 2017, 51(2), 222–229 © 2016 Fauna & Flora International doi:10.1017/S003060531500109X
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226         L. M. González et al.

             electric fences (matched pair data), and applied a Wilcoxon
             test (Z). Only significant results and those that approached
             significance are presented. The analyses were carried out
             using Statistica v. . (StatSoft, ).

             Results

             Monitoring of rice fields

             In Orango we located  flooded rice fields, .% of them
             abandoned. The cause of abandonment was known in 
             cases: entry of hippopotamuses was the most frequent
             cause (%), followed by migration of the human popula-
             tion (%) and low rice production (%). In , the
             year before electric fences were installed, hippopotamuses                            FIG. 2 Mean number of flooded rice fields (with % confidence
             had entered four of the  flooded fields that were cultivated.                       intervals) in the study areas in Guinea-Bissau (Fig. ) where
             On the mainland we located  cultivated flooded fields, and                          hippopotamuses were detected at various periods before, during
             in the year before the electric fences were installed ()                          and after the rainy season.
             hippopotamuses had entered all of these fields.
                 Hippopotamuses were detected in .% of fields
             (n = ), and the rate of detection was significantly higher
             in flooded than in rain-fed fields (%, n =  vs .%,
             n = , χ = ., P = .). The mean distance of flooded
             fields to the nearest running water was  m (.–. m, n
              = ), and the number of flooded fields where hippopot-
             amuses were detected was significantly higher closer to
             running water (r = −., P = .).
                 The number of flooded fields in which hippopotamuses
             were detected decreased significantly over the study period
             (r = ., r = −., P = .), increased significantly
             with the number of times the field was cultivated (r = .,
             r = ., P , .), and was significantly higher in Orango
             than on the mainland (.%, n =  vs .%, n = ,
             χ = ., P = .) and in unfenced vs fenced fields
             (.%, n =  vs .%, n = , χ = ., P = .). The
             number of fields where entry by hippopotamuses was de-
             tected was significantly lower for fenced than unfenced
             fields (.%, n =  vs .%, n = , χ = ., P , .).                       FIG. 3 Mean number of flooded rice fields (with % confidence
                 In flooded fields we detected hippopotamuses                                      intervals) in the study areas in Guinea-Bissau (Fig. ) where
             during September−January (n = ), mostly during                                      hippopotamuses were detected during various stages of crop
                                                                                                   development.
             October−December (.%). In rain-fed fields we detected
             hippopotamuses in August and September (n = ). In all
             areas studied the number of fields where hippopotamuses
                                                                                                   number of cultivated flooded fields increased significantly
             were detected was significantly higher at the end of the
                                                                                                   during the study (. ha in , n = ; . ha in
             rainy season and at the start of the dry season than at
                                                                                                   , n = ; r = ., r = ., P = .) and consequently
             other times (χ = ., P , .; Fig. ), and during the
                                                                                                   rice production increased (from , kg in  to ,
             period of vegetative stem growth than during sowing and
                                                                                                   kg in ). The mean productivity increased significantly
             harvest (χ = ., P = .; Fig. ).
                                                                                                   after the installation of the electric fencing (. vs .
                                                                                                   kg ha−, n = ; Z = ., P = ., n = ).
             Area and productivity of rice fields                                                     The area of rain-fed fields in Orango decreased (. ha
                                                                                                   in , n = ; . ha in , n = ), although not sig-
             The area of flooded fields was .−. ha in Orango and                            nificantly (r = ., r = −., P = .). The number of
             .−. ha on the mainland. The total area and the                                  families working in fenced fields in Orango increased

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Preventing crop raiding by hippopotamus                      227

                                                                                                 installed the hippopotamus passed between the two wires,
                                                                                                 pushing one wire until the tension caused it to break, or
                                                                                                 breaking the wire by standing on it.

                                                                                                 Discussion

                                                                                                 Our results indicate that the frequent access of hippopot-
                                                                                                 amuses to flooded rice fields resulted in significant damage
                                                                                                 to crops, confirming the information provided by farmers in
                                                                                                 previous studies (Lopes, ; Campos et al., ) and as
                                                                                                 occurred in other regions (Clarke, ; Mkanda, ;
                                                                                                 Kendall, ). In our study area, as in other regions
                                                                                                 (Verweij et al., ; Chansa et al., ), the dry and
           PLATE 2 A hippopotamus Hippopotamus amphibius feeding in a                            humid savannah grasslands are the preferred feeding
           flooded rice field in Carantaba, observed by local people.                            grounds of the hippopotamus, and are the habitat that
           (Photograph by Tete Sambu, )                                                      holds the greatest diversity and abundance of grass species.
                                                                                                 In our study area up to  species have been identified in the
                                                                                                 diet of the hippopotamus (Duarte et al., ). Much of the
                                                                                                 diversity and abundance of local grass species is maintained
                                                                                                 in the flooded rice fields (Catarino et al., ), which could
                                                                                                 explain why hippopotamuses are attracted to these areas.
                                                                                                 Hippopotamuses frequently enter flooded fields when the
                                                                                                 stem rice is in the vegetative growth phase, at the end of
                                                                                                 the rainy season and the beginning of the dry season, and
                                                                                                 also when the dry savannah grass begins to dry out and
                                                                                                 humid savannah grasses and stem rice in flooded fields
                                                                                                 are still green and growing (Catarino et al., ). The dry
                                                                                                 savannah grasses are found in the plains and are rarely in-
           PLATE 3 Camera trap images of a hippopotamus crossing an                              undated, whereas the humid or wet savannah are flooded
           electric fence in a rice field in Acagume (Orango), in .                          during the wet season, with the dominant species being
                                                                                                 Anadelphia afzeliana (Duarte et al., ). Thus, the
                                                                                                 humid savannahs and the flooded rice fields established in
           significantly (from  in  to  in , r = ., r =
                                                                                                 them constitute a critical dry-season feeding ground for
            ., P = .).
                                                                                                 hippopotamuses, and could explain the high frequency of
                                                                                                 hippopotamuses entering these rice fields.
           Hippopotamus behaviour                                                                    The lower frequency of hippopotamuses entering
                                                                                                 rain-fed fields compared to flooded fields is probably a re-
           Hippopotamuses entered the fields at night, except in one                             sult of rain-fed fields generally being located far from water-
           case where an individual was observed feeding during the                              courses. In addition, in rain-fed fields vegetative stems grow
           day in an unfenced field (Plate ). Although hippopot-                                earlier during the rainy season, when the greatest abundance
           amuses were detected in  fenced fields, in nine of these                            of pasture is available in the dry savannahs. Thus, rice-
           the electric system was not operating because the surround-                           planting dates, which determine the duration of the
           ing vegetation was in contact with the wire or the wire was                           cultivation in each season, could be a factor in determining
           not connected to the energizer. After resetting the appropri-                         the extent of crop damage by hippopotamuses, and could be
           ate operating conditions we detected no more entries into                             adjusted to minimize the amount of damage caused. In
           these fields, although the hippopotamuses did make further                            flooded fields, as in rain-fed fields, the dates for sowing
           attempts. Only in one field did an individual enter while the                         seeds could be brought forward by using short-cycle seeds,
           electric fencing was operating properly, and this same indi-                          to match the period of vegetative stem growth to that of the
           vidual (as indicated by the marks on their skin, Plate ),                            natural grasses in the hippopotamuses’ feeding grounds.
           an adult male, entered the field .% of the days surveyed                               Electric fencing produced a negative reaction in hippo-
           (n = ). The individual entered for the first time before the                        potamuses because they tended to touch the wire with
           electric fence had been installed, and after installation en-                         their noses, a poorly insulated and highly innervated part
           tered by walking below the wire. After a second wire was                              of the body (Eltringham, ). However, we observed one

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228         L. M. González et al.

             individual that crossed the electric fence while it was oper-                         limited applicability in many countries (Lindsey et al.,
             ating. It touched the wire with its neck, back or chest                               ). In our study there were no malfunctions of the electric
             (Plate ), which are less sensitive areas than the nose, and                          equipment, and the few incidents detected were attributable
             then moved swiftly, crossing before the electrical pulse                              to human factors and were easily resolved. The success of
             was generated. We do not discard the possibility that this                            electric fencing as a deterrent was primarily a result of
             individual may have been tolerant to the level of shock in-                           using a new energizer unit, which is portable, well suited
             tensity. The decrease in detection of hippopotamuses in                               to the various enclosure sizes and moisture conditions in
             fenced fields over the course of the study period may also                            the field, and its operation is simple and easily understood.
             indicate that damage was caused by a few individuals, who                             Thus, based on our results we conclude that the electric fen-
             learned to avoid the fields after the electric fences were in-                        cing system, if used properly, is potentially an efficient and
             stalled. Hippopotamuses may also have detected the electric                           viable method of preventing the access of hippopotamuses
             fence without actually touching it; as has been suggested for                         and other megaherbivores into crop fields. However, electric
             domestic animals, they may have detected the odour, elec-                             fences should be designed to minimize the loss of wild ani-
             tromagnetic field or ozone generated around the wires                                 mals as a result of entanglement or electrocution (Lindsey
             (McKillop et al., ).                                                              et al., ). As the components of electrified fencing are
                 A potential ecological disadvantage of fenced fields is                           valuable and vulnerable to theft, their use should be re-
             that fencing may be regarded as an incentive to transform                             stricted to smaller and well-surveyed protected areas
             humid savannahs to flooded rice fields, which could reduce                            where there are severe human−wildlife conflicts.
             hippopotamuses’ access to dry-season grazing grounds. In
             this context, and considering that the grazing requirement,
             estimated in a subtropical ecosystem, is  ha of grassland per                        Acknowledgements
             individual (Chansa et al., ), the hippopotamus popula-
             tion of Orango, estimated to be  individuals (Silva, ),                        We thank the technicians of the Instituto da Biodiversidade
             requires  ha of grassland, which is less than the area of                          e das Áreas Protegidas of Guinea-Bissau (IBAP), in particu-
             dry and humid savannahs available, estimated to be , ha                           lar the Director Alfredo Simão, and Justino Biai, Cristina
             (Cuq et al., ). The hippopotamus population is probably                           Silva, Aissa Regalla, Antonio Da Silva, Domingozinho,
             below its carrying capacity in Orango. Although hippopot-                             Fernando Biagi, Januario Gomes, Domingos Betunde and
             amus populations are limited by food availability in the dry                          Cecilia Netambo. We are also grateful to the wardens of
             season (Lewison, ; Harrison et al., ), in Orango the                          Orango National Park and Tarrafes do Rio Cacheu
             savannahs are the most extensive habitat, occupying % of                            Natural Park for their invaluable help. We especially
             the surface area (Cuq et al., ), and have increased not-                          thank Jose Eliseo Benante, Teté Sambú and Guerra
             ably since the mid th century (Vasconcelos et al., ).                           Bidonga. We are grateful also to Pierre Campredon,
             This could mitigate the effects of fencing in reducing hippo-                         Nelson Dias, John Sivhik and Kurt Verkauteren for provid-
             potamuses’ access to flooded fields. However, we recom-                               ing technical assistance. Jean Paul Taris, Luc Hoffman,
             mend an assessment of the impact of fencing flooded                                   Ignacio Morillo, Mary Seck, Iris Cardiel, José Jiménez,
             fields on the ecological requirements of hippopotamuses, es-                          Francisco Cantos, Miguel Angel Cedenilla, Sandra
             pecially during the dry season.                                                       Agudín, Fernando Silvestre, Mercedes Muñoz and Cristina
                 In the two protected areas the wildlife authorities do-                           Martínez provided advice and logistical support throughout
             nated the electric fencing to the farmers, who paid a small                           this study. We thank Rubén Moreno-Opo and Jaime Muñoz
             amount annually for maintenance and installation. With                                for comments and advice. Financial support for this study
             the fencing we observed that night watchmen were no long-                             was provided by MAVA Foundation, Fundación
             er necessary to protect the crops. There was a notable in-                            Biodiversidad of Spain, O. A. Parques Nacionales,
             crease in the number of active flooded rice fields, rice                              MAGRAMA and AECID Spain.
             production and the number of families working in the fields.
             For these reasons, in this case mitigation is more cost effect-
             ive than compensation. In this context we recommend that                              References
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           Oryx, 2017, 51(2), 222–229 © 2016 Fauna & Flora International doi:10.1017/S003060531500109X
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