Cultural behavior, which is transmitted among conspecifics through social learning [1], is found across various taxa [2, 3, 4, 5, 6]. Vertical social transmission from parent to offspring [7] is thought to be adaptive because of the parental generation being more skilled than maturing individuals. It is found throughout the animal kingdom, particularly in species with prolonged parental care, e.g., [8, 9]. Social learning can also occur among members of the same generation [4, 10, 11] or between older, non-parental individuals and younger generations [7] via horizontal or oblique transmission, respectively. Extensive work on primate culture has shown that horizontal transmission of foraging behavior is biased toward species with broad cultural repertoires [12] and those with increased levels of social tolerance [13, 14], such as great apes. Vertical social transmission has been established as the primary transmission mechanism of foraging behaviors in the Indo-Pacific bottlenose dolphin (Tursiops aduncus) population of Shark Bay, Western Australia [6, 9, 15, 16]. Here, we investigated the spread of another foraging strategy, “shelling” [17], whereby some dolphins in this population feed on prey trapped inside large marine gastropod shells. Using a multi-network version of “network-based diffusion analysis” (NBDA), we show that shelling behavior spreads primarily through non-vertical social transmission. By statistically accounting for both environmental and genetic influences, our findings thus represent the first evidence of non-vertical transmission of a foraging tactic in toothed whales. This research suggests there are multiple transmission pathways of foraging behaviors in dolphins, highlighting the similarities between cetaceans and great apes in the nature of the transmission of cultural behaviors.

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中文翻译：

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整合遗传、环境和社交网络，揭示海豚觅食创新的传播途径。

文化行为通过社会学习在同种之间传播 [1]，可以在各种分类群中找到 [2, 3, 4, 5, 6]。从父母到后代的垂直社会传播 [7] 被认为是适应性的，因为父母一代比成熟的个体更熟练。它在整个动物界都有发现，特别是在有长期父母照顾的物种中，例如 [8, 9]。社会学习也可能发生在同一代人之间 [4, 10, 11] 或年长、非父母和年轻一代 [7] 之间，分别通过水平或斜向传播。对灵长类动物文化的广泛研究表明，觅食行为的横向传播偏向于具有广泛文化曲目的物种 [12] 和具有更高社会容忍度的物种 [13, 14]，例如类人猿。Tursiops aduncus ) 西澳大利亚鲨鱼湾的种群 [6, 9, 15, 16]。在这里，我们调查了另一种觅食策略“炮击”[17] 的传播，该种群中的一些海豚以被困在大型海洋腹足动物壳内的猎物为食。使用“基于网络的扩散分析”（NBDA）的多网络版本，我们表明炮击行为主要通过非垂直社会传播传播。通过对环境和遗传影响进行统计，我们的研究结果代表了齿鲸觅食策略的非垂直传播的第一个证据。这项研究表明，海豚的觅食行为存在多种传播途径，突出了鲸类和类人猿在文化行为传播性质上的相似之处。

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