Orangutan ranging in Kutai National Park by The Leakey Foundation

Leakey Foundation Final Grant Report (grant May 12-Dec 13) Orangutan ranging in Kutai National Park

Anne E. Russon Psychology Dept., Glendon College of York University Toronto M4N 3M6, CANADA e-mail: arusson@gl.yorku.ca

Putri, our resident orangutan female, with her new baby © Purwo Kuncoro

Overview This project aims to study orangutan ranging in Kutai National Park (KNP), E. Kalimantan. Recent inter-site comparisons show east Bornean orangutans, Pongo pygmaeus morio, as the extreme of orangutan adaptation (Wich et al., 2009). They face the harshest environment and have correspondingly distinct morphology, genetics, behavior, and life history. Most morio-KNP data date from 30-35 years ago, however. They are not of the quality needed to verify these distinctions, suggest other differences, and represent ecological conditions that may no longer exist because natural disasters and development have since damaged and fragmented much of morio’s KNP habitat. New studies are needed, and ranging flexibility is a good place to start. I chose ranging for the good overview it offers of orangutans’ main activities (feed, travel, rest, socialize) and two of its main determinants: feeding ecology, which distinguishes morio, and cognition, which should guide orangutans’ ranging responses to spatio-temporal fluctuations in food availability. The research project aims to reassess morio KNP ranging flexibility by assessing long-term ranging patterns from standard and cognitive perspectives. Study is necessarily longterm because supra-annual climate cycles affect resource availability, which in turn affects orangutan behavior. Goals include assessing variation in ranging and diet with supra-annual cycles (e.g., home range size) and with increasing habituation (e.g., terrestriality), and more sophisticated cognitive studies (spatial and spatio-temporal cognition). To date, we have also broadened the project’s scope through collaborative studies that may help clarify our morio community structure (genetics) and reproduction (nutrition) and hosting compatible studies (e.g., forestry/botany, camera trap survey of KNP wildlife, phenology), student theses (spatial cognition), and application of our findings to morio conservation and management programs in and around Kutai National Park. We established a new research site along the Sangatta River, KNP’s northern border (Figure 1, below) where we have been collecting field data on this project since early January, 2010. We have also been collecting data for collaborative studies of orangutan genetics (PI: Dr. G. Banes) and reproduction (ecology, energetics, life history, and endocrine physiology; PI: Dr. C. Knott). In 2013 we purchased a small research drone and pilot tested its use for monitoring and assessing habitat condition and phenology. Four York University students contributed to data collection and drone pilot testing through internships and theses (A. Bebko, S. Iannicello, B. Jevons, K. Maklan). To date, our research activities have focused on activity budgets and travel, as basic adaptive behavior with implications for energy management, and studies of spatial cognition. Use of Leakey 2012-13 grant funds This Leakey Foundation grant was for $3000 USD, funded personally by Gordon Getty, for the period May/12 – Dec/13. In line with my proposal, I used the grant primarily for our field assistants’ salaries and benefits. This usage is the most effective; it is also in line with this project’s design, long-term study of orangutan ranging and its correlates. Expense item Salaries and benefits Materials and supplies Total

Total cost $ 2,882.86 117.14 $ 3,000.00

I have attached the final financial report that York University’s Research Accounting Department prepared; I believe it was sent directly to the Leakey Foundation when the grant period terminated.

Methods: Study area, data collection. Study area. Figure 1 show Bendili, our orangutan study area. Bendili overlaps with the well known Mentoko orangutan study area, established by Rodman in 1970-71 and extended to the west by Leighton in 1982, and the Mentoko-Sinara orangutan study area, used by Suzuki from 1984 to 88 and which extends the Mentoko study area to include Bukit Sinara (Sinara Hill) to the east. Figure 2 shows Bendili’s location relative to these older study areas.

Figure 1. Bendili study area (above), and Bendili’s location within KNP and relative to other orangutan study sites (below)

Figure 2. Bendili relative to Mentoko and Sinara Data collection. Behavioral and phenological data are collected in standardized fashion, i.e., observation of behavior during full-day focal individual follows, monthly phenology monitoring of trees in permanent vegetation plots and fruit trails. In addition to standard behavioral observations, we record the focal orangutan’s location at 15 minute intervals, as a basis for spatial analyses. Grant period activities Activities during this Leakey grant granting period included (a) regular long-term field data collection (standard orangutan behavior and ranging, monthly phenological monitoring in permanent tree plots and fruit trails), (b) training and pilot testing use of a research drone for habitat monitoring, (c) computerizing behavior, GPS, and phonological data for 2010-2012, (d) analysis and writeup of our research findings on orangutan activity budgets and spatial cognition, (d) reporting/presenting research findings (conferences, publications). Main Findings Activity Budgets and Ranging We have completed activity budget analyses for three consecutive years (2010-2012), calculated for both intra-site (change over time) and inter-site comparisons (geographic variation related to taxonomic and ecological factors). Inter-site comparisons (i.e., geographical variation), are based on yearly activity budget estimates for each age-sex class; intrasite comparisons are based on monthly estimates for each age-sex class (Morrogh-Bernard et al., 2009). Intra-site comparisons. Kutai National Park’s orangutans have been studied in the Mentoko-Bendili area several times over the last 44 years, starting in 1970 (Rodman, 1973; Leighton, 1982, 1993; Mitani, 1989; Suzuki, 1984, 1986, 1988, 1989; Campbell, 1992). Over this period, the habitat has undergone profound changes. Rodman (1973) described Mentoko habitat as “near pristine” in 1970-71. Later in the 1970s, commercial logging made incursions. In 1982-83 an exceptionally severe drought followed by great forest fires damaged much of Kutai National Park, including Mentoko and adjacent areas to the east (Leighton & Wirawan, 1986). In 1997-98, a second severe drought and massive forest fires again damaged much of the area. Small scale poaching (hunting, illegal logging) followed. By the time our study began, January 2010, the area had been recovering from the 1997-98 forest fires for almost 12 years. Accordingly, the data we have analyzed to date offers a picture of how these orangutans have adjusted to Mentoko area habitat that has had 12-14 years to recover from the most recent forest fires.

Main findings on how Mentoko-Bendili area orangutans coped with the series of changes to their habitat since 1970 are shown in Figure 3, in terms of their activity budgets, i.e., the percentage of daily active time devoted to each main activity (feed, rest, travel). Habitat conditions are nearpristine (Rodman 05/1970-07/1971, Mitani 08/1981-01/1983), recent drought-fire damage (2-3 and 3-4 yrs post 1982-83 damage—Suzuki, 1985-86; Campbell, 1986-87), and longer term recovery from drought and fire damage (12-14 yrs post 1997-98 damage, this study, 2010-12).

Rodman 70-‐71 60

Suzuki 85-‐86

Campbell 86-‐87 Russon 10-‐12

40 30 20

Russon 10-‐12 Campbell 86-‐87

Suzuki 85-‐86

0 Feed

researcher Rodman Suzuki Campbell Russon

Rodman 70-‐71 Rest

study area Mentoko Mentoko-Sinara Mentoko Bendili

Travel

period 1970-71 1985-86 1986-87 2010-12

feed 45.9 52.3 32.5 44.2

rest 39.2 37.3 53.3 33.0

travel 11.1 9.2 9.5 18.8

Figure 3. Activity budgets for Mentoko-Bendili with changing habitat conditions. Values represent the percentage of daily active time in each activity (exact figures below the chart). Figure 3 suggests orangutan behavioral changes in response to habitat change: (a) rapid adjustment to the 82-83 damage and short-term recovery—initially (Suzuki) increasing feeding reducing resting, later (Campbell) reducing feeding and increasing resting, (b) longer term tracking habitat recovery so that 10-14 yrs after the most recent damage (1997-98), activity budgets have become very close to those reported in near pristine conditions, except that travel may be slightly higher and offset by slightly lower resting. There is a good possibility that Rodman underestimated travel: much local orangutan is E-W along the Sangatta River and Rodman’s Mentoko site was very narrow east to west, and Rodman’s data were heavily biased to months when travel distances were relatively low. Overall, our findings suggest there may be an ‘optimal’ activity budget for orangutans ranging in this area, possibly as a function of longer term habitat features (e.g., topography, water availability) and the area’s orangutans respond to change immediately and then, with sufficient time, adjust their behavior in this direction. This is relatively consistent with findings that morio and some other primates in North Borneo recover well from selective logging damage after long-term recovery (Berry et al., 2010; Edwards et al., 2010; Johns, 1992). Inter-site comparisons. Comparisons are shown in Figure 4. Most estimates are from Morrogh-Bernard et al. (2009). We added several that have appeared since then: Ketambe, Sumatra (primary vs. recently logged forest: Hardus at al., 2012), Sabangau, Central Kalimantan

(Harrison et al., 2009, Danum Valley, Sabah (Kanamori et al., 2012), and our own new estimates.

% daily ac)ve )me

a) Sumatra: P. abelii 80 SB

KET

KET-‐ P

20 0

b) CW Borneo: P. p. wurmbii

c) E Borneo: P. p. morio

% daily ac)ve )me

60 40 20 0

SAB-‐ 3 SAB-‐ 6 TUA

% daily ac)ve )me

60 40 20 0

KIN US DV-‐6 KNP-‐ MM

Figure 3. Activity Budgets – intersite comparisons. Legend codes: MB-Morrogh-Bernard et al. (2009). SB – Suaq Balimbing; KET/KET-P/KET-L – Ketambe-MB/primary/logged (Hardus et al., 2012); SAB/SAB-6 – Sabangau-MB/Harrison et al. (2009) based on min 6-hr follows; TUA – Tuanan; TP – Tanjung Puting; GP – Gunung Palung; US – Ulu Segama; KIN – Lower Kinabatangan, DV-6 – Danum Valley (Kanamori et al. 2012) based on min 6-hr follows; KNPMM/KNP-B – Kutai NP Mentoko-MB/Bendili Morrogh-Bernard et al. (2009) interpreted these findings as indicating activity budget differences between masting and non-masting (peat swamp) forests. Masting forests are represented by Gunung Palung (P. p. wurmbii) and all P. p. morio sites (Sabah and E Kalimantan: Lower Kinabatangan, Danum Valley, Mentoko-Bendili). Our 2010-12 findings for Bendili and the newer findings we incorporated into the comparison are largely consistent with this pattern. We also compared day travel distances because our 2010-12 activity budgets suggest P. p. morio in the Bendili-Mentoko area devotes more time daily to travel than it did in the past. On the basis of Rodman’s estimates for Mentoko and estimates for sites in Sabah, Singleton et al. (2009) suggested that morio travels less than other orangutan taxa. Table 1 compares day travel distance for adult females. For Mentoko-Bendili area morio, our 2010-12 day travel estimates are considerably higher than earlier ones and within one standard deviation of the G. Palung (P. p. wurmbii) estimate. Methodology and change over time both probably account for differences between older and new values for Mentoko-Bendili morio: early studies probably underestimated their day travel due to sampling bias but they may travel farther now due to habitat changes. Implications are that morio may differ less from wurmbii than Singleton et al. (2009) suggested.

Table 1. Day travel distances, intersite comparisons. Site

island

S. Balimbing Ketambe –old Sabangau Sabangau 5 Tuanan T. Puting G. Palung Kinabatangan KNP: Mentoko1 KNP: Mentoko-Sinara2 KNP: Mentoko3 KNP: Bendili-2010-124

S S B B B B B B B B B B

subspecies abelii abelii wurmbii wumbii wurmbii wurmbii wurmbii morio morio morio morio morio

damage U U L L L U U L, P U DF DF R

SAF mean 1077 722 769 1058 1025 711 690 162 [305]

std 368 293 -365 471 -----

[379] 551

265 145

mothers mean 833 675 809 858 766 ---[305] 373 [379] 504

std 306 282 -246 355 ----220

Table notes Damage: L-logged, U-unlogged, P-plantation, DF-drought/fire damage, R-recovering from drought/fire damage. SAF: sexually active females, females mating and travelling with males when observed). Mothers: females with dependent offspring, not yet mating when observed (Singleton et al., 2009). Sources: 1-Rodman 1977; 2-Suzuki 1988; 3-Campbell 1992, 4-this study, 5-Harrison et al. 2009. All other values are comparative inter-site estimates from Singleton et al. (2009). Spatial cognition Our current spatial cognition studies aim to establish links between orangutan behavior and their spatial movements and location. Orangutan travel routes. Primates’ small-scale travel is increasingly used to assess their spatial cognition, due to recent improvements in accuracy and the commercial availability of Global Positioning System (GPS) technology (Janson & Byrne, 2007). Many now study individual primates’ daily movement paths (travel routes) as a basis for assessing spatial cognitive abilities such as “mental maps” (how individuals store spatial information), navigation, and planning. Several primate species show travel route networks that are used repeatedly over long periods of time (hereafter, habitual route networks) (DiFiore & Suarez, 2007; Hopkins, 2010; Noser & Byrne, 2010; Porter & Garber, 2012). In these networks, several repeatedly used travel routes interconnect at intersections, called nodes (Di Fiore & Suarez, 2007). In species with overlapping individual home ranges, several individuals may share the same route network. Orangutans are primarily arboreal and rely heavily on key fruit species that are patchily distributed (van Schaik, Marshall, & Wich, 2009), so they may develop and use habitual route networks. We recently confirmed the use of a habitual travel route network this by orangutans in our Bendili study area (Figure 4). Analytical work was part of an MA thesis by Adam Bebko, one of my graduate students. Findings are based on a new GPS-based computer mapping method that he developed, which accounts for GPS error by drawing a buffer around every travel route of all orangutans observed to represent uncertainty in the GPS locations. These error-buffers are overlapped one on top of another to determine how many routes overlap in a particular area.

Figure 4: Habitual travel route network identified for wild orangutans in Bendili, Kutai National Park. Findings are based on data for 26 orangutans, Jan 2010 to Jul 2011. What intersections represent. An important question is what intersections (nodes) in route networks represent. They are more often visited and/or passed than other areas in the network, so they presumably represent important areas for users. Studies attempting to assess why nodes are so heavily visited have found there are often important resources at notes (e.g., large fruit trees of key food species, water holes) (Byrne et al., 2009; Porter & Garber, 2012) but have not tested whether the resource-node association is greater than would be expected at a randomly selected location where individuals travelled. For example, individuals may frequently travel past the resources at nodes but may feed on these same resources at other locations; alternatively these resources may be extremely plentiful and occur throughout the route network including non-node locations. Adam Bebko examined the spatial distribution feeding bouts relative to nodes in Bendili orangutans’ habitual route network. He assessed all observed feeding bouts of four key fruit food species (Dracontomelon dao, Castanopsis sp., Diospyros sp., Ficus obpyramidata) to determine whether they are more strongly associated with node locations than randomly selected points where the individuals were not feeding on these four species. He found that indeed, feeding bouts for three of these four species are more likely to occur at nodes than randomly selected points. Figure 5 illustrates this finding for sengkuang (Dracontomelon dao).

9 Â Â

Figure 5. Dracontomelon dao (sengkuang) feeding bouts and network nodes.

10 Â Â

Accomplishments 05/12-12/13 Conference Presentations Bebko, A.O. & Russon, A.E. (2012). New mapping methods to evaluate route-based travel networks and planning in wild orangutans (Pongo pygmaeus morio). International Primatological Congress, Cancun, Mexico, Aug. 12-17. Russon, A.E. & Kuncoro, P. (2012). Orangutans in Kutai National Park: Behavior in recovering habitat. International Primatological Congress, Cancun, Mexico, Aug. 12-17. Russon, A. E., Kuncoro, P. & Sukrismanto, E. (2014). Orangutans in Kutai National Park: An island of forest in a sea of industry. Invited symposium contribution, From forest to plantation: How orangutans adapt to human-induced changes (organized by S. Spehar, G. Campbell-Smith & M. Ancrenaz). Congress of the International Primatological Society, Hanoi, Vietnam, Aug.11-16. Other Presentations Russon A, E. & Kuncoro, P. (2013). Orangutans in Kutai National Park: Behavior in Recovering Natural Habitat. Human-Orangutan Conflict Workshop, Bontang, E Kalimantan, Dec. 10. Russon, A. E. & Kuncoro, P. (2013). Orangutan di TN Kutai. Journalist Field Trip, Kutai National Park Authority, Kutai National Park, Dec. 8, 2013. Conference organization. Balai Taman Nasional Kutai & A. Russon (organizers) (2013). Lokakarya Internasional : Konservasi habitat dan orangutan disekitar Taman Nasional Kutai, Kalimantan Timur. (International Workshop on habitat and orangutan conservation in and around Kutai National Park, East Kalimantan). Hotel Royal Victoria, Sangatta, Kalimantan Timur, Indonesia, June 10-13. Publications Russon, A. E., Compost, A., Kuncoro, P., & Ferisa, A. (revised and resubmitted, Jan. 11, 2014). Orangutan fish eating, primate aquatic fauna eating, and their implications for the origins of ancestral hominin fish eating. Journal of Human Evolution Bebko, A. (2014). Introduction to small-scale GIS analysis: Determining the presence of habitual travel route networks in orangutans (Pongo pygmaeus morio). In C. A. Shaffer, F. Dolins, J. R. Hickey et al. (eds.), GPS and GIS for Primatologists: A Practical Guide to Spatial Analysis. Cambridge, U. K.: Cambridge University Press. Russon, A.E. et al. (in prep.) Pongo pygmaeus morio activity budgets in recovering habitat.

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