Crop Protection 30 (2011) 1263e1268

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Crop Protection
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Understanding of relationships between ground cover and rat abundances: An integrative approach for management of the oil palm agroecosystem
C.L. Puan a, b, *, A.W. Goldizen c, M. Zakaria a, G.S. Baxter d
a

Faculty of Forestry, Universiti Putra Malaysia, 43400 UPM, Serdang, Selangor, Malaysia The University of Queensland, School of Integrative Systems, Gatton QLD 4343, Australia c The University of Queensland, School of Biological Sciences, St Lucia QLD 4072, Australia d The University of Queensland, School of Geography, Planning and Environmental Management, St Lucia QLD 4072, Australia
b

a r t i c l e i n f o
Article history: Received 21 September 2010 Received in revised form 23 May 2011 Accepted 25 May 2011 Keywords: Vegetation cover Rattus argentiventer Oil palms Weeds Spatial associations

a b s t r a c t
Ground cover is an essential element in the selection of habitats by small mammals. It provides shelter, foraging opportunities and a hiding place from predators. In oil palm plantations rodents are abundant yet the relationships between ground cover structure and rats are not well understood. Plantation management emphasizes maximisation of oil palm yield by reducing rat damage and competition from weeds, typically through chemical control. This study examined the relationships between ground cover, including undergrowth and frond piles, and rat abundance. Rat trapping and vegetation assessment were conducted simultaneously in ve study plots in the Labu Estate, Negeri Sembilan, Malaysia. Over 21,000 trap-nights 1190 individual rats were captured and these data were analysed using spatial analysis by distance indices (SADIE). No signicant clustering of rats was observed within any plot over time. Redundancy analyses suggested that the overall occurrence of rats, especially Rattus argentiventer, was positively correlated with vegetation cover and height. This implies that habitat complexity is important to rats, even in a relatively simple agroecosystem. Results of the SADIE analyses were inconsistent, perhaps due to highly variable management practices in the plantations. The ndings from this study suggest that using an integrated approach in the control of both weeds and rats in oil palm plantations may maximise the benets from weeding while reducing the production loss due to rats and the need for chemical control of rats. 2011 Elsevier Ltd. All rights reserved.

1. Introduction In the tropics, there has been a rapid expansion of oil palm plantations (Carter et al., 2007) from 3.6 million ha globally in 1961 to 13.8 million ha in 2007 (FAO, 2009) because of the high demand for oils and fats (Basiron, 2007; Murphy, 2007; Corley, 2009; Lam et al., 2009). At the age of about 2.5 years, oil palms (Elaeis guineensis Jacq.) begin to produce fruit bunches reaching optimum oil production at seven to ten years. Replanting is normally conducted when palms reach about 20 years or sometimes more (Corley and Tinker, 2003). The main reason for replanting is that harvesting, fruit assessment and maintenance activities are more difcult when palms are tall. In such a monoculture some pest species, such as rodents, are common (Wood and Chung, 2003). Stomach analyses (Liau, 1990)

* Corresponding author. Present address: Faculty of Forestry, Universiti Putra Malaysia, 43400 UPM, Serdang, Selangor, Malaysia. Tel.: 603 8946 7183. E-mail address: chongleong@putra.upm.edu.my (C.L. Puan). 0261-2194/$ e see front matter 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.cropro.2011.05.025

of three rat species found in oil palm plantations indicate that oil palm mesocarp comprised a considerable part of their daily diets, i.e. 9.9 g/rat for Rattus rattus diardii (Jentink), 8.6 g/rat for Rattus argentiventer (Robinson and Kloss) and 4.3 g/rat for Rattus tiomanicus (Miller). Another study of R. tiomanicus in an oil palm plantation also revealed that on average about 96% of their stomach contents was made up of oil palm fruit (Wood and Liau, 1984). They feed on fruit both on trees and on the ground. The annual losses associated with rat damage were estimated at over a hundred million Malaysian Ringgit (RM) or US$ 32 million for the oil palm industry in Malaysia over twenty years ago (Basri and Halim, 1985) and these losses are likely to be greater now since these plantations have increased over that time (Carter et al., 2007; FAO, 2009). In oil palm estates, young palms are initially planted with ground cover, such as a legume, which eventually forms a thick blanket. This vegetative cover prevents erosion and nutrient runoff. It also improves soil structure, moderates soil temperature, controls certain insects (Corley and Tinker, 2003) and xes nitrogen. Mixed natural cover may be easier to manage (Hartley, 1988) than cover consisting solely of legumes, but overgrown vegetation may

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compete with palms for nutrients and water, and even cause yield reduction (Gray and Hew, 1968). Ring weeding around the base of each palm and the control of cover at intervals are needed to prevent competition with the oil palms. In addition application of articial fertilizer for the palms is common (Hartley, 1988). As palms grow in height, frond length increases and the canopy closes. In response the composition of the ground cover gradually changes (Dahlan et al., 1993; Wong et al., 2005), to favour shade tolerant plants such as ferns. Throughout the life of the palms older fronds are progressively pruned to ease harvesting and fruit collecting. Pruned fronds are stacked in piles between the palm rows, in an attempt to provide nutrients and mulch. Empty fruit bunches discharged from the mill are similarly deployed on the ground, although the effectiveness of using fronds and fruit in this way is yet to be empirically and economically tested. This management regime maintains a more or less constant supply of frond piles between the rows of palms in addition to undergrowth. Undergrowth may provide shelter and foraging opportunities (Bos and Carthew, 2003) for small mammals. The selection of cover is often associated with increased predation risk (Dickman, 1992; Hughes and Ward, 1993; Moreno et al., 1996; Marin et al., 2003; Stokes et al., 2004). Structurally complex habitats may provide small mammals, like rats, with nest sites and opportunities to avoid predators; thus modifying habitats by reducing structural complexity may reduce the potential of habitats to support small mammal populations (Jacob, 2008; Lambert et al., 2008). Several studies have shown that changes in habitats associated with agricultural practices may alter small mammal assemblages (Sullivan et al., 1998; Michel et al., 2007; Moro and Gadal, 2007). In the case of rodent pests in oil palm plantations, changes in vegetation cover could expose the rats during foraging and increase their predation risk. The resulting predator avoidance behaviour (Abramsky et al., 1996) may force rats to alter their feeding (Wood and Chung, 2003), activity ranges and thus distributions. If routine plantation management increases cover for rats in the form of palm frond piles or undergrowth there may be implications for both weed and pest management in oil palm plantations. Hence, this study examined the relationships between ground cover, including undergrowth vegetation and frond piles, and rat abundance. We hypothesized that; (1) the abundance of rats will increase with the increase of ground cover as undergrowth may act as cover and facilitate the foraging of rats, and (2) frond piles may act as refuges or shelter for rats in oil palm plantations in which changes in ground cover occurred through weeding and cattle grazing. 2. Materials and methods This study was conducted in the Labu Estate (2 450 N, 101490 E) in the southwestern part of Peninsular Malaysia. The 2569 ha oil palm plantation was converted from a rubber estate prior to the 1940s. It has sandy clay soils and undulating terrain. Palms were planted at an average distance of 8e9 m between and within rows. Cattle grazing was incorporated into palm oil production several years ago with an estimated 200 free-ranging animals moving in herds throughout the estate. Rodent trapping was conducted in ve study plots; three plots (Plots A, B and C) were located in older palms (seven years old) while the other two (Plots D and E) were in younger palms (three years old). All plots were located more than 800 m apart to ensure the independence of different plots. The independence of the plots was further conrmed by a complete absence of marked rats recaptured in a neighbouring plot. A total of 200, single catch cage traps (280 140 100 mm) arranged in pairs in a 10 by 10 grid with 30 m spacing were opened at each study plot for three consecutive nights during each of seven trapping sessions.

Trapping sessions were eight weeks apart (from November 2007 to December 2008), providing sufcient time for the rat population to change either through breeding or immigration (Aplin et al., 2003). Animals captured were identied, measured and marked with unique ear-notches. Ground cover materials including vegetation cover (%), vegetation height (mm) and frond piles (mm) were quantied in May, July and September 2007, concurrently with trapping sessions. Vegetation sampling in oil palm plantations was adapted from that of Germer (2003) and Germer and Sauerborn (2004). A total of three subplots of 1 m2 were randomly placed at each trapping point in three pre-determined strata: under the oil palm canopy, along the harvesting path and in the inter-row between the palms. Harvesting paths were roads used by tractors for fruit collection, whereas inter-rows between palms were not on the roads. This means there was a total of 300 subplots in each of the ve plots, with 100 under the oil palm canopy, and 100 each in the harvest path and inter-row positions. A sole surveyor estimated the percentage of vegetation cover to reduce the bias inherent when different observers make these estimates. A Robel pole (Robel et al., 1970) was used to measure the typical heights of ground vegetation and frond piles in each subplot at each trapping point. The typical height represents the highest totally visible band of the marked pole rounded to the nearest 5 cm. Measurements were taken for these variables with the eye level 1 m above the ground and at 4 m from the pole. All site measurements were later combined and means were calculated. As the frond piles had been systematically placed between palms throughout each plot their abundance could only be assessed at each trapping point (instead of each subplot). This allowed the analysis of spatial association of structural features of the habitat with rat abundance using spatial analysis by distance indices (SADIE, Perry et al., 1999; Perry and Dixon, 2002). Redundancy analyses were performed to examine the relationships among multivariate data using CANOCO for Windows,  version 4.5 (ter Braak and Smilauer, 2002). This software used Monte Carlo simulations to arrive at estimates of probability. Relationships were examined rst between rodent captures and environmental variables including vegetation cover, vegetation height, and amount of frond piles, and, second, between rodent species captures and cover and height of different vegetation groups, i.e. grasses and sedges, non-creeping broadleaf weeds, creeping broadleaf weeds, woody broadleaf weeds and ferns. Forward selection with a Monte Carlo permutation test was performed to select and assess the signicance of the explanatory variables. The SADIE procedure uses spatially referenced counts to characterize the spatial pattern of a data set by measuring the minimum distance to regularity, that is the distance that these individual counts would need to move to result in a completely regular or uniform arrangement (Perry et al., 1999). The program produces a number of indices to quantify the degree of non-randomness within the data set. The index of aggregation (Ia) equals 1 for a random arrangement, while indices of greater than or less than 1 indicate clustering or overdispersion, respectively. The probability that the observed counts were arranged randomly among the given sample units (Pa) is used to indicate the signicance of the overall degree of clustering; the 5% level of signicance was adopted. Clusters can be further dened in the form of patches (neighbourhoods of units with counts that are larger than the sample mean) or gaps (neighbourhoods of units with counts that are smaller than the sample mean), indicated with vi and vj clustering indices and the associated probabilities Pi and Pj, respectively. The presence of patches is indicated by a value of vi > 1.5 while gaps are indicated by vj < 1.5.

C.L. Puan et al. / Crop Protection 30 (2011) 1263e1268 Table 1 Distribution of Rattus spp. captured in older and younger palms. Species name Rattus rattus diardii Rattus argentiventer Rattus tiomanicus Others Total All palms Frequency 758 385 40 7 1190 % 63.70 32.35 3.36 0.59 100 Older palms Frequency 613 194 31 5 843 % 72.38 23.07 3.81 0.74 100 Younger palms Frequency 145 191 9 2 347 %

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41.79 55.04 2.59 0.58 100

To measure the spatial association between two data sets (e.g. rat counts and vegetation cover or frond pile data) collected using the same grid, an cp index was generated based on the similarity between the clustering indices obtained for the two variables (Perry and Dixon, 2002). A positive value for cp indicates an association between the two data sets whereas a negative value indicates dissociation and the signicance of an cp value is based on Pa<0.025 or >0.975. Further details on the derivation of the associated statistics from SADIE can be found in Perry et al. (1999) and Perry and Dixon (2002). 3. Results A total of 1190 individual rats were captured in the ve plots in 21,000 trap-nights (Tables 1 and 2) and over seven trapping session (Fig. 1). The majority of the rats comprised R. rattus diardii (63.70%), followed by R. argentiventer (32.35%) and R. tiomanicus (3.36%). A total of 35 plant species belonging to 24 families were identied; these were further categorised into six species of grasses and sedges, 14 non-creeping broadleaf weeds, seven creeping broadleaf weeds, three woody broadleaf weeds and ve ferns. Spatial analysis of the rat numbers in the trapping grids indicated that there was no signicant clustering of rats in any plots during any trapping sessions with the aggregation indices, Ia, less than or only slightly greater than 1. Regardless of the age of palms, all clustering indices (as indicated by the mean values of vj and vi) were also non-signicant, implying that there were no patches (clusters of large counts) or gaps (clusters of small or zero counts) of rat captures within the grids. SADIE analyses indicated that the spatial aggregation of vegetation cover and vegetation height was signicant in some plots and sampling periods (Table 3). For frond pile data, aggregation was observed in several plots but this was not stable over time. In those plots with aggregation of vegetation cover, no signicant associations or dissociations were observed with rat counts. Similar ephemeral associations occurred for vegetation height and there was no signicant relationship between vegetation height and rat numbers (Table 3). Following this a redundancy analysis was performed to visualise the correlations among the abundances of different rat species and all types of ground cover (Fig. 2). The rst two axes of the ordination diagram explained 34.5% and 17.2% of the variance in rat captures, respectively. Species-environment correlations were 0.664 for axis 1 and 0.921 for axis 2. The total variance explained by all variables including vegetation cover (P 0.049), vegetation height (P 0.017), frond pile (P 0.388), mulch (P 0.430), fruit bunch numbers (P 0.061) and age of palms (P 0.276) was 52.2%. The relative

abundance of R. argentiventer was observed to be correlated better with that of R. tiomanicus than that of R. r. diardii (Fig. 2). The relative abundances of R. argentiventer and R. tiomanicus were more positively correlated with vegetation cover and vegetation height than was the abundance of R. r. diardii, which correlated better with frond piles and the age of the palms than the other two rat species. Indeed, there was a negative correlation between the abundance of R. argentiventer and palm age. The amount of mulch was weakly correlated with rat abundance. Partial correlation further indicated a strong correlation between the relative abundance of R. argentiventer and vegetation cover (r 0.892, P < 0.001), while this was not the case for the other two species (R. r. diardii: r 0.300, P 0.175; R. tiomanicus: r 0.066, P 0.772). A separate redundancy analysis investigated correlations between the abundances of different rat species and different groups of vegetation cover, with the rst two axes explaining 15.1% and 3.1% of the variance in rat captures, respectively. Speciesenvironment correlations were 0.881 and 0.197, respectively. The total variance explained by the variables, including non-creepers (P 0.068), creepers (P 0.980), woody weeds (P 0.703), grasses (P 0.441) and ferns (P 0.490), was 18.3%. The biplots (Fig. 3) show that R. argentiventer and R. tiomanicus were more strongly correlated with all types of vegetation cover than R. r. diardii, except for ferns, with which R. r. diardii correlated strongly.

75.00

Rat abundance

50.00

25.00

0.00 1 2 3 4 5 6 7

Trapping session
Fig. 1. Means (SE) of rat abundances across plots by trapping session.

Table 2 Distribution of captures by sexes, age and breeding conditiona (with the respective percentages for each category in parentheses). Species name Rattus rattus diardii Rattus argentiventer Rattus tiomanicus
a

Male 392 (47.92%) 180 (44.33%) 25 (56.82%)

Female 426 (52.08%) 226 (55.67%) 19 (43.18%)

Adult 713 (94.06%) 336 (87.27%) 37 (92.50%)

Juvenile 45 (5.94%) 49 (12.73%) 3 (7.50%)

Breeding 139 (32.63%) 69 (30.53%) 4 (21.05%)

Non-breeding 287 (67.37%) 157 (69.47%) 15 (78.95%)

Breeding individuals include both pregnant and lactating females which were otherwise categorized as non-breeding individuals.

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Table 3 Summary of indices (and the associated probabilities) from spatial analysis of ground material variables (Ia, vj and vi), and between rats and ground material variables (cp) using SADIE. Variables Vegetation cover Month May 2007 Plot A B C D E A B C D E A B C D E A B C D E A B C D E A B C D E Ia 1.45 1.05 0.96 1.31 1.63 0.98 0.98 1.10 1.34 1.30 0.98 1.19 1.34 0.88 1.97 1.06 0.93 1.31 1.36 0.82 1.04 0.92 1.21 1.55 1.22 1.01 1.73 1.51 0.89 1.12 (0.256) (0.310) (0.540) (0.049)a (0.003)a (0.423) (0.471) (0.254) (0.057) (0.083) (0.436) (0.135) (0.052) (0.720) (<0.001)a (0.333) (0.637) (0.051) (0.029)a (0.886) (0.359) (0.606) (0.117) (0.014)a (0.128) (0.397) (0.002)a (0.016)a (0.724) (0.238) Mean vj 1.37 1.01 0.95 1.24 1.53 1.00 0.92 1.11 1.29 1.23 1.07 1.10 1.27 0.88 1.78 1.06 0.88 1.27 1.42 0.77 1.06 0.91 1.22 1.49 1.17 1.01 1.75 1.62 0.88 1.03 (0.013)a (0.394) (0.558) (0.068) (0.007)a (0.449) (0.617) (0.232) (0.064) (0.124) (0.256) (0.257) (0.085) (0.751) (0.004)a (0.385) (0.792) (0.067) (0.016)a (0.967) (0.295) (0.623) (0.104) (0.020)a (0.157) (0.282) (<0.001)a (0.005)a (0.739) (0.355) Mean vi 1.50 1.10 1.06 1.18 1.59 0.98 0.98 0.96 1.15 1.34 0.96 1.23 1.09 1.13 1.80 1.28 0.95 1.34 1.28 0.88 1.01 0.79 1.23 1.49 1.10 1.11 1.68 1.63 0.90 1.07 (0.013)a (0.211) (0.294) (0.111) (0.004)a (0.500) (0.450) (0.514) (0.146) (0.060) (0.500) (0.093) (0.225) (0.179) (0.002)a (0.090) (0.560) (0.033)a (0.051) (0.792) (0.513) (0.910) (0.882) (0.017)a (0.229) (0.141) (0.001)a (0.005)a (0.692) (0.274)

cp
0.23 (0.884) 0.15 (0.373) 0.01 (0.491) 0.16 (0.762) 0.16 (0.381) 0.22 (0.850) 0.203 (0.655) e 0.66 (0.072) 0.32 (0.150) 0.13 (0.368) e e e 0.29 (0.716) 0.08 (0.336) 0.41 (0.193) 0.05 (0.414) 0.05 (0.420) 0.23 (0.294) 0.42 (0.980) 0.40 (0.467) e e 0.37 (0.143) 0.25 (0.175) e e e 0.06 (0.462)

July 2007

Sept. 2007

Vegetation height

May 2007

July 2007

Sept. 2007

Pi and Pj < 0.05; bold cp value indicates Pa > 0.975; dash indicates parameter not analysed due to low numbers or even distribution of counts.

R. argentiventer and R. tiomanicus were more strongly correlated with creeping and non-creeping weeds than with woody weeds or grasses. 4. Discussion In oil palm plantations, vegetation other than the cash crop itself may be useful at some stages of planting, but is often disadvantageous at others. This study suggested that reduction of ground vegetation may possibly aid in rodent control. It is
1.0

probable that any operational activities such as weeding may have associated impacts on the communities of rodents living within oil palm plantations. Redundancy analyses indicated that the occurrence of rats was positively associated with both the percentage cover and height of vegetation (Fig. 2); this suggests that a reduction in those parameters would likely lead to a reduction in rat abundance. In oil palm plantations, regular control of ground cover may inuence the rats feeding behaviour, activity ranges and distribution. Exposed habitat may restrict the distances rats travel while

R. argentiventer fruit

vegetation height

1.0

vegetation cover

fern
R. tiomanicus
Axis 2 (17.2%)

R. r. diardii
Axis 2 (3.1%)

grass

mulch

R. r. diardii
creeper R. argentiventer R. tiomanicus non-creeper woody
-0.4

frond palm age

-1.0 -1.0

Axis 1 (34.5%)

1.0

Fig. 2. Redundancy analysis biplot diagram of rat species captures (arrows with solid line) and other environmental variables (arrows with dotted line). Arrows indicate the direction that species was increasing in abundance whereas the angles between variables approximate their correlations.

-0.4

Axis 1 (15.1%)

1.0

Fig. 3. Redundancy analysis biplot diagram of rat species captures and of cover of different vegetation groups.

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foraging because they must avoid predators in open habitat. The results from this study support those from radio-telemetry. The movements of radio-tagged R. tiomanicus in an oil palm estate in Malaysia (Buckle et al., 1997) revealed a preference for the use of areas with ground vegetation. A similar study on R. argentiventer in rice elds also suggested that the rats used the grassy environment of paddies for foraging, nesting and protection from predation (Tristiani et al., 2003), particularly when the crop became denser (Brown et al., 2001). However, the usage of ground vegetation elements in Buckle et al.s (1997) study, based on radio-xes, was inconsistent, which was also the case in this study. An attempt to further characterize spatial association patterns between vegetation cover and height and rat numbers at predetermined grid points using SADIE was inconclusive (Table 3). Although there was some clustering of ground vegetation in certain plots, this seemed to be ephemeral and changed over time. For the rats, clustering was rather weak and non-signicant. Spatial association between the two variables also varied by plot and was inconsistent. The pattern of clustering of rats may have been unclear as a result of regular weeding practices as well as the introduction of free-ranging cattle to the plantation as part of integrative agricultural practices. The dynamic or rapid turnover of ground vegetation may have caused the non-signicance of clustering of rats over space. Other possible factors that caused temporal variations in the clustering of ground vegetation and/or rats include food abundance (Puan et al., 2011) and rainfall. Frond heaps are another important habitat structure for rats, particularly as shelter (Buckle et al., 1997). It is believed that the frond heaps may favour the rats either by improving foraging or by providing hiding spots from predators (Lambert et al., 2008). However, a redundancy analysis indicated that there was no signicant relationship between the amount of frond piles and rat occurrence (Fig. 2). Similarly the degree of spatial aggregation of frond piles within grids determined using SADIE was also inconsistent for all plots and not signicantly related with the distribution of rats. In addition, this study has suggested that correlations between the occurrence of rats and ground materials may be species-specic and vary according to the ages of palms (Figs. 2 and 3). For example, redundancy analysis indicated a negative correlation between the abundance of R. argentiventer and palm age (Fig. 2). This was expected as this species often invades oil palms at early maturity (Wood, 1976; Corley and Tinker, 2003). As R. argentiventer is known to be a common species of grassy areas, this implies that the occurrence of such areas could be a potential source of colonization for the species (Wood, 1971). This also provides a possible explanation for why this species was still recorded in older palms in which vegetation cover was still available. As indicated by the present study, overall rat numbers were associated with ground vegetation. It is possible that reductions in ground cover would affect the rats by limiting their access to breeding sites and thus their population growth. Due to resource constraints we were unable to experimentally manipulate ground cover over larger spatial and temporal scales. It should be stressed that this study was conducted in a study site in which poison baiting was being practiced. Attempts to assess the effects of poisoning on the uctuations in rat numbers were unsuccessful. This is partly due to the way that each poisoning session was practiced with progressive baiting being done among plantation blocks at a large scale, which could take up to several months. The use of anticoagulant poison that may not have instantly killed the rats that ingested the poison may also have confounded the responses of rat populations to changes in vegetation cover and fruit abundance. Trap success in the present study was relatively lower than in other studies conducted in oil palm plantations (Wood, 1984) and

other farming systems (Brown et al., 2005). The recapture rate was also rather low in this study (i.e. 0.02e0.07% for each plot at each single time period) and correlations of recaptures with ground cover variables could not be obtained, preventing calculation of the trappability of each species. Although recaptures were low, all individuals re-captured were from plots where they were initially marked. One of the recaptures was obtained after 14 months indicating it was likely that some rats, at least, were sedentary residents. We suggest that experimental manipulations of ground cover be conducted in future studies to test the effects of changes of ground cover on rat abundances; it would be ideal to use a large experimental area free of baiting. The results of such experiments will shed light on the costs and the associated benets gained in rodent control by weeding, which could be incorporated into a broader cost/benet analysis of traditional pest control using rodenticide. Such a study would improve our understanding of whether manipulations of ground cover could reduce or avoid the necessity of poison baiting for rats. It may be important to maintain a high standard of clearing with minimal vegetative materials or grassy areas in palm estates. Weeding procedures and schedules should be implemented in a way that reduces the cover for rats as much as possible. This agroecosystem may not be as simple as previously thought and changes in ground vegetation over time may have an inuence on rodent pest communities. This should be taken into consideration by plantation managers when working towards better management practices. Acknowledgements Live trapping and vegetation surveys were conducted in the Labu Estate with permission given by Sime Darby Plantations Sdn Bhd. We appreciate the help and support given by Tey Chin Chong and other personnel of the Labu Estate. We are grateful to Latifah Zainal Abidin and Siti Eryani Suterisno for their kind assistance with vegetation identication and to those who kindly assisted in the trapping sessions. We thank Hadzi Mohd Noor for securing some nancial assistance for the project under Science Fund (Project No: 05-01-04-SF0740) given by the Ministry of Science, Technology and Innovation, Malaysia. The project was carried out with animal ethics approval from the University of Queensland (NRSM/325/08/NRSM). References
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