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.     

A noninvasive demographic assessment of sea lions based on stage-specific abundances

A pressing need exists to develop new approaches for obtaining information on demographic rates without causing further threats to imperiled animal populations. In this paper, we illustrate and apply a data-fitting technique based on quadratic programming that uses stage-specific abundance data to estimate demographic rates and asymptotic population growth rates (lambda). We used data from seven breeding colonies of California sea lions (Zalophus californianus) in the Gulf of California, Mexico. Estimates of lambda were similar to those from previous studies relying on a diffusion approximation using trends in total abundance. On average, predicted abundances were within 24% of the observed value for the inverse estimation method and within 29% of the observed value for the diffusion approximation. Our results suggest that three of the seven populations are declining (lambda < 1), but as many as six may be at risk. Elasticity and sensitivity analyses suggest that population management in most sites should focus on the protection of adults, whose survival generally contributes the most to lambda. The quadratic programming approach is a promising noninvasive technique for estimating demographic rates and assessing the viability of populations of imperiled species.     

Reduction of the impact produced by sea lions on the fisheries in Mexico

Activities of fishermen in the Bay of La Paz, B. C. S. are focused to satisfy the local demand of fish and shellfish by using approximately 300 small outboard crafts equipped with gillnets.Sea lions in this Bay attack the fishes captured damaging both product and gear. We did experimental gillnet throws to determine the frequency and preferences of sea lions in fishing areas. 52 experimental gillnet throws with time averages of 2 hr were performed, rending an average of 30 kg of fish captured and less than 10% of damages to the net.. Traditional fishermen in this Bay usually left the net the whole night, (approximately 7.50 hr, obtaining an average of 50 kg of captured fish, but the damages to the nets is in average of 40%. The cost-benefit balance comparing our alternative fishing method, which includes the use of the gillnets during the afternoon, watching for sea lions and retiring the nets at their arrival, it is more sustainable and profitable than the traditional fishing method currently used by the local fishermen. This paper suggests how to minimize the harmful effects of the sea lions on the fishermen productivity and gear, maximizing the production and reducing the damage. Our alternative method is applicable to other regions where this harmful interaction is taking place. We conclude that the coexistence of sea lions-fisheries is feasible, by applying the simple measures that we propose.     

Age-structured modeling reveals long-term declines in the natality of western Steller sea lions.

Since the mid-1970s, the western Steller sea lion (Eumetopias jubatus), inhabiting Alaskan waters from Prince William Sound west through the Aleutian Islands, has declined by over 80%. Changing oceanographic conditions, competition from fishing operations, direct human-related mortality, and predators have been suggested as factors driving the decline, but the indirect and interactive nature of their effects on sea lions have made it difficult to attribute changes in abundance to specific factors. In part, this is because only changes in abundance, not changes in vital rates, are known. To determine how vital rates of the western Steller sea lion have changed during its 28-year decline, we first estimated the changes in Steller sea lion age structure using measurements of animals in aerial photographs taken during population surveys since 1985 in the central Gulf of Alaska (CGOA). We then fit an age-structured model with temporally varying vital rates to the age-structure data and to total population and pup counts. The model fits indicate that birth rate in the CGOA steadily declined from 1976 to 2004. Over the same period, survivorship first dropped severely in the early 1980s, when the population collapsed, and then survivorship steadily recovered. The best-fitting model indicates that in 2004, the birth rate in the central Gulf of Alaska was 36% lower than in the 1970s, while adult and juvenile survivorship were close to or slightly above 1970s levels. These predictions and other model predictions concerning population structure match independent field data from mark-recapture studies and photometric analyses. The dominant eigenvalue for the estimated 2004 Leslie matrix is 1.0014, indicating a stable population. The stability, however, depends on very high adult survival, and the shift in vital rates results in a population that is more sensitive to changes in adult survivorship. Although our modeling analysis focused exclusively on the central Gulf of Alaska, the western Gulf of Alaska and eastern Aleutians show a similar pattern of declining pup fraction with no increase in the juvenile, or pre-breeding, fraction. This suggests that declining birth rate may be a problem for western Steller sea lions across the Gulf of Alaska and into the Aleutian Islands.     

Assessing competition between Steller sea lions and the commercial groundfishery in Alaska: A bioenergetics modelling approach

In the last three decades the western stock of the Steller sea lion (Eumetopias jubatus) has declined by more than 85%. Nutritional stress resulting in increased juvenile mortality is one of the leading hypotheses to account for this decline. Competition between Steller sea lions and the commercial groundfishery for walleye pollock (Theragra chalcogramma) has been proposed as a mechanism underlying the nutritional stress. In order to examine the competition component of the nutritional stress hypothesis, we developed a bioenergetics-based model to project the population trends of Steller sea lions under various scenarios of continued groundfish harvest. Annual energy budgets were calculated for the Gulf of Alaska population of Steller sea lions, and compared with projected available energy from walleye pollock under a variety of harvest scenarios. Model simulations produced 50-year Steller sea lion population projections consistent with current trends, as well as with published projections for stable and increasing populations from stable age distribution life table models. Model simulations were unable to produce energy deficits sufficient to account for the decline in Steller sea lions, but do suggest areas where existing data need supplementing.     

Sexual reproduction in the corallimorpharian sea anemone Corynactis californica in a central California kelp forest

The seasonal cycle of sexual reproduction in the corallimorpharian sea anemone Corynactis californica (Calgren, 1936) was studied for 18 consecutive months (July 1987 to December 1988) at a subtidal area in the Hopkins Marine Refuge (HMLR), Pacific Grove, Monterey Bay, California. Samples were collected, histological sections were prepared, and gametocytes were examined and measured. C. californica grows by multiple or longitudianl fission to form single-sexed clones of various color and size. In female clones, oocytes appeared in late August and early September; they increased in size steadily in fall, and peaked in early December in both years. In male clones, spermatogenesis was synchronous with the female oogenic cycle, and motile sperm were observed in most testes in early December. Spawned gonads were found in both sexes from late November to early December. Spawning was induced in the laboratory in early December, and external fertilization was followed by development of free-swimming larvae. Gametogenesis and spawning are correlated with seasonal increases in seawater temperature and phytoplankton abundance, and we discuss and compare timing of sexual reproduction in this corallimorpharian to those in several actiniarian and scleractinian species.     

Population trends of Steller sea lions (Eumetopias jubatus) with respect to remote sensing measures of chlorophyll-a in critical habitat

The recovery plan for Steller sea lions (SSL; Eumetopias jubatus) suggests critical habitat should be enhanced to incorporate the spatio-temporal variation in dynamic oceanographic features that influence the prey and survival of SSL. It is necessary, therefore, to determine which features affect SSL. Demographics for sub-regions of the endangered, western stock of SSL were examined with respect to corresponding average, maximum, and variance of chlorophyll-a data (SeaWIFS), a proxy for primary productivity. Overall, SSL trends (2000–2008) and pup productivity (1999–2009) were related to maximum values of chl-a in critical habitat. Additionally, conditions in critical habitat appeared worse in areas of decline (i.e., dispersed patterns of chl-a hotspots and greater distances from SSL sites to productive areas). Although there may be a low feasibility of mitigating the effects of dynamic features on the recovery of SSL, the interactive effects of primary productivity and other stressors should be investigated for safeguarding their prey.     

Effects of predators on sea urchin density and habitat use in a southern California kelp forest

Grazing sea urchins can reduce kelp abundance and therefore strongly affect kelp forest community structure. Despite the ecological importance of sea urchins, direct field studies on the role that urchin predators play in shaping urchin populations are rare for southern California. We conducted surveys and manipulative experiments within kelp forests near San Diego, CA, (32–51′28″N, 117–16′00″W) from 2006 to 2009 to determine whether predators such as sheephead (Semicossyphus pulcher) and spiny lobsters (Panulirus interruptus) may be linked to purple urchin (Strongylocentrotus purpuratus) and red urchin (Strongylocentrotus franciscanus) distribution and habitat use, as well as purple urchin density-dependent mortality. Purple urchins were less dense and more cryptic inside a local marine protected area (MPA) that contained high predator abundance than in nearby heavily fished areas, whereas red urchins rarely were found outside the MPA. Urchin proportional mortality was inversely density dependent during the day when sheephead were active, despite fish aggregations in plots of high urchin density, but was density independent during the night when lobsters were active. Urchin mortality was reduced under understory algal cover during the day, but not during the night. Examining whether urchin mortality from predation is density dependent and how habitat complexity influences this relationship is imperative because behavioral changes and increases in urchin populations can have vast ecological and economic consequences in kelp forest communities.     

Expansion of a central California kelp forest following the mass mortality of sea urchins

The mass mortality by disease of a localized population of sea urchins, Strongylocencrotus franciscanus, on the seaward side of a kelp forest was followed by the rapid seaward expansion of 4 species of brown algae, Macrocystis pyrifera, Laminaria dentigera, Pterygophora california, and to a lesser extent, Nereocystis leutkeana. One other brown alga, Cystoseira osmundacea, failed to become established in the newly available area. Competition among M. pyrifera, L. dentigera, P. californica, and N. Leutkeana apparently was severe, and within 1 year after the demise of the sea urchins, M. pyrifera formed a dense, nearly monospecific stand. Experimental removal of M. pyrifera demonstrated that the canopy of these plants limited light penetration to levels below that necessary for the growth and survival of other brown and red algae.     

The effect of a natural environmental disturbance on maternal investment and pup behavior in the California sea lion

Summary Observed changes in maternal investment due to an environmentally induced decrease in food supply (the 1983 El Ni?o-Southern Oscillation) are compared witha priori predictions for the California sea lion (Zalophus californianus). Changes in behavior, growth and mortality of off-spring were also examined. Data collected in the first two months postpartum for the years before (PRE), during (EN), and the two years after (POST1 and POST2) the 1983 El Ni?o indicate that females initiated postpartum feeding trips earlier during the food shortage, and spent more time away on individual feeding trips in both the El Ni?o year and the year after. Perinatal sex ratios (♀:♂) in the years PRE, EN, POST1 and POST2 were 1:1, 1.4:1, 1.1:1 and 1:1.4, respectively. Fewer copulations were observed during the El Ni?o year, but this difference was not statistically significant. Pups spent less time suckling in the food shortage year and the year following, but attempted to sneak suckle more. Pups were less active and played on land less in the El Ni?o and following year. Finally, maternal investment as measured by milk intake of offspring was decreased, pups grew more slowly, and suffered increased mortality during the food shortage year. Despite expected sex differences in maternal investment and pup behavior in response to food shortage, there were no sex-biased differences in response in either females or pups. As expected, the food shortage did not affect adult males since they migrate north during the non-breeding season where the environmental perturbation was less severe.     

Cooperative hunting in lions: the role of the individual

Summary Individually identified lions (Panthera leo) were observed on the open, semi-arid plains in Namibia. Data from 486 coordinated group hunts were analysed to assess cooperation and individual variation in hunting tactics. Group hunts generally involved a formation whereby some lionesses (wings) circled prey while others (centres) waited for prey to move towards them. Those lionesses that occupied wing stalking roles frequently initiated an attack on the prey, while lionesses in centre roles moved relatively small distances and most often captured prey in flight from other lionesses. Each lioness in a given pride repeatedly occupied the same position in a hunting formation. Hunts where most lionesses present occupied their preferred positions had a high probability of success. Individual hunting behaviour was not inflexible, however, but varied according to different group compositions and to variations in the behaviour of other individuals present. The role of cooperative hunting and its apparent advantages within the semi-arid environment of Etosha National Park, Namibia, are discussed.     

Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

Estimating Sustainable Bycatch Rates for California Sea Lion Populations in the Gulf of California

Abstract:  Commercial and subsistence fisheries pressure is increasing in the Gulf of California, Mexico. One consequence often associated with high levels of fishing pressure is an increase in bycatch of marine mammals and birds. Fisheries bycatch has contributed to declines in several pinniped species and may be affecting the California sea lion ( Zalophus californianus ) population in the Gulf of California. We used data on fisheries and sea lion entanglement in gill nets to estimate current fishing pressure and fishing rates under which viable sea lion populations could be sustained at 11 breeding sites in the Gulf of California. We used 3 models to estimate sustainable bycatch rates: a simple population-growth model, a demographic model, and an estimate of the potential biological removal. All models were based on life history and census data collected for sea lions in the Gulf of California. We estimated the current level of fishing pressure and the acceptable level of fishing required to maintain viable sea lion populations as the number of fishing days (1 fisher/boat setting and retrieving 1 day's worth of nets) per year. Estimates of current fishing pressure ranged from 101 (0–405) fishing days around the Los Machos breeding site to 1887 (842–3140) around the Los Islotes rookery. To maintain viable sea lion populations at each site, the current level of fishing permissible could be augmented at some sites and should be reduced at other sites. For example, the area around San Esteban could support up to 1428 (935–2337) additional fishing days, whereas fishing around Lobos should be reduced by at least 165 days (107–268). Our results provide conservation practitioners with site-specific guidelines for maintaining sustainable sea lion populations and provide a method to estimate fishing pressure and sustainable bycatch rates that could be used for other marine mammals and birds .     

Abalones and sea urchins in an area inhabited by sea otters

Abalones (Haliotis spp.) and sea urchins (Strongylocentrotus spp.) are part of the subtidal fauna in the kelp bed off Hopkins Marine Station, Pacific Grove, California, USA, a protected marine reserve. Although these animals have been preyed upon by sea otters for over 10 years, their densities are gubstantial (Haliotis spp.: 0.21/m²; Strongylocentrotus spp.: 0.22/m²), and two species, H. rufescens (Swainson) and S. purpuratus (Stimpson), have wide size distributions indicating broad age ranges. Most of the adult abalones and sea urchins occur clustered deep in crevices, either as a direct result of sea-otter predation or because ample drift of algal food reduces foraging activities. Abalones tend to occur in larger crevices than sea urchins, and competition for suitable crevice space may exist among these large, invertebrate, algal-drift feeders.     

Growth and mortality of subtidal red sea urchins (Strongylocentrotus franciscanus) at San Nicolas Island,California, USA: problems with models

Red sea urchins, Strongylocentrotus franciscanus, were tagged with tetracycline in 1990 at subtidal sites off San Nicolas Island, California, USA. After one year in the field, the sea urchins were collected and growth increments were measured based on tetracycline marks, which indicated initial slow growth, a maximum rate, and finally a prolonged period of very slow growth. Small red sea urchins (4 cm diam) were estimated to be 3 to 4 yr old, which is much older than has previously been reported. It is estimated that about 12 yr would be required to attain 10 cm diam. Survival has previously been modeled assuming a constant rate. If the population of red sea urchins is assumed to be stable and stationary, annual survival rate was between 71 and 77% yr^-1. Census data for the two years of study have permitted annual survival to be estimated without assuming stable and stationary population structure. Under these conditions, annual survival rate was between 79 and 86% yr^-1. Analysis of transitions in the size distributions from 1990 to 1991 suggested that annual survival may have been sizespecific: 91% yr^-1 for individuals 1.1 to 4.0 cm diam, 82% yr^-1 for individuals 4.1 to 7.0 cm diam, and 63% yr^-1 for those of 7.1 to 10.0 cm diam. An alternative explanation to size-specific survival in our study is sizespecific immigration.