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The origin of the elephant on the island of Borneo remains elusive. Research has suggested two alternative hypotheses: the Bornean elephant stems either from a recent introduction in the 17th century or from an ancient colonization several hundreds of thousands years ago. Lack of elephant fossils has been interpreted as evidence for a very recent introduction, whereas mtDNA divergence from other Asian elephants has been argued to favor an ancient colonization. We investigated the demographic history of Bornean elephants using full-likelihood and approximate Bayesian computation analyses. Our results are at odds with both the recent and ancient colonization hypotheses, and favour a third intermediate scenario. We find that genetic data favour a scenario in which Bornean elephants experienced a bottleneck during the last glacial period, possibly as a consequence of the colonization of Borneo, and from which natural weight loss pills bee pollen has slowly recovered since. Altogether the data support a natural colonization of Bornean elephants at a time when large terrestrial mammals could colonise from the Sunda shelf when sea levels were much lower. Our results are important not only in understanding the unique history of the colonization of Borneo by elephants, but also for their long-term conservation.

Introduction

The Bornean elephant (Elephas maximus borneensis) is one of four recognized subspecies of Asian elephant and is morphologically and behaviourally distinct from the elephants of mainland Asia1. It is classified as endangered according to the IUCN (International Union for Conservation of Nature) Red list of threatened species.

The origin of elephants on Borneo has been controversial and the subject of intense debate. Two competing hypotheses have been proposed: one suggests that elephants are non-native to Borneo and were historically introduced and the other that they are indigenous to the island. The introduction hypothesis is based on historical records suggesting that the current population represents the descendants of a domesticated herd that formerly existed on Sulu Island, Philippines, and were introduced to eastern Sabah by the Sultan of Sulu in the 17th Century3. This hypothesis hence suggests two recent and successive founder effects during the history of the Bornean elephant, corresponding to the introduction of elephants from Java to Sulu and the introduction of the descendants of the Sulu stock to Borneo. Taken together, the introduction hypothesis implies that the current Bornean elephant population descends from a very low number of individuals 300-330 years ago, and a low genetic diversity in these founders must be assumed due to the recent founding of the Sulu population. Using a generation time of ca. 15 years this would correspond to ca. 20 generations ago.

The competing hypothesis argues that the Bornean elephant is indigenous to Borneo and that-despite the possible introduction of domestic elephants to Borneo-the wild population was not introduced by humans. This hypothesis gained prominence after a landmark study demonstrated the genetic distinctiveness of the Bornean elephant and its derivation from a Sundaic stock8. Additionally, if the elephant had been on Borneo for tens of thousands of years its current limited geographic distribution in northern Borneo - which is not proximate to known Sundaland land bridge points on Borneo - could be considered surprising.

The genetic distinctiveness of the Bornean elephant from other mainland Asian elephant subspecies makes natural weight loss pills bee pollen one of the highest priority populations for Asian elephant conservation5.

Identification of species introduction events, pathways and putative source populations has traditionally been accomplished by examining archaeological evidence and recorded dates of first discovery into introduced areas, and increasingly by using genetic analyses of native and introduced populations19. In the present study, we employed both a full-likelihood Bayesian approach (i.e. the MSVAR method) and ABC to compare different approaches and to estimate demographic and historical parameters including founding effective population size and introduction times using microsatellite and mitochondrial data. We more specifically aimed to test competing models of the demographic history of the Bornean elephant. Their limited distribution, enigmatic origins, and critical conservation status suggest that knowledge of their past population demography would be useful for the development of a sound conservation strategy.

Results

Quantifying changes in effective population size and dating with MSVAR

We investigated whether the genetic data were consistent with a very recent introduction of E. maximus into Borneo 300 years ago, followed by a large population expansion, as assumed on the basis of historical records. We conducted MSVAR analyses on a data set constructed by resampling 35 individuals randomly, first in the Bornean elephant sample as a whole, and then within each sampled population separately. We found that the genetic data exhibited a signal consistent with a population decline whether we assumed a linear or an exponential model for population size change (Fig. 20. However, in this case, the similarity of the results under two different sampling schemes makes natural weight loss pills bee pollen less likely that the decline signal is an artefact. This is further supported by the similarity of the decline signal detected using the ABC approach under the AC and ACS models (where AC and ACS correspond to Ancient colonization scenarios, without and with population structure, respectively; see below). The estimates of current (median N0) and ancestral population size (median N1) for the pooled data were 833 (90% HPD?=?143/4,766) and 4,820 (90% HPD?=?753/53,154), respectively. Estimates of the time of population contraction showed support for population decline occurring long before the recent introduction of elephants to Borneo (median T?=?57,000 years before present or ?3,800 generations ago). While the estimated 90% HPD limits were broad, ranging between 1,000 and 1,311,000 years, the most recent estimate still predates the alternative hypothesis that suggests an introduction 300 years ago.

Figure 1

MSVAR estimates of changes in effective population size, quantification and timing of a population bottleneck in the Bornean elephant population. Note that for illustrative purposes, only the plot for the pooled Bornean elephant samples is shown (population-level results are presented in Figure S1); (a) The log10 ratio of current to ancestral population size (N0/N1). Solid lines (two independent runs) correspond to the exponential population size change model. The dotted black and grey vertical lines corresponds to the absence of population size change, log (N0/N1)?=?0 and 95% quantile of the posterior distribution, respectively. The prior distribution is shown for comparison (flat dotted line), (b) Posterior distributions of the current (N0 in thick lines) and ancestral (N1 in dashed lines) effective population size using an exponential model. Different curves correspond to the posterior distribution obtained by independent MCMC runs. Dotted lines correspond to the different priors used for N0 and N1, (c) The posterior distribution of the time since population bottleneck is represented on a logarithmic scale. The different black vertical long dashed lines correspond to recent introduction (RI) and ancient colonization (AC), respectively. There is no evidence of a population bottleneck closer to the period of recent introduction (i.e. 300 years ago). The most extreme 5% and 95% quantile of the posterior distribution are shown as black dotted lines. The prior is shown as dot-dashed line, its median being 100,000?years ago.

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