Gotowa bibliografia na temat „Santalum spicatum”

Natural recruitment of sandalwood (Santalum spicatum) is generally low in pastoral regions of the Midwest and Goldfields, Western Australia. Harvesting of S, spicatum for the aromatic timber occurs in these regions, creating a need to develop management strategies to conserve the species. This paper examines sandalwood recruitment over three years within a natural stand of 32 ha, near Paynes Find, Western Australia. Santalum spzcatum recruitment success was compared between three establishment treatments, and between two fencing treatments (+I-). At age three years, mean survival of S. spicatum seedlings planted next to host trees (25%) was significantly higher than those planted at harvesting spots (2%) and beneath parent trees (0%). In the unfenced treatment, there was evidence of grazing and S, spicatum survival and growth were significantly lower than in the fenced treatment. However, fencing alone did not improve S. spicatum recruitment because natural seed dispersal was poor and survival beneath parent trees was low. De-stocking, combined with seed enriching host trees is recommended to dramatically improve S, spicatum recruitment in the Paynes Find region. Santalum spicatum seedling performance was compared growing next to three N2-fixing species (Acacia burkittii, A. tetragonophylla and A. ramulosa) and one non N2-fixing species (Hakea recurva). At age three years, S. spicatum survival was significantly higher next to A. burkittii (33%) than A. tetragonophylla (1 7%). Santalum spicatum survival next to A. ramulosa and H. recurva was 24-26%. Fencing improved S. spicatum survival next to A. burkittii, and to a lesser extent next to A. tetragonophylla and A. ramulosa. In contrast, survival of S, spicatum seedlings next to H. recurva was unaffected by fencing. Santalum spicatum growth next to each host species was slow and significantly higher in the fencing treatment. Foliar concentrations of N, P, K and Ca were the same across 5'. spicatum treatments, but the concentration of Mg varied. The foliar K:Ca ratio was also similar between S. spicatum treatments, ranging from 1.4 to 2.0. Key words: Santalum spicatum, recruitment, seed enrichment, host species, foliar nutrients

Santalum spicatum (R.Br.) A.DC is a West Australian sandalwood species highly valued for the sesquiterpene-rich oil in mature heartwood. The oil composition, particularly levels of the valuable sesquiterpenoids α- and β-santalol and the allergenic E,E-farnesol, are known to vary across its natural distribution. Our study investigated associations of oil characteristics in 186 S. spicatum trees in semiarid and arid regions of Western Australia with genetic structure, environmental parameters and morphological features. We found associations between oil composition and genetic structure, as well as between oil composition and environmental factors. Analysis of individuals using STRUCTURE revealed two major genetic clusters (K = 2), comprising trees from the arid north clustered together, and the semiarid south-west clustered separately. Mantel tests revealed a significant association between oil characteristics and genetic distance (r = 0.129, P = 0.02). There was considerable variation in the growing environment of S. spicatum. An Adonis test showed a significant association between oil composition and provenance (P = 0.001) and between oil composition and soil type (P = 0.002) but not oil composition and other environmental characters. Soil type was significantly related to santalol and E,E-farnesol content. No significant associations between oil composition and morphological features were identified.

Population size structure of sandalwood (Santalum spicatum) was studied on four pastoral leases near Paynes Find and Menzies, in semi-arid Western Australia. Stem diameter, height, height to crown and the orientation of dry-sided stems were recorded for 1017 individual sandalwood. Populations of S. spicatum at Paynes Find contained only mature trees, indicating no successful recruitment for at least 30 years. In contrast, populations of S, spicatum at Menzies had a high proportion of seedlings and saplings. Crown measurements of mature S. spicatum trees indicated high grazing intensity at Paynes Find: mean height to crown at Paynes Find (147-148 cm) was significantly higher than Menzies (92-94 cm). Dry-side percentage differed significantly between directional faces, consistent with sun damage. Highest mean dry-side percentages were on stem sides facing the sun between midday and late afternoon: west, north-west, south-west and north. This directional pattern was the same between pastoral leases, and there was no interaction between pastoral lease and dry-side direction. Mean percentage of mature trees with a dry-sided stem was also significantly higher at Paynes Find (76-82%) than at Menzies (42-46%). Significantly less foliage low to the ground on mature trees at Paynes Find may have exposed the stems to more sun damage. Land systems did not significantly influence dry-side direction on Burnerbinmah or Goongarrie. No S. spicatum seedlings or saplings had a dry-sided stem. Key words: Santalum spicatum, size structure, dry-sided stems

Seeds of many plant species, including those of sandalwood (Santalum spicatum (R.Br.) A.DC., Santalaceae), are surrounded by a fruit endocarp that is hard and woody (this structure hereafter referred to as a 'nut'). The woody endocarp of S. spicatum provides a physical barrier to germination. This study investigated how this barrier is removed and the mechanism(s) controlling it. Field trials demonstrated that the endocarp cracked naturally and that the time of harvest and the presence of the epicarp affected the percentage of endocarps that were cracked. An investigation of the influence of wetting period and rate of drying on endocarp cracking showed that the rate of drying was most critical in inducing cracking and that the process was not heat-dependent. Field and pot studies showed that germination of sown nuts was improved when the woody endocarp was fractured. Results suggest that a simple wetting and rapid drying procedure can be used to crack large amounts of sandalwood nuts prior to sowing in the field. Results are discussed in relation to S. spicatum seed ecology. The relevance of weakening the woody endocarp of other non-Santalum species through endocarp wetting and rapid drying is discussed.

The density of Santalum spicatum was compared between 'land systems' and between 'land surface types' on four sheep stations in the north-eastern Goldfields: Yakabindie, Tarmoola, Glenorn and Minara. S.A spicatum density was recorded in 4–6A ha transect plots, with a total of 14,090 ha surveyed. Within each transect plot, the S. spicatum were divided into five groups based on stem diameter at 150A mm: < 25 mm, 25–74 mm, 75–124 mm, 125–174 mm and > 174 mm. The proportion of S. spicatum in each of the five size categories was similar between land surface types and between land systems, with the majority in two groups: 75–124 mm and 125–174 mm. S. spicatum recruitment was low, with less than 1.5 % seedlings (< 25 mm) and 7.9 % saplings (25–74 mm). Total density of S. spicatum on hills and ridges (0.65 stems/ha) was significantly higher than any other land surface type. The sandplains (0.05 stems/ha) supported the least. Within land systems, Laverton and Bevon (both hills and ridges) had the highest S. spicatum density. Yakabindie supported higher densities of S. spicatum than the other stations.

The commercially valuable root hemiparasite Santalum spicatum (R.Br.) A.DC. (sandalwood) once grew throughout the medium- to low-rainfall areas of the south-western agricultural region of Australia; however, this resource has been exhausted by over-exploitation and clearing for agriculture. There has been growing interest from the farming community and other investors in the development of a plantation Santalum spicatum industry in southern Western Australia. This study investigated the distribution of remnant S. spicatum within the Pallinup River catchment and assessed the risk of S. spicatum population decline due to salinity. The natural range of host species at different sites (river catchments) across the south coast was also investigated. Remnant populations of S.�spicatum within and adjacent to the Pallinup River catchment were small (1–70 trees) and highly fragmented. The risk of further population decline due to salinity was concluded to be small because remnant trees were generally growing in well drained, sandy soils that were elevated above (median 9 m) their immediate drainage line. Across the seven river catchments surveyed, S. spicatum occurred in a range of vegetation associations and parasitised numerous species (68) from a wide range of genera and families. The suite of species exploited varied within and between catchments. Thirty species, including most monocots and Myrtaceae, were not successfully parasitised. Remnant S. spicatum always occurred on well drained soil types that supported open-woodland or mallee–heath communities. Sandalwood plantations, supported by numerous individuals of a range of host species (10–40 species), were shown to be productive in terms of sandalwood growth. The scale of the developing sandalwood plantation industry is likely to be small and unlikely to cover large areas of catchments. Thus, this industry alone is unlikely to address the salinity crisis through broadscale recharge management. However, additional to on-site recharge reduction, biodiverse host plantations may improve the prospects for biodiversity and rivers in salinising landscapes through the protection and enhancement of natural biodiversity, creation of new habitat, conservation of plant species and by providing a commercial incentive to protect biodiversity.

The seed of Santalum spicatum is rich in a fixed oil (59% by weight), which is characterised by a high percentage of acetylenic, ethylenic ximenynic acid (35% of total fatty acids). A number of important aspects of the seed fixed oil, its composition in developing seeds, its triacylglycerols molecular species in the oil, the nutrition and toxicity of the oil feeding, and the possible bioactivity of ximenynic acid in mice were investigated.The identification of cis and trans isomers of ximenynic acid in the seed oil, and the metabolite of ximenynic acid in mouse liver lipid fractions were achieved using 2-amino-2-methyl-1-propanol to form 2-substituted 4,4-dimethyloxazoline derivatives, which were analysed by gas chromatography with mass spectrometric detection.Changes in proximate and fatty acid composition were investigated in developing seed collected weekly from about seven days after flowering to maturity. It was determined that moisture and carbohydrate contents decreased significantly during the development sequence, while fixed oil content increased from 0.3% to 50% (by weight) with seed development. A corresponding increase in the proportions of both oleic and ximenynic acids occurred suggesting a precursor/product relationship. Mature seed collected from different locations in Western Australia showed minor differences in characteristics and lipid composition, which may have been influenced by geographical origin and harvesting year of samples.The lipid components from the seed oil were separated using thin-layer chromatography and the individual triglyceride bands were characterised by high performance liquid chromatography and gas chromatography using flame ionisation and mass spectrometric detection after removal from the plate. The triximenynin (trisantalbin) band showed no other contaminating fatty acids and was obtained in a relatively pure state.A nutrition and toxicity study was performed by feeding a semi-synthetic diet containing sandalwood seed oil to a level of 15% of total energy content to a group of mice for one month and another group for two months. The most significant effect of sandalwood seed oil ingestion when compared with a standard lab diet (5% fat, by weight) and a canola oil-enriched diet (15% fat, by weight) was an apparent reduction in body weight gain, which may be the effect of ximenynic acid as a growth retardant. Serum aspartate aminotransferase levels were determined in the mice as an indicator of hepatotoxicity. These levels were higher in mice fed the sandalwood seed oil diet than those fed the standard lab diet, suggesting that ximenynic acid may affect liver-specific enzyme activity. Analysis of the total lipid fatty acids of various tissues and organs of mice showed only a low incorporation of ximenynic acid into the general tissues (0.3-3% by weight), and its absence in the brain.This study suggests a few health benefits from consumption of large quantities of sandalwood seed oil in the diet. These include a low lipid content in blood, heart, muscle, increase in the 16:1/16:0 and 18:1/18:0 ratios, production of increased levels of 18:1 (n-9) and docosahexaenoic acid, and decreased levels of arachidonic acid in certain tissues. There were no specific pathological, morphological or mortality changes observed in the mice.Sandalwood seed may be both a food and a medicine.

The seed of Santalum spicatum is rich in a fixed oil (59% by weight), which is characterised by a high percentage of acetylenic, ethylenic ximenynic acid (35% of total fatty acids). A number of important aspects of the seed fixed oil, its composition in developing seeds, its triacylglycerols molecular species in the oil, the nutrition and toxicity of the oil feeding, and the possible bioactivity of ximenynic acid in mice were investigated.The identification of cis and trans isomers of ximenynic acid in the seed oil, and the metabolite of ximenynic acid in mouse liver lipid fractions were achieved using 2-amino-2-methyl-1-propanol to form 2-substituted 4,4-dimethyloxazoline derivatives, which were analysed by gas chromatography with mass spectrometric detection.Changes in proximate and fatty acid composition were investigated in developing seed collected weekly from about seven days after flowering to maturity. It was determined that moisture and carbohydrate contents decreased significantly during the development sequence, while fixed oil content increased from 0.3% to 50% (by weight) with seed development. A corresponding increase in the proportions of both oleic and ximenynic acids occurred suggesting a precursor/product relationship. Mature seed collected from different locations in Western Australia showed minor differences in characteristics and lipid composition, which may have been influenced by geographical origin and harvesting year of samples.The lipid components from the seed oil were separated using thin-layer chromatography and the individual triglyceride bands were characterised by high performance liquid chromatography and gas chromatography using flame ionisation and mass spectrometric detection after removal from the plate. The triximenynin (trisantalbin) band showed no other contaminating fatty acids and was obtained in a relatively pure state.A ++
nutrition and toxicity study was performed by feeding a semi-synthetic diet containing sandalwood seed oil to a level of 15% of total energy content to a group of mice for one month and another group for two months. The most significant effect of sandalwood seed oil ingestion when compared with a standard lab diet (5% fat, by weight) and a canola oil-enriched diet (15% fat, by weight) was an apparent reduction in body weight gain, which may be the effect of ximenynic acid as a growth retardant. Serum aspartate aminotransferase levels were determined in the mice as an indicator of hepatotoxicity. These levels were higher in mice fed the sandalwood seed oil diet than those fed the standard lab diet, suggesting that ximenynic acid may affect liver-specific enzyme activity. Analysis of the total lipid fatty acids of various tissues and organs of mice showed only a low incorporation of ximenynic acid into the general tissues (0.3-3% by weight), and its absence in the brain.This study suggests a few health benefits from consumption of large quantities of sandalwood seed oil in the diet. These include a low lipid content in blood, heart, muscle, increase in the 16:1/16:0 and 18:1/18:0 ratios, production of increased levels of 18:1 (n-9) and docosahexaenoic acid, and decreased levels of arachidonic acid in certain tissues. There were no specific pathological, morphological or mortality changes observed in the mice.Sandalwood seed may be both a food and a medicine.

Several approaches have been applied to develop procedures for genetic improvement of Western Australian sandalwood (Santa/um spicatum) and Indian sandalwood (S. album): conventional crossing, protoplast fusion and Agrobacterium-mediated gene transfer. In the conventional approach, the possibility of producing inter-specific and intra-specific hybrids of these species was investigated. The reproductive biology of the species ie. flower morphology, stigma and ovular receptivity and sexual compatibility between genotypes and species, was studied. The results showed that both sandalwood species are obligate out-crossing species with pre- and postfertilisation barriers preventing self and inter-specific pollination. For both intra- and inter-specific crosses, initial fruit set was low with 70-100% fruit abscission. Reasons for failure of fruit set included lack of fertilisation, lack of an embryo sac in some S. album flowers, delayed endosperm development and necrosis of placenta tissue. An ovary culture technique was developed and which could be used to rescue embryos of intra-specific crosses 4-7 months after pollination, but not putative inter-specific hybrid fruit, which abscised at 1-3 months old. Putrescine spray increased fruit set and delayed fruit abscission for both intra-specific and inter-specific crosses. However, only a few putative hybrid fruits were retained until 4 months after pollination; they were not harvested for ovary culture but allowed to develop to maturity. These mature putative hybrid seeds did not germinate but their endosperms were confirmed to be hybrid by random amplified polymorphic DNA (RAPD) analysis. RAPD results indicated the wide genetic distance between S. spicatum and S. album. A tissue culture approach was conducted in parallel with conventional crossing. Direct somatic embryogenesis of S. album and S. spicatum was obtained by including thidiazuron in the culture medium. The published protocol of S. album regeneration through protoplasts could not be reproduced due to necrosis of the microcalli. An improved method of protoplast culture was developed by including thidiazuron and 2,4-D in the protoplast culture medium to induce direct somatic embryo formation in S. album. Reproducible protoplast fusion for S. album and S. 5picatum was accomplished and a high proportion of binary heterokaryons obtained. The heterokaryons grew to non-morphogenic microcalli while homokaryons of S. album developed to somatic embryos. The efficient in vitro regeneration of S. album via direct somatic embryos was combined with an Agrobacterium-mediated gene transfer procedure. Putative transgenic somatic embryos and plantlets were obtained and confirmed to be transgenic by the gus histochemical assay.

Sandalwood is an important international commodity, recognised for its aromatic oil which is a key ingredient in many fragrances and cosmetics. Western Australian (WA) sandalwood (Santalum spicatum) is known to be a cheaper alternative for the superior Indian sandalwood (Santalum album) as it has a lower oil content and lower quality oil. The natural stocks of S. album have declined due to illegal poaching, mismanagement, and disease. WA sandalwood’s natural stands have also reduced due to historical mismanagement. As a result, WA sandalwood (S. spicatum) has been established in plantations in the southern half of WA to attempt to meet the demands of the sandalwood industry. Plantation WA sandalwood is promoted to farmers as agroforestry, with the promise of economic and environmental benefits. While these benefits are attractive, sandalwood has an estimated 25 year rotation. This research aimed to determine the effect of physical and chemical treatments on oil production and heartwood formation in WA sandalwood, with the aim being to increase oil production, thus allowing the time between establishment and harvesting to be reduced. This study was conducted over three plantations in the Wheatbelt region of Western Australia; ‘Sandawindy’, ‘Kylie Reserve’, and ’Brookton’. At each site, four treatments were applied: a dowel soaked with the plant hormone Methyl Salicylate (MeSA) and inserted into the tree (Treated Dowel treatment), a dowel with no MeSA inserted into the tree (Blank Dowel treatment), a drill hole left empty (Empty Drill treatment), and a section of bark removed from the tree (Bark Removed treatment), as well as a group of trees left as a control for comparison. The Blank Dowel and Empty Drill treatments were established to determine if any significant increases of sandalwood oil in the Treated Dowel treatment were a result of the MeSA, the foreign dowel, or drilling into the tree. The Bark Removed treatment was used to mimic drysidedness; a condition that occurs naturally in the Rangelands of WA as a result of sun scald. The sandalwood trees were measured and treated in November of 2016. Plantations were divided into 30 evenly sized blocks per site, with 6 replicate blocks allocated to each treatment and control group. Two replicate blocks for every treatment and control group at each plantation were harvested in November of 2017, and all trees were remeasured. Of the approximate 300 trees harvested, 150 were cored using a 12 mm auger drill. These core samples were analysed for oil yield and composition at Wescorp’s laboratory. The total oil was measured an analysed, as well as the oil constituents α-santalol, β-santalol, farnesol, nuciferol, and β-bisabalol oil compositions (percentage) and yield (%w/w). All trees that were harvested ii were cut into 8 discs measuring 25 mm each, and the percentage of heartwood area at each height was measured and recorded. All data was statistically analysed using a univariate general linear model. There was no treatment that consistently increased total oil or oil component yields, qualities, or heartwood area percentages. The Empty Drill treatment resulted in more oil production than the control group on the most occasions, however it did not consistently increase oil production. This showed that the presence of MeSA did not have a significant effect on oil production, and the physical wounding of the tree had the overall greatest effect. The Kylie Reserve plantation showed low oil yield and low heartwood area percentages compared to the Sandawindy and Brookton plantations, although also showed the highest oil yields. This research, while not showing significant increases in oil production for the different treatments used, has giving a promising indication that a longer time between treatment and harvesting could influence the oil production. Further research extending this study should be conducted to give more information on the effect of the treatments on oil production.

[Truncated abstract] An investigation into the causes of heartwood and essential oil content of Australian plantation sandalwood, Santalum album was undertaken. Genetic diversity of 233 S. album, five S. austrocaledonicum and fifteen S. macgregorii trees growing in the Forest Products Commission arboretum, Kununurra WA, was assessed using nuclear and chloroplast RFLPs. Santalum spicatum was chosen as an out-group. Nuclear genetic diversity of the S. album collection was very low, with observed and expected heterozygosity levels of 0.047. This was lower than the results previously reported in the literature for trees in India, however a different technique was used. Based on allelic patterns, the collection was able to be categorised into 19 genotypes; each representing some shared genetic origin. Some groups were highly redundant with 56 trees being represented, while others were populated by just one tree. The essential oil yield and heartwood contents of trees from these genetic groups were compared. Yields were highly variable both within and between groups of trees which share a common genetic history, suggesting a significant environmental component was contributing to the observed phenotype, despite identical soil and climatic conditions. Ancestral lineages were tested using chloroplast RFLPs, although a lack of shared mutations between species made this difficult. Only one S. album tree originating from Timor was resolved using nuclear RFLPs, with the other trees being grouped with material sourced from India. There was no resolution of Indian S. album from Timorese using chloroplast RFLPs, however one S. album tree grown from Indian seed possessed a single unique mutation. The low genetic diversity of the Australian S. album collection is likely to be a combination of incomplete seed sourcing and highly restricted gene flow during the evolution of the species. Combined with information gathered on the phylogeny of the genus by other researchers, S. album is postulated to have originated from an over-sea dispersal out of northern Australia or Papua New Guinea 3 to 5 million years ago. Essential oil yield and composition was assessed for 100 S. album trees growing in the collection, ranging in age from 8 to 17 years. Oil content of heartwood ranged from 30 mg g-1 to 60 mg g-1, and the transition zone 36 mg g-1 to 90 mg g-1. Sapwood contained almost no sesquiterpene oils. Despite the highly variable total oil yields, the chemical profile of the oil did not vary, suggesting there was limited genetic diversity within this region of the genome. Strong, positive correlations existed between v sesquiterpenoids in the essential oil of S. album. ... These represent the first TPS genes to be isolated from sandalwood and will enable further elucidation of oil biosynthesis genes. This thesis compiles a three-pronged approach to understanding the underlying causes of oil yield variation in S. album. As a species for which so little is known, the research presented here provides a major leap forward for tree improvement, breeding and silviculture. Hence the best of Santalum album research is presented.

Links between species and the effect they have on ecosystem function is becoming increasingly recognised. Examples of such links include the complex, often multi-staged process of animal mediated seed dispersal, seed-caching by mammals and the tripartite relationship that occurs between many fungus-eating (mycophageous) mammal species, mycorrhizal fungi and woody plants. In this thesis, the role a small omnivorous marsupial, Bettongia penicillata ogilbyi (woylie), plays in ecosystem function, using its interaction with Santalum spicatum (Western Australian sandalwood) as a model, was examined. The study was conducted in a semi-arid open wandoo (Eucalyptus wandoo) woodland, in Western Australia. Dryandra Woodland, (32°48'8, l16°54'E) 160 km southeast of Perth, is one of the largest and most diverse remnants supporting over 800 native plant species and 24 mammal species, seven of which are threatened. Dryandra Woodland experiences a Mediterranean climate with warm to hot, dry summers and mild, wet winters. Bettongia penicillata Grey 1837 is a small, nocturnal marsupial within the family Potoroidae. Since European settlement, the distribution and numbers of woylies have decreased dramatically. Factors attributed to this decline include habitat loss, the introduction of feral predators such as the cat (Felis catus), the European red fox (Vulpes vulpes) and competing herbivores. From its original distribution across the south-western third of the continent, only three remnant natural populations remain in the south-west of Western Australia at Perup Nature Reserve, Tutanning Nature Reserve and Dryandra Woodland representing a reduction in range of approximately 97%. Santalum spicatum (R. Br) DC (Western Australian sandalwood), family Santalaceae, is a small, hemiparasitic tree which has virtually disappeared from the 300-600 mm rainfall zone due to widespread clearing of natural woodland and excessive unregulated harvesting. Furthermore, it has been suggested that seed dispersal is limited in areas where woylies have become extinct. Four broad objectives were addressed in this thesis: 1. To determine the population and dietary requirements of woylies in Dryandra Woodland 2. To determine the impact woylies have on the regeneration of sandalwood, by measuring recruitment of sandalwood in an area where woylies are present compared with an area where they are absent 3. To examine, in detail, seed dispersal and seed caching behaviour in the woylie by radiolabelling seeds with scandium 46. This will include the types of seeds cached, how caches are located, whether secondary or tertiary caching occurs, seed preference, germination rate from caches and seedling predation 4. To develop a rationale for woylie conservation and reintroduction based on an understanding on the woylie's role in ecosystem function and species' coexistence. Two experimental sites, with both woylies and sandalwood, and a control site, with sandalwood but no woylies were used in this study. Site A is outside the main block of Dryandra Woodland, 1.65 km from the main entrance to the Woodland. This site lies parallel to the main Wandering-Narrogin road and across from farmland situated next to woodland. The sandalwood was planted in the 1950s and has numerous mature sandalwood trees, saplings and seedlings. Site B is within the woodland, 3.2 km from the Dryandra village. Site B was planted in the late 1970s to early 1980s and has very few new sandalwood recruits and the few seedlings that did occur were found on soil mounds along the fenced area parallel to and across from a dirt road. The control site, with sandalwood but no woylies was at the Wickepen Water Reserve, 40 km south-east of Dryandra Woodland. This site had many mature sandalwood trees but very little recruitment growing away from the parent crown. Trapping sessions, lasting four nights per experimental site, were commenced in May 2002 and repeated at regular intervals until December 2005 giving a total of 1300 trap nights at site A and 1400 at site B. The 2005 trapping sessions were carried out four times a year to cover the seasons for dietary analysis for Bettongia penicillata ogilbyi (woylie) and Trichosurus vulpecula (brushtail possum). Because the number of trapped and re-trapped woylies were so low, the Jolly-Seber method, used to estimate possum numbers, could not be used to estimate woylie populations at either site. Instead, the minimum number of animals known to be alive (KTBA) was calculated by counting the number of times an individual woylie was trapped over at least three sessions. Findings from this study indicate that woylie numbers are declining in Dryandra. Site A had a very low number of woylies KTBA with only two females trapped once in June 2002 and one individual male, trapped intermittently until February 2004. Subsequent trapping failed to catch woylies until December 2005. Because of the low number of woylie captures at this site no population data could be analysed. The number of woylies KTBA at site B was consistently higher than site A, although still low. A total of 11 individual females and 17 individual males were trapped between May 2002 and December 2005 giving a sex ratio of 1:1.5 females to males. The trapping effort in the spring of 2003 resulted in an estimated population of brushtail possums ranging from 17 at site A to 70 at site B thus giving a density of approximately 1.4 and 5.8 brush tail possums ha- 1, respectively. The trapping effort in spring 2005 resulted in an estimated population of brushtail possums of 25 at site A and 35 at site B, thus giving a density of 2.1 and 2.9 brushtail possums ha -1, respectively. The trap success per 100 trap nights for brushtail possums was significantly (p <0.0001) higher than that of woylies at both experimental sites during 2003 and 2005. Total trap success was compared between sites for woylies and brushtail possums. There was a significant ex;= 16.41, p <0.01) difference in the trap success between the sites for woylies, and a highly significant ex;= 42.04, p <0.001) difference for brushtail possums for the 2003 trapping. The trap success for woylies, per 100 trap nights, was not significantly ex;= 4.1358, p >0.100) different between seasons for site B in 2005. Similarly, the trap success for brushtail possums, per 100 trap nights, was not significantly ex; =2.8565, p >0.200) different. However, brushtail possums, for all seasons, had a significantly (ex12 =103.9, p <0.0001) higher trap success compared with woylies for site B. Site A was not analysed due to the lack of woylies at this site from 2004 onwards. Trapping at the control site failed to catch woylies, and only one possum was caught during the trapping effort. Fungal spores occurred in 100% en= 16) of woylie scats for winter, spring and summer. During the three seasons analysed, woylies consumed an equal balance of spore types from hypogeal (n = 7) and epigeal en= 7) fungi. Overall, woylies ate eight spore types in winter and 12 spore types in spring. Fungi appeared to be important in the diet of brushtail possums during autumn and summer as a higher percentage of brushtail possums had fungal spores in their scats during these seasons compared with spring (x12= 13.94,p <0.001) and winter (x12 =8.65, p <0.005). The percentage of brushtail possums that consumed fungi during winter and spring did not (x12 = 1.128, 0.250 > p <0.500) differ, nor did the percentage of brushtail possums that ate fungi in autumn and summer (X12 = 0.00046, 0.925> p <0.99). The major spore type consumed throughout the year by both the brushtail possum and the woylie was Mesophellia. However, the amount of Mesophellia consumed by the brushtail possum differed between seasons (F = 83.472, df=1, p <0.001) as it was only dominant in the diet during the hotter months (i.e. summer and autumn). Austrogautieria and Mesophellia were the only spore types present in both brushtail possum and woylie scat samples in summer. In contrast to Mesophellia, spores of Gastrotylopilus dominated the brushtail possum and woylie scat samples during the cooler seasons (winter and spring) and were absent in summer and autumn. They occurred in significantly higher densities in winter than spring (F= 10.390, df = 1 p = 0.002) in brushtail possum scats. Overall, woylie scats contained a higher density of spores for all spore types than in brushtail possum scats. The total spore densities per gram of scat for woylies were 348.4 x 104 ± SE 277.1 x 104 for winter, 170.7 x 104 ± SE 110.3 x 104 for spring and 1353.4 x 104 ± SE 450.6 x 104 for summer. At site A the spatial distribution of adult sandalwood trees, adult to juvenile, adult to seedling and juvenile to juvenile were aggregated (p <0.05), whilst seedling to seedling distribution was highly aggregated (p <0.005). The density of adults was approximately 24 trees ha-1, there were approximately 107 saplings ha-1 and approximately 128 seedlings ha- 1. The furthest sandalwood seedling found growing away from an adult was 91 m. At site B the distribution of mature adult trees, adults to saplings and saplings to saplings were all highly aggregated (p <0.005). Adults occurred at an approximate density of 81 trees ha-1, and saplings at 38 trees ha-1. At the control site, mature trees were highly aggregated (p <0.005) with a density of 15.7 trees ha-1. There were very few seedlings and saplings at the site, all of which were clumped under or around the crown of the parent tree. Log-linear analysis indicated a significant two-way interaction between the presence of woylies and the distance the offspring were found in relation to adult trees (x; = 288.4,p <0.0001) and between the presence of woylies and the age of the offspring (saplings and seedlings) (x; = 34, p <0.0001). Thus, in Dryandra, where woylies were present, there were higher numbers of both seedlings and saplings growing more than 1 m from adult trees, compared with Wickepin Water Reserve, where woylies were not present. Conversely, in the absence of woylies, more seedlings and saplings grew less than 1 m from adult trees. For the three year survival rates of seedlings surveyed for the three sites, log-linear analysis indicated a significant two-way interaction between site and seedling survival (x23 = 218, p <0.0001) and between seedling survival and under/away from parent crown (x23 = 107, p <0.0001). Site A experienced significantly higher mortality of seedlings growing away from the crown after three years compared with site B, whilst the control site suffered significant seedling mortality under the crown. There was a highly significant (one-way ANOVA, F(2,99)= 125.58, p <0.0001) difference between Dryandra Woodland and Wickepin Water Reserve in the mean number of whole seeds found under adult sandalwood trees. The mean number of whole seeds under the parent crown at site A was 0.97 ± 0.16, range 1 to 2 (n = 34). At site B the mean was 0.76 ± 1.67, range 1 to 9 (n = 34), while at the control site the mean was 59.97 ± 11.19, range 10 to 330 (n = 34). A posteriori analysis indicated that the mean number of whole seeds under the parent crown at Wickepin Water Reserve, where woylies were extinct, was significantly greater than those at Dryandra Woodland where woylies were still present (p <0.0001). Log-linear analysis indicated a significant two-way interaction between the presence of woylies and the distance the offspring were found in relation to adult trees (x22 = 288.4, p <0.0001) and between the presence of woylies and the age of the offspring (saplings and seedlings) (x22= 34, p <0.0001). To enable large numbers of seeds to be tracked effectively over a period of time, two experiments were undertaken using a labelling technique with the isotope scandium-46 (Sc46). Scandium-46 is a moderate beta and a high level gamma-emitting radionuclide with a half life of 83.6 days and a maximum photon energy of 1.12 Mega electron volts (MeV) 100%. It is the high gamma emissions that allow seeds labelled with Sc46 to be tracked using a Geiger Muller (GM) counter to determine their fate. Scandium-46 is absorbed by the seed hull and each seed typically received ~ 1 microcurie (uCi) [37 ki1obecquerel (kBq)] of activity, sufficiently strong to allow detection of buried seeds from about 30 cm with a GM counter. The seeds labelled for the May 2005 experiment were sandalwood, S. acuminatum (quandong), Acacia acuminata (jam) and Gastrolobium microcarpum whilst in the February 2006 experiment only sandalwood seeds were used. In the May 2005 experiment, woylies ate or cached all the sandalwood seeds before any interaction with either S. acuminatum (quandong), Acacia acuminata (jam) or Gastrolobium microcarpum seeds ocurred. In the first night all the sandalwood seeds had been removed with 26 cached and 24 seeds eaten in situ under or within 1 m from the source tree. The woylies took three nights to remove all the quandong. On the third night the rate of removal increased which resulted in 6 cached seeds and 39 seeds were consumed in situ. The next type of seed to be removed was the Acacia acuminata which were all consumed by one woylie. No Acacia seed caches were found. On the fifth night of the experiment the majority of Gastrolobium seeds were observed to be eaten in situ by one woylie. Two caches of Gastrolobium seeds were located, one cache with 22 seeds and one with 15 seeds buried within 8 m of each other. In the February 2006 experiment, of the 500 seeds deployed under the source tree over four nights, 211 (42.2%) were eaten in situ, 185 (37%) were cached and 104 (20.8%) had an unknown fate. Individual seeds were buried between 1 and 6 cm deep (mean 3.15 ± SE 0.11 em). By November 2006, 185 primary, 120 secondary, 52 tertiary, 35 quaternary and 17 quintic caches had been located. Of the 185 seeds initially buried, 40 (21.6%) seeds had been dug-up and eaten in situ, 5 (2.7%) were left undisturbed in the caches, 20 (10.8%) were removed from the area, their fate unknown, and the remaining 120 (65%) seeds were re-cached into secondary caches. Of the 120 secondary caches, 38 (32%) seeds were dug up and eaten in situ, 12 (10%) were left undisturbed, 18 (15%) were removed from the area, their fate unknown, and 52 (43%) had been re-cached into tertiary caches. Subsequently, seven (13.5%) of the seeds were consumed from these caches, 10 (19.2%) were left undisturbed and 35 (67%) seeds were re-cached to quaternary caches. From the quaternary caches, 12 (34.2%) seeds were dug-up and eaten in situ, 6 (17.1%) caches were left undisturbed and 17 seeds were dug-up and recached for a fifth time. Of these quintic caches, 4 were dug-up and eaten, 5 (29.4%). The most common site for primary through to quintic caches was A. acuminata. The next most common area for primary to quaternary caches was out in the open not near any particular vegetation type. The exception was the location of quintic caches which had a higher percentage of caches under fallen logs (24% n = 4) compared with out in the open (18% n = 3). There were a relatively high number of seeds cached within 30 em of a sandalwood tree in all types of caches. The fate of 17 individual seeds that were cached and re-cached in five different caches was mapped, however, it remains unknown whether the same woylie that made the primary cache went on to move the seeds five times or whether it was several different woylies moving the seeds around. The fate of cached seeds was monitored for germination rates in situ. All the seeds were eventually consumed from the May 2005 experiment before any germination could take place. For the 2006 experiment, overall germination rates were low with only 6 (15.8%) of seeds from the 38 undisturbed caches germinating. Of these, two from the secondary caches germinated out in the open and did not survive. The single seed from an undisturbed tertiary cache germinated beneath a fallen log as did one of the seeds that germinated and survived from the undisturbed quintic cache while another successfully germinated beneath a G. microcarpum bush. Olfaction appeared to be the principle method used by woylies to locate buried seeds, a process which they are very efficient at. There was a significant (x12 = 6.5, 0.025> p <0.01) difference in the number of artificially cached seeds dug-up by woylies compared with disturbance of the control 'caches'. Of the 80 caches, 65 (81.2%) were located by woylies over three nights. During the same time period, 39 (48.7%) of the control 'caches' showed signs of being disturbed, thus suggesting that woylies were, at least in part, responding to the smell of disturbed soil as well as the actual seed. Furthermore, there was also a highly significant (x12 = 7.5, 0.01> p <0.005) difference between the number of seeds located by woylies under leaf litter compared to the control. All of the 20 seeds under the leaf litter were located within two nights compared with only six (30%) of the control 'caches' in the leaf litter showing signs of disturbance. The predation rates of buried seeds and emerging seedlings was measured. After distributing 100 sandalwood seeds under a random sandalwood tree at each site, there was significantly (x12 = 12.25, p <0.0001) less seed disturbance at Site A on the first night compared to site B. Conversely, on the third night there was significantly (x12 = 10.96, p <0.0001) more seed disturbance at site A compared with site B. All of the 100 seeds had been eaten in situ or removed by woylies at the end of the third night at site B whilst at site A the 100 seeds were eaten in situ or removed by woylies by the end of the fourth night. It is without doubt that woylies are prolific seed cachers and that their relationship with sandalwood is complex and mutual in nature. The woylies benefit by consuming some of the large nutritious seeds sandalwood produces. In-turn, sandalwood benefits by having a vector, the woylie, to disperse their seeds many of which are buried in areas conducive to germination, for example, under fallen logs and near sandalwood's host species. Woylies are able to very efficiently locate buried seeds by using olfaction and possibly visual clues of disturbed earth during random foraging for hypogeous fungi, although there was no evidence to suggest they use spatial memory to locate seeds. To determine if woylies use emerging seedlings as markers, seedlings were grown in the greenhouse and transplanted in Dryandra. Of the 46 seedlings transplanted at site A, 14 (30%) were intact and growing after one month. A total of 23 seedlings (50%) were dug-up by woylies. Of these, 17 (37%) were discarded whilst the remainder of the endosperm was eaten in situ leaving the endocarp on the ground. In the remaining six (13%) seedlings the endocarps were removed from the area, presumably to be eaten elsewhere rather than re-cached as they would have been split open during the germination process and, therefore, no longer suitable for storage. Similar numbers and fates were recorded at site B with 10 ( 43%) left intact, 22 (51%) of the seedlings had been dug-up by woylies and discarded whilst the remainder of the epicarp was eaten in situ leaving the endocarp on the ground. Three (7%) of the seedlings were dug-up by woylies but again the endocarps were removed from the area. Six (14%) seedlings at site B were also grazed and two (5%) seedlings died. There was no significant (x12 = 3.96, p <0.25) difference between sites for the fate of seedlings. The majority of seeds that were retrieved by woylies from the seedlings were eaten in situ at both sites. There was no significant (x12 = 1.583, 0.25 < p > 0.10) difference between sites for seeds eaten in situ or removed from the area. The fate of the seeds removed from the area was unknown. It was concluded that woylies have both a mutualistic and antagonistic relationship with sandalwood and its propagules and as a result may strongly influence the recruitment and spatial distribution of sandalwood by reducing the number of seeds and seedlings available for dispersal and regeneration, respectively. Woylies are both pre and postdispersal predators consuming seeds under the parent trees, seeds that have been dispersed away from the parent trees and buried and germinating seeds. Evidence has been provided to support the early anecdotal reports that woylies use emerging seedlings as 'markers' to locate buried seeds. This is the first study of its type in Australia to demonstrate, unequivocally, the consequences of the loss of a key seed-dispersal vector, the woylie, on sandalwood recruitment and regeneration in Dryandra Woodland. Through seed-caching and seed-predation, woylies have been shown to substantially alter the fate and distribution of sandalwood seeds and seedlings. The loss of woylies from 97% of their former range will undoubtedly have serious implications for sandalwood and possibly other plant species. In losing this seed-dispersing and seed-caching animal from our ecosystems we have lost a keystone species. The woylie sandalwood interaction seems to have shaped the morphology of the sandalwood seed and fruiting phenology by making the propagules attractive and rewarding to the woylie. In this way both the woylie and sandalwood benefit from this interaction. Such close interactions have a positive impact on the health and functioning of ecosystems and landscapes.