Migratory orientation of red-eyed vireos,Vireo olivaceus,in relation to energetic condition and ecological context

How and when migrants integrate directional information from different sources may depend not only on the bird’s internal state, including fat stores, but also on the ecological context during passage. We designed experiments to (1) examine the influence of stored fat on the decision to migrate and on the choice of migratory direction and (2) investigate how the integration of orientation cue information is tied to energetic status in relation to migration across an ecological barrier. Migratory orientation of red-eyed vireos (Vireo olivaceus) at twilight was recorded using two different techniques, orientation cage experiments and free-flight release tests, during both fall and spring migration. During fall migration, the amount of stored fat proved decisive for directional selections of the vireos. Fat birds chose directions in accordance with migration across the Gulf of Mexico. Lean birds oriented either parallel to the coast line (cage tests) or moved inland (free-flight releases). Whereas only fat birds showed significant responses to experimental deflections of the geomagnetic field, lean birds displayed a tendency to shift their activity in the expected direction, making it difficult to evaluate the prediction that use of the magnetic compass is context dependent. Fat loads also had a significant effect on the decision to migrate, i.e., fat individuals were more likely to embark on migration than were lean birds (true for both cage and release experiments). During spring migration, a majority of experimental subjects were classified as lean, following their arrival after crossing the Gulf of Mexico, and oriented in seasonally appropriate directions. The vireos also showed significant responses to experimental deflections of the geomagnetic field regardless of their energetic status. Free-flight release experiments during spring migration revealed a significant difference in mean directions between clear sky and overcast tests. The difference may indicate a compensatory response to wind drift or possibly a need for celestial cues to calibrate the magnetic compass. Finally, this is the first demonstration of magnetic compass orientation in a North American vireo. Received: 15 December 1995/Accepted after revision: 24 March 1996     

Pigeon homing: sun compass use in the southern hemisphere

Although the sun compass of birds is based on learning the sun's arc during development, it was unclear whether birds can use the sun when its apparent movement is reversed, in particular, whether northern birds that have been introduced into the southern hemisphere can use the southern sun. To answer this question, clock-shift experiments were performed with local homing pigeons in Auckland, New Zealand (37°S). In three fast-shift tests and two slow-shift tests, the experimental birds showed deflections from the untreated controls that were the mirror images of those observed in the northern hemisphere. These results clearly show that homing pigeons in New Zealand use a sun compass that is adapted to the situation in the southern hemisphere. The learning processes establishing the compensation mechanisms thus appear to be free of constraints concerning the direction of the sun's movement. Differences from recent findings with migratory birds, where the direction of celestial rotation proved of crucial importance for establishing the migratory direction, are discussed: the differences may arise from the different orientation tasks, in particular, from the involvement of innate information in establishing the migratory direction. Received: 13 November 1997 / Accepted after revision: 28 February 1998     

Autumn migratory fuelling: a response to simulated magnetic displacements in juvenile wheatears, Oenanthe oenanthe

Recent experiments exposing migratory birds to altered magnetic fields simulating geographical displacements have shown that the geomagnetic field acts as an external cue affecting migratory fuelling behaviour. This is the first study investigating fuel deposition in relation to geomagnetic cues in long-distance migrants using the western passage of the Mediterranean region. Juvenile wheatears (Oenanthe oenanthe) were exposed to a magnetically simulated autumn migration from southern Sweden to West Africa. Birds displaced parallel to the west of their natural migration route, simulating an unnatural flight over the Atlantic Ocean, increased their fuel deposition compared to birds experiencing a simulated migration along the natural route. These birds, on the other hand, showed relatively low fuel loads in agreement with earlier data on wheatears trapped during stopover. The experimental displacement to the west, corresponding to novel sites in the Atlantic Ocean, led to a simulated longer distance to the wintering area, probably explaining the observed larger fuel loads. Our data verify previous results suggesting that migratory birds use geomagnetic cues for fuelling decisions and, for the first time, show that birds, on their first migration, can use geomagnetic cues to compensate for a displacement outside their normal migratory route, by adjusting fuel deposition.     

Effect of modified magnetic field on the ocean migration of maturing chum salmon, Oncorhynchus keta

To investigate the role of magnetic compass orientation in oceanic migrating chum salmon, Oncorhynchus keta, an ultrasonic telemetry study was carried out in the western North Pacific off the coast of Kushiro, Hokkaido. Four salmon were fitted with a tag which generated an artificial magnetic field and modified the geomagnetic field around the head of the fish. Initially, the free-ranging salmon with stomach-implanted ultrasonic transmitters were tracked for a period of several hours before the magnetic field was altered for a period of 16 h. The generator produced an alternating magnetic field intensity of about 6 gauss, with polarity which reversed every 11.25 min. There was no observable effect on the horizontal and vertical movements of the salmon when the magnetic field was modified. However, it was noted that salmon slowed their swimming speed significantly before changing direction, regardless of whether the fish were swimming under the normal geomagnetic field or whether they were swimming under the modified field. Received: 6 April 1997 / Accepted: 29 April 1997     

Orientation of the pied flycatcher Ficedula hypoleuca: cue-conflict experiments during spring migration

Hierarchical relationships among different compass systems in long-distance migrants are still a matter for discussion because different studies have led to highly variable and apparently contradictory results. We carried out cue-conflict experiments during spring migration on pied flycatchers Ficedula hypoleuca (Passeriformes, Muscicapidae). Birds were exposed to a conflict between celestial and magnetic information by altering the polarized light pattern or magnetic field. The polarization pattern was shifted (±90°) with filters, whereas the magnetic field was altered (+90°) through Helmholtz coils. Birds were tested in modified Emlen funnels both before and after the cue conflict; during the tests, only the natural magnetic field was available. This protocol was designed to test whether the experimental birds recalibrated their magnetic compass on the directional information derived from the light polarization pattern when the region near the horizon was visible during the conflict. Contrary to this expectation, we did not record any significant shift in magnetic orientation after one or repeated exposures to the cue conflict. Our results support earlier studies, which suggest that the magnetic field is the primary compass cue during the migratory period.     

Night sky orientation of migratory pied flycatchers raised in different magnetic fields

Summary To test whether the initial night sky orientation response of migratory pied flycatchers (Ficedula hypoleuca) is calibrated from the ambient magnetic field experienced by birds during their first summer, three groups of pied flycatchers were hand-reared and then held under different magnetic field conditions during the course of the summer. All groups were held outdoors and given full exposure to the day and night sky. One group was exposed to the local earth's magnetic field. A second group was exposed to a magnetic field of local earth strength, local earth inclination shifted 105° counter-clockwise relative to the local earth's field. The last group was exposed to a vertical, and thus nondirectional magnetic field.In autumn, the birds were tested for their orientation under the night sky in the absence of a directional magnetic field. When tested, all three groups were oriented with mean directions varying from south to southeast. No statistical differences emerged in any between group comparisons. The data indicate that earth's magnetism does not serve as a calibrating reference in the development of a pied flycatcher's initial orientation response to the night sky.     

Calibration of the sun compass by sunset polarized light patterns in a migratory bird

Summary Migrating birds derive compass information from the sun, stars, geomagnetic field and polarized light, but relatively little is known about how these multiple sources of directional information are integrated into a functional orientation system. We found that migratory warblers exposed to a rotated polarized light pattern at sunset oriented at a constant angle to the axis of polarization. When polarized light cues were eliminated, this shifted orientation was maintained relative to the setting sun. Polarized light patterns, thus, appear to provide a calibration reference for the sun compass in nocturnal migrants, and may also play a role in calibrating other compass systems. Correspondence to: J.B. Phillips     

Magnetic compass mediates nocturnal homing by the alpine newt, Triturus alpestris

Experiments were carried out to investigate the use of magnetic compass cues in the nocturnal homing orientation of the alpine newt Triturus alpestris. Tests were carried out at a site 9 km to the east–northeast of the breeding pond. Newts were tested at night in an outdoor circular arena that provided an unimpeded view of celestial cues, in one of four symmetrical alignments of an earth-strength magnetic field. In tests carried out under partly cloudy skies newts exhibited homeward magnetic compass orientation. Because the moon was visible in some trials, but obscured by clouds in others, we investigated whether the presence of the moon contributed to the scatter in the distribution of magnetic bearings. When the moon was visible, the distribution of magnetic bearings was more scattered than when the moon was obscured by clouds, although in neither case was the distribution significant due, in part, to the small sample sizes. Moreover, when the moon was visible, newts oriented along a bimodal axis perpendicular to the moon azimuth, suggesting that the presence of the moon may have affected the newts behavior. To provide a more rigorous test of the role of magnetic compass cues when celestial cues were unavailable, nocturnal tests were carried out during the following migratory season under total overcast. In the absence of celestial compass cues, the distribution of magnetic bearings exhibited highly significant orientation in the homeward direction. These findings indicate that newts are able to orient in the homeward direction at night using the magnetic compass as the sole source of directional information. Moon light altered the newts behavior. However, this apparently resulted from the asymmetrical distribution of moon light in the testing arena, rather than the use of an alternative compass.     

Clock-shift experiments with Savannah sparrows, Passerculus sandwichensis, at high northern latitudes

Orientation can be difficult for nocturnal bird migrants at high northern latitudes because of the large changes of magnetic declinations, rapid longitudinal time-shifts experienced during a long-distance flight and the invisibility of stars during the polar summer. Both sunset cues as well as geomagnetic cues have been shown to be of great importance in the orientation system of Savannah sparrows, Passerculus sandwichensis. We used clock-shift experiments to investigate whether geomagnetic and sunset cues were used for migratory orientation by wild-caught young Savannah sparrows at high geomagnetic latitudes in Northern Canada. We exposed birds to a 4-h slow clock-shift, expecting a 60° clock-wise shift in orientation after the treatment. Under natural clear skies in the local geomagnetic field, the birds responded by showing a significant axial mean orientation directed towards the position of the setting sun in the NW and towards their preferred migratory direction in the SE. After exposure to the clock-shift for 6 days and nights the birds showed a clear response to the treatment and shifted significantly towards NNE. Birds that first oriented towards NW in the experiments before clock-shift tended to shift clock-wise, thus reacted to the clock-shift in the expected way. The reaction of the individual birds that originally oriented towards SE seems to vary. In summary, our birds did not select a constant angle (menotaxis) in relation to the sun's position during the experiments, but presumably were affected by the sun showing phototaxis or followed their magnetic compass. Possible explanations of the unexpected experimental results are discussed. Electronic Publication     

Exaggerated orientation scatter of nocturnal passerine migrants close to breeding grounds: comparisons between seasons and latitudes

Using tracking radars, we investigated the variability of flight directions of long-distance nocturnal passerine migrants across seasons (spring versus autumn migration) and sites at the southern (56° N) and northern (68° N) ends of the Scandinavian Peninsula (Lund versus Abisko). Whilst most migrants at Lund are on passage to and from breeding sites in Fennoscandia, the majority of the migrants at Abisko are close to their breeding sites, and migration at Abisko thus to a large degree reflects initial departure from breeding sites (autumn) or final approach to breeding destinations (spring). The radar data were used to test predictions about differences in orientation and wind drift effects between adult and juvenile birds (a large proportion of autumn migrants consists of juvenile birds on their first journey), between situations far away from or near the goals and between different phases of migration (initial departure, en route passage, final approach to goal). The concentrations (both total and within-night concentrations) of flight directions differed significantly between seasons as well as sites, with the highest concentration at Lund in spring (mean vector length of track directions, r = 0.79) and lowest at Abisko during spring (r = 0.35). Partial wind drift and partial compensation were recorded at Lund, with a similar effect size in spring and autumn, whilst possible wind drift effects at Abisko were obscured by the large directional scatter at this site. The results from Lund support the prediction that the high proportion of juveniles in autumn contributes to increase the directional scatter during this season, whilst there was no support for predictions of differential wind drift effects between seasons and situations with different goal distances. The most striking and surprising result was the exceedingly large scatter of flight directions at Abisko, particularly in spring. We suggest that such an exaggerated scatter may be associated with final approach orientation, where migrants reach their specific goals from all various directions by final navigation within a more wide-ranging goal region. The larger scatter of autumn flight directions at Abisko compared to Lund may be due to exploratory flights in variable directions being more common at initial departure from breeding sites than later during migratory passage. These surprising results highlight the importance of studying and analysing orientation during final approach to (and initial departure from) migratory goals for understanding the orientation systems of migratory birds.     

Magnetic compass orientation in the yellow-faced honeyeater,Lichenostomus chrysops,a day migrating bird from Australia

Summary In Australia, the southern populations of the yellow-faced honeyeater, Lichenostomus chrysops (Meliphagidae), perform annual migrations, with routes following the eastern coastline. In order to assess the role of magnetic cues in the migratory orientation of this diurnal migrant, its directional behaviour was recorded in recording cages under natural and experimentally manipulated magnetic-field conditions. During autumn the birds tested indoors in the local geomagnetic field showed a directional change from north initially to northwest later in the season (Fig. 1 a, b), which corresponds well with the general pattern of movement of this species in the field. Deflecting magnetic north to ESE resulted in a clockwise shift of the mean direction by 77° and 71°, respectively (Fig. 1 c, d), while no significant directional tendencies were observed in a magnetic field with a compensated horizontal component (Fig. 1 e, f; see Table 1). In outdoor tests in spring, the birds preferred southerly directions when tested in the local geo-magnetic field. In a magnetic field with a reversed vertical component (i.e. with an inclination pointing down instead of upwards) the birds reversed their directional tendencies and oriented northward (Fig. 2, Table 2). These results clearly show: (1) that yellow-faced honeyeaters can use the magnetic field for direction finding, and (2) that their magnetic compass functions as an inclination compass, as has been shown for several holarctic migrants.Correspondence to: W. Wiltschko     

Orientation of juvenile barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) tested in Emlen funnels during autumn migration

Although hirundines have been used extensively in homing experiments, to date no investigation of their migratory orientation has been carried out, despite the well-known migratory habits of many species of this family. This paper reports on a study of the orientation of the barn swallow (Hirundo rustica), a typical diurnal trans-Saharan migrant. Modified Emlen funnels were used to verify the suitability of this species for cage experiments and investigate the role of visual and magnetic cues during the birds first migratory journey. Juvenile swallows were mist-netted at a roost site in central Italy and then tested in a site 19 km apart. Orientation experiments were performed under four experimental conditions: natural clear sky and simulated overcast, in both local and shifted magnetic fields (magnetic North=geographical West). Under clear sky, the swallows tended to orient phototactically toward the best-lit part of the funnel and failed to respond to the magnetic field shift. Under overcast conditions, they oriented northward and modified their directional choices as expected in response to the shifted magnetic North. On the whole, our data indicate that swallows can use magnetic information for compass orientation. Possible explanations for the northward orientation of birds tested under overcast conditions are discussed.Communicated by W. Wiltschko     

Sunset orientation of robins,Erithacus rubecula,with different fields of sky vision

Summary Migratory orientation of robins (Erithacus rubecula) at sunset was recorded using orientation cages, under clear autumn skies. The aim of the experiments was to examine the importance of different visible sky sections for the orientation of robins. I obtained the following results: (1) Robins tested with the visible sky section limited to 90° around zenith (45° above the horizon) showed a mean orientation that coincided with the average sunset azimuth, with little scatter around the mean angle (Fig. 2). (2) When the birds were allowed a more extensive field of sky vision (maximum 160°), they chose headings on an approximate north-south axis, significantly different from tests with a restricted view of the sky (Fig. 3). (3) Experiments were also performed in which the response of robins to a mirror deflection (about 120° counterclockwise) of visual cues in the lower parts of the sunset sky was examined. The outcome indicated that visual information in the lower part of the sky may be critical for the orientation of robins (Fig. 5). These results, together with recent findings that robins captured and tested at two nearby sites show distinctly different orientation behavior in relation to experimental manipulations of the magnetic field, suggest that priorities among orientation cues may differ depending on the migratory situation encountered.     

The magnetic compass mechanism of birds and its possible association with the shifting course directions of migrants

Summary Many birds of the northern hemisphere shift their migratory course to more southerly directions when moving from northern to southern latitudes. Birds from Central Europe, for example, change their course from SW to S or from SE to S respectively (Fig. 1). This also seems to apply to some other animals.The hypothesis presented here explains the observed shifts in migratory direction on the basis of changes in the parameters of the earth's magnetic field and hence would make a genetic fixation of shifts in the migratory direction unnecessary.To determine the direction of migration birds do not refer to the polarity of the magnetic field but to its dip (=). According to the hypothesis presented here, the birds, however, do not refer to the direction of dip as previously believed but to the individual apparent angle of dip (=), this angle changes depending on the heading of the bird (see Fig. 3 and Eq. 1). Maintaining a species specific or population specific the bird will move in its predetermined migratory direction. Changes in the dip of the earth's magnetic field correspond to changes in latitude. According to the hypothesis with fixed, the migratory direction will change when the dip changes. Given the hypothesis and the parameters of the earth's field theoretical migratory paths of birds between summer and winter quarters may be calculated (Figs. 8–11). The calculated tracks and the actually observed migratory routes agree well. This is also confirmed by radar and other observations of migratory directions in areas of different dip angles (Fig. 13). Displacing migrating birds to areas of smaller dip angles (= lower magnetic latitudes) results in predeterminable shifts in the birds migratory direction (Figs. 5, 6). The hypothesis also accounts for the so far unexplained orientation behaviour of transequatorial migrants under the magnetic equator.A very simple model of this hypothetical compass mechanism may be based on the assumption of the sensor axis is supposed to correspond to the apparent angle of dip when moving in the migratory direction. In this position the difference between the apparent angle of dip and the angle of the sensor is zero. Any change in the direction of movement, however, will result in a difference leading to a response of an assumed receptor. When maintaining the zero difference the bird invariably sticks to its migratory course. The proposed mechanism is a null instrument unaffected by changes in field intensity and not depending on the measurement of absolute values.     

Geomagnetic field affects spring migratory direction in a long distance migrant

Night-migrating song birds travel to and from their wintering and breeding areas often separated thousands of kilometers apart and are clearly capable of finding intended goal areas from a distant location. Displacement experiments provide a useful way to highlight orientation and navigational skills in migrants. To investigate which cues birds actually use to compensate for displacement and the exact mechanism of each cue, experiments with manipulation of single cues are required. We conducted a simulated displacement of lesser whitethroats Sylvia curruca on spring migration. Birds were displaced not geographically but in geomagnetic space only, north and south of their breeding area to test whether they incorporate information from the geomagnetic field to find their breeding area. Lesser whitethroats held in southeast Sweden but experiencing a simulated displacement north of their breeding area (Norway) failed to show a consistent direction of orientation, whereas birds displaced south of their breeding area (Czech Republic) exhibited consistent northerly orientation, close to the expected seasonally appropriate direction, after displacement toward the trapping location. The absence of a clear compensatory direction in birds displaced north might be due to unfamiliar magnetic information or lack of sufficient information such as a magnetic gradient when moving around. By isolating one orientation cue, the geomagnetic field, we have been able to show that lesser whitethroats might incorporate geomagnetic field information to determine latitude during spring migration.     

Homing behaviour in the sleepy lizard (Tiliqua rugosa): the role of visual cues and the parietal eye

To investigate how visual cues are integrated into a navigational strategy for homing in the Australian sleepy lizard (Tiliqua rugosa), lizards were displaced beyond their home range, either with full access to visual cues or with no access to visual cues during the displacement. Homeward orientation was significantly worse when lizards were denied visual cues during the displacement than when they were not. However when lizards were displaced with their field of view restricted to the sky, their homeward orientation was equally as good as that of lizards displaced with no visual restriction. These experiments suggest that sleepy lizards use celestial cues to determine the compass bearing of the outward journey, and reverse this bearing to orient in the homeward direction (course reversal). In a subsequent experiment, lizards oriented randomly with respect to home when the parietal eye was entirely covered with a patch during the displacement and return, while control lizards fitted with a sham parietal eye patch were well oriented towards home. In both groups, the lateral eyes were unobstructed and had complete access to visual cues including celestial cues and landmarks. These results suggest that the parietal eye plays a highly significant role in sleepy lizard homing, perhaps mediating a sky polarization compass sense.     

Influence of body condition and weather on departures of first-year European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>, from an autumn migratory stopover site

How migratory birds decide when to leave a stopover site is important to the understanding of bird migration strategies. Our study looks at how body condition and the weather affect the decision to depart on nocturnal migratory flight. During two autumn migration seasons (2002–2003), we radio tracked 51 first-year European robins, Erithacus rubecula, at a stopover site on the Courish Spit (Eastern Baltic) from the first day after landing until their migratory departure. The tagged robins stopped over for 1–14 days. There was no clear relationship between stopover duration and energetic condition on arrival. Weather conditions (wind, precipitation, and cloud cover) on departure differed measurably between years. In 2002, robins took off mainly under following winds and clear skies. In 2003, there were mainly light head winds and partially cloudy or overcast skies. This could be explained by the year-specific role of weather factors in making the decision to depart. In both years, robins making short (1–2 days) stopovers took off in more varied weather situations than those individuals with long stopovers. This suggests that robins from the former group were more inclined to continue with migration than longer-stay birds that, apart from re-fuelling, could be waiting for favourable weather. The lack of a relationship between stopover duration and body condition and some departures under unfavourable weather conditions suggest that endogenous spatiotemporal programmes may play an important role in controlling stopover duration in robins.     

Northern magnetic displacements trigger endogenous fuelling responses in a naive bird migrant

In a previous study, we found that juvenile northern wheatears (Oenanthe oenanthe) exposed to a magnetic displacement to the west of their natural migration route increased their body mass. The total intensity and inclination used for the western displacement may also have been interpreted as northern compared to the experimental site (stronger total field intensity and steeper inclination angle). In order to investigate whether the fuelling response was a response to an unexpected magnetic field or specific to the northern magnetic field, we conducted a new experiment. Juvenile wheatears from the same study population were magnetically displaced to southwestern magnetic fields, exposing the birds to unexpected magnetic combinations, but eliminating the possible effect of a northern magnetic field. A control group was kept in the local geomagnetic field in Sweden for comparison. There was no difference in body mass increase between treatments, suggesting that the fuelling response previously found was not a simple response to an unexpected magnetic field, but rather a specific response to the northern magnetic field. Juvenile wheatears may have developed a fuelling response to northern magnetic fields in order to enable a successful flight towards the migration goal.     

Astronomical, landscape and climatic factors influencing oriented movements of Talitrus saltator in nature

 The present study was conducted on a Mediterranean beach (Burano, southern Tuscany, Italy) to examine the timing, orientation, and motivating and directing factors of the spontaneous movements of the sandhopper Talitrus saltator (Montagu, 1808). In April 1994, October 1994 and June 1995, during different moon phases, traps were positioned in the eulittoral zone, which intercepted sandhoppers when walking on the sand surface. At the same time environmental parameters were registered. Contemporaneously, orientation tests were carried out on active individuals using two arenas, one of which permitted a view of both sky and landscape and one which prohibited the landscape view. The results show landward migration after sunset, for juveniles later than for adults, nonoriented activity for 2 or 3 h after midnight and seawards zonal recovery before and after sunrise. The main microclimatic factor modulating activity was sand temperature, while the nonoriented activity between the two migratory movements seems to be endogenously determined. Both sky and landscape cues are used by sandhoppers for orienting their course, but the landscape view is sufficient and necessary on new moon nights. These results contribute to solution of the controversy on the mechanisms actually motivating sandhoppers in nature. Received: 24 September 1996 / Accepted: 25 October 1996     

Song sharing in two populations of song sparrows (Melospiza melodia)

Sharing song types with immediate neighbors is widespread in birds with song repertoires, and sharing songs may confer a selective advantage in some cases. Levels of song sharing vary between different geographical populations of several bird species, and ecological differences often correlate with differences in singing behavior; in particular, males in migratory subspecies often share fewer songs than males in resident subspecies. The song sparrow (Melospiza melodia) appears to fit this pattern: resident song sparrows in western North America generally share 20–40% of their repertoire (of about eight songs) with each neighbor, while migratory subspecies from eastern North America often share 10% or less. We compared song sharing in two populations within a single subspecies of song sparrow (M. m. morphna) in Washington State. These populations, separated by only 120 km, nonetheless differ in migratory tendencies and several other ecological and life history variables. We recorded complete song repertoires from 11 male song sparrows in a high-elevation, migrating population at Gold Creek in west-central Washington, and compared them to two samples (n = 15 and n = 36) from a coastal, resident population at Discovery Park, Seattle, Washington. Despite major differences in habitat, population density, and migratory tendencies, song sharing among Gold Creek males was as high as that among Discovery Park males. In both populations, sharing was highest between immediate neighbors, and declined with distance. We conclude that at the within-subspecies level, neither migration nor population density affect song sharing in song sparrows, a song repertoire species. Received: 26 November 1998 / Received in revised form: 1 May 1999 / Accepted: 29 May 1999