It is widely accepted that pollination services, provided by a diversity of pollinators, are important for global food security, ecosystem integrity, nutrition and health security, and human well-being (Chaplin-Kramer et al., 2014; Huttenhower et al., 2012; IPBES, 2016; Klein et al., 2018; Ollerton, 2021; Potts et al., 2016). Pollination is an indisputably critical ecosystem service, with approximately 90% of our wild-growing plant communities (Ollerton et al., 2011) and 75% of our leading global crop types dependent on pollinators, to varying degrees (IPBES, 2016; Klein et al., 2007). This is no exception for India, which, together with China and the USA, accounts for approximately 30% of the total global cropland (FAO, 2020); important crops for India include mangoes, fresh vegetables, tomatoes, soybeans, eggplant, cardamom, coffee, and others — which are all pollinator-dependent (Abraham et al., 2013; Agarwal et al., 2013; Basu et al., 2011; Bhattacharya & Basu, 2018; Cooley & Vallejo-Marín, 2021; Garibaldi et al., 2021; Huda et al., 2015; Krishnan et al., 2012).
Studies from various parts of the world have suggested that bees are on the decline (Biesmeijer et al., 2006; Cameron & Sadd, 2020; Gallai et al., 2009; IPBES, 2016; Koh et al., 2016; Orr et al., 2021; Potts et al., 2016). Yet, data and evidence are sparse from some regions. The largest drivers of biodiversity loss are land- and sea-use change, direct exploitation of nature, climate change, invasive alien species and pollution (IPBES, 2016), which unsurprisingly, match the drivers of pollinator decline (Dicks et al., 2021; Goulson et al., 2015). Dicks et al. (2021) found changes in land cover and configuration to be important drivers of pollinator loss in most regions worldwide; for the Asia-Pacific region, the study found pesticides to be among the top three drivers of pollinator loss, which differs from the global ranking of drivers for biodiversity loss overall (i.e., not the same relative rank of importance). In terms of land configuration, native forests are known to play a beneficial role in wild bee populations (Aizen & Feinsinger, 2003; Ghazoul et al., 1998; Schrader et al., 2018), largely because native trees and older trees provide suitable nesting substrates for bees such as Apis dorsata (Thomas et al., 2009) and Tetragonula species (Bhatta et al., 2019). Proximity to forests or semi-natural sites increases pollinator species richness and abundance within agricultural systems (Blanche et al., 2006; Blanche & Cunningham, 2005; Chacoff & Aizen, 2006; Free, 1993; Klein et al., 2003; Kremen et al., 2002, 2004; Morandin & Winston, 2006; Ricketts, 2004; Viswanathan et al., 2020, 2020; Widhiono & Sudiana, 2016). Natural forests and semi-natural plots play an important role for pollinators and their delivery of pollination services in several important cash crops, including coffee and spices.
One of the most important global agricultural commodities is coffee (Khoury et al., 2014), worth approximately $40 billion/yr in production value alone and significantly more when considering retail value (Rhiney et al., 2021); coffee prices reached an average of $190.82 US cents/lb (July 2022). The coffee industry involves upwards of twenty-five million coffee farmers and producers (Donald, 2004; Jha et al., 2011; Pham et al., 2019). Coffee production occupies over 10 million hectares of land (FAO, 2021). India is the seventh top coffee producer in the world (Sänger, 2018), making coffee an important cash crop for India (Bhattacharya, 2014), being the seventh top coffee grower in the world, India produces approximately 3% of the total global coffee export, equivalent to approximately 334,000 metric tons (estimated for 2020–2021, Coffee Board of India), which mainly comes from smallholder farmers (Chengappa et al., 2017; DaMatta et al., 2007; Ranjan Jena & Grote, 2017), some of whom are Indigenous forest-dwelling tribes. In the Western Ghats of India, coffee is traditionally grown under an agroforestry landscape with a diversity of native shade trees or intercropped with pepper, but in recent years, some coffee farms have shifted to a more intensified cultivation system (Chengappa et al., 2017). Specifically, in the coffee-growing slopes of Nilgiri Biosphere Reserve, which spans across the States of Tamil Nadu, Karnataka, and Kerala, the characteristic coffee agro-forest is densely planted with coffee bushes under an evergreen forest patch close to human habitation. However, some coffee growers have shifted to using an exotic fast-growing monocrop of silver oak (Grevillea robusta) trees (Bali et al., 2007) to shade their coffee crops. The impact of this shift on pollinator communities is unclear.
For coffee, fruit set is known to increase with cross-pollination by bees – as shown in the Indonesian coffee agroecosystems (Klein et al., 2003); fruit set increased by approximately 12.3% in C. arabica and 16% in C. robusta (Klein et al., 2003) with “open pollination” treatment where all insects could access coffee flowers compared to “control” where insect pollinators were excluded from the treatment. Pollination is important not only for the quantity of coffee production but also the quality; sufficient coffee pollination results in double-seeded full berries (fruit), while a pollination deficit leads to peaberries (i.e., the development of one ovule into a single-seeded fruit) (Classen et al., 2014; Ghazoul & Krishnan, 2012; Krishnan et al., 2012). Previous studies on pollinators and coffee production in other regions have shown that coffee fruit set increases when coffee is grown near rainforests, even fragments (De Marco & Coelho, 2004; Klein et al., 2003; Ricketts, 2004; Ricketts et al., 2004), since many wild bees nest in natural habitats – large tree branches and cavities, rocks, etc. (Aizen & Feinsinger, 2003; Cunningham, 2000; Ghazoul et al., 1998). In a recent global meta-analysis of Coffea arabica systems, Moreaux et al. (2022) found animal pollination to be important for increasing fruit set by approximately 18%, yet when looking at habitat features such as forest cover, distance to nearby forest and forest canopy density, the study found that forest cover and distance to open forest were not to be correlated to bee species richness and fruit set – which does not support previous research (e.g., Klein, 2009; Ricketts, 2004) highlighting the need for further research on this topic.
The Western Ghats of India is a biodiversity hotspot (Myers et al., 2000) and a UNESCO World Heritage Site with “high geological, cultural and aesthetic values” but has undergone extensive loss in forest cover and biodiversity owing to land-use change (Ambinakudige & Choi, 2009; Daniels et al., 1995; Gupta et al., 2014; Kale et al., 2016). Data has been documented the last 60 years for honey bee stocks (i.e., hives) for India and the trend has been increasing (approx. 5 million hives in 1961 to almost 13 million hives in 2021) at the same time, data are lacking on the status and trends of wild bees, which are important for coffee production (Pannure, 2016; Veddeler et al., 2008). Previous studies have cited Apis dorsata, the giant Asian honey bee, as the main pollinator of coffee in the Kodagu district (Karnataka); Krishnan et al. (2012) found A. dorsata individuals comprising almost 60% of floral visits recorded on coffee (C. canephora), which was associated with increased coffee fruit production by 50% (Krishnan et al., 2012).
There is a knowledge gap regarding the impacts of reducing shade tree diversity or changing tree cover type on bee pollinators and their pollination services to coffee crops in the Western Ghats. In a study in Kodagu, "a coffee-growing region where a high percentage of native tree cover is still intact to provide shade for coffee plants” (Bhagwat et al., 2005) shade tree diversity has been shown to enhance coffee production and quality of C. canephora while reducing incidences of the coffee berry borer Hypothenemus hampei (Nesper et al., 2017).
We assessed bee abundance in small holdings of Coffea arabica grown under two types of tree cover – native forest trees and silver oak (G. robusta). Based on previous work (Barrios et al., 2018; Nesper et al., 2017; Prado et al., 2018), we predicted that coffee farms with native tree cover would offer better delivery of pollination services compared to those with only silver oak (G. robusta); bee abundance was used as proxy indicator of pollination services. In order to determine bee abundance, only insect visitors observed foraging on pollen or nectar were considered potential pollinators. We quantified bee abundance as a function of (a) tree cover type, (b) distance from the nearest forest edge, and (c) area under coffee cultivation. We examined the influence of these three factors on abundance of Apis dorsata, A. cerana, A. florea, Tetragonula iridipennis and Xylocopa sp., the dominant pollinators of coffee. Although Xylocopa has been reported as a minor pollinator in several studies (Krishnan et al., 2012; Prado et al., 2019; Samnegård et al., 2016), their efficiency in pollination is believed to be high in comparison to social bees (Klein et al., 2003).