Threatened stick-nest rats preferentially eat invasive boxthorn rather than native vegetation on Australia’s Reevesby Island

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Many of these are degraded novel ecosystems invaded by African boxthorn (Lycium ferocissimum), a weed of national significance. However, L. conditor does not appear to be negatively impacted by the presence of boxthorn, raising the question of how the two species co-exist. Aims: To understand how L. conditor uses African boxthorn, we evaluated dietary composition of L. conditor on parts of Reevesby Island by comparing consumption of invasive boxthorn with that of native vegetation. Methods: We identified three key vegetation types on the centre of the island and used point-intercept vegetation surveys to estimate relative availability of plant species in each. We then used micro-histological faecal analysis to estimate the proportions of each species in the diet of L. conditor, and quantified plant species selection using selection ratios (use/availability). Key results : Qualitative evidence of L. conditor activity suggested it was mostly confined to vegetation with greater abundance of boxthorn than the other vegetation types (13.5%, compared to 5.7% total sampled vegetation). Furthermore, African boxthorn comprised of 51.7% of the faecal plant content and 11.8% of total sampled vegetation, resulting in a selection ration for boxthorn of 4.4. Native species that appeared to be favoured food sources of L. conditor included Olearia axillaris, Myoporum insulare and Enchylaena tomentosa. Conclusions : Stick-nest rats of Revesby Island demonstrate a clear preference for African boxthorn, both in terms of diet (tested quantitatively) and nesting (from previous research and field observations). Implications : The strong preference of stick-nest rats for a declared noxious weed as its main food source and persistence of stick-nest rats on Reevesby Island requires consideration with regards to vegetation management on all islands where L. conditor occurs. More broadly, it highlights that some elements of novel ecosystems may have unexpected positive impacts on parts of original ecosystems. Figures Figure 1 Figure 2 Figure 3 Figure 4 Summary Text Greater stick-nest rats are extinct on the Australian mainland but persist on small islands, some of which have been invaded by noxious African boxthorn. Using vegetation surveys and faecal analysis on Reevesby Island, South Australia, we found that the species preferences boxthorn for feeding and probably also for nesting. This highlights the need for nuanced appraisals of vegetation in novel ecosystems in the context of small mammal conservation. Introduction Novel ecosystems are an ensemble of organisms that are both self-sustaining and without historical precedent (Hobbs et al. 2013). They develop naturally as a result of biotic or abiotic changes, such as climate change, dispersal events, and the merging of continents. Humans have also caused the development of novel ecosystems, and at an unprecedented rate (Hobbs et al. 2009). As novel ecosystems are defined by the presence of new (and usually invasive) organisms, they are synonymous with loss of biodiversity and reduced ecosystem value and function. This has certainly been the case in Australia (Carr 1993; Centre for Agriculture and Bioscience International 2022; Dickman 1996; Gross 1995), where invasions are known to negatively impact on ecosystem function, with high economic costs (Bradshaw et al. 2021; Carneiro et al. 2024). Unfortunately, returning ecosystems to their pre-invasion state is not always possible (Hobbs et al. 2009). However, invasive species may be useful to supplement natural ecosystems in cases where functionally equivalent native organisms and processes have been impacted (Packer et al. 2016). This is particularly the case where invasive plant species dominate novel ecosystems yet play important roles in maintaining ecosystem function. Examples of this include the problem of how to manage the invasive tamarisk shrub in the US, where biocontrol measures have impacted on bird and insect fauna (Carothers et al. 2020); the dune-stabilising function of invasive sedges (Wootton et al. 2005); or the documented benefit of invasive plants on pollinating insects (Kovács-Hostyánszki et al. 2022). Because many invasive plants outcompete the surrounding vegetation through dense growth, they can benefit vertebrates by providing vegetation cover (Dutra et al. 2011). This is also the case in Australia, for example where dense native but invasively expanding shrublands can be beneficial for small native mammals, possibly due to providing suitable undergrowth cover (Hradsky et al. 2015; Law et al. 2016). Some of these impacts can be restricted to just a single species; an example is the case of the highly invasive blackberry which aiding bandicoots and other small mammals by providing a functional equivalent to the vegetation structure present in intact native ecosystems (Packer et al. 2016). Despite the well-known negative impacts of invasive weeds to biodiversity and ecosystem values (Hoffmann and Broadhurst 2016; Pimentel et al. 2005; Pyšek and Richardson 2010; Williams and West 2000), it is therefore important to assess if – and how – an invasive species benefits a native ecosystem or species. A potential new case of a single species benefiting from a plant invasion is the Near-threatened native Australian greater stick-nest rat (genus Leporillus conditor ). L. conditor is the only survivor of two Leporillus species. Both went extinct on the Australian mainland soon after European invasion due to the introduction of feral predators, habitat degradation caused by livestock grazing, and competition with introduced herbivores (Copley 1999a). However, a single natural population of L. conditor was discovered on the Franklin Islands, located in the Nuyt’s Archipelago off the west coast of Eyre Peninsula (Jones 1922). This was subsequently used as a source population for ten translocation efforts, as part of the stick-nest rat recovery plan (Copley 1995; Short et al. 2019). Successful translocations occurred in novel ecosystems on St Peter Island and Reevesby Island in South Australia, and Salutation Island in Western Australia, which are free from feral predators such as cats and foxes (Copley 1999a). The success of this recovery plan resulted in the down-listing of this species’ conservation status from ‘Endangered’ in 1996 to ‘Vulnerable’ in 2008 to ‘Near Threatened’ in 2016, under the IUCN Red List of Threatened Species (Short et al. 2018). The St Peter Island and Reevesby Island translocation sites are novel ecosystems that have been changed substantially by the introduction of cats (eradicated on Reevesby Island in the 1980s), over a century of sheep grazing, farming, and introduction of a range of weeds (Robinson et al. 1996b). The change in vegetation and legacy impacts of grazing are a concern for the survival of the species on the island (Robinson et al. 1996a). One of the most successful invasive weeds on Reevesby Island is African boxthorn, Lycium ferocissimum (Robinson et al. 1996b), which is classed as a Weed of National Significance due to its invasiveness, potential for spread, and economic and environmental impacts (Noble et al. 2014). L. ferocissimum forms dense thickets that can block access to water points and provide shelter for invasive animals, such as rabbits and foxes. Additionally, its invasive nature allows it to encroach on the nesting sites of native birds and mammals and displace native plant species that provide essential ecosystem functions (Centre for Agriculture and Bioscience International 2022). It is known to interfere with seal breeding on beaches in the Recherche Archipelago, Western Australia (Gross 1995), as well as penguin breeding on Motunau Island in New Zealand (Centre for Agriculture and Bioscience International 2022). However, the species is also a potential case for benefitting a single vertebrate species because it was shown to provide better protection for little Penguins against cats and foxes than available native vegetation for (Woehler et al. 2021). There is some evidence that L. conditor might be another beneficiary of L. ferocissimum infestation on the islands it inhabits. is so extensive that more stick-nest rats now live in boxthorn-infested novel ecosystems than in areas still dominated by native trees and shrubs (Short et al. 2019). Copley (1999b) found that stick-nests were most numerous around the bases of L. ferocissimum bushes, and, when the population reached its peak in 1997, multiple stick platforms were evident amongst the higher branches of many of these bushes (Copley 1999b; Short et al. 2019). This may be a strong sign of preference, as raised shelters are likely to be more difficult to build and more exposed to the elements and avian predators. Stick-nest rats may also favour L. ferocissimum on St Peter Island, as a 2006 survey of pitfall traps and Elliott traps at St Peter found that L. conditor were most abundant in localised patches of L. ferocissimum and Atriplex paludosa , despite this being a small minority of the island's vegetation (Copley 1999b; Short et al. 2019; Stewart 1996). While the extensive benefit of L. ferocissimum to L. conditor as a shelter is indicated by previous studies, it is important to also assess if it provides as suitable source of nutrition. This is particularly important in small areas such as Reevesby Island, where food is a critical resource and choices are likely to be limited. Any introduced species of herbivorous animal will need to adapt its diet to take advantage of the plant species available and this may differ considerably from the preferences in its original habitat. Given the uncertainty if the outcome of such introductions, it is therefore important to be able to quantify the food selections made. In attempting to reconstruct the diet, including the potential use of invasive plant species, of L. conditor , microhistological faecal analysis is an ideal method. This is an established technique for rodent diet analysis (Bergstrom 2013; Castleberry et al. 2002; Khanam et al. 2015; McIntire 1989; McMurry et al. 1993; Newmaster et al. 2013; Soininen et al. 2013), which relies on the identification of species-specific characteristics of plant cuticle in faecal samples. Cuticle is the waxy, non-cellular layer that conforms to the surface of a terrestrial leaf epidermis, protecting it from desiccation. While most leaf cells are denatured by the digestion process, much of the cuticle remains, still bearing the imprint of the leaf epidermis (Jones and Krockenberger 2007) which can be diagnostic for the plant material eaten. Microhistological faecal analysis has the benefit of being inexpensive and non-invasive to the animals. It has some risk of bias due to the differential digestion of plants with cuticles with varying capacities for preservation (Jones and Krockenberger 2007; Norbury 1988). However, it provides strong positive evidence of consumption, and is generally considered to be a reliable technique for most plant species (Anthony and Smith 1974; Ellis et al. 1999; Khanam et al. 2015; Norbury 1988; Pareja et al. 2021; Todd and Hansen 1973). In this study, we combine point intercept vegetation surveys with microhistological investigations of the degree to which L. ferocissimum is used by the Reevesby Island L. conditor population. This allows us to ask what proportions of native plants and L. ferocissimum are available in areas of L. conditor activity, and whether this availability is reflected in the feeding patterns of L. conditor and hence to understand the risks, and potential opportunities, of boxthorn infestation on this Vulnerable Australian native. Material and Methods Study site and history Reevesby Island is located at 34.534°S and 136.281°E in off the South Australian coast (refer to figure 1). It features four rocky lobes linked by narrow sandbars. L. conditor was originally present on the island, but went extinct before its reintroduction (Pedler and Copley 1993). While the cause and timing of this population's extinction remain unclear, it is thought to be linked to the introduction of feral cats ( Felis catus ) in the 1830s (Dickman 1996). The island was used for agricultural purposes and sheep grazing from 1838, until it was designated a conservation park in 1974. By this time the landscape had been extensively altered (Robinson et al. 1996a). Today, Reevesby Island is littered with abandoned farming equipment and populated by invasive plant species, including pasture grasses and Lycium ferocissimum . However, natives such as Myoporum insulare , Atriplex paludosa , and Threlkeldia diffusa , are reclaiming the deserted pastures (Robinson et al., 1996). The eradication of the feral cat population in the 1980s facilitated the translocation of an L. conditor population in 1990. Annual monitoring of the population then took place from 1992 to 2000. This revealed that the population peaked in 1998, with numbers fluctuating in boom-bust cycles between 1000 and 5000 individuals (Copley 1999b). Figure 1 here Field Design We use the word “nest” to describe any bush (or dense vegetation cluster) showing signs of frequent and sustained use or occupancy by L. conditor (for examples, see Fig. 2) These signs include: an abundance of faecal matter and multiple sets of fresh tracks, in addition to signs of gardening, such as the maintenance of an entrance and the construction and maintenance of “highways” (many sets of tracks, following a distinct path that has, in places, been carved, like a tunnel, through the branches of surrounding bushes). Sampling was carried out under Government of South Australia Permit to Undertake Scientific Research Number A26856-1. Figure 2 here To understand where L. conditor were living, we searched the area 2.3km north of the old homestead (-34.539°S; 136.276°E) in search of signs of stick-nest rat activity and marked the coordinates of all the nests we found (see figure 1). This methodology allowed us to rapidly identify the areas that L. conditor favoured and the areas they did not. We found that L. conditor activity was highest on a patch of vegetation sheltered in the sand dunes on the sand bar connecting the southernmost lobes of the island, and a patch of very similar vegetation on the sandbar connecting the southern lobes to the northern lobes. We focused our efforts on the more accessible, densely populated area between the two southern lobes. Vegetation Survey To assess the vegetation available to L. conditor on Reevesby Island, the island was divided into three distinct vegetation communities by identifying visual differences in satellite imagery. During the survey, it was then confirmed that each vegetation type contained a distinctive subset of plant species (Table 1). The vegetation types were then mapped using satellite imagery using the labels “Vegetation type 1-3” (see Fig. 1; description of their composition is in the results). Within each community, vegetation cover was measured as a proxy for availability using point-intercept plots, based on the Ausplots manual (White et al. 2012) as a guide. GPS points were located and marked using google maps on a Google Pixel 2 mobile device. These data points are accurate to approximately 5-10 meters. Therefore, to ensure accurate length and distances of transect lines on the ground, we relied on measuring tape as described below. Plot 1 was designed to cover an area of high Leporillus activity. Using Google Maps satellite imagery, we placed a 50x50 meter plot over this patch of vegetation and generated GPS coordinates for each corner. We then located each corner and marked out the plot using pegs and 100-meter tapes. We then used 50 meter tapes to create six parallel transects within this plot, each 50 meters long and spaced 10 meters apart, oriented in a north-south direction. Note that, while the work was done within the 50x50 GPS-based plot, the distances of the transect lines over the ground were shorter because the landscape relief shortened the distances over-ground relative to the two-dimensional satellite image (as visible by the rectangular, rather than square, shape of the transect area in Fig. 1). Measurement points or “intersects” were established at 1-meter intervals along each transect. At each “intersect”, a vertical 5-meter staff was positioned, and every plant species touching the staff was recorded. The height at which each species made contact was also noted, recording only the maximum height in instances of multiple contacts by one species. Plots 2 and 3 contained vegetation communities near Plot 1, but with no evidence of L. conditor activity. These were 20x50 metres in size, and consisted of three parallel 50 metre transects, spaced ten metres apart. Plot 2 ran parallel with the beach in the north-south direction, on the eastern side of Plot 1, for the purpose of sampling Vegetation type 2. Plot 3 ran in an east-west direction, on the southern side of Plot 1, and was selected for the purpose of sampling Vegetation type 3. Specimen samples To identify plant cuticle in L. conditor faecal pellets, we created a reference dataset of cuticle morphologies from plants found on Reevesby island. Plant vouchers for the reference data set were collected, labelled, and pressed according to the vouchering method described in Ausplots (White et al. 2012). Plants were identified using standard identification books (Berkinshaw 2010; Saunders 2018) and a list of plant species on Reevesby Island was generated using NatureMaps 3.0. Additional tissue samples for each plant species (leaves and stems, as well as fruits and/or flowers where available) were labelled and stored in vials at the University of Adelaide in 100% ethanol. Leaf and flower tissues were processed and made into slides using the protocol outlined in Storr (1961). A minimum of ten faecal pellets were collected per nest. At each nest, fresh faecal pellets were collected using forceps, labelled with the nest number and location and stored dry in paper envelopes. Pellets were considered fresh if they were moist, malleable, and dark in appearance. The first step was to wash faecal pellets in cold reverse-osmosis treated water to minimise contamination. The disaggregation procedure conventionally dissolves the faecal pellets in warm water (McCarthy et al. 1996; Pearson and Dodson 1993). However, this method proved ineffective for fresh faecal pellets, resulting in breakage of cuticle, as well as being slow and labour intensive. Instead, we developed a more reliable protocol that resembled the treatment of the reference cuticles, as follows: Faecal pellets were placed into labelled test tubes with a mixture of two parts 35% hydrogen peroxide (w/v) and one part 80% ethanol (v/v) at 90°C until the pellet dissolved. It was often necessary to use forceps to gently pull the pellet apart, breaking the mucous membrane coating. Any remaining aggregation, usually due to adhesion to mucous membrane, is easily dealt with using light water pressure and a small brush over a sieve. The remaining material is a combination of plant cuticle, seeds, small pieces of wood and bark, lignin coils, small rocks, and other plant material, such as mesophyll. This is then washed and stained using toluidine blue for one-two minutes and rinsed again. The dissolved scat is then placed in a petri dish to be manually cleaned and sorted under a dissecting microscope. Seeds and woody material were separated, labelled, and placed in long-term storage. Once cleaned and sorted, all cuticle was transferred to microscope slides, smoothed flat with a brush, covered with a drop of phenol glycerine jelly warmed to 60°C and then covered with glass coverslips, and inspected under a AX70 (Olympus, Australia) microscope with a UC50 (Olympus, Australia) camera attached, and CellSens (Olympus, Australia) image software. Any image stacking was done using Helicon Focus 7.6.4. Plant cuticle from the scat material was prepared and identified with reference to plant voucher slides. It was quantified by counting square millimetres covered of each species on each slide using a microscopic grid. This was then calculated into percentage values for further analysis. Identification of cuticle To determine what plant species L. conditor were consuming, it was necessary to have a systematic method for comparing the fragments of cuticle in the faecal pellets with the reference cuticle, for identification. Due to the fragmented nature of the digested cuticle, we sought multiple identification characters to ensure that we correctly identified as much of the cuticle as possible. The key identifiers used in this study were differential absorption of stain, unspecialised epidermal cells, stomata, subsidiary cells and trichomes. The methods for identifying cuticle based on differential absorption of stain and unspecialised epidermal cells are outlined in Storr (1961), and Metcalfe and Chalk (1950), respectively. We measured identifiers for each species and created an identification table for key plant species on the island (Supplementary Material 1). Statistics To understand the selective consumption of plants by L. conditor on Reevesby Island, we used Design I, described by Manly (2002) as a statistical design for quantifying habitat use and availability at the population level. The function for analysing Design I data, widesI, tests resource selection using Pearson's chi-squared test (χ2), and a log-likelihood chi-squared test. Pearson's chi-squared test tests a null hypothesis stating that that L. conditor select plants at random, meaning the portion of plant species represented in scats is consistent with that in the environment (allowing for a degree of error that is dependent on variation and sample size). The log-likelihood chi-squared test computes the goodness of fit of the null hypothesis vs the alternative hypothesis based on the ratio of their likelihoods for each plant species. If the null hypothesis is supported by the observed data, the two likelihoods should not differ by more than sampling error. Estimation of selective bias was estimated using the Manly selectivity measure (wi) with Bonferroni confidence intervals proposed by Manly et al. (2003): w i = o i /π i where oi is the proportion of plant species present in scats and πi is the proportion of available plant species in category i (plant species). The Selection ratio for a species is therefore: If the plant species is used (and preserved) in proportion to their availability, then w = 1. If wi > 1, the plant species is consumed or preserved more than expected by unbiased selection and preservation (preferred). The resource selection function (wi) was computed using adehabitatHS version 0.3.14 package with design I data type (Calenge 2006)in the R computing environment 3.6.1 (R Development Core Team 2019). Results For all raw data and code, refer to Supplementary Material 2 and 3, respectively. Three vegetation types were present in the areas we surveyed (see Methods). Vegetation type 1 is the least abundant within the study area. It is located where there is some protection from the strong ocean winds and salt spray. The dominant species are Myoporum insulare and Lycium ferocissimum (as medium to large bushes) and Muehlenbeckia gunnii and Tetragonia implexicoma (as ground cover and in the canopy as creepers/vines). Leporillus faecal pellets and tracks are a common occurrence within patches of this vegetation type, but rarely found outside of them. Vegetation type 2 is the most prevalent in the study area, occupying the exposed dunes. It is defined by the dominance of Leucophyta brownii and Olearia axillaris (as low shrubs) and Spinifex hirsutus (tufty grass). There is a substantial variation in density within this group, however, dominant species remain a fixed parameter. Due to accessibility and time constraints this vegetation type was sampled in a low-density area. Vegetation type 3 occupies low-lying salt and mud flats on the island in areas that are likely to become hypersaline due to inundation and subsequent evaporation of sea water. The dominant species are Salicornia quinqueflora (a small succulent shrub), Atriplex paludosa (marsh saltbush), Tetragonia implexicoma and Olearia axillaris . Table 1 here Plant species availability, use and selection by L. conditor Vegetation availability and use are shown in Figure 3, and the selection ratio is presented in Figure 4. L. ferocissimum is by far the most used resource, comprising 51.7% used material despite only representing 11.8% of the available vegetation in plot 1 and 10.43% of total sampled vegetation, as measured in the point intercept survey (Table 2). This gives it a selection ratio of 4.4 (Table 3). Figures 3&4 here Table 2 here Plant use Due to its abundant representation in the scat material, L. ferocissimum was broken into two subclasses based on cuticle type: “Leaf”, and “Flower”. Leaf cuticle comprised 62.2% of L. ferocissimum cuticle, and the remaining 37.9% can be attributed to flowers (Supplementary Materials 4). Atriplex paludosa (marsh saltbush), Carpobrotus rossii (Australian pig face), L. ferocissimum, Enchylaena tomentosa (ruby saltbush), Olearia axillaris (coastal daisy-bush), and Myoporum insulare (native juniper) have selection ratios >1, although this is not always statistically significant (Figure 4, Table 2). Tetragonia implexicoma (bower spinach) and Muehlenbeckia gunnii (coastal lignum) are the only species that seem to be selected against. They comprise 23.7% and 20.2% of the total available vegetation in plot 1, but only 1.8% and 0% of the used resources (Figure 3). Of all the available species, the succulents have the most fragile cuticle, and are the most susceptible to damage during digestion and lab processes. As such, it may be reasonable to class the “used” data for these species as missing, or incomplete. However, studies show that the effects of differential preservation are usually minor (Anthony and Smith 1974; Ellis et al. 1999; Khanam et al. 2015; Norbury 1988; Pareja et al. 2021; Todd and Hansen 1973) and, as the other plant cuticles are more robust, they are unlikely to be impacted by this secondary selective process. Discussion We found clear evidence of high use of the invasive Lycium ferocissimum (African boxthorn) by L. conditor (stick-nest rat), which the animals appear to strongly preference as a food source over other native vegetation. Although it was not quantified, the density of L. conditor activity in vegetation dominated by L. ferocissimum suggests that it is also the preferred habitat for L. conditor as already shown by previous research (Copley 1995; Short et al. 2019). Importantly, L. ferocissimum was the most strongly selected plant species and preferred even over succulent species such as the ruby saltbush, which form the majority of L. conditor’s diet in the comparatively natural or restored habitats such as the Franklin Islands and the Arid Recovery native mammal re-introduction site of northern South Australia (Copley 1999a; Read 1984; Robinson et al. 1996b; Ryan et al. 2003). It is possible that flowers are more palatable and could have increased selective use of L. ferocissimum . Flowers were abundant despite the collection period occurring in winter, outside the species’ usual summer flowering period in summer (Erkelenz 1993). However, it seems more likely that this preference is driven by the habitat choice of L. conditor , as the species seems to be a dietary generalist and consumes a variety of different plants, depending on location and season (Read 1984). . The benefit of L. ferocissimum as a food plant is probably accompanied by the further preference as shelter by L. conditor, although this was not quantified here . At the Arid Recovery native mammal re-introduction site (Moseby et al. 2011), L. conditor demonstrated a preference for low, dense shrubs as the base for nest sites (O’Neill 1999; Short et al. 2019), likely due to the protection they offer against predators, especially avian predators (Copley 1988; O’Neill 1999). While L. ferocissimum provides L. conditor with low, dense cover on Reevesby Island, so could other plants that grow within the same microhabitat. However, L. conditor may favour L. ferocissimum bushes because the large thorns provide more protection from predators than the plants that are native to the animal’s offshore island refuges. A preference for thorns and spines by L. conditor is supported by Short et al. (2018) who found that very prickly Acacia tetragonophylla bushes were the primary nesting shrubs used by L. conditor at Heirisson Prong (Introduced in 1999, last sighting in 2007), in Shark Bay, Western Australia. Despite the successful adaptation of L. conditor to boxthorn invasion, is difficult to judge the degree to which L. ferocissimum is beneficial in replacing pre-disturbance structural complexity and food resources on Reevesby Island, as there is no historical reference for precolonial vegetation on the island. However, the location of the only natural Leporillus population, on the Franklin Islands, offers an interesting comparison. Descriptions of the vegetation there appear to be very similar to Reevesby Island, except that they do not contain L. ferocissimum (Osborn 1922; Read 1984; Robinson et al. 1996b). Interestingly, on the Franklin Islands, stick-nest rats also show a clear preference for a habitat that is not part of the native vegetation, as population density of L. conditor appears to be highest among the granite slabs along the coastline, rather than more heavily vegetated areas. Areas of dense vegetation cover are also used, but appear to have lower population densities (Read 1984). It therefore remains unclear if L. conditor ’s preference for nesting in L. ferocissimum thickets is due to its efficiency as a shelter, as it is for other vertebrates (Dutra et al. 2011; Hradsky et al. 2015; Law et al. 2016; Packer et al. 2016); or whether it relates to a strong preference for L. ferocissimum as a food source. Evidence of stick-nest rats using native vegetation for food and habitat on Reevesby Island and the Franklin Islands (Read 1984) suggests that L. conditor on Reevesby Island are unlikely to require boxthorn for their survival. However, the fact that they appear not to favour native vegetation as habitat at either location emphasises the lack of information we have on this species. Part of the issue is that all translocation sites, and remnant natural populations, are located on the very fringe of L. conditor’s former distribution (Copley 1999a) and unlikely to represent their optimal environment. In addition, there is no evidence of a preference for thorned plants preserved in fossilised stick-nest rat middens, suggesting that more research is required as to the original habitat preferences of this species (Allen et al. 2000; McCarthy 1999; McCarthy and Head 2001; McCarthy et al. 1996; Pearson 1999; Pearson et al. 2001; Pearson and Betancourt 2002; Pearson and Dodson 1993; Webeck and Pearson 2005). Lack of good understanding of habitat preference is an important issue in translocations, particularly in Australia and surrounding Oceanian areas where translocation sites are often subject to extensive weed invasions (Morris et al. 2021). Another recent Australian example is the single remaining population of Shark Bay mouse ( Pseudomys gouldi) , which also has a broader range of suitable habitats than previously assumed (Palmer et al. 2024). The apparent beneficial effects for African boxthorn infestation on stick-nest rats are consistent with other observations of beneficial interactions between invasives with native ecosystem components (Azmi and Jennings 2013; Carothers et al. 2020; Packer et al. 2016). This is an important consideration when assessing the economic impact of invasions and their control, particularly when expensive but inefficient measures result in a removal of beneficial effects (Boltovskoy et al. 2022). In the case of stick-nest rats, this might be a particular issue in times of warming climates, which makes it particularly important for the species to have sufficient cover (Onley et al. 2022). However, the beneficial effects of improved plant cover from invasive species (Dutra et al. 2011) need to be balanced against the otherwise detrimental effect of these often very destructive species. It is also not clear if the benefits will persist when a novel ecosystem may has reached a stable state. African boxthorn is highly invasive and, if left unchecked, may continue to spread, displacing native vegetation, and resulting in decreased ecosystem diversity and function that might ultimately also affect stick-nest rats themselves. It is even possible that the feeding preferences of L. conditor facilitate the invasion of boxthorn because of its potential to spread the seed through its faeces, similar to observations that re-introductions of burrowing mammals can aid the spread of invasive plants (Palmer et al. 2020). This highlights the importance of considering the potential negative outcomes of translocation efforts in cases where translocated species interact with novel ecosystems in a way that is detrimental to overall ecosystem function (Palmer et al. 2020). Our study has shown an unexpected preference for invasive L. ferocissimum on Reevesby Island. Together with the species’ apparent preference for granite slabs on Franklin Island, this provide important clues as to the importance of vegetation type, terrain, and geology in identifying potential future translocation sites. Our lack of understanding on the historical habitat of stick-nest rats suggests that assessments of the pre-colonial landscapes of previous occurrence sites (such as including midden sites and record maps) will be important for future translocation success. Declarations Acknowledgments We thank field volunteers Baillie Trenwith, Holly Kraehe, Czes Klopotowski, and two anonymous reviewers. Declaration of Funding VW was funded by the Australian Research Council Centre of Excellence for Australian Biodiversity and Heritage (CE170100015) and Future Fellowship (FT180100634). Conflict of Interest The authors have no conflict of interest to declare Data availability statement The raw vegetation survey and cuticle identification data for this manuscript, and the code to replicate Figures 5 and 6, are available in the supplementary materials. 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The percentages in Plot 1 are the same as the percentages of available vegetation in Fig. 3. Species Plot 1 Plot 2 Plot 3 Total sampled vegetation Atriplex paludosa 0.54% 0.00% 17.53% 4.08% Carpobrotus rossii 1.08% 6.33% 0.00% 2.72% Dionella revoluta 0.54% 2.53% 2.06% 1.59% Enchylaena tomentosa 1.61% 0.00% 10.31% 2.95% Euphorbia paralias 0.00% 0.00% 0.00% 0.00% Ficinia nodosa 2.15% 0.00% 0.00% 0.91% Leucophyta brownii 5.38% 0.00% 1.03% 2.49% Lycium ferocissimum 11.83% 15.19% 0.00% 10.43% Muehlenbeckia gunnii 34.41% 3.16% 3.09% 16.33% Myoporum insulare 8.60% 24.05% 10.31% 14.51% Nitraria billardierei 0.00% 0.63% 0.00% 0.23% Olearia axillaris 7.53% 12.03% 13.40% 10.43% Rhagodia crassifolia 1.08% 0.63% 0.00% 0.68% Salicornia quinqueflora 0.00% 0.00% 20.62% 4.54% Senecio lautus 5.91% 0.63% 0.00% 2.72% Spinifex hirsutus 1.61% 1.27% 0.00% 1.13% Tetragonia implexicoma 17.20% 31.01% 16.49% 22.00% Threlkeldia diffusa 0.54% 1.90% 0.00% 0.91% Table 2: Outcome of Manly selection ratio design I in Plot I “Used”, total proportion of each species in the total faecal plant material; “available” represents the total proportion of each species in the point intercept surveys. Selection ratio is the product of “used” divided by “available”. Species Used available selection ratio standard error p-value Atriplex paludosa 0.007 0.005 1.355 1.433 0.804 Carpobrotus rossii 0.013 0.011 1.186 0.891 0.835 Dionella revoluta 0.000 0.005 0 - - Enchylaena tomentosa 0.034 0.016 2.089 1.243 0.381 Ficinia nodosa 0 0.022 0 - - Leucophyta brownii 0 0.054 0 - - Lycium ferocissimum 0.517 0.118 4.374 0.885 0.000 Muehlenbeckia gunnii 0 0.344 0 - - Myoporum insulare 0.161 0.086 1.874 0.466 0.061 Olearia axillaris 0.235 0.075 3.122 0.82 0.01 Rhagodia crassifolia 0 0.011 0 - - Senecio lautus 0.013 0.059 0.216 0.085 0.000 Spinifex hirsutus 0 0.016 0 - - Tetragonia implexicoma 0.02 0.172 0.116 0.031 0.000 Threlkeldia diffusa 0 0.005 0 - - Additional Declarations The authors declare no competing interests. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-1963287","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":136979429,"identity":"0dca9be1-6bc8-4e54-932b-0055b5a97da3","order_by":0,"name":"Annie Grace Kraehe","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA+klEQVRIiWNgGAWjYJCCAwxsElBmBRAzMzeQouUMSAsjYS0MDGxQmrENTOLXwj+79+CBD2UW9vLRhx9++DivNpq/HajlR8U2nFok7pxLODjjnETixnNpxpIztx3PnXGYsYGx58xt3NbcyDE4zNsmkWDYw2DGzLvtWG4DUAszYxtuLfIgLX/bJOwNe9i/Mf+dcyx3PiEtBiAtjG0SjPN5eMyAYVWTu4GQFkOgloM9QL9s4OEpluw5diB3I1DLQXx+kbuRY/zhR1mdvXwP+8YPP2rqcuedP3zwwY8KPN6Hu/AAmDoMJg8QVg8E8g1gqo4oxaNgFIyCUTCyAAA89F37Y7903QAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0003-3900-006X","institution":"University of Adelaide","correspondingAuthor":true,"prefix":"","firstName":"Annie","middleName":"Grace","lastName":"Kraehe","suffix":""},{"id":136979430,"identity":"2436beec-165a-404e-a222-33a7b41578f6","order_by":1,"name":"Vera Weisbecker","email":"","orcid":"","institution":"Flinders University","correspondingAuthor":false,"prefix":"","firstName":"Vera","middleName":"","lastName":"Weisbecker","suffix":""},{"id":136979431,"identity":"081bee03-4376-4fab-bfa9-63b0e7b29f6c","order_by":2,"name":"Robert Hill","email":"","orcid":"","institution":"The University of Adelaide","correspondingAuthor":false,"prefix":"","firstName":"Robert","middleName":"","lastName":"Hill","suffix":""},{"id":136979432,"identity":"c5e4e42d-f61b-4b1f-bf2c-d3c2879aae03","order_by":3,"name":"Kathryn Hill","email":"","orcid":"","institution":"The University of Adelaide","correspondingAuthor":false,"prefix":"","firstName":"Kathryn","middleName":"","lastName":"Hill","suffix":""}],"badges":[],"createdAt":"2022-08-15 09:02:30","currentVersionCode":2,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-1963287/v2","doiUrl":"https://doi.org/10.21203/rs.3.rs-1963287/v2","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1071/WR23140","type":"published","date":"2024-07-25T00:46:20+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":57024973,"identity":"bbf67f60-26dd-4f1f-ab06-d4a27473d384","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"bmp","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":278661,"visible":true,"origin":"","legend":"\u003cp\u003eMap of Reevesby Island showing Transect Lines, Nest Locations, and Vegetation Zones. Areas that were not explored have been left blank. Made in ArcGIS 3.1.1\u003c/p\u003e","description":"","filename":"Fig1map.bmp","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/4c6e15f348e9d155c386f7bd.bmp"},{"id":57024979,"identity":"c57b48be-b6b2-4d2f-a3fe-a2d20d638bf9","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2230899,"visible":true,"origin":"","legend":"\u003cp\u003eExample of dense\u003cem\u003e Leporillus\u003c/em\u003e nests in large \u003cem\u003eLycium ferocissimum \u003c/em\u003ebush\u003c/p\u003e","description":"","filename":"Fig2nests.png","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/1ca323c087fe7b550909b20a.png"},{"id":57024980,"identity":"3f1c8408-3859-4a3b-aa8b-1ae07ae5a7df","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":93527,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage of Available Vegetation (as per relative abundance, see methods; green) vs Use (purple) by\u003cem\u003e Leporillus conditor\u003c/em\u003e. Standard error shown in black. Data not shown for species not found in Plot 1.\u003c/p\u003e","description":"","filename":"Fig3percentages.png","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/42189937c57ab45c86286f5f.png"},{"id":57025445,"identity":"e324a6ed-212a-4174-a392-2bf8cb18145f","added_by":"auto","created_at":"2024-05-23 14:51:02","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":69502,"visible":true,"origin":"","legend":"\u003cp\u003eSelective use of plant species by\u003cem\u003e Leporillus conditor\u003c/em\u003e in Plot 1. Graph showing selection ratios on the X-axis and plant species on the Y-axis, with 95% confidence intervals as horizontal lines. The dotted vertical line marks a selection ratio of 1. Species on the right side of the dotted line have a selection ratio \u0026gt;1, which means they statistically more Used than they are Available. This indicates selective use by\u003cem\u003eLeporillus conditor.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"Fig4selectionRatio.png","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/fd7b73d780990648a1f8535d.png"},{"id":61039493,"identity":"e46171a4-66ce-4d6c-8ff9-95c9f90feb09","added_by":"auto","created_at":"2024-07-25 00:46:27","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4329575,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/d193ef0c-d366-4fa9-b9fd-394673fdb560.pdf"},{"id":57024972,"identity":"5f147b96-4b95-4571-a039-8d32702f0ab1","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":503258,"visible":true,"origin":"","legend":"","description":"","filename":"Supp1ID.docx","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/9a48bc661c618ee86c01cf8a.docx"},{"id":57024974,"identity":"842a5784-87d8-432d-99c5-f639a875e992","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"xlsx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":56243,"visible":true,"origin":"","legend":"","description":"","filename":"Supp2Reevesbyvegetationandsticknestrats.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/0b8da02d0a635e1815362f1f.xlsx"},{"id":57024977,"identity":"9cf2472a-590b-46f3-9852-1a60c747122b","added_by":"auto","created_at":"2024-05-23 14:43:02","extension":"r","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":4812,"visible":true,"origin":"","legend":"","description":"","filename":"Supp3Kraeheetalcode.r","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/ce16fd47e8c247b513a55d56.r"},{"id":57025444,"identity":"c829dc11-e5d5-4bb2-b1d3-716e5f16580d","added_by":"auto","created_at":"2024-05-23 14:51:02","extension":"docx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":15961,"visible":true,"origin":"","legend":"","description":"","filename":"Supp4surfaceleafbreakdown.docx","url":"https://assets-eu.researchsquare.com/files/rs-1963287/v2/4ff3ebbd15111f946268cf3d.docx"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eThreatened stick-nest rats preferentially eat invasive boxthorn rather than native vegetation on Australia’s Reevesby Island\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Summary Text","content":"\u003cp\u003eGreater stick-nest rats are extinct on the Australian mainland but persist on small islands, some of which have been invaded by noxious African boxthorn. Using vegetation surveys and faecal analysis on Reevesby Island, South Australia, we found that the species preferences boxthorn for feeding and probably also for nesting. This highlights the need for nuanced appraisals of vegetation in novel ecosystems in the context of small mammal conservation.\u0026nbsp;\u003c/p\u003e"},{"header":"Introduction","content":"\u003cp\u003eNovel ecosystems are an ensemble of organisms that are both self-sustaining and without historical precedent\u0026nbsp;(Hobbs\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2013). They develop naturally as a result of biotic or abiotic changes, such as climate change, dispersal events, and the merging of continents. Humans have also caused the development of novel ecosystems, and at an unprecedented rate\u0026nbsp;(Hobbs\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2009). As novel ecosystems are defined by the presence of new (and usually invasive) organisms, they are synonymous with loss of biodiversity and reduced ecosystem value and function. This has certainly been the case in Australia\u0026nbsp;(Carr 1993; Centre for Agriculture and Bioscience International 2022; Dickman 1996; Gross 1995), where invasions are known to negatively impact on ecosystem function, with high economic costs\u0026nbsp;(Bradshaw\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2021; Carneiro\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2024). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eUnfortunately, returning ecosystems to their pre-invasion state is not always possible\u0026nbsp;(Hobbs\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2009). However, invasive species may be useful to supplement natural ecosystems in cases where functionally equivalent native organisms and processes have been impacted\u0026nbsp;(Packer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016). This is particularly the case where invasive plant species dominate novel ecosystems yet play important roles in maintaining ecosystem function. Examples of this include the problem of how to manage the invasive tamarisk shrub in the US, where biocontrol measures have impacted on bird and insect fauna\u0026nbsp;(Carothers\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2020); the dune-stabilising function of invasive sedges\u0026nbsp;(Wootton\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2005); or the documented benefit of invasive plants on pollinating insects\u0026nbsp;(Kov\u0026aacute;cs-Hosty\u0026aacute;nszki\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2022).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eBecause many invasive plants outcompete the surrounding vegetation through dense growth, they can benefit vertebrates by providing vegetation cover\u0026nbsp;(Dutra\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2011). This is also the case in Australia, for example where dense native but invasively expanding shrublands can be beneficial for small native mammals, possibly due to providing suitable undergrowth cover\u0026nbsp;(Hradsky\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2015; Law\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016). \u0026nbsp;Some of these impacts can be restricted to just a single species; an example is the case of the highly invasive blackberry which aiding bandicoots and other small mammals by providing a functional equivalent to the vegetation structure present in intact native ecosystems\u0026nbsp;(Packer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016). Despite the well-known negative impacts of invasive weeds to biodiversity and ecosystem values\u0026nbsp;(Hoffmann and Broadhurst 2016; Pimentel\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2005; Py\u0026scaron;ek and Richardson 2010; Williams and West 2000), it is therefore important to assess if \u0026ndash; and how \u0026ndash; an invasive species benefits a native ecosystem or species.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA potential\u0026nbsp;new case of a single species benefiting from a plant invasion is the Near-threatened native Australian greater stick-nest rat (genus\u003cem\u003e\u0026nbsp;Leporillus conditor\u003c/em\u003e). \u003cem\u003eL. conditor\u003c/em\u003e is the only survivor of two \u003cem\u003eLeporillus\u003c/em\u003e species. Both went extinct on the Australian mainland soon after European invasion due to the introduction of feral predators, habitat degradation caused by livestock grazing, and competition with introduced herbivores\u0026nbsp;(Copley 1999a). However, a single natural population of \u003cem\u003eL. conditor\u003c/em\u003e was discovered on the Franklin Islands, located in the Nuyt\u0026rsquo;s Archipelago off the west coast of Eyre Peninsula (Jones 1922). This was subsequently used as a source population for ten translocation efforts, as part of the stick-nest rat recovery plan\u0026nbsp;(Copley 1995; Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2019). Successful translocations occurred in novel ecosystems on St Peter Island and Reevesby Island in South Australia, and Salutation Island in Western Australia, which are free from feral predators such as cats and foxes\u0026nbsp;(Copley 1999a). The success of this recovery plan resulted in the down-listing of this species\u0026rsquo; conservation status from \u0026lsquo;Endangered\u0026rsquo; in 1996 to \u0026lsquo;Vulnerable\u0026rsquo; in 2008 to \u0026lsquo;Near Threatened\u0026rsquo; in 2016, under the IUCN Red List of Threatened Species\u0026nbsp;(Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2018).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe St Peter Island and Reevesby Island translocation sites are novel ecosystems that have been changed substantially by the introduction of cats (eradicated on Reevesby Island in the 1980s), over a century of sheep grazing, farming, and introduction of a range of weeds\u0026nbsp;(Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996b). The change in vegetation and legacy impacts of grazing are a concern for the survival of the species on the island\u0026nbsp;(Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996a). One of the most successful invasive weeds on Reevesby Island is African boxthorn, \u003cem\u003eLycium ferocissimum\u003c/em\u003e (Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996b), which is classed as a Weed of National Significance due to its invasiveness, potential for spread, and economic and environmental impacts\u0026nbsp;(Noble\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2014).\u003cem\u003e\u0026nbsp;L. ferocissimum\u0026nbsp;\u003c/em\u003eforms\u0026nbsp;dense thickets that can block access to water points and provide shelter for invasive animals, such as rabbits and foxes. Additionally, its invasive nature allows it to encroach on the nesting sites of native birds and mammals and displace native plant species that provide essential ecosystem functions\u0026nbsp;(Centre for Agriculture and Bioscience International 2022). It is known to interfere with seal breeding on beaches in the Recherche Archipelago, Western Australia\u0026nbsp;(Gross 1995), as well as penguin breeding on Motunau Island in New Zealand\u0026nbsp;(Centre for Agriculture and Bioscience International 2022). However, the species is also a potential case for benefitting a single vertebrate species because it was shown to provide better protection for little Penguins against cats and foxes than available native vegetation for\u0026nbsp;(Woehler\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThere is some evidence that \u003cem\u003eL. conditor\u003c/em\u003e might be another beneficiary of \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003einfestation on the islands it inhabits. \u0026nbsp;is so extensive that more stick-nest rats now live in boxthorn-infested novel ecosystems than in areas still dominated by native trees and shrubs\u0026nbsp;(Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2019). Copley (1999b) found that stick-nests were most numerous around the bases of \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003ebushes, and, when the population reached its peak in 1997, multiple stick platforms were evident amongst the higher branches of many of these bushes (Copley 1999b; Short \u003cem\u003eet al.\u003c/em\u003e 2019). This may be a strong sign of preference, as raised shelters are likely to be more difficult to build and more exposed to the elements and avian predators. Stick-nest rats may also favour \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003eon St Peter Island, as a 2006 survey of pitfall traps and Elliott traps at St Peter found that \u003cem\u003eL. conditor\u003c/em\u003e were most abundant in localised patches of \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003eand \u003cem\u003eAtriplex paludosa\u003c/em\u003e, despite this being a small minority of the island\u0026apos;s vegetation\u0026nbsp;(Copley 1999b; Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2019; Stewart 1996). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWhile the extensive benefit of \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003eto \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eas a shelter is indicated by previous studies, it is important to also assess if it provides as suitable source of nutrition. This is particularly important\u0026nbsp;in small areas such as Reevesby Island, where food is a critical resource and choices are likely to be limited. Any introduced species of herbivorous animal will need to adapt its diet to take advantage of the plant species available and this may differ considerably from the preferences in its original habitat. Given the uncertainty if the outcome of such introductions, it is therefore important to be able to quantify the food selections made.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn attempting to reconstruct the diet, including the potential use of invasive plant species, of \u003cem\u003eL. conditor\u003c/em\u003e, microhistological faecal analysis is an ideal method. This is an established technique for rodent diet analysis\u0026nbsp;(Bergstrom 2013; Castleberry\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2002; Khanam\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2015; McIntire 1989; McMurry\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1993; Newmaster\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2013; Soininen\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2013), which relies on the identification of species-specific characteristics of plant cuticle in faecal samples. Cuticle is the waxy, non-cellular layer that conforms to the surface of a terrestrial leaf epidermis, protecting it from desiccation. While most leaf cells are denatured by the digestion process, much of the cuticle remains, still bearing the imprint of the leaf epidermis\u0026nbsp;(Jones and Krockenberger 2007)\u0026nbsp;which can be diagnostic for the plant material eaten. Microhistological faecal analysis has the benefit of being inexpensive and non-invasive to the animals. It has some risk of bias due to the differential digestion of plants with cuticles with varying capacities for preservation\u0026nbsp;(Jones and Krockenberger 2007; Norbury 1988).\u0026nbsp;However,\u0026nbsp;it provides strong positive evidence of consumption, and\u0026nbsp;is generally considered to be a reliable technique\u0026nbsp;for most plant species\u0026nbsp;(Anthony and Smith 1974; Ellis\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1999; Khanam\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2015; Norbury 1988; Pareja\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2021; Todd and Hansen 1973).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn this study, we combine point intercept vegetation surveys with microhistological investigations of the degree to which \u003cem\u003eL. ferocissimum\u003c/em\u003e is used by the Reevesby Island \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003epopulation. This allows us to ask what proportions of native plants and \u003cem\u003eL. ferocissimum\u003c/em\u003e are available in areas of \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eactivity, and whether this availability\u003cem\u003e\u0026nbsp;\u003c/em\u003eis reflected in the feeding patterns of \u003cem\u003eL. conditor\u003c/em\u003e and hence to understand the risks, and potential opportunities, of boxthorn infestation on this Vulnerable Australian native.\u0026nbsp;\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003ch2\u003eStudy site\u0026nbsp;and history\u003c/h2\u003e\n\u003cp\u003eReevesby Island\u0026nbsp;is located at 34.534\u0026deg;S and 136.281\u0026deg;E in off the South Australian coast (refer to figure 1). It features four rocky lobes linked by narrow sandbars. \u003cem\u003eL. conditor\u003c/em\u003e was originally present on the island, but went extinct before its reintroduction\u0026nbsp;(Pedler and Copley 1993). While the cause and timing of this population\u0026apos;s extinction remain unclear, it is thought to be linked to the introduction of feral cats (\u003cem\u003eFelis catus\u003c/em\u003e) in the 1830s\u0026nbsp;(Dickman 1996). The island was used for agricultural purposes and sheep grazing from 1838, until it was designated a conservation park in 1974. By this time the landscape had been extensively altered\u0026nbsp;(Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996a). Today, Reevesby Island is littered with abandoned farming equipment and populated by invasive plant species, including pasture grasses and \u003cem\u003eLycium ferocissimum\u003c/em\u003e. However, natives such as \u003cem\u003eMyoporum insulare\u003c/em\u003e, \u003cem\u003eAtriplex paludosa\u003c/em\u003e, and \u003cem\u003eThrelkeldia diffusa\u003c/em\u003e, are reclaiming the deserted pastures (Robinson et al., 1996). The eradication of the feral cat population in the 1980s facilitated the translocation of an \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003epopulation in 1990. Annual monitoring of the population then took place from 1992 to 2000. This revealed that the population peaked in 1998, with numbers fluctuating in boom-bust cycles between 1000 and 5000 individuals\u0026nbsp;(Copley 1999b).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 1 here\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eField Design\u003c/h2\u003e\n\u003cp\u003eWe use the word \u0026ldquo;nest\u0026rdquo; to describe any bush (or dense vegetation cluster) showing signs of frequent and sustained use or occupancy by \u003cem\u003eL. conditor\u003c/em\u003e (for examples, see Fig. 2) These signs include: an abundance of faecal matter and multiple sets of fresh tracks, in addition to signs of gardening, such as the maintenance of an entrance and the construction and maintenance of \u0026ldquo;highways\u0026rdquo; (many sets of tracks, following a distinct path that has, in places, been carved, like a tunnel, through the branches of surrounding bushes). Sampling was carried out under Government of South Australia Permit to Undertake Scientific Research Number A26856-1.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 2 here\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTo understand where \u003cem\u003eL. conditor\u003c/em\u003e were living,\u0026nbsp;we searched the area 2.3km north of the old homestead (-34.539\u0026deg;S; 136.276\u0026deg;E) in search of signs of stick-nest rat activity\u0026nbsp;and marked the coordinates of all the nests we found (see figure 1). This methodology allowed us to rapidly identify the areas that \u003cem\u003eL. conditor\u003c/em\u003e favoured and the areas they did not.\u0026nbsp;We found that \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eactivity was highest on a patch of vegetation sheltered in the sand dunes on the sand bar connecting the southernmost lobes of the island, and a patch of very similar vegetation on the sandbar connecting the southern lobes to the northern lobes. We focused our efforts on the more accessible, densely populated area between the two southern lobes.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003ch4\u003eVegetation Survey\u003c/h4\u003e\n\u003cp\u003eTo assess the vegetation available to \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eon Reevesby Island, the island was divided into three distinct vegetation communities by identifying visual differences in satellite imagery. During the survey, it was then confirmed that each vegetation type\u0026nbsp;contained a distinctive subset of plant species (Table 1). The vegetation types were then mapped using satellite imagery using the labels \u0026ldquo;Vegetation type 1-3\u0026rdquo; (see Fig. 1; description of their composition is in the results). Within each community, vegetation cover was measured as a proxy for availability using point-intercept plots, based on the Ausplots manual\u0026nbsp;(White et al. 2012)\u0026nbsp;as a guide. GPS points were located and marked using google maps on a Google Pixel 2 mobile device. These data points are accurate to approximately 5-10 meters. Therefore, to ensure accurate length and distances of transect lines on the ground, we relied on measuring tape as described below.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePlot 1\u003c/strong\u003e was designed to cover an area of high \u003cem\u003eLeporillus\u003c/em\u003e activity. Using Google Maps satellite imagery, we placed a 50x50 meter plot over this patch of vegetation and generated GPS coordinates for each corner. We then located each corner and marked out the plot using pegs and 100-meter tapes. We then used 50 meter tapes to create six parallel transects within this plot, each 50 meters long and spaced 10 meters apart, oriented in a north-south direction. Note that, while the work was done within the 50x50 GPS-based plot, the distances of the transect lines over the ground were shorter because the landscape relief shortened the distances over-ground relative to the two-dimensional satellite image (as visible by the rectangular, rather than square, shape of the transect area in Fig. 1). Measurement points or \u0026ldquo;intersects\u0026rdquo; were established at 1-meter intervals along each transect. At each \u0026ldquo;intersect\u0026rdquo;, a vertical 5-meter staff was positioned, and every plant species touching the staff was recorded. The height at which each species made contact was also noted, recording only the maximum height in instances of multiple contacts by one species.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePlots 2 and 3\u0026nbsp;\u003c/strong\u003econtained vegetation communities near Plot 1, but with no evidence of \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eactivity. These were 20x50 metres in size, and consisted of three parallel 50 metre transects, spaced ten metres apart. Plot 2 ran parallel with the beach in the north-south direction, on the eastern side of Plot 1, for the purpose of sampling Vegetation type 2. Plot 3 ran in an east-west direction, on the southern side of Plot 1, and was selected for the purpose of sampling Vegetation type 3.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003ch4\u003eSpecimen samples\u003c/h4\u003e\n\u003cp\u003eTo identify plant cuticle in \u003cem\u003eL. conditor\u003c/em\u003e faecal pellets, we created a reference dataset of cuticle morphologies from plants found on Reevesby island. Plant vouchers for the reference data set were collected, labelled, and pressed according to the vouchering method described in Ausplots (White \u003cem\u003eet al.\u003c/em\u003e 2012). Plants were identified using standard identification books\u0026nbsp;(Berkinshaw 2010; Saunders 2018)\u0026nbsp;and a list of plant species on Reevesby Island was generated using NatureMaps 3.0. Additional tissue samples for each plant species (leaves and stems, as well as fruits and/or flowers where available) were labelled and stored in vials at the University of Adelaide in 100% ethanol. Leaf and flower tissues were processed and made into slides using the protocol outlined in Storr (1961).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA minimum of ten faecal pellets were collected per nest. At each nest, fresh faecal pellets were collected using forceps, labelled with the nest number and location and stored dry in paper envelopes. Pellets were considered fresh if they were moist, malleable, and dark in appearance. The first step was to wash faecal pellets in cold reverse-osmosis treated water to minimise contamination. The disaggregation procedure conventionally dissolves the faecal pellets in warm water\u0026nbsp;(McCarthy\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996; Pearson and Dodson 1993). However, this method proved ineffective for fresh faecal pellets, resulting in breakage of cuticle, as well as being slow and labour intensive. Instead, we developed a more reliable protocol that resembled the treatment of the reference cuticles, as follows: Faecal pellets were placed into labelled test tubes with a mixture of two parts 35% hydrogen peroxide (w/v) and one part 80% ethanol (v/v) at 90\u0026deg;C until the pellet dissolved. It was often necessary to use forceps to gently pull the pellet apart, breaking the mucous membrane coating. Any remaining aggregation, usually due to adhesion to mucous membrane, is easily dealt with using light water pressure and a small brush over a sieve. The remaining material is a combination of plant cuticle, seeds, small pieces of wood and bark, lignin coils, small rocks, and other plant material, such as mesophyll. This is then washed and stained using toluidine blue for one-two minutes and rinsed again. The dissolved scat is then placed in a petri dish to be manually cleaned and sorted under a dissecting microscope. Seeds and woody material were separated, labelled, and placed in long-term storage. Once cleaned and sorted, all cuticle was transferred to microscope slides, smoothed flat with a brush, covered with a drop of phenol glycerine jelly warmed to 60\u0026deg;C and then covered with glass coverslips, and inspected under a AX70 (Olympus, Australia) microscope with a UC50 (Olympus, Australia) camera attached, and CellSens (Olympus, Australia) image software. Any image stacking was done using Helicon Focus 7.6.4.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003ePlant cuticle from the scat material was prepared and identified with reference to plant voucher slides. It was quantified by counting square millimetres covered of each species on each slide using a microscopic grid. This was then calculated into percentage values for further analysis.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eIdentification of cuticle\u0026nbsp;\u003c/h2\u003e\n\u003cp\u003eTo determine what plant species \u003cem\u003eL. conditor\u003c/em\u003e were consuming, it was necessary to have a systematic method for comparing the fragments of cuticle in the faecal pellets with the reference cuticle, for identification. Due to the fragmented nature of the digested cuticle, we sought multiple identification characters to ensure that we correctly identified as much of the cuticle as possible. The key identifiers used in this study were differential absorption of stain, unspecialised epidermal cells, stomata, subsidiary cells and trichomes. The methods for identifying cuticle based on differential absorption of stain and unspecialised epidermal cells are outlined in Storr\u0026nbsp;(1961), and Metcalfe and Chalk\u0026nbsp;(1950), \u0026nbsp;respectively. We measured identifiers for each species and created an identification table for key plant species on the island (Supplementary Material 1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003ch2\u003eStatistics\u003c/h2\u003e\n\u003cp\u003eTo understand the selective consumption of plants by \u003cem\u003eL. conditor\u003c/em\u003e on Reevesby Island, we used Design I, described by Manly\u0026nbsp;(2002)\u0026nbsp;as a statistical design for quantifying habitat use and availability at the population level. The function for analysing Design I data, widesI, tests resource selection using Pearson\u0026apos;s chi-squared test (\u0026chi;2), and a log-likelihood chi-squared test. Pearson\u0026apos;s chi-squared test tests a null hypothesis stating that that \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eselect plants at random, meaning the portion of plant species represented in scats is consistent with that in the environment (allowing for a degree of error that is dependent on variation and sample size). The log-likelihood chi-squared test computes the goodness of fit of the null hypothesis vs the alternative hypothesis based on the ratio of their likelihoods for each plant species. If the null hypothesis is supported by the observed data, the two likelihoods should not differ by more than sampling error.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eEstimation of selective bias was estimated using the Manly selectivity measure (wi) with Bonferroni confidence intervals proposed by Manly et al. (2003):\u0026nbsp;w\u003csub\u003ei\u003c/sub\u003e = o\u003csub\u003ei\u0026nbsp;\u003c/sub\u003e/\u0026pi;\u003csub\u003ei\u003c/sub\u003e\u003c/p\u003e\n\u003cp\u003ewhere oi is the proportion of plant species present in scats and \u0026pi;i is the proportion of available plant species in category i (plant species). The Selection ratio for a species is therefore:\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/58854_b38fc7f3db2c487f/58854_custom_files/img1716475067.png\" width=\"856\" height=\"96\"\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003eIf the plant species is used (and preserved) in proportion to their availability, then w = 1. If wi \u0026gt; 1, the plant species is consumed or preserved more than expected by unbiased selection and preservation (preferred). The resource selection function (wi) was computed using adehabitatHS version 0.3.14 package with design I data type (Calenge 2006)in the R computing environment 3.6.1 (R Development Core Team 2019).\u0026nbsp;\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eFor all raw data and code, refer to Supplementary Material 2 and 3, respectively. Three vegetation types were present in the areas we surveyed (see Methods).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetation type 1\u003c/strong\u003e is the least abundant within the study area. It is located where there is some protection from the strong ocean winds and salt spray. The dominant species are \u003cem\u003eMyoporum insulare\u003c/em\u003e and \u003cem\u003eLycium ferocissimum\u003c/em\u003e (as medium to large bushes) and \u003cem\u003eMuehlenbeckia gunnii\u0026nbsp;\u003c/em\u003eand \u003cem\u003eTetragonia implexicoma\u0026nbsp;\u003c/em\u003e(as ground cover and in the canopy as creepers/vines).\u003cem\u003e\u0026nbsp;Leporillus\u003c/em\u003e faecal pellets and tracks are a common occurrence within patches of this vegetation type, but rarely found outside of them.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetation type 2\u003c/strong\u003e is the most prevalent in the study area, occupying the exposed dunes. It is defined by the dominance of \u003cem\u003eLeucophyta brownii\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;Olearia axillaris\u003c/em\u003e (as low shrubs) and\u0026nbsp;\u003cem\u003eSpinifex hirsutus\u003c/em\u003e (tufty grass). There is a substantial variation in density within this group, however, dominant species remain a fixed parameter. Due to accessibility and time constraints this vegetation type was sampled in a low-density area.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetation type 3\u003c/strong\u003e occupies low-lying salt and mud flats on the island in areas that are likely to become hypersaline due to inundation and subsequent evaporation of sea water. The dominant species are \u003cem\u003eSalicornia quinqueflora\u0026nbsp;\u003c/em\u003e(a small succulent shrub), \u003cem\u003eAtriplex paludosa\u003c/em\u003e (marsh saltbush), \u003cem\u003eTetragonia implexicoma\u003c/em\u003e and \u003cem\u003eOlearia axillaris\u003c/em\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003ch4\u003e\u003cstrong\u003eTable 1 here\u003c/strong\u003e\u003c/h4\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003ch4\u003ePlant species availability, use and selection by\u0026nbsp;L. conditor\u003c/h4\u003e\n\u003cp\u003eVegetation availability and use are shown in Figure 3, and the selection ratio is presented in Figure 4.\u003cem\u003e\u0026nbsp;L. ferocissimum\u003c/em\u003e is by far the most used resource, comprising 51.7% used material despite only representing 11.8% of the available vegetation in plot 1 and 10.43% of total sampled vegetation, as measured in the point intercept survey (Table 2). This gives it a selection ratio of 4.4 (Table 3).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigures 3\u0026amp;4 here\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2 here\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003ch4\u003ePlant use\u003c/h4\u003e\n\u003cp\u003eDue to its abundant representation in the scat material, \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003ewas broken into two subclasses based on cuticle type: \u0026ldquo;Leaf\u0026rdquo;, and \u0026ldquo;Flower\u0026rdquo;. \u003cem\u003e\u0026nbsp;\u003c/em\u003eLeaf cuticle comprised 62.2% of \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003ecuticle, and the remaining 37.9% can be attributed to flowers (Supplementary Materials 4).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAtriplex paludosa\u003c/em\u003e (marsh saltbush), \u003cem\u003eCarpobrotus rossii\u0026nbsp;\u003c/em\u003e(Australian pig face),\u0026nbsp;\u003cem\u003eL. ferocissimum,\u003c/em\u003e \u003cem\u003eEnchylaena tomentosa\u0026nbsp;\u003c/em\u003e(ruby saltbush),\u0026nbsp;\u003cem\u003eOlearia axillaris\u0026nbsp;\u003c/em\u003e(coastal daisy-bush), and\u0026nbsp;\u003cem\u003eMyoporum insulare\u003c/em\u003e (native juniper) have selection ratios \u0026gt;1, although this is not always statistically significant (Figure 4, Table 2). \u003cem\u003eTetragonia implexicoma\u003c/em\u003e (bower spinach) and \u003cem\u003eMuehlenbeckia gunnii\u003c/em\u003e (coastal lignum) are the only species that seem to be selected against. They comprise 23.7% and 20.2% of the total available vegetation in plot 1, but only 1.8% and 0% of the used resources (Figure 3).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eOf all the available species, the succulents have the most fragile cuticle, and are the most susceptible to damage during digestion and lab processes. As such, it may be reasonable to class the \u0026ldquo;used\u0026rdquo; data for these species as missing, or incomplete. However, studies show that the effects of differential preservation are usually minor\u0026nbsp;(Anthony and Smith 1974; Ellis\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1999; Khanam\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2015; Norbury 1988; Pareja\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2021; Todd and Hansen 1973) and, as the other plant cuticles are more robust, they are unlikely to be impacted by this secondary selective process.\u0026nbsp;\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWe found clear evidence of high use of the invasive \u003cem\u003eLycium ferocissimum\u0026nbsp;\u003c/em\u003e(African boxthorn) by\u0026nbsp;\u003cem\u003eL. conditor\u003c/em\u003e (stick-nest rat), which the animals appear to strongly preference as a food source over other native vegetation. Although it was not quantified, the density of \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eactivity in vegetation dominated by \u003cem\u003eL. ferocissimum\u003c/em\u003e suggests that it is also the preferred habitat for \u003cem\u003eL. conditor\u003c/em\u003e as already shown by previous research (Copley 1995; Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2019). Importantly, \u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003ewas the most strongly selected plant species and preferred even over succulent species such as the ruby saltbush, which form the majority of \u003cem\u003eL. conditor\u0026rsquo;s\u003c/em\u003e diet in the comparatively natural or restored habitats such as the Franklin Islands and the Arid Recovery native mammal re-introduction site of northern South Australia\u0026nbsp;(Copley 1999a; Read 1984; Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996b; Ryan\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2003).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIt is possible that flowers are more palatable and could have increased selective use of \u003cem\u003eL. ferocissimum\u003c/em\u003e. Flowers were abundant despite the collection period occurring in winter, outside the species\u0026rsquo; usual summer flowering period in summer\u0026nbsp;(Erkelenz 1993). \u0026nbsp;However, it seems more likely that this preference is driven by the habitat choice of \u003cem\u003eL. conditor\u003c/em\u003e, as the species seems to be a dietary generalist and consumes a variety of different plants, depending on location and season\u0026nbsp;(Read 1984). \u0026nbsp;.\u003c/p\u003e\n\u003cp\u003e\u003cu\u003e\u0026nbsp;\u003c/u\u003e\u003c/p\u003e\n\u003cp\u003eThe benefit of \u003cem\u003eL. ferocissimum\u003c/em\u003e as a food plant is probably accompanied by the further preference as shelter by \u003cem\u003eL. conditor,\u0026nbsp;\u003c/em\u003ealthough this was not quantified here\u003cem\u003e.\u0026nbsp;\u003c/em\u003eAt the Arid Recovery native mammal re-introduction site\u0026nbsp;(Moseby\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2011), \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003edemonstrated a preference for low, dense shrubs as the base for nest sites\u0026nbsp;(O\u0026rsquo;Neill 1999; Short\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2019), likely due to the protection they offer against predators, especially avian predators\u0026nbsp;(Copley 1988; O\u0026rsquo;Neill 1999). While \u003cem\u003eL. ferocissimum\u003c/em\u003e provides\u0026nbsp;\u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003ewith low, dense cover on Reevesby Island, so could other plants that grow within the same microhabitat. However,\u0026nbsp;\u003cem\u003eL. conditor\u003c/em\u003e may favour\u0026nbsp;\u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003ebushes because the large thorns provide more protection from predators than the plants that are native to the animal\u0026rsquo;s offshore island refuges. A preference for thorns and spines by \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eis supported by Short \u003cem\u003eet al.\u003c/em\u003e (2018)\u0026nbsp;who found that very prickly \u003cem\u003eAcacia tetragonophylla\u003c/em\u003e bushes were the primary nesting shrubs used by \u003cem\u003eL. conditor\u0026nbsp;\u003c/em\u003eat Heirisson Prong (Introduced in 1999, last sighting in 2007), in Shark Bay, Western Australia.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eDespite the successful adaptation of \u003cem\u003eL. conditor\u003c/em\u003e to boxthorn invasion, is difficult to judge the degree to which \u003cem\u003eL. ferocissimum\u003c/em\u003e is beneficial in replacing pre-disturbance structural complexity and food resources on Reevesby Island, as there is no historical reference for precolonial vegetation on the island. However, the location of the only natural \u003cem\u003eLeporillus\u0026nbsp;\u003c/em\u003epopulation, on the Franklin Islands, offers an interesting comparison. Descriptions of the vegetation there appear to be very similar to Reevesby Island, except that they do not contain \u003cem\u003eL. ferocissimum\u003c/em\u003e (Osborn 1922; Read 1984; Robinson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996b). Interestingly, on the Franklin Islands, stick-nest rats also show a clear preference for a habitat that is not part of the native vegetation, as population density of\u0026nbsp;\u003cem\u003eL. conditor\u003c/em\u003e appears to be highest among the granite slabs along the coastline, rather than more heavily vegetated areas. Areas of dense vegetation cover are also used, but appear to have lower population densities\u0026nbsp;(Read 1984). It therefore remains unclear if\u0026nbsp;\u003cem\u003eL. conditor\u003c/em\u003e\u0026rsquo;s preference for nesting in\u0026nbsp;\u003cem\u003eL. ferocissimum\u003c/em\u003e thickets is due to its efficiency as a shelter, as it is for other vertebrates\u0026nbsp;(Dutra\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2011; Hradsky\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2015; Law\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016; Packer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016); or whether it relates to a strong preference for\u0026nbsp;\u003cem\u003eL. ferocissimum\u0026nbsp;\u003c/em\u003eas a food source.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eEvidence of stick-nest rats using native vegetation for food and habitat on Reevesby Island and the Franklin Islands\u0026nbsp;(Read 1984)\u0026nbsp;suggests that \u003cem\u003eL. conditor\u003c/em\u003e on Reevesby Island are unlikely to require boxthorn for their survival. However, the fact that they appear not to favour native vegetation as habitat at either location emphasises the lack of information we have on this species. Part of the issue is that all translocation sites, and remnant natural populations, are located on the very fringe of \u003cem\u003eL. conditor\u0026rsquo;s\u003c/em\u003e former distribution\u0026nbsp;(Copley 1999a)\u0026nbsp;and unlikely to represent their optimal environment. In addition, there is no evidence of a preference for thorned plants preserved in fossilised stick-nest rat middens, suggesting that more research is required as to the original habitat preferences of this species\u0026nbsp;(Allen\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2000; McCarthy 1999; McCarthy and Head 2001; McCarthy\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 1996; Pearson 1999; Pearson\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2001; Pearson and Betancourt 2002; Pearson and Dodson 1993; Webeck and Pearson 2005). Lack of good understanding of habitat preference is an important issue in translocations, particularly in Australia and surrounding Oceanian areas where translocation sites are often subject to extensive weed invasions\u0026nbsp;(Morris\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2021). Another recent Australian example is the single remaining population of Shark Bay mouse (\u003cem\u003ePseudomys gouldi)\u003c/em\u003e, which\u003cem\u003e\u0026nbsp;\u003c/em\u003ealso has a broader range of suitable habitats than previously assumed\u0026nbsp;(Palmer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2024).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe apparent beneficial effects for African boxthorn infestation on stick-nest rats are consistent with other observations of beneficial interactions between invasives with native ecosystem components\u0026nbsp;(Azmi and Jennings 2013; Carothers\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2020; Packer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2016). This is an important consideration when assessing the economic impact of invasions and their control, particularly when expensive but inefficient measures result in a removal of beneficial effects\u0026nbsp;(Boltovskoy\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2022). In the case of stick-nest rats, this might be a particular issue in times of warming climates, which makes it particularly important for the species to have sufficient cover\u0026nbsp;(Onley\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2022). However, the beneficial effects of improved plant cover from invasive species\u0026nbsp;(Dutra\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2011)\u0026nbsp;need to be balanced against the otherwise detrimental effect of these often very destructive species. It is also not clear if the benefits will persist when a novel ecosystem may has reached a stable state. African boxthorn is highly invasive and, if left unchecked, may continue to spread, displacing native vegetation, and resulting in decreased ecosystem diversity and function that might ultimately also affect stick-nest rats themselves. It is even possible that the feeding preferences of \u003cem\u003eL. conditor\u003c/em\u003e facilitate the invasion of boxthorn because of its potential to spread the seed through its faeces, similar to observations that re-introductions of burrowing mammals can aid the spread of invasive plants\u0026nbsp;(Palmer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2020). This highlights the importance of considering the potential negative outcomes of translocation efforts in cases where translocated species interact with novel ecosystems in a way that is detrimental to overall ecosystem function\u0026nbsp;(Palmer\u003cem\u003e\u0026nbsp;et al.\u003c/em\u003e 2020).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eOur study has shown an unexpected preference for invasive \u003cem\u003eL. ferocissimum\u003c/em\u003e on Reevesby Island. Together with the species\u0026rsquo; apparent preference for granite slabs on Franklin Island, this provide important clues as to the importance of vegetation type, terrain, and geology in identifying potential future translocation sites. Our lack of understanding on the historical habitat of stick-nest rats suggests that assessments of the pre-colonial landscapes of previous occurrence sites (such as including midden sites and record maps) will be important for future translocation success.\u0026nbsp;\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank\u0026nbsp;field volunteers Baillie Trenwith, Holly Kraehe, Czes Klopotowski, and two anonymous reviewers.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Funding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eVW was funded by the Australian Research Council Centre of Excellence for Australian Biodiversity and Heritage (CE170100015) and Future Fellowship (FT180100634).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no conflict of interest to declare\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe raw vegetation survey and cuticle identification data for this manuscript, and the code to replicate Figures 5 and 6, are available in the supplementary materials.\u0026nbsp;A preprint version of of this article is available at\u0026nbsp;\u003ca href=\"https://url.au.m.mimecastprotect.com/s/xnesC3QNYgTGWYZ7s2v_Zr?domain=researchsquare.com\"\u003ehttps://www.researchsquare.com/article/rs-1963287/v1\u003c/a\u003e\u003c/p\u003e"},{"header":"References","content":"\u003cp\u003eAllen V, Head L, Medlin G, and Witter D (2000) Palaeo‐ecology of the Gap and Coturaundee Ranges, western New South Wales, using stick‐nest rat (\u003cem\u003eLeporillus spp\u003c/em\u003e.)(Muridae) middens. \u003cem\u003eAustral Ecology\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e, 333-343.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAnthony RG and Smith NS (1974) Comparison of rumen and fecal analysis to describe deer diets. \u003cem\u003eThe Journal of Wildlife Management\u003c/em\u003e, 535-540. doi: \u003ca href=\"https://doi.org/10.2307/3800886\"\u003ehttps://doi.org/10.2307/3800886\u003c/a\u003e.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAzmi WA and Jennings J (2013) The impact of management practices of exotic willows (Salix spp.) on aquatic invertebrate communities in South Australian freshwater streams. \u003cem\u003eJournal of Sustainability Science and Management\u003c/em\u003e \u003cstrong\u003e8\u003c/strong\u003e, 43-52.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eBergstrom BJ (2013) Would East African savanna rodents inhibit woody encroachment? 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(Australian Goverrunent Publishing Service: Canberra)\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eRobinson T, Canty P, Mooney T, and Rudduck P (1996b) South Australia\u0026rsquo;s offshore islands. \u003cem\u003eAustralian Heritage Commission: Canberra\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eRyan S, Moseby K, and Paton D (2003) Comparative foraging preferences of the greater stick-nest rat Leporillus conditor and the European rabbit Oryctolagus cuniculus: implications for regeneration of arid lands. \u003cem\u003eAustralian Mammalogy\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e, 135-146. doi: \u003ca href=\"https://doi.org/10.1071/AM03135\"\u003ehttps://doi.org/10.1071/AM03135\u003c/a\u003e.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSaunders B (2018) \u0026apos;Flowering Plants of Lower Eyre Peninsula: An Illustrated Tour of the Native Flora.\u0026apos; (Lane Print \u0026amp; Post: 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\u003cstrong\u003e15\u003c/strong\u003e, 466-471.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWhite A, Sparrow B, Leitch E, Foulkes J, Flitton R, Lowe AJ, and Caddy-Retalic S (2012) \u0026apos;AUSPLOTS rangelands survey protocols manual.\u0026apos; (University of Adelaide Press)\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWilliams JA and West CJ (2000) Environmental weeds in Australia and New Zealand: issues and approaches to management. \u003cem\u003eAustral Ecology\u003c/em\u003e \u003cstrong\u003e25\u003c/strong\u003e, 425-444.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWoehler EJ, Glencross JS, and Riley MA (2021) Survey of Little Penguins \u003cem\u003eEudyptula minor\u003c/em\u003e at Low Head 2019-2021.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWootton L, Halsey S, Bevaart K, McGough A, Ondreicka J, and Patel P (2005) When invasive species have benefits as well as costs: managing Carex kobomugi (Asiatic sand sedge) in New Jersey\u0026rsquo;s coastal dunes. \u003cem\u003eBiological Invasions\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e, 1017-1027.\u003c/p\u003e"},{"header":"Tables","content":"\u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"color:black;\"\u003eTable 1:\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cspan style=\"color:black;\"\u003eSummary of surveyed vegetation on Reevesby Island, showing percentage of vegetation cover as informed by each of the 3 plots the point intercept survey. \u0026ldquo;Total sampled vegetation\u0026rdquo; is the summary of the percentage cover for each plot, divided by the number of plots. \u0026nbsp;The percentages in Plot 1 are the same as the percentages of available vegetation in Fig. 3.\u0026nbsp;\u003c/span\u003e\u003c/p\u003e\n\u003ctable style=\"width:454.3pt;border-collapse:collapse;border:none;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;\"\u003eSpecies\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;\"\u003ePlot 1\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;\"\u003ePlot 2\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;\"\u003ePlot 3\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;\"\u003eTotal sampled vegetation\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;border: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eAtriplex paludosa\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;border: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.54%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;border: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;border: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e17.53%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;border: none;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e4.08%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eCarpobrotus rossii\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.08%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e6.33%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e2.72%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eDionella revoluta\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan 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15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.63%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.23%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eOlearia axillaris\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e7.53%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e12.03%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e13.40%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e10.43%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eRhagodia crassifolia\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.08%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.63%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.68%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eSalicornia quinqueflora\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e20.62%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e4.54%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eSenecio lautus\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e5.91%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.63%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e2.72%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eSpinifex hirsutus\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.61%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.27%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.13%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eTetragonia implexicoma\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e17.20%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e31.01%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e16.49%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e22.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 166.85pt;border-top: none;border-right: none;border-left: none;border-image: initial;border-bottom: 2.25pt solid windowtext;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;\"\u003eThrelkeldia diffusa\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 46.05pt;border-top: none;border-right: none;border-left: none;border-image: initial;border-bottom: 2.25pt solid windowtext;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.54%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 53.3pt;border-top: none;border-right: none;border-left: none;border-image: initial;border-bottom: 2.25pt solid windowtext;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e1.90%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 62.7pt;border-top: none;border-right: none;border-left: none;border-image: initial;border-bottom: 2.25pt solid windowtext;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;color:black;\"\u003e0.00%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 125.4pt;border-top: none;border-right: none;border-left: none;border-image: initial;border-bottom: 2.25pt solid windowtext;padding: 0in 5.4pt;height: 15.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:normal;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;\"\u003e0.91%\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;margin-bottom:8.0pt;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e\u0026nbsp;\u003c/span\u003e\u003c/p\u003e\n\u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"color:black;\"\u003eTable 2: Outcome of Manly selection ratio design I in Plot I\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u0026ldquo;Used\u0026rdquo;, total proportion of each species in the total faecal plant material; \u0026ldquo;available\u0026rdquo; represents the total proportion of each species in the point intercept surveys. Selection ratio is the product of \u0026ldquo;used\u0026rdquo; divided by \u0026ldquo;available\u0026rdquo;.\u0026nbsp;\u003c/p\u003e\n\u003ctable style=\"width:408.5pt;border-collapse:collapse;border:none;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 121pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eSpecies\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.8pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eUsed\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eavailable\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eselection ratio\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003estandard error\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.6pt;border-top: 1pt solid windowtext;border-left: none;border-bottom: 1pt solid windowtext;border-right: none;padding: 0in 5.4pt;height: 15pt;vertical-align: bottom;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003ep-value\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 121pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eAtriplex paludosa\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.8pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.007\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.005\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e1.355\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e1.433\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.6pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.804\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 121pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eCarpobrotus rossii\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.8pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.013\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.011\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e1.186\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.891\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.6pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.835\u003c/span\u003e\u003c/p\u003e\n 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style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.005\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:center;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e-\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n 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style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.034\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.016\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e2.089\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e1.243\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.6pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.381\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 121pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eFicinia nodosa\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.8pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.022\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:center;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e-\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.6pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:center;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e-\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 121pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;'\u003e\u003cem\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003eLeucophyta brownii\u003c/span\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.8pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49.15pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0.054\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 73pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp style='margin:0in;line-height:200%;background:white;font-size:16px;font-family:\"Times New Roman\",serif;color:#222222;text-align:right;'\u003e\u003cspan style=\"font-size:13px;line-height:200%;\"\u003e0\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 77.95pt;border: none;padding: 0in 5.4pt;height: 15pt;vertical-align: top;\"\u003e\n \u003cp 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style=\"color:windowtext;\"\u003e\u0026nbsp;\u003c/span\u003e\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-1963287/v2","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-1963287/v2","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eContext:\u003c/strong\u003e The threatened native rodent species Leporillus conditor (greater stick-nest rat) is extinct on the Australian mainland and now lives primarily on small islands off the coast of southern Australia. Many of these are degraded novel ecosystems invaded by African boxthorn (Lycium ferocissimum), a weed of national significance. However, L. conditor does not appear to be negatively impacted by the presence of boxthorn, raising the question of how the two species co-exist.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAims: \u003c/strong\u003eTo understand how L. conditor uses African boxthorn, we evaluated dietary composition of L. conditor on parts of Reevesby Island by comparing consumption of invasive boxthorn with that of native vegetation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMethods: \u003c/strong\u003eWe identified three key vegetation types on the centre of the island and used point-intercept vegetation surveys to estimate relative availability of plant species in each. We then used micro-histological faecal analysis to estimate the proportions of each species in the diet of L. conditor, and quantified plant species selection using selection ratios (use/availability).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eKey results\u003c/strong\u003e: Qualitative evidence of L. conditor activity suggested it was mostly confined to vegetation with greater abundance of boxthorn than the other vegetation types (13.5%, compared to 5.7% total sampled vegetation). Furthermore, African boxthorn comprised of 51.7% of the faecal plant content and 11.8% of total sampled vegetation, resulting in a selection ration for boxthorn of 4.4. Native species that appeared to be favoured food sources of L. conditor included Olearia axillaris, Myoporum insulare and Enchylaena tomentosa.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConclusions\u003c/strong\u003e: Stick-nest rats of Revesby Island demonstrate a clear preference for African boxthorn, both in terms of diet (tested quantitatively) and nesting (from previous research and field observations).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eImplications\u003c/strong\u003e: The strong preference of stick-nest rats for a declared noxious weed as its main food source and persistence of stick-nest rats on Reevesby Island requires consideration with regards to vegetation management on all islands where L. conditor occurs. More broadly, it highlights that some elements of novel ecosystems may have unexpected positive impacts on parts of original ecosystems.\u003c/p\u003e","manuscriptTitle":"Threatened stick-nest rats preferentially eat invasive boxthorn rather than native vegetation on Australia’s Reevesby Island","msid":"","msnumber":"","nonDraftVersions":[{"code":2,"date":"2024-05-23 14:42:56","doi":"10.21203/rs.3.rs-1963287/v2","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}},{"code":1,"date":"2022-09-15 14:34:56","doi":"10.21203/rs.3.rs-1963287/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"9dfabe05-1d78-4903-bc67-897d63c27888","owner":[],"postedDate":"May 23rd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-07-25T00:46:20+00:00","versionOfRecord":{"articleIdentity":"rs-1963287","link":"https://doi.org/10.1071/WR23140","journal":{"identity":"wildlife-research","isVorOnly":true,"title":"Wildlife Research"},"publishedOn":"2024-07-25 00:46:20","publishedOnDateReadable":"July 25th, 2024"},"versionCreatedAt":"2024-05-23 14:42:56","video":"","vorDoi":"10.1071/WR23140","vorDoiUrl":"https://doi.org/10.1071/WR23140","workflowStages":[]},"version":"v2","identity":"rs-1963287","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-1963287","identity":"rs-1963287","version":["v2"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}