Only a handful of bird species are known to use foraging tools in the wild. Amongst them, the New Caledonian crow (Corvus moneduloides) stands out with its sophisticated tool-making skills. Despite considerable speculation, the evolutionary origins of this species' remarkable tool behaviour remain largely unknown, not least because no naturally tool-using congeners have yet been identified that would enable informative comparisons. Here we show that another tropical corvid, the 'Alalā (C. hawaiiensis; Hawaiian crow), is a highly dexterous tool user. Although the 'Alalā became extinct in the wild in the early 2000s, and currently survives only in captivity, at least two lines of evidence suggest that tool use is part of the species' natural behavioural repertoire: juveniles develop functional tool use without training, or social input from adults; and proficient tool use is a species-wide capacity. 'Alalā and New Caledonian crows evolved in similar environments on remote tropical islands, yet are only distantly related, suggesting that their technical abilities arose convergently. This supports the idea that avian foraging tool use is facilitated by ecological conditions typical of islands, such as reduced competition for embedded prey and low predation risk. Our discovery creates exciting opportunities for comparative research on multiple tool-using and non-tool-using corvid species. Such work will in turn pave the way for replicated cross-taxonomic comparisons with the primate lineage, enabling valuable insights into the evolutionary origins of tool-using behaviour.

Abstract. Species loss can lead to cascading effects on communities, including the disruption of ecological processes such as seed dispersal. The endangered 'Alalā (Corvus hawaiiensis), the largest remaining species of native Hawaiian forest bird, was once common in mesic and dry forests on the Big Island of Hawai'i, but today it exists solely in captivity. Prior to its extinction in the wild, the 'Alalā may have helped to establish and maintain native Hawaiian forest communities by dispersing seeds of a wide variety of native plants. In the absence of 'Alalā, the structure and composition of Hawai'i's forests may be changing, and some large-fruited plants may be dispersal limited, persisting primarily as ecological anachronisms. We fed captive 'Alalā a variety of native fruits, documented behaviors relating to seed dispersal, and measured the germination success of seeds that passed through the gut of 'Alalā relative to the germination success of seeds in control groups. 'Alalā ate and carried 14 native fruits and provided germination benefits to several species by ingesting their seeds. Our results suggest that some plants rely heavily on 'Alalā for these services. In captivity, juvenile birds displayed seed dispersal behaviors more often than adult birds for most fruiting plants in our study. We introduced captive 'Alalā to two large-fruited, dry-forest plants, not previously recorded as 'Alalā food resources, but which may once have been part of their natural diet. The seed dispersal behavior that 'Alalā displayed toward these species supports the inclusion of dry and mesic forests in 'Alalā habitat restoration plans and adds weight to the idea that plant dispersal limitation may contribute to the rarity of these plants. Our study provides evidence that 'Alalā have the capacity to play a vital role in maintaining the diversity of fruiting plants in native Hawaiian forests through seed dispersal and enhanced seed germination, thus adding greater urgency to efforts to restore 'Alalā to their former range.

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