BIOLOGICAL CONSERVATION Biological Conservation 118 (2004) 195–204 www.elsevier.com/locate/biocon Winter flocking behaviour of speckled warblers and the Allee effect Janet L. Gardner School of Botany and Zoology, Australian National University, Australia Received 25 May 2003; received in revised form 8 August 2003; accepted 11 August 2003 Abstract The aggregation of individuals into foraging flocks is one behavioural trait that, if disrupted, can cause the Allee effect, which is a slowing in population growth at low density or small population size, and this can greatly increase the risk of extinction. Here, I describe intraspecific flocking behaviour of a colour-banded population of speckled warblers, Chthonicola sagittata, a species that has declined across a large part of its range in the fragmented temperate woodlands of Australia. I make predictions about the context in which the Allee effect might be expressed and the consequences for the viability of populations living in small habitat remnants. Speckled warblers lived in discrete, stable social groups throughout winter, the nucleus of which was the residents from one or more adjacent breeding territories. The timing and mode of flock formation and the size of flocks varied between two winters, apparently in response to the severity of conditions; thus flocking probably facilitates increased foraging efficiency and predator detection, potentially leading to increased survival in harsh conditions. Because flock territories were up to 30 ha each, and larger territories are likely, birds living in remnants smaller than 40 ha may suffer increased mortality if there are too few birds available to form flocks of an appropriate size to facilitate the benefits of grouping when conditions are most extreme. Further, in small remnants where survival is reduced, dominance behaviour and male–male competition may act to compound the Allee effect by reducing reproductive success. Regardless of these predictions, speckled warbler populations may only be viable in remnants that are large enough to support multiple flocks, to enable rapid recruitment to breeding vacancies and thus provide adequate numbers of birds for flocking. Ó 2003 Elsevier Ltd. All rights reserved. Keywords: Habitat fragmentation; Allee effect; Flocking; Behaviour and conservation; Birds 1. Introduction group size, up to a point, and group size is varied in response to the severity of conditions (Elgar, 1989). A A variety of behavioural traits, subject to particular reduction in population size that depletes the number of conditions, can cause the Allee effect, which is a slowing individuals available for grouping may therefore lead to in population growth at low density or small population a reduction in survival if groups become so small that size (Allee, 1931; Sæther et al., 1996; Courchamp et al., foraging and anti-predator strategies become inefficient. 1999; Stephens and Sutherland, 1999). One of the best- Thus, the Allee effect may be an important mechanism known examples of such behaviour is the aggregation of for population regulation, especially in species that show individuals into foraging groups (Sæther et al., 1996; a high degree of sociality (Sæther et al., 1996; Cour- Courchamp et al., 1999; Reed, 1999; Stephens and champ et al., 1999; Reed, 1999; Stephens and Suther- Sutherland, 1999; Stephens et al., 1999). Grouping is land, 1999). thought to facilitate increased foraging efficiency and There is evidence that the Allee effect can occur in a predator detection and therefore acts to increase sur- wide range of taxa and can greatly increase the likeli- vival in harsh conditions (Pulliam, 1973; Ekman, 1987; hood of local and global extinction, but despite this, its Hogstad, 1988; Lima and Dill, 1990; Bednekoff and importance in population ecology has traditionally been Lima, 1998). The benefits of grouping increase with underestimated (Creel, 1998; Courchamp et al., 1999; Reed, 1999; Stephens and Sutherland, 1999). This may E-mail address: [email protected] (J.L. Gardner). be due to the inherent difficulty of demonstrating an 0006-3207/$ - see front matter Ó 2003 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2003.08.018 196 J.L. Gardner / Biological Conservation 118 (2004) 195–204 imbalance between births and deaths in small popula- lation size in fragmented landscapes and thus make it tions (Courchamp et al., 1999; Stephens et al., 1999). In prone to extinction (May, 1973; Soule, 1987; Caugh- addition, the consequences of such an imbalance for ley, 1994; Mac Nally and Bennett, 1997; Reed, 1999). population viability may be difficult to predict because One aspect of behaviour that may have consequences outcomes will depend on the prevailing environmental for survival as a result of the potential for the Allee conditions. Nevertheless, recent advances in statistical effect is the inclination to form intraspecific flocks as modelling have led to a better understanding of such well as join mixed species flocks in winter. Habitat interactions and have highlighted the importance of the fragmentation can influence the dynamics of winter Allee effect in the dynamics of small populations (e.g., flocks because it can affect the numbers of individuals McCarthy, 1997; Amarasekare, 1998). available for flocking, as well as predator abundance Habitat fragmentation is one form of anthropogenic and the availability of food and suitable habitat disturbance that has driven many populations to small (Telleria et al., 2001). sizes or low densities, potentially placing them at risk Here, I describe the formation, structure and com- from the Allee effect (Courchamp et al., 1999; Reed, position of intraspecific flocks of speckled warblers liv- 1999). Habitat fragmentation alters the amount, con- ing in a large (>1000 ha) habitat remnant, based on figuration in the landscape and quality of suitable hab- observations of a colour-banded population over two itat available for species, thereby having potential to winters. I conclude that flocking is likely to increase affect the distribution and abundance of species (Saun- individual fitness and thus, under specific conditions, ders et al., 1991; Fahrig, 1997). Studies of habitat frag- trigger the Allee effect. I make predictions about the mentation have tended to focus on description of the context in which the Allee effect might be expressed and loss of biodiversity and changes to species composition discuss probable consequences for populations living in in remnants. Much less is known about mechanisms of small remnants of habitat. decline and this makes it difficult to predict the conse- quences of environmental change (Sutherland, 1998; Courchamp et al., 1999; Reed, 1999; Anthony and 2. Methods Blumstein, 2000). This limitation has led to a call for behavioural studies, including those that elucidate the 2.1. Study species and site causal mechanisms of the Allee effect (Courchamp et al., 1999; Stephens and Sutherland, 1999). The speckled warbler is a small (11–15 g) insectivo- The speckled warbler Chthonicola sagittata is one of a rous passerine which is a sedentary resident of the number of declining species from the highly fragmented temperate woodlands of eastern Australia (Blakers temperate woodlands of Australia (Garnett, 1992; et al., 1984). It is a member of the Acanthizinae within Robinson and Traill, 1996; Recher, 1999; Reid, 1999). the Pardalotidae (Christidis and Boles, 1994) or Acan- Since European settlement, up to 80% of this habitat thizidae, in which case the species name has reverted to type has been cleared to make way for grazing of live- Pyrrholaemus sagittatus (Schodde and Mason, 1999). I stock and agriculture, and this has had considerable use Acanthizinae hereafter. Males are slightly larger impact on the status of species within the region (Ford than females and can be distinguished on the basis of a et al., 2001). Speckled warblers have declined across a black rather than rufous crown stripe (Blakers et al., large part of their range, and in districts where no 1984). In the population studied, speckled warblers bred habitat fragments larger than 100 ha remain, they ap- in the austral spring and summer (Gardner, 2002). Most pear to be locally extinct (Barrett et al., 1994; Robinson birds bred as a simple pair but up to 30% of breeding and Traill, 1996; Traill and Duncan, 2000). In addition units were trios, consisting of a female with two males, to area, habitat quality is likely to affect distribution in in which case the second or beta male was subordinate remnants, since presence of the species appears to be to the alpha (Gardner et al., in press; Gardner et al., associated with a high degree of habitat complexity 2003). Females laid clutches in up to 7 months of each (Watson et al., 2001). year, were multi-brooded and fledged up to three broods Specific causes of the decline of speckled warblers per season, but despite this, reproductive success was are unknown but thought to be linked with processes low with each female producing about 1.0 fledgling per associated with the fragmentation of their habitat, season (Gardner, 2002). All juveniles dispersed from the rather than a direct result of habitat loss, since pop- natal territory about 7 weeks after fledging, soon after ulations have continued to decline in areas where land reaching foraging independence (Gardner et al., 2003). clearing has stopped (Reid, 1999). The species occurs The population studied occurred in open eucalypt at low population density and is a specialist in terms woodland, dominated by Eucalyptus rossii, Eucalyptus of foraging and nesting behaviour (Ford et al., 1986; mannifera and Eucalyptus macrorhyncha, on the lower Tzaros, 1996; Gardner, 2002; Gardner et al., 2003), slopes of Mt. Anslie in the Canberra Nature Park, characteristics that are likely to result in small popu- Australian Capital Territory (149°90E, 35°160S). The J.L. Gardner / Biological Conservation 118 (2004) 195–204 197 study area was about 300 ha in size, within the larger finish of each watch, and these, along with the initial 1500 ha reserve.