Age-related multi-year associations in female humpback whales (Megaptera novaeangliae)

Analyses of social structures in baleen whales are rare, and so far, they are thought to consist of mostly short and unstable associations. We investigated the association patterns of individual humpback whales from a summer feeding aggregation in the Gulf of St. Lawrence from 1997 to 2005. Photo-identified animals were sexed using genetic methods and were grouped into five categories: juvenile males/females, mature males and lactating/non-lactating females. We calculated half-weight association indices within and between the groups and found that 45% of the observation showed single animals and another 45% small groups (two to three) consisting mainly of mature animals besides lactating females. Using permutation tests, we found evidence for long-term associations between mature males and non-lactating females as well as among non-lactating females. Standardised lagged association rates revealed that these male–female groups disassociated quickly over about 2 weeks, whereas associations increased again towards the beginning of the breeding season. Non-lactating females of similar age engaged in multi-seasonal stable pairs for up to six consecutive feeding seasons; no mature male–female association was observed in consecutive years. The females with the most stable and long-term associations also had the highest reproductive output. While the risk of predation could not explain these long-term bonds, feeding cooperation seemed the most plausible explanation for group forming behaviour during the summer months.     

Sounds of two entrapped humpback whales (Megaptera novaeangliae) in Newfoundland

A wide variety of sounds were recorded from two entrapped humpback whales (Megaptera novaeangliae), a female and a male, in Newfoundland, Canada, during the summers of 1975 and 1976. The sounds included pulsed moans, moans, yups, cries, chirps and clicks. Comparisons are made between the sounds of the two whales (male and female). The most frequently produced sounds, pulsed moans and moans, are compared with similar sounds that compose part of the winter song. The sounds from the New-foundland whales were not put together into songs.     

Effect of stage-specific vital rates on population growth rates and effective population sizes in an endangered iteroparous plant

Effective population size (N(e)) determines the strength of genetic drift and can influence the level of genetic diversity a population can maintain. Assessing how changes in demographic rates associated with environmental variables and management actions affect N(e) thus can be crucial to the conservation of endangered species. Calculation of N(e) through demographic models makes it possible to use elasticity analyses to study this issue. The elasticity of N(e) to a given vital rate is the proportional change in N(e) associated with a proportional increase in that vital rate. In addition, demographic models can be used to study N(e) and population growth rate (λ) simultaneously. Simultaneous examination is important because some vital rates differ diametrically in their associations with λ and N(e). For example, in some cases increasing these vital rates increases λ and decreases N(e). We used elasticity analysis to study the effect of stage-specific survival and flowering rates on N(e), annual effective population size (N(a)), and λ in seven populations of the endangered plant Austrian dragonhead (Dracocephalum austriacum). In populations with λ ≥ 1, the elasticities of N(e) and N(a) were similar to those of λ. Survival rates of adults were associated with greater elasticities than survival rates of juveniles, flowering rates, or fecundity. In populations with λ < 1, N(e) and N(a) exhibited greater elasticities to juvenile than to adult vital rates. These patterns are similar to those observed in other species with similar life histories. We did not observe contrasting effects of any vital rate on λ and N(e); thus, management actions that increase the λ of populations of Austrian dragonhead will not increase genetic drift. Our results show that elasticity analyses of N(e) and N(a) can complement elasticity analysis of λ. Moreover, such analyses do not require more data than standard matrix models of population dynamics.     

Extinction-effective population index: incorporating life-history variations in population viability analysis

Viability status of populations is a commonly used measure for decision-making in the management of populations. One of the challenges faced by managers is the need to consistently allocate management effort among populations. This allocation should in part be based on comparison of extinction risks among populations. Unfortunately, common criteria that use minimum viable population size or count-based population viability analysis (PVA) often do not provide results that are comparable among populations, primarily because they lack consistency in determining population size measures and threshold levels of population size (e.g., minimum viable population size and quasi-extinction threshold). Here I introduce a new index called the "extinction-effective population index," which accounts for differential effects of demographic stochasticity among organisms with different life-history strategies and among individuals in different life stages. This index is expected to become a new way of determining minimum viable population size criteria and also complement the count-based PVA. The index accounts for the difference in life-history strategies of organisms, which are modeled using matrix population models. The extinction-effective population index, sensitivity, and elasticity are demonstrated in three species of Pacific salmonids. The interpretation of the index is also provided by comparing them with existing demographic indices. Finally, a measure of life-history-specific effect of demographic stochasticity is derived.     

Population structure of humpback whales in the western and central South Pacific Ocean as determined by vocal exchange among populations

For cetaceans, population structure is traditionally determined by molecular genetics or photographically identified individuals. Acoustic data, however, has provided information on movement and population structure with less effort and cost than traditional methods in an array of taxa. Male humpback whales (Megaptera novaeangliae) produce a continually evolving vocal sexual display, or song, that is similar among all males in a population. The rapid cultural transmission (the transfer of information or behavior between conspecifics through social learning) of different versions of this display between distinct but interconnected populations in the western and central South Pacific region presents a unique way to investigate population structure based on the movement dynamics of a song (acoustic) display. Using 11 years of data, we investigated an acoustically based population structure for the region by comparing stereotyped song sequences among populations and years. We used the Levenshtein distance technique to group previously defined populations into (vocally based) clusters based on the overall similarity of their song display in space and time. We identified the following distinct vocal clusters: western cluster, 1 population off eastern Australia; central cluster, populations around New Caledonia, Tonga, and American Samoa; and eastern region, either a single cluster or 2 clusters, one around the Cook Islands and the other off French Polynesia. These results are consistent with the hypothesis that each breeding aggregation represents a distinct population (each occupied a single, terminal node) in a metapopulation, similar to the current understanding of population structure based on genetic and photo‐identification studies. However, the central vocal cluster had higher levels of song‐sharing among populations than the other clusters, indicating that levels of vocal connectivity varied within the region. Our results demonstrate the utility and value of using culturally transmitted vocal patterns as a way of defining connectivity to infer population structure. We suggest vocal patterns be incorporated by the International Whaling Commission in conjunction with traditional methods in the assessment of structure.     

Responses of wintering humpback whales (Megaptera novaeangliae) to playback of recordings of winter and summer vocalizations and of synthetic sound

Summary Three natural sounds and one synthetic sound were played back to humpback whales during their 1985 and 1986 winter residency in Hawaiian waters. Playback was conducted from a vessel positioned within visual range of an elevated shorestation equipped with a high-precision surveyor's theodolite, used to determine the positions and movements of observed whale and of the playback vessel. A playback session consisted of 20 min of pretest observation with the vessel in place and underwater speaker deployed, followed by a 20-min test phase during which sound, or a blank tape control, was introduced. A total of 143 playback sessions, involving a total of 338 pods (a single whale or a group of whales), were completed over the two winter seasons. The major response observed during playback was a rapid approach to the playback vessel, characterized in some cases by velocities up to 9 km/h and approaches to within 50 m or less. Whales approaching were mainly singletons and, secondly, apparent adult pairs. No cow-calf pair ever approached. The approach was selective: 21.6% of targeted pods approached in response to a feeding sound recorded in summer feeding grounds in Alaska; 8.3% approached in response to social sounds recorded in the Hawaiian winter grounds in the presence of large surface-active pods; 3.4% responded to playback of winter song; and 4.1% responded to playback of synthetic sound. There were no approach responses to the blank tape control. Singing whales have been identified as males by many researchers. Data from Alaska suggested that the feeding sound was produced by a female; data from Hawaii suggests that the social sounds were produced by males. The different rates of response were attributed to the behavior of sexually active males seeking to affiliate with sexually mature females. Although a female may be present in pods producing social sounds, the presence of multiple males exhibiting aggression may inhibit the approach of other males. Song did not serve as an attractant for females, as measured by direct approach, but may still serve as a basis for female choice.     

Population size affects vital rates but not population growth rate of a perennial plant

Negative effects of habitat fragmentation on individual performance have been widely documented, but relatively little is known about how simultaneous effects on multiple vital rates translate into effects on population viability in long-lived species. In this study, we examined relationships between population size, individual growth, survival and reproduction, and population growth rate in the perennial plant Phyteuma spicatum. Population size positively affected the growth of seedlings, the survival of juveniles, the proportion of adults flowering, and potential seed production. Analyses with integral projection models, however, showed no relationship between population size and population growth rate. This was due to the fact that herbivores and pathogens eliminated the relationship between population size and seed production, and that population growth rate was not sensitive to changes in the vital rates that varied with population size. We conclude that effects of population size on vital rates must not translate into effects on population growth rate, and that populations of long-lived organisms may partly be able to buffer negative effects of small population size on vital rates that have a relatively small influence on population growth rate. Our study illustrates that we need to be cautious when assessing the consequences of habitat fragmentation for population viability based on effects on only one or a few vital rates.     

Translating effects of inbreeding depression on component vital rates to overall population growth in endangered bighorn sheep

Evidence of inbreeding depression is commonly detected from the fitness traits of animals, yet its effects on population growth rates of endangered species are rarely assessed. We examined whether inbreeding depression was affecting Sierra Nevada bighorn sheep (Ovis canadensis sierrae), a subspecies listed as endangered under the U.S. Endangered Species Act. Our objectives were to characterize genetic variation in this subspecies; test whether inbreeding depression affects bighorn sheep vital rates (adult survival and female fecundity); evaluate whether inbreeding depression may limit subspecies recovery; and examine the potential for genetic management to increase population growth rates. Genetic variation in 4 populations of Sierra Nevada bighorn sheep was among the lowest reported for any wild bighorn sheep population, and our results suggest that inbreeding depression has reduced adult female fecundity. Despite this population sizes and growth rates predicted from matrix-based projection models demonstrated that inbreeding depression would not substantially inhibit the recovery of Sierra Nevada bighorn sheep populations in the next approximately 8 bighorn sheep generations (48 years). Furthermore, simulations of genetic rescue within the subspecies did not suggest that such activities would appreciably increase population sizes or growth rates during the period we modeled (10 bighorn sheep generations, 60 years). Only simulations that augmented the Mono Basin population with genetic variation from other subspecies, which is not currently a management option, predicted significant increases in population size. Although we recommend that recovery activities should minimize future losses of genetic variation, genetic effects within these endangered populations-either negative (inbreeding depression) or positive (within subspecies genetic rescue)-appear unlikely to dramatically compromise or stimulate short-term conservation efforts. The distinction between detecting the effects of inbreeding depression on a component vital rate (e.g., fecundity) and the effects of inbreeding depression on population growth underscores the importance of quantifying inbreeding costs relative to population dynamics to effectively manage endangered populations.     

Assessing the effects of disease and bleaching on Florida Keys corals by fitting population models to data

Coral diseases have increased in frequency over the past few decades and have important influences on the structure and composition of coral reef communities. However, there is limited information on the etiologies of many coral diseases, and pathways through which coral diseases are acquired and transmitted are still in question. Furthermore, it is difficult to assess the impacts of disease on coral populations because outbreaks often co-occur with temperature-induced bleaching and anthropogenic stressors. We developed spatially explicit population models of coral disease and bleaching dynamics to quantify the impact of six common diseases on Florida Keys corals, including aspergillosis, dark spots, white band, white plague, white patch, and Caribbean yellow band. Models were fit to an 8-year data set of coral abundance, disease prevalence, and bleaching prevalence. Model selection was used to assess alternative pathways for disease transmission, and the influence of environmental stressors, including sea temperature and human population density, on disease prevalence and coral mortality. Classic disease transmission from contagious to susceptible colonies provided the best-fit model only for aspergillosis. For other diseases, external disease forcing, such as through a vector or directly from pathogens in the environment, provided the best fit to observed data. Estimates of disease reproductive ratio values (R₀) were less than one for each disease, indicating coral colonies were below densities required for diseases to become established through contagious spread alone. Incidences of white band and white patch disease were associated with greater susceptibility or slower recovery of bleached colonies, and no disease outbreaks were associated with periods of elevated sea temperatures alone. Projections of best-fit models indicated that, atleast during the period of this study, disease and bleaching did not have substantial impacts on populations and impaired rates of population growth appeared to be attributable to other stressors. By applying epidemiological models to field data, our study gives qualitative insights into the dynamics of coral diseases, relative stressor impacts, and directions for future research.     

Temperature effects on vital rates of different life stages and implications for population growth of Baltic sprat

Baltic sprat (Sprattus sprattus balticus S.) is a key species in the pelagic ecosystem of the Baltic Sea. Most stocks of small pelagic species are characterized by natural, fishery-independent fluctuations, which make it difficult to predict stock development. Baltic sprat recruitment is highly variable, which can partly be related to climate-driven variability in hydrographic conditions. Results from experimental studies and field observations demonstrate that a number of important life history traits of sprat are affected by temperature, especially the survival and growth of early life stages. Projected climate-driven warming may impact important processes affecting various life stages of sprat, from survival and development during the egg and larval phases to the reproductive output of adults. This study presents a stage-based matrix model approach to simulate sprat population dynamics in relation to different climate change scenarios. Data obtained from experimental studies and field observations were used to estimate and incorporate stage-specific growth and survival rates into the model. Model-based estimates of population growth rate were affected most by changes in the transition probability of the feeding larval stage at all temperatures (+0, +2, +4, +6?°C). The maximum increase in population growth rate was expected when ambient temperature was elevated by 4?°C. Coupling our stage-based model and more complex, biophysical individual-based models may reveal the processes driving these expected climate-driven changes in Baltic Sea sprat population dynamics.     

Age structure,growth rates,movement patterns and feeding in an estuarine population of the cardinalfish Apogon rueppellii

The biology of a population of the cardinalfish Apogon rueppellii has been studied over several years (1977–1983) in the Swan Estuary in south-western Australia, using ramples collected monthly from the shallows by beach seine and from various depths by otter trawl. While the life cycle of this species typically lasts for one year, at the end of which time the mean length is 50 to 60 mm, some individuals survive for a further year and attain lengths up to 104 mm. A. rueppellii shows a marked tendency to move offshore into deeper water during the winter months. This tendency is more pronounced in the 1+ than in the 0+ year class and in larger than smaller 0+ individuals. An inshore movement of A. rueppellii in the spring is followed by spawning and by oral brooding by the males, which leads to the recruitment of large numbers of a new 0+ year class on to the banks during the summer. The offshore movement is correlated with changes in salinity and temperature. The larger catches taken by otter trawl during the day than at night indicate that A. rueppellii exhibits a diel pattern of activity. Mean fecundity ranged from 70 in the 45 to 49 mm size class to 345 in the 90 to 94 mm size class. Measurements of fecundity and the number of oral-brooded eggs demonstrated that the majority of the eggs released by the female are collected and incubated by the males. Copepods are ingested in relatively greater amounts by small than by large A. rueppellii, whereas the reverse situation occurs with larger crustaceans, polychaetes and small fish. The presence of greater amounts of copepods in the diet during the day and of amphipods at night probably reflects the diel activity patterns of the prey.     

Blubber fatty acids of minke whales: stratification,population identification and relation to diet

We determined the fatty acids in the blubber of 10 minke whales (Balaenoptera acutorostrata) from the Norwegian Sea and 12 minke whales from the North Sea. The composition of the fatty acids differed between the inner and outer blubber layer. The saturated, the monounsaturated with 20 and 22 carbon atoms and the polyunsaturated fatty acids had higher relative amounts in the inner than in the outer blubber, and the short-chain monounsaturated acids, i.e. with 14, 16 and 18 carbon atoms, had the highest relative amounts in the outer layer. The change in relative concentration was steeper in the outer parts of the blubber than in the inner parts. Whales from the two areas had different fatty-acid profiles in both inner and outer blubber layer, although these differences were smaller than the difference between inner and outer layer. The difference between the whales from the two areas corresponds to the management stocks defined by the International Whaling Commission. Determination of the fatty-acid profile may be used for identification of whale populations. The composition of the fatty acids in the blubber differed from that of the main prey of the whales: sand eel for the North Sea whales and herring for the Norwegian Sea whales. These differences between prey and blubber fatty acids were larger for the outer blubber layer than for the inner.     

Lack of population subdivision among the minke whales (Balaenoptera acutorostrata) from Icelandic and Norwegian waters based on mitochondrial DNA sequences

The minke whale (Balaenoptera acutorostrata) is subject to commercial whaling, but stock identification and assessment are still uncertain. Mitochondrial DNA (mtDNA) sequences were determined to examine the population structure of minke whales from the central and northeastern parts of the North Atlantic, as well as the Antarctic regions IV and V. The analyses include 345 nucleotide positions of the control region of 110 individuals, and 250 nucleotide positions of the NADH dehydrogenase subunit 2 gene for a representative selection of North Atlantic minke whales. Maximum parsimony analyses and sequence divergence calculations did not reveal any genetic differentiation between individuals from the central and northeastern parts of the North Atlantic. These results do not support the International Whaling Commission's separation of minke whales in this area into different management units, and they are in conflict with previously reported results from allozyme analyses. Comparison of minke whale control region sequences showed that the sequence diversity of North Atlantic minke whales is substantially lower (0.0065) than that of Antarctic minke whales (0.0166), and clearly demonstrated that individuals from these two areas represent genetically distinct populations.     

Growth and average growth rates of tanner crab zoeae collected from the plankton

Growth of the two zoeal stages of Tanner crabs, Chionoecetes bairdi and C. opilio, was estimated by changes in total dry weight of individuals sampled from plankton of the southeastern Bering Sea during the springs of 1977, 1978, 1980 and 1981. Since the age of larvae within a stage cannot be determined, approximate beginning and end weights of each stage were used. Epidermal retraction, conspicuous under low magnification of a dissecting microscope, was used to identify larvae presumed to be in advanced stages of growth (biomass) for their respective larval stage. This assumption was corroborated by holding retracted larvae that subsequently molted on board ship, by comparison of dry weight measurements from extensive field sampling, and by the low proportion of larvae showing such conspicuous retraction. An exponential growth model was used to calculate the average daily growth rate based on estimates of stage duration in the field. The adequacy of this model for describing the average rate, rather than the pattern, of growth is discussed. Carbon-specific rates of respiration from discrete measurements and of growth averaged for each zoeal stage indicate average net growth efficiencies comparable to laboratory results for other species. The method used to estimate growth should be applicable to other larval decapods and offers the advantage of estimating this parameter from data gathered from natural populations.Contribution No. 652 of the School of Fisheries, University of Washington     

Estimation of population density by spatially explicit capture-recapture analysis of data from area searches

The recent development of capture-recapture methods for estimating animal population density has focused on passive detection using devices such as traps or automatic cameras. Some species lend themselves more to active searching: a polygonal plot may be searched repeatedly and the locations of detected individuals recorded, or a plot may be searched just once and multiple cues (feces or other sign) identified as belonging to particular individuals. This report presents new likelihood-based spatially explicit capture-recapture (SECR) methods for such data. The methods are shown to be at least as robust in simulations as an equivalent Bayesian analysis, and to have negligible bias and near-nominal confidence interval coverage with parameter values from a lizard data set. It is recommended on the basis of simulation that plots for SECR should be at least as large as the home range of the target species. The R package "secr" may be used to fit the models. The likelihood-based implementation extends the spatially explicit analyses available for search data to include binary data (animal detected or not detected on each occasion) or count data (multiple detections per occasion) from multiple irregular polygons, with or without dependence among polygons. It is also shown how the method may be adapted for detections along a linear transect.     

Local population dynamics of an invasive tree species with a complex life-history cycle: A stochastic matrix model

Biological invasions are widely accepted as having a major impact on ecosystem functioning worldwide, giving urgency to a better understanding of the factors that control their spread. Modelling tools have been developed for this purpose but are often discrete-space, discrete-time spatial-mechanistic models that adopt a computer simulation approach and resist mathematical analysis. We constructed a simple demographic matrix model to explore the local population dynamics of an invasive species with a complex life history and whose invasive success depends on resource availability, which occurs stochastically. As a case study we focused on the American black cherry (Prunus serotina Ehrh.), a gap-dependent tree able both to constitute a long-living seedling bank under unfavourable light conditions and to resprout vigorously once cut-down, which is invading European temperate forests. The model used was a stage-classified matrix population model (i.e., Lefkovitch matrix), integrating environmental stochasticity. Stochastic matrix projection analysis was combined with elasticity analysis and stochastic simulations to search for the species’ ‘Achille heel’. As expected, the population growth rate (i.e., Lyapunov exponent), which measures the risk of P. serotina invasion at the stand scale, increased with light frequency. There was a critical value above which the population of P. serotina explodes and below which it locally goes extinct. The resprouting capacity usually speed up the invasion but appeared to play a minor role. The mean duration of stand invasion was measured and important life stage transitions that mostly contribute to the local stochastic growth rate were identified. Some relevant management implications are discussed and the interest of such models for the understanding of demographic characteristics of invasive species is stressed.     

Estimating age from recapture data: integrating incremental growth measures with ancillary data to infer age-at-length

Estimating the age of individuals in wild populations can be of fundamental importance for answering ecological questions, modeling population demographics, and managing exploited or threatened species. Significant effort has been devoted to determining age through the use of growth annuli, secondary physical characteristics related to age, and growth models. Many species, however, either do not exhibit physical characteristics useful for independent age validation or are too rare to justify sacrificing a large number of individuals to establish the relationship between size and age. Length-at-age models are well represented in the fisheries and other wildlife management literature. Many of these models overlook variation in growth rates of individuals and consider growth parameters as population parameters. More recent models have taken advantage of hierarchical structuring of parameters and Bayesian inference methods to allow for variation among individuals as functions of environmental covariates or individual-specific random effects. Here, we describe hierarchical models in which growth curves vary as individual-specific stochastic processes, and we show how these models can be fit using capture-recapture data for animals of unknown age along with data for animals of known age. We combine these independent data sources in a Bayesian analysis, distinguishing natural variation (among and within individuals) from measurement error. We illustrate using data for African dwarf crocodiles, comparing von Bertalanffy and logistic growth models. The analysis provides the means of predicting crocodile age, given a single measurement of head length. The von Bertalanffy was much better supported than the logistic growth model and predicted that dwarf crocodiles grow from 19.4 cm total length at birth to 32.9 cm in the first year and 45.3 cm by the end of their second year. Based on the minimum size of females observed with hatchlings, reproductive maturity was estimated to be at nine years. These size benchmarks are believed to represent thresholds for important demographic parameters; improved estimates of age, therefore, will increase the precision of population projection models. The modeling approach that we present can be applied to other species and offers significant advantages when multiple sources of data are available and traditional aging techniques are not practical.