Understorey to canopy vertebrate fauna of a lowland evergreen forest in Mt. Makiling Forest Reserve, Philippines.

We examined the vertical stratification of forest wildlife, from the ground up to the canopy layer, within a 2-hectare permanent plot of lowland evergreen rainforest on the Mt. Makiling Forest Reserve. Our aim was to determine the species richness of the different forest layers and evaluate their ecosystem services. Understorey, sub-canopy and canopy sampling were conducted during July 2016, March to April 2017 and February to March 2018, respectively. We were able to record a total of 68 species, consisting of 11 amphibians, 15 reptiles, 25 birds and 17 mammals. Increasing species richness with increasing vertical stratification was observed for both reptiles and mammals. For birds, the peak richness was observed in the sub-canopy and then decreased in the canopy. A decreasing trend was observed with amphibians wherein the peak species richness was observed in the understorey. Increasing vertical stratification influenced vertical habitat use and species richness. For the similarity index, the same pattern was observed for all species groups. Highest similarity was observed between the sub-canopy and the canopy and the least similarity was observed between the understorey and canopy. These results indicate that the understorey and the canopy host different species groups, thus, sampling of the understorey alone, often done in biodiversity surveys, may lead to the underestimation of species richness in an area.

Description and comparison of Philippine hornbill (Bucerotidae) vocalizations.

The role of vocalisation for the Philippine hornbills' ecology and speciation and their implication in understanding speciation is not well understood. We described and compared recorded calls of seven hornbill taxa in captivity namely Mindanao Wrinkled hornbill (Rhabdotorrhinus leucocephalus), Rufous-headed hornbill (Rhabdotorrhinus waldeni), Luzon Rufous hornbill (Buceros hydrocorax hydrocorax), Samar Rufous hornbill (Buceros hydrocorax semigaleatus), Mindanao Rufous hornbill (Buceros hydrocorax mindanensis), Mindanao Tarictic hornbill (Penelopides affinis), Samar Tarictic hornbill (Penelopides samarensis), Visayan Tarictic hornbill (Penelopides panini) and Luzon Tarictic hornbill (Penelopides manillae), as well as comparison with the non-native Papuan hornbill (Rhyticeros plicatus). Vocalisation analysis included call duration, minimum frequency, maximum frequency, bandwidth and peak frequency. For each species in the sample, the mean and standard deviation were used to calculate the Cohen's d statistic by using an effect size calculator. Results showed that the effect size for minimum frequency was small for P. panini vs. P. samarensis and B. hydrocorax vs. B. h. mindanensis. However, bandwidth, duration, minimum frequency, maximum frequency and peak frequency have large effect sizes for the rest of the allopatric species pairs. Hornbills' conspicuous resonating calls are sufficiently quantifiable for bioacoustic analysis and may provide new insights for their taxonomic review.

Distribution and diversity patterns of herpetofauna in the Pantabangan-Carranglan Watershed, Nueva Ecija, Caraballo Mountain Range, Philippines.

The recent extensive survey conducted in the Pantabangan-Carranglan Watershed, located in the Caraballo Mountain Range during the rainy season (October to November) resulted in the recording of fifty-nine (59) species of amphibians and reptiles (17 frogs, 14 skinks, 3 agamids, 6 gekkonid lizards, 2 varanids and 17 snakes). Out of 59 species, 42 species were Philippine endemics and 25 species are recorded only from the Luzon faunal region. Habitat analysis and mapping showed seven habitat types including lowland dipterocarp forest, grassland, lower montane forest, upper montane forest, pine forest, agricultural areas and riparian habitats. GIS-based distribution mapping showed that the number of individuals and species are high in forest and associated riparian habitats at mid-elevation (1,000-1,250 m a.s.l.). The distribution pattern in the area is influenced by similarity in microclimatic conditions, availability of resources and niches which species can utilise. Species diversity is centred in mid- to high elevation forest and riparian habitats and in less disturbed areas. Snake diversity is adversely affected by increased disturbance, making them good indicators of the health of an area. The abundance-based Jaccard's Similarity Index showed that connected habitats and elevation gradients have higher species similarity.

Environmental stability and the evolution of cooperative breeding in hornbills.

Reproductive cooperation in social animals has been the focus of intensive research, yet the role of environmental factors in promoting such cooperation remains uncertain. A recent global analysis suggested that cooperative breeding in birds is a 'bet-hedging' strategy associated with climatic uncertainty, but it is unclear whether this mechanism applies generally or is restricted to the insectivorous passerines that predominate as cooperative breeders at the global scale. Here, we use a phylogenetic framework to assess the effect of climate on the evolution of cooperation in hornbills (Bucerotidae), an avian family characterized by frugivory and carnivory. We show that, in contrast to the global pattern, cooperative reproduction is positively associated with both inter- and intra-annual climatic stability. This reversed relationship implies that hornbills are relatively insensitive to climatic fluctuations, perhaps because of their dietary niche or increased body mass, both of which may remove the need for bet-hedging. We conclude that the relationship between climatic variability and cooperative breeding is inconsistent across taxa, and potentially mediated by life-history variation. These findings help to explain the mixed results of previous studies and highlight the likely shortcomings of global datasets inherently biased towards particular categories.

A comprehensive molecular phylogeny for the hornbills (Aves: Bucerotidae).

The hornbills comprise a group of morphologically and behaviorally distinct Palaeotropical bird species that feature prominently in studies of ecology and conservation biology. Although the monophyly of hornbills is well established, previous phylogenetic hypotheses were based solely on mtDNA and limited sampling of species diversity. We used parsimony, maximum likelihood and Bayesian methods to reconstruct relationships among all 61 extant hornbill species, based on nuclear and mtDNA gene sequences extracted largely from historical samples. The resulting phylogenetic trees closely match vocal variation across the family but conflict with current taxonomic treatments. In particular, they highlight a new arrangement for the six major clades of hornbills and reveal that three groups traditionally treated as genera (Tockus, Aceros, Penelopides) are non-monophyletic. In addition, two other genera (Anthracoceros, Ocyceros) were non-monophyletic in the mtDNA gene tree. Our findings resolve some longstanding problems in hornbill systematics, including the placement of 'Penelopides exharatus' (embedded in Aceros) and 'Tockus hartlaubi' (sister to Tropicranus albocristatus). We also confirm that an Asiatic lineage (Berenicornis) is sister to a trio of Afrotropical genera (Tropicranus [including 'Tockus hartlaubi'], Ceratogymna, Bycanistes). We present a summary phylogeny as a robust basis for further studies of hornbill ecology, evolution and historical biogeography.