Modelling the effect of El Niño on the persistence of small populations: The Galápagos penguin as a case study

Small populations are vulnerable to long-term declines, even where short-term censuses indicate increasing trends in numbers. Census data for the Galápagos penguin (Spheniscus mendiculus) collected between 1970 and 2004 provide evidence that despite year-to-year population increases detected in most of the annual censuses, the strong El Niño events of 1982-83 and 1997-98 were followed by population declines of more than 60% from which the species has yet to recover. Such large declines raise concerns about the future viability of the species because the frequency and severity of El Niño events are predicted to increase. We used the simulation software VORTEX to evaluate the potential effects of El Niño on the risk of extinction of the Galápagos penguin population and its four constituent subpopulations. Weak and strong El Niño events were treated as catastrophes, with varying frequencies, which simulated past, current and future effects on the penguin population. The “Current El Niño” scenario, based on the frequency of El Niño events recorded in the Galápagos between 1965 and 2004, indicated an approximately 30% probability of extinction within the next 100 years for the penguin population. More ominously, the species may be at a greater risk if the frequency of strong El Niño episodes increases only marginally. A probability of extinction greater than 80% was predicted when the current frequency (5%) of strong El Niño events was doubled (to 10%). The probabilities of extinctions were higher for each subpopulation treated individually, ranging from 34% for Isabela and Fernandina, 64% for Bartolomé-Santiago to 78% for the smallest subpopulation on Floreana. Sensitivity analyses identified survival of penguins during El Niño events and sex ratio as influential parameters. The estimates of extinction risk may be conservative as other threats associated with increased human activities on the islands may further compromise species persistence.     

Assessing the risks of introduced chickens and their pathogens to native birds in the Galápagos Archipelago

Poultry production is an important economic activity on inhabited islands of the Galápagos archipelago. There has been a recent surge in both small-scale backyard chickens and larger scale broiler production associated with growth in the human population and the tourist industry. With increased poultry production, concerns have been expressed about the increasing risk of transfer of disease from chickens to native Galápagos bird species that may have little resistance to introduced pathogens [Wikelski, M., Foufopoulos, J., Vargas, H., Snell, H., 2004. Galápagos birds and diseases: invasive pathogens as threats for island species. Ecology and Society 9(5). Available from: URL:http://www.ecologyandsociety.org/vol9/iss1/art5]. This study evaluates risks posed by chicken disease to endemic and native Galápagos bird species, based on empirical evidence of pathogens present in chickens on the islands and a literature review of effects of these pathogens in wild species. Pathogens identified in domestic chicken populations of immediate avian conservation concern are Newcastle disease, Mycoplasma gallisepticum, and the proventricular parasite Dispharynx sp. Newcastle disease (avian paramyxovirus-1) poses an imminent threat to Galápagos penguins (Spheniscus mendiculus), flightless cormorants (Phalacrocorax harrisi), and lava gulls (Larus fuliginosus), species with very small population sizes (less than 1500 animals each). Additionally, litter from broiler farms could affect ecological processes in local ecosystems. Improved poultry biosecurity measures are urgently needed on the Galápagos Islands for avian disease management, yet developing these strategies presents political, social, and economic challenges.     

Evaluating effects of deforestation, hunting, and El Niño events on a threatened lemur

Madagascar ranks as one of the world’s top extinction hotspots because of its high endemism and high rate of habitat degradation. Global climate phenomena such as El Niño Southern Oscillations may have confounding impacts on the island’s threatened biota but these effects are less well known. We performed a demographic study of Propithecus edwardsi, a lemur inhabiting the eastern rainforest of Madagascar, to evaluate the impact of deforestation, hunting, and El Niño on its population and to re-evaluate present endangerment categorization under the IUCN. Over 18 years of demographic data, including survival and fecundity rates were used to parameterize a stochastic population model structured with three stage classes (yearlings, juveniles, and adults). Results demonstrate that hunting and deforestation are the most significant threats to the population. Analysis of several plausible scenarios and combinations of threat revealed that a 50% population decline within three generations was very likely, supporting current IUCN classification. However, the analysis also suggested that changing global cycles may pose further threat. The average fecundity of lemurs was over 65% lower during El Niño years. While not as severe as deforestation or hunting, if El Niño events remain at the current high frequency there may be negative consequences for the population. We suggest that it is most critical for this species continued survival to create more protected areas, not only to thwart hunting and deforestation, but also to give this endangered lemur a better chance to recover from and adapt to altered climate cycles in the future.     

Effective population size in a bottlenecked fur seal population

The census population size (N) is usually the only information available for most threatened species. For evolutionary matters, the effective population size (N_e), not the census number, is a prime concern. Factors such as variation in the sex ratio of breeding individuals, variation of population size in different generations and mating system are important. The South American fur seal, Arctocephalus australis, has been exploited in Peru by humans since ca. 2000 BC and now the original population declined 72%, as a result of low food availability during the severe El Niño in 1997-1998. In this sense A. australis is now classified as in danger of extinction in Peru. We present the first estimate of N_e of the Peruvian population of A. australis that takes into account the effects of mating system and variation in population size caused by the 1997-1998 El Niño. The resulting N_e was 2153 specimens. We believe that the estimated N_e for the Peruvian population is a critical value, because it is significantly lower than the mean minimum viable population for vertebrates (7000 breeding age adults). This estimated N_e is of critical importance because combined with the current El Niño events are reasons of great concern for the survival of the species and should be taken into account in future management plans to ensure the conservation and protection of the species in the Peruvian coast.     

Plasmodium blood parasite found in endangered Galapagos penguins (Spheniscus mendiculus)

This is the first report of a Plasmodium blood parasite found in the Galapagos Archipelago. Phylogenetic analyses place this parasite, recovered from endangered Galapagos penguins (Spheniscus mendiculus), within the genus Plasmodium, and suggest a close relationship to some of the most dangerous lineages of Plasmodium that have been known to cause severe mortality and morbidity in captive penguin populations. Infectious disease is an increasingly important cause of global species extinctions, and extinctions due to avian pox and avian malaria (Plasmodium relictum) have been well documented in Hawaiian avifauna. Plasmodium blood parasites had not been detected in Galapagos birds until now, despite previous microscopic and molecular screening of many of the species, including the Galapagos penguin. While penguin populations now appear healthy, it is unclear whether this parasite will have an obvious impact on their survival and reproduction, particularly during El Niño events, which cause stress due to reduced food availability. It is possible that this parasite arrived with or shortly after the recent arrival of an introduced mosquito, Culex quinquefasciatus, known elsewhere as a competent vector of Plasmodium blood parasites.     

An experimental study on the effects of an introduced parasite in Darwin’s finches

It is widely recognised that the main threats to the biota of the Galápagos Islands stem from newly introduced diseases, plants and animals. Introduced organisms may lead to the rapid extinction of naïve insular populations. Previous research on Galápagos showed that the parasitic larvae of the fly Philornis downsi occurred in virtually all nests of land bird species examined, with high nestling mortality (27% of all broods) in relation to high parasite infestation. In this study, we used an experimental approach to measure the fitness impacts of the fly larvae in two species of Darwin’s finch (Geospizinae). We created parasite-reduced nests by applying a 1% pyrethrin solution to the nest during the early nesting phase, and measured: (1) nestling growth, (2) nestling haemoglobin (Hb) concentration and (3) fledging success. A single insecticide treatment was sufficient to reduce parasite intensity to nearly zero, and revealed reduced mass gain and reduced fledging success in parasitized nests compared to parasite-reduced nests. This is the first experimental study of fitness impacts by Philornis on its host species. The results are discussed in the light of short-term protection measures for nests of rare finch species in the Galápagos archipelago.     

Extinction behind our backs: the possible fate of one of the Darwin’s finch species on Isla Floreana, Galápagos

Without regular monitoring a rare species may slip into extinction unnoticed. We report a possible case from the Galápagos archipelago. The warbler finch (Certhidea fusca) has not been recorded by scientists on Isla Floreana in recent years, and we have been concerned about its status. On a two-day visit to prime habitat in the breeding season of 2004 we used playback of warbler finch song and calls recorded on another island to stimulate an approach of local birds. We failed to find a single warbler finch, whereas we encountered numerous small tree finches (Camarhynchus parvulus), medium tree finches (C. pauper) and yellow warblers (Dendroica petechia). Searches in 1979, 1983, 1997 and 1999 were also unsuccessful. Our continuing failure to find any warbler finches indicates the species must be extremely rare on the island, if not actually extinct. We discuss possible reasons for their demise and draw attention to other species that could be heading in the same direction.     

厄尔尼诺/拉尼娜事件对长江流域气候的影响研究

通过对长江流域近60 a来气象资料和自然灾害资料的搜集和整理,从金沙江/雅砻江水系、岷沱江水系、上游干流区间、嘉陵江水系、乌江水系、汉江水系、洞庭湖水系、鄱阳湖水系和中下游干流区间等9个不同的区域研究了厄尔尼诺/拉尼娜（ElNio—Southern Oscillation,ENSO）事件对长江流域气候的影响。通过对降水量、气温、自然灾害事件等气候因子与ENSO事件的特征进行对比统计分析得出,ENSO事件对长江流域降水量和气温的影响十分显著。厄尔尼诺事件年大多表现为温度升高和降水减少,且发生干旱灾害的概率升高。拉尼娜事件年一般表现为降水增多,洪涝灾害发生率相对增加。建国60 a来长江两次全流域性特大洪涝灾害都发生在厄尔尼诺事件向强拉尼娜事件转换年。厄尔尼诺事件对长江流域气候的影响要强于拉尼娜事件。2009—2010年的ENSO事件气象条件与1998年发生大洪水时的气象条件状况极其相似,2010—2011年和以后强ENSO冷暖事件转换年长江发生全流域性大洪水的可能性大,一定要做好气象监测和气象灾害防治。     

Exposure to tourism reduces stress-induced corticosterone levels in Galápagos marine iguanas

Unstressed and stress-induced plasma corticosterone levels in Galápagos marine iguanas (Amblyrhynchus cristatus) were compared in animals from a site heavily exposed to tourism to animals from a site undisturbed by humans. Initial corticosterone levels not only did not differ between the two groups, but they were 50% of levels in iguanas known to be chronically stressed. These data suggest that iguanas in tourist areas are not chronically stressed. Both groups of iguanas exhibited elevated corticosterone levels after 30 min of capture and restraint, indicating that they can physiologically respond to stressful stimuli. The stress response was lower, however, at the tourist site. This result indicates that iguanas are physiologically affected by tourism, although it is presently unknown whether these changes are ultimately beneficial or harmful.     

Hazard,risk and agrarian adaptations in a hyperarid watershed: El Niño floods,streambank erosion,and the cultural bounds of vulnerability in the Andean Middle Horizon

Peru's Osmore drainage, also known as the Moquegua Valley, is one of the driest regions on Earth, yet agricultural development has supported complex societies in the basin for almost 4000 yrs because of canal construction and irrigation. We compare the distinct agrarian and settlement systems of three coeval archaeological cultures in this arid region, Huaracane, Wari and Tiwanaku, and how each adapted—or failed to adapt—to geomorphic and climatic hazards. Systematic settlement pattern survey and radiocarbon dating along with geomorphological analysis of flood history and riverine processes permit detailed discussion of agrarian strategies during the Formative (1800 BC–AD 500) and Middle Horizon (AD 500–1000) periods, with distinct settlement “niches” in terms of agricultural practices and longitudinal position in the drainage, and lateral location relative to the floodplain. These adaptive strategies each manifest distinct “bounds of adaptability” to natural hazard, climate events and social stressors, and thus varying risk profiles. Besides the continual risk associated with sustained droughts, the Atacama and Peruvian Coastal Desert and culturally connected highland Altiplano are also vulnerable to the vagaries of the El Niño-Southern Oscillation (ENSO) leading to catastrophic floods in the mid-valley during warm phase ENSOs, in tandem with simultaneous hazards in other regions inhabited by these transregional cultures. By comparing the archaeological record of Wari, Huaracane, and Tiwanaku culture settlements with geomorphic signatures of catastrophic El Niños, we show that the viability of each cultural sequence depended on specific relationships to floodplain streambank erosion, construction and reworking over multiple time scales.     

Relations between El Nino/Southern Oscillation anomalies and wildland fire activity in the United States

The 1982–1983 El Nino resulted in climatic anomalies on a global scale, including record high wildland fire activity in Indonesia and record low activity in the United States. This paper describes the El Nino/Southern Oscillation phenomena and possible teleconnections to United States weather. Because precursors of an El Nino may be evident several months before the onset of an event, the phenomena has potential for long-range fire activity predictions.Using 53 years of data, we tested the hypothesis that El Nino events affect annual fire occurrence and area burned in the United States. We found a statistically significant relation between El Nino events and decreased fire activity in the South. Results for the North-Central and Eastern states are weak or inconsistent. There is no evidence for any relation with the Pacific Coast or Rocky Mountain states. Despite the coarse, exploratory nature of this study, the results are sufficiently encouraging to warrant more detailed examination.     

Conservation action in the Galàpagos: feral pig (Sus scrofa) eradication from Santiago Island

Introduced mammals are major drivers of extinction and ecosystem change. As omnivores, feral pigs (Sus scrofa) are responsible for wholesale adverse effects on islands. Here, we report on the eradication of feral pigs from Santiago Island in the Galápagos Archipelago, Ecuador, which is the largest insular pig removal to date. Using a combination of ground hunting and poisoning, over 18,000 pigs were removed during this 30-year eradication campaign. A sustained effort, an effective poisoning campaign concurrent with the hunting program, access to animals by cutting more trails, and an intensive monitoring program all proved critical to the successful eradication. While low and fluctuating control efforts may help protect select native species, current eradication methods, limited conservation funds, and the potential negative non-target impacts of sustained control efforts all favor an intense eradication effort, rather than a sustained control program. The successful removal of pigs from Santiago Island sets a new precedent, nearly doubling the current size of a successful eradication, and is leading to more ambitious projects. However, now we must turn toward increasing eradication efficiency. Given limited conservation funds, we can no longer afford to spend decades removing introduced mammals from islands.     

Assessing the impact of fisheries, climate and disease on the dynamics of the Indian yellow-nosed Albatross

Many seabird populations are currently decreasing, especially albatrosses for which the primary threat is recognised to be mortality in fisheries. Introduced predators, climate change and other factors such as diseases can also have large impacts on seabirds. Here, we assessed the relative effect of three potential threats: climate, fisheries and diseases on the demography of an endangered marine predator and modelled its population dynamics to project its size under different scenarios. We based our study on a long-term monitoring of a colony of individually marked Indian yellow-nosed albatrosses at Amsterdam Island, subtropical Indian Ocean, that has declined during the past twenty years. We found no evidence for an impact of legal tuna longlining on demographic parameters. Hatching success was lower during El Niño years but survival (0.902 ± 0.011) was not affected by climatic factors. Avian cholera caused high chick mortality (0.808 ± 0.181) which in turn probably triggered the high emigration rate (0.038 ± 0.011) through dispersal of failed breeders. This colony has a high risk of extinction. However, the rest of the population at Amsterdam Island seemingly not affected to the same extent, declined but stabilised since 1998. Matrix models indicated that lowered adult survival and the very low breeding success, resulting in low recruitment, have both contributed to the decline of the yellow-nosed albatross colony until the mid-1990s, but that more recent decline was primarily caused by low fledging success. Our results highlight that potential threats such as fisheries, diseases or climate have to be considered simultaneously to disentangle their roles when assessing the conservation status of a marine predator species.     

ENSO classification indices and summer crop yields in the Southeastern USA

This research uses a quantitative methodology for directly comparing the responses of observed crop yields in the SE USA to ENSO phenomena classified using dissimilar ENSO indices. ENSO condition has been related to agricultural yields in many parts of the world. It has been generally recognized that the strongest effects on spring and summer crops occur during the boreal winter, and therefore most directly affect spring-summer field crops in the southern hemisphere. However, some ENSO effects on northern hemisphere spring and summer crops have been found, particularly when researchers have used sub-annual indicators of ENSO conditions that, unlike annual ENSO indices, distinguish between continuity and change prior to or during the crop season. To evaluate the utility of such sub-annual ENSO indicators for agriculture in the SE USA, a tercile-based scoring system was devised to compare four distinct ENSO indices: three monthly ENSO indices and the JMA annual ENSO index. Each index was scored in its ability to predict crop yields differing from the historically normal tercile for corn (Zea mays L.), cotton (Gossypiumhirsutum L.), and peanut (Arachis hypogaea L.). Annual crop yield data were used from selected counties in five Southeastern USA states. No geographic differentiation among the data was included in the analysis. This aggregation of county data increased the sample size for each crop, to address the limitation of a short time-series (47 years) distributed among up to 9 ENSO categories. Statistical significance was compared using contingency tables and the Fisher exact test. Peanut and corn yield response matched best to the Multivariate ENSO Index (MEI) and cotton, to the Oceanic Niño Index (ONI). The MEI and ONI are quantitative indices, while the lower-scoring JMA and Modified-JMA indices are categorical. Therefore, future efforts may reduce the focus on categorical (Niño, Niña, Neutral) classification, and explore the response of crop yields to quantitative ENSO data.     

Effects of the introduced parasite Philornis downsi on nestling growth and mortality in the medium ground finch (Geospiza fortis)

Invasive species have the potential to detrimentally affect native ecosystems by out competing or directly preying upon native organisms, and have been implicated in the extinction of endemic populations. One potentially devastating introduced species in the Galápagos Islands is the parasitic fly Philornis downsi. As larvae, P. downsi parasitize nestling birds and have been associated with high nestling mortality and reduced growth rates. Here I document nestling growth and mortality in a bimodal population of the medium ground finch, Geospiza fortis. Observations were conducted over three years, and under variable ecological conditions. Annual parasite prevalence in nests ranged from 64% to 98%, and nestling mortality in nests with parasites ranged from 16% to 37%. Parasite load and parasite load per nestling follow a skewed distribution with many nests having relatively few parasites, and few nest having many. Parasite load, however, was not correlated with onset of breeding, clutch size, the number of nestlings, nestling survival or fledgling success. Parasite load per nestling, on the other hand, was correlated with clutch initiation date and the proportion of nestlings that died in parasitized nests. Neither nestling size nor growth rate differed between parasitized and unparasitized nests. In addition, male and female beak morphology was not correlated with parasite load, breeding variables or nestling survival. Thus, while overall mortality due to parasitism is high, ecological conditions and possible host defenses may potentially counter some of the detrimental affects of P. downsi on nestling size and growth. These results taken together suggest that parasitism of P. downsi larvae on nestling G. fortis has the potential to lead to large population declines.     

Environmental uncertainty and commercial fishing: Effects on Peruvian guano birds

The roles of the El Niño phenomenon, a cessation of upwelling, and of human fishing in limiting populations of Peruvian guano birds (Guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata, and Peruvian brown pelican Pelecanus occidentalis thagus) were investigated using historical records and the scientific literature. In normal years, the combined populations of the three species showed an 18% increase per year. El Niños occurred at approximately five-year intervals and caused a population decrease of 17% and 35% desertion of nesting areas. In severe El Niños at approximately 12-year intervals, there was a mean decrease of 47% and complete nesting failure. El Niños varied in severity and a possible explanation is presented.In non-El Niño years, commercial fishing did not directly control bird populations through starvation of adults or drowning in nets. Increased commercial fishing apparently led to decreased nesting success: the greater the quantity of fish landed, the lower the percentage increase of the bird population that year.Short-lived, heavily-fished species such as anchoveta are in constant danger of population collapse during years of occasional, unpredictable reproductive failure. Bird populations could be maintained as buffers to prevent terminally-destructive overfishing by humans.     

The Galápagos giant tortoises (Geochelone elephantopus) Part I: Status of the surviving populations

Eleven of the original 15 races of Galápagos tortoises are known to survive. The populations of three of these may be self-replacing. The other eight are threatened with extinction, each by one or more of the following: (1) greatly decreased population size, (2) predation on nests and/or young by introduced pigs, dogs, cats, or black rats, and (3) competition for food resources with feral populations of goats or donkeys. Human exploitation of the tortoise populations has become relatively unimportant in the past few years for a number of reasons, including more wide-spread acceptance of protective laws and the increasing activity of the Servicio del Parque Nacional Galápagos and the Charles Darwin Research Station. Data on population sizes and sex structures, reproductive potential, and natural mortality rates suggest that all but two of the endangered populations could remain stable or even increase in size if the effects of introduced mammals were eliminated or greatly lessened.     

El Niño induced local extinction of coral reef bryozoan species from Northern Bahia, Brazil

The 1997-1998 El-Niño Southern Oscillation was the most severe on record and dramatically impacted corals worldwide. However, the effect of this event on the associated community of reef organisms has received much less attention. The composition of the bryozoan assemblage from the coral reefs of Northern Bahia, Brazil were monitored annually from 1995 to 2000, allowing the investigation of the effects of this large-scale stressor on an important, diverse, yet understudied component of the coral reef system. Bryozoan samples (35 replicates/reef) were collected during April/May from four shallow bank reefs (10-40 m depth) located a few kilometres off the coast, together with measurements of the associated environmental parameters. Currently 157 species have been recorded from the study area, but significant reductions in density and diversity were apparent between pre- and post El-Niño Southern Oscillation (ENSO) years, multivariate analysis denoting significant changes in assemblage composition. A total of 61 species were unrecorded following the 1997-1998 ENSO event (22 species from 1997; 25 further species from 1998 and 14 more from 1999). These included several species endemic to Brazil, suggesting that the 1997-1998 ENSO has had a marked influence on the reef bryozoan community, resulting in the local extinction of several species. Bryozoan mortalities were probably initiated by elevated temperatures, but continued disappearance of species for 2 years after ENSO suggests other indirect factors are also influential. These results demonstrate that ENSO events can have severe long-term impacts on the biodiversity of coral reefs, with important conservation consequences.     

Death from anthropogenic causes is partially compensatory in recovering wolf populations

There is substantial interest in how mortality rates affect animal populations, but mechanisms explaining when and under what circumstances particular causes of death incur demographic responses are far from clear. In theory, small or expanding populations should experience additive mortality from anthropogenic causes of death, but whether such effects are homogenous across a population or expressed only in certain high-risk individuals is open for debate. We used competing risks models to analyze mortality patterns among radio-collared wolves (Canis lupus, n = 711) from three populations in northwestern United States (1982-2004), and evaluated the degree to which anthropogenic mortality was additive vs. compensatory to natural demographic processes. Almost 80% (n = 320) of wolves dying of known fates were killed by anthropogenic causes (legal control, illegal killing, harvest in Canada, vehicle collision), and additive effects of anthropogenic mortality were most pronounced in northwestern Montana where wolf exposure to humans and livestock was high compared to either the Greater Yellowstone Area or central Idaho, where anthropogenic risk was lower. In contrast, risk from natural hazards was lower in northwestern Montana than in the other areas, implying some degree of compensatory mortality from anthropogenic risk. Animals recruited to the study following human-wolf conflict had markedly higher anthropogenic risk than those recruited for standard monitoring purposes, and juvenile wolves as well as dispersers, succumbed to higher anthropogenic risk. Multivariate models revealed that increasing wolf population density promoted higher anthropogenic risk and reduced natural risk, indicating that partially-compensatory effects of anthropogenic mortality actually became increasingly additive with population density. The observed compensatory mortality and hazard heterogeneity in our study implies that demographic responses to mortality risk may be complex and more subtle than previously thought; the density-dependent effect of anthropogenic mortality portends a stabilizing influence of humans on recovering wolf populations. We conclude that future assessment of the role of anthropogenic mortality should include individual-based hazard estimation as a complement to traditional population-level approaches.     

Modeling the reintroduction of lynx to the southern portion of its range

We modeled populations of lynx (Lynx canadensis) and snowshoe hares (Lepus americanus) to determine prey densities required for persistence of lynx translocated to the southern portion of the species' range. The models suggested that a density of 1.1-1.8 hares/h is required for lynx persistence; these densities are higher than those reported for most hare populations across the USA. We found that lynx dispersal and density-independent mortality substantially increased the hare density required for lynx persistence. Reintroduction success was associated with number of release events, total number of animals released, and timing of release relative to the phase of the hare population cycle. However, no release protocol could override the negative effects of low prey density or high population losses. We conclude that successful lynx reintroduction requires high hare densities and minimal anthropogenic disturbance; few areas in the contiguous USA currently posses such qualities.