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Autumn Migration Speed of Juvenile Reed and Sedge Warblers in Relation to Date and Fat Loads’

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Autumn Migration Speed of Juvenile Reed and Sedge Warblers in Relation to Date and Fat Loads’ SHORT COMMUNICATIONS 153 The Condor 101:153-156 0 The Cooper Ornithological Society 1999 AUTUMN MIGRATION SPEED OF JUVENILE REED AND SEDGE WARBLERS IN RELATION TO DATE AND FAT LOADS’ STAFFAN BENSCH Department of Animal Ecology, Ecology Building, Lund Universit;v, S-223 62 Lund, Sweden. e-mail: [email protected] Bo NIELSEN Linderotksgatan 28, 703 67 iirebro, Sweden Abstract: We analyzed speed of migration in two fattening, and the quality of these fattening sites do congeneric warblers, the Reed and the Sedge Warbler, not deteriorate in the course of the season as much as Acrocepkalus scirpaceus and A. sckoenobaenus. Sedge is the case for the aphid sites used by Sedge Warblers Warblers migrated at a higher speed than Reed War- (Bibby and Green 1981). Because Sedge Warblers blers. The two species showed similar rates of fat dep- seem more time stressed, we expected them to migrate osition at our Swedish study site, although Sedge War- at a higher speed than Reed Warblers. blers departed with lower fat loads. The higher speed Apart from the difference in diet, the two species of migration in Sedge Warblers and their lower depar- exhibit other ecological differences that might be ture fat loads suggest that they encounter stopover sites linked to different migratory adaptations: Sedge War- which offer higher relative fat deposition rates farther blers lay more eggs per clutch (Bibby 1978) and have south. The amount of visible fat at the time of banding a shorter life-snan (Peach et al. 1990) than Reed War- was positively related to the speed of migration. Esti- blers. The two’studies that used banding recoveries did mates of speed of migration for the two species sug- not confirm statistically that Sedge Warblers migrated gest that the recoveries were situated on average 76- at a higher speed than Reed Warblers, although both 111 km farther south per increase in fat score, corre- studies reported higher average migration speeds in sponding to 58-85% of the expected distance a bird Sedge Warblers (Hildtn and Saurola 1982, Ellcgren can cover by using the fuel of one unit of fat score. 1993). Here, we use banding recoveries and employ mul- Key words: Acrocephalus schoenobaenus, Acro- tiple regression analyses to examine whether Reed cephalus scirpaceus, fat loads, Reed Warbler, ringing Warblers and Sedge Warblers show different speeds of recoveries, Sedge Warbler, speed of migration. migration. In contrast to earlier studies, WC USCa more homogeneous data set because we limit the analyses For migratory birds, the speed of migration is set by to juvenile birds banded at one site in south central the total time it takes to cover the distance between Sweden. We also examine whether fat load at the time summer and winter destinations, including stopover of banding is correlated with the speed of migration. time. Speed of migration is expected to vary between different ecological situations depending upon food METHODS availability and whether the bird is minimizing the We used banding recoveries of birds banded at Kvis- time, total energy expenditure, cost of transport, or risk mare Bird Observatory (59”10’N, 15”25’E) between of predation (Lindstriim and Alerstam 1992). 1982 and 1995. Each year the banding was conducted Reed Warblers (Acrocepkalus scirpaceus) and between late June and late September at two to four Sedge Warblers (A. sckoenobaenus) are two closely different sites in the Kvismaren-Hjalmaren area (max- related species with similar morphology and geograph- imum distance between sites 15 km). Birds were aged ical distribution of both breeding and winter quarters according to Svensson (1992). The amount of visible (Cramp 1992). It has been suggested that Sedge War- fat in the furcular cavity and belly was scored on a blers migrate from Northern Europe at a higher speed scale between 0 and 6 (Pettersson and Hasselquist than Reed Warblers in order to reach important sites 1985). for fattening (Bibby and Green 1981). At these sites, In total, 20,661 juvenile Reed Warblers and 8,354 situated north of the Mediterranean Sea, high abun- juvenile Sedge Warblers were banded. Our banding ef- dance of reed aphids enable Sedge Warblers to build fort resulted in 111 and 36 recoveries, respectively, up fat stores sufficient for nonstop flights to tropical during the birds’ first autumn migration through Eu- Africa (Bibby and Green 1981). In contrast, Reed War- rope. In order to exclude birds not yet migrating, we blers, which feed on Diptera, use areas closer to the only included birds recovered south of the banding Sahara including southern Europe and north Africa for site, at a distance of at least 50 km and with a migra- tion speed exceeding 10 km day-’ (Hilden and Saurola 1982, Ellegren 1993). We also restrict analyses to birds I Received 29 December 1997. Accepted 24 Sep- that were scored for fat loads at the time of banding. tember 1998. This reduced data set consists of 85 Reed Warbler and 154 SHORT COMMUNICATIONS 3 1 Sedge Warblers. Most recoveries (95%) involve re- vation probably arises due to two circumstances capturesby other banders. (Fransson 1995). Many birds banded early in the sea- Statistical tests were done in SYSTAT (Wilkinson son may not have startedmigration and will thus con- 1992), and all P-values are two-tailed tests. When we tribute to the initially low recordedspeed of migration. applied multivariate statistics,residuals were approxi- Late in the season,on the other hand, many birds will mately normally distributed. We used distance from be capturedwith filled fat loads, and if recoveredsoon the banding site as the dependentvariable and date of after banding, they may have covered a considerable banding and recapture as independent variables. Re- distance. The recorded asymptotic speed of migration sidual distance should hence be analogousto speed of will be reached when one or several full migratory migration. We chose to use distance as the dependent episodes (stopover time plus migratory flight) have variable rather than speed becausethe latter is a ratio been completed. (distance/time) showing nonlinear relations with both The multiple regressionmodel described above can banding and recapturedate. If not otherwise stated,we be used to control for the effect of banding and recap- report mean ? SD. ture date on how far south the birds were recovered (residual distance). By adding fat score to this model, RESULTS we can examine whether the amount of fuel the birds The average speed of migration was 55.3 2 30.7 km carry at the time of banding may further explain the day-’ for Sedge Warblers (n = 31) and 38.8 ? 24.4 distance to the recovery site. For Reed Warblers, fat km day-’ for Reed Warblers (n = 85), and the differ- score significantly increased(P = 0.03) the proportion ence is significant (ANOVA, F, ,,4 = 8.94, P < 0.01). of variance explained (R* = 0.47 for the model in- The two species did not differ ‘in date of banding (7 cluding fat score). The slope of the regression(mean versus 11 August, t-test, t,,, = 1.24, P > 0.2) or in the 2 SE) suggeststhat the distance to the recovery site number of days elapsed between banding and recap- increasedby 76.0 + 34.1 km per fat score. Similarly, ture (20.1 +- 8.7 versus 20.5 2 9.4 days, t,,, = 0.19, when we added fat score of Sedge Warblers to the P > 0.5). However, the distance to the recovery site multiple regressionmodel, it tended (P = 0.07) to in- differed between species (Sedge Warblers: 1,078 2 crease the explanation of the variance in distance (R* 607 km versus 770 5 519 km for Reed Warblers, t,,, = 0.59 for the model including fat score). The slope suggeststhat the birds were recovered 110.9 + 59.3 = 2.71, P < 0.01). For both Sedge and Reed Warblers, we found that km farther south per fat score. the date of banding and the date of recovery had sig- DISCUSSION nificant effects on the distanceto the recovery site (all Ps < 0.001, based on multiple regressionanalysis). In SPEEDOF MIGRATION neither of the specieswas the interactionterm between The present study confirms the suggestionthat Sedge date of banding and date of recovery significant, sug- Warblers migrate at a higher speedthan Reed Warblers gesting that the effect of recovery date on distance is (Bibby and Green 1981). The reason why Ellegren the same, independentof when the banding was done, (1993) did not find a significant difference in speed of and vice versa. migration between the two specieseven though he an- In Reed Warblers, the distance to the recovery site alyzed a larger data set, which partly included the one decreasedby 32.7 km day-’ of banding and increased we examined here, might be that he combined recov- by 37.8 km day-’ of recovery. In other words, for a eries from several banding sites. This possibly increas- given recapturedate, birds will on average be encoun- es the variance of the means, becauselocales may dif- tered 32.7 km closer to the banding site for each day fer in the proportion of birds molting/staging and mi- later in the seasonthey were banded. Similarly, for a grating. given banding date, birds will on average be encoun- In general, the speed of migration increaseswith the tered 37.8 km farther south of the banding site for each migratory distance (Ellegren 1993). At Kvismaren, ju- day later in the seasonthey were recaptured.The cor- venile Sedge Warblers have a more restrictedfirst pre- responding values for Sedge Warblers were 43.4 and basic molt than Reed Warblers which allows them to 63.1 km day-‘, respectively.
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