Farmland ecosystem services frequently result from different ecological functions simultaneous provided by specific biodiversity groups such as birds. These bundles of ecosystem services may be approached from inter-specific interaction networks, which inform about the structure of interactions while identifying the species more relevant for combining ecosystem services. Here, we studied how birds provide pest control in apple orchards, and seed dispersal in orchard-adjacent hedgerows and forests. For this, we used field data and DNA-metabarcoding of bird fecal samples, obtained across a whole year from three orchards in northern Spain, to build interaction networks between birds and arthropod apple pests, and between birds and non-crop fleshy-fruited plants. We addressed the structure of the different networks underpinning pest control and seed dispersal. We also combined both ecosystem services through a hybrid network to assess, by means of centrality measures, the topological roles of individual bird species and their ecological determinants. Interaction networks differed in structure, with pest-control network showing higher modularity and specialization than seed-dispersal network, otherwise characterized by high nestedness. These differences emerged from the different typologies, i.e. antagonistic vs. mutualistic, of the bird-pest and bird-plant interactions. The hybrid network integrating pest control and seed dispersal evidenced strong variability across birds in their role to connect ecosystem services, depending on their central or peripheral positions. Bird centrality was positively related with species abundance and independent of body size or diet diversity. This finding suggests the relevance of neutral processes (i.e. the random encounters of organisms) for driving inter-specific interactions, and it highlights the relevance of common birds for sustaining ecosystem service bundles in agroecosystems. The preservation of fruit-rich hedgerows and forest patches around apple orchards is an action recommended to maintain the populations of common bird species, responsible of both agricultural pest control and the natural regeneration of these habitats.

中文翻译：

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常见鸟类通过混合互动网络将苹果园中的害虫防治和种子传播结合起来

农田生态系统服务通常是由鸟类等特定生物多样性群体同时提供的不同生态功能产生的。这些生态系统服务包可以通过特定的相互作用网络来获取，该网络可以了解相互作用的结构，同时识别与组合生态系统服务更相关的物种。在这里，我们研究了鸟类如何在苹果园中进行害虫防治，以及如何在果园附近的树篱和森林中传播种子。为此，我们使用从西班牙北部三个果园全年获得的鸟类粪便样本的现场数据和 DNA 元条形码，建立鸟类与节肢动物苹果害虫之间以及鸟类与非农作物肉质果植物之间的相互作用网络。我们讨论了支持害虫控制和种子传播的不同网络的结构。我们还通过混合网络将两种生态系统服务结合起来，通过中心性度量来评估单个鸟类的拓扑作用及其生态决定因素。交互网络在结构上有所不同，害虫控制网络比种子传播网络表现出更高的模块化和专业化，否则具有高嵌套性。这些差异源于鸟类与害虫以及鸟类与植物相互作用的不同类型，即拮抗与互惠。结合害虫防治和种子传播的混合网络证明，鸟类在连接生态系统服务方面的作用存在很大差异，具体取决于它们的中心或外围位置。鸟类中心性与物种丰富度呈正相关，与体型或饮食多样性无关。这一发现表明中性过程（即生物体的随机遭遇）对于驱动种间相互作用的相关性，并且强调了常见鸟类对于维持农业生态系统中的生态系统服务束的相关性。保护苹果园周围水果丰富的树篱和森林斑块是一项建议采取的行动，以维持常见鸟类的种群，负责农业害虫控制和这些栖息地的自然更新。