COMMUNITY AND ECOSYSTEM ECOLOGY

Effect of Deforestation on a Southwest Ghana Dung Beetle Assemblage (Coleoptera: Scarabaeidae) at the Periphery of Ankasa Conservation Area
ADRIAN L. V. DAVIS1
AND

T. KEITH PHILIPS2

Environ. Entomol. 34(5): 10811088 (2005)

ABSTRACT During June 2003, an exploratory dung beetle survey (Coleoptera: Scarabaeidae: Scarabaeinae) was conducted in southwest Ghana at the southern edge of Ankasa Conservation Area, which is a 500-km2 conserved region of the threatened Eastern Upper Guinean rainforest. The survey provided a quantitative assessment of peripheral deforestation effects and an inventory for species on omnivore dung. Pitfall traps baited with pig dung were placed in both selectively logged and unlogged rainforest as well as in nearby oil palm and cacao plantations. The Ankasa forest inventory was mostly comprised of species previously trapped by human dung in the Western Upper Guinean rainforest of Ta National Park, Cote dIvoire. However, there was relatively low species abundance similarity between the June/July assemblages on omnivore dung in Ta and Ankasa rainforests, and even lower similarity between assemblages of Ankasa and the adjacent plantations. Of 29 species recorded in Ankasa forest, most were absent or rare in plantations where assemblages were numerically dominated by species previously recorded from the West African savannas. This replacement plantation fauna showed signicantly lower species richness and diversity, but signicantly higher abundance, compared with Ankasa forest assemblages. The results reiterate that forest reserves are essential for the conservation of specialist rainforest dung beetle species. KEY WORDS deforestation, dung, Ghana, plantation, rainforest

THE HIGHLY DIVERSE, EASTERN Upper Guinean rainforest extends from southeast Cote dIvoire across southern Ghana to the Togo Hills (Lebbie 2001). There is a rainfall gradient across this region from a maximum of 1,500 mm/yr in the Togo Hills to a maximum of 2,100 2,500 mm/yr in southwest Ghana and southeast Cote dIvoire. This latter region of wet evergreen forest constitutes one of ve centers of rainforest plant diversity occurring across tropical Africa (Hamilton 1989). Like the rest of the West African rainforest, much of this center of biodiversity has been exploited by agricultural interests (Gyasi et al. 1995, Osei 1996) or by the forestry (Hill and Curran 2003) and bush meat industries (Bowen-Jones and Pendry 1999, Barnes 2002, Fa et al. 2005). Although 500 km2 is conserved in the Ankasa Conservation Area, comprised of the Ankasa Resources Reserve and the adjoining Nini Suhien National Park, the Ghana gold mining industry (Adimado and Baah 2002) is exerting renewed pressure to exploit some existing Ghana rainforest reserves for gold deposits, which are known to exist in the underlying strata (Anane 2003). This includes the
1 Corresponding author: Department of Zoology and Entomology, University of Pretoria, 1 Lynnwood Rd., Pretoria 0002, South Africa (e-mail: adavis@zoology.up.ac.za). 2 Department of Biology, Western Kentucky University, Bowling Green, KY 42101-3576. E-mail: keith.philips@wku.edu.

Cape Three Points Reserve in the wet evergreen forest belt close to Ankasa. In 2003, an exploratory study was conducted to assess some effects of previous exploitation in this wet evergreen forest zone of southwest Ghana. Dung beetles were used as indicators (Halffter and Favila 1993, McGeoch et al. 2002) because of their specialist associations with both vegetation and dung type (Cambefort 1982, 1991, Cambefort and Walter 1991, Davis 1994, 1996). As part of the ground fauna, dung beetle species assemblages are associated with the inuence of plant physiognomy on microclimatic factors such as insolation, temperature, and light intensity (Davis 1996, Davis et al. 2002), rather than directly to plant species composition. Nevertheless, they are strongly inuenced by forest modication (Nummelin and Hanski 1989), deforestation (Howden and Nealis 1975, Klein 1989, Halffter et al. 1992, Estrada et al. 1998, Estrada and Coates-Estrada 2002), and replacement of forest trees by vegetation types that offer shade but are microclimatically dissimilar, such as in early succession after deforestation (Davis et al. 2002) or in plantations (Davis et al. 2000). Similarly, disappearance of particular mammals and their dung types, through hunting, strongly inuences the geographical occurrence of dung beetles specialized to those dung types (Davis 1997). Although many dung beetle species found in forests are essentially endemic to forest
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ecotypes, there is a surprising paucity of formal quantitative comparison between assemblages of forest and adjacent partially or entirely deforested localities in Africa. Therefore, the survey was designed to document some effects of deforestation through plantation development, to inventory dung beetles occurring on omnivore dung in the wet evergreen forest, and to document some specic biodiversity losses that might be expected if agricultural activities and gold mining are permitted in forest reserves. Materials and Methods This study was conducted on alluvial sandy clay loam soils at an altitude of 6.8 109.5 m, primarily within the wet, evergreen, southern forest of the Ankasa Resources Reserve, southwest Ghana, and in peripheral oil palm plantations. Whereas some parts of the reserve have never been logged, other parts were selectively logged 30 40 yr ago (S. O. Kwarteng, personal communication). Outside of the reserve, there is partial but extensive deforestation caused by the establishment of several villages and numerous small farms. The farms are interspersed between patches of degraded forest and support mixed crops, including small oil palm and cacao plantations. Twelve study sites were selected to make an exploratory study on how logging and the establishment of plantations affect local dung beetle assemblages. Four sites were selected in oil palm plantations, four sites in selectively logged forest, and four in unlogged forest. To be able to conduct quantitative trapping of dung beetles within an acceptable period of time, six of the eight trapping sites within Ankasa Resources Reserve were placed alongside footpaths branching from the two main access roads. These roads extend to the exploration camp from the Ankasa Gate to the south (8 km) and from the Elubo gate to the west (10 km). Other than a track alongside the pylons of the power line that crosses the southern part of Ankasa, all other access was by a system of footpaths. Outside of the reserve, trapping sites were placed on farms along the access road to the Ankasa Gate and near the Elubo gate along the main tarmac road from Accra to Elubo (5 17 N, 2 46 W) on the border with Cote dIvoire. Trapping sites were arranged from the southern edge toward the center of the reserve. Specically sites were placed in plantations a short distance from the gates (two), in plantations just outside of the gates (two), in the forest just within the gates (one logged and one unlogged), in forest halfway along the access roads (one logged and one unlogged), further into the forest reserve near the exploration camp (one logged and one unlogged), and along the Dyers Camp footpath that leads toward the center of the forest reserve (one logged and one unlogged). These latter sites were the closest to Nini-Suhien National Park, which borders the northern edge of the Ankasa Resources Reserve. Three traps were placed at least 10 m apart at each site. The traps were 2-liter plastic buckets (top diameter: 17 cm, depth: 14 cm) sunk into the soil so that the

rim was level with the surface. Pig dung was used for baits because it was readily available and acted as a surrogate for the dung of forest primates and pigs. Each bait was 250 ml of dung wrapped in chiffon and was supported over the center of each trap using two strong wires. Trap catches were immobilized using a little water and detergent placed in the base of the traps. An average of 350 mm of rain falls during June in Ankasa. Therefore, most days were predominantly cloudy with frequent rain. However, traps were baited in the morning on two 24-h trapping occasions (23 and 3 4 June) when there was little rain and numerous sunny periods, although day 2 was more cloudy than day 1. Trap catches were removed and preserved in alcohol after each 24-h period to make 24 samples for each of the three habitat categories. Voucher specimens of recorded species were placed in the T. K. Philips collection, the Museum of Natural History, Northern Flagship Institution, Pretoria, and the Museum National dHistoire Naturelle, Paris. During the rst baiting at oil palm plantations, it was discovered that site 3 was waterlogged. Because there were no refrigeration facilities and no local sources for large quantities of pig dung, it was decided not to risk bait deterioration by restarting the following day, but rather to move the traps to higher ground and trap from the edge of the oil palms into a cacao plantation, which differs physiognomically. Oil palm plantations show higher light intensity and a diverse understory of herbs and shrubs that requires frequent clearance. Cacao plantations show low light intensity and an understory comprised only of a thin groundcover of leaves. Although this low light intensity rivals that in natural rainforest, the forest differs in showing a diverse shrub understory with the ground covered by a litter of leaves, deadwood, and humus. Paired comparisons of species abundance structure were made between forest categories in Ankasa, between combined results for Ankasa and the adjoining plantations, and between Ankasa and June/July trap data previously recorded from the Western Upper Guinean rainforest of Ta National Park, Cote dIvoire (5 26 N, 6 55 W) (Cambefort and Walter 1991). These comparisons were made using the Steinhaus similarity coefcient ( 100). This is expressed as S 2W/(A B) (Legendre and Legendre 1983), in which W is the sum of the smaller numbers in pairs of species abundance data converted to decimal proportions for the species in each habitat and in which A ( 1) and B ( 1) are the total sums of the decimalized abundances in the two habitats (A B 2). Further comparisons were made solely between Ankasa and adjoining plantations. Completeness of the species richness record in plantations and each forest category were assessed by comparing observed values with values predicted using the abundancebased coverage estimator (ACE) method of Chadzon et al. (1998) contained in the EstimateS v. 6, computer package (Colwell 2000). Relative contribution of habitat, sites, and trap day to variance in species richness, abundance, and diversity was assessed using multifac-

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Regional occurrence and abundance of 40 dung beetle species in three habitats of southwest Ghana Mean abundance/trap SD Plantation 0.28 0.00 0.00 0.20 11.97 37.98 0.00 0.00 0.00 0.69 0.20 0.95 0.00 9.70 0.00 0.33 0.61 0.00 0.00 0.28 1.34 0.00 0.28 0.00 0.50 0.00 1.00 3.88 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.28 0.40 7.57 192.27 Logged forest 0.20 3.67 6.68 3.21 0.00 0.20 0.47 0.86 0.44 0.00 0.00 0.90 3.13 0.00 2.51 0.00 0.00 6.92 0.00 3.19 0.00 0.44 8.80 5.32 1.04 0.47 0.86 0.00 2.86 0.60 1.38 0.84 0.33 0.28 0.00 0.00 0.00 0.00 0.00 0.00 Unlogged forest 1.76 2.57 4.14 2.00 0.00 0.00 0.70 0.70 0.47 0.00 0.00 1.86 2.04 0.00 1.22 0.00 0.00 4.51 0.55 5.35 0.00 0.00 9.45 5.18 0.82 0.20 0.00 0.00 1.21 0.33 0.44 0.80 0.20 0.00 0.20 0.20 0.00 0.00 0.20 0.00

Species and regional associationa Anachalcos cupreus (Fabricius) Neosisyphus angulicollis (Felsche) Sisyphus latus Boucomont Sisyphus eburneus Cambefort Sisyphus sp. nr gazanus Arrowb Sisyphus seminulum Gerstaecker Pseudopedaria grossa (Thomson) Heliocopris dianae Hope Copris phungae Cambefortc Copris carmelita (Fabricius) Catharsius sp. Diastellopalpus tridens (Fabricius) Diastellopalpus noctis (Thomson) Onthophagus atridorsis dOrbigny Onthophagus atronitidus dOrbigny Onthophagus bidens (Olivier) Onthophagus curvifrons dOrbigny Onthophagus densepilis dOrbigny Onthophagus depilis dOrbigny Onthophagus deplanatus Lansberge Onthophagus feai dOrbigny Onthophagus fouillouxi Cambefort Onthophagus fuscatus dOrbigny Onthophagus fuscidorsis dOrbigny Onthophagus ieti Cambefort Onthophagus laeviceps dOrbigny Onthophagus liberianus Lansberge Onthophagus mucronatus Thomson Onthophagus rufopygus Frey Onthophagus semiviridis dOrbigny Onthophagus strictestriatus dOrbigny Onthophagus synceri Cambefort Onthophagus sp. a Onthophagus sp. b Onthophagus sp. c Onthophagus sp. d Onthophagus sp. e Onthophagus sp. f Drepanocerus caelatus (Gerstacker) Tiniocellus spinipes (Roth)d SFP FTA FTA FTA SP SP FTA FTA FTA SP ? FTA FTA SP FTA SP FTP FTA FTA FTA FTP FTA FTA FTA FTA FTA FTA SP FTA FTA FTA FTA ? ? ? ? ? ? SP SP

Plantation 0.08 0.00 0.00 0.04 12.50 47.33 0.00 0.00 0.00 0.33 0.04 0.63 0.00 8.79 0.00 0.13 0.29 0.00 0.00 0.08 0.96 0.00 0.08 0.00 0.21 0.00 0.54 2.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.08 0.08 9.88 253.33

Logged forest 0.04 4.13 4.00 2.63 0.00 0.04 0.17 0.63 0.13 0.00 0.00 0.33 4.67 0.00 2.29 0.00 0.00 7.13 0.00 0.88 0.00 0.13 15.58 11.75 0.79 0.17 0.58 0.00 3.04 0.13 0.92 0.29 0.13 0.08 0.00 0.00 0.00 0.00 0.00 0.00

Unlogged forest 1.00 3.29 3.00 1.46 0.00 0.00 0.38 0.42 0.17 0.00 0.00 1.17 1.63 0.00 1.38 0.00 0.00 5.92 0.33 2.46 0.00 0.00 13.38 5.50 0.54 0.04 0.00 0.00 1.17 0.13 0.13 0.17 0.04 0.00 0.04 0.04 0.00 0.00 0.04 0.00

a Regional associations from Cambefort (1991), Cambefort and Walter (1991), and Ankasa data. FTA, recorded in both Ta and Ankasa forests, entirely or primarily restricted to forests on omnivore dung (human or pig; 22 species); FTP, recorded as rarities in Ta forest but only in plantations at Ankasa (2 species); SP, recorded in savannas of Cote dIvoire and primarily in plantations at Ankasa (8 species): SFP, recorded in savannas of Cote dIvoire and in both rainforest and plantations at Ankasa (1 species); ?, unnamed and, therefore, unclassied (7 species). b Cited by Cambefort (1991) as Sisyphus gazanus Arrow. c Cited by Cambefort and Walter (1991) as Copris amabilis Kolbe. d Currently considered a pan-African savanna species, but there are subtle differences between the smaller-bodied populations of nergrained soils in moist West and East African savannas compared with those of sandier soils in dryer southern African savannas.

tor analysis of variance (MANOVA) of log10-transformed data (total values in each of three traps at each of four sites in each of three habitats on each of 2 trap d). For the MANOVA design, sites were nested in habitats and days were treated as repeated measures. Similarity in overall species representation in Ankasa and plantations was measured using the percentage disagreement distance measure (StatSoft 1995). This is expressed as Distance (x, y) 100 ( xi yi/i), in which i is the total number of species, and xi yi is the number of instances in which a species is present in one habitat and absent from the other. This measure was converted to percentage agreement (100 Distance [x, y]). Species abundance similarities between study sites were compared using cluster analysis and ordination. For the cluster analysis, a nonmetric Bray-

Curtis similarity matrix was calculated from an untransformed, site mean species abundance data matrix (12 sites 40 species), and subjected to analysis by the agglomerative clustering technique, group average linking. For the ordination, the site species abundance data matrix was fourth root-transformed. A correlation matrix was calculated from the data matrix and subjected to factor analysis using principal components as the method of factor extraction. Results A total of 40 species were recorded (Table 1), with forest showing greater overall species richness (29 species in 48 samples) than plantations (20 species in 24 samples). If trapping intensity is equalized (24

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Table 2. Species richness, abundance, and Shannon-Wiener diversity of southwest Ghana dung beetles Plantation Totals in 24 traps Observed species (N) Predicted species (N) Abundance (N) Diversity (H) Mean/trap SD Observed species (N) Day 1 Day 2 Predicted species (N) Day 1 Day 2 Abundance (N) Day 1 Day 2 Diversity Day 1 Day 2 Logged forest 25 25.9 1,455 2.31 10.9 10.6 13.4 14.4 62.8 70.3 1.99 1.86 1.9 3.2 3.9 7.3 24.8 26.2 0.15 0.27 Unlogged forest 25 30.1 1,051 2.34 10.3 8.8 14.8 11.6 44.5 43.1 1.91 1.72 2.2 3.5 3.9 4.3 16.9 28.5 0.20 0.38

20 21.3 8,098 0.91 7.6 7.7 10.2 10.7 432.5 242.3 0.95 0.92 1.3 2.9 2.7 5.6 276.2 148.6 0.26 0.23

samples per habitat), species richness remains greater in each of the two forest categories compared with plantations (Table 2). Forest and plantation assemblages showed extreme dissimilarity in species abundance structure (Tables 13; Figs. 1 and 2). Only eight species were shared between forest and plantations (Table 1). Relative abundances between habitats were extremely unbalanced in six of these species, whereas those of the other two were more evenly shared but contributed little to overall abundance (Table 1). Tiniocellus spinipes (Roth) was superabundant in plantations (Table 1) and showed extreme numerical dominance. This species was the primary contributor to the greater overall abundance in plantations and the lower Shannon-Wiener diversity than in either logged or unlogged forest. This was a consistent trend as mean values per site showed signicantly greater abundance and signicantly lower species richness and diversity
Table 3. Results for MANOVA on log10-transformed species diversity of southwest Ghana dung richness, abundance, and beetles MS effect Observed species (N) Habitat Sites Days Habitat days Sites days Abundance Habitat Sites Days Habitat days Sites days Diversity Habitat Sites Days Habitat days Sites days 0.145 0.070 0.041 0.013 0.013 3.506 0.814 0.188 0.181 0.047 0.786 0.030 0.017 0.004 0.008 df 2,24 9,24 1,24 2,24 9,24 2,24 9,24 1,24 2,24 9,24 2,24 9,24 1,24 2,24 9,24 F value 10.77 5.19 5.42 1.72 1.75 44.71 10.38 18.01 17.31 4.49 115.46 4.39 4.31 1.00 2.10 P 0.001 0.001 0.05 0.20 0.13 0.001 0.001 0.001 0.001 0.01 0.001 0.02 0.05 0.38 0.07

in plantations (Table 2). Although between-habitat differences were the largest contributor to variance (Table 3), there was also signicant within-habitat site variability, and signicant between-trap days variability. Post hoc tests showed that abundance differed signicantly between all three habitats (plantation to both forest types, P 0.001; unlogged to logged forest, P 0.05; Tukeys honestly signicant difference [HSD]). However, observed species richness and diversity only differed between plantations and forest (P 0.001 or P 0.05 in the case of plantation versus unlogged forest species richness; Tukeys HSD). There was slightly greater dung beetle species richness and abundance deeper into the conservation area at both unlogged and selectively logged sites, but trends were uneven. Moving from the reserve periphery, at unlogged sites, there were 18, 14, and 14 species, and 308, 104, and 205 individuals at sites 13 (each in 3 traps 2 d), respectively, whereas beyond the access roads, site 4 showed 23 species comprised of 434 individuals. Moving from the periphery, at selectively logged sites, there were 17 species each and 242, 448, and 236 individuals at sites 13, respectively, whereas beyond the access roads, there were 22 species comprised of 529 individuals at site 4. There were also differences between the dung beetle assemblages of the cacao and oil palm plantations (Fig. 1), which differed in physiognomy and microclimate. The differences in cacao largely result from lower species richness (10) and abundance (347) of the principal representatives recorded in the oil palm plantations where light intensity is higher (species richness at sites 1, 2, and 4, respectively: 11, 12, and 16; abundance, respectively: 2,788, 2,144, and 2,847). Only one species recorded in the cacao plantation (Onthophagus sp. f) was not also recorded under oil palms. Of the 29 species comprising the forest assemblage (Table 1), 22 of 25 named species (88%), were also recorded on human dung in the Western Upper Guinean forest of Ta National Park, Cote dIvoire, although 7 of these (28%) were not recorded during June/July. A further six human dung species recorded in Ta during June/July were not recorded in Ankasa, although it should be noted that four of the Ankasa species remain unnamed. In comparison, there was only 22.5% species similarity between Ankasa and the peripheral plantations (100 percentage disagreement measure). Although Steinhaus species abundance similarity ( 100) for June/July assemblages on omnivore dung was 82.1% between forest categories in Ankasa and 35.1% between Ankasa and the more distant Ta, it was an extremely low (0.01%) between Ankasa and adjacent plantations. All of the numerically dominant species in plantations were recorded only in savannas in Cote dIvoire (Table 1). Discussion This exploratory study was designed primarily to quantify some effects of deforestation on dung beetle faunas in West Africa. The species inventory was a

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Fig. 1. Dendrogram showing similarities between plantation and rainforest dung beetle faunas along the southern edge of Ankasa Conservation Area, southwest Ghana.

secondary consideration because it was necessarily limited to beetles recorded on a single dung type within a single month. During 1980 1981, a more extensive inventory study was conducted in the Western

Upper Guinean rainforest of Ta National Park, Cote dIvoire (Cambefort and Walter 1991). These Cote dIvoire data are probably a good guide to the overall annual species inventory and species richness that

Fig. 2. Ordination plot showing the statistical distance between plantation and rainforest dung beetle faunas along the southern edge of Ankasa Conservation Area, southwest Ghana.

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might be expected in Ankasa because the two reserves occur in the same climatic region (Type I2) (Walter and Lieth 1964). In Ta, Cambefort and Walter (1991) trapped scar abaeine dung beetles over six occasions, once every 2 mo, using seven different bait types. They recorded a total of 75 species. Most individuals of 66 species were recorded primarily on human (43 species) and/or elephant dung (44 species), whereas the balance was comprised of carrion or rotting vegetation specialists and rarities recorded on other dung types (all three species each). It might be predicted that a similar trapping effort would record a similar number in Ankasa, given that primates, elephants, carrion, and rotting vegetation all occur in the reserve. Of the 43 species trapped by human feces in Ta, only 24 species were recorded during JuneJuly 1980, and only 29 species were trapped by pig dung in Ankasa during June 2003. However, within these limited samples, 88% of the identied Ankasa species were in common between the two reserves and different omnivore dung types. These similarities suggest that differences in the species inventory and species richness caused by seasonal uctuations would be comparable between the two reserves and characteristic of the tropical bimodal rainfall climate type I2 (Walter and Lieth 1964). This climate type is wet from March to July (peaking in June) and from September to November (peaking in September in Ta and in October in Ankasa) ( 100 mm rain/mo on average). It is dryer from December to February and in August ( 100 mm rain/mo on average). If seasonal uctuations in the Ankasa fauna largely parallel those in Ta, one would predict that the driest period during December to January will yield the lowest annual values for species richness and abundance as in Ta (Cambe fort and Walter 1991), that the greatest of the two peaks in annual rainfall in June will yield the next lowest annual value for species richness as for June July in Ta, and that the preceding wet (AprilMay) and succeeding moist to wet (AugustSeptember) periods will produce the annual high points in species richness and abundance as in Ta. There are various threats to the dung beetle fauna of the Eastern Upper Guinean rainforests. One major threat derives from deforestation and habitat fragmentation by forestry (FAO 1988), agriculture (Gyasi et al. 1995), and mining interests (Anane 2003). A further threat derives from the removal of dung-producing mammals by the bush meat trade (Fa et al. 2005). According to Lebbie (2001), the greatest threat to regional forest conservation remains slash-and-burn agriculture. Although this study examines only one aspect of agricultural deforestation, creation of plantations clearly causes the virtual disappearance of the specialist rainforest dung beetle fauna attracted to omnivore dung and its replacement by species that are abundant in savannas (Cambefort 1991) but absent from rainforest (Cambefort and Walter 1991). This suggests that the plantation faunas are largely derived from range expansion by shade tolerant species centered in the savannas. The change is presumably as-

sociated primarily with the differences in microclimate and surface cover between natural and agricultural habitats because it is probable that omnivore dung is equally available in both forest (wild porcines, nonhuman primates) and plantations (domestic pigs, humans). Although differences have been shown between dung beetle assemblages of unlogged and selectively logged rainforests (Nummelin and Hanski 1989, Davis et al. 2000, 2001), there were no consistent differences in species richness or diversity between rainforest categories in this study. Because some 30 40 yr has elapsed since disturbance at Ankasa, the results may imply a great deal of ecological recovery over this period. However, this assumes that selective logging was initially disruptive. Although this cannot now be ascertained, there is, currently, an absence of obvious differences in oral physiognomy between the two forest categories, and this is reected by their largely similar forest faunas. In contrast, oil palm and cacao plantations, with their obvious physiognomic differences, show signicant reductions in dung beetle species richness and diversity, which parallel the patterns in clear cuts after partial deforestation within Brazilian rainforest (Klein 1989) or in an agricultural landscape in Mexico (Estrada et al. 1998). However, whereas transformed Neotropical habitats showed lower abundance compared with rainforests, West African plantations showed signicantly higher abundance than in rainforest. This difference may be caused by the absence of a nearby source of adaptable savanna dung beetles in the Neotropical localities (Davis et al. 2004). The bush meat trade decreases mammal and dung density and may nally remove many of the mammals on which dung beetles rely for their survival. Even with only partial removal of the mammal fauna, species richness may be affected because some species of forest dung beetles are recorded primarily on the dung characteristic of particular groups such as omnivores (primates, wild pigs) or large nonruminant (monogastric) herbivores (elephants) (Cambefort and Walter 1991). Figures on density and seasonal movements of different mammal species in Ankasa were unavailable to the authors, so it was not possible to predict the spatial distribution of dung and its effect on species abundance composition of dung beetles. However, if edge effects and bush meat hunting for forest omnivores (primates, wild pigs) have inuenced these results, these might be predicted to be greatest peripherally in Ankasa. Although there was slightly greater dung beetle species richness and abundance deeper into the conservation area, trends were uneven. In unlogged forest, site 2, the particularly low species richness and abundance may be symptomatic of roadside environmental stress because there was a large amount of dead wood littering the surface. However, further work is necessary to conrm if higher species richness and abundance beyond the access roads is a consistent trend attributable to lower disturbance and/or greater density of dung. In conclusion, plantation development is clearly detrimental to the specialized rainforest omnivore

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dung beetle assemblages of West Africa. It may be predicted that deforestation through agricultural and industrial practices would result in the rarity or loss of most species recorded in rainforest. Forest reserves, protected from deforestation and the bush meat trade, are essential for the survival of these specialist assemblages.

Acknowledgments
We thank M. Adu-Asiah, Senior Wildlife Ofcer with the Ghana Wildlife Department, for permits to sample dung beetles. Pig dung for baits was kindly supplied by E. A. Canacoo and A. Naazie of the Experimental Farm, University of Accra, and by S. Tete and I. Atinga of the adjoining private farm. The local knowledge and technical assistance of our guide, S. O. Kwarteng (Lobito), proved invaluable for securing introductions, selecting study sites, and setting traps in the Ankasa area. D. Foster, graduate student at West Kentucky University, also provided helpful technical assistance in Ghana. In addition, we thank K. Asamoah, K. Lodo, B. Quame, and M. Armah for permission to sample dung beetles on their farms. Y. Cambefort of the University of Paris-7 was instrumental in facilitating the production of this manuscript by kindly providing dung beetle species names using a reference collection drawn from the samples. We sincerely thank the National Science FoundationEpscor Research Enhancement Grant Program for funding this study. Support provided by the Faculty Scholarship Council at Western Kentucky University is also gratefully acknowledged.

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