Long-term response of spring flora to chronic herbivory and deer exclusion in Great Smoky Mountains National Park, USA

Chronic herbivory by white-tailed deer (Odocoileus virginianus Zimmerman) can have profound impacts on the function and structure of forest ecosystems. We examined the combined influence of intense herbivory associated with a deer population eruption and chronic herbivory by the post eruption population on the spring flora of Cades Cove, Great Smoky Mountains National Park. During the 1970s the deer population reached a peak of 43 deer per km², from which it has slowly declined in recent decades. To examine the influence of intense herbivory, we compared the abundance and flowering rates of early flowering plants in Cades Cove to a nearby reference site with similar bedrock geology, vegetation, and disturbance history but contrasting history of deer abundance. Our results suggest that significant changes (p ? 0.05) in the diversity, evenness, and species richness of the spring flora occurred during the eruptive phase. Trillium spp. and other liliaceous species appeared to be disproportionately impacted. Comparisons between control and exclosure plots established after the deer population eruption indicate that recovery has been largely restricted to species that were able to persist under intense herbivory. These species have increased in number in exclosures, suggesting continued impacts by deer on the plant community outside the exclosures. Little to no recolonization by browse sensitive species was observed. Consequently, to restore the natural diversity of early flowering plants once present in Cades Cove, active restoration may be necessary in addition to maintaining deer densities below current levels.     

A natural experiment on the effects of high deer densities on the native flora of coastal temperate rain forests

The introduction of Sitka black-tailed deer (Odocoileus hemionus sitkensis Merriam) to Haida Gwaii (Queen Charlotte Islands, BC, Canada) in the late 19th century, provided an opportunity to understand the long-term effects of deer populations on the vegetation of temperate rain forests in the absence of their natural predators wolves (Canis lupus L.), and cougars (Puma concolor L.). Using seven small islands with different browsing histories (no deer, deer for <20 years, deer for >50 years), we tested the long-term effects of high deer densities on plant cover and species richness in the understorey of forest interior and forest edge habitats. Overall vegetation cover exceeded 80% in the lower vegetation layers on islands without deer and was less than 10% on the islands with deer for more than 50 years. Although overall plant species richness was similar on islands with or without deer, plant species richness at the plot scale (314 m²) was reduced by 20-50% on islands with deer for >50 years. The differences were most pronounced for the species-rich edge communities and among herb and shrub species. These results suggest that in the absence of predators, deer have the potential to greatly simplify the forest ecosystem.     

Seeing the forest for the deer: Do reductions in deer-disturbance lead to forest recovery?

High levels of deer browsing can prevent canopy tree regeneration, but little is known about changes to forest size-structure following long-term deer herd reductions. We monitored changes in forest stand structure and composition in southwestern Ontario, Canada, over 28-years using permanent plots. Our study site was the largest remaining tract of Carolinian (deciduous) forest in Canada (11 km²), a habitat type that contains up to a fifth of Canada’s species at risk and is under intense anthropogenic pressures. We recorded declines in all tree size classes between 1981 and 1996, during which densities of white-tailed deer (Odocoileus virginianus) reached a peak of 55 deer km⁻². Despite significant and sustained deer herd reductions between 1996 and 2009, which reduced deer densities to 7 deer km⁻², there was limited recruitment of small trees and declines in basal area of tree species that were sensitive to deer browsing. Our results suggest that recovery from herbivory is a protracted process during which canopy tree regeneration may continue to decline despite a reduction in browsing pressure due to deer culling. Large declines in canopy-tree densities in Carolinian forests may lead to forest size-structures and herbaceous plant communities that resemble rare oak savanna habitat, creating difficult decisions for conservation managers aiming to protect rare and endangered species within native ecosystems. We recommend that managers protect Carolinian forest stands and encourage canopy tree regeneration by increasing seed sources of native trees. While deer control is essential in reducing forest damage, our results highlight the need to explore other forms of active management to expedite otherwise slow increases in tree density.     

Long-term effects of deer browsing and trampling on the vegetation of peatlands

Overabundance of wild ungulates, especially exotic species, is a major threat to several ecosystems worldwide. While the response of forest vegetation to high density of herbivores has been well studied, far less is known about peatland vegetation. In this paper, we assessed the long term impact of white-tailed deer (Odocoileus virginianus) on plant communities of ombrotrophic (bog) and minerotrophic (fen) peatlands in eastern North America. Vegetation of five peatlands that have experienced high deer densities for at least 75 years was compared with that of five peatlands situated at proximity but on deer-free islands. We investigated deer impacts on plant species composition and cover, shrub height and cloudberry (Rubus chamaemorus) fruit/flower production and morphology. In bogs, white-tailed deer had no long-term impact on plant species assemblages, but reduced lichen cover and increased sedges and grasses cover as well as the surface area of bare peat. On the other hand, the floristic composition of fens differed significantly between sites where deer were present or absent. Plant diversity was greater in undisturbed fens than in disturbed fens, especially for shrubs, sedges and liverworts. No detrimental effects of browsing on shrub height were observed. Conversely, deer browsing seemed to have deleterious impacts on cloudberry fruit/flower production as well as on the number of leaves per individual. Overall, our results suggest that white-tailed deer had some important impacts on the vegetation of peatlands that could be harmful for the long-term conservation of peatland plant diversity.     

Restoring forest herb communities through landscape-level deer herd reductions: Is recovery limited by legacy effects?

White-tailed deer (Odocoileus virginianus) overbrowsing has altered plant species diversity throughout deciduous forest understories in eastern North America. Here we report on a landscape-level (306 km²) project in Pennsylvania, USA that tracked the herbaceous community response to deer herd reductions. From 2001 to 2007, we estimated deer densities, browse impact on woody seedlings, and censused the herbaceous flora in permanent plots throughout the area. We assessed herb layer species richness, abundance, and dominance and measured three known phytoindicators of deer impact: Trillium spp., Maianthemum canadense, and Medeola virginiana. We predicted that browse-sensitive taxa would increase in abundance, size, and flowering as would overall species diversity following deer culls and browse impact that declined by an order of magnitude by 2007. Following intensified deer harvests, we observed a limited recovery of the herbaceous community. Trillium spp. abundance, height, and flowering; M. canadense cover; and M. virginiana height all increased following herd reductions. Similarly, forb and shrub cover increased by 130% and 300%, respectively. Nevertheless, species diversity (i.e., richness and dominance) did not vary. Our work demonstrates that reducing deer densities can provide rapid morphological and population-level benefits to palatable species without a concomitant recovery in diversity. We suggest that decreasing deer populations alone may not promote plant diversity in overbrowsed, depauperate forests without additional restoration strategies to mitigate a browse-legacy layer dominated by browse-resistant species.     

Fifty years of change in northern upland forest understories: Identity and traits of “winner” and “loser” plant species

Resurveys of plant communities provide valuable information on changes in species composition over time and clues about how species respond to environmental change. We report results from resurveys of 62 upland forest stands in northern Wisconsin and the western Upper Peninsula of Michigan first surveyed around 1950. We identify plant species that have significantly increased or decreased in frequency in 1 m² quadrats (‘winners’ and ‘losers’) and evaluate the traits that distinguish these groups. Twenty-one winner species increased across sites (by 25-400%), while 21 loser species decreased (by 21-95%). Winners include both common, native species and five invading exotics. Many are grasses or sedges and most are abiotically pollinated or dispersed (e.g., Carex, which increased 286%). Losers are mostly rarer native forbs that rely on animals for pollination and/or dispersal. Losers appear sensitive to desiccation, anthropogenic disturbance, and/or herbivory by white-tailed deer (e.g., Streptopus roseus, which decreased 73%). Declines in loser species are heterogeneous and stochastic across sites whereas increases in winners are more uniform and deterministic. Increases in common widespread native species account for most of the community change we observe across sites. The fact that winners resist or tolerate deer herbivory while many losers are sensitive to deer suggests that deer may be a key driver of the shifts we observe in these forests.     

Browsing down our natural heritage: Deer impacts on vegetation structure and songbird populations across an island archipelago

Declines in forest and woodland birds have largely been attributed to habitat loss and fragmentation. In the past decade, however, the potential for herbivores to influence bird species abundance and community composition via their direct impact on vegetation structure has also been recognised. We tested the hypothesis that deer influence vegetation structure and bird assemblages in a large island archipelago in western North America using surveys of 18 islands with deer densities ranging from 0 to over 1 deer/ha. Amongst these islands, reduced predation and hunting pressure has allowed deer populations to increase above those likely to have existed in pre-European times. Our results support a growing body of evidence that deer regulate both the cover and architecture of understory vegetation which in turn profoundly affects island bird assemblages. Deer-free islands supported the most abundant and diverse bird fauna. Iconic songbirds such as the rufous hummingbird, song and fox sparrow were abundant on islands with no deer but substantially reduced on islands with high deer densities. Only one bird species, the dark-eyed junco, preferred moderate and high density deer islands. Our observations suggest that current cohorts of palatable shrubs on islands with high deer densities are relatively old and potentially represent an impending extinction debt, where the full effects of high deer density on island biota may take decades to fully unfold. Our results suggest that deer densities below a threshold of 0.1 deer/ha should allow native vegetation to recover and a rich and diverse bird species assemblage to persist. We suggest that adaptive management be used to test the validity of this threshold, and that without active management of deer abundance, local extinctions of native flora and fauna appear likely to accelerate.     

Disperser communities and legacies of goat grazing determine forest succession on the remote Three Kings Islands, New Zealand

Many remote islands are degraded as a result of deforestation and browsing of vegetation by introduced goats. Goat eradication is therefore a focus for island restoration, but there are few long-term records of change on islands after eradications. In 1946, three permanent plots were established immediately after goats were eradicated from Great Island (Manawa Tawhi), 60 km from northern New Zealand, and provide a 57-year record of change across a sequence of forest succession. Since 1946, the native and non-native bird communities that disperse 75% of the woody flora have increased from six to eight species and bird-dispersed woody plants in plots have increased from 7 to 11 species. After 1946, palatable trees were recruited in the plots. Unpalatable understorey sedges, present when goats were abundant, have persisted and may impede tree seedling establishment. Of the bird-dispersed woody plant species, 41% occur in the plots compared with 67% of the non-bird-dispersed species. Large-seeded species were unable to germinate away from parents until native pigeons Hemiphaga novaeseelandiae were present during the last decade. Forest succession is a consequence of interactions between the legacy of goat grazing and current disperser communities. Survival of seed-limited rare plants is not guaranteed in these circumstances. Although non-native goats no longer influence succession directly, non-native birds have been and remain important components of the disperser community. Our study supports the view that a whole-ecosystem understanding of the interactions between native and non-native species is needed to predict the consequences of eradications on islands worldwide.     

The effect of habitat fragmentation on communities of mutualists: Amazonian ants and their host plants

The consequences of fragmentation for communities of mutualist partners are for the most part unknown; moreover, most studies addressing this issue have been conducted on plant-pollinator communities. We evaluated how the experimental fragmentation of lowland Amazonian rain forest influenced a community of ant-plant mutualists. We inventoried a total of 1057 myrmecophytes in four fragments and four continuous forest sites; the twelve plant species recorded were occupied by 33 ant morphospecies, of which 11 were obligate plant inhabitants. Neither plant species richness, ant species richness, nor total ant-plant density were significantly lower in forest fragments. However, eight of the plant species, including three of the four most common, had higher mean densities in continuous forest than fragments. Of these four species, only one (Cordia nodosa) had significantly different colonization rates between habitat types, with higher colonization rates of plants in fragments. This may be because the Azteca species it is associated with increases in abundance in forest isolates. Although our results suggest that communities of ant-plant mutualists are likely to persist in fragmented tropical landscapes 25 years after fragment isolation, most species are rare and populations sizes in fragments are extremely low. Environmental and demographic stochasticity could therefore limit long-term population viability. We suggest future studies focus on evaluating how fragmentation has altered herbivore pressure and the dispersal of ants and plants to fragments, since the interaction of these factors is likely to have the greatest impact on long-term patterns of population persistence.     

Impacts of urbanization on Florida Key deer behavior and population dynamics

Rapid human population growth and urbanization have had a negative impact on species biodiversity. As competition for resources between man and wildlife continues, it is important to understand the effects of urbanization on species. Endangered Key deer (Odocoileus virginianus clavium) are endemic to the Florida Keys which have undergone rapid human population growth and development over the past 30 years. Our study objectives were to evaluate the impacts of urban development on Key deer habitat use, population dynamics, behavior, and body mass. We used data from two comprehensive studies on Key deer spanning 30 years to evaluate these changes. Our results suggest that Key deer have become more urbanized, using urban areas more today than they did 30 years ago. Contrary to our predictions, survival was higher for more urban deer than for less urban deer. Problems still exist with mortality factors heavily impacting some portions of the deer population including lower survival associated with less urban male deer. Analysis of Key deer body mass also was converse to our predictions as deer weights appear to have increased over time. Collectively, our results suggest that over the past 30 years Key deer have become more urbanized and that deer plasticity has allowed them to adapt and persist in an urbanizing environment. However, the future ability of Key deer to persist in an environment with continued urban development is unknown. At some threshold, urban development would become unsustainable and unlike other forms of habitat change or environmental disturbances, urban development is in most cases irreversible requiring careful planning in habitat conservation strategies.     

Husking stations provide evidence of seed predation by introduced rodents in Tongan rain forests

The islands of Tonga, in Western Polynesia, lacked rodents prior to human colonization. During the past 3000 years, people have introduced three species of rats (Rattus). Seed predation by rodents structures plant communities in many parts of the world. Therefore, there is the potential for rats to have a significant impact on the regeneration of plant communities in Tonga. We found 53 husking stations—sheltered sites used by rats to strip inedible protective structures (husks) from the seeds they eat—on eight islands in Tonga. The stations contained 13,555 empty husks from at least 18 plant species, but only 165 viable seeds and seedlings. These husking stations provide evidence of seed predation by rats. Determining the extent to which seed predation threatens plant regeneration will be important for conservation of native forest remnants.     

Relationship between soil enzyme activities, nutrient cycling and soil fungal communities in a northern hardwood forest

Soil fungi are highly diverse and act as the primary agents of nutrient cycling in forests. These fungal communities are often dominated by mycorrhizal fungi that form mutually beneficial relationships with plant roots and some mycorrhizal fungi produce extracellular and cell-bound enzymes that catalyze the hydrolysis of nitrogen (N)- and phosphorus (P)- containing compounds in soil organic matter. Here we investigated whether the community structure of different types of mycorrhizal fungi (arbuscular and ectomycorrhizal fungi) is correlated with soil chemistry and enzyme activity in a northern hardwood forest and whether these correlations change over the growing season. We quantified these relationships in an experimental paired plot study where white-tailed deer (access or excluded 4.5 yrs) treatment was crossed with garlic mustard (presence or removal 1 yr). We collected soil samples early and late in the growing season and analyzed them for soil chemistry, extracellular enzyme activity and molecular analysis of both arbuscular mycorrhizal (AM) and ectomycorrhizal/saprotrophic fungal communities using terminal restriction fragment length polymorphism (TRFLP). AM fungal communities did not change seasonally but were positively correlated with the activities of urease and leucine aminopeptidase (LAP), enzymes involved in N cycling. The density of garlic mustard was correlated with the presence of specific AM fungal species, while deer exclusion or access had no effect on either fungal community after 4.5 yrs. Ectomycorrhizal/saprotrophic fungal communities changed seasonally and were positively correlated with most soil enzymes, including enzymes involved in carbon (C), N and P cycling, but only during late summer sampling. Our results suggest that fine scale temporal and spatial changes in soil fungal communities may affect soil nutrient and carbon cycling. Although AM fungi are not generally considered capable of producing extracellular enzymes, the correlation between some AM taxa and the activity of N acquisition enzymes suggests that these fungi may play a role in forest understory N cycling.     

Soil arthropods beneficially rather than detrimentally impact plant performance in experimental grassland systems of different diversity

Impacts of belowground insecticide application on plant performance and changes in plant community structure almost uniformly have been ascribed to reduced belowground herbivory, although recent studies reported distinct side effects on detritivore soil animals, particularly on Collembola. Consequently, it remains controversial if the resulting soil feedbacks on plants are due to alterations in arthropod herbivory or to changes in the activity of detritivores. We investigated the impacts of the application of a commonly used belowground insecticide (chlorpyrifos) on soil animals and soil feedbacks on model plant species representing two main plant functional groups of grassland communities, the grass Lolium perenne and the forb Centaurea jacea.Insecticide application decreased soil insect herbivore densities considerably. However, also Collembola densities and diversity decreased markedly due to insecticide application and this was most pronounced in Entomobryidae, Isotomidae, Hypogastruridae, and Sminthuridae. While densities of other detritivore taxa were not affected or even increased (Oribatida) in insecticide subplots, that of predators mostly decreased.Both model plant species built considerably more biomass in control subplots than in insecticide subplots irrespective of characteristics of the resident plant community. This suggests that soil feedbacks on plants were not due to belowground herbivory and highlights the significance of alternative mechanisms responsible for insecticide-mediated soil feedbacks on plants. The deterioration of model plant species’ performances in insecticide subplots most likely was due to decreased densities of Collembola resulting in the deceleration of nutrient cycling and plant nutrition. The results suggest that it is oversimplistic to only ascribe insecticide-mediated soil feedbacks on plants to belowground herbivores. The results further indicate that in the present study the impact of arthropod detritivores on plant productivity was more important than that of belowground herbivores. This emphasizes that plant-soil arthropod interactions in grassland might be based on both facilitative and antagonistic interrelationships.     

Effects of sika deer on vegetation in Japan: A review

The Japanese sika deer Cervus nippon has expanded its range by nearly 70% during the last two decades. Browsing by sika deer affects vegetation in both agricultural and forested habitats. Effects of sika deer on vegetation are conspicuous on deer-inhabited islands: forest structure and composition are altered by deer grazing and browsing, and consequently regeneration is prevented. By felling of old trees, forest gaps are formed, but since sapling recruitment is prevented, shade intolerant plants invade. Unpalatable forbs like ragwort Senecio cannabifolius and ferns like bracken Pteridium aquilinum (Dennstaedtiaceae) become dominants in open habitats. At the places of highest deer density, the Zoysia japonica community, a low growing grass mat, develops. Sika deer function as a seed dispersal agent of this grass. Indirect effects of sika deer are not well studied, but some studies have shown reduction of understory bamboo cover, which in turn improves the survival of tree seedlings and declines of wood mice Apodemus spp. Japanese forests in lower mountainous areas were widely logged during the 1940s and 1950s. These areas were not well planted during the war and until 1950, but thereafter intensively planted to alter to conifer plantation as a nationwide campaign. For several decades after the war, rodents and hares grazed planted trees. According to growth of the planted trees to form canopy, which is unfavorable for rodents and hares, their damage declined. After the 1960s, old-growth forests in high mountainous areas were logged, and conifers were planted. Animals causing forestry damage were replaced by sika deer during the 1980s. Sika deer eat a wide variety of foods, and are gregarious, which causes heavy impacts on vegetation. Effects of sika deer are expanding to natural forests, alpine vegetation, and marshes. To reduce damages on forestry and natural vegetation, as many as 100,000 sika deer are culled every year. However, the hunter population is rapidly declining, and it is expected population control by culling will be insufficient. Although sika deer are an important component of Japanese forests, current population densities exceed the capacity of many ecosystems to tolerate herbivory.     

COLONIZATION PATTERNS OF WOODY SPECIES ON LANDS MINED FOR COAL IN SPAIN: PRELIMINARY INSIGHTS FOR FOREST EXPANSION

In this study, we combined the analysis of a coenocline with Huisman–Olff–Fresco (HOF) modelling of species behaviour to describe colonization patterns of woody species on reclaimed coal wastes in northern Spain. We hypothesize that fine‐scale variation in abiotic factors along the forest‐grassland gradient that appears after mining restoration affects colonization pattern of woody species. To understand these factors, we analysed changes in floristic composition and abundance of the main woody colonizers according to the spatial distance from forest limit, as well as the spatial variability of some edaphic and microclimatic parameters. Our results showed how the primary coenocline extracted from DCA1 reflected a gradient in environmental conditions influencing the species composition of the woody plant communities, from the more shade, moist and dense forest on acid and rich soils to the open, dry and oligotrophic grassland. The colonization pattern of woody species is affected by fine‐scale variations in abiotic factors, such as the increase in pH, P, light, and the decrease in K, N, C/N, organic matter and soil moisture, from the forest to the restored mine. Most species HOF modelled showed monotone responses with decreasing trends along the environmental gradient from forest to mine, being Quercus petraea the species clearly dominant in the forest. On the contrary, Cytisus scoparius and Genista florida colonize intensely in the mine area, reaching the maximum abundance around 15 and 10 m distance from the forest limit, respectively. We conclude that the composition of woody species in the forest is not a good predictor of the colonizing intensity, not even of the species composition in the mine area. Copyright © 2011 John Wiley & Sons, Ltd.     

Endangered crayfish in northern Japan: Distribution, abundance and microhabitat specificity in relation to stream and riparian environment

Japan’s only native crayfish species Cambaroides japonicus has been declining dramatically in the past few decades. For the purpose of conservation planning, twenty-two coastal streams were surveyed to investigate summer distributions of crayfish in relation to stream and riparian environment. Classification and regression trees were used to predict the occurrence and abundance of crayfish. The classification tree model with stream variables as predictors showed that crayfish would occur in swift or high gradient streams (correct classification rate = 91%). Within those streams, however, crayfish only inhabited depositional microhabitats, in which the areas are limited in availability. Crayfish were not found in gentle, low gradient streams containing abundant depositional microhabitats. This paradoxical distribution pattern was attributed to availability of boulder substrates in swift or high gradient streams. The regression tree model indicated that crayfish abundance was determined primarily by the percentage of boulder substrates and the presence of fish (observed vs. predicted r = 0.64).The classification tree model using only riparian variables indicated that the total woody plant (mainly broadleaf species) density followed by the percentage of early successional species such as alder and willow determined the splits of the tree model (correct classification rate = 95%). A leaf processing experiment on 10 riparian plant species suggested that crayfish preferred high nitrogen (or low C/N) leaves.These results suggest that swift or high gradient fishless streams associated with abundant cover in dense broadleaf forest serve conservation areas for this endangered crayfish, and that consideration of riparian composition may facilitate conservation efforts.     

Conservation in a city: Do the same principles apply to different taxa?

Factors shaping overall species richness and representation of endangered species of vascular flora and butterflies were determined in 48 nature reserves in Prague, Czech Republic. Total species richness of both groups, and the presence of endangered butterflies, reflect the present status of habitats, while that of endangered plants reflects habitat composition at the time of reserve establishment. Reserve area has a much stronger effect on the species richness of sessile plants than mobile butterflies which, especially endangered species, respond more positively to heterogeneity than to area. Both species richness and endangered species peak in reserves half covered with forest, likely because they harbor species of both woodland and non-woodland habitats, and edge specialists. Solely relying on area and disregarding habitat quality, or a failure to conserve small but high quality sites, may be counterproductive for conserving endangered plant and butterfly species. To conserve diversity of plants and butterflies current management practices, often aimed at maintaining large blocks of pre-selected plant communities, should change to include maintenance of fallows, edges and transient zones.     

Impacts of insect herbivores on rare plant populations

Population-level impacts of insect herbivory on rare and endangered plants are poorly understood, being frequently suggested but seldom quantified. We compiled 37 studies from the primary literature encompassing 35 rare plant species and at least 63 insect herbivore species, and assessed patterns in the effects of insect herbivores on rare plant populations and vital rates. Only three studies directly quantified effects of insect herbivory on population size or growth rate, with the vast majority instead extrapolating from vital rates to population size. Few studies (eight) studied herbivory experimentally, and feeding guilds other than seed predators have been relatively neglected. Estimates of population-level impacts vary extremely widely, but for many rare plants insect herbivores cause major reductions in survivorship or fecundity or can even drive local extinctions. Four studies documented positive (plant-) density dependence in insect attack, suggesting that herbivores may have a stabilizing influence on plant populations and thus may play a role in regulating rare plant population size. Most reported herbivores of rare plants are oligophagous or polyphagous, but monophagous herbivores of rare plants do exist, and there was no detectable difference in impact among specialization classes. Attack on rare plants by escaped biocontrol agents can sometimes have significant population-level effects, but such cases appear uncommon. Because insect herbivory can strongly suppress rare plant populations or vital rates, we need to know much more about what ecological and evolutionary factors determine the population-level impacts of insect herbivores. We suggest several ways in which the results of such research could inform conservation practices for rare plant species – for example, the possibility of different management regimes for plants under attack by monophagous, oligophagous, and polyphagous herbivores.     

Population demise and extinction threat of the Angel de la Guarda deer mouse (Peromyscus guardia)

We analyze the status of the Angel de la Guarda deer mouse (Peromyscus guardia), a species endemic to Mexico, based on our own fieldwork, bibliographic records and information from colleagues. This species, with different subspecies on three islands and a population of an undetermined subspecies on a fourth, was apparently common until the mid-1960s. Currently the species is critically endangered, if not already extinct. One of the subspecies and the undetermined population are presumed extinct. Of the two other subspecies, one is at least in critical condition, and the other at least reduced. The demise of the species can be attributed to the introduction of domestic cats. The case of P. guardia is a good example of the vulnerability of other taxa of endemic rodents on islands in the Gulf of California.     

黄土高原主要树(草)种热值试验

能源林主要是生产热能,树（草）种燃烧值的高低,是评选能源林树（草）种的重要指标。通过对黄土高原29种树（草）种燃烧值的测定表明,木本植物的燃烧值大于草本植物,这种能量差异主要是草本植物含有较多的灰分,结构物质纤维素含量较高,木本植物灰分含量较低,脂类物质相对较多。同时发现10种树种的热值随着年龄的变化多以5年生时的热值为最高,这对确定树种的平茬期,在生产上具有重要意义。