DNA Isolation and Amplification from Cacti

The cacti family is a morphologically heterogeneous group comprising 100 genera and about 1500 species (Hernandez and Barcenas, 1996). With the exception of one genus, all members of this family are native to America (Hernandez and Barcenas, 1996). There are three subfamilies, Opuntioideae, Cactoideae, and Pereskioideae (Gibson and Nobel, 1986). DNA isolation from cacti is notoriously difficult because they contain high amounts of polysaccharides and secondary metabolites which form insoluble complexes with nucleic acids during extraction (Guillemaut and Marechal-Drouard, 1992). Like in other groups of plants, the secondary metabolites and polysaccharides in cacti inhibit enzyme action (Porebski et al., 1997). The polysaccharides are visually evident by their viscous, glue-like texture and they make the DNA unmanageable when pipeting and hard to amplify by the polymerase chain reaction (PCR) (Poresbski et al., 1997). We report an easy and inexpensive protocol to isolate DNA from cacti. We used this method to isolate DNA from 85 species (170 individuals) of 39 genera of the subfamilies Pereskioideae, Opuntioidea, and Cactoideae. This procedure is a modification of a protocol described by De la Cruz et al. (1995) for the Cacti family. It requires only a few grams of tissue and does not require destruction of the whole plant to produce high molecular weight genomic DNA. The DNA from this procedure can be amplified consistently by PCR and used for RAPD analysis.     

Cactus height increases the modularity of a plant–frugivore network in the Caatinga dry forest

Cacti fruits are key resources to many frugivorous animals in Neotropical arid and semiarid regions. However, most studies have focused on a particular animal group or cacti species, but few have explored the overall interactions of such species at the community level. Here we monitored frugivory on five cacti species using camera traps that sampled diurnal and nocturnal interactions. We investigated the structure of interactions with bird, mammal, and reptile frugivores in the Brazilian Caatinga dry forest. We hypothesized that the height of cacti limit interactions with different types of frugivores, which would result in highly structured and modular interaction networks. In 2929 camera-days, we recorded 23 vertebrate species feeding on cacti fruits, including seven new records, all determined to be primary seed dispersers. As predicted, the cacti-frugivore network was modular and non-nested, with the two shortest cacti species grouped in a module dominated by interactions with reptiles and non-flying mammals. The tallest cacti species were dominated by frugivory interactions with birds and had comparatively less interaction diversity than shorter cacti species. Our results support the contention that cacti are keystone species in semiarid ecosystems where they produce small-seeded fleshy fruits year-round.     

Protection from uv radiation in the economic crop, <Emphasis Type="Italic">Opuntia</Emphasis> SPP.

Cacti of the genus puntia are an economically important crop. Understanding the mechanisms they possess to protect against UV radiation is important for assessing their possible response to climatic change. Measurements of the concentrations of UV-screening compounds and epidermal transmittance for two species of platyopuntia, Opuntia engelmannii Salm-Dyck. and O. phaeacantha Engelm. during 1998 and 1999 were used to investigate the UV-protection afforded by the cactus epidermis. A UV-radiative transfer model was used to investigate the interception of UV radiation on differently oriented surfaces. We show that vertical morphology itself confers significant protection against UV radiation compared to a horizontal surface. Concentrations of UV-screening flavonoids were found to vary depending on the UV exposure of different surfaces. West-facing surfaces had lower concentrations than east-facing surfaces, although theoretically they should be identical. This might be explained by the higher mean temperatures on west-facing surfaces. Although UV-absorbing soluble flavonoids in the epidermis block both UV-B and UV-A, the structure of the epidermis alone may be sufficient to remove up to 94% of the UV-B portion of the spectrum. These data yield insights into possible mechanisms of recent declines in cacti populations.     

Structure-function relationships in highly modified shoots of cactaceae

BACKGROUND AND AIMS: Cacti are extremely diverse structurally and ecologically, and so modified as to be intimidating to many biologists. Yet all have the same organization as most dicots, none differs fundamentally from Arabidopsis or other model plants. This review explains cactus shoot structure, discusses relationships between structure, ecology, development and evolution, and indicates areas where research on cacti is necessary to test general theories of morphogenesis. SCOPE: Cactus leaves are diverse; all cacti have foliage leaves; many intermediate stages in evolutionary reduction of leaves are still present; floral shoots often have large, complex leaves whereas vegetative shoots have microscopic leaves. Spines are modified bud scales, some secrete sugar as extra-floral nectaries. Many cacti have juvenile/adult phases in which the flowering adult phase (a cephalium) differs greatly from the juvenile; in some, one side of a shoot becomes adult, all other sides continue to grow as the juvenile phase. Flowers are inverted: the exterior of a cactus 'flower' is a hollow vegetative shoot with internodes, nodes, leaves and spines, whereas floral organs occur inside, with petals physically above stamens. Many cacti have cortical bundles vascularizing the cortex, however broad it evolves to be, thus keeping surface tissues alive. Great width results in great weight of weak parenchymatous shoots, correlated with reduced branching. Reduced numbers of shoot apices is compensated by great increases in number of meristematic cells within individual SAMs. Ribs and tubercles allow shoots to swell without tearing during wet seasons. Shoot epidermis and cortex cells live and function for decades then convert to cork cambium. Many modifications permit water storage within cactus wood itself, adjacent to vessels.     

Spiny invaders – Patterns and determinants of cacti invasion in Europe

In this paper, we analyse the patterns and determinants of cacti invasion in 22 European countries. We compiled a checklist for each country. Cacti were classified for each country according to their invasion status as casuals, locally established (1–5 localities of small population size) and widely established (>5 localities of considerable population size).We used generalised linear models (GLM) from the Poisson family with a log-link function and a set of seven country-specific explanatory variables to account for geographical, climatic, habitat-related and economic determinants to test which features of the recipient area determine invasion success and if distribution patterns of species at different invasion stages are governed by the same interplay of explanatory variables. Separate models were fitted with the same predictor variables for casual, locally established, widely established and all cacti. Further, we analysed the temporal invasion trend, and tested if niche breadth (expressed as the number of habitat types colonized) is influenced by the range size (measured as the number of countries invaded). Finally, we reviewed the consequences of cacti invasion for nature conservation.In total, 26 cacti species have been recorded in Europe. Ten species are more widespread and occur in at least three countries, Opuntia humifusa (six countries) being the most widespread species. The country with most cacti is Spain (21 species), whereas in 13 countries no cactus species have been recorded. By far the most important genus is Opuntia with 20 alien species. The temporal invasion pattern shows an exponential increase of the cumulative number of invasion events, increasing from three (1801–50) to nine (1951–2000) invasion events over a 50-year period.Regardless of the invasion stage, the factor explaining most of the variance in the models is the presence of the Mediterranean biogeographic region, and a significant positive effect of the country size on species numbers was identified.Considering the invasion stage, some interesting deviations in the models can be observed. Invasion of casual cacti is only influenced by the presence of the Mediterranean biogeographic region. For locally established cacti, precipitation is negatively correlated with the invasion rate, and the presence of the Alpine biogeographic region is positively correlated; the latter is due the local occurrence of few hardy cacti (Opuntia phaeacantha, O. humifusa) in low-lying valleys of the Alps. As all widely established cactus species are restricted to the Mediterranean region, only this factor was included in the model.All cacti are confined to dry, open habitats on acid siliceous bedrock. Thus, the predominant habitats invaded are rock vegetation, dry grassland, open Mediterranean scrub and dry ruderal habitats. The niche breadth of cacti increases with the numbers of countries colonized. Further, the niche breadth of cacti exhibits a geographic gradient towards the Mediterranean region.Until the 19th century, the dominant pathway of invasion was agriculture, as some cactus species had been introduced for the production of forage and fruits. However, in the last decades horticulture and deliberate planting in the wild have become the dominant pathways.The invasion of cacti in natural and semi-natural habitats in the Mediterranean region changes habitat structure and species composition. However, dense and extensive stands of cacti are restricted to few species (e.g. Opuntia ficus-indica).     

Ecohealth and Aboriginal Testimony of the Nexus Between Human Health and Place

The spread of industrial civilizations has been particularly traumatic for the last remaining hunter-gatherer societies. Manifestations of this include expatriation from ancestral lands, sickness, poverty, and environmental degradation. Northern Australia has been no exception despite remaining a stronghold of Aboriginal cultures and still containing vast areas of relatively intact landscapes. Most Aboriginal people reside in remote settlements where they remain on the negative extreme of basic indicators such as life expectancy and educational attainment. In addition, biodiversity declines are being documented from loss of Aboriginal fire management and invasion by feral species. There has been little consideration of potential health, social, economic, or environmental benefits of routinely hunting, gathering or being on their land. This reflects a Western philosophical position that segregates land management and health policy, a view at odds with Aboriginal peoples’ testimony of the indivisibility of people and land. Here we report perspectives from Arnhemland gathered through observation and unstructured and semistructured interviews. Themes that emerged included the high level of detailed, complex knowledge of their traditionally owned lands, the perceived urgency about passing this on to younger people, and the need that both land and people have for each other for the well-being of both. Primary motivations for returning to traditional lands were gathering food, escaping from stresses, and educating young people. The many barriers included no transport, family problems, frequent funerals, and other cultural or family obligations. This work forms part of a larger transdisciplinary research program that aims to inform policy about sustainable futures in northern Australia.     

An evaluation of the global potential of bioenergy production on degraded lands

In this article the global potential of energy crop production on degraded lands was estimated using detailed, spatially explicit data about the area, type and extent of degradation derived from the Global Assessment of Land Degradation Dataset, and by combining this dataset with various spatially explicit data sets. Next, an estimate was made of the possible yield of perennial energy crops on the degraded areas as a function of the type and degree of degradation. Lightly degraded areas were not included, as these areas might be suitable for conventional food production. The total global potential energy production on degraded lands was assessed to be slightly above 150 and 190 EJ yr^?1, for grassy and woody energy crops, respectively. Most of this potential, however, is on areas currently classified as forest, cropland or pastoral land, leaving a potential of around 25 and 32 EJ yr^?1 on other land cover categories. Most of the potential energy crop production on degraded land is located in developing regions. China has a total potential of 30 EJ yr^?1, of which 4 EJ yr^?1 from areas classified as other land. Also USA, Brazil, West Africa, East Africa, Russia and India have substantial potentials of 12–18 EJ yr^?1, with up to 30% of the potential from areas classified as other land.     

Stem surface injuries of 20 species of succulent Euphorbia (Euphorbiacae) from South Africa

Cactus and succulent Euphorbia plants have green stems that are exposed to sunlight over long periods. Previous research demonstrated that barking injuries occur to stem surfaces of 20 species of cacti in the Americas. For these 20 cactus species the amounts of injuries were related to annual amounts of sunlight exposure on surfaces. Cacti with many injuries had high mortality rates. Data herein show that stem surfaces of 20 species of Euphorbia in South Africa have identical surface injuries as for cacti. Euphorbia plants at 30°S show a mean ratio of scale and bark injuries of 3.5:1 of equatorial-facing to polar-facing surfaces, a ratio is similar to the 4:1 of annual sunlight exposure ratio at that latitude. Results show that Euphorbia stems have injuries on younger tissues than cactus species. Thus, Euphorbia stems are more sensitive than cactus stems. Euphorbia species with most stem injuries had smooth (non-undulate) surfaces, a one-celled epidermal cell layer only, cortex cells that abutted the epidermis, and initial injuries that involved only epidermal cells. This is the first report of identical stem surface injuries to Euphorbia species that are identical to surface injuries to cactus species worldwide. Moreover, the amounts on stem injuries on Euphorbia and long-lived, columnar cactus species are highly correlated with relative amounts of sunlight on stem surfaces over the annual cycle. These results suggest that these species are good bio-indicators of annual sunlight exposure.     

Selecting plant species for practical restoration of degraded lands using a multiple‐trait approach

Ecological restoration is essential in rehabilitating degraded areas and safeguarding biodiversity, ecosystem services and human welfare. Using functional traits to plan restoration strategies has been suggested as they are the main ecological attributes that underlie ecosystem processes and services. However, few studies have translated ecological theory into actual restoration practices that can be easily used by different stakeholders. In this article, we applied a multiple‐trait approach to select plant species for the restoration of degraded lands inside the Brazilian Amazon Forests. We selected 10 traits encompassing ease of management, geographical distribution and interactions with animals and other ecosystem services and scored these traits using 118 native species. Then, we ranked all species according to the total number of traits that they exhibited to obtain a list of 53 highly ranked species. In addition, we employed non‐metric multidimensional scaling (NMDS) to assess the variation in these traits across the entire group of species. Based on the results, we selected a subset of species that maximizes functional diversity (high variability). We performed a sparse linear discriminant analysis (SLDA) to highlight a minimum set of traits to effectively discriminate botanical families. The final list of species and their traits highlight the importance of preserving not only the historical reference of a focused ecosystem but also its functional diversity to restore the interaction with local fauna, enrich the food chain and guarantee ecosystem services for local communities.     

Phylogenetic analysis and classification of insect achaete-scute complex genes

Achaete-scute complex (ASC) genes play essential roles in neurogenesis of eukaryotes. Insect ASC genes are currently grouped into two distinct families, ASCa and ASCb. However, the identified insect ASCb family genes are not orthologous to known ASCb family genes from other animals such as nematode and mammal. Besides, many ASC genes have been annotated differently in GenBank. In this study, ASC genes from 41 insect species were obtained in silico and analyzed to clarify their evolutionary relationships with known ASC genes from other animals. Based on our analyses, a new naming system is suggested for classification of ASC genes. Firstly, a new family (ASCc) is added into the already existing two families and the gene of this family is named Asl (Achaete-scute-like). Secondly, Ash4, Ash5 and Ash6 are used to name insect ASC genes that are not orthologous to ac, sc, l'sc and ase genes of Drosophila melanogaster and Ash1 to Ash3 genes of Bombyx mori. This naming system can be used to correct inappropriate annotations of insect ASC genes in GenBank and to classify newly identified insect ASC genes. Additionally, a vast difference was found in ASC gene composition among various insect species, suggesting that different insect species might have originated from various ancestral lineages.     

Isolation and characterization of chicken DNA homologous to the two putative oncogenes of avian erythroblastosis virus

The genome of avian erythroblastosis virus contains two independently expressed genetic loci (v-erbA and v-erbB) whose activities are probably responsible for oncogenesis by the virus. Both loci are closely related to nucleotide sequences found in the DNA and RNA of chickens and other vertebrates. We have isolated and characterized chicken DNA homologous to v-erbA and v-erbB. The two viral genes are represented by separate domains within chicken DNA (c-erbA and c-erbB), which are separated by a minimum of 12 kilobases (kb) of DNA and may not be linked at all. The nucleotide sequences shared by the viral and cellular erb loci are colinear, but the cellular loci are interrupted by multiple intervening sequences of various lengths. Polyribosomes prepared from normal chicken embryos contain two polyadenylated RNAs transcribed from c-erbA and two transcribed from c-erbB. The evident coding regions of these RNAs represent an unusually small fraction of the lengths of the RNAs, as if the 3′ untranslated domains of the RNAs might be exceptionally large (3–11 kb). These findings indicate that the c-erb loci are normal vertebrate genes rather than genes of cryptic endogenous retroviruses, and that they may have a role in the metabolism of normal cells. It appears that the viral erb genes, like most other retrovirus oncogenes, have been copied from cellular genes. In the viral genome, the two genes are devoid of introns, but they remain independently expressed loci, and they remain colinear with the coding domains of their cellular progenitors.     

Endophytic bacteria of the rock-dwelling cactus Mammillaria fraileana affect plant growth and mobilization of elements from rocks

Mammillaria fraileana is a major pioneer, small cactus that harbors endophytic bacteria that have plant growth-promoting traits, including rock-weathering capacity. Our working hypothesis was that this functional group of endophytic bacteria assists in establishing pioneer plants on rocks. When these endophytic bacteria were inoculated on seedlings grown in rock substrate, mobilization of elements from the substrate increased at variable levels across combinations of substrates and inoculants. In plants grown in the rhyodacite substrate, where these cacti naturally grow, increased mobilization occurred in plants inoculated with several strains. Promotion of plant growth, manifested as an increase in dry weight, was greater in cacti inoculated with Enterobacter sakazakii M2PFe. Accumulation of nocturnal acids, indicating photosynthesis by crassulacean acid metabolism, was superior in plants inoculated with the endophytes Azotobacter vinelandii M2Per and Pseudomonas putida M5TSA. Inoculation with endophytes can stimulate plant growth of M. fraileana by mobilizing elements from rock, which can lead to higher photosynthetic activity and accumulation of biomass. Inoculation with P. putida M5TSA also led to accumulation of more total nitrogen than plants inoculated with a control nitrogen-fixing bacteria. Evidence of endophytic colonization is provided after initial inoculation of seedlings and re-isolation and sequencing of 16S DNA of recovered bacteria from developing disinfected plants. The associative interaction between pioneer cacti and their bacterial endophytes enable the host plants to grow in places where plants do not normally grow. Through colonization and establishment of pioneer plants, soil is created, which facilitates colonization by other desert species and contributes to the diversity of dry lands.     

Oviposition preference and life history traits in cactophilic Drosophila koepferae and D. buzzatii in association with their natural hosts

Drosophila koepferae and D. buzzatii are two closely related cactophilic species inhabiting the arid lands of southern South America. Previous studies have shown that D. buzzatii breeds primarily on the necrotic cladodes of several Opuntia cacti and D. koepferae on the rotting stems of columnar cacti of the genera Trichocereus and Cereus. In this paper, we analyze the patterns of host plant utilization in a locality where both Drosophila species are sympatric. Field studies showed an absence of differential attraction of adult flies to the rots of two major host cacti: O. sulphurea and T. terschekii. However, the proportion of D. buzzatii flies emerged from the rotting cladodes of O. sulphurea was significantly higher than in T. terschekii. In laboratory experiments, egg to adult viability in single species cultures varied when both Drosophila species were reared in media prepared with O. sulphurea or T. terschekii. In addition, between-species comparisons of flies emerged from single species cultures showed that D. buzzatii adults were smaller and developed faster than D. koepferae. Furthermore, analysis of flies emerged in mixed species cultures showed differences in oviposition preference and oviposition behavior. We discuss the observed between-species differences and suggest that these traits are the result of adaptation to specific patterns of spatial and temporal predictability of their respective preferred host plants: columnar are less dense and less ephemeral resources, whereas the opuntias are more abundant, and fast rotting cacti. This revised version was published online in July 2006 with corrections to the Cover Date.     

Trophic interactions between two medium-sized mammals: the case of the native Dolichotis patagonum and the exotic Lepus europaeus in a hyper-arid ecosystem

As medium-sized herbivores, the exotic Lepus europaeus (European hare) and the native Dolichotis patagonum (mara) have been considered ecological equivalents. These species coexist in Ischigualasto Provincial Park, a hyper-arid ecosystem with scarce food resources. Our objective was to evaluate diet composition, relationship between diets and food availability, and trophic relationships between both herbivores. Collection of feces and vegetation sampling were made in the Mesquite woodland community. Diet composition was analyzed by microhistological analysis of feces. In both seasons, shrub species represented the most abundant cover type in the area, and annual forbs and grasses appeared in the wet season. Herbivores showed similar dietary ecology: shrubs were the main food items along the year, showing a higher plasticity compared to their diets in other ecosystems, where they selected mostly grasses. The mara selected shrubs such as Atriplex sp. and Prosopis torquata, whereas the European hare selected Cyclolepis genistoides, Atriplex sp., and Bulnesia retama. During the wet season, both herbivores supplemented their diets with grasses and annual forbs. In the dry season, there was increased consumption of cacti, such as Tephrocactus sp. The mara and the European hare are likely close ecological equivalents, in terms of dietary similarity, and they showed strong dietary overlap across the dry season (over 60 %). Thus, we can assume the existence of a potential trophic competition between mara and European hare, especially during the season when food resources are scarce. These results can be important for the management of drylands in South America, where populations of threatened herbivorous species, such as the mara, coexist with exotic animals, sharing spatial and trophic resources even in protected areas.     

Induced climate change impairs photosynthetic performance in Echinocactus platyacanthus, an especially protected Mexican cactus species

The responses of desert plants to climate warming have been poorly assessed, perhaps due to the overall expectation that desert vegetation will expand as a consequence of this component of climate change. However, determining what plant species will tolerate the expected increase in temperature is a question that remains unanswered. The Chihuahuan Desert is the largest warm desert of North America, and predictive models of climate change indicate that summer temperatures in this desert will increase by 1–2 °C in the next decade. This study experimentally assessed the performance of an endangered cacti species from the Chihuahuan Desert under simulated warming conditions. Hexagonal open top-chambers (OTCs) were used to simulate the effects of global warming on five-years-old individuals of the specially protected species Echinocactus platyacanthus. Temperature was 1.9 °C higher in open top-chambers than in control plots. In contrast, relative humidity was 3.1% higher in control plots than in open top-chambers. E. platyacanthus showed 100% survival for 14 weeks in both OTC and control plots. However, induced warming negatively affected the photosynthetic performance of this species. Cacti located within OTCs displayed lower maximum quantum efficiency of photosystem II (F_v/F_m), effective quantum yield of photosystem II (Φ_PSII), and electron transport rate (ETR) values, but higher non-photochemical quenching (NPQ) values, than cacti from control plots. This is the first study focused on the potential impact of climate warming on survival and photosynthetic performance of young individuals of a succulent species from American deserts. Induced warming negatively affected the photosynthetic performance of young E. platyacanthus, but it also increased non-photochemical quenching, a mechanism for avoiding photoinhibition.     

Giant shoot apical meristems in cacti have ordinary leaf primordia but altered phyllotaxy and shoot diameter

BACKGROUND AND AIMS: Shoot apical meristems (SAMs) in most seed plants are quite uniform in size and zonation, and molecular genetic studies of Arabidopsis and other model plants are revealing details of SAM morphogenesis. Some cacti have SAMs much larger than those of A. thaliana and other seed plants. This study examined how SAM size affects leaf primordium (LP) size, phyllotaxy and shoot diameter. METHODS: Apices from 183 species of cacti were fixed, microtomed and studied by light microscopy. KEY RESULTS: Cactus SAM diameter varies from 93 to 2565 microm, the latter being 36 times wider than SAMs of A. thaliana and having a volume 45 thousand times larger. Phyllotaxy ranges from distichous to having 56 rows of leaves and is not restricted to Fibonacci numbers. Leaf primordium diameter ranges from 44 to 402 microm, each encompassing many more cells than do LP of other plants. Species with high phyllotaxy have smaller LP, although the correlation is weak. There is almost no correlation between SAM diameter and LP size, but SAM diameter is strongly correlated with shoot diameter, with shoots being about 189.5 times wider than SAMs. CONCLUSIONS: Presumably, genes such as SHOOT-MERISTEMLESS, WUSCHEL and CLAVATA must control much larger volumes of SAM tissue in cacti than they do in A. thaliana, and genes such as PERIANTHIA might establish much more extensive fields of inhibition around LP. These giant SAMs should make it possible to more accurately map gene expression patterns relative to SAM zonation and LP sites.     

Ethnobotany of Ferocactus histrix and echinocactus platyacanthus (Cactaceae) in the Semiarid Central Mexico: Past,Present and Future

Ferocactus histrix and Echinocactus platyacanthus are two common barrel cacti of the semiarid highlands of Central Mexico. In pre-Columbian times, these plants were sacred, used for sacrifices, food and medicine. Today, these cacti are used in the candy industry, as medicine, as a source of food and water for goats and as ornamental plants. The floral buds and fruits ofF. histrix are consumed by the local people; the latter are sold in local markets. Until recently, the apical indument ofE. platyacanthus was used for padding and weaving. The importance of these species increases with the aridity of the land and its inadequacy for agriculture. Habitat protection and artificial propagation are needed to avoid the extinction of these species.     

The Role of Endangered Species Reintroduction in Ecosystem Restoration: Tortoise–Cactus Interactions on Española Island, Galápagos

We evaluated the role that endangered species reintroduction efforts can play in the larger context of ecosystem restoration. To do so, we examined interactions between endangered giant tortoises (Geochelone nigra hoodensis), currently being reintroduced to Isla Española, Galápagos, and an arboreal cactus (Opuntia megasperma var. megasperma), which is itself endangered and a keystone resource for many animals on the island. We collected information on spatial patterns of occurrence of cacti, tortoises, and woody vegetation and compared recruitment of juvenile cacti in areas occupied versus unoccupied by tortoises. Reintroduced tortoises appeared to suppress cactus recruitment near the few remaining adult cacti at the study site, but facilitate it at longer distances, with tortoise–cactus interactions mediated by the presence of woody vegetation, which likely alters tortoise movements and thereby patterns of cactus seed dispersal. The net effect of tortoises on cacti appeared to be positive insofar as tortoise presence was associated with greater recruitment of juveniles into cactus populations. Our study provides support for reintroducing endangered reptiles and other animals to aid ecosystem restoration in areas where they might once have played an important role in grazing upon and dispersing plants.