Some aspects of the biology of the red bass, Lutjanus bohar (Forsskal), from the Tigak Islands, Papua New Guinea

The growth rate, food, reproduction and mortality rates of the red bass, Lutjanus bohar , are described for fish associated with coral reefs in the Tigak Islands of Papua New Guinea. Spawning intensity by month was investigated by examination of gonad maturation stages. The length–at–maturity of female L. bohar was estimated from the relationship between changes in gonad index with increasing size. Female fish matured at about 45–0 cm fork length. The Petersen method was used to obtain age–at–length data from length frequency data of L. bohar captured by hook–and–line fishing. The growth of L. bohar was described by a von Bertalanffy growth curve with the parameters AT=0.27 and L_∞ = 81.7 cm. There was a positive correlation between water depth (D) and the length of L. bohar (L) that was described by L = 318–8+ 3.050 D . The mortality rates ( Z ) of L. bohar from shallow (≤30m) and deep (≤80m) water were estimated from length converted catch curves and were 1–181 and 0–64 respectively. The high Z value for the shallow water population of L. bohar was ascribed to the migration of larger specimens into deeper water. The deep water population of L. bohar remains unfished and the estimate of Z was believed to be equivalent to the natural mortality rate ( M ). An empirical method of calculating M gave an estimate of 0.59. Lutjanus bohar is a carnivorous piscivore that feeds throughout the water column.     

Fluctuating recruitment and variable life history of migratory brown trout, Salmo trutta L., in a small, unstable stream

The recruitment dynamics and life history of migratory brown trout, Sulmo trutta L., were investigated in a small Baltic coast stream subject to recurring drought. Spawning males consisted of both mature male parr (101–206 mm t.l. ) and migrant males (205–780 mm t.l. ). Spawning females were all migrants which delayed maturity until reaching a significantly greater size on average (424–805 mm t.l. ) than migrant males. Male: female ratios were very high in spawning aggregations (9–12 males: 1 female) with males representing up to five year-classes or more. Gametes from several generations of males per spawning event may be important for maintaining the genetic viability of this population with few female spawners per year. The amount of spawning was dependent on precipitation just prior to and during the spawning period since migrants could not enter the stream under drought conditions. Migrants did not overwinter in the stream.
Drought also caused variable fry mortality following emergence in early summer. Recruitment of 0+ parr in autumn varied from c . 175 to 3000 during 3 years. Smolts were relatively young (ages 1–2) and small (≥8 cm), and were significantly longer on average than sibling parr. Yet emigration of 1-year-olds was not related to 0+ parr size the previous autumn because of overlapping growth rates.
Persistence of the migratory brown trout in this unstable environment may be the consequence of (i) life history adaptation (e.g. short freshwater residence of both juveniles and spawners), and (ii) a complementary set of individual life histories where variation in age of migrant spawners and the occurrence of mature male parr result in a stable spawner population despite inconsistent recruitment of migrants to the sea.     

Distribution and biology of Alepocephalus rostvatus from the Mediterranean Sea

In the Catalan Sea (NW Mediterranean) A. rostratus was found on the bottom below 500 m and reached its maximum abundance and biomass at 1100–1300 m depth, being captured down to 2209 m. Mean fish size increased progressively with depth. Larger fish were predominantly females, and 50% of maturity was reached at 23–6 and 27–7 cm in males and females, respectively. Although a high percentage of the population was mature all year round, a peak was attained in autumn, coinciding with the slowest growth of the species. Size-at-age indicated a slow growth rate. The greatest observed age was 23 years, although most of individuals were 7–12 years old. Overall, A. rostratus can be considered as a typical K-strategist, showing a good adaptation to the oligotropic deepsea environment.     

Reproduction,growth and mortality of the exploited sillaginid,Sillago ciliata Cuvier, 1829

The goal of this study was to examine the age and size composition, growth, reproductive biology and mortality of Sillago ciliata Cuvier, 1829 in one of the largest estuarine commercial fisheries in south‐eastern Australia. The study also aimed to present a qualitative comparison of latitudinal variations in some of these characteristics along the eastern Australian coastline. The sampled population contained fish aged up to 10 years with a maximum size of 39.2 cm fork length (L_F), and was dominated by 1–5 year olds. Sexual divergence in both the age and size structure of the population was recorded. Female S. ciliata grew slightly faster and attained a greater maximum size (L_∞ = 33.79 cm L_F, k = 0.50 year^?1 and t₀ = ?0.57 years) than males (L_∞ = 29.73 cm L_F, k = 0.49 year^?1 and t₀ = ?0.67 years). Females also matured at a significantly larger size (19.13 cm) and older age (1.63 years) than males (size: 17.07 cm, age: 1.10 years). Reproductive activity was highest between September and March. There were no differences between males and females in terms of mortality rate; the estimated total population, natural and fishing mortality rates were Z = 0.64, M = 0.42 and F = 0.22, respectively. Although these mortality rates suggest that S. ciliata in the Clarence River are relatively resilient to current rates of exploitation, regular monitoring of their commercial and recreational catch as well as their population structure is recommended in order to maintain sustainable fisheries. Potential latitudinal shifts in the spawning period, age structure and growth of S. ciliata along eastern Australia were also revealed.     

The ecology of the Mediterranean stony coral Cladocora caespitosa (Linnaeus, 1767) in the Gulf of Trieste (northern Adriatic Sea): a 30-year long story

Cladocora caespitosa is an endemic coral of the Mediterranean Sea and an important carbonate bioconstructor that adds 3D complexity to the habitat, thus increasing marine biodiversity. Despite its important role in the ecosystem, the real status of the population along most of the Mediterranean coastline is still poorly investigated and very little is known about the resilience of the species. Using non-destructive visual surveys, colonies of C. caespitosa were investigated by SCUBA diving in 2013 and 2015 at seven sites of the northern Adriatic Sea (southern part of the Gulf of Trieste). Data about colony size, index of sphericity and corallite diameter were collected. Almost all biometrical parameters differed significantly among sampling sites, showing low occurrence of the larger size classes compared to the abundance of small-sized colonies. This pattern of distribution is typical of long-lived organisms. The positively skewed colony size distribution could be due to both a high mortality rate of small colonies unable to reach larger size classes, and to a high fragmentation rate of colonies, related to a strong hydrodynamic forces. The northern Adriatic population of C. caespitosa has previously been investigated by Schiller, who reported size and abundance data of colonies from one site, at a depth range of 2–5?m. We compared these data with our findings from the same sampling site, adding new information about the ecology of C. caespitosa. After a 30-year period, the comparison shows a change in the size distribution of colonies, with a decrease of the small class and an increase of the medium class of colonies. In view of these conclusions, further assessments are required in order to evaluate the trend of the northernmost C. caespitosa population in the Mediterranean Sea.     

Food habits, growth rates, and reproductive biology of grass snakes, Natrix natrix (Colubridae) in the Italian Alps

A five-year mark-recapture study at Sella Nevea, a montane (1100 m a.s.1.) site in the Carnic Alps, provided information on diets, growth rates, and reproductive output in an Italian population of the wide-ranging grass snake, Natrix natrix. Our snakes resembled a previously-studied population in lowland Sweden in terms of body size at sexual maturation in females (70 cm) and mean adult female body length (82 cm). However, growth rates were lower in our population, and sexual maturation was delayed (6–8 years, versus 4–5 years in Sweden), perhaps because of the cool climate and relatively brief growing period each year. Females produced a single clutch of 4–24 eggs in late July each year. Larger females produced larger clutches, but clutch size relative to maternal size was lower than in Swedish grass snakes. Hatchling sizes and Relative Clutch Masses (RCMs) did not shift with increasing female size. RCMs may provide a useful index of 'costs of reproduction' in this population, because females with high RCMs were very emaciated after oviposition, and hence may experience a greater risk of mortality, as well as a high energy expenditure. Prolonged incubation gave rise to longer, thinner hatchlings, but the low environmental temperatures at the study site may favour early hatching (and hence, result in a shorter fatter hatchling emerging from the egg, with more of its energy stores unused). Compared to sympatric viviparous snakes ( Coronella austriaca and Vipera berus ), the oviparous grass snakes can achieve a much higher reproductive output owing to a larger clutch size and more frequent reproduction (annual, rather than biennial or triennial). The abundant prey resource used by grass snakes (amphibians) may also enable them to recoup energy more rapidly after reproduction; dietary composition shifts ontogenetically in both sexes, with the largest prey (mice and adult toads) taken primarily by large female snakes.     

Reproduction, age, growth and mortality of axillary seabream, Pagellus acarne (Sparidae), from the Canarian archipelago

Axillary seabream Pagellus acarne (Risso 1826) caught off the Canary Islands from January 1991 to December 1994 were studied. The length range of the catches was between 11 and 31 cm, with a modal distribution between 17 and 21 cm. The overall ratio of males to females was 1:1.74. Males were observed up to a length of 24 cm. Hermaphrodites were recorded at lengths between 15 and 23 cm. The species was characterized by protandric hermaphroditism. The reproductive season extended from October to March, with a peak in spawning activity in December–January. The size at sexual maturity was 15.8 cm total length (2 years old) for males and 19.4 cm total length (3 years old) for females. The total length–total weight relationship for the entire population is described by the parameters a = 0.0068, and b = 3.2401. Otolith age readings showed that the population exploited consisted of 10 age groups (1–10 years), including a very high proportion of individuals between 1 and 4 years old. The von Bertalanffy growth parameters for all individuals were L_∞ = 32.98 cm, k = 0.22 years⁻¹, and t₀ = −0.87 years. Males grew comparatively slower than females. The instantaneous rates of mortality for all fish were Z = 0.96 years⁻¹, M = 0.30 years⁻¹, and F = 0.66 years⁻¹. The exploitation ratio and the length at first capture were, respectively, E = 0.69 and LC₅₀ = 16.1 cm. The stock is overexploited, therefore measures such as closed seasons or changes in fishing patterns would be desirable to safeguard the spawning stock and recruits.     

Forest responses to simulated elevated CO2 under alternate hypotheses of size‐ and age‐dependent mortality

Elevated atmospheric carbon dioxide (eCO₂) is predicted to increase growth rates of forest trees. The extent to which increased growth translates to changes in biomass is dependent on the turnover time of the carbon, and thus tree mortality rates. Size‐ or age‐dependent mortality combined with increased growth rates could result in either decreased carbon turnover from a speeding up of tree life cycles, or increased biomass from trees reaching larger sizes, respectively. However, most vegetation models currently lack any representation of size‐ or age‐dependent mortality and the effect of eCO₂ on changes in biomass and carbon turnover times is thus a major source of uncertainty in predictions of future vegetation dynamics. Using a reduced‐complexity form of the vegetation demographic model the Functionally Assembled Terrestrial Ecosystem Simulator to simulate an idealised tropical forest, we find increases in biomass despite reductions in carbon turnover time in both size‐ and age‐dependent mortality scenarios in response to a hypothetical eCO₂‐driven 25% increase in woody net primary productivity (wNPP). Carbon turnover times decreased by 9.6% in size‐dependent mortality scenarios due to a speeding up of tree life cycles, but also by 2.0% when mortality was age‐dependent, as larger crowns led to increased light competition. Increases in aboveground biomass (AGB) were much larger when mortality was age‐dependent (24.3%) compared with size‐dependent (13.4%) as trees reached larger sizes before death. In simulations with a constant background mortality rate, carbon turnover time decreased by 2.1% and AGB increased by 24.0%, however, absolute values of AGB and carbon turnover were higher than in either size‐ or age‐dependent mortality scenario. The extent to which AGB increases and carbon turnover decreases will thus depend on the mechanisms of large tree mortality: if increased size itself results in elevated mortality rates, then this could reduce by about half the increase in AGB relative to the increase in wNPP.     

Sexual and geographical variation in life history parameters of the shorthorn sculpin

A total of 293 shorthorn sculpins Myoxocephalus scorpius from Tromsø, northern Norway, were sampled between November 1998 and April 1999 to determine sex, total length, age, growth, maturity and mortality. Females grew to larger sizes ( L _∞=26·9 v. 18·5 cm), matured later (2 v. 1 year of age) at larger size (maturation length=16 v. 14 cm L _T), and had lower instantaneous mortality rates (0·93 v. 1·20 year⁻¹) than males. The life history parameters of shorthorn sculpins in northern Norway were more similar to the parameters of short-lived central European populations than to the parameters of the long-lived population of Newfoundland. This study confirms that northern Norwegian shorthorn sculpins exhibit sexual dimorphism as in other shorthorn sculpin populations. The relationships between growth pattern, age at maturity and mortality rates observed in the Tromsø population and in other shorthorn sculpin populations, correspond well with the predictions from a published life history model.     

REBUILDING SEAL STOCKS IN THE KATTEGAT-SKAGERRAK

The harbor seal ( Phoca vitulina ) population in the Kattegat-Skagerrak area has been dwindling for several centuries due to excessive hunting pressure. Corrected hunting statistics during 1890–1976 are used to estimate changes in population size over the past century. After protection was introduced in the 1960s and 1970s the harbor seal population in the area increased at an exponential rate of 0.12 and exceeded 5,000 animals in 1986. The present rate of population growth is used for modelling the influence of fertility and age-specific mortality. It is found that the observed high rate of increase is only realistic if female fertility rate is very high, the range of juvenile mortality rate is 0.33–0.52 and adult mortality is less than 0.15. Commonly cited higher mortality rates are not realistic in the Kattegat-Skagerrak area.     

Size matters when it comes to the survival of transplanted yellow gorgonian fragments

The yellow gorgonian Eunicella cavolini is a Mediterranean endemic coral. Mediterranean populations underwent sharp decreases in the last decades due to climatic anomalies. In the meanwhile, populations in the Sea of Marmara thrive under conditions exempted from significant thermal variations thanks to the peculiar oceanography of the basin. However, they are vulnerable to anthropogenic activities, such as fishing, coastal constructions, dredging and landfilling. Major infrastructure projects at two remote islands, Yassıada and Sivriada, put healthy gorgonian communities at risk. We observed many dead colonies when Yassıada was constructed and before operations commenced on Sivriada, we transplanted gorgonian fragments 13 km away to a site at lower risk from construction. The transplantation site already hosts yellow gorgonian as sparse colonies; but also, it sits within an important fishing area, and entangled nets over rocks are common. Therefore, together with a local NGO, we initiated conservation efforts simultaneously with the transplantation works. 300 gorgonian fragments in the 4.5–15.4 cm size range were transplanted in two years (2017 – 2019). At six months, the survival of the colonies was significantly size dependent. Fragments of 10 – 15.4 cm initial sizes had the highest survival rate (up to 90 %), while small fragments yielded substantially lower survival. Despite severe impacts from fishing nets and mucilaginous entanglement, 50 % of fragments larger than 10 cm survived after 4.5 years of plantation. Besides, several recruits were observed among 4.5 years old transplants. Therefore, when restoration by raw transplantation actions is planned for gorgonians with thin scleraxis, a minimum fragment size of 10 cm is recommended. The study was a notable example of science/NGO collaboration with substantial local conservation success by using extensive media coverage of conservation efforts as leverage to declare the recipient area as a Marine Protected Area.     

Testes development and maturity classification of albacore (Thunnus alalunga ( Bonaterre, 1788)) from the Eastern Mediterranean Sea

The criteria of testicular maturity during the spawning season and for maturity rates are described according to age for 140 male albacore (Thunnus alalunga, Bonnaterre, 1788) sampled between April and August, 2002–2008 in the Eastern Mediterranean Sea. Histology of gonad samples was used to determine maturity status. Spermatogenesis and its relation to reproductive biology are reported. Fish ranged in fork length (FL) from 63 to 90 cm. The reproductive classes were based upon changes in the testicular morphology and stages of the germinative cells, i.e. immature, developing, maturing, spawning, and spent. Mature stages of testes including the smallest sample of a 63 cm male were observed from May to August of each year during the study. The reported evidence indicates that the size and age of first sexual maturity values of males of the Eastern Mediterranean population are nearly the same as in the Western Mediterranean population, but lower than in the Atlantic stock. The results of this histological investigation, conducted for the first time on this species in the Mediterranean Sea, will enable more precise future estimations for sustainability of the male albacore stock in the Eastern Mediterranean.     

The population density, growth rate and production of roach Rutilus mtilus (L.) in Tjeukemeer, The Netherlands

A method of estimating the population density of roach in Tjeukemeer (21.3 km²) using 20 529 introduced fin-clipped fish is described. Fyke nets proved to be an effective method of sampling the population for marked fish during the spawning season. A total of 20 277 roach were processed during the recapturing period. The population density of roach (⋝ 14 cm) was estimated to be 1 246 458. The growth rate of roach in the lake although relatively poor (von Bertalanffy's L ∞ for males and females, 22 and 26 cm respectively) for the species was similar to that recorded in three other Friesian lakes. The mortality rate of males was higher than that of females. Logarithmic length-weight regression analyses showed that the value of the coefficient varied both within and between the sexes, that of females being higher (range 3.03–3.375) than that of males (range 2.76–3.254). Seasonal changes in the size of the coefficient were due to a disproportionate loss of gonad weight in larger fish. The fecundity of the population was comparatively high for the species.
The total production of the population was estimated to be 95 hg ha⁻¹ of which 39 kg ha⁻¹ was contributed by fry. In older (⋝ IV) fish the production of females (12 kg ha⁻¹) exceeded that (2 kg ha⁻¹) of males, due to differences in their growth and mortality rates. The relatively poor performance of roach in Tjeukemeer, in terms of biomass and production was due to a scarcity of zoobenthos and competition from other species offish. There is no evidence either from this study or the literature that the productive potential of roach in lakes is high, even though macrophytes and detritus can be consumed in significant amounts.     

AGE and SEX COMPOSITION OF THE STRIPED DOLPHIN DIE-OFF IN THE WESTERN MEDITERRANEAN

During the morbillivirus epizootic that affected the Mediterranean striped dolphins ( Stenella coeruleoalba )in 1990–1992, several thous and individuals are believed to have died. In order to determine the segments of the population that were affected by the event, the sex- and age-frequency distributions of the animals that were washed ashore on the Spanish coasts were studied. The age of the animals was determined by counting dentine growth layer groups in the teeth. The results obtained indicate that both sexes were affected similarly by the disease and that the population component that suffered the largest mortality was that of sexually mature individuals, although substantial mortality of calves, probably still dependent on their mothers, was also observed. Juveniles, in contrast, appeared to be only slightly affected by the epizootic. This pattern of mortality is not consistent with the epidemiology of morbillivirus infections previously observed in other mammals. Possible explanations of this particular age distribution include behavioral factors, increased susceptibility to the disease in adults due to the effects of pollution, high temperatures and decreased food availability, or allopatric geographical distribution of population components and, therefore, of their associated mortality.     

东灵山辽东栎林主要树种种群11年动态变化

 该研究在北京东灵山区一个30 m×40 m的纯辽东栎(Quercus liaotungensis)林永久性样地内进行,林龄为80年左右。样地中高于2 m的成树密度在1991、1997和2002年3次调查中呈逐年递减趋势,这种降低主要是由辽东栎个体的死亡引起的;而树木总胸径面积呈逐年增加趋势,主要是存活个体生长的结果。从1991年到2002年,辽东栎种群的优势度变化不大,且种群径级结构为单峰曲线,差异不显著。为了探讨这种径级分布格局形成的机制,应用离散模型对辽东栎种群的胸径年平均生长率G(t, x)、年平均死亡率M(t, x)及年平均生长率方差D(t, x)进行了计算,发现辽东栎种群的死亡率曲线呈倒J-型,个体的年平均生长率与胸径呈显著的线性正相关（p<0.01）。离散模型的系数分析和G(t, x)-D(t, x)的关系曲线分析均表明辽东栎种群内存在非对称种内竞争,即大胸径个体对小胸径个体的生长有抑制作用,这种作用主要是由相对较高的种群密度引起的对光资源的竞争。非对称种内竞争有利于维持辽东栎种群和整个群落的稳定状态。     

The spatial scale of competition from recruits on an older cohort in Atlantic salmon

Competitive effects of younger cohorts on older ones are frequently assumed to be negligible in species where older, larger individuals dominate in pairwise behavioural interactions. Here, we provide field estimates of such competition by recruits on an older age class in Atlantic salmon (Salmo salar), a species where observational studies have documented strong body size advantages which should favour older individuals in direct interactions. By creating realistic levels of spatial variation in the density of underyearling (YOY) recruits over a 1-km stretch of a stream, and obtaining accurate measurements of individual growth rates of overyearlings (parr) from capture–mark–recapture data on a fine spatial scale, we demonstrate that high YOY density can substantially decrease parr growth. Models integrating multiple spatial scales indicated that parr were influenced by YOY density within 16 m. The preferred model suggested parr daily mass increase to be reduced by 39% when increasing YOY density from 0.0 to 1.0 m⁻², which is well within the range of naturally occurring densities. Reduced juvenile growth rates will in general be expected to reduce juvenile survival (via increased length of exposure to freshwater mortality) and increase generation times (via increased age at seaward migrations). Thus, increased recruitment can significantly affect the performance of older cohorts, with important implications for population dynamics. Our results highlight that, even for the wide range of organisms that rely on defendable resources, the direction of competition among age classes cannot be assumed a priori or be inferred from behavioural observations alone.     

Optimal reproductive strategies in age-structured populations of zooplankton

SUMMARY. We describe a model of zooplankton population dynamics that accounts for differences in mortality and physiology among animals of different ages or sizes. The model follows changes in numbers of individuals and changes in individual and egg biomass through time and it expresses mortality and net assimilation as functions of animal size.
We investigated the effect of egg size, age at first reproduction, and size at first reproduction on the per capita growth rates of populations growing under different conditions. In the absence of predation or when exposed to vertebrate predators that prefer large prey, populations achieve maximum growth rates when animals hatch from small eggs and reach maturity quickly at small sizes. Populations exposed to invertebrate predators that concentrate on small animals may increase r in two different ways. One way is for animals to increase juvenile survivorship by hatching from large eggs and by shortening the juvenile period. An alternative strategy is for animals to hatch from small eggs and to postpone maturity until they grow beyond the range of sizes available to their predators. Certain life history strategies maximize r if animals continue to grow after they reach maturity. By growing larger, non-primiparous females are able to hatch larger clutches and thereby increase the overall rate of population growth.
The model analysis shows how to assess age-dependent mortality rates from field data. The net rate of population increase and the age distribution of eggs together provide specific, quantitative information about mortality.     

Population structure, growth and reproduction of roughhead grenadier on the Flemish Cap and Flemish Pass

Roughhead grenadier Macrourus berglax were collected during the Flemish Cap bottom survey (1991–2001, NAFO division 3M, depth of 200–720 m) and on commercial vessels (1998–2000, NAFO divisions 3L, M and N, depth of 700–1700 m). The abundance of roughhead grenadier, as well as their average size, increased with depth. Age and length composition of the catches showed clear differences between the sexes. The proportion of males in the catch declined in the larger length classes and there were no males in the largest length classes (25 cm). There were differences in the growth trajectories of the two sexes; both sexes grew similarly up to 9 years, but the male growth was slower thereafter. The pre-anal fin length at first maturity ( L _PA50) for roughhead grenadier females was 28·5 cm, whilst the age at 50% maturity was 15–16 years. Individual total fecundity varied between 8522 and 61 844 oocytes. Macrourus berglax had a prolonged life cycle and multi-aged population structure, with differences in growth and mortality between males and females, slow maturation and low fecundity.     

Dynamics,productivity and species richness of tropical rainforests along elevational and edaphic gradients on Mount Kinabalu,Borneo

We studied the dynamics of nine tropical rainforests on Mount Kinabalu, Borneo, at four elevations (700, 1,700, 2,700 and 3,100 m) on various edaphic conditions for four 2-year periods over 8 years (1995–2003), and examined the relationships with above-ground productivity. Mean growth rate of stem diameter, basal area turnover rate and estimated recruitment rate (using growth rate and size distribution) correlated with productivity among the nine forests in all periods. These rates based on growth rates of surviving stems appeared to be good measures of stand turnover. However, observed recruitment rate and mortality (and turnover rate as mean of these rates) based on direct observation of recruits and deaths did not correlate with productivity in some periods. These rates may not be useful as measures of stand turnover given small sample size and short census interval because they were highly influenced by stochastic fluctuation. A severe drought associated with the 1997–1998 El Niño event inflated mortality and depressed mean growth rate, recruitment rate and basal area turnover rate, but had little effect on the correlations between these rates (except mortality) and productivity. Across broad elevational and edaphic gradients on Mount Kinabalu, forest turnover, productivity and species richness correlated with each other, but the causal interpretation is difficult given the different histories and species pools among forests at different elevations.     

Nuthatches (Sitta europaea: Aves) in forest fragments: demography of a patchy population

Breeding density, local survival and summer recruitment of nuthatches were evaluated in a population scattered over many small (1–30 ha) forest fragments, and compared with study plots inside larger forests. Since most young birds settle outside the fragment in which they were born this population corresponds to the “patchy population” concept implying that patterns in abundance may be better explained by processes at the population level than by metapopulation processes. Mean breeding density was c. 50% lower in fragments and decreased with regional isolation (distance from larger forests) but not with local isolation (distance to nearby fragments). Local survival of adults and established (i.e. territorial) 1st-year birds was not related to forest size or isolation. However, fewer young birds settled in summer in the fragments compared with a large forest. This difference probably reflects high mortality during the sensitive dispersal phase. Moreover, the observed number of recruits and their estimated survival was insufficient to maintain the breeding population, suggesting significant net immigration from larger forests. This “rescue effect” explains why densities are affected by regional, but not local isolation. Received: 14 December 1998 / Accepted: 1 March 1999