Effects of Artisanal Fishing on Marine Communities in the Galápagos Islands

Abstract: The Galápagos Islands harbor some of the least impacted marine ecosystems in the tropics, but there are indications that local artisanal fishing is affecting exploited marine communities. To quantify these effects, I sampled communities of fishes and sea urchins at a number of heavily fished and lightly fished sites throughout the central islands of the archipelago. Sites were selected based on information collected as part of a local fisheries monitoring study and standardized across a number of abiotic factors. Abundance and biomass of the primary target species were significantly lower in the heavily fished sites than in the lightly fished sites. Community structure also differed between heavily and lightly fished sites. Cluster analyses of the full community of fishes and a subset of nontarget fishes revealed that sites within a treatment were more similar to one another than sites between treatments. Herbivorous fishes tended to be lower in abundance and sea urchins tended to be higher in abundance in heavily fished sites, but these differences were not significant. My results are encouraging in that the direct effects of artisanal fishing are limited to the primary target species, which probably results from a high specificity of fishing gear. The differences in community structure, however, suggest that artisanal fishing also has cascading effects on noncommercial species throughout the community. An improved understanding of important ecological interactions, increased ecological and fishery monitoring, and effective precautionary management are needed to ensure that human effects in these waters remain minimal.     

Heavy metals in the Sea-Skater Halobates robustus from the Galápagos Islands: Concentrations in nature and uptake experiments,with special reference to cadmium

Samples of Halobates robustus Barber (Heteroptera: Gerridae) from the Galápagos Islands were analysed by optical emission spectrometry. The levels (in g g^-1 dry weight) of Zn (134), Cu (155), Pb (< 1), Cd (7), and Cr (3) were not significantly different among insects of different sexes or developmental stages. The low natural levels of Cd in H. robustus from the relatively unpolluted environment of the Galápagos Islands are compared to the high concentrations of Cd in Halobates spp. from relatively polluted regions. Since the measured levels of Cd in their natural zooplankton food rarely exceed 10 g g^-1, and very little of the Cd is found in the soft tissues, the high Cd concentrations (100 to 200 g g^–1) in some seaskater species have evidently been derived by drinking from the surface microlayer of the seawater.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and Galápagos Islands (Ecuador). II. Poritidae

A comparative study of the reproductive ecology of the zooxanthellate, scleractinian corals Porites lobata Dana and P. panamensis Verrill was conducted from 1985 to 1991 in eastern Pacific reef environments that were severly impacted by the 1982–1983 El Niño warming events. P. lobata, a presumed broadcast spawner of large colony size, is widely distributed in the equatorial eastern Pacific, whereas P. panamensis, a brooder of small colony size, is abundant only on some reefs in Panamá. Both species were gonochoric with nearly 1:1 sex ratios in large study populations except for P. lobata at Caño Island that had 14% hermaphroditic colonies. Mature, unfertilized oocytes contained numerous zooxanthellae in both Porites species, and all planula developmental stages contained zooxanthellae in P. panamensis. Year-round sampling revealed high proportions of colonies with gonads, ranging from 30 to 68% in P. lobata and from 60 to 68% in P. panamensis. No clear relationship between numbers of reproductive colonies and the thermal stability of the habitat was evident in P. lobata: percent colonies with gonads at non-upwelling sites was 48 to 68% at Caño Island (Costa Rica) and Uva Island (Panamá), and at upwelling sites 30 to 50% at Saboga Island and Taboga Island (Panamá), and the Galápagos Islands (Ecuador). Similarly, 90% of all P. panamensis colonies were reproductive at Uva Island (a non-upwelling site), and 86% were reproductive at Taboga Island (an upwelling site). Upwelling at Taboga Island is seasonal, nevertheless P. panamensis produced mature gonads or planulae over most of the year (11 mo), whereas P. lobata exhibited reproductive activity during only 2 mo (May and June). No clear lunar periodicity was observed in P. panamensis (Taboga Island), but a high proportion of P. lobata showed increased gonadal development around full and new moon, especially at Caño and Uva Islands. Estimated fecundities were relatively high for P. lobata at Caño (4000 eggs cm^-2 yr^-1) and Uva (5200 eggs cm^-2 yr^-1) Islands, and notably low (70 to 110 eggs cm^-2 yr^-1) in the Galápagos Islands. P. panamensis mean fecundity at Taboga Island was 720 planulae cm^-2 yr^-1 or 4.0 mm³ cm^-2 yr^-1, which was lower than the egg volume production of P. lobata at Caño and Uva Islands (7.0 to 10.0 mm³ cm^-2 yr^-1). The capacity of P. lobata and P. panamensis to reproduce sexually supports the notion that eastern Pacific coral reef recovery may not be dependent on long-distance dispersal from central Pacific areas. However, sexual recruits of P. lobata are absent or uncommon at all eastern Pacific study sites while recruits of P. panamensis were common to abundant only at the Uva Island study site. Asexual fragmentation in P. lobata augments recruitment locally, but plays no role in P. panamensis recruitment.     

Predominance of clade D <Emphasis Type="Italic">Symbiodinium</Emphasis> in shallow-water reef-building corals off Kish and Larak Islands (Persian Gulf,Iran)

Scleractinian coral species harbour communities of photosynthetic taxa of the genus Symbiodinium. As many as eight genetic clades (A, B, C, D, E, F, G and H) of Symbiodinium have been discovered using molecular biology. These clades may differ from each other in their physiology, and thus influence the ecological distribution and resilience of their host corals to environmental stresses. Corals of the Persian Gulf are normally subject to extreme environmental conditions including high salinity and seasonal variation in temperature. This study is the first to use molecular techniques to identify the Symbiodinium of the Iranian coral reefs to the level of phylogenetic clades. Samples of eight coral species were collected at two different depths from the eastern part of Kish Island in the northern Persian Gulf, and Larak Island in the Strait of Hormuz. Partial 28S nuclear ribosomal (nr) DNA of Symbiodinium (D1/D2 domains) were amplified by polymerase chain reaction (PCR). PCR products were analyzed using single stranded conformational polymorphism and phylogenetic analyses of the LSU DNA sequences from a subset of the samples. The results showed that Symbiodinium populations were generally uniform among and within the populations of eight coral species studied, and there are at least two clades of Symbiodinium from Kish and Larak islands. Clade D was detected from eight of the coral species while clade C was found in two of species only (one species hosted two clades simultaneously). The dominance of clade D might be explained by high temperatures or the extreme temperature variation, typical of the Persian Gulf. Publication of this article was held up owing to technical problems. The publisher apologizes sincerely for this lengthy delay.     

Male reproductive biology of the red frog crab,Ranina ranina,off Hachijojima,Izu Islands,Japan

Male red frog crabs, Ranina ranina, were collected year round in 1990 and 1991 off Hachijojima for histological study of the reproductive system and cycle. The testis containing the lobules and seminiferous ducts is surrounded by connective tissue. The seminiferous duct connects to the anterior end of the vas deferens, which can be histologically divided into three portions similar to one another in appearance. It was surrounded by fibrous connective tissue, muscle fibrils and columnar epithelium. Muscle fibrils were absent in the anterior portion. Multiple sperm masses were not formed in the vas deferens and ejaculatory duct, but the sperm mass was covered with a capsule composed of two layers. The outer layer of the capsule was periodic acid Schiff (PAS)-positive, but the inner layer was negative. Both layers were Alcian Blue negative, except the vacuoles in the outer layer that were stained blue. The small round androgenic gland was attached to the posterior end of the vas deferens of the coxa of the eighth thoracic appendage. The ejaculatory duct was distinguishable from the vas deferens by the absence of columnar epithelium and the presence of thick longitudinal muscle fibers. Spermatogenesis was histochemically examined. The acrosomal vesicle appeared to be derived from PAS-positive vesicles in the cytoplasm of the spermatid at the early stage of spermiogenesis. The arms were positive to the Feulgen reaction and the subacrosomal region was negative to PAS. Seasonal changes in reproductive cycle were inconspicuous histologically and microscopically. Sperm were always present in the testis and vas deferens throughout the year and occupied 5.1 to 19.6% of testis observed in cross sections. The minimum size of maturity is less than 39 mm carapace length, but the minimum size capable of successful mating was estimated to be ca. 55 mm.     

Long‐Range Movement of Humpback Whales and Their Overlap with Anthropogenic Activity in the South Atlantic Ocean

Humpback whales (Megaptera novaeangliae) are managed by the International Whaling Commission as 7 primary populations that breed in the tropics and migrate to 6 feeding areas around the Antarctic. There is little information on individual movements within breeding areas or migratory connections to feeding grounds. We sought to better understand humpback whale habitat use and movements at breeding areas off West Africa, and during the annual migration to Antarctic feeding areas. We also assessed potential overlap between whale habitat and anthropogenic activities. We used Argos satellite‐monitored radio tags to collect data on 13 animals off Gabon, a primary humpback whale breeding area. We quantified habitat use for 3 cohorts of whales and used a state‐space model to determine transitions in the movement behavior of individuals. We developed a spatial metric of overlap between whale habitat and models of cumulative human activities, including oil platforms, toxicants, and shipping. We detected strong heterogeneity in movement behavior over time that is consistent with previous genetic evidence of multiple populations in the region. Breeding areas for humpback whales in the eastern Atlantic were extensive and extended north of Gabon late in the breeding season. We also observed, for the first time, direct migration between West Africa and sub‐Antarctic feeding areas. Potential overlap of whale habitat with human activities was the highest in exclusive economic zones close to shore, particularly in areas used by both individual whales and the hydrocarbon industry. Whales potentially overlapped with different activities during each stage of their migration, which makes it difficult to implement mitigation measures over their entire range. Our results and existing population‐level data may inform delimitation of populations and actions to mitigate potential threats to whales as part of local, regional, and international management of highly migratory marine species. Cuantificación de los Movimientos de Gran Amplitud y el Traslape Potencial con Actividad Antropogénica y las Ballenas Jorobadas en el Océano Atlántico Sur     

Female reproductive biology and oocyte development of the red frog crab,Ranina ranina,off Hachijojima,Izu Islands,Japan

Speciments of the red frog crab, Ranina ranina, examined in this study were collected off Hachijojima, Japan from 1987–1988 and 1990–1991. Ovigerous females occurred from May to September, and their carapace length ranged from 61.8 to 102.1 mm. Minimum size at the maturity was estimated to be 50 to 60 mm carapace length. The lobule-type ovary had a longitudinal ovarian cavity at the center. Germinal zones were located along the sides of the cavity. Oogenesis was divided into five main and ten detailed stages: proliferation (oogonium); previtellogenesis (bouquet, early chromatin nucleolus, late chromatin nucleolus); primary vitellogenesis (oil globule and yolk granule); secondary vitellogenesis (primary yolk platelet, secondary yolk platelet and prematuration); and maturation (maturation) stages. Vitellogenesis started at the oil globule stage when weakly positive periodic acid-Schiff granules appear in the cytoplasm. Based on the gonadosomatic index and histological changes in ovary, the reproductive cycle was divided into five stages: multiplication, from December to January; developing, from February to March; ripe in April and early May; spawning, from late May to August; and recovery, from August to November. Each crab spawned more than once during the spawning season.     

Citation Rate and Perceived Subject Bias in the Amphibian‐Decline Literature

Abstract: As a result of global declines in amphibian populations, interest in the conservation of amphibians has grown. This growth has been fueled partially by the recent discovery of other potential causes of declines, including chytridiomycosis (the amphibian chytrid, an infectious disease) and climate change. It has been proposed that researchers have shifted their focus to these novel stressors and that other threats to amphibians, such as habitat loss, are not being studied in proportion to their potential effects. We tested the validity of this proposal by reviewing the literature on amphibian declines, categorizing the primary topic of articles within this literature (e.g., habitat loss or UV‐B radiation) and comparing citation rates among articles on these topics and impact factors of journals in which the articles were published. From 1990 to 2009, the proportion of papers on habitat loss remained fairly constant, and although the number of papers on chytridiomycosis increased after the disease was described in 1998, the number of published papers on amphibian declines also increased. Nevertheless, papers on chytridiomycosis were more highly cited than papers not on chytridiomycosis and were published in journals with higher impact factors on average, which may indicate this research topic is more popular in the literature. Our results were not consistent with a shift in the research agenda on amphibians. We believe the perception of such a shift has been supported by the higher citation rates of papers on chytridiomycosis.     

Phylogeography of California and Galápagos sea lions and population structure within the California sea lion

We investigate the phylogeography of California (Zalophus californianus) and Galápagos (Z. wollebaeki) sea lions and describe within-population structure for the California sea lion based on mitochondrial DNA. Fifty control-region haplotypes were found, 41 from Z. californianus and 9 from Z. wollebaeki, with three fixed differences between the two species. Ranked population boundaries along the range of Z. californianus were defined based on the Monmonier Maximum Difference Algorithm, resulting in five genetically distinct populations, two in the Pacific Ocean and three inside the Gulf of California. A Minimum Spanning Network showed a strong phylogeographic signal with two well-defined clusters, Z. californianus and Z. wollebaeki, separated by six base-pair differences, supporting the existence of two genetically distinct species with an estimated divergence time of ~0.8 Ma. Results are discussed in the context of the historical geologic and paleoceanographic events of the last 1 Ma in the eastern Pacific.     

Chemical composition of midwater fishes as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

The variation with depth in water, lipid, protein, carbon and nitrogen contents (% wet weight) of 42 species of midwater fishes, collected in November 1976 off the west coast of Oahu in the Hawaiian Archipelago, was measured. The Hawaiian fishes show significant relationships between these components and depth of occurrence. The slopes of these relationships are not significantly different from those reported for midwater fishes from off California, USA. However, the fishes from Hawaii have significantly lower lipid levels and higher protein levels than do the species from off California. The deep-living Hawaiian species (500 m and deeper) have significantly lower lipid (% wet weight), but there is no significant difference in protein (% wet weight). The difference in lipid contents at all depths appears to be an evolved characteristic, with the greater lipid levels off California being selected for by greater spatial and temporal variation in the food supply for these fishes off the California coast than off Hawaii. The higher protein contents in the shallow-living Hawaiian fishes appear to reflect greater muscle power selected for in these fishes by the greater water clarity, and therefore greater reactive distances, in the surface layers off Hawaii. These conclusions support the general hypothesis that the lower protein contents of bathypelagic fishes are not directly selected by food limitation at depth, but rather result from the relaxation of selection for rapid-swimming abilities at greater depths due to the great reduction at greater depths in the distance over which visual predator-prey interactions can take place. The lower lipid levels in the deeper-living species are apparently made possible by the reduced metabolic rates of these species which reduces their need for energy stores.     

Metabolic rates of midwater crustaceans as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

During July of 1983, 1986, and 1987, we measured rates of oxygen consumption of 234 individuals of 17 species of midwater crustaceans (orders Decapoda, Mysidacea, and Euphausiacea) off the Hawaiian islands at depths from the surface to greater than 1200 m. The routine metabolic rates declined with increasing depths of the species' occurrence to an extent greater than could be accounted for by depth-related changes in body size or water temperature. Most species appeared able to regulate their oxygen consumption down to the lowest oxygen partial pressures found in their depth range (20 mm Hg O₂), but did not regulate to such low oxygen partial pressures as did similar midwater crustaceans off California, where oxygen levels reach as low as 6 mm Hg. Metabolic rates of the shallower-living, but not the deepest-living Hawaiian crustaceans were significantly higher than those of Californian crustaceans. This is interpreted as indicating that the metabolic rates of midwater crustaceans are not adapted specifically to differing levels of primary production and that the decline with depth of metabolic rates in these species is not the result of food limitation at depth. The data are, however, consistent with the hypothesis that lower metabolic rates at depth are due to the relaxation of selection pressures relating to visual predation near the surface.     

Gray Whales and the Value of Monitoring Data in Implementing the U.S. Endangered Species Act

Abstract: Many scientists lament the absence of data for endangered species and argue that more funds should be spent acquiring basic information about population trends. Using 19 years of abundance estimates for the eastern North Pacific gray whale ( Eschrichtius robustus ), we sampled subsets of the original survey data to identify the number of years of data required to remove the population from the U.S. Endangered Species Act's (ESA) list of endangered and threatened wildlife. For any given duration of monitoring, we selected all possible combinations of consecutive counts. To incorporate variability in growth rates, we extracted a maximum likelihood estimator of growth rate and confidence interval about that growth rate on the assumption that the population changes can be approximated by a simple diffusion process with drift. We then applied a new approach to determine ESA status for each subset of survey data and found that a quantitative decision to delist is unambiguously supported by 11 years of data but is precariously uncertain with fewer than 10 years of data. The data needed to produce an unequivocal decision to delist gray whales cost the National Marine Fisheries Service an estimated U.S. $660,000, a surprisingly modest expense given the fact that delisting can greatly simplify regulatory constraints. This example highlights the value of population monitoring in administering the ESA and provides a compelling example of the utility of such information in identifying both imperiled and recovered species. The economic value of such data is that they provide the foundation for delisting, which could ultimately save much more money than the collection of the data would ever cost.     

Trophic Cascades Following the Disease‐Induced Decline of an Apex Predator,the Tasmanian Devil

As apex predators disappear worldwide, there is escalating evidence of their importance in maintaining the integrity and diversity of the ecosystems they inhabit. The largest extant marsupial carnivore, the Tasmanian devil (Sarcophilus harrisii) is threatened with extinction from a transmissible cancer, devil facial tumor disease (DFTD). The disease, first observed in 1996, has led to apparent population declines in excess of 95% in some areas and has spread to more than 80% of their range. We analyzed a long‐term Tasmania‐wide data set derived from wildlife spotlighting surveys to assess the effects of DFTD‐induced devil decline on populations of other mammals and to examine the relative strength of top–down and bottom–up control of mesopredators between 2 regions with different environmental conditions. Collection of the data began >10 years before DFTD was first observed. A decrease in devil populations was immediate across diseased regions following DFTD arrival, and there has been no indication of population recovery. Feral cats (Felis catus) increased in areas where the disease was present the longest, and feral cat occurrence was significantly and negatively associated with devils. The smallest mesopredator, the eastern quoll (Dasyurus viverrinus), declined rapidly following DFTD arrival. This result suggests the species was indirectly protected by devils through the suppression of larger predators. Rainfall deficiency was also a significant predictor of their decline. Environmental variables determined the relative importance of top–down control in the population regulation of mesopredators. In landscapes of low rainfall and relatively higher proportions of agriculture and human settlement, top–down forces were dampened and bottom–up forces had the most effect on mesopredators. For herbivore prey species, there was evidence of population differences after DFTD arrival, but undetected environmental factors had greater effects. The unique opportunity to assess population changes over extensive temporal and spatial scales following apex predator loss further demonstrated their role in structuring ecosystems and of productivity in determining the strength of top–down control. Cascadas Tróficas Después de la Declinación Inducida por Enfermedad de un Depredador Apical, el Demonio de Tasmania     

A Retrospective Evaluation of the Global Decline of Carnivores and Ungulates

Assessing temporal changes in species extinction risk is necessary for measuring conservation success or failure and for directing conservation resources toward species or regions that would benefit most. Yet, there is no long‐term picture of genuine change that allows one to associate species extinction risk trends with drivers of change or conservation actions. Through a review of 40 years of IUCN‐related literature sources on species conservation status (e.g., action plans, red‐data books), we assigned retrospective red‐list categories to the world's carnivores and ungulates (2 groups with relatively long generation times) to examine how their extinction risk has changed since the 1970s. We then aggregated species’ categories to calculate a global trend in their extinction risk over time. A decline in the conservation status of carnivores and ungulates was underway 40 years ago and has since accelerated. One quarter of all species (n = 498) moved one or more categories closer to extinction globally, while almost half of the species moved closer to extinction in Southeast Asia. The conservation status of some species improved (toward less threatened categories), but for each species that improved in status 8 deteriorated. The status of large‐bodied species, particularly those above 100 kg (including many iconic taxa), deteriorated significantly more than small‐bodied species (below 10 kg). The trends we found are likely related to geopolitical events (such as the collapse of Soviet Union), international regulations (such as CITES), shifting cultural values, and natural resource exploitation (e.g., in Southeast Asia). Retrospective assessments of global species extinction risk reduce the risk of a shifting baseline syndrome, which can affect decisions on the desirable conservation status of species. Such assessments can help conservationists identify which conservation policies and strategies are or are not helping safeguard biodiversity and thus can improve future strategies. Una Evaluación Retrospectiva de la Declinación Global de Carnívoros y Ungulados     

Epidemic Disease and the Catastrophic Decline of Australian Rain Forest Frogs

In the montane rain forests of eastern Australia at least 14 species of endemic, stream-dwelling frogs have disappeared or declined sharply (by more than 90%) during the past 15 years. We review available information on these declines and present eight lines of evidence that collectively suggest that a rapidly spreading, epidemic disease is the most likely responsible agent. The extreme virulence of the putative frog patbogen suggests it is likely exotic to Australian rain forests. We propose that exotic pathogens may be responsible for some recent declines of amphibian populations on other continents and that the intercontinental spread of such pathogens is greatly facilitated by human activities such as the thriving international trade in aquarium fish. Our hypothesis may help explain why some amphibian populations in seemingly pristine environments have mysteriously declined.     

Climate Change,Multiple Stressors,and the Decline of Ectotherms

Climate change is believed to be causing declines of ectothermic vertebrates, but there is little evidence that climatic conditions associated with declines have exceeded critical (i.e., acutely lethal) maxima or minima, and most relevant studies are correlative, anecdotal, or short‐term (hours). We conducted an 11‐week factorial experiment to examine the effects of temperature (22 °C or 27 °C), moisture (wet or dry), and atrazine (an herbicide; 0, 4, 40, 400 μg/L exposure as embryos and larvae) on the survival, growth, behavior, and foraging rates of postmetamorphic streamside salamanders (Ambystoma barbouri), a species of conservation concern. The tested climatic conditions were between the critical maxima and minima of streamside salamanders; thus, this experiment quantified the long‐term effects of climate change within the noncritical range of this species. Despite a suite of behavioral adaptations to warm and dry conditions (e.g., burrowing, refuge use, huddling with conspecifics, and a reduction in activity), streamside salamanders exhibited significant loss of mass and significant mortality in all but the cool and moist conditions, which were closest to the climatic conditions in which they are most active in nature. A temperature of 27 °C represented a greater mortality risk than dry conditions; death occurred rapidly at this temperature and more gradually under cool and dry conditions. Foraging decreased under dry conditions, which suggests there were opportunity costs to water conservation. Exposure to the herbicide atrazine additively decreased water‐conserving behaviors, foraging efficiency, mass, and time to death. Hence, the hypothesis that moderate climate change can cause population declines is even more plausible under scenarios with multiple stressors. These results suggest that climate change within the noncritical range of species and pollution may reduce individual performance by altering metabolic demands, hydration, and foraging effort and may facilitate population declines of amphibians and perhaps other ectothermic vertebrates. Cambio Climático, Estresantes Múltiples y la Declinación de Ectotermos     

Globalization of the Cashmere Market and the Decline of Large Mammals in Central Asia

As drivers of terrestrial ecosystems, humans have replaced large carnivores in most areas, and human influence not only exerts striking ecological pressures on biodiversity at local scales but also has indirect effects in distant corners of the world. We suggest that the multibillion dollar cashmere industry creates economic motivations that link western fashion preferences for cashmere to land use in Central Asia. This penchant for stylish clothing, in turn, encourages herders to increase livestock production which affects persistence of over 6 endangered large mammals in these remote, arid ecosystems. We hypothesized that global trade in cashmere has strong negative effects on native large mammals of deserts and grassland where cashmere‐producing goats are raised. We used time series data, ecological snapshots of the biomass of native and domestic ungulates, and ecologically and behaviorally based fieldwork to test our hypothesis. In Mongolia increases in domestic goat production were associated with a 3‐fold increase in local profits for herders coexisting with endangered saiga (Saiga tatarica).That increasing domestic grazing pressure carries fitness consequences was inferred on the basis of an approximately 4‐fold difference in juvenile recruitment among blue sheep (Pseudois nayaur)in trans‐Himalayan India. Across 7 study areas in Mongolia, India, and China's Tibetan Plateau, native ungulate biomass is now <5% that of domestic species. Such trends suggest ecosystem degradation and decreased capacity for the persistence of native species, including at least 8 Asian endemic species: saiga, chiru (Pantholops hodgsoni), Bactrian camel (Camelus bactrianus), snow leopard(Panthera uncia), khulan(Equus hemionus), kiang (E. kiang), takhi (E. przewalski), and wild yak (Bos mutus). Our results suggest striking yet indirect and unintended actions that link trophic‐level effects to markets induced by the trade for cashmere. Globalización del Mercado de Cachemira y la Declinación de Mamíferos Mayores en Asia Central