Changes in bird‐migration patterns associated with human‐induced mortality

Many bird populations have recently changed their migratory behavior in response to alterations of the environment. We collected data over 16 years on male Great Bustards (Otis tarda), a species showing a partial migratory pattern (sedentary and migratory birds coexisting in the same breeding groups). We conducted population counts and radio tracked 180 individuals to examine differences in survival rates between migratory and sedentary individuals and evaluate possible effects of these differences on the migratory pattern of the population. Overall, 65% of individuals migrated and 35% did not. The average distance between breeding and postbreeding areas of migrant individuals was 89.9 km, and the longest average movement of sedentary males was 3.8 km. Breeding group and migration distance had no effect on survival. However, mortality of migrants was 2.4 to 3.5 times higher than mortality of sedentary birds. For marked males, collision with power lines was the main cause of death from unnatural causes (37.6% of all deaths), and migratory birds died in collisions with power lines more frequently than sedentary birds (21.3% vs 6.3%). The percentage of sedentary individuals increased from 17% in 1997 to 45% in 2012. These results were consistent with data collected from radio‐tracked individuals: The proportion of migratory individuals decreased from 86% in 1997–1999 to 44% in 2006–2010. The observed decrease in the migratory tendency was not related to climatic changes (temperatures did not change over the study period) or improvements in habitat quality (dry cereal farmland area decreased in the main study area). Our findings suggest that human‐induced mortality during migration may be an important factor shaping the migration patterns of species inhabiting humanized landscapes.     

Estimating the influence of overhead transmission power lines and landscape context on the density of little bustard Tetrax tetrax breeding populations

Collision with conductors and earth cables is a known impact generated by transmission power lines, however there is virtually no information on how these infrastructures might affect bird distribution in a landscape context. With this work we specifically hypothesise that transmission power lines may affect the occurrence of a threatened bird, the little bustard (Tetrax tetrax). To test this hypothesis we used a Stochastic Dynamic Methodology (StDM), analysing the effects of power lines in a landscape perspective and simulating population trends as a response to power line installation and habitat changes induced by agricultural shifts in southern Portugal. The data used in the dynamic model construction included relevant gradients of environmental conditions and was sampled during the breeding seasons of 2003-2006. Transmission power lines were significantly avoided by the little bustard and the developed StDM model showed that the distance to these utility structures is the most important factor determining breeding densities in sites with suitable habitat for the species, which possibly leads to displacement of populations and habitat fragmentation. The model simulations also provided the base to analyse the cumulative effects caused by the habitat degradation that can ultimately lead to the extinction of local populations. Within priority conservation sites, the dismantling of existing transmission lines should be considered whenever possible, in order to ensure adequate breeding habitat. The model is considered useful as an auxiliary tool to be used in environmental impact assessments, management and conservation studies.     

Interpopulational variation in chemosensory responses to selected steroids from femoral secretions of male lizards, Podarcis hispanica, mirrors population differences in chemical signals

Chemical signals are important for mate and species recognition. If variation in chemical signals occurs between populations of the same species, these differences could later preclude mating between populations and lead to speciation. In the Iberian wall lizard, Podarcis hispanica, the lipophilic fraction of femoral secretions of males is mainly a mix of steroids and fatty acids. Among steroids, the most abundant compounds are cholesterol and cholesta-5,7-dien-3-ol, which are implicated in intraspecific communication and sexual selection. Interpopulational differences in chemical signals of males, and in response to these chemicals, could contribute to reproductive isolation between populations, which would explain the known genetic differences between these populations. Chemical analyses indicated that five distinct populations of this lizard from Madrid (Central Spain) differed in the proportions of two steroids (cholesterol and cholesta-5,7-dien-3-ol) in femoral secretions. Moreover, lizards discriminated and had high chemosensory responses (i.e., high tongue-flick rates) to these steroids, but showed interpopulational differences. Lizards from populations with cold temperatures and high relative humidity (i.e., northern Madrid) elicited higher responses to these steroids, whereas the converse occurred for lizards from populations occupying dry and hot habitats (i.e., southern Madrid). Interestingly, the magnitude of the chemosensory responses to cholesta-5,7-dien-3-ol in each population mirrored the abundance of this compound in secretions of males of that population. These results suggest that the importance of cholesta-5,7-dien-3-ol in males’ secretions might be lower for lizards from the southern populations. These differences in the relative importance of chemical signals could explain reproductive isolation and cryptic speciation between populations of this lizard.     

Spatio-temporal distribution and migration of adult male loggerhead sea turtles (Caretta caretta) in the Mediterranean Sea: further evidence of the importance of neritic habitats off North Africa

Ten adult male loggerhead sea turtles, captured by trawlers or dip nets, were satellite-tracked from a neritic foraging ground in the Mediterranean in order to investigate adult spatio-temporal distribution and breeding migration. Five individuals migrated to potential breeding sites in Libya and one to Greece. The results complement previous studies and show that: (1) the Tunisian shelf may be more important for turtles from Libyan rookeries than previously thought; (2) male tracks corroborate a conservation hotspot previously identified for juveniles; (3) the north African coast represents a preferred migratory corridor, unless open sea routes are more direct; (4) adult males may exhibit high fidelity to relatively small areas, without evident seasonal differences; (5) adults home ranges were smaller and more neritic than juveniles frequenting the same area; (6) males may frequent multiple courtship areas; (7) the average remigration interval of males frequenting this region is longer than 1 year.     

Density dependence and habitat quality modulate the intensity of display territory defence in an exploded lekking species

We evaluated the effect of conspecific abundance and habitat quality of leks on the territorial behaviour of males in an exploded lekking species, the Little Bustard (Tetrax tetrax). The hypothesis that males more intensely defend territories with higher conspecific abundance and better habitat quality was evaluated experimentally analysing the agonistic response of experimental males to male decoys placed on their displaying areas. Decoy experiments showed that the intensity of display territory defence by little bustard males is density dependent. The time experimental males took to return to their display sites after decoy placement decreased with abundance of both males and females. The strength of their final response was positively associated to local male and female abundance in the vicinity of their display sites. Habitat quality also influenced males’ display territory defence since the intensity of male response increased with the degree of natural vegetation cover. Habitat quality was particularly relevant in explaining variation of experimental males’ snort call rate, which decreased with the degree in plough cover and increased with the number of fields in the lekking area. Snort call rate decreased with the level of male aggregation and was lowest in males exhibiting the strongest aggressive response to decoys. These results add new evidence for the density dependence of species’ breeding territorial behaviour, supporting density-dependent models of lek formation and reinforcing the role of resources defence in exploded lek mating systems.     

Male's return rate, rather than territory fidelity and breeding dispersal, explains geographic variation in song sharing in two populations of an oscine passerine (Oreothlypis celata)

Males of some oscine passerines learn and share songs of neighboring males. This process can lead to the formation of song pattern neighborhoods or microhabitat song dialects. The degree to which song sharing occurs between populations and the spatial scale over which neighboring males share songs, however, can vary widely, and interpopulation comparisons have suggested that song sharing is more common in residents than in migrants. Here, we examine two populations of the orange-crowned warbler (Oreothlypis celata) to quantify patterns of song sharing at the northern (long-distance migrant) and southern (short-distance migrant) edges of the breeding distribution and to test if return rate, territory fidelity, and breeding dispersal explain the patterns found in the two populations. The southern population (O. celata sordida breeding on Santa Catalina Island, California; 33°N) had a higher annual return rate to their territories and exhibited higher song sharing in neighborhoods than their counterparts (O. celata celata breeding in Fairbanks, Alaska; 64°N). Year-to-year patterns of territory fidelity and breeding dispersal distances were similar between populations. Our results suggest that if migratory distance generally covaries with the proportion of returning males, this could explain different levels of song sharing between the short- and long-distance migrants.     

Bottoms up: great bustards use the sun to maximise signal efficacy

Visual displays are signals that may be selected to increase visibility. Light is a crucial component in the transmission of visual signals, and white colour is very conspicuous when illuminated by sun and exhibited against darker backgrounds. Here we tested the hypothesis that orientation of sexual displays in male great bustard (Otis tarda) depends upon position of the sun, i.e., males direct their uplifted white tails towards the sun in order to maximise signal detectability to distant females. We recorded the orientation of 405 male displays in relation to the sun and to females at seven leks. Great bustard males signalled towards the sun more often than expected by chance in early morning, although this pattern was not obvious at other times of day, when males displayed more towards females. Our hypothesis was further supported by the fact that displays were more directed towards the sun when the sun was most visible. Males were more likely to direct their displays towards females during the most elaborate components of their courtship display and when there were fewer males on the lek. Pointing white plumage to the sun may be a behaviour selected in species living in steppe-like open landscapes if individuals obtain net fitness benefit by increasing the likelihood of mating.     

Monarch butterfly migration and parasite transmission in eastern North America

Seasonal migration occurs in many animal systems and is likely to influence interactions between animals and their parasites. Here, we focus on monarch butterflies (Danaus plexippus) and a protozoan parasite (Ophryocystis elektroscirrha) to investigate how host migration affects infectious disease processes. Previous work showed that parasite prevalence was lower among migratory than nonmigratory monarch populations; two explanations for this pattern are that (1) migration allows animals to periodically escape contaminated habitats (i.e., migratory escape), and (2) long-distance migration weeds out infected animals (i.e., migratory culling). We combined field-sampling and analysis of citizen science data to examine spatiotemporal trends of parasite prevalence and evaluate evidence for these two mechanisms. Analysis of within-breeding-season variation in eastern North America showed that parasite prevalence increased from early to late in the breeding season, consistent with the hypothesis of migratory escape. Prevalence was also positively related to monarch breeding activity, as indexed by larval density. Among adult monarchs captured at different points along the east coast fall migratory flyway, parasite prevalence declined as monarchs progressed southward, consistent with the hypothesis of migratory culling. Parasite prevalence was also lower among monarchs sampled at two overwintering sites in Mexico than among monarchs sampled during the summer breeding period. Collectively, these results indicate that seasonal migration can affect parasite transmission in wild animal populations, with implications for predicting disease risks for species with threatened migrations.     

Winter migration of magellanic penguins (Spheniscus magellanicus) from the southernmost distributional range

A total of 22 magellanic penguins (Spheniscus magellanicus) from Isla Martillo in the Beagle Channel, Argentina, were successfully satellite tracked in 2004 (n = 7), 2005 (n = 7) and 2006 (n = 8) to monitor their winter migration after moult. Only one magellanic penguin migrated northwards into the Pacific Ocean, whereas all others remained in the Atlantic Ocean. In general, these birds left the island in an easterly direction, rounded Cabo San Diego, the southeasterly tip of South America, and continued northwards occupying inshore waters mostly less than 50 km from the coast, only occasionally venturing further offshore. By the end of the transmission period, birds were still travelling northwards and the most northerly positions were obtained from birds located in the area of Peninsula Valdés, Argentina, at a latitude of around 42°S, some 1,500 km from their breeding site on Isla Martillo. The mean maximum distance to the breeding site was, however, only 624 ± 460 km. The mean minimum distance covered during the study period was 1,440 ± 685 km, which corresponded to a mean distance of 23.2 ± 6.6 km covered per day. The northbound migration of the penguins could be separated into periods of rapid movement, interspersed with periods during which the birds remained for some time in particular coastal regions. Areas with a high density of daily penguin positions were observed in three distinct areas: at the northeastern coast of Tierra del Fuego, at the southern entrance of Golfo San Jorge and to the northeast of the Peninsula Valdez. The observed migration pattern is presumably driven by the formation and subsequent dispersal of areas of enhanced productivity as the season progressed. Our findings also suggest that magellanic penguins are increasingly threatened by human activities in coastal areas as penguins migrate northwards.     

Pleistocene Reindeer and Global Warming

Abstract:  Current concerns for the future of reindeer and caribou ( Rangifer tarandus ) in the far north under conditions of global warming focus on the increased energetic and predation costs associated with warmer winters and on vegetation change and increased insect harassment caused by warmer summers. At the Grotte XVI archaeological site (Dordogne, southwestern France), episodes of summer warming between about 36,000 and 12,000 radiocarbon years ago appear to be associated with lowered relative abundances of reindeer. As the Pleistocene ended and summer temperatures climbed higher, reindeer were extirpated from southern France. A similar phenomenon appears to have occurred here during the prior Eemian interglacial. These records suggest that increased summer temperatures under conditions of global warming may have a direct negative effect on reindeer and caribou populations, including a northward displacement of their southern distributional boundary.     

Life history benefits of residency in a partially migrating pond-breeding amphibian

Species with partial migration, where a portion of a population migrates and the other remains residential, provide the opportunity to evaluate conditions for migration and test mechanisms influencing migratory decisions. We conducted a five-year study of two populations of red-spotted newts (Notophthalmus viridescens), composed of individuals that either remain as residents in the breeding pond over the winter or migrate to the terrestrial habitat. We used multistate mark-recapture methods to (1) test for differences in survival probability between migrants and residents, (2) determine if migrants breed every year or skip opportunities for reproduction, and (3) estimate the frequency of individuals switching migratory tactic. We used estimates of life history parameters from the natural populations in combination with previous experimental work to evaluate processes maintaining partial migration at the population level and to assess mechanisms influencing the decision to migrate. Based on capture-recapture information on over 3000 individuals, we found that newts can switch migratory tactics over their lifetime. We conclude that migrants and residents coexist through conditional asymmetries, with residents having higher fitness and inferior individuals adopting the migrant tactic. We found that newts are more likely to switch from residency to migrating than the reverse and males were more likely to remain as residents. Migration differences between the sexes are likely driven by reproduction benefits of residency for males and high energetic costs of breeding resulting in lower breeding frequencies for females. Environmental conditions also influence partial migration within a population; we found support for density-dependent processes in the pond strongly influencing the probability of migrating. Our work illustrates how migration can be influenced by a complex range of individual and environmental factors and enhances our understanding of the conditions necessary for the evolution and maintenance of partial migration within populations.     

Role of temperature in triggering migratory behavior of the adult tautog Tautoga onitis under laboratory conditions

Adult tautog Tautoga onitis, a marine temperate-water labrid, migrate each fall from inshore summering grounds to offshore overwintering sites. In a previous laboratory study, when adults were held under natural fall-winter cycles of temperature and photoperiod, changes in behavior, indicative of migration, occurred at the same time as their fall migration in nature. In the present study when fish were held under natural photoperiod in summer but with decreasing fall-winter temperature, while highly variable and transient components of migratory behavior were observed, no sustained trend was evident. Then, in the fall, under a naturally changing photoperiod but at simulated summer temperature, no trace of migratory behavior was observed. These findings indicate that temperature appears to play a leading role in triggering the fall migration of the species. The significance of these results and their ecological implications is discussed.     

Phenotypic differentiation in female preference related to geographic variation in male predation risk in the Trinidad guppy (Poecilia reticulata)

Summary Populations of the Trinidad guppy range from areas with high levels of predation by other species of fish to areas with little or no piscine predation. Previous studies have shown that variation among populations in male coloration can be explained by a balance between female preference for brighter males and natural selection against bright males. High levels of male courtship activity may also increase both predation risk and mating success. Therefore, in high-predation areas, females that mate with males that court frequently as well as those that choose bright males would presumably produce male offspring with low survivorship. Consistent with this variation in natural selection, we observed that females from high-predation populations were less likely to choose bright and frequently courting males than females from low-predation populations. This result supports the hypothesis that female preference is evolving as a character genetically correlated with the preferred male character, in which case higher levels of natural selection acting against the male character should be related to lower levels of female preference.     

Weather effects on avian breeding performance and implications of climate change

The influence of recent climate change on the world's biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting (Calamospiza melanocorys) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation; yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 3 degrees C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.     

Testis size variation in the greenfinch Carduelis chloris : relevance for some recent models of sexual selection

Interspecific evidence that testis size responds to selection caused by sperm competition has been obtained from many taxa. However, little is known about the sources of intraspecific variation in testis size, although such variation may have functional significance. Variation in testis size and asymmetry was studied within and between eight geographically separated (and genetically differentiated) populations of greenfinches Carduelis chloris. The relationships between testis size and plumage brightness (degree of yellowness) and the prevalence of haematozoan infections were also investigated in three of these populations, as they related to the predictions of the immunocompetence handicap hypothesis, and Møller's hypothesis relating directional testis asymmetry to phenotypic quality. There were large differences between populations in testis size, with males from northern populations having larger testes than those from southern populations. Within populations, large testes were associated with larger body size and greater age. When the influence of these factors was removed statistically, males with large testes were more likely to be infected with haematozoan parasites, and had brighter yellow plumage. No evidence was found that directional asymmetry in testis size was related to either of these measures of phenotypic quality. These results are consistent with the hypothesis that males with large testes, while signalling higher phenotypic quality as revealed by increased plumage brightness, also pay a cost in terms of reduced immunocompetence, revealed by the increased probability of infection in these males. That these patterns were similar in three different populations adds further strength to these conclusions. Our results suggest that studying the sources of variation in testis size among individuals can reveal interesting processes in sexual selection.     

Assessment of the Conservation Efforts to Prevent Extinction of the Iberian Lynx

Abstract: The Iberian lynx (Lynx pardinus) may be the first charismatic felid to become extinct in a high‐income country, despite decades of study and much data that show extinction is highly probable. The International Union for Conservation of Nature categorizes it as critically endangered; about 200 free‐ranging individuals remain in two populations in southern Spain. Conservation measures aimed at averting extirpation have been extensively undertaken with 4 of the former 10 Iberian lynx populations recorded 25 years ago. Two of the four populations have been extirpated. The number of individuals in the third population have declined by 83%, and in the fourth the probability of extirpation has increased from 34% to 95%. Major drivers of the pending extinction are the small areas to which conservation measures have been applied; lack of incorporation of evidence‐based conservation, scientific monitoring, and adaptive management into conservation efforts; a lack of continuity in recovery efforts, and distrust by conservation agencies of scientific information. In contrast to situations in which conservation and economic objectives conflict, in the case of the Iberian lynx all stakeholders desire the species to be conserved.     

Female-female competition is influenced by forehead patch expression in pied flycatcher females

There is increasing evidence that sexual selection operates in females and not only in males. However, the function of female signals in intrasexual competition has been little studied in species with conventional sex roles. In the Iberian populations of the pied flycatcher (Ficedula hypoleuca), some females express a white forehead patch, a trait that in other European populations, only males exhibit and has become a classical example in studies of sexual selection. Here, we investigated whether the expression of this trait plays a role in female-female competition during early breeding stages. To test this hypothesis, we simulated territorial intrusions by challenging resident females with stuffed female decoys expressing or not a forehead patch. We found that resident females directed more attacks per trial and maintained closer distances to non-patched decoys than to patched ones. Also, patched females were more likely to attack the decoy than non-patched females. Interestingly, females were more aggressive against the decoys when their mate was absent. This may indicate that females relax territory vigilance in the presence of their mate or that males interfere in the interaction between competing females. The behavior of resident males was also observed, although it was not affected by decoy’s patch expression. Our findings suggest that the forehead patch plays a role in female intrasexual competition. If the forehead patch signals fighting ability, as it does in males, we may interpret that non-patched females probably avoided repeating costly agonistic encounters with the most dominant rivals.     

Annual migrations,diving behavior,and thermal biology of Atlantic bluefin tuna, <Emphasis Type="Italic">Thunnus thynnus</Emphasis>, on their Gulf of Mexico breeding grounds

Electronic tags were used to examine the biology of Atlantic bluefin tuna (Thunnus thynnus L.) on their breeding grounds in the Gulf of Mexico (GOM). The hypothesis that movement patterns, diving behavior, and thermal biology change during different stages of the breeding migration was tested. Mature Atlantic bluefin tuna tagged in the western Atlantic and the GOM, were on their breeding grounds from February to June for an average of 39 ± 11 days. The bluefin tuna experienced significantly warmer mean sea surface temperatures (SSTs) within the GOM (26.4 ± 1.6°C) than outside the GOM (20.2 ± 1.9°C). As the bluefin tuna entered and exited the GOM, the fish dove to daily maximum depths of 568 ± 50 and 580 ± 144 m, respectively, and exhibited directed movement paths to and from the localized breeding areas. During the putative breeding phase, the bluefin tuna had significantly shallower daily maximum depths (203 ± 76 m), and exhibited shallow oscillatory dives during the night. The movement paths of the bluefin tuna during the breeding phase were significantly more residential and sinuous. The heat transfer coefficients (K) were calculated for a bluefin tuna in the GOM using the recorded ambient and body temperatures. The K for this fish increased rapidly at the high ambient temperatures encountered in the GOM, and was significantly higher at night in the breeding phase when the fish was exhibiting shallow oscillatory dives. This suggests that the fish were behaviorally and physiologically thermoregulating in the Gulf of Mexico. This study demonstrates that the movement patterns, diving behavior, and thermal biology of Atlantic bluefin tuna change significantly at different stages of the breeding migration and can be used to define spawning location and timing. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Breeding of Euterpina acutifrons,a harpacticid copepod,with special reference to dimorphic males

The breeding of Euterpina acutifrons, a marine harpacticid copepod, was studied both in the laboratory and in the natural population round Anglesey, England, with special reference to dimorphic males. As a result of successful rearing of the species under laboratory conditions, rates of development at different temperatures, and survival and maturation of all types of adults were studied. The dimorphic males developed faster than the females, and the small males had a shorter development time than the larger individuals. Laboratory observations clearly indicated that the males differ markedly in sexual behaviour, the small male being more potent, readily copulating with females, and always very active in breeding. In natural populations, temperature was found to have a limiting influence on the breeding period. Breeding began when the temperature reached 15° to 16°C, and ceased when it fell to 8°C. The most significant feature was the production of small males in nature; they increased in number with the onset of breeding, and always formed a high proportion of the actively breeding population. Both laboratory and field investigations have shown that the small male is a breeding form, and is apparently an adaptation of the species for carrying out successful breeding in the colder areas of its distribution.     

Vigilance in wild Thomas's langurs (Presbytis thomasi ): the importance of infanticide risk

This study examines vigilance as a behavioural indicator of the importance of infanticide risk by comparing the infanticide avoidance hypothesis with the predation avoidance and mate defence hypotheses for wild Thomas's langurs (Presbytis thomasi) in Sumatra. We found that all individuals were more vigilant in situations of high predation risk, i.e. lower in the trees and in the absence of neighbours. Females were also more vigilant on the periphery of the group. However, there were variations in vigilance levels that could not be accounted for by the predation avoidance hypothesis. Males without infants showed higher levels of vigilance in areas of home range overlap than in non-overlap areas during the early phase of their tenure, strongly suggesting mate defence. In these areas of home range overlap where Thomas's langur groups can interact, males may attack females and infants, and so the infanticide risk for males and females with infants is likely to be high in these areas. Only females with infants, but not males with infants or females without infants, showed higher vigilance levels in overlap areas than in non-overlap areas; in addition, in overlap areas, females with an infant were more vigilant than females without an infant, while this was not the case in non-overlap areas. Both females and males with infants were more vigilant high in the trees than at medium heights in overlap areas but not elsewhere. These findings can only be explained by the infanticide avoidance hypothesis. In contrast to predator attacks, infanticidal male attacks come from high in the canopy, and only occur in overlap areas. There was a significant sex difference in vigilance, but males were only more vigilant than females without an infant, and not more vigilant than females with an infant. We conclude that vigilance varied mainly in relation to the risk of predation and infanticide. Mate competition only played a role for males during the early phase of their tenure. Predation risk seems to offer the best explanation for vigilance for all individuals in the absence of infants. Both predation risk and infanticide risk played a role for females and males with infants. Received: 4 April 1998 / Accepted after revision: 6 September 1998