Estimating the Potential Effects of Sudden Oak Death on Oak-Dependent Birds
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BIOLOGICAL CONSERVATION 127 (2006) 146– 157 available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/biocon Estimating the potential effects of sudden oak death on oak-dependent birds William B. Monahan*, Walter D. Koenig Museum of Vertebrate Zoology, Integrative Biology, 3101 Valley Life Sciences Building, University of California, Berkeley, CA 94720-3160, USA ARTICLE INFO ABSTRACT Article history: Sudden oak death (SOD), a disease induced by the fungus-like pathogen Phytophthora ramo- Received 9 June 2004 rum, threatens to seriously reduce or eliminate several oak species endemic to the west Available online 13 September 2005 coast of North America. We investigated how the disappearance of one of these species, coast live oak (Quercus agrifolia), may affect populations of five resident oak-affiliated Cali- Keywords: fornia birds – acorn woodpecker (Melanerpes formicivorus), NuttallÕs woodpecker (Picoides nut- California birds tallii), HuttonÕs vireo (Vireo huttoni), western scrub-jay (Aphelocoma californica), and oak Sudden oak death titmouse (Baeolophus inornatus) – using geocoded data from Audubon Christmas Bird Quercus agrifolia Counts, North American Breeding Bird Surveys, and the California Gap Analysis. Capitaliz- Oak woodlands ing on observed relationships between the focal bird species and both oak species diversity and areal extent, we modeled relative bird abundance while assuming complete loss of Q. agrifolia and complete, partial, or no loss of oak habitat following a disease sweep. Post-SOD projections of bird populations occurring within the range of coast live oak were on average 25–68% smaller and 13–49% more variable relative to pre-SOD estimates. SOD effects were greatest for habitats with low initial oak species diversity. Climatic SOD models predicted that the disease stands to negatively impact populations of all five focal bird species throughout 20% of CaliforniaÕs coast live oak habitats. This study provides the first spatially explicit insights into the potential effects of SOD on avian distribution and abundance. Results may be used to help prioritize conservation plans aimed at minimizing overall community level disturbances resulting from the disease. Ó 2005 Elsevier Ltd. All rights reserved. 1. Introduction and severe, as exemplified by the reduction of the American chestnut by the chestnut blight fungus (Cryphonectria parasi- Highly virulent forest pathogens have the potential to greatly tica), which virtually exterminated the tree throughout much reduce or even eliminate tree species at large spatial scales. of its original range within 50 years (McKeen, 1995). Unfortu- For example, within forests of the eastern United States, dis- nately, even with knowledge on how tree diseases affect can- eases caused by fungal pathogens introduced over the past opy composition, it is often difficult to predict a priori the century have significantly impacted the native American cascading effects on the larger community. chestnut (Castanea dentata), American beech (Fagus grandifo- Sudden oak death (SOD) is a disease that currently lia), and American elm (Ulmus americana), in some cases dra- threatens to reduce or eliminate several species of red oaks matically changing forest composition (Houston, 1975; (family Fagaceae) endemic to the west coast of North Amer- Karnosky, 1979; Anagnostakis, 1987). Declines can be rapid ica. Since its detection in 1995, SOD has caused significant * Corresponding author: Tel.: +1 510 642 7888; fax: +1 510 643 8238. E-mail addresses: [email protected], [email protected] (W.B. Monahan). 0006-3207/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2005.08.005 BIOLOGICAL CONSERVATION 127 (2006) 146– 157 147 Fig. 1 – Southern distribution of P. ramorum (a, black buffer) and locations of BBS routes (b, bold black lines) and CBC circles (c, black circles) used in the study. Three Jepson ecological regions (NW, CW, SW) bounded by 40°N encompass the entire statewide distribution of Q. agrifolia (gray areas). mortality of coast live oak (Quercus agrifolia) within primar- We focus on five bird species, acorn woodpecker (Melaner- ily coastal areas of central California (Garbelotto et al., 2001; pes formicivorus), NuttallÕs woodpecker (Picoides nuttallii), Hut- Rizzo et al., 2002; Swiecki and Bernhardt, 2002; Garbelotto tonÕs vireo (Vireo huttoni), western scrub-jay (Aphelocoma et al., 2003; Rizzo and Garbelotto, 2003)(Fig. 1(a)). The dis- californica), and oak titmouse (Baeolophus inornatus), chosen ease is triggered by the fungus-like pathogen Phytophthora for their high dependence on coastal oak habitats for survival ramorum and promotes the rapid development of bleeding and reproduction (Grinnell and Miller, 1944; Pitelka, 1951; bark cankers and foliar pigment degeneration in mature American OrnithologistsÕ Union, 1957; Short, 1971; Miller trees, often resulting in tree death (McPherson et al., 2002; and Bock, 1972; Verner, 1979; Koenig and Haydock, 1999; Cic- Rizzo and Garbelotto, 2003). Known mortality due to P. ero, 2000)(Fig. 2). While SOD clearly stands to negatively impact ramorum is recent, and thus the wildlife impacts are un- populations of such oak-dependent bird species, our goal here known (CalPIF, 2002). is to estimate where in California these SOD-induced losses State and federal wildlife agencies as well as the general will be greatest for both birds and coast live oak. public are becoming increasingly interested in knowing In this study we (1) compare post-SOD relative abundance how SOD stands to negatively impact native species of predictions (mean and CV) for two independent bird census oak-dependent vertebrates. Birds are particularly useful indi- datasets and three different habitat loss scenarios, (2) esti- cators in this regard since excellent survey databases are mate model accuracies with respect to known P. ramorum readily available for both wintering and breeding populations occurrences and climatic SOD models identifying the poten- throughout North America. The most extensive of these are tial current geographical extent of the disease, and (3) con- the Audubon Christmas Bird Count (CBC) and North Ameri- sider how our findings might be further impacted by a can Breeding Bird Survey (BBS), both of which have been continued P. ramorum range expansion coupled with anthro- used extensively and highly successful to study avian distri- pogenic climate change by modeling the future geographical bution and abundance (Bock and Root, 1981; Root, 1988; Price extent of the disease in the year 2050. et al., 1995; Koenig, 1998), as well as model the potential community-level effects of SOD on California birds (Monahan and Koenig, 2005). Our approach here estimates 2. Methods the population-level consequences of SOD using the two independent datasets whose seasonal timing and methodol- 2.1. Data overview ogies are quite different and thus to some degree complementary. Bird distribution and abundance data were obtained from We use the statistical relationship between relative bird Audubon Christmas Bird Counts (CBC) (Butcher, 1990) and abundance and oak species diversity and oak areal extent to the North American Breeding Bird Survey (BBS) (Sauer et al., quantify expected avian population declines and census size 2003). A single CBC involves between a few and over 100 peo- fluctuations resulting from a reduction in oak diversity and ple counting all the birds they can during a single day within either complete, partial, or no loss of oak habitat. Permanent the last two weeks of December within a 24 km diameter cir- change to coast live oak habitats is probable since Q. agrifolia cle (total area approximately 450 km2). Sites are usually cen- suffers extensive SOD mortality (Rizzo and Garbelotto, 2003), tered on some geographic landmark and are in many cases and because large portions of its geographic range are ex- surveyed year after year. CBC analyses utilized data from pected to experience disease sweeps as P. ramorum continues the 38 years between 1960 and 1997, inclusive. For each spe- to expand throughout California (Meentemeyer et al., 2004; cies x count combination, we standardized observer effort Guo et al., 2005). by using the log-transformed (log(x + 1)) number of birds 148 BIOLOGICAL CONSERVATION 127 (2006) 146– 157 Fig. 2 – Statewide range maps of five oak-dependent bird species (gray + black areas), acorn woodpecker (a), NuttallÕs woodpecker (b), HuttonÕs vireo (c), western scrub-jay (d), and oak titmouse (e), intersected with coast live oak habitats (black areas only). Intersections encompass 10% (acorn woodpecker) to 17% (NuttallÕs woodpecker and HuttonÕs vireo) of the bird speciesÕ California distributions. counted divided by the number of Ôparty-hoursÕ of observa- (Q. garryana), valley oak (Q. lobata), interior live oak (Q. wislize- tion. We then determined the mean and coefficient of nii), and tanoak (Lithocarpus densiflorus). The analyses thus variation (CV; SD/mean · 100) of this value (Ôbirds per party considered all tree species of oaks in mainland California; hourÕ) across all years the count was performed as an esti- shrub oaks were not included. mate of overall relative abundance and variability of the spe- Distributions were generated for Q. agrifolia occurring in cies at that site. Breeding Bird Surveys consist of 3-min the absence of all other focal oaks and for all possible combi- censuses at a series of 50 stops, 0.8 km apart, conducted once nations of 1, 2, and P3 additional oak species.