综述：世界草原上的社会性、穴居性、食草性哺乳动物的生态作用和保护挑战

 

frontiers in ecology and the environment（13.780/Q1）Ecological roles and conservation challenges of social, burrowing, herbivorous mammals in the world’s grasslands

摘要The world’s grassland ecosystems are shaped in part by a key functional group of social, burrowing, herbivorous mammals. Through herbivory and ecosystem engineering they create distinctive and important habitats for many other species, thereby increasing biodiversity and habitat heterogeneity across the landscape. They also help maintain grassland presence and serve as important prey for many predators. However, these burrowing mammals are facing myriad threats, which have caused marked decreases in populations of the best-studied species, as well as cascading declines in dependent species and in grassland habitat. To prevent or mitigate such losses, we recommend that grasslands be managed to promote the compatibility of burrowing mammals with human activities. Here, we highlight the important and often overlooked ecological roles of these burrowing mammals, the threats they face, and future management efforts needed to enhance their populations and grassland ecosystems.

世界上的草原生态系统部分是由一群社会性的、穴居的、食草的哺乳动物组成的关键功能群所塑造的通过食草和生态系统工程，它们为许多其他物种创造了独特而重要的栖息地，从而增加了整个景观的生物多样性和栖息地的异质性。

它们还帮助维持草原的存在，并作为许多捕食者的重要猎物然而，这些穴居哺乳动物正面临着无数的威胁，这些威胁造成了研究得最好的物种种群的明显减少，以及依赖物种和草原生境的连带减少为了防止或减轻这种损失，我们建议对草原进行管理，以促进穴居哺乳动物与人类活动的兼容。

在此，我们强调了这些穴居哺乳动物重要而又经常被忽视的生态作用，它们所面临的威胁，以及为提高它们的种群数量和草原生态系统所需的未来管理努力In a nutshell:简而言之：• Social, burrowing, herbivorous mammals play important functional roles in grasslands around the world

社会性的、穴居的、食草的哺乳动物在世界各地的草原上发挥着重要的功能作用• These mammals face many threats, including intentional poisoning, exotic diseases and pests, overhunting, habitat loss, and climate change。

这些哺乳动物面临许多威胁，包括故意投毒、外来疾病和害虫、过度捕猎、栖息地丧失和气候变化• Grassland management must include promoting sufficient numbers of burrowing mammals to fulfill their landscape-scale functional roles, so as to maintain the health and biodiversity of grassland systems and the ecosystem services they provide。

草地管理必须包括促进足够数量的穴居哺乳动物，以发挥它们在景观尺度上的功能作用，从而保持草地系统的健康和生物多样性以及它们提供的生态系统服务Grassland ecosystems worldwide are fundamentally shaped by an underappreciated but key functional group of social, semi-fossorial (adapted to burrowing and living underground), herbivorous mammals (hereafter, burrowing mammals). Examples include not only the phylogenetically similar species of prairie dogs of North America (Cynomys spp); ground squirrels (Family Sciuridae) of North America, Eurasia, and Africa; and marmots (Marmota spp) of North America and Eurasia, but also the more distantly related but functionally similar plains vizcachas (Lagostomus maximus), Patagonian maras (Dolichotis patagonum), and degus (Octodon degus) of South America; pikas (Ochotona spp) of Asia; ice rats (Otomys sloggetti) and springhares (Pedetes capensis) of Africa; and burrowing bettongs (Bettongia lesueur) and southern hairy-nosed wombats (Lasiorhinus latifrons) of Australia (。

Figure 1). Often living in colonies ranging from tens to thousands of individuals, these mammals collectively transform grassland landscapes through their burrowing and feeding activity. By grouping together socially, they also create distinctive habitat patches that serve as areas of concentrated prey for many predators. Their ecosystem engineering and trophic effects help maintain grassland biodiversity and, consequently, they frequently play keystone roles in these ecosystems (

Figure 2).世界范围内的草原生态系统从根本上说是由一个未被重视但却具有关键功能的社会性、半化石性（适应于钻洞和生活在地下）的食草哺乳动物（以下简称钻洞哺乳动物）所形成这方面的例子不仅包括系统发育相似的北美。

草原土拨鼠（Cynomys spp）；北美、欧亚和非洲的地鼠（Sciuridae科）；北美和欧亚的旱獭（Marmota spp），还包括关系较远但功能相似的南美平原鼠（Lagostomus maximus）、

巴塔哥尼亚鼠（Dolichotis patagonum）和八齿鼠（Octodon degus）；亚洲的鼠兔（Ochotona spp）；非洲的冰鼠（Otomys sloggetti）和跳兔（Pedetes capensis）；以及

澳大利亚的穴居袋鼠（Bettongia lesueur）和南澳毛吻袋熊（Lasiorhinus latifrons）（图1）这些哺乳动物通常生活在几十到几千只的群落中，通过它们的钻洞和觅食活动，共同改变草原景观。

。通过社会性的聚集，它们也创造了独特的生境斑块，成为许多捕食者集中捕食的区域。它们的生态系统工程和营养效应有助于维持草原的生物多样性，因此，它们经常在这些生态系统中发挥关键作用（图2）。

Figure 1. Examples of social, burrowing, herbivorous mammals from grasslands around the world. North America: California ground squirrels, black-tailed and Gunnison’s prairie dogs; Eurasia: European ground squirrels, Himalayan and Siberian marmots, plateau pikas; Australia: burrowing bettongs, southern hairy-nosed wombats; Africa: ice rats, Cape ground squirrels, springhares; South America: degus, Patagonian maras, plains vizcachas. Map from World Resources Institute (White et al. 2000). (See

WebTable 1 for species names and WebPanel 1 for photo credits.)图1. 世界各地草原上的社会性、穴居性、食草性哺乳动物的例子北美洲：加利福尼亚地松鼠，黑尾和冈尼森草原犬；。

欧亚大陆：欧洲地松鼠，喜马拉雅和西伯利亚旱獭/土拨鼠，高原鼠兔；澳大利亚：钻地鼠，南澳毛吻袋熊；非洲：冰鼠，南非地松鼠，跳兔；南美洲：八齿鼠，巴塔哥尼亚鼠，南美平原鼠地图来自世界资源研究所（White等人，2000）。

(物种名称见WebTable 1，图片来源见WebPanel 1）

Figure 2. Conceptual diagram showing the trophic (herbivory, prey) and ecosystem engineering (clipping, burrow construction, and mound building) effects of burrowing mammals on grassland ecosystems, based on the best-studied species: the black-tailed prairie dog in North America. Plus signs indicate an increase; minus signs indicate a decrease. Black arrows depict the effects of burrowing mammals (eg prairie dogs), green arrows depict the impacts of megaherbivores (eg bison), and the red arrow indicates the role of predators. (Drawings provided by SN Davidson.)

图2. 概念图显示了穴居哺乳动物对草原生态系统的营养（食草，猎物）和生态系统工程（剪枝，洞穴建造和土丘建造）的影响，基于研究的最好的物种：北美的黑尾土拨鼠正号表示增加；负号表示减少黑色箭头表示穴居哺乳动物（如草原土拨鼠）的影响，绿色箭头表示大型食草动物（如野牛）的影响，红色箭头表示捕食者的作用。

(图纸由SN Davidson提供）Despite their importance to grassland ecosystems, burrowing mammal populations have declined dramatically, primarily as a result of human impacts; indeed, because grasslands provide the world’s most important habitat for agricultural and livestock production, burrowing mammals are often in direct conflict with human activities (Smith et al. 2006; Miller et al. 2007; Delibes-Mateos et al. 2011). Human-mediated introductions of exotic species, disease agents, and overhunting are also reducing their populations (Branch et al. 2002; Gage and Kosoy 2005; Wingard and Zahler 2006; Rodriguez 2009). The population dynamics and ecological roles of most burrowing mammal species remain poorly understood, however. The patterns discussed here apply to most, but not necessarily all, of the species mentioned. What is known about the few well-studied species suggests that burrowing mammals likely play widespread and important ecological roles, and that their loss can have cascading detrimental effects on the grassland ecosystems on which both humans and wildlife depend.

尽管穴居哺乳动物对草原生态系统很重要，但它们的数量已经急剧下降，这主要是由于人类的影响；事实上，由于草原为农业和畜牧业生产提供了世界上最重要的生境，穴居哺乳动物经常与人类活动发生直接冲突(Smith等人, 2006; Miller等人, 2007; Delibes-Mateos等人, 2011)。

人类介导的外来物种的引入、疾病媒介和过度捕猎也在减少它们的数量(Branch等人，2002；Gage和Kosoy，2005；Wingard和Zahler，2006；Rodriguez，2009)然而，大多数穴居哺乳动物物种的种群动态和生态作用仍然鲜为人知。

这里讨论的模式适用于大多数，但不一定是所有提到的物种对少数研究良好的物种的了解表明，穴居哺乳动物可能发挥着广泛而重要的生态作用，它们的消失会对人类和野生动物赖以生存的草原生态系统产生连带的不利影响An important challenge facing grassland managers is maintaining the important functional roles of these burrowing mammals in ways that are compatible with human activities. Here, we present a conceptual model (。

Figure 2) that illustrates the common but underappreciated roles that burrowing mammals play in grasslands and highlight the widespread threats facing these species. We also outline future directions required for their conservation and management.

草原管理者面临的一个重要挑战是以与人类活动相适应的方式维持这些穴居哺乳动物的重要功能作用在此，我们提出了一个概念模型（图2），说明了穴居哺乳动物在草原上发挥的常见但未被重视的作用，并强调了这些物种面临的广泛威胁。

我们还概述了保护和管理它们所需的未来方向Ecological roles of burrowing mammals穴居哺乳动物的生态作用Because they tend to cluster in social groups or colonies, burrowing mammals create islands of open grassland habitat that differ from the surrounding landscape and that attract numerous animals (

Figure 3). Many burrowing mammals preferentially forage on grasses, thereby facilitating the establishment of forbs; this foraging activity also creates a low mat of grazing-tolerant grasses and forbs within their colonies (

Figure 3; Whicker and Detling 1988; Branch et al. 1996b; Yoshihara et al. 2009). Burrowing mammals also dig underground tunnels, which provide dens and shelter for many animals. This digging

activity creates aboveground patches of disturbed soil that can vary in size and type: from open areas surrounding pika burrows (≤ 0.5 m

2); to aboveground soil mounds of 0.5–3 m2 around prairie dog, marmot, and ground squirrel burrows; to 15–65 m

2 soil mounds created by wombats and bettongs; and to vizcacha mounds that are 300–700 m2with up to 100 burrow entrances (

Figure 4; Steele and Temple-Smith 1998; Noble et al. 2007b; Wesche et al. 2007; A Smith and L Branch pers comm). Some burrowing mammals, such as prairie dogs, create numerous small mounds, while others, such as vizcachas and bettongs,

construct one large mound that houses the entire colony or family group (Figure 4). Each mound, and often also the colony, provides distinctive habitat that supports plant and animal assemblages that differ from those in the surrounding grassland (Branch et al. 2002; Komonen et al. 2003; Davidson and Lightfoot 2006). Although their colonies and mounds may be more or less species-rich than adjacent grassland (Lenihan 2007; Noble et al. 2007b; Yoshihara et al. 2009), these distinctive habitat patches increase overall heterogeneity and biodiversity at multiple scales across the landscape (

Figure 5; Whicker and Detling 1988; Davidson et al. 2008; Hogan 2010).由于它们倾向于聚集在社会群体或群落中，穴居哺乳动物创造了与周围景观不同的开放草原生境岛屿，并吸引了许多动物

（图3）许多穴居哺乳动物喜欢在草地上觅食，从而促进了草本植物的建立；这种觅食活动也在它们的群落中创造了低矮的耐牧草和草本植物垫（图3；Whicker and Detling 1988; Branch等人1996b; Yoshihara等人2009）。

穴居哺乳动物还挖掘地下隧道，为许多动物提供巢穴和庇护所这种挖掘活动创造了地上的干扰土壤斑块，其大小和类型各不相同：从鼠兔洞穴周围的开放区域（≤0.5平方米）；到地上土堆的0. 5-3平方米的地上土堆，围绕着草原土拨鼠、旱獭和地鼠的洞穴；15-65平方米的土堆，由袋熊和地鼠创造；还有300-700平方米的兔鼠土堆，有多达100个洞穴入口(图4；Steele 和 Temple-Smith 1998; Noble等人 2007b; Wesche等人 2007; A Smith和L Branch pers comm)。

一些穴居哺乳动物，如草原土拨鼠，创造许多小土丘，而其他的，如兔鼠和地鼠，建造一个大土丘，容纳整个族群或家庭群体（图4）每个土丘，通常也是族群，提供独特的栖息地，支持与周围草原不同的植物和动物组合(Branch等人, 2002; Komonen等人, 2003; Davidson and Lightfoot, 2006)。

虽然他们的殖民地和土丘可能比邻近的草地更多或更少的物种丰富(Lenihan 2007; Noble et al. 2007b; Yoshihara et al. 2009)，这些独特的生境斑块增加了整个景观多个规模的整体异质性和生物多样性

(图5; Whicker and Detling 1988; Davidson et al. 2008; Hogan 2010)。

Figure 3. Photos showing the open habitats that burrowing mammals help create. (a) Black-tailed prairie dogs, (b) European ground squirrel, (c) plateau pika, (d) ice rat, (e) Patagonian maras and their habitat, and (f) grass understory heavily grazed by vizcachas. (See WebPanel 1 for photo credits.)

图3. 显示穴居哺乳动物帮助创造的开放栖息地的照片(a)黑尾土拨鼠，(b)欧洲地鼠，(c)高原鼠兔，(d)冰鼠，(e)巴塔哥尼亚马拉斯及其栖息地，以及(f)被兔鼠过渡啃食的草丛(照片来源见网络面板1）

Figure 4. Photos showing the different sizes and types of soil disturbances that burrowing mammals create when building their underground burrows. (a) Plateau pika burrows, (b) black-tailed prairie dog mounds, and (c) a southern hairy-nosed wombat warren. (See WebPanel 1 for photo credits.)

图4. 照片显示了穴居哺乳动物在建造地下洞穴时造成的不同大小和类型的土壤扰动。(a) 高原鼠兔的洞穴，(b) 黑尾土拨鼠的土堆，和(c) 南澳毛吻袋熊的巢穴。(照片来源见网络面板1）。

Figure 5. Diagram illustrating the distinctive islands of habitat that burrowing mammals create across multiple spatial scales with their (a) mounds, (b) individual colonies, and (c) colony complexes. These features enhance habitat heterogeneity and, consequently, increase biodiversity across grassland landscapes. This illustration is based on black-tailed prairie dogs in the Great Plains grasslands of North America. (Drawing provided by SN Davidson.)

图5. 图中说明了穴居哺乳动物用它们的（a）土墩，（b）单个殖民地，和（c）殖民地复合体在多个空间尺度上创造的独特的生境岛这些特征增强了生境的异质性，从而增加了整个草原景观的生物多样性这幅插图是基于北美大平原草原上的黑尾土拨鼠。

(绘图由SN Davidson提供）Burrowing mammals often move large amounts of soil during burrow construction. For instance, Arctic ground squirrels (Spermophilus parryii) and wombats can move as much as 20 metric tons and 1.3–6.0 metric tons, respectively, of soil per hectare (Price 1971; James and Eldridge 2007). The underground systems of burrowing mammals

provide an important ecosystem service by facilitating water infiltration (eg Kotliar et al. 2006; Hogan 2010). Soil mixing and urine and fecal deposition around mounds can also increase soil organic matter and inorganic nutrients (Noble et al. 2007b; Wesche et al. 2007; Villarreal et al. 2008).

穴居哺乳动物在建造洞穴时经常移动大量的土壤例如，北极地松鼠(Spermophilus parryii)和袋熊每公顷可以移动多达20公吨和1.3-6.0公吨的土壤(Price 1971; James and Eldridge 2007)。

穴居哺乳动物的地下系统通过促进水的渗透，提供了一个重要的生态系统服务（如Kotliar等人，2006；Hogan 2010）土丘周围的土壤混合和尿液和粪便沉积，也可以增加土壤有机物和无机营养物质(Noble等人, 2007b; Wesche等人, 2007; Villarreal等人, 2008)。

Burrowing mammals help maintain grasslands and open habitat. Species like vizcachas, burrowing bettongs, and prairie dogs prevent invasion and establishment of shrubs through their foraging and clipping (ie pruning) (Weltzin et al. 1997; Branch et al. 2002; Noble et al. 2007a). Shrubland expansion in semi-arid regions of North America and Australia has been attributed, in part, to population reductions of prairie dogs and bettongs, respectively (Weltzin et al. 1997; Noble et al. 2007a).

穴居哺乳动物有助于维持草地和开放的生境像兔鼠，钻地鼠和草原土拨鼠，通过它们的觅食和修剪，防止灌木的入侵和建立(Weltzin等人, 1997; Branch等人, 2002; Noble等人, 2007a). 北美和澳大利亚的半干旱地区的灌木地扩张，部分原因是草原土拨鼠和兔鼠的数量减少(Weltzin等人, 1997; Noble等人, 2007a)。

Herbivory and burrowing activities can reduce overall plant biomass associated with burrowing mammal colonies, but the higher levels of soil nutrients and greater degree of water infiltration that occurs around their mounds can result in elevated foliar nutrient concentrations and greater plant biomass surrounding their burrows (Retzer 2007; Van Staalduinen and Werger 2007; Hogan 2010). Grazing by burrowing mammals also enhances plant nitrogen uptake, resulting in increased forage quality on their colonies (Whicker and Detling 1988; Wesche et al. 2007; Villarreal et al. 2008). Increased forage quality apparently attracts megaherbivores, such as bison (Bison bison) and cattle (Bos taurus), to prairie dog colonies in North America (Whicker and Detling 1988; Davidson et al. 2010); grazing and defecation by such megaherbivores further increases forage quality and decreases vegetation height (Whicker and Detling 1988). Consequently, megaherbivores and small burrowing mammals can have mutualistic relationships (Krueger 1986; Davidson et al. 2010). Indeed, grazing by megaherbivores facilitates increases in population densities of burrowing mammal species that prefer more open grassland, thereby increasing their overall impact on the ecosystem. This relationship has been observed among native megaherbivores and many burrowing mammals, such as that between bison and prairie dogs, as well as among livestock species and prairie dogs, vizcachas, Patagonian maras, springhares, marmots, pikas, Brandt’s voles (Lasiopodomys brandtii), and long-tailed ground squirrels (Urocitellus undulatus; Jackson 1988; Augustine et al. 1995; Ronkin et al. 2009). Although burrowing mammals often benefit from grazing by megaherbivores, overgrazing by the latter may result in food limitation and desertification, frequently leading to population declines in burrowing mammals (Komonen et al. 2003; Read et al. 2008; Rodriguez 2009).

食草和钻洞活动可以减少与穴居哺乳动物群落有关的整体植物生物量，但它们的土丘周围有较高的土壤养分水平和更大程度的水渗入，可以导致它们的洞穴周围的叶子养分浓度升高，植物生物量增加(Retzer 2007; Van Staalduinen and Werger 2007; Hogan 2010)。

穴居哺乳动物的放牧，也会加强植物氮的吸收，导致其殖民地的饲料质量增加(Whicker and Detling 1988; Wesche等人 2007; Villarreal等人 2008)饲料质量的提高，显然吸引大型食草动物，如野牛(Bison bison)和牛(Bos taurus)，到北美的草原土拨鼠殖民地(Whicker and Detling 1988; Davidson et al. 2010)；这些。

大型食草动物的放牧和排泄，进一步增加饲料质量，降低植被高度(Whicker and Detling 1988)因此，大型食草动物和小型穴居哺乳动物可以有相互的关系(Krueger 1986; Davidson等人, 2010)。

事实上，大型食草动物的放牧促进了喜欢更开放草地的穴居哺乳动物种群密度的增加，从而增加了它们对生态系统的整体影响这种关系在本地大型食草动物和许多穴居哺乳动物之间已经被观察到，例如野牛和草原土拨鼠之间的关系，以及牲畜物种和草原土拨鼠、兔鼠、巴塔哥尼亚豚鼠（Patagonian maras）、跳兔、旱獭、鼠兔、布氏田鼠（Lasiopodomys brandtii）和长尾地松鼠（Urocitellus undulatus; Jackson 1988; Augustine等人1995; Ronkin等人2009）。

虽然穴居哺乳动物经常受益于大型食草动物的放牧，但后者的过度放牧可能导致食物限制和沙漠化，经常导致穴居哺乳动物的数量下降(Komonen等人, 2003; Read等人, 2008; Rodriguez 2009)。

The excavations created by burrowing mammals provide important belowground habitat for many grassland animals. For example, burrows can increase overall arthropod diversity and abundance by two- to threefold (Davidson and Lightfoot 2007; Read et al. 2008), and herpetofauna commonly use such burrows to escape from predators and as breeding sites (WebTable 1; Smith and Foggin 1999; Davidson et al. 2008; Hogan 2010). These burrows are also important for other rodents, such as spotted ground squirrels (Xerospermophilus spilosoma), northern grasshopper mice (Onychomys leucogaster), and Bennett’s chinchilla rats (Abrocoma bennettii), and groundnesting birds, such as burrowing owls (Athene cunicularia), miners (Geositta cunicularia), swallows (Notiochelidon cyanoleuca), ground jays (Pseudopodoces humilis), snowfinchs (Montifringilla spp and Pyrgilauda spp), and ant-eating chats (Myrmecocichla formicivora) (

WebTable 1). Several mustelids, herpestids, felids, and canids that prey on burrowing mammals also inhabit their burrows (Branch et al. 2002; Waterman and Roth 2007; Murdoch et al. 2009).

穴居哺乳动物创造的洞穴，为许多草原动物提供重要的地下栖息地例如，洞穴可以增加整体节肢动物的多样性和丰度2至3倍(Davidson and Lightfoot 2007; Read et al. 2008)，爬行动物通常使用这种洞穴来逃避捕食者和作为繁殖地(WebTable 1; Smith and Foggin 1999; Davidson et al. 2008; Hogan 2010)。

这些洞穴对其他啮齿类动物也很重要，如斑点地松鼠(Xerospermophilus spilosoma)，白腹食蝗鼠(Onychomys leucogaster)，和班尼特龙猫鼠(Abrocoma bennettii)，和地巢鸟。

如穴居猫头鹰（Athene cunicularia），掘穴雀（Geositta cunicularia），燕子（Notiochelidon cyanoleuca），地鸦（Pseudopodoces humilis），雪雀（Montifringilla spp和Pyrgilauda spp），和食蚁鸟（Myrmecocichla formicivora）（WebTable 1）。

一些捕食穴居哺乳动物的鼬科动物、牧羊犬科动物、猫科动物和犬科动物也居住在它们的洞穴中(Branch等人, 2002; Waterman and Roth 2007; Murdoch等人, 2009)Aboveground, burrow mounds attract many animals, including unique grasshopper and ground-dwelling arthropod species assemblages, lizards that use mounds for basking, and megaherbivores like bison that wallow in the disturbed soil (Coppock et al. 1983; Davidson and Lightfoot 2007; Davidson et al. 2008). At the landscape scale, colonies of burrowing mammals attract species that prefer open grassland habitat, such as lesser earless lizards (Holbrookia maculata) on prairie dog colonies (Davidson et al. 2008), and bird species, such as mountain plovers (Charadrius montanus), long-billed curlews (Numenius americanus), and burrowing owls (Branch et al. 2002; Kotliar et al. 2006). Open grassland habitat increases the ability of animals to detect predators. This benefits coexisting burrowing species like Patagonian maras, yellow mongoose (Cynictis penicillata), meerkats (Suricata suricatta), and spotted ground squirrels that also profit from the sociality of, and predator detection by, the burrowing mammals with which they associate (Waterman and Roth 2007; Villarreal et al. 2008; AD Davidson pers observ). The greater abundance of forbs and dwarf shrubs associated with colonies also attracts ungulates like pronghorn (Antilocapra americana) (Krueger 1986), and the high floral densities and open soil habitat (used for nesting) on colonies increase the abundance and diversity of insect pollinators (Hardwicke 2006; Yoshihara et al. 2010a).

在地面上，洞穴土丘吸引许多动物，包括独特的蚱蜢和地栖节肢动物物种组合，利用土丘晒太阳的蜥蜴，以及像野牛这样的大型食草动物，在被扰乱的土壤中打滚(Coppock等人, 1983; Davidson and Lightfoot, 2007; Davidson等人, 2008)。

在景观尺度上，穴居哺乳动物群吸引喜欢开放草原生境的物种，如土拨鼠移殖地的小无耳蜥蜴(Holbrookia maculata)(Davidson et al. 2008)，以及鸟类物种，如岩鸻(Charadrius montanus)，长嘴杓鹬(Numenius americanus）和穴居猫头鹰（Branch等人，2002；Kotliar等人，2006）。

开放的草地栖息地增加了动物探测捕食者的能力这有利于共存的穴居物种，如巴塔哥尼亚豚鼠、笔尾獴(Cynictis penicillata)、狐獴(Suricata suricatta)和斑点地松鼠，它们也从与它们有联系的穴居哺乳动物的社会性和捕食者探测中受益(Waterman and Roth 2007; Villarreal et al. 2008; AD Davidson pers observ)。

与殖民地有关的更多的草本植物和矮小的灌木，也吸引了有蹄类动物，如美洲羚羊(Antilocapra americana)(Krueger 1986)，殖民地的高花卉密度和开放的土壤生境(用于筑巢)增加了昆虫授粉者的丰度和多样性(Hardwicke 2006; Yoshihara et al. 2010a)。

Because their colonies represent high-density, localized patches of reliable prey, burrowing mammals also attract predators (

Figure 6; WebTable 1). Indeed, the abundance and richness of carnivorous mammalian and avian fauna is often greater in areas where burrowing mammal colonies are located (eg Lai and Smith 2003; Kotliar et al. 2006; Lenihan 2007; Ceballos et al. 2010). Raptors, canids, felids, herpestids, mustelids, and some snakes are common predators of many burrowing mammals, and the predator communities that associate with them are similar across different grassland ecosystems (

Figure 6). For instance, black-backed jackals (Canis mesomelas) and honey badgers (Mellivora capensis) are common predators of springhares and Cape ground squirrels (Xerus inauris) in Africa, whereas coyotes (Canis latrans) and American badgers (Taxidea taxus) are important predators of North American prairie dogs (

WebTable 1). Small cats also prey on burrowing mammals, including bobcats (Lynx rufus) on California ground squirrels (Otospermophilus beecheyi) and prairie dogs, Geoffroy’s cats (Leopardus geoffroyi) on vizcachas, Pallas’s cats (Felis manul) on pikas and long-tailed ground squirrels, and African wild cats (Felis silvestris lybica) on springhares and Cape ground squirrels (

WebTable 1). Raptors like giant eagle owls (Bubo lacteus) prey on springhares (Skinner and Chimimba 2005), while Daurian pikas (Ochotona dauurica) often make up more than 70% of the Eurasian eagle owl’s (Bubo bubo) diet (Smith and Foggin 1999). Wolves (Canis lupus), brown bears (Ursus arctos), mountain lions (Puma concolor), and snow leopards (Uncia uncia) also prey on burrowing mammals (

WebTable 1); for example, mountain lions in South America often rely on vizcachas for up to 50–85% of their diets, while on the Tibetan Plateau, plateau pikas (Ochotona curzoniae) can represent as much as 50% of Tibetan wolf (C lupus chanco) and 78% of brown bear diets (Branch et al. 1996a; Smith and Foggin 1999; Xu et al. 2006).

因为它们的群落代表着高密度、局部的可靠猎物斑块，穴居哺乳动物也会吸引捕食者（图6；WebTable 1）事实上，食肉哺乳动物和鸟类动物群的丰度和丰富度通常在穴居哺乳动物群所在的地区更大（例如Lai和Smith 2003；Kotliar等人2006；Lenihan 2007；Ceballos等人2010）。

猛禽、犬科动物、猫科动物、蛇科动物、鼬科动物和一些蛇类是许多穴居哺乳动物的常见捕食者，与它们相关的捕食者群落在不同的草原生态系统中是相似的（图6）例如，黑背豺（Canis mesomelas）和蜜獾（Mellivora capensis）是非洲跳兔和南非地松鼠（Xerus inauris）的常见捕食者，而郊狼（Canis latrans）和美洲獾（Taxidea taxus）是北美草原土拨鼠的重要捕食者（WebTable 1）。

小型猫科动物也捕食穴居哺乳动物，包括短尾猫（Lynx rufus）捕食加州地松鼠（Otospermophilus beecheyi）和草原土拨鼠，乔氏虎猫（Leopardus geoffroyi）捕食兔鼠，兔狲（Felis manul）捕食鼠兔和长尾地松鼠，以及非洲野猫（Felis silvestris lybica）捕食跳兔和南非地松鼠（网络表1）。

像黄雕鸮(Bubo lacteus)这样的猛禽捕食跳兔(Skinner and Chimimba 2005)，而达乌尔鼠兔(Ochotona dauurica)通常占雕鸮(Bubo bubo)食物的70%以上(Smith and Foggin 1999)。

狼（Canis lupus）、棕熊（Ursus arctos）、美洲狮（Puma concolor）和雪豹（Uncia uncia）也捕食穴居哺乳动物（WebTable 1）例如，南美洲的山狮经常依赖兔鼠，占其食物的50-85%，而在青藏高原，高原鼠兔（Ochotona curzoniae）可占藏狼（C lupus chanco）食物的50%和棕熊食物的78%（Branch et al. 1996a; Smith and Foggin 1999; Xu et al. 2006）。

Figure 6. Broadly similar predator communities associated with burrowing mammals around the world. 图6. 世界各地与穴居哺乳动物相关的大体相似的捕食者群落。

Examples of predators associated with prairie dogs of North America, pikas of Eurasia, springhares of Africa, and plains vizcachas of South America are shown. Map from World Resources Institute (White et al. 2000). (See

WebTable 1 for predator species names and predator communities associated with these and other burrowing mammal species. See

WebPanel 1 for photo credits.)图中显示了与北美洲的草原土拨鼠、欧亚大陆的鼠兔、非洲的跳兔和南美洲的平原兔鼠有关的捕食者的例子地图来自世界资源研究所（White等人，2000）。

(与这些和其他穴居哺乳动物物种相关的捕食者物种名称和捕食者群落见WebTable 1照片来源见网络面板1）Ecosystem engineers and keystone species 生态系统工程师和关键物种

Species considered to be ecosystem engineers physically create, maintain, and modify their environment (Jones et al. 1994), whereas those that have irreplaceable ecological impacts that are large relative to their abundance are referred to as keystone species (Power et al. 1996). Because of the large, transformative effects that burrowing mammals have on grassland ecosystems, they are often considered ecosystem engineers, keystone species, or both (Samjaa et al. 2000; Branch et al. 2002; Lai and Smith 2003; Kotliar et al. 2006). Although the relative importance of burrowing mammal impacts may vary across species, space, and time, these animals play unique and non-substitutable keystone roles (Davidson and Lightfoot 2006). Moreover, although population densities of burrowing mammals can be quite high within their colonies, they are not necessarily abundant across the greater landscape. Consequently, because they create unique patches of important grassland habitat, increase biodiversity across the landscape, and are needed in large numbers to support associated species, the ecological roles of burrowing mammals must be examined from a landscape perspective (Figures 2, 5, and 6; WebTable 1).

被认为是生态系统工程师的物种在物理上创造、维持和改变他们的环境(Jones等人，1994)，而那些具有不可替代的生态影响的物种，相对于他们的丰度来说是很大的，被称为关键物种(Power等人，1996)由于穴居哺乳动物对草原生态系统有很大的改造作用，它们通常被认为是生态系统的工程师，关键物种，或两者都是

(Samjaa等人, 2000; Branch等人, 2002; Lai and Smith 2003; Kotliar等人, 2006)虽然穴居哺乳动物影响的相对重要性可能因物种、空间和时间而异，但这些动物发挥着独特的、不可替代的关键作用（Davidson和Lightfoot，2006）。

此外，尽管穴居哺乳动物的种群密度在它们的群落中可能相当高，但它们在整个大景观中不一定很丰富因此，由于它们创造了独特的重要草原生境斑块，增加了整个景观的生物多样性，并且需要大量的数量来支持相关的物种，所以必须从景观的角度来研究穴居哺乳动物的生态作用

（图2，5和6；WebTable 1）Conservation status and current threats保护状况和当前威胁The conservation status of burrowing mammals varies greatly across species, but populations of most species are heavily and negatively impacted by humans (。

WebTable 1). Some, like Siberian marmots (Marmota sibirica), are listed as “Endangered” on the International Union for Conservation of Nature (IUCN) Red List, whereas others, such as the Patagonian mara and European ground squirrel (Spermophilus citellus), are listed as “Near Threatened” and “Vulnerable”, respectively, and are declining at a rate of 30% or more per decade. Despite being nearly eradicated on Australia’s mainland, burrowing bettongs are considered “Near Threatened” because their populations are now stable or improving. Many others are listed in the category of “Least Concern” by the IUCN, but several of those species – such as prairie dogs, bobak marmots (Marmota bobak), southern hairy-nosed wombats, and vizcachas – have also experienced dramatic population declines across much of their former ranges. Some species considered of “Least Concern”, such as ice rats and degus, are thought to be common and have stable populations, but others, including several species of pikas, are in decline because of extensive, ongoing poisoning campaigns. For many other species of burrowing mammals, population trends are poorly known. Below, we describe the primary threats impacting burrowing mammals around the world.

穴居哺乳动物的保护状况在不同的物种中差异很大，但大多数物种的种群都受到了人类的严重负面影响（网络表1）一些物种，如西伯利亚旱獭（Marmota sibirica），在国际自然保护联盟（IUCN）红色名录中被列为 "濒危"，而其他物种，如巴塔哥尼亚豚鼠和欧洲地松鼠（Spermophilus citellus），分别被列为 "近危 "和 "易危"，并以每十年30%以上的速度在下降。

尽管在澳大利亚大陆几乎被根除，但钻地鼠被认为是 "近危 "的，因为它们的种群现在已经稳定或正在改善其他许多物种被世界自然保护联盟列为 "最不关注 "类别，但其中一些物种--如草原土拨鼠、草原旱獭（Marmota bobak）。

、南澳毛吻袋熊和兔鼠--在它们以前的大部分地区也经历了急剧的数量下降一些被认为是 "最低关注 "的物种，如冰鼠和八齿鼠，被认为是常见的，而且种群数量稳定，但其他物种，包括几种鼠兔，由于广泛的、持续的毒害活动而正在下降。

对于许多其他种类的穴居哺乳动物，其种群趋势也不甚明了下面，我们描述了影响世界各地穴居哺乳动物的主要威胁Habitat loss栖息地丧失Grasslands cover 41% of the world’s land surface and are the primary environments used for growing crops and grazing livestock (White et al. 2000). Widespread conversion of grassland to cropland and extensive overgrazing and desertification have resulted in 20–80% declines in grassland area across all continents (White et al. 2000), leading to substantial reductions in the amount of suitable habitat for burrowing mammals and consequent population declines in many species (eg Hoogland 2006; Noble et al. 2007b; Ceballos et al. 2010).。

草地占世界陆地表面的41%，是用于种植作物和放牧的主要环境（White等人，2000）草地广泛转化为耕地和广泛的过度放牧和沙漠化，导致各大洲的草地面积减少20-80%(White等人, 2000)，导致适合穴居哺乳动物的栖息地数量大幅减少，许多物种的数量也随之减少

(如Hoogland 2006; Noble等人, 2007b; Ceballos等人, 2010)Conflicts with livestock与牲畜的矛盾Heavy reliance on grasslands for livestock production has resulted in major conflicts between the livestock industry and burrowing mammals (Miller et al. 2007; Delibes-Mateos et al. 2011). A common threat facing burrowing mammals in many parts of the world is widespread persecution because of their perceived competition with livestock; for instance, the extensive poisoning and shooting of prairie dogs in North America during the past century is largely responsible for the 98% decline in their populations (Hoogland 2006). Poisoning is also prevalent throughout Asia (eg pikas and Brandt’s voles), South America (eg vizcachas), and Africa (eg ice rats) (Branch et al. 2002; Bagchi et al. 2006; Mokotjomela et al. 2009). Governments spend millions of dollars each year “controlling” burrowing mammal populations to benefit the livestock industry, despite research repeatedly demonstrating that such campaigns are not cost effective and result in the indiscriminant poisoning of other types of wildlife (Hoogland 2006; Wesche et al. 2007; Delibes-Mateos et al. 2011). In fact, burrowing mammals, particularly at low to moderate densities, can have beneficial effects by increasing forage quality and productivity (Whicker and Detling 1988; Smith and Foggin 1999; Noble et al. 2007b). Livestock weight gains are minimally affected when burrowing mammal colonies cover less than 30% of the landscape (Derner et al. 2006), and livestock mass has been shown to decline over the same period that burrowing mammals are poisoned (Smith and Foggin 1999). Still, burrowing mammals are often blamed for grassland degradation, despite having co-existed with free-ranging, native megaherbivores for millions of years. Many grasslands are simply overstocked with livestock (Smith and Foggin 1999; Bagchi et al. 2006; Mokotjomela et al. 2009); for example, Ceballos et al. (2010) described a grassland in Mexico that could sustainably support 200 head of cattle but was being grazed by。

2000 cattle. Concomitant with this excessive livestock grazing was the collapse of one of the largest remaining black-tailed prairie dog (Cynomys ludovicianus) colonies (Ceballos et al. 2010). However, where they co-exist, the effects of livestock on grasslands are not independent of burrowing mammals. Indeed, relationships between livestock and burrowing mammals can be mutualistic, and their combined effects on the grassland ecosystem can be synergistic (Davidson et al. 2010). Burrowing mammals also tend to have more intensive localized effects due to their sedentary, colonial behavior, whereas megaherbivores have more diffuse impacts across larger spatial scales (Davidson et al. 2010; Yoshihara et al. 2010b).

畜牧业对草原的严重依赖导致了畜牧业与穴居哺乳动物之间的重大矛盾（Miller等人，2007；Delibes-Mateos等人，2011）世界上许多地方的穴居哺乳动物面临的一个共同威胁是广泛的迫害，因为它们被认为是与牲畜竞争；例如，在过去的一个世纪里，北美的草原土拨鼠被广泛地毒杀和射杀，这在很大程度上导致了它们的数量下降98%（Hoogland 2006）。

在亚洲（如鼠兔和布氏田鼠）、南美洲（如兔鼠）和非洲（如冰鼠）也普遍存在中毒现象（Branch等人，2002；Bagchi等人，2006；Mokotjomela等人，2009）政府每年花费数百万美元 "控制 "穴居哺乳动物的数量，以利于畜牧业，尽管研究一再表明，这种活动不符合成本效益，并导致其他类型的野生动物被滥杀（Hoogland 2006；Wesche等人，2007；Delibes-Mateos等人，2011）。

事实上，穴居哺乳动物，特别是在低到中等密度的情况下，可以通过增加饲料质量和生产力而产生有益的影响(Whicker and Detling 1988; Smith and Foggin 1999; Noble et al. 2007b)。

当穴居哺乳动物群落覆盖不到景观的30%时，牲畜的体重增加受到最小的影响(Derner等人, 2006)，而且已经证明在穴居哺乳动物被毒害的同一时期，牲畜质量会下降(Smith and Foggin 1999)。

然而，尽管穴居哺乳动物与自由放养的本地大型食草动物共存了数百万年，但仍经常被指责为草原退化的罪魁祸首许多草原仅仅是过度饲养牲畜（Smith和Foggin 1999；Bagchi等人，2006；Mokotjomela等人，2009）；例如，。

塞巴洛斯等人(2010)描述了墨西哥的一个草原，该草原可以持续养活200头牛，但正在被放牧2000头牛伴随这种过度放牧的是现存最大的黑尾土拨鼠(Cynomys ludovicianus)群落之一的崩溃(Ceballos et al. 2010)。

然而，在它们共存的地方，牲畜对草原的影响并不是独立于穴居哺乳动物的事实上，牲畜和穴居哺乳动物之间的关系可能是相互的，它们对草原生态系统的综合影响可能是协同的(Davidson等人, 2010)穴居哺乳动物。

由于其定居的殖民行为，往往有更密集的局部影响，而大型食草动物在更大的空间范围内有更分散的影响（Davidson等人，2010；Yoshihara等人，2010b）Overexploitation过度开发。

Many burrowing mammal species are overhunted, often for their pelts. Siberian marmots of the Mongolian steppe, for example, once numbered around 40 million individuals; overhunting reduced their populations to 20 million by 1990, and fewer than 5 million remained by 2002 – a 70% decline in little over a decade (Wingard and Zahler 2006; IUCN 2011). Plains vizcachas, Patagonian maras, European ground squirrels, long-tailed ground squirrels, gray marmots (Marmota baibacina), Siberian marmots, bobak marmots, Himalayan marmots (Marmota himalayana), and Arctic ground squirrels, among others, are similarly threatened by overexploitation (

WebTable 1).许多穴居哺乳动物物种被过度猎杀，往往是为了它们的皮毛例如，蒙古草原上的西伯利亚旱獭，曾经有大约4000万只；到1990年，过度捕猎使它们的数量减少到2000万只，到2002年，只剩下不到500万只--在短短十年多的时间里，减少了70%（Wingard和Zahler 2006；IUCN 2011）。

平原兔鼠、巴塔哥尼亚豚鼠、欧洲地松鼠、长尾地松鼠、灰旱獭（Marmota baibacina）、西伯利亚旱獭、草原旱獭、喜马拉雅旱獭（Marmota himalayana）和北极地松鼠等，同样受到过度开发的威胁（网络表1）。

Introduced species and disease引进的物种和疾病Exotic species also pose major threats to many burrowing mammals. Plague, a disease caused by the bacterium Yersinia pestis, is native to rodents in Asia but has been introduced to Africa and the Americas (Gage and Kosoy 2005). Since its introduction to these parts of the world, this exotic pathogen has been devastating populations of some species, such as prairie dogs in North America, that lack immunity (Gage and Kosoy 2005, but see Rocke et al. 2012). Even in its native range, plague can cause massive, episodic die-offs in Siberian marmots, exacerbating declines of this already endangered species (Wingard and Zahler 2006). Sarcoptic mange (Sarcoptes scabiei) is an emerging disease threatening the highly fragmented populations of southern hairy-nosed wombats in Australia (IUCN 2011). Overgrazing by introduced European rabbits (Oryctolagus cuniculus) and domestic livestock, as well as predation by introduced predators, are also important threats to Australia’s bettongs and wombats (Noble et al. 2007b; IUCN 2011).

外来物种也对许多穴居哺乳动物构成主要威胁鼠疫是一种由鼠疫耶尔森氏杆菌（Yersinia pestis）细菌引起的疾病，原产于亚洲的啮齿动物，但已被引入非洲和美洲(Gage and Kosoy 2005)。

自从引入世界这些地区，这种外来的病原体一直在破坏一些缺乏免疫力的物种的种群，如北美的草原土拨鼠Gage and Kosoy 2005, but see Rocke et al. 2012)即使在它的原生地，鼠疫可以造成西伯利亚旱獭的大规模、偶发的死亡，加剧了这个已经濒临灭绝的物种的衰退(Wingard and Zahler 2006)。

疥螨病 (Sarcoptes scabiei)是一种新出现的疾病，威胁着澳大利亚南澳毛吻袋熊高度分散的种群(IUCN 2011)引进的欧洲穴兔(Oryctolagus cuniculus)和家畜的过度放牧，以及引进的捕食者的捕食。

，也是澳大利亚地鼠和袋熊的重要威胁(Noble et al. 2007b; IUCN 2011)Climate change气候变化The impacts of climate change on burrowing mammals are only beginning to be detected. For instance, prairie dog populations have declined dramatically in parts of the southern distribution of their North American range after periods of drought, which are projected to increase considerably in this region over the coming decades (Ceballos et al. 2010). Plague epizootics correlate positively with precipitation (Gage and Kosoy 2005), so a changing climate may lead to shifts in areas impacted by plague. Recently, plague has expanded eastward in the US, possibly due to a changing climate, occurring recently and for the first time in the Conata Basin of South Dakota, where the largest remaining complex of black-tailed prairie dog colonies in the US occurs. This has caused large die-offs in prairie dogs and endangered black-footed ferrets (Mustela nigripes) (USFWS 2009).。

气候变化对穴居哺乳动物的影响才开始被发现例如，草原土拨鼠在其北美分布区南部的部分地区，在干旱期后，种群数量急剧下降，预计在未来几十年，该地区的种群数量将大大增加（Ceballos等人，2010）鼠疫发作与降水呈正相关。

(Gage and Kosoy 2005)，因此不断变化的气候可能会导致受鼠疫影响的地区发生变化最近，可能由于气候的变化，鼠疫在美国向东扩展，最近首次发生在南达科他州的科纳塔盆地，那里有美国最大的剩余黑尾土拨鼠群落。

这导致了草原土拨鼠和濒危的黑足鼬(Mustela nigripes)的大量死亡(USFWS 2009)Ecological consequences of burrowing mammal declines

穴居哺乳动物减少的生态后果Consistent with the loss of keystone species (Power et al. 1996), the impacts of burrowing mammal declines can cascade throughout ecosystems (

WebTable 1). Not only can their loss facilitate woody plant invasion (Weltzin etal. 1997; Noble et al. 2007a; Ceballos et al. 2010), but animals that rely on their colonies for nesting habitat are also negatively affected, such as mountain plovers that have declined with the loss of prairie dogs, burrowing owls that have declined with losses of both prairie dogs and vizcachas, and ground-nesting birds that have

declined following pika poisoning (Branch et al. 2002; Lai and Smith 2003; Kotliar et al. 2006). Predators dependent on burrowing mammals for prey have also shown dramatic declines. Black-footed ferrets, for example, rely on prairie dogs for about 90% of their diet; largely because of the extensive decline in prairie dogs, black-footed ferrets have become one of the most endangered mammals in North America (Kotliar et al. 2006). Interestingly, the US Fish and Wildlife Service’s multi-million-dollar breeding program to recover black-footed ferret populations is running out of suitable reintroduction habitat because ferrets require extensive prairie dog colony complexes to support them, but such complexes are now extremely rare as a result of habitat fragmentation, introduced plague, and government-funded extermination programs. Similarly, European ground squirrel populations have experienced major declines, which may be causing an associated decline in one of their predators, the steppe polecat (Mustela eversmanii) (IUCN 2011). Ferruginous hawks (Buteo regalis), which are highly reliant on Gunnison’s prairie dogs (Cynomys gunnisoni) for prey during their winter migration, are now threatened, mainly because of the fall in prairie dog populations (Cartron et al. 2004). Comparably, where plateau pikas have been poisoned in the Tibetan Plateau, bird species abundance and richness have declined considerably, reflecting declines in species that nest in pika burrows or prey on pikas. In fact, many predators that rely on plateau pikas as key prey – including upland buzzards (Buteo hemilasius), saker falcons (Falco cherrug), brown bears, Pallas’s cats, Tibetan foxes (Vulpes ferrilata), and steppe polecats – have nearly disappeared from areas where pikas have been poisoned (Lai and Smith 2003; Delibes-Mateos et al. 2011).

与关键物种的损失一致(Power等人, 1996)，穴居哺乳动物减少的影响可以在整个生态系统中层层递减(WebTable 1)不仅它们的损失会促进木本植物的入侵(Weltzin etal. 1997; Noble et al. 2007a; Ceballos et al. 2010)，。

而且依赖它们群落筑巢的动物也会受到负面影响，例如岩鸻因草原土拨鼠的损失而减少，穴居猫头鹰因草原土拨鼠和兔鼠的损失而减少，以及地面巢的鸟类因鼠兔中毒而减少（Branch等人，2002；Lai和Smith 2003；Kotliar等人，2006）。

依赖于穴居哺乳动物作为猎物的捕食者也显示出急剧的下降例如，黑足鼬，大约90%的食物依赖草原土拨鼠；主要是因为草原土拨鼠的广泛减少，黑足鼬已经成为北美最濒危的哺乳动物之一(Kotliar等人, 2006)。

有趣的是，美国鱼类和野生动物管理局的数百万美元的繁殖计划，以恢复黑足鼬的种群，正在耗尽合适的重新引入的栖息地，因为雪貂需要广泛的草原土拨鼠群落来支持它们，但由于栖息地的破碎化，引入的瘟疫和政府资助的灭绝计划，这种群落现在极为罕见。

同样地，欧洲地松鼠种群经历了严重的下降，这可能会导致它们的天敌之一，艾鼬(Mustela eversmanii)相应的下降(IUCN 2011)王鵟(Buteo regalis)在冬季迁徙时高度依赖古氏土拨鼠(Cynomys gunnisoni)作为猎物，现在受到威胁，主要是因为草原土拨鼠数量的下降(Cartron等人, 2004)。

相比较而言，在青藏高原高原鼠兔被毒死的地方，鸟类物种丰度和丰富度都大大下降，反映了在鼠兔洞穴中筑巢或捕食鼠兔的物种的下降事实上，许多依靠高原鼠兔作为主要猎物的捕食者--包括大鵟（Buteo hemilasius）、猎隼（Falco cherrug）、棕熊、兔、藏狐（Vulpes ferrilata）和艾鼬--在鼠兔被毒死的地区几乎消失了（Lai和Smith 2003；Delibes-Mateos等人 2011）。

Research, policy, and management implications 研究、政策和管理方面的影响Burrowing mammals play important functional roles in grasslands and a variety of other ecosystems around the world (Kelt 2011). However, their requirements and impacts vary by species, which have been shaped by their evolutionary histories and extrinsic environmental conditions, such as climate, soils, vegetation, and associated animal species. The distinctive impacts that each burrowing mammal species has play out across broad landscapes, where they facilitate associated

species and perform key ecosystem functions. However, because many burrowing mammal populations have undergone severe numerical reductions, their key ecological roles have been greatly diminished throughout much of their geographic range. To support the ecosystems associated with burrowing mammals, we recommend that conservation and management efforts include maintaining or re-establishing these populations and their functional roles at the landscape scale.

穴居哺乳动物在草原和世界各地的各种其他生态系统中发挥着重要的功能作用(Kelt 2011)然而，它们的需求和影响因物种而异，这取决于它们的进化史和外部环境条件，如气候、土壤、植被和相关的动物物种每个穴居哺乳动物物种的独特影响在广阔的景观中发挥着作用，它们促进了相关物种的生长，并发挥了关键的生态系统功能。

然而，由于许多穴居哺乳动物种群的数量急剧减少，它们在其大部分地理范围内的关键生态作用已大大减弱为了支持与穴居哺乳动物相关的生态系统，我们建议保护和管理工作包括维持或重建这些种群及其在景观尺度上的功能作用。

Research needs研究需求Because most species of burrowing mammals remain poorly studied, considerable additional research is necessary to understand their impacts, conserve their populations, and preserve or restore their ecological roles. Among the most critical research needs are to:

由于对大多数穴居哺乳动物物种的研究仍然不足，因此需要大量的额外研究来了解它们的影响，保护它们的种群，并保护或恢复它们的生态作用其中最关键的研究需求是：(1) Quantify and understand their roles in maintaining biodiversity, and identify those species with which they are strongly associated. We also need to better understand their population trends and conservation needs, because declines in burrowing mammals are likely to have cascading effects throughout the ecosystems in which they occur.。

（1）量化和了解它们在维持生物多样性方面的作用，并确定那些与它们有密切联系的物种我们还需要更好地了解它们的种群趋势和保护需求，因为穴居哺乳动物的减少可能会对它们所在的整个生态系统产生连带影响(2) Determine under what conditions their interactions with associated species can be positive, neutral, or negative, and how this translates into overall grassland health. This includes quantifying the ecosystem services they provide and their potential economic value.。

（2）确定在什么条件下它们与相关物种的相互作用可以是积极的、中性的或消极的，以及这如何转化为整体草原健康这包括量化它们提供的生态系统服务及其潜在的经济价值(3) Study the relationships between burrowing mammals and livestock to determine how they can co-exist, from ecological, economic, and social perspectives.。

（3）研究穴居哺乳动物和牲畜之间的关系，从生态、经济和社会的角度确定它们如何共存(4) Determine sustainable harvest rates for those species threatened by overhunting.。

（4）确定那些受到过度狩猎威胁的物种的可持续捕获率(5) Develop the capability to predict and help prevent disease outbreaks, such as plague and mange, possibly through the use of vaccines like those being developed to protect critical populations of North American prairie dogs from plague.。

（5）发展预测和帮助预防疾病爆发的能力，如鼠疫和疥疮，可能通过使用正在开发的疫苗来保护北美草原土拨鼠的关键种群免受鼠疫的影响Policy and management implications 政策和管理影响。

As the demand for food production continues to grow over the next decades, conflicts between burrowing mammals and people are bound to increase. Although we have cited evidence that these animals have positive impacts on their environment, and that eradication programs are typically costly and ineffective, burrowing mammals remain highly misunderstood and heavily persecuted. The traditional assumption that they compete and have other net negative interactions with livestock needs to be re-evaluated. Results of recent research should be used to educate managers and the public on their diverse ecological roles and positive impacts. Such efforts could help change public attitudes, correct misconceptions, and reverse government policies that continue to fuel eradication programs (Smith and Foggin 1999; Miller et al. 2007; Delibes-Mateos et al. 2011). In areas where their populations must be controlled, managers may be able to reduce burrowing mammal populations by simply reducing livestock grazing and allowing grass to grow tall (Smith et al. 2006; Davidson et al. 2010). Overall, however, grassland management must be more holistic, man- aging not only for livestock production but also for preserving burrowing mammal populations that are essential for maintaining healthy grasslands over the long term.

随着未来几十年对食物生产的需求继续增长，穴居哺乳动物和人之间的矛盾必然会增加尽管我们已经列举了这些动物对其环境有积极影响的证据，而且根除计划通常是昂贵而无效的，但穴居哺乳动物仍然被高度误解，并受到严重迫害。

需要重新评估传统的假设，即它们与牲畜竞争和有其他净负面的互动最近的研究结果应被用来教育管理人员和公众，使他们了解它们的不同生态作用和积极影响这种努力可以帮助改变公众的态度，纠正错误的观念，并扭转政府继续助长根除计划的政策。

(Smith and Foggin 1999; Miller et al. 2007; Delibes-Mateos et al. 2011)在必须控制其种群的地区，管理者可能能够通过简单地减少牲畜放牧和让草长得更高来减少穴居哺乳动物的数量(Smith等人, 2006; Davidson等人, 2010)。

然而，总的来说，草原管理必须更加全面，不仅要为畜牧业生产进行管理，还要保护穴居哺乳动物种群，这对长期保持健康的草原是至关重要的Finally, conservation plans are needed to reverse the declines in burrowing mammal populations and associated species. Reintroductions are currently an important component of such conservation efforts but are expensive, intensive, and small in scale. Consequently, managers should focus on maintaining and increasing existing populations wherever possible, including the creation of protected areas, engagement of local communities, and provision of economic incentives whereby landowners receive financial compensation for supporting burrowing mammals and the ecosystem services they provide (Hoogland 2006). Bolstering populations is also important to mitigate against future losses of these species as a result of disease and climate change. Without such actions, there is serious concern as to how species threatened by multiple, compounding human impacts today will be able to withstand a rapidly changing environment.。

最后，需要制定保护计划来扭转穴居哺乳动物种群和相关物种的下降重新引种是目前这种保护工作的一个重要组成部分，但成本高，强度大，规模小因此，管理者应尽可能地集中精力维持和增加现有种群，包括建立保护区，让当地社区参与进来，并提供经济激励措施。

，让土地所有者因支持穴居哺乳动物及其提供的生态系统服务而获得经济补偿（Hoogland 2006）增加种群数量也很重要，可以缓解这些物种未来因疾病和气候变化而造成的损失如果没有这样的行动，人们会很担心今天受到人类多重影响威胁的物种将如何抵御快速变化的环境。

免责声明：本站所有信息均搜集自互联网，并不代表本站观点，本站不对其真实合法性负责。如有信息侵犯了您的权益，请告知，本站将立刻处理。联系QQ：1640731186