The vulnerable Osprey breeding population of the Al Hoceima National Park,Morocco: present status and threats

In the Mediterranean, most areas belonging to the initial distribution range of the Osprey Pandion haliaetus have been lost and local populations have disappeared in recent decades because of persecution. Even though direct management actions have allowed local partial recovery, the Mediterranean population currently only holds a few tens of breeding pairs and is still exposed to local extinction risks. One of the last Mediterranean Osprey breeding areas lies along the North African coast between Morocco and Algeria. In this paper, we report new information on the Osprey population within the Al Hoceima National Park, Morocco. The status of the population for 2012 and 2013 is reported and compared with data collected during the period 1983–1990. A reduction in number of nests and breeding pairs was observed and a 35.7% decrease in the population size recorded. In addition, we discuss the main identified threats to Osprey habitats (e.g. dynamite and poison fishing) that affect the Osprey breeding population in this area. In this context, we stress the necessity for urgent measures to be adopted at the local scale for the protection of this vulnerable population in the light of a sound conservation strategy also at the scale of the Mediterranean.     

Density dependence,population persistence,and largely futile arguments

Wolda and Dennis (1993) suggest that no valid conclusions about population regulation can be drawn on the basis of statistical tests of density dependence in time series data of population abundance. They give some examples in which a population persists even if it is not regulated by a density-dependent process: a sequence of independent, identically distributed random variables, the numbers of the migrant moth Autographa gamma in Britain, annual rainfall data. We suggest that such time series data may show persistence because of a static constraint, which compels the numbers to remain within finite, positive limits, or to fit some prescribed distribution. But this mechanism can explain persistence in a biological population only when the population represents a sample from a regulated population (the case of A. gamma). We also comment on some suggestions made by Wolda and Dennis (1993) concerning the general value of statistical tests of density dependence, frequency of delayed versus non-delayed density dependence in natural populations, relative performance of different kinds of insect traps in sampling local populations, and the wider issue of how ecologists are likely to make progress in the study of population dynamics.     

Investigating patterns and processes of demographic variation: environmental correlates of pre-breeding survival in red-billed choughs Pyrrhocorax pyrrhocorax

1. Quantifying the pattern of temporal and spatial variation in demography, and identifying the factors that cause this variation, are essential steps towards understanding the structure and dynamics of any population. 2. One critical but understudied demographic rate is pre-breeding survival. We used long-term colour-ringing data to quantify temporal (among-year) and spatial (among-nest site) variation in pre-breeding survival in red-billed choughs (Pyrrhocorax pyrrhocorax) inhabiting Islay, Scotland, and identified environmental correlates of this variation. 3. Random-effects capture-mark-recapture models demonstrated substantial temporal and spatial process variance in first-year survival; survival from fledging to age 1 year varied markedly among choughs fledged in different years and fledged from different nest sites. Spatial variance exceeded temporal variance across choughs fledged from well-studied nest sites. 4. The best-supported models of temporal variation suggested that first-year survival was higher in years following high tipulid larvae abundance and when weather conditions favoured increased invertebrate productivity and/or availability to foraging choughs. These variables explained up to 80% of estimated temporal process variance. 5. The best-supported models of spatial variation suggested that first-year survival was higher in choughs fledged from nest sites that were further from exposed coasts and closer to flocking areas, and surrounded by better habitat and higher chough density. These variables explained up to 40% of estimated spatial process variance. 6. Importantly, spatio-temporal models indicated interactive effects of weather, tipulid abundance, local habitat and local chough density on first-year survival, suggesting that detrimental effects of poor weather and low tipulid abundance may be reduced in choughs fledged from nest sites surrounded by better foraging habitat and lower chough density. 7. These analyses demonstrate substantial temporal and small-scale spatial variation in pre-breeding survival, a key demographic rate, and indicate that this variation may reflect interactive effects of weather, prey abundance, habitat and geography. These patterns illustrate the value of holistic models of demographic variation, and indicate environmental factors that may limit the growth rate of Islay's protected chough population.     

Density-dependent population dynamics in clonal organisms: a modelling approach

1. Density dependence may act at several stages in an organisms life-cycle (e.g. on mortality, fecundity, etc.), but not all density-dependent processes necessarily regulate population size. In this paper I use a density manipulation experiment to determine the effects of density on the transition rates between different size classes of the clonal zoanthid Palythoa caesia Dana 1846. I then formulate a density-dependent matrix model of population dynamics of Palythoa , and perform a series of sensitivity analyses on the model to determine at what stage in the life-cycle regulation acts.
2. Seven of the 16 transition probabilities decreased with density, most of them being shrinkage (due to loss of tissue or fission) and stasis (the self–self transition) of medium and large colonies. The only probability to increase was for the stasis of large colonies. Recruitment was quadratically dependent on density, peaking at intermediate densities.
3. Equilibrium cover in the model was 84% and was reached in ≈40 years. To determine which density-dependent transitions were involved in population regulation, the strength of density dependence was varied in each independently. This sensitivity analysis showed that only changes in the probabilities of large colonies remaining large and producing medium colonies, were regulating.
4. These results suggest that regulation is primarily acting on fission of large colonies to produce intermediate-sized colonies, in combination with size specific growth rates. Fission rates decrease greatly with density, resulting in a greater proportion of large colonies at high densities and large colonies grow more slowly than small. Overall, this behaviour is very similar to that of clonal plants which have a phalanx type life history.     

Density dependence in resource exploitation: empirical test of Levins' metapopulation model

Levins' model of metapopulation dynamics is modified to incorporate variable degrees of density dependence in the per capita exploitation of resource patches. We demonstrate a simple means of testing for this density dependence in a sample of metapopulations, each at its equilibrium balance of local colonization to extinction. The fraction of habitable unoccupied patches equilibrates to a constant number under the null model of density independent colonization, and to a constant proportion under strong density dependence. We compare the null model to two density dependent alternatives, using data on exploitation of nest boxes by collared flycatchers Ficedula albicollis . The analysis shows how predicted trends in the equilibrium unoccupied fraction are similar for both spatial interference and net immigration. This needs to be recognized, since the null hypothesis of a constant unused resource applies also to the dynamics of consumable resources, where it is expressed in a constant stock of uneaten prey at the dynamic equilibrium of predators to prey.     

Cycles, stochasticity and density dependence in pink salmon population dynamics

Complex dynamics of animal populations often involve deterministic and stochastic components. A fascinating example is the variation in magnitude of 2-year cycles in abundances of pink salmon (Oncorhynchus gorbuscha) stocks along the North Pacific rim. Pink salmon have a 2-year anadromous and semelparous life cycle, resulting in odd- and even-year lineages that occupy the same habitats but are reproductively isolated in time. One lineage is often much more abundant than the other in a given river, and there are phase switches in dominance between odd- and even-year lines. In some regions, the weak line is absent and in others both lines are abundant. Our analysis of 33 stocks indicates that these patterns probably result from stochastic perturbations of damped oscillations owing to density-dependent mortality caused by interactions between lineages. Possible mechanisms are cannibalism, disease transmission, food depletion and habitat degradation by which one lineage affects the other, although no mechanism has been well-studied. Our results provide comprehensive empirical estimates of lagged density-dependent mortality in salmon populations and suggest that a combination of stochasticity and density dependence drives cyclical dynamics of pink salmon stocks.     

Population regulation of territorial species: both site dependence and interference mechanisms matter

Spatial patterns of site occupancy are commonly driven by habitat heterogeneity and are thought to shape population dynamics through a site-dependent regulatory mechanism. When examining this, however, most studies have only focused on a single vital rate (reproduction), and little is known about how space effectively contributes to the regulation of population dynamics. We investigated the underlying mechanisms driving density-dependent processes in vital rates in a Mauritius kestrel population where almost every individual was monitored. Different mechanisms acted on different vital rates, with breeding success regulated by site dependence (differential use of space) and juvenile survival by interference (density-dependent competition for resources). Although territorial species are frequently assumed to be regulated through site dependence, we show that interference was the key regulatory mechanism in this population. Our integrated approach demonstrates that the presence of spatial processes regarding one trait does not mean that they necessarily play an important role in regulating population growth, and demonstrates the complexity of the regulatory process.     

Patterns of density‐dependent growth in juvenile stream‐dwelling salmonids

Literature data for 19 populations from 16 different studies of six species of stream‐dwelling salmonids were analysed to test the hypothesis that density‐dependent growth is stronger at low rather than at high population density. Fifteen of 19 populations showed evidence of a significant decrease in growth rate with increasing density. In 11 of these 15 populations, the pattern of density‐dependent growth was better described by a negative power curve than by a linear regression ( i.e . Akaike Information Criterion, AIC _linear −  AIC _power > 2), whereas only one population was better described by a linear regression than by a negative power curve; three populations were adequately described by both models ( AIC  < 2). In 10 of the 11 populations that were best described by a negative power curve, most of the decrease in growth rate occurred at population densities <1 fish m⁻², when space limitation is unlikely. This analysis provides broad support for the hypothesis that density‐dependent growth in stream salmonids occurs primarily at low population densities, probably due to exploitative competition.     

Density dependence in cereal aphid populations

Long sequences of data on the incidence of cereal aphids from five European countries were analysed for evidence of density dependent processes occurring between years. Using a randomisation test (Pollard, Lakhani & Rotheray, 1987), density dependence was revealed in all (16) population censuses of Metopolo-phium dirhodum , 60% of Rhopalosiphum padi censuses (10 of 17) but only 25% of Sitobion avenae population censuses (4 of 16). Correcting for density independent effects of weather revealed the existence of significant direct density dependence in some populations censuses where it was previously undetected. The implications of density dependence in cereal aphid populations are considered.     

Density dependence depends on scale; at larval resource patch and at whole population

A previous study (Tuda and Shimada, 1993) has shown that the equilibrium population size of the azuki bean beetle was lower at 32°C than at 30°C and that this difference was due to a reduced maximum population size of emerged progeny through inside-bean process. In this paper, these results were analyzed further on the scale of the individual bean where interaction among larvae took place. Per-bean numbers of deposited eggs, hatched eggs, and emerged adults have been recorded at seven different parental densities under the two temperature conditions. Three individual-bean-scale process hypotheses that may explain the reduced maximum emergence density on the whole population scale are suggested: (1) a lower maximum emergence per bean at 32°C than at 30°C, if the bean scale and the wholepopulation scale share the same density-dependent pattern in adult emergence, (2) a limited range of hatched egg number per bean at 32°C, resulting from the adult oviposition process outside beans, and (3) different patterns of density-dependent emergence between the two different scales. This study showed that the inside-bean pattern of responses on the bean scale was a simple saturated curve at 30°C, but one with a discontinuous decline at higher hatched egg densities at 32°C. On the contrary, during outside-bean process, the peak number of hatched eggs decreased on this scale as observed on the wholepopulation scale. I discuss why the extracted factor of inside-bean process on the whole-population in the previous study could not be applied to the bean-scale pattern.     

Population dynamics of coral-reef fishes: Controversial concepts and hypotheses

Abstract Knowledge of processes that drive the local population dynamics of coral-reef fishes is important for managing reef fisheries, and for using these species as models for understanding the ecology of demersal marine fishes in general. However, the reef-fish literature is replete with poorly defined concepts and vague hypotheses regarding the issue of population dynamics. Dichotomous arguments, such as whether or not recruitment drives population dynamics, are misdirected because they fail to incorporate several important concepts. First, changes in local population size are driven by four demographic rates (birth, death, immigration and emigration), all of which must be studied to understand population dynamics. Second, all populations that persist do so because at least one of these demographic rates operates in a density-dependent way that is both sufficiently strong and appropriately time-lagged. Therefore, identifying the source(s) of direct density dependence is critical for understanding the limits to variation in population size (i.e. population regulation). Third, regulation does not imply a simple point equilibrium in population size; density dependence in populations of reef fishes is bound to lie within a field of stochastic variation, and thus be difficult to detect. Since its formal origin in 1981, the ‘recruitment limitation’ hypothesis for explaining local population dynamics in reef fishes has undergone ambiguous changes in definition that threaten its usefulness. ‘Recruitment, ‘originally defined as the appearance of newly settled fish on a reef, more recently is often measured months after settlement, thus confounding pre- and post-settlement processes. ‘Limitation, ‘ which originally referred to recruitment being so low as to preclude local populations from reaching densities where resources were limiting, is more recently defined as an absence of any form of density dependence after settlement. The most effective means of testing whether post-settlement mortality is in fact density-independent is to examine patterns of mortality directly, rather than indirectly by interpreting the shape of the relationship between initial recruit density and subsequent adult density within a cohort (the recruit-adult function). Understanding the population dynamics of coral-reef fishes will require a more equitable focus on all four demographic rates, be they density dependent or not, as well as greater attention to identifying sources of density dependence. Such a pluralistic focus necessitates integrated studies of both pre- and post-settlement processes conducted at multiple spatial and temporal scales. For example, recent evidence suggests that density-dependent pre-dation on new recruits that have settled among reefs at different densities may prove to be an important source of local population regulation, especially via the aggregative response of transient piscivores.