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The Relationship Between Wing Area and Raggedness During Molt in the Willow Warbler and Other Passerines

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The Relationship Between Wing Area and Raggedness During Molt in the Willow Warbler and Other Passerines J. Field Ornithol., 69(1):103-108 THE RELATIONSHIP BETWEEN WING AREA AND RAGGEDNESS DURING MOLT IN THE WILLOW WARBI.ER AND OTHER PASSERINES ANDERS HEDENSTROM Department of Animal Ecology Lund University,Ecology Building S-223 62 Lund, Sweden Abstract.--Molt involvesincreased metabolic cost due to the energy required to synthesize new feathers and increasedcost of thermoregulation becausethe insulation capacityis re- duced. Flight cost is also increasedbecause wing span and wing area are reduced when remigesare missingor growing. However,data on the reduction of wing area during molt (necessaryfor calculatingaerodynamic performance) is lacking.I showhow standardmolt scores(converted into a raggednessscore) can be used to estimate the reduction of wing area due to molt in Willow Warblers (Phylloscopustrochilus) and selectedother passerines. Maximum wing area reduction associatedwith maximum raggednessscore of the birds ex- amined wasapproximately 10%. Wing loadingwill thereforebecome increased during molt and flight performance will be reduced in similar waysas during migratory fattening. LA RELACION ENTRE EL J,REA DEL ALA Y LA DESALII•EZ DURANTE LA MUDA DE PHYLLOSCOPUS TROCHILUS Y DE OTRAS PASERINAS Sinopsis.--Lamuda incluyeaumentos en costosmetab61icos pot la energia requeridapara sintetizar plumas nuevasy al costo de termoregular, porque la capacidadde insulaci6nse reduce. Los costosde volar tambi6n aumentan porque la expansi6ny el largo del ala se reducen cuando las remeras faltan o estrin creciendo. Sin embargo, faltan datos sobre la reducci6ndel •irea del ala (necesariospara poder calcularla ejecuci6naerodin•imica) dur- ante la muda. Aquf muestro como registrosregulares de muda (convertidosen datos de desalifiez) pueden usarsepara estimar la reducci6n del •irea del ala debido a la muda en Phylloscopustrochilus. yen otras avespaserinas seleccionadas. La reducci6n mJ_ximadel •irea del ala asociadacon la mayor desalifiezen las avesexaminadas fu6 de aproximadamenteun 10%. Por lo tanto, el empujedel ala aumentar5durante la muda y la ejecuci6nde vuelose reducir•ien forma similarcomo durante el engordamientomigratorio. Molt in birds is associatedwith elevatedenergy costs because (1) there is an energycost to synthesizenew feathers (reviewedby Lindstr/Smet al. 1993), (2) the insulation capacityof the plumage is reduced during molt which increasesthe energy costsof thermoregulation (Lustick 1970, Payne 1972) and (3) the cost of flight increasesdue to a reduction of wing area and/or wing span (Ginn and Melville 1983). The latter costis difficult to estimatesince current aerodynamictheory for bird flight (e.g., Pennycuick1989) is basedon momentum-jettheory of flight and requires only the wing span as the morphologicalcharacteristic of the wing. How- ever,some researchers studying life-history trade-offs of parent birdsfeed- ing young (reviewedby Mauck and Grubb 1995) and song-flightperfor- mance (Mather and Robertson 1992, Moller 1991) have experimentally reduced the wing area in order to induce elevatedflight costs. Gaps in the wing will affect the lift distribution and hence flight per- formance. Therefore, in addition to the position of gaps it is important to know the relative wing area during molt in order to estimatethe as- sociatedflight costsaccurately, although at present no straightforward 103 104] A. Hedenstr6m J.Field Ornithol. Winter 1998 method is availableof calculatingthese costsunless using so-calledpanel methods (cf. Katz and Plotkin 1991). However, wing areas of birds are rarely reported and wing areas during molt are virtually non-existentin the literature. In the present short paper I show that the standardmolt scoringtechnique may be used to estimate the relative wing area during molt in the Willow Warbler (Phylloscopustrochilus) and a few other pas- serine species. METHODS Passerineswere captured in a subalpine birch forest at Lake TjultrSsk (520 m elevation)near AmmarnSs(65ø58'N, 16ø07'E) in SwedishLapland during their prebasicmolt (July 1992), as part of the LUVRE project (Enemar et al. 1984). Birdswere aged and sexedon the basisof plumage and molt characteristics and, in the Willow Warbler, on the basis of size (Svensson1984). All birds were in their second calendar year or older. Molt in each bird's left wing wasrecorded accordingto Ginn and Melville (1983): old featherswere scored0; new, fully grown featherswere scored 5; missingand growing featherswere scoredbetween I and 4. The out- ermost reduced primary was excluded in the analysisbecause its contri- bution to the total wing area is negligible. By using these molt scores,an estimate of the gaps causedby missingor growing feathers can be rained from the wing raggednessscore introduced by Haukioja (1971). Raggednessis calculatedas 5 minus the molt scorefor missingand grow- ing feathers; old and new (fully grown) feathers receive raggednessscore of 0. The total raggednessscore for one wing can theoreticallyvary be- tween 0 (completewing) and 60 (nine primariesand six secondariesshed simultaneously). I traced the contour of one wing onto paper (cf. Pennycuick1989) in order to obtain the wing area. For aerodynamicpurposes, the area of the body between the wings is included in the total wing area (Pennycuick 1989). Therefore, I measuredthe width of the body betweenthe wings at the centerline of the wing chord (chord is here used in the aeronau- tical sense,i.e. the distancefrom leading to trailing edge of the wing), to the nearest1 mm with Vernier calipers.This measurement,multiplied by the chord of the wing at the wing root gavethe area of the body between the wings.The wing tracingswere later copied onto high quality paper, cut out and the area was determined gravimetricallyby weighing the "wings" on a high precision balance. I also determined the wing area with a planimeter (computer digitizer), and the results from the two methodswere practicallyidentical (r -- 0.99, P < 0.001, n = 41), and so I used the data obtained by the gravimetricmethod in the analyses.The areasof complete wingswere constructedfor each bird in molt, by con- structing the outline of the wing had it been complete on the basisof wing tracingsfor birds of the samespecies which were not in molt. Hence, the relative wing area is the wing area during molt divided by the wing area of the complete wing. Birds are only representedonce in the data set. I also measuredthe wing span to the nearest I mm when the wings Vol.69, No. 1 WingArea and Molt [105 TABLE1. Summarystatistics for each speciesof data usedin analyses:sample size (n), body mass (m), raggednessvalue (R; mean and range), wing area before molt (S), wing area during molt (Smoot),relative wing area (Smoot/S;mean and range) and wing loading during molt (Nmo•t;mean and range). S Smolt Smolt/S Nmolt Species n m (g) R (cm•) (cm2) (%) (Nm -2) Motacillaflava 4 16.4 17 105.2 95.8 91.0 16.8 (8-21) (88.3-93.9) (15.2-19.0) Prunella modularis 3 19.3 10.3 95.7 91.9 96.1 20.7 (8-12) (93.6-98.5) (19.6-21.4) Luscinia svecica 5 18.3 10.6 108.3 102.0 94.1 17.6 (7-15) (90.7-97.0) (16.0-18.6) Phylloscopustrochilus 21 9.1 11.2 78.1 73.5 94.2 12.2 (2-19) (88.6-98.1) (10.3-13.9) Ficedula hypoleuca I 11.8 11 88.1 84.9 96.4 13.6 Parus major I 16.5 8 115.7 111.5 96.4 14.5 Fringilla montifringilla 3 21 5 125.6 123.2 98.0 16.8 (2-7) (96.5-99.4) (15.3-18.9) Carduelisflammea 2 12.4 6 83.4 82.2 98.6 14.8 (5-7) (97.5-99.7) (14.3-15.4) Emberiza schoeniclus I 19.9 16 117.0 102.4 87.5 19.1 were stretched out to represent a normal flight position (Pennycuick 1989). Finally, the body masswas measuredto the nearest0.1 g using a Pesolaspring balance. When calculatingthe wing loadingI usedthe body massmeasurement recorded when the bird wascaught, i.e., during molt. RESULTS AND DISCUSSION I obtained data on wing area for 41 passerinesof which 21 were Willow Warblers(Table 1). The maximumwing area reductionwas 12.5% asso- ciated with a raggednessscore of 16 in a Reed Bunting (Emberizaschoen- iclus) (Table 1). Maximum raggednessscores are usually around 20 (Bensch and Grahn 1993) and scoresof 16 or more yielded an average wing area reduction of 10% (n = 5). The relationshipbetween relative wing area and raggednessscore was highly significantfor the pooled sam- ple of nine species(linear regression;r = -0.72, P < 0.001, df = 39; Fig. 1); for the Willow Warbler alone the relationship was also significant (r = -0.60, P < 0.01, df = 19; Fig. 1). Hence, the raggednessscore pre- dicted the relative wing area and therefore also the molt gap size. The variation found wasdue to the fact that the samemolt scorecan be given to a range of feather lengths (e.g., score 3 is when the new feather is between one- and two-thirdsgrown; Ginn and Melville 1983). The fact that the primaries are longer than the secondariesalso contributesto the variation. There wasalso a significantrelationship between relative wing area and primary molt score (the sum of individualprimary scores)(r = 0.57, P < 0.001, df = 39). This correlation arose becausethere was a significant correlation betweenprimary molt scoreand raggednessscore (r = 0.54, 106] A. Hedenstr6m j. FieldOrnithol. Winter 1998 ß ß o <= 95 ß ß ß ß ß 85 0 5 10 1'5 20 Raggedness score FIGURE1. The relationshipbetween relative wing area (%; ordinate) and raggednessscore (abscissa)for nine passerinespecies. Regression for all speciespooled is Y = 100.2 - 0.52 X, r• = 0.51, P < 0.001, n = 41, and for the Willow Warbler only (open symbols) Y = 99.5 - 0.47 X, r• = 0.37, P < 0.01, n = 21. Speciesincluded are (sample size separatedon sex given within parentheses):Motacilla flava (4F), Prunella modularis (3M), Luscinia svecica(5M), Phylloscopustrochilus (10M; 11F), Ficedulahypoleuca (1F), Parus major (1F), Fringilla montifringilla(1M; 2F), Carduelisflammea(1M; 1F), and Emberiza schoeniclus(1M).
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