Modelling changes in migration pattern of herring: collective behaviour and numerical domination

According to the adopted-migrant hypothesis, first time spawning herring Clupea harengus learn their migration pattern from schooling with older individuals. Changes in migration pattern of Norwegian spring-spawning herring co-occur with the recruitment of abundant year classes to the spawning stock. It is argued that this is due to the abundant year classes being unable to learn from the older ones since most of the recruiting year class only experience their own naïve class-mates. An individual-based school simulation study is presented to explore the hypothesis. Behaviour is simulated using individual rules that take into account the movements of nearby individuals. After establishing a schooling pattern, a small proportion of the individuals is directed to move towards a given position while the remaining individuals keep using their behaviour rules. When <4% of the individuals are so directed there is no response seen in the remaining school. But when ≥7% of the total school is directed to a specific location, the remaining individuals always respond and follow the directed individuals. The simulations thus show that determined individuals can have a substantial influence on the collective behaviour of schools. Furthermore, the knife-edge response in school behaviour suggests that the relative abundance of determined individuals in a school plays an important part in the collective behaviour. This mechanism can explain the observed changes in migration pattern of Norwegian spring-spawning herring associated with the recruitment of abundant year classes to the adult Stock.     

Ovigerity,selective predation,and variable diel vertical migration in Euchaeta elongata (Copepoda: Calanoida)

Summary We present a statistical analysis of a previously published (Yen, 1983) but heretofore unanalyzed data set on the vertical distributions and diel vertical migration (DVM) of adult females of the marine planktonic copepod Euchaeta elongata in Dabob Bay, Washington, USA. Non-ovigerous females were strongly migratory on all four dates sampled, residing between 75–175 m during the day and at shallower depths during the night, commonly entering the upper 50 m of the water column. In contrast, ovigerous females were non-migratory or weakly migratory, largely remaining between 100–175 m both day and night, and entering the upper 50 m of the water column only rarely. Thus non-ovigerous females always migrated much more strongly, as measured by both amplitude of migration and the proportion of animals migrating, than did ovigerous females. These results led us to hypothesize that differential susceptibility to visually orienting predators was the cause of these differences in DVM behavior in female E. elongata, and we subsequently undertook an experimental study of the feeding selectivity of the copepod's natural predator, Pacific herring (Clupea harengus pallasi). Pacific herring exhibited a highly significant preference for ovigerous over nonovigerous adult female E. elongata. The demographic consequences of variable DVM in adult female E. elongata were investigated by way of life table analyses. Results indicated that under conditions of thermal stratification of the water column there is a distinct demographic disadvantage (reduced rate of realized population growth) incurred by non-migratory or weakly migratory ovigerous females due to delayed egg development at cooler subsurface temperatures. We conclude that ovigerous female E. elongata remain at depth both day and night to avoid visually orienting predators, and that such behavior must afford the copepod a demographic advantage of no less than a 26% reduction in adult mortality to offset the demographic cost of delayed egg development.     

The Dynamics of a Metapopulation: Changes in Life-History Traits in Resident Herring that Co-Occur with Oceanic Herring during Spawning

Different populations of Atlantic herring are regarded as forming a metapopulation, but we know little about the dynamics of the connectivity and degree of interbreeding between the populations. Based on data from three periods between 1962 and 2011, we identified the presence of two components of herring in a small semi-enclosed coastal marine ecosystem based on different somatic growth patterns and mean vertebrae sum (VS). The two components were interpreted as belonging to a resident herring population and the migratory, oceanic Norwegian spring spawning (NSS) herring population, and they co-occurred during spawning. In the 1960s, resident herring characterized by slow growth and low VS co-occurred with rapid growth, high VS oceanic NSS herring. Similar slow-growing resident and rapid-growing NSS herring were found in the 1970–80s, but both populations now had low VS suggesting similar origins. Finally, in the 2000s both populations showed rapid growth. The changes coincided with the NSS herring going from a state of high abundance and oceanic distribution to a collapse in the late 1960s that resulted in a coastal distribution closer to resident herring populations, before full recovery and resumption of the migratory, oceanic pattern in the 1990s. During all three periods, NSS herring were only present in the local system up to an age of about five years, but the synchronous spawning of the populations supports mixed spawning and interbreeding. During the investigation period both longevity, length at age (growth) and length-at-first maturity increased markedly for the resident herring, which then became more similar to the NSS herring. Genetic and/or cultural factors are believed to be the main causes of the observed changes in life history traits, although some effect of changes in environmental factors cannot be excluded. Our study suggests that relationships among populations in a metapopulation can be highly dynamic.     

Cultural transmission and flexibility of partial migration patterns in a long‐lived bird,the great bustard Otis tarda

Factors responsible for individual variation in partial migration patterns are poorly known, and identifying possible causes of these changes is essential for understanding the flexibility in migratory behavior. Analyzing 190 life histories of great bustards Otis tarda radio‐tagged in central Spain, we investigated the changes in migratory tendency across lifetime in this long‐lived bird, and how migratory flexibility is related to individual condition. In females migratory behavior was not fixed individually. For every age class there was a fraction of ca 15–30% of females that changed their migratory pattern between consecutive years. Migrant females tended to remain sedentary in years when they had dependent young to attend. These findings show that the female migratory tendency is a behaviorally flexible, condition‐dependent trait. Immature females usually acquired their migratory behavior by learning from the mother in their first winter or by social transmission from other migratory females in their second winter. As for immature males, their summer migratory behavior was not related to mother–offspring transmission, but learned from adult males. We found that their age‐related increase in migratory tendency was associated to a greater integration in flocks of migrant adult males. These results show that within the partial migration system, cultural transmission mechanisms, either mediated by kin or not, and individual condition, may contribute to shape the migratory tendency. Our study reinforces the view that the migratory behavior is an evolutionary complex trait conditioned by the interaction of individual, social and environmental factors. Particularly in long‐lived species with extended parental care, the inherited migration program may be shaped by mother–offspring and social transmission of migratory patterns.     

Ovarian dynamics and fecundity regulation in blueback herring,Alosa aestivalis,from the Connecticut River,US

We analyzed ovarian dynamics of anadromous blueback herring, Alosa aestivalis, in Connecticut River with the principal aim of exploring oocyte recruitment and how it shapes the fecundity pattern. We examined the oocyte release strategy and analyzed spawning cyclicity by linking oocyte growth to the degeneration of postovulatory follicles. Females were accordingly classified as pre-spawners, early and late active spawners, and oocyte recruitment intensity was compared among the different spawning phases. Oocyte recruitment occurred continuously and in parallel with spawning activity, a pattern which is diagnostic of indeterminate fecundity. However, both fecundity and oocyte recruitment intensity progressively decreased (tapered) throughout spawning, until the ovary was depleted of vitellogenic oocytes. There was no massive atresia of vitellogenic oocytes at the end of the spawning season, which is atypical of indeterminate spawners. We propose that tapering in oocyte recruitment and fecundity is an adaptation to the high energetic expenditure of the upstream spawning migration.     

Olfactory gene expression in migrating adult sockeye salmon Oncorhynchus nerka

Expression of 12 olfactory genes was analysed in adult sockeye salmon Oncorhynchus nerka nearing spawning grounds and O. nerka that had strayed from their natal migration route. Variation was found in six of these genes, all of which were olfc olfactory receptors and had lower expression levels in salmon nearing spawning grounds. The results may reflect decreased sensitivity to natal water olfactory cues as these fish are no longer seeking the correct migratory route. The expression of olfactory genes during the olfactory‐mediated spawning migration of Pacific salmon Oncorhynchus spp. is largely unexplored and these findings demonstrate a link between migratory behaviours and olfactory plasticity that provides a basis for future molecular research on salmon homing.     

Migratory behaviour of adult pikeperch (Stizostedion lucioperca) in a lowland river

The behaviour of radio-tagged adult pikeperch (Stizostedion lucioperca (L.)) from two areas in the Danish River Gudenaa were recorded prior to, and during the spawning period. Eight of 13 tagged fish in the lower reaches of the river were located throughout the whole study. Five of these fish moved upstream to various sites in the river prior to spawning, which occurred from late April to June. The three remaining fish moved to the fjord. These movements were interpreted as a spawning migration, and it is suggested that the lower reaches of the River Gudenaa constitute an over-wintering area for pikeperch, which use different spawning areas. Ten pikeperch caught just downstream of an impassable hydropower plant were radio-tagged and translocated upstream of the dam to a reservoir. Within a week, half of the fish moved to a lake situated more than 30 km upstream the reservoir. This behaviour is hypothesised to be a homing response. The study reveals that the pikeperch is a highly mobile species with a complicated migration pattern, even in relatively small river systems.     

Gulls can change their migratory behavior during lifetime

Migration is a widespread phenomenon among birds and is likely to be subject to strong selective pressures. Birds' annual routines and behaviors might be expected to change during their different life history stages, resulting in different, age‐related migration patterns. However, although migration has been the subject of many publications, age‐related differences in migration have received little attention. The present study examined age‐related changes in individual migration habits in lesser black‐backed gulls, Larus fuscus. We analyzed data from 10‐year (1998–2007) color‐ringing project in NW England, comprising more than 10 000 ringed individuals. Our results showed a latitudinal cline in age structure across the wintering range, with adults and gulls in their first breeding year wintering closer to the breeding grounds. Supporting this result we observed that individuals, as they get older, changed the migration behavior and winter closer to the breeding areas. Interestingly, we found no differences in survival rates across the wintering grounds. Thus differences in survival rates can not account for the latitudinal cline in age structure, and the observed findings seem to be best explained by the arrival time hypothesis, based on a mechanism whereby individuals are able to change their migratory behavior as result of the onset of sexual maturity and associated mating pressures.     

Migratory history of the spotted flagtail, <Emphasis Type="Italic">Kuhlia marginata</Emphasis>

The spotted flagtail, Kuhlia marginata (Kuhliidae) is commonly found in streams throughout the Ryukyu Archipelago, including Okinawa Island, Japan. Although it has been suggested that they spawn at sea, little is known about their migratory history. The aim of the study was to clarify their migration history based on otolith microchemistry analysis, size composition and the gonadal development of fish collected from the Genka River on Okinawa Island. All strontium/calcium (Sr/Ca) ratios of otoliths remained high around their cores, and then dropped rapidly at some distance from the core. The estimated standard length corresponding to decreases correlated with the size of new recruits collected in the river (ca. 20 mm in standard length: SL). Multiple increases, the sign for migrations into the high saline area, were recognized intermittently around the otolith margin. As the estimated SL at these increases in the Sr/Ca ratio occurred above the minimum maturation size (female: 95.5 mm SL; male: 83.5 mm SL), such multiple increases in Sr/Ca ratios were caused by spawning migration. Therefore, we conclude that the catadromous pattern of K. marginata is as follows; this species grows in the sea during the early life stage until ca. 20 mm SL and then grow in the freshwater area until maturation, before they migrate again to the sea for spawning. Sr/Ca ratio profiles also suggested that a large number of males did not return to the freshwater area after spawning, whereas females might spawn several times during their lives around the inshore area.     

The survival of <Emphasis Type="Italic">Sicyopterus stimpsoni</Emphasis>, an endemic amphidromous Hawaiian gobiid fish,relies on the hydrological cycles of streams: evidence from changes in algal composition of diet through growth stages fish

Gut contents of larval, juvenile, and adult specimens of the Hawaiian gobiid fish Sicyopterus stimpsoni were examined to catalog the algal flora ingested by this species. The developmental stages of S. stimpsoni examined represented hallmark points in the fish’s life cycle corresponding with major migratory and metamorphic transitions. The algal flora was dominated by diatom species and shifted from taxa representative of a marine, planktonic community in larval fish to a freshwater, benthic community in juvenile and adult fish. This change in diet corresponds with the migration of larval fish to freshwater streams just prior to juvenile development in which rapid modification in mouth anatomy makes ingestion of planktonic algal species difficult. Benthic diatoms from juvenile and adult fish assemblages represented multiple genera that live in a narrow set of environmental conditions. These algae grow during a specific period in the development of the benthic algal community in Hawaiian streams. This suggests a highly specialized dietary behavior that depends heavily on continually restarting the benthic algal successional pattern, which appears to be regulated by the hydrological cycles of streams on the island.     

Factors affecting migration patterns of juvenile river herring in a coastal Massachusetts stream

Juvenile anadromous river herring (alewives, Alosa pseudoharengus, and blueback herring, Alosa aestivalis) spend 3–7 months in freshwater before migrating to the ocean, often exhibiting waves of early and late migrations. Migratory patterns and associated abiotic and biotic factors were examined for both species migrating in 2003 from Herring River in Bourne Massachusetts. Migrating herring were grouped into two temporal periods and separated by species. Relationships between abiotic (water temperature, lunar phase, and precipitation) and biotic (size, age, growth rate, and hatch date) factors and the timing of out-migration (transition to seawater) were examined. Blueback herring migration coincided with a sharp decrease in temperature consistent with the time of year. Peaks in alewive migration may reflect adaptive traits maximizing likelihood of survival through seasonal marginal stream habitats and also avoiding lethal water temperatures in winter. Migrating bluebacks exhibited a single migratory period (late September–October) which occurred between the early and late migratory pulses of the alewives. These patterns may also represent distinct migration strategies between early and late migrating alewives and between the two species of river herring. Early migration may be a mechanism by which factors such as competition and low food availability are mitigated. In contrast, environmental conditions during the post hatching time of late migrators may allow for a size maximizing strategy that promotes migration from the nursery area as late as possible.     

Geographic and clonal variation in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae), for winged morph production,life history,and morphology in relation to host plant permanence

Summary Populations of the milkweed-oleander aphid,Aphis nerii, were sampled in California, Iowa and Puerto Rico. Among these localities the aphid's host plants differ greatly in permanence. I compared populations for migratory potential, measured as the proportion of winged offspring produced in response to being crowded, and for life history and morphometric traits of the subsequent adult winged aphids. I predicted a negative correlation between degree of host plant permanence and migratory potential. As predicted, aphids from Iowa, where migration on to temporary hosts must occur each year, produce a greater proportion of winged offspring (37.7%) than those from California (25.7%) or Puerto Rico (31.6%) where hosts are more permanent. However, hosts in Puerto Rico appear to be more permanent than those in California, yet the difference between populations for migratory potential was opposite to that predicted. Within California the prediction again held: aphids collected from the most impermanent sites produce the greatest proportion of winged offspring. There were no population differences for any life history or morphometric traits of winged aphids that are important contributors to fitness or migratory ability such as time to reproductive maturity, fecundity or wing length. Nor did any traits covary with migratory potential. Thus, there does not appear to be an association of life history and morphology with migratory potential that could enhance the colonizing ability of migrant aphids. I was unable to detect population differentiation for life history and morphology even though there is ample genetic variation within populations on which selection could act and an absence of constraints arising from genetic correlations that could prevent appropriate evolution of traits within populations. The exploitation of temporary host plants therefore occurs by an increase in the number of colonists produced and not by change in life history or morphology of those colonists.     

Return migration of Atlantic Salmon in the River Tana: Phases of migratory behaviour

From a total of 174 multi-sea-winter Atlantic salmon radio tagged in the Tanafjord (northern Norway, 70°N) during 1992 and 1993, 48 Atlantic salmon were followed from entering the River Tana until spawning. Three phases were identified: (1) migratory, direct or stepwise migration to, or close to the position held at spawning; (2) search, movements both up and down river at or close to the position held at spawning; (3) holding, a period without movements prior to spawning. During the migratory phase, Atlantic salmon migrated directly to near the spawning area, or stopped between one and nine shorter periods during the upstream migration. Number of stops increased with increasing migratory distance in 1993, but no such correlation was found in 1992. The highest migratory speeds were recorded in the lower parts of the river. A distinct change in migratory pattern was found in 67% of the Atlantic salmon near or at the area held at spawning. Most common was a search phase of erratic movements with more than one down river movement. After the movement terminated, 96% of the Atlantic salmon had a period when no or little movement was recorded until spawning (on average 55 days in 1992 and 51 days in 1993). There was no preference for staying at, up or down river from the spawning area during this holding period. Early ascending Atlantic salmon migrated to spawning areas further from the mouth than the later arriving Atlantic salmon in 1993, but not in 1992. The proportion of time spent on the migratory phase increased, while the proportion of time spent on the holding phase decreased with increasing distance to the spawning area.     

Evolutionary aspects of migratory behavior in European warblers

During the past 20 years, European Sylvia warblers have been used for a model study of the control mechanisms of bird migration and of evolutionary aspects of migratory behavior. Endogenous annual rhythms (‘circannual’ rhythms) and photo-period have proved to be the essential internal and external controlling factors. It is unknown whether this basic system, that also controls migration in other bird species, is currently evolutionarily stable or is instead adapting birds to the present slightly changing environmental conditions. Using the Blackcap, the control of partial migration in a bird species was analyzed. Two-way selective breeding experiments demonstrated a large selection response and high heritability values. These experiments have also indicated that a partially migratory population can become either almost completely migratory, or sedentary, in two to five generations. Hence, genetic influences are very important and presumably dominant over environmental factors in the expression of migratory or sedentary behavior. The large selection response implies a strikingly high evolutionary potential with respect to strong selection pressures. Further, in the Blackcap, migratory orientation behavior (in addition to migratory activity) was immediately transmitted into a F₁-generation when a cross-breeding experiment was performed using birds from a migratory and a resident population. The hybrids displayed their migratory activity along an axis that is used by their migratory parents. Finally, a rapidly developing novel migratory habit (new migratory direction to new wintering areas) in the Blackcap is discussed with respect to a positive feedback-mechanism, possibly including a series of advantages leading to above-average fitness.     

Effect of spatial differences in growth on distribution of seasonally co‐occurring herring Clupea harengus stocks

The mechanisms most likely to determine the distribution of the two major herring Clupea harengus stocks in their common early summer feeding ground in the eastern North Sea, Skagerrak and Kattegat were investigated through analysis of acoustic survey data from six consecutive years. No change was detected in biomass of North Sea autumn spawning C. harengus (NSAS) over time, whereas the biomass of western Baltic spring spawning C. harengus (WBSS) declined severely. Analyses of centre of abundance by stock showed no change in NSAS distribution, whereas the WBSS changed to a more western distribution over time. Contrary to previous perception of the juvenile migration, NSAS were found to leave the study area at the age between 1 and 2 years and WBSS 1 year olds were encountered in the Skagerrak. The estimated parameters of von Bertalanffy growth equations showed marked differences between areas with fish in the eastern part of the area having the lowest size at age at all ages. Further, their growth conditions appeared to deteriorate progressively over the period studied. Both NSAS and WBSS showed the highest condition in the North Sea and Skagerrak while condition was substantially lower in age Kattegat. The westward movement of spring spawners over time suggests that growth rate and possibly density of conspecifics influence the migration pattern and distribution of C. harengus in the area. In contrast, there was no evidence to suggest that distribution was constant over time within stocks or that distribution reflected size‐dependent limitations on migration distance.     

Climate change and condition of herring (Clupea harengus) explain long-term trends in extent of skipped reproduction

It is commonly assumed that iteroparous fish, once mature, normally reproduce in all consecutive seasons. Recent work has suggested, however, that in Norwegian spring-spawning herring—a population that undertakes extensive spawning migrations—almost one in two adults may skip their second spawning migration. Why should herring not return to spawn the year after first spawning, but instead wait an extra year? For herring, participation in distant, energetically costly, and risky spawning migrations will only pay off in terms of fitness if individuals are sufficiently large, and in sufficient condition, to both successfully migrate and spawn. Changes in the environment and individual condition should therefore affect the likelihood of skipped spawning. This paper describes long-term changes in the extent to which the second reproductive season is skipped in this herring population. These are shown to be linked to the size and condition of herring as first-time spawners, and to climatic factors possibly related to food availability. The findings corroborate the hypothesis that skipped reproduction results from trade-offs between current and future reproduction, growth and survival.     

Biocomplexity in a highly migratory pelagic marine fish, Atlantic herring

The existence of biologically differentiated populations has been credited with a major role in conferring sustainability and in buffering overall productivity of anadromous fish population complexes where evidence for spatial structure is uncontroversial. Here, we describe evidence of correlated genetic and life history (spawning season linked to spawning location) differentiation in an abundant and highly migratory pelagic fish, Atlantic herring, Clupea harengus, in the North Sea (NS) and adjacent areas. The existence of genetically and phenotypically diverse stocks in this region despite intense seasonal mixing strongly implicates natal homing in this species. Based on information from genetic markers and otolith morphology, we estimate the proportional contribution by NS, Skagerrak (SKG) and Kattegat and western Baltic (WBS) fish to mixed aggregations targeted by the NS fishery. We use these estimates to identify spatial and temporal differences in life history (migratory behaviour) and habitat use among genetically differentiated migratory populations that mix seasonally. Our study suggests the existence of more complex patterns of intraspecific diversity than was previously recognized. Sustainability may be compromised if such complex patterns are reduced through generalized management (e.g. area closures) that overlooks population differences in spatial use throughout the life cycle.     

Mismatch between fishway operation and timing of fish movements: a risk for cascading effects in partial migration systems

Habitat fragmentation is a growing problem worldwide. Particularly in river systems, numerous dams and weirs hamper the movement of a wide variety of species. With the aim to preserve connectivity for fish, many barriers in river systems are equipped with fishways (also called fish passages or fish ladders). However, few fishways provide full connectivity. Here we hypothesized that restricted seasonal opening times of fishways can importantly reduce their effectiveness by interfering with the timing of fish migration, for both spring‐ and autumn‐spawning species. We empirically tested our hypothesis, and discuss the possible eco‐evolutionary consequences of affected migration timing. We analyzed movements of two salmonid fishes, spring‐spawning European grayling (Thymallus thymallus) and autumn‐spawning brown trout (Salmo trutta), in Norway's two largest river systems. We compared their timing of upstream passage through four fishways collected over 28 years with the timing of fish movements in unfragmented river sections as monitored by radiotelemetry. Confirming our hypothesis, late opening of fishways delayed the migration of European grayling in spring, and early closure of fishways blocked migration for brown trout on their way to spawning locations during late autumn. We show in a theoretical framework how restricted opening times of fishways can induce shifts from migratory to resident behavior in potamodromous partial migration systems, and propose that this can induce density‐dependent effects among fish accumulating in lower regions of rivers. Hence, fragmentation may not only directly affect the migratory individuals in the population, but may also have effects that cascade downstream and alter circumstances for resident fish. Fishway functionality is inadequate if there is a mismatch between natural fish movements and fishway opening times in the same river system, with ecological and possibly evolutionary consequences for fish populations.     

Coastal zone use and migratory behaviour of the southern population of Mugil liza in Brazil

We analysed the ratios Sr:Ca and Ba:Ca in the otoliths of 55 adults of the southern population of Mugil liza in Brazil (Paraná to Rio Grande do Sul) to investigate its coastal zone use and migratory behaviour. All individual M. liza analysed had Sr:Ca and Ba:Ca values indicating that their birth was in the marine environment, which is consistent with the southern population migration to spawn in the ocean,. Juveniles exhibited at least three coastal use and recruitment strategies (contingents): the majority (89%) of M. liza juveniles migrated toward brackish water. They entered the estuary before completing the first year of life (64%) or after (25%) their first year of life. The remaining 11% did not appear to enter brackish or freshwater water as a nursery or at any point in their life cycle. Some adults returned to the estuary after spawning in the ocean but others (of both sexes) never returned to the estuary after spawning, remaining in the marine environment. The pattern of juvenile habitat use in the Brazilian southern population of M. liza seems to be recurrent throughout the extent of its distribution as a consequence of the reproductive spawning aggregation behaviour, which mixes all contingents (with marine or estuarine preferences).     

“Closer‐to‐home” strategy benefits juvenile survival in a long‐distance migratory bird

Human‐induced changes in the climate and environment that occur at an unprecedented speed are challenging the existence of migratory species. Faced with these new challenges, species with diverse and flexible migratory behaviors may suffer less from population decline, as they may be better at responding to these changes by altering their migratory behavior. At the individual level, variations in migratory behavior may lead to differences in fitness and subsequently influence the population's demographic dynamics. Using lifetime GPS bio‐logging data from 169 white storks (Ciconia ciconia), we explore whether the recently shortened migration distance of storks affects their survival during different stages of their juvenile life. We also explore how other variations in migratory decisions (i.e., time, destination), movement activity (measured using overall body dynamic acceleration), and early life conditions influence juvenile survival. We observed that their first autumn migration was the riskiest period for juvenile white storks. Individuals that migrated shorter distances and fledged earlier experienced lower mortality risks. In addition, higher movement activity and overwintering “closer‐to‐home” (with 84.21% of the tracked individuals stayed Europe or North Africa) were associated with higher survival. Our study shows how avian migrants can change life history decisions over only a few decades, and thus it helps us to understand and predict how migrants respond to the rapidly changing world.