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Response of Passerine Birds to an Irruption of a Pine Processionary Moth Thaumetopoea Pityocampa Population with a Shifted Phenology

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Response of Passerine Birds to an Irruption of a Pine Processionary Moth Thaumetopoea Pityocampa Population with a Shifted Phenology Ardeola 56(2), 2009, 189-203 RESPONSE OF PASSERINE BIRDS TO AN IRRUPTION OF A PINE PROCESSIONARY MOTH THAUMETOPOEA PITYOCAMPA POPULATION WITH A SHIFTED PHENOLOGY RESPUESTA DE LAS PASERIFORMES A LA IRRUPCIÓN DE LA PROCESIONARIA DEL PINO THAUMETOPOEA PITYOCAMPA CON FENOLOGÍA VARIADA Carla PIMENTEL* 1 and Jan-Åke NILSSON** SUMMARY.—Response of passerine birds to an irruption of a pine processionary moth Thaumetopoea pityocampa population with a shifted phenology. The main purpose of the present work was to investigate if the passerine community was able to re- spond to a localized irruption of a temporally shifted population of the pine processionary moth Thaumetopoea pityocampa, the larval development of which takes place during the summer, called sum- mer population (SP). The work was conducted in the National Pine Forest of Leiria, a highly organised production forest, essentially consisting of maritime pine, Pinus pinaster, and located in the central west coast of Portugal (39º 50` N, 8º 57` W, 30 - 50 m a.s.l.). The SP irrupted in a homogeneous area consist- ing of young pines. Bird abundances and richness were estimated by point counts in the infested area and in two similar non-infested areas. The counts were made during all four seasons, covering all stages of the moth annual cycle and were repeated during two years. The overall abundance of birds was similar in both infested and non-infested areas. However, the richness of canopy gleaners was higher in the SP area than in a nearby non-infested area. Only two bird species responded numerically to the increased insect abun- dance: the blackbird Turdus merula, and the great tit Parus major. Thus the passerine community respond- ed only to a limited degree to the irruption of the SP. Three factors might have accounted for the results: (i) the bird community consists mostly of territorial residents or short distance migrants and their low range of dispersal may decrease the probability of a response to a localized insect outbreak. (ii) The lar- vae of the pine processionary moth has urticating hairs which are considered to be an effective repellent defence against vertebrate predators, thus reducing the number of species that could react to the high cater- pillar densities. (iii) The simple vegetation structure of the forest, homogeneous stands of small young trees, precludes the establishment of cavity nesters that constitute a large part of the canopy gleaners and most of the species which have been reported as potential T. pityocampa predators. However, results in- dicate that this important Mediterranean defoliator may have a positive effect on the canopy gleaners and on some species that are able to act as its predators. Key words: bird community, insect defoliator, insect outbreak, life-cycle shift, Mediterranean pine for- est, numerical response, point count. * DCEA/FCT, Universidade Nova de Lisboa. PT-2825-516 Campus de Caparica, Portugal. ** Department of Animal Ecology, Lund University. S-223 62 Lund, Sweden. 1 Corresponding author: [email protected] 190 PIMENTEL, C. and NILSSON, J. Å. RESUMEN.—Respuesta de las paseriformes a la irrupción de la procesionaria del pino Thaumetopo- ea pityocampa con fenología variada. El propósito principal de este trabajo fue investigar si la comunidad de paseriformes respondería a una irrupción local de la procesionaria del pino Thaumetopea pityocampa en una población con cambios temporales, cuyo desarrollo de larvas ocurre en verano (llamada población de verano, PV). El trabajo se desarrolló en el Bosque Nacional de Pino de Leiria (Portugal), un bosque productor altamente organiza- do, localizado en la costa centro-occidental de Portugal (39° 50’ N, 8° 57’W, 30 - 50 m.s.n.m.) y que está compuesto esencialmente por el pino marítimo Pinus pinaster. La PV irrumpió en un área homogénea poblada por pinos juveniles. La abundancia y riqueza de aves fue estimada por medio de conteos por pun- tos en el área infestada y en dos áreas similares no infestadas. Los conteos fueron realizados en las cua- tro estaciones, cubriendo todas las etapas del ciclo anual de la polilla y fueron repetidos durante dos años. La riqueza de recogedores del dosel fue mayor en el área de la PV que en un área adyacente no in- festada. Sólo dos especies respondieron numéricamente al incremento de abundancia de insectos: el mirlo común Turdus merula y el carbonero común Parus major. Así, la comunidad de paseriformes res- pondió sólo en menor grado a la irrupción de la PV. Tres factores podrían explicar los resultados: (i) la co- munidad de aves consiste, en su mayoría, en residentes territoriales o migratorios de corta distancia en las cuales su escaso rango de dispersión podría limitar la probabilidad de respuesta a una irrupción localiza- da. (ii) Las larvas de la procesionaria del pino tienen pelos urticantes que son considerados un mecanis- mo de defensa repelente contra depredadores vertebrados, reduciendo así el número de especies que po- drían reaccionar a altas densidades de orugas. (iii) La estructura simple de la vegetación del bosque, con parcelas homogéneas de pinos juveniles pequeños, excluye el establecimiento de las aves que anidan en cavidades que constituyen gran parte de los recogedores del dosel, además de la mayoría de las especies que han sido registradas como depredadores de T. pityocampa. Los resultados indican que este impor- tante defoliador mediterráneo podría tener efectos positivos en los recogedores del dosel y en algunas aves con capacidad de funcionar como depredadores de sus larvas. Palabras clave: comunidad de aves, insecto defoliador, irrupción de insectos, cambios en el ciclo de vida, bosque de pino mediterráneo, respuesta numérica, conteo por puntos. INTRODUCTION having a major impact on the bird communi- ty structure (Holmes et al., 1986). This phe- Tree canopy insects are generally consid- nomenon was also observed in North Euro- ered as an important food item for most forest pean deciduous forests, although at a somewhat birds (e.g. Buckner and Turnock, 1965; Gui- smaller scale (Hogstad, 2005). However, in tián, 1985; Poulin and Lefebvre, 1996; Hogstad, forests from more southern latitudes, includ- 2005). In temperate forest ecosystems, some ing the Mediterranean area, there is limited in- of these folivorous species are able to cycli- formation on how a bird community responds cally reach outbreak numbers and cause ex- to irruptions of insect defoliators. tensive defoliation (Myers, 1998; Bjørnstad et The pine processionary moth (Thaume- al., 2002; Tenow et al., 2007), a phenomenon topoea pityocampa, Lepidoptera: Notodonti- which is also known in other forest ecosys- dae) is an important pine defoliator in the tems, although not so well studied (Van Bael Mediterranean region, and there is evidence of et al., 2004). In temperate deciduous forests cyclic outbreaks (Hódar and Zamora, 2004; in North America, the sudden impulse of food Robinet, 2006). This univoltine species spends caused by Lepidoptera outbreaks significant- the pupal stage in the ground during the spring. ly affects abundances of many bird species, Adults emerge in the summer and live for a few Ardeola 56(2), 2009, 189-203 PASSERINE BIRDS AND THE PINE PROCESSIONARY MOTH 191 days, just enough for reproduction. Female liators through severe defoliation by late instar moths of this species lay a single batch of eggs, aposematic caterpillars (Pelech and Hannon, and larvae hatch about one month later, devel- 1995). However, in spite of the importance of oping throughout the winter (between Septem- T. pityocampa as a widespread forest defolia- ber and March) (OEPP/EPPO, 2004). Howev- tor, studies on the responses of the bird com- er, in 1997, a population of T. pityocampa with munity to high and low densities of this pest, a temporally shifted life cycle was recorded for are still scarce (but see Barbaro et al., 2008). the first time in a restricted area of the Nation- The irruption of the SP in a localized patch al Pine Forest of Leiria. Larval development of the forest, in an area where previous out- of this novel population occurs during the sum- breaks by defoliators are unknown (informa- mer instead of during the winter, and so this tion provided by the National Forest Services), population will be referred to as the Summer offered us an opportunity to assess how the bird Population (SP). Since its discovery, the SP has community of Mediterranean pine forests re- been observed every year in the same re- sponds to outbreaks of a common insect defo- stricted area, causing visibly high levels of de- liator. Thus, in the present paper we character- foliation (own observations) and attaining high- ize the passerine community in the pine er densities than the Winter Population (WP) plantations where the SP established, and in- in the same area (Pimentel et al., 2006). vestigate if birds are able to respond to the ir- The pine processionary moth has aposemat- ruption of the shifted population of T. pityocam- ic gregarious caterpillars, with conspicuous pa. Accordingly, we compare bird richness and hairs and urticant setae in the last two instars abundance in the area of the irruption of the of development. These characteristics are con- SP, with similar uninfested forest areas. Pre- sidered as effective repellent defences dictably, the unexpected food source and de- against most bird predators (e.g. Bowers and foliation caused by a T. pityocampa outbreak Farley, 1990; Lindström et al., 2001; Exnerová should affect some, but not all, of the guilds et al., 2007). This way, it is expected that T. comprising the community, and therefore we pityocampa early instar larvae will constitute examine the effects of the SP outbreak sepa- a food source for avian predators, but preda- rately for each foraging guild. tion of later urticant instar larvae and pupae should be restricted to a few species that are able to overcome its defences.
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