Bio-Economics of Large-Scale Eradication of Feral Goats From Santiago Island,Galápagos

ABSTRACT Invasive mammals are premier drivers of extinction and ecosystem change, particularly on islands. In the 1960s, conservation practitioners started developing techniques to eradicate invasive mammal populations from islands. Larger and more biologically complex islands are being targeted for restoration worldwide. We conducted a feral goat (Capra hircus) eradication campaign on Santiago Island in the Galápagos archipelago, which was an unprecedented advance in the ability to reverse biodiversity impacts by invasive species. We removed >79,000 goats from Santiago Island (58,465 ha) in <4.5 years, at an approximate cost of US$6.1 million. An eradication ethic combined with a suite of techniques and technologies made eradication possible. A field-based Geographic Information System facilitated an adaptive management strategy, including adjustment and integration of hunting methods. Specialized ground hunting techniques with dogs removed most of the goat population. Aerial hunting by helicopter and Judas goat techniques were also critical. Mata Hari goats, sterilized female Judas goats induced into a long-term estrus, removed males from the remnant feral population at an elevated rate, which likely decreased the length and cost of the eradication campaign. The last 1,000 goats cost US$2.0 million to remove; we spent an additional US$467,064 on monitoring to confirm eradication. Aerial hunting is cost-effective even in countries where labor is inexpensive. Local sociopolitical environments and best practices emerging from large-scale, fast-paced eradications should drive future strategies. For nonnative ungulate eradications, island size is arguably no longer the limiting factor. Future challenges will involve removing invasive mammals from large inhabited islands while increasing cost-effectiveness of removing low-density populations and confirming eradication. Those challenges will require leveraging technology and applying theory from other disciplines, along with conservation practitioners working alongside sociologists and educators.     

Feral donkey (<Emphasis Type="Italic">Equus asinus</Emphasis>) eradications in the Galápagos

Introduced herbivores are major drivers of ecosystem change and biodiversity loss, particularly on islands. Tools and techniques now exist to routinely remove introduced herbivores from islands, providing a powerful conservation tool. Here, we summarize the few documented feral donkey removals on islands worldwide, and report on the removal of populations from the Galápagos archipelago, Ecuador. After decades of sporadic control programs on Santiago Island and Alcedo Volcano, Isabela Island, donkey populations were removed from both areas, concurrent with a goat eradication program. Both ground and aerial hunting programs were utilized. The latter method was highly efficient; donkeys were removed from Santiago Island with less than 80 h of aerial hunting. Given the clear impacts of introduced herbivores on islands worldwide, feral donkey populations should be routinely removed from islands.     

Modelling the recovery of resident shorebirds following a fox eradication program using citizen science data

Historically, the land-based threat to shorebird colonies on Phillip Island, Victoria, Australia, was fox predation. As a result, a fox eradication programme consisting of three phases: knock-down (i.e., 2006), clean-up (i.e., 2011), and post-eradication. In 2011 an effective knock-down was declared, signalling the beginning of the clean-up phase. The purpose of this research is to assess the recovery of six resident shorebird species on Phillip Island following fox removal. The statistical methodologies used are novel for assessing bird species population recovery following a successful predator eradication program. We used citizen science data from 2003 to 2017, extracted from the Atlas of Living Australia. The first analysis method used INLA modelling, which relied on a Negative Binomial distribution for bird counts to look for upward trends in shorebird populations during the fox eradication operation. The second method use changepoint analysis techniques to see whether successive phases of the eradication process were associated with changes in bird population numbers. Four of the six shorebird species investigated responded positively to reduced fox populations over the 15-year study, and all changepoint approaches consistently recognised the start of the clean-up phase, with less consistency identifying the start of the knock-down phase. Since 2006, the INLA models indicate a significant increase in the upward trend of shorebird populations for three of the six shorebird species investigated. Agreement across the four changepoint techniques indicates that changes in bird numbers were associated with the date of the eradication program's clean-up phase for all of these shorebird species. These results demonstrate some promise for these methods to monitor native species recovery during eradication programs.     

Predicted population trends for Cozumel Curassows (Crax rubra griscomi): empirical evidence and predictive models in the face of climate change

ABSTRACT We assessed the current conservation status and population trends of Cozumel Curassows (Crax rubra griscomi), a critically endangered and endemic cracid of Cozumel Island, Mexico. Distance sampling was used to estimate the population size in 2005 and compare it with a previous estimate in 1995. Population viability analysis (PVA) was used to model population trends, considering ecological and environmental conditions prevailing in 1995 and 2005. Additional PVA models were constructed using different hypothetical scenarios to assess the effects of particular factors and management strategies on population trends. We estimated a population size of 372 ± 155 curassows in the island's tropical semideciduous forest, before two hurricanes hit Cozumel Island in 2005. PVA modeling predicted a steep decline in population size in the ensuing decades as a result of an increased frequency and intensity of hurricanes related to global climate change, and an observed female bias in the sex ratio likely caused by a higher mortality rate for adult males. We recommend urgent management actions for the long‐term conservation of Cozumel Curassows, including a ban on hunting, eradication of the feral fauna, particularly dogs, and implementation of a captive‐breeding program to supplement the wild population.     

Population Response of Giant Galápagos Tortoises to Feral Goat Removal

Efforts to eradicate nonnative mammals to restore oceanic island ecosystems have become increasingly successful but parallel tracking of response by native species for which control efforts are intended to benefit has been rare. A major campaign to eradicate nonnative goats and burros was initiated in 1995 on Alcedo Volcano in the Galápagos Archipelago that ultimately removed 62,868 goats and eliminated them by 2006. Planners of the eradication program had the foresight to invest in intensive monitoring of the status of the volcano's giant tortoise (Chelonoidis nigra) population whose welfare was a primary motivation for the eradication effort. Monitoring revealed an increase in the proportion of juveniles among all tortoises as well as increased growth rates of individual tortoises on Alcedo Volcano from earlier to later phases of the eradication campaign. Over the same time frame in a control population on nearby Santa Cruz Island (where goats and donkeys were not removed) juvenile fraction and individual growth rates remained unchanged. Although goat removal coincided with occurrence of a rare climatic event that simultaneously boosted forage availability for tortoises, failure to observe a comparable improvement in the control population implies that removal of goats and burros was the primary causative factor of improving population status of tortoises on Alcedo Volcano.     

Paleochemistry of manganese in corals from the Galapagos Islands

Approximately 550 measurements of Mn/Ca ratios in three corals from the western Galapagos Islands have been performed to reconstruct a 380-year history of surface ocean variability with respect to this trace element. The time period studied encompasses 1600 A.D. to 1978. Manganese is inferred to be lattice-bound in coralline aragonite at 10–50% of its seawater proportion to calcium; uncertainty about the distribution coefficient stems from inherent variability of oceanic Mn in nearshore settings. Interannual variations at Urvina Bay, Isabela Island are generally small, with the exception of a few decades during the nineteenth century. A large positive Mn/Ca anomaly found between 1821–1830 is hypothesized to have resulted from a major volcanic eruption on nearby Fernandina Island in 1825. On intrannual timescales a pronounced cycle occurs in response to seasonal upwelling. Quarterly changes in Mn/Ca are six months out-of-phase with Cd/Ca variations-a reflection of the opposite distributions of these metals in the upper waters of the eastern Pacific. High frequency reconstructions over brief time intervals from the 17th, 18th, and 20th century reveal that the seasonal onset of warm and cool phases near Galapagos has persisted for at least 340 years. A quantitative assessment of historical changes in upwelling intensity is complicated by offsets in background Mn levels recorded by different corals. One apparent longterm feature is an overall decline in skeletal Mn concentrations from 1600–1978 which results in a net decrease of 20–30%. Several possible explanations exist for this trend, ranging from accumulation of a persistent diagenetic Mn phase in fossil aragonite to a temporal shift in oceanic/atmospheric Mn fluxes reaching the surface waters of the Galapagos Islands.     

First direct, site-wide penguin survey at Deception Island, Antarctica, suggests significant declines in breeding chinstrap penguins

Deception Island (62°57′S, 60°38′W) is one of the most frequently visited locations in Antarctica, prompting speculation that tourism may have a negative impact on the island’s breeding chinstrap penguins (Pygoscelis antarctica). Discussions regarding appropriate management of Deception Island and its largest penguin colony at Baily Head have thus far operated in the absence of concrete information regarding the current size of the penguin population at Deception Island or long-term changes in abundance. In the first ever field census of individual penguin nests at Deception Island (December 2–14, 2011), we find 79,849 breeding pairs of chinstrap penguins, including 50,408 breeding pairs at Baily Head and 19,177 breeding pairs at Vapour Col. Our field census, combined with a simulation designed to capture uncertainty in an earlier population estimate by Shuford and Spear (Br Antarct Surv Bull 81:19–30, 1988), suggests a significant (>50?%) decline in the abundance of chinstraps breeding at Baily Head since 1986/1987. A comparative analysis of high-resolution satellite imagery for the 2002/2003 and the 2009/2010 seasons suggests a 39?% (95th percentile CI?=?6–71?%) decline (from 85,473?±?23,352 to 52,372?±?14,309 breeding pairs) over that 7-year period and provides independent confirmation of population decline in the abundance of breeding chinstrap penguins at Baily Head. The decline in chinstrap penguins at Baily Head is consistent with declines in this species throughout the region, including sites that receive little or no tourism; as a consequence of regional environmental changes that currently represent the dominant influence on penguin dynamics, we cannot ascribe any direct link between chinstrap declines and tourism from this study.     

Population size and trends of southern giant petrels (<Emphasis Type="Italic">Macronectes giganteus</Emphasis>) nesting at Signy Island,South Orkney Islands

The southern giant petrel (Macronectes giganteus) has a circumpolar distribution and breeds on subantarctic islands and a few continental Antarctic sites. Although this species has recently been down-listed to “Least Concern” by the World Conservation Union (IUCN), there are strong fluctuations in abundance and variable long-term trends recorded at different sites. Systematic, long-term monitoring is essential to determine drivers underlying its population dynamics. Here, we examine long-term changes in population size and productivity of southern giant petrels at Signy Island, South Orkney Islands. Comparing estimated numbers of breeding pairs over the whole island in 2000/2001, 2005/2006, 2009/2010 and 2014/2015 with historical data revealed several phases of population change: a 64 % decline (6.2 % per annum) from 1968/1969 to 1984/1985, a 162 % increase (6.2 % per annum) to 2000/2001, stability until 2005/2006, a 56 % decline (18.3 % per annum) to 2009/2010 and stability until 2014/2015. This represents a 1.8 % decline per annum between 1968/1969 and 2014/2015. Annual counts within focal study areas suggested a more rapid increase from 1996/1997 to 2006/2007, but the same downward trend from 2006/2007 to present, underlining potential pitfalls in inferring trends from part-island counts. There was also a 20 % decline in breeding success from 1996/1997 to 2014/2015. Our results indicate substantial fluctuations in southern giant petrel abundance at Signy Island over 4–5 decades and a recent decline in breeding numbers and success. As the southern giant petrels breeding at the South Orkney Islands represents ~5–10 % of the global population, continuation of these declines would be of high conservation concern.     

Health evaluation of Galapagos Hawks (Buteo galapagoensis) on Santiago Island, Galapagos

Galapagos Hawks (Buteo galapagoensis), the only endemic, diurnal raptor species in Galapagos, are currently distributed on eight Galapagos Islands having been extirpated from three of the human-inhabited islands. In January 2009, we performed health assessments of 89 Galapagos Hawks on Santiago Island, Galapagos. Four of the 89 Galapagos Hawks (4%) evaluated had physical abnormalities. Blood parameters did not differ between males and females, except for aspartate transaminase values, which were significantly higher in females than males. No Galapagos Hawks tested positive for antibodies to avian encephalitis virus, Marek virus, and paramyxovirus-1 or to haemosporidian antigen. Chlamydophila psittaci antigen was detected in 2 of 86 Galapagos Hawks (2%), with 24 of 43 Galapagos Hawks (56%) antibody-positive for avian adenovirus-1 and 1 of 48 Galapagos Hawks (2%) antibody positive for Toxoplasma gondii. There were no significant differences in infectious disease results based on sex. This study contributes to the understanding of the health status of the Galapagos Hawk and to the establishment of baseline information for the species.     

Long-term variability and density dependence in Hudson River Dreissena populations

1. We used a 27-year record of Dreissena populations in the freshwater tidal Hudson River to describe interannual variation in population density, body size, and body condition; estimate long-term variation in recruitment, survivorship, and shell growth; and assess possible controls on the populations.
2. Dreissena populations in the Hudson have been highly variable, with interannual ranges of c. 100-fold in abundance and biomass, and 7-fold in mean body mass. This large interannual variation arises from both long-term trends and 2–5-year cycles.
3. Long-term trends include the 2008 appearance of the quagga mussel (Dreissena rostriformis), which still forms a small part (<10%) of the dreissenid community, and a decline in zebra mussel body size. The decline in body size was caused by a long-term decline in adult survivorship rather than a decline in rates of shell growth. We could detect no long-term trends in adult abundance or spread of Dreissena onto soft sediments in the Hudson.
4. We observed persistent, strong cycles in adult abundance and body size. These were driven by the appearance and decay of eight dominant year classes over the 27 years of our study, and were a result of temporal variation in recruitment rather than temporal variation in survivorship. The observed strongly irregular recruitment appears to arise from strong adult–larval interactions, and is consistent with previous simulation model results showing that interactions between adults and larvae can drive persistent cycling.
5. We found evidence that negative density dependence affects recruitment, somatic growth, and body condition of Dreissena in the Hudson. Warm summers may also cause high adult mortality.
6. We put our results into the context of a conceptual model of Dreissena population dynamics, and argue that neither the dynamics nor the controls of populations of these important invaders is known satisfactorily.

     

Habitat as a potential factor limiting the recovery of a population of nocturnal seabirds

We asked whether the lack of a population response by ancient murrelets (Synthliboramphus antiquus) to eradication of rats (Rattus spp.) at Langara Island could be due to a change in vegetative cover. We quantified ancient murrelet habitat associations on 12 islands and assessed changes in vegetation at Langara Island between 1981 and 2007. We found that ancient murrelets exhibit a high degree of flexibility in their use of available breeding habitats, and we noted no changes over time. Thus, recovery of ancient murrelets at Langara Island is unlikely to be limited by habitat quality. We propose artificial social attraction as a method to speed recovery. © 2011 The Wildlife Society.     

Decline in abundance and apparent survival rates of fin whales (Balaenoptera physalus) in the northern Gulf of St. Lawrence

Estimates of abundance and survivorship provide quantifiable measures to monitor populations and to define and understand their conservation status. This study investigated changes in abundance and survival rates of fin whales (Balaenoptera physalus) in the northern Gulf of St. Lawrence in the context of anthropogenic pressures and changing environmental conditions. A long‐term data set, consisting of 35 years of photo‐identification surveys and comprising more than 5,000 identifications of 507 individuals, formed the basis of this mark–recapture study. Based on model selection using corrected Akaike Information Criterion, the most parsimonious Cormack–Jolly–Seber model included a linear temporal trend in noncalf apparent survival rates with a sharp decline in the last 5 years of the study and a median survival rate of 0.946 (95% confidence interval (CI) 0.910–0.967). To account for capture heterogeneity due to divergent patterns of site fidelity, agglomerative hierarchical cluster analysis was employed to categorize individuals based on their annual and survey site fidelity indices. However, the negative trend in survivorship remained and was corroborated by a significant decline in the estimated super‐population size from 335 (95% CI 321–348) individuals in 2004–2010 to 291 (95% CI 270–312) individuals in 2010–2016. Concurrently, a negative trend was estimated in recruitment to the population, supported by a sharp decrease in the number of observed calves. Ship strikes and changes in prey availability are potential drivers of the observed decline in fin whale abundance. The combination of clustering methods with mark–recapture represents a flexible way to investigate the effects of site fidelity on demographic variables and is broadly applicable to other individual‐based studies.     

Colonization history of Galapagos giant tortoises: Insights from mitogenomes support the progression rule

Galapagos giant tortoises (Chelonoidis spp.) are a group of large, long-lived reptiles that includes 14 species, 11 of which are extant and threatened by human activities and introductions of non-native species. Here, we evaluated the phylogenetic relationships of all extant and two extinct species (Chelonoidis abingdonii from the island of Pinta and Chelonoidis niger from the island of Floreana) using Bayesian and maximum likelihood analysis of complete or nearly complete mitochondrial genomes. We also provide an updated phylogeographic scenario of their colonization of the Galapagos Islands using chrono-phylogenetic and biogeographic approaches. The resulting phylogenetic trees show three major groups of species: one from the southern, central, and western Galapagos Islands; the second from the northwestern islands; and the third group from the northern, central, and eastern Galapagos Islands. The time-calibrated phylogenetic and ancestral area reconstructions generally align with the geologic ages of the islands. The divergence of the Galapagos giant tortoises from their South American ancestor likely occurred in the upper Miocene. Their diversification on the Galapagos adheres to the island progression rule, starting in the Pleistocene with the dispersal of the ancestral form from the two oldest islands (San Cristóbal and Española) to Santa Cruz, Santiago, and Pinta, followed by multiple colonizations from different sources within the archipelago. Our work provides an example of how to reconstruct the history of endangered taxa in spite of extinctions and human-mediated dispersal events and provides a framework for evaluating the contribution of colonization and in situ speciation to the diversity of other Galapagos lineages.     

Evaluating Genetic Diversity for the Conservation of the Threatened Galapagos Endemic Calandrinia galapagosa (Portulacaceae)

Calandrinia galapagosa (Portulacaceae) is a threatened plant endemic to San Cristobal Island, Galapagos. The species is threatened by goat herbivory: less than 1300 individuals remain in nine populations, three of which have been fenced for protection. It is not known whether these three populations are genetically representative of the species at large or whether the three distinct color morphs exhibited by the species represent different taxonomic units. We measured the frequency of 113 polymorphic amplified fragment length polymorphism markers in 189 specimens representing three different phenotypes from all nine populations to relate the genetic structure of the species to both its geological history and morphological variation. The analysis revealed high diversity within populations and low, but significant, differentiation between populations. In addition, no genetic diversification was observed across color morphs. We conclude that (1) regular gene flow has historically occurred between populations, and (2) individual color morphs are not separate taxa. We identify another population that should be fenced to ensure maximal protection of genetic variation. Our research serves as a case study of using population molecular genetics for optimizing protection strategies for rare plants in tropical archipelagoes affected by introduced herbivores.     

Population dynamics of the West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire): a simulation analysis

The West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) is a major pest of the sweet potato Ipomoea batatas (L.) Lam. and this weevil is a target of an eradication program using the Sterile Insect Technique in Okinawa Prefecture, Japan. Understanding the population ecology is essential in the planning of an eradication program; hence, a host‐plant infestation survey and light trap survey have been conducted to monitor the population dynamics of the weevil on Kume Island (Okinawa Prefecture), which is the target area of the trial weevil eradication project. Seasonal tendencies of weevil density were found in these field surveys, but the tendency found in the host‐plant infestation survey was not seen every year, and the effectiveness of the light trap is somewhat suspect. To confirm the reliability of the tendency observed in these field surveys, the present study attempted to explain the tendency by a seasonal temperature change using a temperature‐based model of weevil population dynamics. The seasonal changes of weevil density differed according to host plants and host‐plant fields. The seasonal changes of weevil density inside the host plant Ipomoea indica and outside the host plants in I. indica fields were consistent with those predicted by the model. However, those inside the host plant Ipomoea pes‐caprae in the host‐plant infestation survey were contrary to the predicted ones, and those observed outside host plants in I. pes‐caprae fields by the light trap survey were not in good agreement with the predicted ones. It was concluded that the seasonal change of the weevil density observed in I. indica and I. indica fields can be explained by a seasonal temperature change, but factors other than seasonal temperature change are needed to explain those in I. pes‐caprae and I. pes‐caprae fields.     

Residency of the scalloped hammerhead shark (<Emphasis Type="Italic">Sphyrna lewini</Emphasis>) at Malpelo Island and evidence of migration to other islands in the Eastern Tropical Pacific

Sixty nine hammerhead sharks, Sphyrna lewini, were tagged at Malpelo Island (Colombia) with ultrasonic transmitters during March 2006, 2007 and 2008, as part of a study to understand their residency at the island and their horizontal and vertical movements. Five sharks visited Cocos Island, 627 km distant from Malpelo. One of the sharks that appeared at Cocos Island also visited the Galapagos Islands, 710 km from Cocos, a month later. There is connectivity of Sphyrna lewini between Malpelo, Cocos and the Galapagos Islands, but the frequency of movements between the islands appears to be relatively low (<7% of the tagged sharks). The most common depth at which the sharks swam coincided with the thermocline (rs = 0.72, p < 0.01). The depth of the thermocline varied depending on the time of the year. Nocturnal detections of the sharks were more frequent during the cold season than during the warm season (W = 60, p < 0.01). We also found that hammerheads spent significantly more time on the up-current side of the island (Kruskal-Wallis = 31.1008; p < 0.01). This study contributes to the knowledge of hammerhead sharks not only in Malpelo Island but also at a regional level in the Eastern Tropical Pacific.     

Declines in summer bat activity in central New England 4 years following the initial detection of white-nose syndrome

White-nose syndrome (WNS) was first reported in a hibernating bat population in central New York State in February 2006. Since 2006, WNS has been reported from bat hibernacula across much of eastern United States and adjacent Canada and has been associated with a dramatic decline in the populations of hibernating bats in the northeastern U.S. We are only beginning to discover how these declines are manifest in changes in summer bat abundance and activity at local scales. A 3-year (2004–2006) acoustic survey showed that the forested watershed of the Quabbin Reservoir in central Massachusetts supported an abundant and species-rich summer bat community. In 2010, 4-years following the initial occurrence of WNS, a re-survey of the same habitats and sites found a 72% reduction in bat activity on the watershed. This is the identical rate of decline reported from cave hibernacula surveys (73%). This decline in summer activity levels is most likely a consequence of WNS-caused mortality. The impacts of population losses of this magnitude of a once widespread and abundant taxa are unknown but are presumed to be ecologically significant.     

Cats and seabirds: effects of feral Domestic Cat Felis silvestris catus eradication on the population of Sooty Terns Onychoprion fuscata on Ascension Island, South Atlantic

The population of Sooty Terns Onychoprion fuscata breeding on Ascension Island in the Atlantic Ocean was monitored over 17 years (1990–2007). This period spanned the programme of feral Domestic Cat Felis silvestris catus eradication from the island, which commenced in 2001 with the last Cat recorded in 2004. We report on the abundance of Sooty Terns and Black Rats Rattus rattus before and after Cat eradication. The Sooty Tern breeding population in the 1990s averaged 368 000 and Cats were killing Terns at an average rate of 33 adults per night. Following Cat eradication, adult Terns are no longer predated. However, egg predation by both Rats and Common Mynas Acridotheres tristis has continued with Mynas destroying more eggs than Rats. Unexpectedly, we observed a change in Rat predatory behaviour. Following Cat eradication, Rats have become a major predator of Sooty Tern chicks. Despite this change, the Tern population has shown a season-on-season increase since Cat eradication, 48.8% in 2005, 8.2% in 2006 and 6.1% in 2007, and the breeding population increased to 420 000 birds in 2007. Incubation success improved from 66.0 to 84.4% during Cat eradication, before dropping down again to 67.9% after Cats were eradicated and Rat control measures were introduced. Index traplines were set for Rats and Rat numbers fluctuated widely immediately after Cats were eradicated but there were no significant differences that could be attributed to changes in Cat numbers. Ascension Island Sooty Terns breed every 9.6 months and juveniles defer breeding for seven seasons. Hence 2008 is the first year in which an increase in the breeding Sooty Tern population directly attributable to Cat eradication is likely to be detected. We conclude that long-term monitoring is essential to guide conservation practice even in this relatively simple predator–prey system.     

Reproductive success and survivorship of the Galapagos Hawk Buteo galapagoensis: potential costs and benefits of cooperative polyandry

The Galapagos Hawk Buteo galapagoensis is one of few species showing cooperative polyandry. As many as four males may mate with one female and share a territory and nest, Such groups, on average, produce more young than monogamous pairs, but on a per male basis (assuming that each male of a group has an equal chance to fertilize each young) polyandrous males produce fewer young per year than monogamous males. Territorial breeding birds show higher yearly survivorship (90%) than non-territorial, non-breeding birds (50% or less). All territorial birds seem to remain on their territories for life, and monogamous pairs actually constitute the remnants of polyandrous groups. Some potential factors producing such an unusual system are discussed. Limited breeding space and high mortality among non-territorial birds seem to work in favour of group breeding in this and other cooperative species. Polyandry may be favoured because of the reduced value of extra females in hawk breeding behaviour. Equal-status males may occur because of the lack of kinship among cooperating males and the high potential genetic cost of being a helper in a long-lived, low fecundity species. Other possible factors favouring group breeding are territory acquisition and the variable nature of the Galapagos climate.     

The population decline of the New Zealand sea lion Phocarctos hookeri: a review of possible causes

• 1 The New Zealand (NZ) sea lion Phocarctos hookeri is NZ's only endemic pinniped and is listed as ‘nationally critical’. The species breeds in the NZ sub‐Antarctic: 71% of the population at the Auckland Islands (2010 pup production: 1814 ± 39) and the remaining 29% on Campbell Island (726 pups in 2010).
• 2 Pup production at the Auckland Islands has declined by 40% since 1998 (1998: 3021 pups produced): only 1501 pups were born in 2009. This decline is directly linked to philopatric females not returning to breeding areas. While the Auckland Island population has declined, the Campbell Island population appears to be increasing slowly.
• 3 Potential reasons for the decline in the Auckland Island population, but not in the Campbell Island population, include non‐anthropogenic factors: (i) disease epizootics, (ii) predation, (iii) permanent dispersal or migration, (iv) environmental change; and anthropogenic impacts: (v) population ‘overshoot’, (vi) genetic effects, (vii) effects of contaminants, (viii) indirect effects of fisheries (i.e. resource competition) and (ix) direct effects of fisheries (i.e. by‐catch deaths). Of the nine potential reasons examined here, six can be discounted (ii–vii). Bacterial epizootics (i) occur in the NZ sea lion population, but their impact has predominantly increased pup mortality, which is unlikely to cause the severe decline observed, as pup mortality throughout the species is naturally high and variable.
• 4 The most plausible hypotheses, based on available evidence, are that the observed decline, in particular, the decreasing number of breeding females in the Auckland Island population, is caused by (viii) fisheries‐induced resource competition and (ix) fisheries‐related by‐catch. By‐catch is the main known anthropogenic cause of mortality in the species. Competition with fisheries resulting in resource competition, nutrient stress and decreased reproductive ability in NZ sea lions should be a priority area for future research.