Multi-scale habitat selection modeling identifies threats and conservation opportunities for the Sunda clouded leopard (Neofelis diardi)
Introduction
Accelerating rates of deforestation, fire, and land conversion (e.g., to large-scale oil palm plantations and Acacia monocultures), have rapidly transformed the natural landscapes of Southeast Asia, with dire consequences for native fauna (Tacconi, 2003; Cushman et al., 2017). Clouded leopards are among Asia's most broadly ranging and charismatic felids, but also among its rarest, least understood, and most vulnerable (Ross et al., 2013; Hearn et al., 2016). Clouded leopards occur throughout Southeast Asia, from the Himalayan foothills in the west to southern China in the north and east, and extend south, in the form of the Sunda species, into the islands of Borneo and Sumatra. In 2006, genetic and morphological analyses led to splitting the species, with Neofelis nebulosa occupying mainland Southeast Asia, and Sunda clouded leopards (N. diardi) occurring on Borneo and Sumatra (Buckley-Beason et al., 2006; Kitchener et al., 2006; Wilting et al., 2007). The elusive nature of these two species poses a research challenge, and there remains sparse information on the ecologies of either species.
Both the mainland and the Sunda clouded leopard occur across broad geographical ranges, prompting questions of how their habitat uses, population densities, and intra-guild interactions vary between regions. Considering that clouded leopards occur in the most rapidly disappearing forests in the world (Miettinen et al., 2011; Gaveau et al., 2016), understanding the habitat preferences of these threatened felids is critical for mitigating their likely deteriorating conservation status. Clouded leopards are listed as Vulnerable on the IUCN Red List of Threatened Species in accordance with their high extinction risk (Hearn et al., 2015; Grassman et al., 2016; Hearn et al., 2016). While at least mainland clouded leopards are threatened by direct exploitation of pelts and bones for medicine (D'Cruze and Macdonald, 2015; Min et al., 2018), both species face grave indirect threats from habitat loss and poaching of their prey. In particular, very high rates of deforestation across their range drive loss, fragmentation and degradation of habitat quality (Cushman et al., 2017; Macdonald et al., in press). In this paper we aim to identify the primary habitat requirements as well as limiting factors and sources of threat for the Sunda clouded leopard across its range. Furthermore, the iconic charisma of these cats combined with their forest-dwelling habits make them ambassador species (Macdonald et al., 2017) whose conservation provides an influential co-benefit to broader forest conservation programmes (Collins et al., 2011).
There have been several local estimates of the abundance of Sunda clouded leopards based on camera trapping (Wilting et al., 2012; Cheyne et al., 2013; Sollmann et al., 2014), and we recently published the first robust estimate of their population size for the Malaysian state of Sabah (Hearn et al., 2017). Low detection rates observed in a recent survey of Indonesian Borneo (Cheyne et al., 2016) suggest N. diardi may be less abundant in south and eastern Kalimantan compared to Sabah in the north. Additionally, Hearn et al. (2018a) conducted a multi-scale path level analysis of N. diardi movement in an anthropogenically disturbed landscape in Sabah, which showed that clouded leopard movements are closely associated with forest habitats with closed canopies and resisted by plantation habitats with open canopies. Yet a comprehensive, range-wide assessment is lacking.
To address this need, we developed a multi-scale habitat model that encompasses the full range of N. diardi across Borneo and Sumatra. Multi-scale habitat modeling is critical to correctly identify the drivers and meaningful scales of relationship with patterns of species distribution (Levin, 1992; Thompson and McGarigal, 2002; Grand and Mello, 2004), yet a recent review showed that <5% of all recent habitat modeling papers used robust approaches that optimize multivariate scale relationships (McGarigal et al., 2016). Given N. diardi's relatively large home range estimates (Hearn et al., 2013, Hearn et al., 2018a), we expected to find that clouded leopards are utilizing key habitat components (e.g., closed canopy forest) at the broadest scales included in our study (8–10 km). The primary goals of this paper were therefore to (1) identify key environmental and anthropogenic variables contributing to Sunda clouded leopard habitat use, (2) determine the spatial scale at which each variable most strongly influences clouded leopard detection, and (3) investigate differences between clouded leopard presence and habitat associations on Sumatra and Borneo. These findings provide critical information to assist in prioritizing habitat conservation networks for the protection of this vulnerable felid, and more broadly demonstrate the value of multi-scale modeling for characterizing the requirements of protected areas.
Section snippets
Data collection
From 2007 to 2016, our field teams conducted intensive and systematic camera trap surveys of N. diardi in 22 sampling locations across the islands of Borneo and Sumatra (Fig. 1, Tables 1 & S1). Sampling locations include sites previously selected for intensive autecological research focused on clouded leopards and their associated guild. Additional sites were chosen to encompass a representative sample of the spectrum of forest types, elevations, and anthropogenic disturbances present
Results
We obtained a combined sampling effort of 138,516 trap nights from 1544 camera stations, with number of trap nights per station averaging 89.7 ± 1.17 SE (Table 1). Records of N. diardi were obtained from 17 of the 22 sampling locations, and at 283 (or 18.3%) of the camera stations, with station-level detection rates of 13/317 (or 4.1%) in Kalimantan, 169/484 (or 34.9%) in Sabah and 101/743 (or 13.6%) in Sumatra. Average detection rates per camera station varied across regions, with 0.04 (±0.02)
Discussion
Megafauna throughout Southeast Asia are threatened by rapid deforestation and land conversion. The strong association between clouded leopards and extensive regions of high forest cover makes them potent ambassadors for conserving broader forest biodiversity (Macdonald et al., 2017). Yet clouded leopards are enigmatic, their habitat use poorly known, and therefore the opportunities to plan their conservation and utilize their potential as umbrella and ambassador species have been limited. Here,
Acknowledgments
The data reported here were gathered by three field teams, each supported under arduous conditions by dedicated local staff. Without their efforts in Sabah, Kalimantan and Sumatra, this work would have been impossible. This study is part of a larger integrated range-wide study of both Sunda and mainland clouded leopards supported by a grant to [confidential for review] from the Robertson Foundation.
Author contributions
[Confidential for double blind review.]
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