High resilience of Pacific Island forests to a category- 5 cyclone.

Assessing how forests respond to, and recuperate from, cyclones is critical to understanding forest dynamics and planning for the impacts of climate change. Projected increases in the intensity and frequency of severe cyclones can threaten both forests and forest-dependent communities. The Pacific Islands are subject to frequent low-intensity cyclones, but there is little information on the effects of high intensity cyclones, or on how forest stewardship practices may affect outcomes. We assess the resistance and resilience of forests in three community-stewarded sites on the island of Tanna, Vanuatu, to the wind-related effects of 2015 Category-5 Cyclone Pam, one of the most intense cyclones to make landfall globally. Drawing on transect data established pre-and post-cyclone, we (1) test whether windspeed and tree structural traits predict survival and damage intensity, and whether this varies across sites; (2) assess post-cyclone regeneration of canopy, ground cover, seedlings, and saplings, and how community composition shifts over time and across sites. In sites that sustained a direct hit, 88 % of trees were defoliated, 34 % sustained severe damage, and immediate mortality was 13 %. Initial mortality, but not severe damage, was lower in areas that received an indirect hit and had lower windspeed. Larger trees and those with lighter wood had a higher probability of uprooting and snapping, respectively. Canopy and ground cover regenerated within three years and seedling and sapling regeneration was widespread across life histories, from pioneer to mature forest species. Three species of non-native vines recruited post-cyclone but within 5 years had largely declined or disappeared with canopy closure. Tanna's historical cyclone frequency, combined with customary stewardship practices that actively maintain a diversity of species and multiplicity of regeneration pathways, are likely responsible for the island's resistance and resilience to an intense tropical cyclone.

Dillenia (Dilleniaceae) pollen heteromorphy and presentation, and implications for pollination by bats.

Bat pollination of Dillenia in Fiji, a genus that was presumed to be pollinated by bees, posits that other Dillenia species may be bat-pollinated, with implications for conservation and the understanding of angiosperm evolution. Botanical descriptions of some corolla behaviours ('falling as a whole') suggest bat removal of permanently closed corollas, as in D. biflora. Considering the remoteness of species of interest, we reviewed some Dillenia floral traits to hypothesise what they may mean for bat pollination of the genus. We investigated D. biflora pollen grains apertures and reviewed Dillenia literature concerning corolla behaviour and colour, and pollen apertures and presentation, including pores and staminodes. Our samples had dramatically different ratios of tricolpate to tetracolpate pollen grains, a trait that does not exclude pollination by bees. Petal colour polymorphism occurs, with mixed colours proportionately less common in flowers with corollas that open. The proportion of species with staminodes did not differ between those presumed to be pollinated by bats and others, but anthers of the former were significantly more likely to have apical pores, and stamens all had similar length or were slightly longer in the middle, whereas stamens in two distinct groups occurred in 55% of bee-pollinated species. Pollen heteromorphy may facilitate pollination by different taxa in tropical environments. However, anther apical pores and stamen uniformity are more likely to be associated with bat-pollinated species than are other morphologies. Dillenia could be a useful model to examine evolutionary aspects of colour, heteranthery, staminodes and pollen heteromorphy. Only field work will verify bat pollination and the implications of bat dependence for Dillenia species.

Molecular Evidence of Leptospira spp. in Isolated Fijian Bats.

Bats host several zoonotic pathogens. Island biogeography and epidemiologic theory predict small remote islands have lower infection diversity. Molecular studies of urine and feces from three species at 10 sites from three islands suggest multiple pathogenic Leptospira, but not coronavirus, paramyxovirus, or Histoplasma, circulate in isolated Pacific Fijian bat populations.

A novel bat pollination system involving obligate flower corolla removal has implications for global Dillenia conservation.

The Dilleniaceae is known to produce nectarless flowers pollinated by bees, but the fact that bats ingest Dillenia biflora pollen led us to question pollination assumptions for these trees. We aimed to identify the pollinators of D. biflora, check for nectar presence, and investigate potential for cleistogamy and global prevalence of this pollination system. We examined aspects of the pollination of D. biflora on two Fijian islands using video recordings, direct observations, hand pollination, measurements (flowers, bite marks, nectar), and monitoring. The flowers, receptive for one night, contained copious nectar and had permanently closed globose corollas that required removal by bats for pollination. All the 101 flowers that retained their corolla died and did not produce seeds by cleistogamy. The bat Notopteris macdonaldi was well adapted to corolla removal. Keeping corollas closed until bats manipulate the nectar-rich flowers is a beneficial strategy in high-rainfall environments with many flower parasites. We propose to name a pollination system reliant exclusively on bats "chiropteropisteusis." From clues in the literature, other species in the geographical range of Dillenia are probably chiropteropisunous. Chiropteropisteusis should be investigated in the Old-World range of Dillenia, many species of which are threatened. The remarkable "fall" of the entire corolla observed by an earlier botanist for several species in the genus is most likely attributable to bats. This discovery has important implications for the conservation of bat-dependent trees and their associated fauna, particularly considering the high level of threat faced by flying-foxes globally.

Genome-wide transcriptome signatures of ant-farmed Squamellaria epiphytes reveal key functions in a unique symbiosis.

Farming of fungi by ants, termites, or beetles has led to ecologically successful societies fueled by industrial-scale food production. Another type of obligate insect agriculture in Fiji involves the symbiosis between the ant Philidris nagasau and epiphytes in the genus Squamellaria (Rubiaceae) that the ants fertilize, defend, harvest, and depend on for nesting. All farmed Squamellaria form tubers (domatia) with preformed entrance holes and complex cavity networks occupied by P. nagasau. The inner surface of the domatia consists of smooth-surfaced walls where the ants nest and rear their brood, and warty-surfaced walls where they fertilize their crop by defecation. Here, we use RNA sequencing to identify gene expression patterns associated with the smooth versus warty wall types. Since wall differentiation occurred in the most recent common ancestor of all farmed species of Squamellaria, our study also identifies genetic pathways co-opted following the emergence of agriculture. Warty-surfaced walls show many upregulated genes linked to auxin transport, root development, and nitrogen transport consistent with their root-like function; their defense-related genes are also upregulated, probably to protect these permeable areas from pathogen entry. In smooth-surfaced walls, genes functioning in suberin and wax biosynthesis are upregulated, contributing to the formation of an impermeable ant-nesting area in the domatium. This study throws light on a number of functional characteristics of plant farming by ants and illustrates the power of genomic studies of symbiosis.

Inter- and intra-archipelago dynamics of population structure and gene flow in a Polynesian bird.

Islands are separated by natural barriers that prevent gene flow between terrestrial populations and promote allopatric diversification. Birds in the South Pacific are an excellent model to explore the interplay between isolation and gene flow due to the region's numerous archipelagos and well-characterized avian communities. The wattled honeyeater complex (Foulehaio spp.) comprises three allopatric species that are widespread and common across Fiji, Tonga, Samoa, and Wallis and Futuna. Here, we explored patterns of diversification within and among these lineages using genomic and morphometric data. We found support for three clades of Foulehaio corresponding to three recognized species. Within F. carunculatus, population genetic analyses identified nine major lineages, most of which were composed of sub-lineages that aligned nearly perfectly to individual island populations. Despite genetic structure and great geographic distance between populations, we found low levels of gene flow between populations in adjacent archipelagos. Additionally, body size of F. carunculatus varied randomly with respect to evolutionary history (as Ernst Mayr predicted), but correlated negatively with island size, consistent with the island rule. Our findings support a hypothesis that widespread taxa can show population structure between immediately adjacent islands, and likely represent many independent lineages loosely connected by gene flow.

A test of island biogeographic theory applied to estimates of gene flow in a Fijian bird is largely consistent with neutral expectations.

Islands were key to the development of allopatric speciation theory because they are a natural laboratory of repeated barriers to gene flow caused by open water gaps. Despite their proclivity for promoting divergence, little empirical work has quantified the extent of gene flow among island populations. Following classic island biogeographic theory, two metrics of interest are relative island size and distance. Fiji presents an ideal system for studying these dynamics, with four main islands that form two large-small pairs. We sequenced thousands of ultraconserved elements (UCEs) of the Fiji bush-warbler Horornis ruficapilla, a passerine distributed on these four Fijian islands, and performed a demographic analysis to test hypotheses of the effects of island size and distance on rates of gene flow. Our demographic analysis inferred low levels of gene flow from each large island to its small counterpart and little or none in the opposite direction. The difference in the distance between these two island pairs manifested itself in lower levels of gene flow between more distant islands. Both findings are generally concordant with classic island biogeography. The amount of reduction in gene flow based on distance was consistent with predictions from island biogeographic equations, while the reduction from small to large islands was possibly greater than expected. These findings offer a hypothesis and framework to guide future study of interisland gene flow in archipelagos as the study of island biogeography progresses into the genomic era.

Linkages between measures of biodiversity and community resilience in Pacific Island agroforests.

Designing agroecosystems that are compatible with the conservation of biodiversity is a top conservation priority. However, the social variables that drive native biodiversity conservation in these systems are poorly understood. We devised a new approach to identify social-ecological linkages that affect conservation outcomes in agroecosystems and in social-ecological systems more broadly. We focused on coastal agroforests in Fiji, which, like agroforests across other small Pacific Islands, are critical to food security, contain much of the country's remaining lowland forests, and have rapidly declining levels of native biodiversity. We tested the relationships among social variables and native tree species richness in agroforests with structural equation models. The models were built with data from ecological and social surveys in 100 agroforests and associated households. The agroforests hosted 95 native tree species of which almost one-third were endemic. Fifty-eight percent of farms had at least one species considered threatened at the national or international level. The best-fit structural equation model (R2 = 47.8%) showed that social variables important for community resilience-local ecological knowledge, social network connectivity, and livelihood diversity-had direct and indirect positive effects on native tree species richness. Cash-crop intensification, a driver of biodiversity loss elsewhere, did not negatively affect native tree richness within parcels. Joining efforts to build community resilience, specifically by increasing livelihood diversity, local ecological knowledge, and social network connectivity, may help conservation agencies conserve the rapidly declining biodiversity in the region.

Response of primary and secondary rainforest flowers and fruits to a cyclone, and implications for plant-servicing bats.

The response of primary (PF) and secondary (SF) rainforests to cyclones has broad implications for servicing fauna and the resilience of forest functions. We collected fine-scale data on the reproductive phenology of plant communities in Fijian PF and SF in 12 monthly surveys before and after Cyclone Tomas (2010). We generated a resource index from the reproductive loads of 2218 trees and 1150 non-trees (>190 species) and trunk and stem diameter to assess patterns in resource abundance for nectarivores and frugivores (hereafter NF resources). We aimed to determine (i) whether species richness of NF resources differed between forests; (ii) the patterns of resilience of NF resources at community level in both forests after a cyclone; and (iii) the effect of response on NF resources for plant-servicing bats (Pteropodidae). In 12 months preceding the cyclone, NF resources were greater in PF trees; non-tree resources fluctuated and were greater in SF. Lower species richness of NF resources in SF indicated that fewer opportunities exist there for exploitation by a diverse fauna. More resources were available for bats in PF. In 12 months following the cyclone, PF flowers and fruits, and SF fruits specifically used by pteropodid bats decreased for trees. Non-tree resources were especially susceptible to the cyclone. No universal pattern of decline was associated with the cyclone; instead, some NF resources declined and others were resilient or responded rapidly to a post-cyclone environment. Both PF and SF demonstrated resilience at the community level via increased flower survival (PF) and rapid flower production (SF). Reduced species richness of NF resources in SF will compromise future resilience and response to disturbance, including for threatened pteropodid bat species. These findings are critical for long-term management of forests, given predicted increases in cyclone frequency and intensity associated with anthropogenic climate change.

A molecular phylogeny of Pacific honeyeaters (Aves: Meliphagidae) reveals extensive paraphyly and an isolated Polynesian radiation.

We investigated the molecular phylogenetic placement of 14 species of Pacific island honeyeaters (Aves: Meliphagidae) in the broader context of an existing family-level phylogeny. We examined the evolutionary history of Pacific honeyeater lineages to assess the accuracy of current taxonomies and to evaluate their biogeographic history. We compare these biogeographic patterns to other Pacific birds to identify emergent patterns across lineages. We found strong support for a previously unknown endemic radiation in central Polynesia, which comprises five genera: Meliarchus, Guadalcanaria, Gymnomyza, Xanthotis, and Foulehaio. Conversely, other Pacific lineages were found to be strongly allied with continental radiations (e.g., Philemon eichhorni, P. cockerelli, and Lichmera incana). Our results necessitated taxonomic changes, both at the generic level (e.g., Xanthotis, Melidectes/Vosea, and Glycifohia/Gliciphila) and regarding species limits within polytypic species. Here, we discuss species limits in Foulehaio and Gymnomyza and recommend elevating three nominal subspecies of Foulehaio to species status, each of which forms well-differentiated clades.