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Diet of the Cantabrian Capercaillie: Geographic Variation and Energetic Content

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Diet of the Cantabrian Capercaillie: Geographic Variation and Energetic Content Ardeola 47(1), 2000, 77-83 DIET OF THE CANTABRIAN CAPERCAILLIE: GEOGRAPHIC VARIATION AND ENERGETIC CONTENT Ana Esther RODRÍGUEZ* & José Ramón OBESO* SUMMARY.—Diet of the Cantabrian Capercaillie: geographic variation and energetic content. The diet of the Cantabrian Capercaillie Tetrao urogallus cantabricus was examined by fecal analysis. Droppings were sampled from April 1998 to May 1999 around 23 leks that were grouped in five zones. Winter diet consisted of beech buds, pine needles, buds and catkins of birch and holly leaves. In each zone, the birds fed almost ex- clusively on one of these species, so winter diet depended on local availability. During spring the consump- tion of beech buds increased in all sites, and later in the season a high diversity of herbaceous plants was in- cluded in the diet. The most important food during summer and autumn was bilberry. Considering the resources that were used as monodiet, beech buds showed both the lowest caloric content and highest the fiber content. The energetic problems derived from winter and spring consumption of beech buds are dis- cussed, and we propose that under certain ecological condition this monodiet might be a limiting factor. Key words: Cantabrian range, diet, digestibility, Tetrao urogallus cantabricus, Tetraonidae. RESUMEN.—Dieta del Urogallo Común cantábrico: composición, variabilidad espacio-temporal y ren- dimiento energético. La dieta del Urogallo Común cantábrico Tetrao urogallus cantabricus fue estudiada me- diante el análisis de heces recolectadas entre abril de 1998 y mayo de 1999 en las proximidades de 23 canta- deros que agrupamos en cinco zonas. La dieta invernal consistió en el uso de diferentes especies arbóreas: brotes de haya, brotes y amentos de abedul, acículas de pino y hojas de acebo fueron los recursos más utili- zados. En cada zona los Urogallos Comunes utilizaron de forma casi exclusiva una de estas especies, de ma- nera que la dieta dependió de la disponibilidad local. Entre la vegetación del suelo destacan la brecina y al- gunos helechos. La dieta primaveral se caracterizó por un aumento generalizado del uso de brotes de haya y posteriormente por la inclusión de diversas especies herbáceas. El arándano fue el alimento principal duran- te el verano y el otoño. De todos los recursos utilizados en monodieta, los brotes de haya fueron los de menor contenido calórico y los de mayor contenido en fibra. Se discuten las repercusiones energéticas que tiene el consumo casi exclusivo de brotes de haya en invierno y primavera, indicando que la dieta podría ser un fac- tor limitante bajo determinadas condiciones ambientales. Palabras clave: cordillera Cantábrica, dieta, digestibilidad, Tetrao urogallus cantabricus, Tetraonidae. INTRODUCTION dators, which might increase in fragmented ha- bitats (Kurki et al., 1997); competition with Diet of birds determines not only resource ungulates (Klaus & Bergmann, 1994; Moss & acquisition, but also habitat selection (Wiens, Picozzi, 1994; Picozzi et al., 1996) and climatic 1989). The diet of threatened species has parti- change (Moss, 1985; Moss & Picozzi, 1994). cular interest because it might be a limiting fac- The effects of these factors are likely to vary tor and, at the same time, supplies information according to geographical location, and other about habitat selection, which is essential to an possibilities, such as diet as a limiting factor, appropriate management of the species. should also be considered. Capercaillie populations have been declining The Cantabrian Capercaillie T. u. cantabri- and their ranges have been contracting during cus has been declining during the last two de- recent decades but the causes are still unknown cades and, although there are few data on Can- (Storch, 1997). The most frequently reported tabrian Capercaillie population trends in the limiting factors for capercaillie are: habitat re- literature, Purroy (1997, 1999) estimated de- duction and fragmentation (Menoni et al., creases between 25 and 50% during the last 1997; Storch, 1991, 1997); the effects of pre- twenty years. * Departamento Biología Organismos y Sistemas, Unidad de Ecología, Universidad de Oviedo. E- 33071 Oviedo. 78 RODRIGUEZ, A. E. & OBESO, J. R. The diet of Capercaillie has been studied hern Cantabrian range: Peloño (Ponga), Bedra- throughout its range and the general characte- món (Allande), Fontiñoso and Llanazanes ristics of food composition are known (see re- (Aller), Muniellos and Hermo (Cangas de Nar- views by Jacob, 1988; Storch et al., 1991). Du- cea) and Lago Bueno, La Pornacal and Salien- ring winter, when the snow covers the ground, cia (Somiedo). These localities were grouped in they feed preferably on trees and the main fo- five zones: 1. Ponga: beech forest with Ilex ods are needles, leaves and buds. During spring aquifolium and Sorbus aucuparia in the under- and summer several ground plants are inclu- growth. 2. Allande: A plantation of Pinus syl- ded in the diet, and during summer and autumn vestris. 3. Aller: beech Fagus sylvatica forest. bilberry Vaccinium myrtillus (shoots, leaves, 4. Somiedo: beech forest. 5. Cangas de Nar- flowers and berries) is usually the main food. cea: Quercus petrea forests with scattered be- Pinus sylvestris is absent from the Cantabrian ech. In the north face of the Cantabrian range range, where winter diets are based on holly the leks are located within beech or oak forests leaves, beech buds and some ferns (Castroviejo, between 1300 and 1500 m a.s.l. (del Campo & 1975; Martínez, 1993). García-Gaona, 1983). The Capercaillie is a typical folivore of the Intestinal droppings were collected as fresh Palearctic boreal coniferous forest and most po- as possible and only one dropping per group of pulations feed almost entirely on conifer needles pellets (e.g. groups of pellets from overnight during most of the year. Folivorous diets are and rest places) was collected. They were sto- rare in birds and demand particular digestive red in paper envelopes and oven-dried at 60° C abilities (Moss, 1983; Sedinger, 1997). In com- for 72 hours. Samples containing a single drop- parison to other diets (animal food and seeds), ping were hydrated in a Petri dish before ob- shoots and leaves are coarse and fibrous; howe- servation under the microscope to identify food ver, the Tetraonidae show two digestive strate- remains. As a reference, a catalogue of cuticu- gies to deal with fibrous foods. One portion of lae from potential food plants from the study the digesta has a rapid passage time through the areas was prepared (Jacob, 1988; Storch, 1991; gut and is excreted as intestinal droppings; the Picozzi et al., 1996). For each sample data of other portion is separated at the end of the intes- presence-absence were taken. tine and introduced in the caeca which retain the Samples were grouped according to the life digesta for long periods and might digest the ce- cycle of the Capercaillie: Spring (April, May llulose (Sybly, 1981; but see Remington 1989; and June; n = 72 samples), summer- autumn Sedinger 1997). The caecal contents are excreted (July, August and September; n = 18 samples) separately as caecal droppings. and winter (November, December, January, Fe- The diet of Cantabrian Capercaillie has pre- bruary and March; n = 137 samples). For each viously been studied by Castroviejo (1975) and period we used the mean of monthly frequen- Martínez (1993); however, it is still a challenge cies. to determine how an eater of needles from the To assess whether statistically significant boreal coniferous forest can survive during differences occurred between seasons and lo- winter in the deciduous forests of the Canta- calities these variables were used as explana- brian range. The objectives of the present study tory variables and presence or absence of food are: 1) to document spatial and temporal va- remains as dependent variables in logistic re- riation in diet composition; 2) to compare the gression analyses. diet in the Cantabrian mountains with diets in To obtain information about food quality, other parts of its range; and 3) to examine the the main food plants were sampled in the study energetic content of its main foods and discuss localities at the time they were included in the whether diet may be a limiting factor in the diet. Each sample included subsamples of at Cantabrian mountains. least ten plants combined; however, the results are only tentative because they refer to single samples. Acid detergent fiber (ADF), crude STUDY AREAS AND METHODS protein (P) and caloric content (cal g–1) were determined according to Van Es & Van Der Droppings were collected around 23 leks at Meer (1980), Van Soest (1963) and Van Soest several localities within Asturias province, nort- & Robertson (1980). TABLE 1 Frequency of food remains in 227 droppings of Capercaillie from five zones within the Cantabrian range. S = April-June (display season), A= July-October (summer and autumn), W= November to March (winter). DIET OFTHECANTABRIANCAPERCAILLIE:GEOGRAPHICVARIATIONANDENERGETICCONTENT [Frecuencias de aparición de los componentes encontrados en 227 excrementos de T. u. cantabricus en la cordillera Cantábrica durante el periodo de estudio. S: celo (abril-junio), A: verano-otoño (julio-octubre), W: invierno (noviembre-marzo)]. SOMIEDO CANGAS ALLANDE ALLER PONGA S 98 A 98 W 98-99 S 99 S 98 A 98 W 98-99 S 99 S 98 A 98 W 98-99 S 99 S 98 A 98 W 98-99 S 99 S 98 A 98 W 98-99 S 99 TREES [árboles] Fagus sylvatica 0.2 0.36 0.33 1 0.14 0.6 0.5 0.7 0.33 0.85 Leaves [hojas] 0.3 Stems [tallos] 0.25
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