4.1 Land-use and biodiversity
In this study we observed that land use changes in the highlands of La Antigua watershed have resulted in loss of species richness and biodiversity (Martınez et al 2009). Landscape condition has usually been used as a proxy for habitat and species diversity, with disturbed land cover signifying a deterioration in habitat condition and diversity (Boakye et al. 2015; DeWalt et. al. 2003; Garcia-Montiel et al. 2007; Liang and Liu 2017). Though habitat loss is due to vegetation clearing, the habitat modification and fragmentation of residual habitats also exert serious secondary deleterious effect (Feurer 2013; Oy 2014; Pambudi and Rahayu 2017). The results of this study indicate that land-use changes in the two watersheds have resulted in the decline of species richness and diversity. Biodiversity loss increased with the land-use transformation, from no effect in Open forest to more than 95 per cent loss in Food Crop landuse. This finding from the study is similar to other studies that also found a reduction in species richness also found reduction in species richness of woody species from 40 in Forest to 19 in Farmland in the northern region of Ghana (Boakye et al, 2015) and reduction from 137 and 121 species in Forest to 14 and 2 species in Cocoa agroforest and rubber plantation in the Tano Offin Forest Reserve and Cape Three Points Forest Reseve respectively in Ghana (Ayesu, 2018). Species diversity in most cases are higher in less disturbed forest than in more intensively managed lands like Cropland (Tree crop and Food crop) where land preparation including burning leads to depletion of trees and other wood plants (Boakye et al. 2015; Frank, 2012; Mensah et al. 2016; Rudmann-Maurer et al. 2008). In contrast, several studies also found that species diversity in agricultural systems in the tropics was improved by the retention of trees on croplands by farmers (Boakye et al. 2015, 2016; Käyhkö et al. 2011; Kotoky et. al., 2012; Tendaupenyu et. al., 2017).
The decrease in species richness and diversity with increasing forest degradation and deforestation may reduce resilience of such socio-ecological systems. This in the long term would result in climate change induced impacts such as spread of invasive species (Ruf et. al., 2015; Sandifer et. al., 2015; Tilman 1999). Also, the loss of plant diversity may result in poor habitat conditions for species that facilitate provision of important ecosystem services, including crop pollination, nutrient cycling and seed dispersal (Addai and Baidoo 2013). Species diversity has been reported for making ecosystems more resistant and resilient to disturbances. Diverse plant communities are more likely to adapt to environmental change. This would facilitate the provision of essential goods and services within an ecosystem in order to continuously promote and maintain a diverse composition of species (Asante-Yeboah 2010). Landscape connectivity have been found to relate strongly with a biodiversity (Asase et. al. 2012; Bowen et al. 2007; Attua et. al. 2018; Castro et al. 2016; Martínez et al. 2009; Taylor et. al. 2014). Though the findings relate to other recent research works which regulated the effects of ecological variables (Boakye et al. 2015, 2016; Polasky et al. 2011), it is consistent with the generally accepted pattern in very diverse natural forests (Asase et. al. 2012; Pappoe et al. 2010).
4.2 Stand structure
Stand structural characteristics helps estimate forest biomass and could be used to generate geographical information on factors that influence species distributions (Couteron et al., 2005). In this work, stand structure refers to tree basal area, tree density and tree diameter-size class distribution. The results indicate that large diameter classes were generally absent in intensively managed landuses except in the case of agroforest tree crop systems that was characterized by retention of trees. In the Barekese watershed, large tree > 110 cm bdh were even absent in Closed Forestand open forest but were present in tree crop systems. The situation was different for the Owabi watershed where Open forest and Closed Foresthad representation in all the diameter class both small and large diameter classes whiles diameter classes in tree crops systems were below 130 cm dbh. This variation in diameter representation could be due to the level of protection accorded the two sites (Forestry Commission, 2014). Most of the diameter class recorded in the Owabi sites were found within the Wildlife Sanctuary which is continuously patrolled by the staff of the Wildlife Division.
The Barekese watershed on the other hand is not adequately protected and therefore has over the years been encroached by illegal chainsaw operators who harvest the large sized trees (Forestry Commission, 2014; Atwima Nwabiagya District Assembly, 2017). These activities have over the years contributed to the depletion of large sized trees within Barekese watershed. Basal area estimates were relatively higher for both close and open forests as compared to other landuses. The presence of large trees in close and open forests contributed to the higher basal area estimates for the Closed Forestand open forest at both sites. The mean tree basal area of 47.62m2/ha and 36.81m2/ha for Closed Forestat Owabi and Barekese respectively were higher than that estimated for a similar forest type in the Kakum National Park of 33.76m2/ha (Pappoe et. al. 2010). The higher basal area estimates for close and open forest compared with the tree crop, food crop and grassland landuse suggests that the conversion of natural forest to other landuse use affect the stand structure in respect of the total covered by trees per unit area. High basal area estimates are typical of established forest stands and reflect growth performance of trees.
4.3 Land use change effects on carbon stocks
The overall mean density for all carbon pools recorded for Owabi watershed (276.1 tC ha − 1) and Barekese watershed (168 tC ha − 1) were higher than the global default value of 141 MgC ha − 1 as stated by Aalde et al. (2006) in his studies in tropical forests. Baccini et al. (2012) indicated that smaller carbon densities were reported for tropical Africa (82 MgC ha − 1) compared with other tropical areas with 116 Mg C ha − 1 for America and 119 Mg C ha − 1 for Asia. The conversion of natural forests to other land-use systems has resulted in a significant reduction in mean carbon stocks by 65–79% (food crop system) and 73–84% (Grassland) for Barekese and Owabi watersheds respectively. Also, conversion Closed Forestto open forest and Tree crop system resulted 36% − 39% and 55–69% carbon loss respectively. The outcome of this study also clearly indicates that land-use change has significant influence on mean carbon stocks per hectare. Carbon stock was found to decrease with increasing intensity in the use of land for cultivation and this findings coroborate other recent studies that reported similar results (Pasco, 2013; Ensslin et al. 2015; Feurer 2013; Grieco et. al., 2012; Makafui and Agboadoh 2011; Pambudi and Rahayu 2017). Adu-Bredu et al. 2008 observed carbon loss with increased with conversion of forest to other land uses; Natural forest (326.75 MgCha-1), Teak plantation (138.33 MgCha-1), Fallow (95.46 MgCha-1) and Cultivated areas (75.12 MgCha-1). Also, a decrease in mean carbon stocks in the dense forests (181.78 ± 27.06 MgC ha − 1, open forests (104.83 ± 12.35 MgC ha − 1), Grasslands (108.77 ± 6.77 MgC ha − 1 and cultivated lands (83.11 ± 8.53 MgC ha − 1) for Afromontane forest in Ethiopia (Solomon et al. 2018). Other studies examining the effects of land-use change on carbon storage found a decrease in carbon stocks with increasing land cover transformation (Pellikka et al. 2018; Kuusipalo et al. 1996; Arets, 2005; Swaine and Agyeman 2008). In most cases, forest conversion lead to the removal of a significant quantity of large trees which are considered to store the greatest amount of carbon (Kirby and Potvin 2007). Mean carbon stocks were significantly higher for Owabi compared to the Barekese watershed. This was attributed to the high incidence of illegal logging, wildfire incidences and intensive farming activities that have denuded the landscape of large diameter size class trees (Atwima Nwabiagya and District Assembly 2017; Ghana Statistical Service 2014). For instance, trees greater than 90 cm DBH are virtually missing in the case of Barekese watershed. These results are also consistent with the Forest Transition Model (Reed et al 2017; Pellikka et al., 2018).
However, in this investigation, higher carbon densities were recorded for tree croplands than the food crops systems and grassland. This observation is due to the agroforestry practices in tree crop systems through the deliberate retention of indigenous trees on farms. The retained trees are characteristically of large diameter size which contribuite substantially to carbon storage. This indicates greater tree volume in cropland system, in cases when agroforestry is practiced. This emphasizes the importance of retained trees on farms in increasing carbon storage (Dawoe et. al., 2016; Mohammed et. al., 2016; Pellikka et al. 2018). Although land-use change resulted in forest degradation, deforestation had a negative effect on carbon storage. However, the findings of this study suggest that Open forest and tree crop land-use systems were less detrimental as compared to Food crop systems and Grasslands. The findings of this study suggest that Food crops and Grasslands are less favourable ecological systems for carbon storage. This supports the research findings of Luedeling et al. (2011) and (Ensslin et al. 2015) who suggested that carbon stocks in agricultural landscapes depend mostly on density and diameter size of trees.There absence of significant difference in mean carbon stocks between Closed Forestand open forest for both watersheds indicates that selective removal of large individual trees did not affect carbon stocks. This observation supports other findings in West, East and Central Africa that selective removal of trees did not affect carbon stocks (Asase et. al., 2012; Ensslin et al. 2015). Though biomass regenerate rapidly, degraded forests may vary in species richness, composition and structure. The results stress on the negative effect of land-use change on carbon stock accumulation and the need to minimize natural forest disturbances to less complex systems.
Implications for watershed management
Generally, there is a positive relationship between biodiversity and carbon stocks globally (Midgley et al., 2010). For instance, natural tropical moist forests unaffected by anthropogenic disturbances are rich in both carbon stocks and biodiversity. Within tropical forests, there were less correlation between spatial patterns of carbon stocks and biodiversity in undisturbed areas and these patterns are considered complex (Talbot, 2010). In this study, a positive and significant effect was found between species richness, diversity and amount of carbon stocks for the two sites. Whilst the findings of this study are consistent with other studies (Asante-yeboah 2010; Asase et. al., 2012; Dawoe et al. 2016), it also supports the commonly described pattern in highly diverse natural forests; thus, biomass and carbon stocks increase with increasing diversity. Several studies on forest ecosystems have shown a positive relationship between species richness and forest biomass and/or carbon at the local, regional and global level (Asase et. al., 2012; Dawoe et al. 2016). In particular, studies across various ecosystems such as boreal (Paquette & Messier, 2011), temperate (Paquette & Messier, 2011; Vilà et al., 2007), and tropical forests (Barrufol et al., 2013) have reported an increase in productivity with increasing diversity. Increasing species richness and diversity usually facilitate increased carbon storage since higher taxonomic diversity lead to higher stem density and forest productivity (Ruiz- Benito et al., 2014). The positive effects of species diversity can be explained through the benefits of plant–plant interactions through facilitation, a process where by some species could enhance soil fertility (by fixing nitrogen) for the productivity of other species. This fact is often used to support the reason that mixed species communities of plantations are far more productive than mono-specific stands. However, it is possible that increasing species richness increases the chances of inclusion of highly productive and naturally favoured dominant species (Ruiz- Benito et al., 2014) as shown by Mensah et. al., (2016) on the influence of most dominant species on carbon stocks in Mistbelt tropical forests in Cameroun.
Whilst the findings support the positive species richness–diversity–carbon relationship, however other studies provide evidence of the inverse effect also exists. For instance, studies by Ruiz- Jaen and Potvin (2011) in the natural forest of Barro, in Central Panama and Szwagrzyk and Gazda (2007) in natural forests of central Europe, reported a negative relationship of species diversity and plant biomass. Furthermore, other studies concluded that such relationships were significant (Gairola et al., 2011). These opposing findings suggest that the effects of diversity on forest carbon may be explained by other environmental factors (Lasky et al., 2014; Wu et al., 2015) and also may dependent on the type of diversity measure used (Con et al., 2013; Lasky et al., 2014; Ouyang et al., 2016). The outcome also supports the idea that complementarity and selection effects do not exclusively affect carbon storage (Ruiz- Benito et al., 2014; Wu et al., 2015). That diversity promotes carbon stock through effects of functional diversity and dominance, and this is because these components of diversity are based on specific functional traits, which reflect functional differences among species (Song et al., 2014). This could also be due to the fact that increased species richness indirectly accounted for differences among species, in terms of ecological niche and resource use.