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Uncontrolled hunting and habitat degradation decimate and extirpate forest hornbills in Ghana, West Africa

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Abstract

Tropical forests are on the front lines of the current global extinction crisis. Species with restricted habitat requirements and slow reproductive rates, such as the spectacular hornbills (Bucerotidae) of the Paleotropics, are particularly vulnerable. We present the first long-term quantitative population assessment of nine forest hornbill species in Ghana, part of the Upper Guinea forest biodiversity hotspot in West Africa. From 1990 to 2009, hornbill encounter rates declined with 32–88% across eight species found in the region. Seven separate surveys between 1990 and 2014 indicated declines in at least six of eight species detected, with large-bodied species hardest hit. Depleted remnant populations of large hornbills mainly persist in two large and relatively well-protected wildlife reserves, Ankasa Resource Reserve and Kakum National Park. Contrastingly, the five largest species of the nine hornbills known to Bia Biosphere Reserve, one of Ghana's few forest wildlife reserves, apparently vanished completely since the 1990s, mainly due to uncontrolled hunting. Similarly, several large hornbills have disappeared from forest reserves where hunting is widespread. We conclude that uncontrolled hunting is the major driver of the recent drastic declines and population extirpations of large hornbills, while reductions in small insectivorous species may be related to extensive fragmentation and habitat disturbances of the Ghanaian forest biome. We call for urgent conservation action to prevent further declines and impending extirpations of forest hornbills and other wildlife in West Africa.

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We provide the first documented case of the extinction in the twentieth century of a widely recognized primate taxon. During surveys in Ghana and Côte d'Ivoire in 1993–1999, we were unable to find any surviving populations of Miss Waldron's red colobus monkey ( Procolobus badius waldroni), a primate taxon endemic to the forests of this part of West Africa. We conclude that this monkey, which at least one authority considers worthy of species status, is probably extinct. Hunting by humans appears to be the ultimate cause of the extinction. Until our surveys began, little attention had been paid to the plight of this red colobus monkey, despite its listing as endangered by the World Conservation Union. The extinction of other large animals in the Upper Guinea rainforest region is likely to follow soon unless more attention is paid to the full range of endangered forms and more resources are devoted to their rigorous protection.
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Conservation of tropical forests requires an understanding of the seasonal patterns of abundance and distribution of important seed dispersers. In the forests of Central Africa, three species of hornbills in the genus Ceratogymna (Aves: Bucerotidae) provide dispersal for over 50 species of trees and lianas, yet their ecology is poorly known. We present the first empirical evidence for resource tracking and large-scale movements by two of these species, which were previously considered resident. Hornbill numbers and fruit production were estimated over a one-year period in the Dja Reserve, Cameroon. Monthly abundances of C. atrata and C. cylindricus varied four- and 12-fold, respectively, and were predicted by community fruit availability as measured by fruitfall on a 4.3 km trail. In contrast, C. fistulator abundances varied little and were not related to fruit availability. Synchronous population fluctuations in sites 22 km distant were observed for atrata and cylindricus, but not for fistulator. Together, these results indicate that atrata and cylindricus populations track fruit resources via seasonal emigrations, while the fistulator population appears to be relatively sedentary. Comparisons of hornbill densities in primary and secondary forests suggest that they are important dispersers in both habitats, and will likely play larger roles in central African forest regeneration as more vulnerable seed dispersers are removed by hunting. If large seasonal movements of hornbills turn out to be a general phenomenon, effective management of central African forest reserves will require greater emphasis on reserve connectivity and on seasonal and spatial patterns of fruit production.
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Nine species were edge sensitive, 7 of which were also area sensitive. However, forest structure near patch edges was not consistent with bird responses to canopy and tree density, suggesting that mechanisms other than microclimate or structural changes ( perhaps predation or nest parasitism) underlie the response of most species that are sensitive to both edge and area. Regression of critical patch size (the smallest patch size in which a species was detected) on logarithm of body mass (an index of home range size) for 22 area-sensitive species suggests that area-sensitive species are unlikely to occur in patches smaller than several home-range areas. Canopy density influenced 13 species (11 positively, 2 negatively), and abundance of large trees influenced 8 species (3 positively, 5 negatively). Forest birds did not respond to isolation (distance from a patch to a large forest) for isolation distances of 1–25 km, suggesting that island biogeographic mechanisms had less influence on birds than other potential mechanisms of area sensitivity. Although small patches contributed little to the conservation of forest birds—species found in small patches were well represented in large patches—small patches are probably important for supporting generalist bird species that provide ecological services in the agricultural matrix and serving as nuclei for future ecosystem recovery.Resumen: A pesar de que los bosques tropicales de África Occidental son los más fragmentados en África, no existe información publicada sobre la biodiversidad de esos fragmentos. Para determinar como responden las aves de bosque a cinco factores (tamaño del fragmento, aislamiento del fragmento, densidad del dosel, abundancia de árboles grandes y proximidad al borde bosque) en estos bosques, tomamos muestras de 60 especies a lo largo de 121 transectos (2.5 Ha cada uno) en 35 fragmentos de bosque en la zona de bosque semidecíduo en Ghana. La riqueza de especies por transecto incrementó con el tamaño del fragmento en todo el rango de tamaños de fragmento (3 a 30,000 Ha). La diversidad de aves de bosque (22 especies) a lo largo de un solo transecto fue comparable a la diversidad acumulada (25 especies) en 17 transectos en 13 fragmentos pequeños. Veintidós de 60 especies fueron sensibles al tamaño del fragmento, de las cuales 15 nunca fueron encontradas en fragmentos pequeños. Estos resultados sugieren que muchas especies de aves de bosque en África Occidental se conservarán sólo en fragmentos grandes. Nueve especies fueron sensibles al borde del bosque, de las cuales 7 también fueron sensibles al tamaño del fragmento. Sin embargo, la estructura del bosque cerca del borde de los fragmentos no se correlacionó con las respuestas de las aves a los cinco factores, lo que sugiere que en la respuesta de muchas especies sensibles tanto al borde como al área subyacen mecanismos distintos al microclima o cambios estructurales ( probablemente depredación o parasitismo de nidos). La regresión del tamaño crítico del fragmento (el fragmento más pequeño en que se detectó la especie) en el logaritmo de la masa corporal (un índice del rango de hogar) en 22 especies sensibles al área sugiere que las especies sensibles al área probablemente no ocurran en fragmentos de menor tamaño que varias áreas de rango de hogar. La densidad del dosel influyó a 13 especies (11 positivamente, 2 negativamente), y la abundancia de árboles grandes influyó a 8 especies (3 positivamente, 5 negativamente). Las aves de bosque no respondieron al aislamiento (distancia del fragmento a un bosque extenso) en distancias de 1 a 25 km, lo que sugiere que los mecanismos de biogeografía de islas tuvieron menos influencia en aves que otros mecanismos potenciales de sensibilidad al área. Aunque los fragmentos pequeños contribuyeron poco a la conservación de las aves de bosque (especies encontradas en fragmentos pequeños estuvieron bien representadas en fragmentos grandes), los fragmentos pequeños probablemente son importantes para aves generalistas que proporcionan servicios ecológicos en la matriz agrícola y funcionan como núcleos para la futura recuperación de ecosistemas.
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We studied two species of Ceratogymna hornbills, the black-casqued hornbill, C. atrata, and the white-thighed hornbill, C. cylindricus, in the tropical forests of Cameroon, to understand their movement patterns and evaluate their effectiveness as seed dispersers. To estimate hornbill contribution to a particular tree species' seed shadow we combined data from movements, determined by radio-tracking, with data from seed passage trials. For 13 individuals tracked over 12 months, home range varied between 925 and 4,472 ha, a much larger area than reported for other African avian frugivores. Seed passage times ranged from 51 to 765 min, with C. atrata showing longer passage times than C. cylindricus, and larger seeds having longer gut retention times than smaller seeds. Combining these data, we estimated that seed shadows were extensive for the eight tree species examined, with approximately 80% of seeds moved more than 500 m from the parent plant. Maximum estimated dispersal distances for larger seeds were 6,919 and 3,558 m for C. atrata and C. cylindricus, respectively. The extent of hornbill seed shadows suggests that their influence in determining forest structure will likely increase as other larger mammalian dispersers are exterminated.
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Global biodiversity is threatened by unsustainable exploitation for subsistence and commerce, and tropical forests are facing a hunting crisis. In Central African forests, hunting pressure has been quantified by monitoring changes in the abundance of affected species and by studying wild meat consumption, trade and hunter behaviour. However, a proportion of offtake is also discarded as bycatch or consumed by hunters when working, which can be overlooked by these methods. For example, remains of hornbills and raptors are found regularly in hunting camps but relatively few birds are consumed in households or traded in markets. Hornbill and raptor populations are sensitive to small increases in mortality because of their low intrinsic population growth rates, however, the scale and socioeconomic drivers of the cryptic hunting pressure affecting these species have not been quantified. We used direct and indirect questioning and mixed-effects models to quantify the socioeconomic predictors, scale and seasonality of illegal bird hunting and consumption in Littoral Region, Cameroon. We predicted that younger, unemployed men with low educational attainment (i.e. hunters) would consume birds more often than other demographics, and that relative offtake would be higher than expected based on results from village and market-based studies. We found that birds were primarily hunted and consumed by unemployed men during the dry season but, in contrast to expectations, we found that hunting prevalence increased with educational attainment. Within unemployed men educated to primary level (240 of 675 respondents in 19 villages), we estimated an average of 29 hornbills and eight raptors (compared with 19 pangolins) were consumed per month during the study period (Feb-Jun 2015) in a catchment of c.1135 km 2. We conclude that large forest birds face greater hunting pressure than previously recognised, and birds are a regular source of protein for men during unemployment. Offtake levels may be unsustainable for some raptors and hornbills based on life history traits but in the absence of sufficient baseline ecological and population data we recommend that a social-ecological modeling approach is used in future to quantify hunting sustainability.
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Hunting is a major driver of biodiversity loss, but a systematic large-scale estimate of huntinginduced defaunation is lacking.We synthesized 176 studies to quantify hunting-induced declines of mammal and bird populations across the tropics. Bird and mammal abundances declined by 58% (25 to 76%) and by 83% (72 to 90%) in hunted compared with unhunted areas. Bird and mammal populations were depleted within 7 and 40 kilometers from hunters’ access points (roads and settlements). Additionally, hunting pressure was higher in areas with better accessibility to major towns where wild meat could be traded. Mammal population densities were lower outside protected areas, particularly because of commercial hunting. Strategies to sustainablymanage wildmeat hunting in both protected and unprotected tropical ecosystems are urgently needed to avoid further defaunation.
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Elephant populations are in peril everywhere, but forest elephants in Central Africa have sustained alarming losses in the last decade [1]. Large, remote protected areas are thought to best safeguard forest elephants by supporting large populations buffered from habitat fragmentation, edge effects and human pressures. One such area, the Minkébé National Park (MNP), Gabon, was created chiefly for its reputation of harboring a large elephant population. MNP held the highest densities of elephants in Central Africa at the turn of the century, and was considered a critical sanctuary for forest elephants because of its relatively large size and isolation. We assessed population change in the park and its surroundings between 2004 and 2014. Using two independent modeling approaches, we estimated a 78–81% decline in elephant numbers over ten years — a loss of more than 25,000 elephants. While poaching occurs from within Gabon, cross-border poaching largely drove the precipitous drop in elephant numbers. With nearly 50% of forest elephants in Central Africa thought to reside in Gabon [1], their loss from the park is a considerable setback for the preservation of the species.
Thesis
Agriculture is arguably the greatest threat to tropical forest species. Conservation scientists disagree over the relative importance of two opposing strategies for minimising this threat: enhancing on-farm biodiversity, through wildlife-friendly farming practices, or sparing land for nature by using high-yielding farming methods on the smallest possible area to reduce the need to convert natural habitats. Previous theoretical work shows that understanding the relationship between population density and yield for individual species is crucial for determining whether one of these strategies, or a mixed strategy, will maximise their populations for a given food production target. In this thesis, I aim to identify what land-use strategy will permit increases in food production with least impact on species in the forest zone of Ghana. Farm-fallow mosaic landscapes with shifting cultivation and native canopy trees produced only around 15% as much food energy per hectare as the highest-yielding oil palm plantations. In farm mosaics where perennial tree crops dominate, food production and profits were higher, but did not reach those of oil palm plantations. I surveyed birds and trees in forest, farm mosaic, and oil palm plantation, and combined these data with information on yields to assess the likely consequences of plausible future scenarios of land-use change. My results provide evidence of a strong trade-off between wildlife value and agricultural yield. Species richness was high in low-yielding farming systems, but there was considerable turnover between these systems and forests, with widespread generalists replacing narrowly endemic forest-dependent species. Species most dependent on forest as a natural habitat, those with smaller global ranges and those of conservation concern showed least tolerance of habitat modification. For virtually all species, including even widespread generalists, future land-use strategies based on land sparing are likely to support higher populations of most species and minimise their risk of extinction compared to land-use strategies based on wildlife-friendly farming. If food production is to increase in line with Ghana‘s population growth, a combination of efforts to improve forest protection and to increase yields on current farmed land is likely to achieve this at least cost to forest species. Efforts to better protect forests, which require further restrictions on human use, might be most effective if they can be closely linked to support for farmers to improve their yields. In the long term however, this strategy will only delay and not avert biodiversity loss, unless global society can limit its consumption.
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Despite all the attention the problem of illegal logging has recently attracted, the empirical base remains remarkably weak in many countries including Ghana. Using both official reported figures and actual harvest figures estimated from domestic and export consumption statistics, the annual average illegal logging for the period 2000–2011 among the formal sector in Ghana is estimated at 0.78 million m³ or 104% of the legal harvest. The results indicate that illegal logging was much more prevalent in the forest reserves than outside forest reserves, and among the traditionally endangered timber species more than the lesser known timber species. The implications of these findings on forest law compliance, enforcement, and the Voluntary Partnership Agreement (VPA) between the European Community and Ghana are discussed and recommendations proffered. Spanish A pesar de todo el interés que ha atraído recientemente el problema de la tala ilegal, la base empírica sigue siendo notablemente escasa en muchos países, como por ejemplo Ghana. Mediante la utilización de cifras publicadas oficiales y cifras de aprovechamientos efectuados estimadas a partir de las estadísticas de consumo nacionales y de exportación, se estimó que la tala ilegal para el período 2000–2011 dentro del sector formal en Ghana fue de un promedio anual de 0,78 millones de m³, equivalente al 104% de los aprovechamientos legales. Los resultados indican que la tala ilegal es mucho más prevalente en las reservas forestales que fuera de las mismas, así como entre las especies de madera tradicionalmente en peligro en comparación con las especies maderables menos conocidas. Se discuten las implicaciones de estos resultados en el cumplimiento de la legislación forestal, su aplicación y el Acuerdo Voluntario de Asociación (AVA) entre la Unión Europea y Ghana y se ofrecen recomendaciones. French Malgré toute l'attention que le problème de la coupe de bois illégale a récemment attiré, la base empirique demeure singulièrement faible dans nombre de pays, Ghana inclus. En utilisant à la fois les chiffres rapportés officiellement et les chiffres de récolte réelle estimés par les statistiques de consommation domestique et à l'exportation, la coupe de bois illégale annuelle moyenne pour la période 2000–2011 dans le secteur formel du Ghana est estimée à 0.78 millions de m³, soit 104% de la récolte légale. Ces résultats indiquent que la coupe de bois illégale était bien plus dominante dans les réserves forestières qu'en dehors de ces dernières, et davantage auprès des espèces traditionnellement en danger que parmi celles moins connues. Les implications de ces découvertes sur l'observation de la loi et sa mise en application et sur l'Accord de partenariat volontaire (VPA) entre la Communauté Européenne et le Ghana sont examinées, et de recommandations sont offertes.
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The introduction of nationally-managed "western"-style protected area systems in Africa often led to antagonism between government authorities and local communities, because rural communities, which were largely excluded and displaced, often resorted to unsustainable activities like encroachment for settlement, illegal hunting and farming. It is the view of most biodiversity conservationists, however, that the management of natural resources must take into consideration both the introduced "western" model and traditional or local knowledge. The study aimed at assessing the Boabeng-Fiema Monkey Sanctuary in Ghana, as an example of blending traditional and introduced wildlife conservation systems, using interviews and questionnaire to obtain information from a cross-section of inhabitants of the area. The results indicated that the dual management system in operation at present has been largely successful, as the inhabitants appear to support the mechanisms put in place to manage the Sanctuary. Policy recommendations proposed to further enhance this success are in the areas of research and monitoring, education and awareness, tourism improvement, and introduction of community incentive schemes.
Article
Despite several decades of research on the effects of fragmentation and habitat change on biodiversity, there remain strong biases in the geographical regions and taxonomic species studied. The knowledge gaps resulting from these biases are of particular concern if the forests most threatened with modification are also those for which the effects of such change are most poorly understood. To quantify the nature and magnitude of such biases, we conducted a systematic review of the published literature on forest fragmentation in the tropics for the period 1980-2012. Studies included focused on any type of response of single species, communities, or assemblages of any taxonomic group to tropical forest fragmentation and on fragmentation-related changes to forests. Of the 853 studies we found in the SCOPUS database, 64% were conducted in the Neotropics, 13% in Asia, 10% in the Afrotropics, and 5% in Australasia. Thus, although the Afrotropics is subject to the highest rates of deforestation globally, it was the most disproportionately poorly studied biome. Significant taxonomic biases were identified. Of the taxonomic groups considered, herpetofauna was the least studied in the tropics, particularly in Africa. Research examining patterns of species distribution was by far the most common type (72%), and work focused on ecological processes (28%) was rare in all biomes, but particularly in the Afrotropics and for fauna. We suggest research efforts be directed toward less-studied biogeographic regions, particularly where the threat of forest fragmentation continues to be high. Increased research investment in the Afrotropics will be important to build knowledge of threats and inform responses in a region where almost no efforts to restore its fragmented landscapes have yet begun and forest protection is arguably most tenuous. Sesgos Biogeográficos y Taxonómicos en la Investigación de la Fragmentación de Bosques Tropicales.
Article
Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make pre- dictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. Tis was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non-hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera- ctions, triggered by the loss of seed dispersers. Te results predict a rarity of primate-dispersed trees in future tropical forest canopies; a forest less diverse in timber and non-timber resources.
Article
The urgent global objective of developing sustainable tropical forestry management practices, which also target biodiversity conservation, requires rapid comparative studies that address both biogeography and logging practices. This paper examines avifaunal implications (focusing on species diversity and composition) of forest fragmentation and selective logging, by comparing 15 selected reserved forests in south-west Ghana. The regeneration time since last logging varied from 0 (logging continuing) to 21 years, with one unlogged site as control. Extraction levels ranged between 0.20 and 3.75 trees per ha (mean 0.90). Bird sampling involved understorey mist-netting and standardized line-transect walks, which respectively accumulated 8,348 captures and 22,452 single records of 147 species. Mist-net data showed that understorey bird diversity was positively correlated with logging intensity up to c. 3 trees per ha, reflecting increased influx of open-land species and a persistence of forest obligates during the first 5 years after logging. The overall abundance of forest interior species tended to decrease 5–10 years after logging, but recovered fairly well thereafter. Canopy birds were generally more logging-resilient. Rare forest obligates with high conservation importance were found to be equally abundant in virgin and logged, large forests, whereas these birds were poorly represented in small heavily logged forests. The results are compared with findings from other regions, and conservation implications and constraints are discussed in a global perspective. Finally, recommendations on size-related sustainable extraction levels and regeneration time are presented for the Upper Guinea Forest.
Article
Long-distance seed dispersal influences many critical ecological processes by improving chances of gene flow and maintaining genetic diversity among plant populations. Accordingly, large-scale movements by frugivores may have important conservation implications as they provide an opportunity for long-distance seed dispersal. We studied movement patterns, resource tracking, and potential long-distance seed dispersal by two species of Ceratogymna hornbills, the black-casqued hornbill C. atrata, and the white-thighed hornbill C. cylindricus, in lowland tropical forests of Cameroon. We determined fruiting phenology of 24 tree species important in hornbill diet at monthly intervals and compared these patterns to monthly hornbill census data. After capture and radio-tagging of 16 hornbills, we used radio telemetry by vehicle and fixed wing aircraft to determine the extent of long-distance movements. Hornbills exhibited up to 20-fold changes in numbers in response to fruit availability in our 25 km(2) study area. Also, hornbills made large-scale movements up to 290 km, which are larger than any movement previously reported for large avian frugivores. Together, these observations provide direct evidence that hornbills are not resident and that hornbills track available fruit resources. Our results suggest that Ceratogymna hornbills embark on long-distance movements, potentially dispersing seeds and contributing to rain forest regeneration and diversity.
Article
The biodiversity of West African rainforests is globally significant but poorly described, little understood in terms of its functional significance, and under threat from forest loss and degradation. Estimates suggest that about 10 million ha of forest may have been lost in the 20th Century, and around 80% of the original forest area is now an agriculture-forest mosaic. These highly modified forests provide food, fuel, fibre and a range of ecosystem services for over 200 million people. As a consequence, the future of biodiversity in the region is intimately linked with the lives and livelihoods of local people. The available evidence suggests that forest loss and degradation has been caused primarily by agricultural expansion, sometimes facilitated by other human activities such as wood extraction. This expansion is a response to the demand generated by a growing and increasingly urbanised human population, but has been exacerbated by small increases in crop yields over recent decades. We synthesize and review our state of knowledge on the value of human-modified habitats for forest biodiversity in the region. Data on biodiversity are patchy, but we show that across plant, invertebrate and vertebrate groups, there is a significant loss of forest species as tree cover is reduced and vegetation structure simplified. Agricultural expansion clearly causes significant local biodiversity loss. We argue that replicated landscape-scale studies are now needed that describe changes across a range of biodiversity groups (above and below ground) in relation to land-use and landscape context to address knowledge gaps and biases. Such descriptive studies need to be complemented by a deeper understanding of the causes of species turnover patterns, together with work on the consequences of biodiversity loss for ecosystem function and services. Biodiversity conservation in the region is becoming increasingly embedded within a more multi-functional view of agriculture-forest mosaics that attempts to recognise and value the range of services provided by tree cover and other land-uses. This relatively new perspective has the potential, at least in principle, to re-shape the drivers of land-use change because tree cover can generate additional revenue through carbon trading or certification schemes. It will become clearer over the coming decades whether this potential can be realised, but the implications for biodiversity conservation in the region are potentially profound. The socio-economic processes that have driven forest loss and degradation in West Africa are having an increasing impact on the relatively undisturbed rainforest of the Congo Basin. We conclude by pointing out that while West African forest landscapes are a potent reminder of what might happen in Central Africa in the near future, they also provide insights for developing policies and practice that might avoid comparable levels of forest loss and degradation.
Article
The reproductive biology of two species of African hornbill, the Black-casqued Hornbill, Ceratogymna atrata, and the Whitethighed Hornbill, Ceratogymna cylindricus, was investigated over a four-year period (1994–1997) on a 25km 2 site in lowland rainforest in south-central Cameroon. Nesting attempts varied considerably among years, with the percentage of successful nests highest in 1995, with 64% and 54% of Black-casqued and White-thighed Hornbill fledging offspring, respectively. There were no nesting attempts in 1994, despite the fact that hornbills were present in the study area. Large differences in fruit availability were also noted across years, suggesting that reproductive activity and success are related to fruit availability. Data collected from 38 nests, over four breeding seasons (1994–1997), showed a preference for nest cavities in larger trees within areas of the forest containing larger trees. Hornbills did not show preferences for particular tree species, with the possible exception of Petersianthus macrocarpus, in which nine of the active nest cavities were found. Comparisons showed few significant differences in cavity characteristics between the two species . While cavities may have been a limiting factor in nesting in 1995, the year with the highest fruit availability, cavities were not limiting during other years when fruit availability was lower. Hornbill diets, as determined from seed traps at cavities, showed significant year-to-year variation. Although courtship and exploratory behaviour of cavities by pairs took place in most years, females did not wall themselves into cavities unless fruit was plentiful. Hornbills appear to time reproduction to coincide with peak food supply and successfully reproduce only when food is plentiful, and may curtail or forego nesting in years when fruit availability is low.
Article
Distance sampling, line transects and radio-tracking have been used previously to estimate population abundances and movements of hornbills. We used a large dataset of audio recordings made over 10 years to estimate seasonal variation in abundance of three species: Black-casqued Hornbill Ceratogymna atrata, Yellow-casqued Hornbill C. elata and Brown-cheeked Hornbill Bycanistes cylindricus, in Tai National Park, Cote d'Ivoire. The latter two species are Near Threatened. Recordings of hornbill wing-beats were used to control for potential seasonal variation in call production. Yellow- and Black-casqued Hornbill calls were 10 times as abundant from July to February as they were from March to June. Brown-cheeked Hornbills showed a similar trend, but less pronounced. From these data we concluded that large numbers of Yellow- and Black-casqued Hornbills had left the Tai National Park during this period. The significance of these findings is discussed in relation to the fragmentation and regeneration of the Upper Guinea forests and the relatively small size of the remaining forest habitat in the region.
Article
Almost nothing is known of the effects of forest fragmentation on bird diversity within the heavily degraded and fragmented forest remnants in West Africa. We examined the effects of edge, fragment size and isolation on bird species richness in southwestern Nigeria where forest fragmentation is pronounced. In total, 122 km of line transects were used to survey birds and vegetation within 45 forest patches between January 2000 and March 2002: 197 species were recorded. Avian species number and total counts in forest patches were unrelated to fragment area (within the observed range of 14–445 ha), but were negatively influenced by degree of isolation and increasing distance from the edge. As the total area of forested land within 15 km of a patch fell from 4 to 0%, so 21% of species were lost. In total, six and zero species (of 154 recorded more than once) were consistently recorded in the larger and smaller forest fragments, respectively, and four and two bird species were consistently recorded in unisolated and isolated forest fragments, respectively, suggesting that the addition of ‘edge’ species did not compensate for loss of species sensitive to fragmentation. Diversity index was not affected by either fragment area or degree of isolation, but decreased with distance from the edge. When individual species counts were considered, 68% of species (n = 62) showed no significant effect of distance to edge. Of those 20 species which showed an effect, 12 were less common close to the edge. Most species (65%) did not respond significantly to increasing isolation but of those 22 species that did, 20 were less common in more isolated fragments. Ninety-seven per cent of species showed no significant response to area. As avian diversity and species composition, but not species number, were apparently insensitive to forest fragmentation, our findings suggest that fragmentation reduces the probability of occurrence of a wide range of West African bird species, rather than a subset of fragmentation-sensitive species. The greater apparent sensitivity of present-day West African forest bird communities to fragmentation rather than patch size might reflect previous extinctions of area-sensitive species. Minimizing further forest fragmentation might be the most effective means of conserving avian diversity in current West African landscapes where most remaining forest patches are small (i.e. < 500 ha).
Article
Ghanaians have a long history in ivory, both for export and for carving. From the 1970s to the early 1990s, however, most of Ghana's elephant were killed either by local farmers in retribution for human-elephant conflict or by poachers for the ivory trade. Ghanaian ivory craftsmen used the tusks primarily to make jewellery and figurines over this time. These curios were mostly sold in Accra, the capital, but due to lack of market surveys, very little data are available. In July 2010 I surveyed the retail outlets selling ivory in Accra and counted only 10 items on display in an art gallery and 85 items brought to me in five of 186 souvenir shops and stalls I visited. The reason there were so few items was that the Ghana Police Service and Wildlife Division of the Forestry Commission had carried out a raid in November 2008 in the main curio market confiscating several hundred kilos of ivory items, fining and imprisoning the dealers. Since then the vendors in Accra have feared to sell ivory. Elephant poaching declined at the start of the 21st century thanks to improved law enforcement. In 2004 a new system was introduced that involves performance and adaptive management through the monitoring of patrol effort and observations by the field-staff in the Wildlife Division. The combined effect of performance and adaptive management was that the number of effective days spent in the field by an average Wildlife Guard doubled, which dramatically lowered the number of elephants killed illegally. In addition, governance improved, Ghanaians developed greater respect for the law and there was less corruption, which reduced elephant poaching and the sale of ivory objects. This paper concludes that Ghanaian examples of greater patrol staff performance through improved monitoring—and of successfully raiding outlets selling illegal ivory—are a sound approach to reducing elephant poaching. While improving anti-poaching exercises is more difficult in some African range States, the Ghanaian example of shop raids is easier to implement and has also worked in countries such as Cameroon and Ethiopia. Other countries in Africa, especially Cote d'Ivoire, Democratic Republic of Congo, Nigeria and Senegal, need to follow the example of Ghana in carrying out official raids on ivory in retail outlets.
Article
The exceptionally rapid degradation and fragmentation of the Upper Guinea Forest, caused by slash-and-burn farming and selective logging, oblige bird conservationists to examine critically the conservation value of unprotected areas, which may provide buffer zones or connecting corridors to protected forests. This paper assesses the utilisation of various tree-crop plantations by mainly lower storey forest birds in south-west Ghana, through avifaunal comparisons of shaded cash crops (rustic cocoa; neglected coconut) and exotic tree plantations (Cedrela, Gmelina) with nearby closed forest. Plantations adjacent to, or within, large (.200 km 2), biodiverse forests are superior bird habitats compared to similar plantations within small (,50 km 2) and species-poor forests. A relatively high forest tree density in cash crop (15–20 ha À1) and exotic (15–35 ha À1) tree plantations, combined with a luxuriant woody undergrowth (not slashed for .5–10 years), may additionally explain the presence of many forest specialists, including regionally 'Vulnerable' and locally 'Endangered' species. Overall, 50% of species of conservation importance found in forests were represented in plantations. These findings highlight the importance of shaded plantations with long periods between understorey weeding, as appropriate land-use systems that enhance the area under effective conservation and improve the connectivity of protected forest fragments. Results are compared to similar studies in the Old and New World tropics, and implications for off-reserve land-use management are compared and discussed in regional and global contexts.
Article
We estimated densities of parrot and hornbill species in primary and selectively logged forest and forest gardens at two lowland sites on New Britain, PNG. We related differences in abundance to food and nest-site availability in the different habitats and determined whether nest-site availability might limit local breeding populations. Blue-eyed Cockatoo Cacatua ophthalmica and Blyth's Hornbill Rhyticeros plicatus were usually rarer in forest gardens than in primary forest, but both fared well in logged forest. Eclectus Parrot Eclectus roratus was more common in all human-altered forests than in primary forest, and Eastern Black-capped Lory Lorius hypoinochrous was reasonably common throughout but extremely abundant in forest gardens at one site. Parrots and hornbills were recorded eating fruits of 15 tree species and flowers of nine species. Densities of these fruiting and flowering trees were highest in logged forest and forest gardens, respectively, indicating the importance of these anthropogenic habitats as feeding grounds for the assemblage. Active nest cavities were found in large individuals of 12 tree species. Densities of potential nest cavities were highest in primary forest and lowest in forest gardens. At both sites, estimates of potential nest-site density were significantly lower than estimates of the density of pairs of all species of parrots and hornbills: there may be 10–20 parrot/hornbill individuals per nest-hole. Continuing forest alteration, whilst further reducing nest-site availability, may allow large populations of parrots and hornbills to persist due to increased availability of food in some anthropogenic habitats. However, current abundance of such bird species may be a poor correlate of future extinction risk as long-lived taxa may remain common for some period even when annual recruitment has declined to critically low levels.
Article
We investigated the effects of forest fragmentation on the disappearance of fruit-eating animals and the recruitment of animal-, wind-, and gravity-dispersed trees in 80-year-old forest patches in the East Us- ambara Mountains of Tanzania. We compared adult and juvenile trees in forest transects in a 3500-ha submontane forest with those in four forest fragments of 521,30,9, and 0.5 ha. Preliminary results show that recruitment of seedlings and juveniles of 31 animal-dispersed tree species was more than three times greater in continuous forest and large forest fragments (greater than or equal to 30 ha) than in small forest fragments (less than or equal to 59 ha), whereas recruitment of eight wind- and gravity-dispersed trees of the forest interior was unaffected. Recruitment of 10 endemic, animal-dispersed tree species was 40 times lower in small fragments than in continuous forest or large fragments. Counts of diurnal primates and birds in all five sites indicated that frugivorous species have declined with decreasing fragment size. These results are consistent with the idea that loss of dispersal agents depresses tree recruitment in the course of forest fragmentation.
Article
This article explores spatial and temporal changes in diurnal primate abundance and behavior in response to hunting, logging, and conservation at the Dzanga Sangha Dense Forest Reserve (RDS), Central African Republic over time. We use a combination of line-transect surveys in 2002 and 2009 (N = 540 km) and ethnographic interviews (N = 210) to investigate changes in the status of cercopithecines and colobines at RDS, with additional comparisons to earlier work. This protected area was lightly logged in the 1970s and the park was gazetted in 1990, with multiple-use reserve sectors allocated. Since the park's inception, hunting and the trade of primates have increased, along with human migration, greater accessibility of arms, and reduction of preferred ungulate prey. Primates have declined in both the park and reserve sectors. Our data further suggest that at RDS hunting has had a greater impact on primate diversity and abundance than logging. We have identified changes in species-specific vulnerability to hunting over time, with Cercopithecus nictitans and Lophocebus albigena initially having appeared to be relatively resistant to hunting pressure in 2002. However, subsequently as gun hunting has increased at RDS, these species have become vulnerable. Although monkeys at RDS have been responding behaviorally to increased gun hunting, they are not able to keep pace with changing hunting practices. This study allows us to begin to understand synergistic impacts of hunting and logging, necessary if we are to recommend strategies to better secure the future of primates in multiuse protected areas.