Climate Change Refugia for Biodiversity in the Klamath-Siskiyou Ecoregion Author(s) :David Olson, Dominick A. DellaSala, Reed F. Noss, James R. Strittholt, Jamie Kass, Marni E. Koopman and Thomas F. Allnutt Source: Natural Areas Journal, 32(1):65-74. 2012. Published By: Natural Areas Association DOI: http://dx.doi.org/10.3375/043.032.0108 URL: http://www.bioone.org/doi/full/10.3375/043.032.0108
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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. C O N S E R V A T I O N I S S U E S ABSTRACT: The Klamath-Siskiyou Ecoregion has been a refuge for species during past climate change events, but current anthropogenic stressors are likely compromising its effectiveness as a refugium for this century’s projected changes. Reducing non-climate stressors and securing protection for large, complex landscapes are important long-term actions to alleviate climate change impacts on biodiversity. Equally • important is the immediate protection of a network of climate change microrefugia, particularly old growth and intact forests on north-facing slopes and in canyon bottoms, lower- and middle-elevations, Climate Change wetter coastal mountains, and along elevational gradients. Such areas provide local opportunities for vulnerable species to persist within the ecoregion. We identify a provisional set of 22 highest-priority and 40 high-priority microrefugia that occur mostly outside of existing protected areas and along wetter and Refugia for lower elevations of the ecoregion. Proposed reserve designs, if fully implemented, would capture most of the recommended microrefugia, although we found 11 important gaps. Most of the region’s biodiversity, Biodiversity in the endemic species, and species vulnerable to climate change are invertebrates, non-vascular plants, and fungi that are largely restricted to persistently cool and moist late-successional forests. Opportunities for Klamath-Siskiyou climate change response for vulnerable taxa will necessarily be local due to a limited capacity of many species to move to new habitat, even over relatively small distances where land use practices create Ecoregion inhospitable conditions. The ecoregion’s distinctive and endemic serpentine-substrate flora also is at risk and possible refugia are sites that will retain wet soil conditions, such as seeps and bogs.
1 David Olson Index terms: climate change, ecoregion, Klamath-Siskiyou, microrefugia, refugia 1 Conservation Earth Consulting 4234 McFarlane Ave. Burbank, CA 91505 INTRODUCTION millennia. Impacts include loss of contigu- ous habitat along intact elevational and 2,6 Dominick A. DellaSala The Klamath-Siskyou Ecoregion (KSE) other environmental gradients that may Reed F. Noss3 contains globally important biodiver- facilitate climate-related shifts in natural James R. Strittholt4 sity–only five other temperate forests communities and loss and degradation of regions are as diverse or home to as many most of the mature and old-growth for- Jamie Kass endemic species and ancient lineages (e.g., ests (e.g., only about 28% of the historic Marni E. Koopman2 Caucasus, Southwestern China, Southeast- old-growth forests remain; Strittholt et Thomas F. Allnutt5 ern United States, Coastal Plain/Southern al. 2006), particularly mesic lowland and Appalachians, Valdivia rainforests of Chile mid-elevation habitats (Staus et al. 2002). and Argentina; Olson et al. 2001; Tecklin Increasing prevalence of invasive plants 2 Geos Institute 84 Fourth Street et al. 2011). The special location (latitude and pathogens facilitated by road building Ashland, OR 97520 and coastal proximity), rugged terrain, cli- and land use practices poses an additional matic stability, and complexity of soils and threat to native species and communities 3 Department of Biology microclimates have allowed the region to (DellaSala et al. 1999). University of Central Florida act as a refuge from past climatic changes 4000 Central Florida Blvd. for species and natural communities requir- The existing protected area system (i.e., Orlando, FL 32816-2368 ing cool and moist conditions (Whittaker National and State Parks, Wilderness Areas, 1960, 1961; Stebbins and Major 1965; National Monuments, Botanical Areas) is 4 Conservation Biology Institute Wagner 1997; Coleman and Kruckeberg inadequate for ensuring the persistence 260 SW Madison Avenue, Suite 106 1999; Sawyer 2007). of most of the ecoregion’s vulnerable Corvallis, OR 97333 biodiversity (DellaSala et al. 1999; Noss 5 Department of Environmental Science, One might expect that the KSE will con- et al. 1999; Carroll et al. 2010). Existing Policy and Management tinue to function well as a climate change reserves largely protect higher-elevation 130 Mulford Hall refugium as human-caused climate change communities, while the lower-elevation University of California progresses. However, cumulative land use reserves are limited in their geographic Berkeley, CA 94720-3114 impacts combined with projected climate extent, thereby missing many distinct change could have a profound impact on lowland species assemblages and areas • the ecoregion’s species and ecosystems. that may act as potential microrefugia. In the KSE, over a century of land use We define microrefugia as sites with cool 6 Corresponding author: activities (e.g., logging, mining, livestock and moist conditions conducive to the [email protected] grazing, damming of rivers, mining, and persistence of species vulnerable to climate human-caused alterations of fire) have change. Thus, our conservation strategy for resulted in loss or degradation of mesic the KSE builds on prior reserve propos- Natural Areas Journal 32:65–74 habitats (DellaSala et al. 1999) that may als (Noss et al. 1999; Carroll et al. 2010; have previously functioned as refugia over KS Wild 2010; Siskiyou Project 2010) by
Volume 32 (1), 2012 Natural Areas Journal 65 adding microrefugia and other elements protection of large, complex landscapes California (KS Wild 2010) and a ~445,000 to create a reserve design more robust to with diverse terrains, soils, microclimates ha proposed Siskiyou Wild Rivers National anticipated increases in temperature and and other environmental gradients. In Salmon and Botanical Area in southwest changes in precipitation over the century particular, low and mid-elevation habitats Oregon, a hotspot of serpentine flora and (Koopman et al. 2009). in higher precipitation areas (e.g., along wild rivers (Siskiyou Project 2010; Figure the coast) will provide multiple local op- 1). Protection of these areas will greatly im- portunities for persistence of vulnerable prove the chances for persistence of a large CORE CONSERVATION ELEMENTS species. In the KSE, conservation groups portion of the ecoregion’s terrestrial and FOR ROBUST RESERVE DESIGN IN A have identified two areas having these freshwater biota even if we are uncertain CHANGING CLIMATE characteristics: a 243,000 ha land bridge of the magnitude, timing, and distribution known as the proposed Siskiyou Crest of changes in temperature and precipitation Fundamental to the development of a robust National Monument (considered a climate at sub-ecoregional scales (e.g., Murphy et conservation design are three core-plan- refuge) in southwest Oregon and northern al. 2004; Moilanen et al. 2006). ning elements: (1) reduction of non-cli- mate stressors; (2) protection of complex landscapes; and (3) protection of climate change microrefugia. Taken together, they are the foundation for guiding reserve design and conservation implementation in the KSE.
Reduction of Non-Climate Stressors
Reducing non-climate stressors across the landscape, such as curtailing or greatly reducing logging and road building, is the single most important action that land managers can take to help the regional biota and ecosystems persist in the face of a changing climate. The release from stressors should be strategically targeted to critical core habitats, old-growth forest mi- crorefugia, and adaptation corridors along environmental gradients (sensu, Olson et al. 2009). For example, if large complex landscapes were off-limits to logging (only about 13% of the region is strictly protected; DellaSala et al. 1999), and all of the predicted local climate refugia, old-growth forests, and priority corridors in the KSE (e.g., Noss et al. 1999) were effectively protected, this would have a much more positive effect for biodiversity than if most of the area released from log- ging was in highly degraded, mid-elevation production forests. The release of strategic areas from land use stressors would need to allow maturing forests to once again dominate the landscape.
Protection of Complex Landscapes
Securing a high level of protection and undertaking ecologically based restoration Figure 1. Climate change vulnerability zones (coastal, transition, dry, high elevation) of the Klamath- in degraded areas is important, as well as Siskiyou Ecoregion, southwest Oregon and northern California, used in the analysis to inform priority site selection for conservation action.
66 Natural Areas Journal Volume 32 (1), 2012 Protection of Climate Change tively mature and old growth) that can act better able to retain more moisture and Microrefugia as climatic buffers. cooler conditions than lower precipita- tion zones. This is due to more abundant In order to maintain pockets of habitat Many extant microrefugia and the species water and greater canopy, understory for climate-vulnerable species, conserva- and populations they contain may be lost vegetation, litter biomass, and complexity tion attention should be aimed at securing or degraded within a few decades due to in these forests. Late-seral forests within microrefugia that may uniquely provide ongoing exploitation of forests and land- watersheds are also superior to degraded, opportunities for many species to persist scapes within the ecoregion, particularly logged, roaded, and burned vegetation for and are particularly threatened due to ongo- at low and mid elevations, slow pace of providing cooler stream temperatures and ing habitat degradation and rapid warming. change that is typical for forest manage- robust aquatic ecosystems (Strittholt and The importance of microrefugia for the ment and protected areas practices, and DellaSala 2001; Staus et al. 2010). long-term persistence of species that are rapid changes being documented in climate sensitive to climate change is increasingly and natural communities (Damschen et al. Storm tracks, regional rainfall, and fog pat- being recognized (Noss 2001; Loarie et 2010). Although the long-term efficacy of terns may shift due to climate change (Det- al. 2008, 2009; Rull 2009, 2010; Ashcroft microrefugia is still uncertain (Carroll et al. tinger et al. 1998; Salathé et al. 2008; Mote 2010; Dobrowski et al. 2010). In temper- 2010; Dobrowski et al. 2010), especially and Salathé 2009; Johnstone and Dawson ate regions, terrain positions and habitat if they remain embedded within largely 2010), but coastal mountains are expected types that maintain persistent cool and degraded landscapes, it remains a prudent, to continue to receive Pacific storms first moist conditions favorable for effective bet-hedging strategy in the face of uncer- and much of the region’s rainfall into the microrefugia are increasingly well defined tainty to protect a network of microrefugia future (Daly et al. 1994). Certainly, vul- (e.g., Dobrowski et al. 2010). representative of the ecoregion’s distinct nerable species and communities occur at species assemblages. higher elevations and in drier areas towards Because of the rapid speed of climate the eastern portion of the ecoregion, but the change (Loarie et al. 2008, 2009), includ- vast majority of distinctive biodiversity for Microrefugia Site Features ing warmer temperatures (Koopman et al. the ecoregion (all taxa being considered) 2009) and diminishment of fog (Johnstone occur within the coastal fog and transition Site features for effective microrefugia in and Dawson 2010) in the KSE, opportu- zones (Figure 1; Sawyer 2007). The latter the KSE include north-facing slopes, valley nities for long-term persistence for many zone includes more mesic forests along bottoms and steep canyons, and sinks and species will be local, likely within a scale the Siskiyou Crest (Oregon/California), basins because they are shadier and exist of a few kilometers, from the location of Eddy Mountains (northwest California), where cool air predictably pools in the present populations. Many species will be Scott Mountains (northwest California), lower sites (Dobrowski et al. 2010). Such unable to shift rapidly enough to areas with and Yolla Bolly’s (southern limits of the sites are likely to have climate states and more favorable conditions. Moreover, most ecoregion) that are relatively far from trends that are decoupled from regional of KSE’s species, distinctive (endemic) the coast. In general, the larger and more averages, a requisite for microrefugia to species, and those vulnerable to climate round a forest block, the greater the core persist through time. Forests with a north- change are mesophilic, old-growth forest habitat area–internal habitat that does not east- and north-facing aspect also have a specialists, largely lesser known taxa (by experience the drying effects of forest edges lower frequency of wildfires that can alter the public) such as invertebrates, fungi, (Chen et al. 1999). the capacity of habitats to retain cool and bryophytes, and other non-vascular plants moist conditions (Taylor and Skinner 2003; (Olson 1992; Lattin 1993; Olson 2010; Natural communities and vulnerable spe- Alexander et al. 2006). Vicente 2010). The majority of these taxa cies within refugia also will have improved cannot cross even small distances of terrain opportunities for persistence if microrefu- Habitat types that will function well as with unfavorable conditions (e.g., light, gia span broad elevational gradients, allow- microrefugia for climate change-sensitive hot, and dry; Frest and Johannes 1993; ing populations to shift locally over time species include late-seral forests, although Niwa and Peck 2002). Thus, protection through contiguous mesic habitat (Noss the greater litter, understory vegetation, and restoration of microrefugia around 2001; Olson et al. 2009). North-South cor- and canopy complexity and biomass of extant populations is essential for the long- ridors of contiguous natural vegetation are old-growth forests (> 150 yrs) makes them term perpetuation of the vast majority of important for many reasons, such as dis- superior at retaining moisture (Chen et al. the KSE biota. The ecoregion’s endemic persing vertebrates, but a swiftly changing 1999). Late-seral forests that occur in areas serpentine flora (Kruckeberg 1984; Har- climate will likely limit the ability of most with high-precipitation and fog, such as rison et al. 2006; Sawyer 2007) is also slowly dispersing organisms to move long in coastal mountains (Loarie et al. 2008; highly vulnerable to projected increases distances northwards over generations. in temperature and drying (Damschen et Ackerly et al. 2010; Carroll et al. 2010) al. 2010) and some taxa may only persist or other areas that experience significant within persistently wet pockets and seeps orographic precipitation (e.g., > 1143 mm REPRESENTATION OF BIODIVERSITY surrounded by late-seral forests (collec- annual precipitation) will, on average, be WITHIN MICROREFUGIA:
Volume 32 (1), 2012 Natural Areas Journal 67 MESOREFUGIA AS A PROXY
Until patterns of local endemism and beta-diversity for speciose groups, such as invertebrates, are better known, prox- ies for mapping distinct assemblages can be used to assess how well a network of microrefugia provides refuge to KSE’s diverse biota. Useful proxies for assess- ing representation of biodiversity within and among microrefugia are mesorefugia. We define mesorefugia as large areas that contain nested clusters of microrefugia with similar species assemblages that have functioned as a refugium over millennia (Rull [2009] defines mesorefugia as larger regions to which temperate biotas shifted during glacial maxima). Mesorefugia typically occur at the scale of mountain ranges or watershed complexes along coastlines, and their location along river canyons (e.g., Rogue, Umpqua, Klamath, Eel rivers) may facilitate future expansions and enable vagile species to move more freely across landscapes. Careful selection and protection of microrefugia of varying species assemblages (e.g., plant association groups) within and among mesorefugia would help to achieve representation goals while maximizing the number of extant species that will persist in emerging novel ecosystems.
Mesorefugia analyses complement exist- ing representation analyses that focus on vegetation types and other communities derived from combinations of biophysi- cal features (e.g., Vance-Borland 1999; Staus et al. 2001; Carroll et al. 2010). As such, candidate mesorefugia (Figure 2) Figure 2. Provisional mesorefugia (ovals) within the Klamath-Siskiyou Ecoregion, southwest Oregon and northern California, approximated from large-scale predictors (e.g., coastal mountains in areas of for the KSE were initially identified from relatively high precipitation) and an overlay of the distribution of mesophilic, restricted-range species large-scale biophysical features and loca- including Plethodontid and Dicamptodon salamanders, Caseyid millipedes, Pentanychid harvestman, tions that predict effective refugia–coastal endemism zones for vascular plants, and relict conifers. Mesorefugia likely contain concentrations of mountains with complex topography and restricted-range species due to their persistently wet conditions and long-term stability. Dashed ovals areas of high precipitation (Loarie et al. represent high-elevation refugia that may, or may not (depending on the severity of warming tem- peratures at higher elevations), function well under current and future human-caused climate change. 2009; Rull 2009, 2010; Dobrowski 2010). Numbering refers to locations discussed in the text. Areas with concentrations of restricted- range (i.e., local endemic) species or relict taxa dependent on cool and moist habitats Plethodon and Dicamptodon salamander 2005), and land snails (Frest and Johannes were also evaluated to refine candidate species and subgoups (Bury 1973; Mead 1993). We stress the mesorefugia proposed mesorefugia locations and boundaries et al. 2005; Steele and Storfer 2006), and here are provisional and will benefit from (i.e., where multiple species boundaries numerous other plants (Sawyer 2007) and more rigorous analyses of biophysical overlap). These include the distribution of invertebrates (Olson 1992), such as harvest- predictors and species distributions. Brewer spruce (Picea breweriana), Engel- man (Briggs 1969, 1971ab), millipedes mann spruce (Picea engelmanni), foxtail (Gardner and Shelley 1989; Olson 1992), Based on these criteria and species distribu- pine (Pinus balfouriana) (Sawyer 2007), trapdoor spiders (Cokendolopher et al. tion maps, important mesorefugia for the
68 Natural Areas Journal Volume 32 (1), 2012 KSE include: (1) Kalmiopsis; (2) North habitat types (e.g., serpentine barrens) and Rock Creek area, California); northern Siskiyou Mountains; (3) East Siskiyous; (4) were identified as candidate microrefugia Siskiyou Mountains to western Siskiyou north of the southern bend of the Klamath (Figure 3a). Many important old-growth Crest region, California; and a network River; (5) West Siskiyous; (6) Lower Scott forest microrefugia occur in close prox- of serpentine-substrate areas representing Bar River; (7) Russian Wilderness; (8) imity to existing protected areas, such assemblages of endemic plant species and Lower Trinity River (multiple locations); as the Kalmiopsis Wilderness. Most of their surrounding forest buffers mainly in and (9) Middle Eel/Yolla Bolly (multiple the candidate microrefugia lie towards southwest Oregon. locations, numbers correspond to Figure the western, wetter part of the ecoregion 2). The Russian Wilderness was selected and are generally located at mid and low This provisional network of priority due to the extraordinary sympatric assem- elevations. Remnants of late-seral forest in climate change microrefugia outside the blage of conifer species whose presence the fog zone are particularly important to existing reserve system should be targeted could be due to mesorefugia conditions. protect and restore, as they likely contain for immediate protection and restoration. The mesorefugia located in the coastal a sizable proportion of vulnerable species. A variety of conservation approaches is zone experiences the highest rainfall in Some old-growth forest blocks at higher required because candidate sites are in the KSE and is likely to have the highest elevation in the eastern part of the ecore- diverse locations, habitat types, tenures, concentrations of restricted-range and cli- gion were also recommended, as they span and land use pressures. Some are located mate change-vulnerable species (contrast a broad elevational range and are among within active federal and state forestry Figures 1 and 2). We acknowledge that the largest remaining old-growth fragments zones, and some are on private lands. The much KSE biodiversity occurs outside in the ecoregion. Data on late-seral forests priority areas identified here would not, by of these mesorefugia, but suggest that were unavailable for some portions of the themselves, constitute a comprehensive ensuring adequate protection of habitats ecoregion, such as the southwestern coastal conservation strategy as they are intended buffered from warming in these zones is hills and the foothills of the Central Valley primarily to buffer a good portion of the an important first step. that may contain additional microrefugia KSE biota from extinction and extirpation (Figure 3a). due to changing climate, and they would not The current protected areas system does necessarily address a wide range of other a poor job of representing the provisional Finer-resolution analyses and field sur- conservation goals and objectives. mesorefugia. Only the Kalmiopsis, Sis- veys within priority areas (Figure 3a) are kiyou, Russian, and Middle Eel/Yolla required to identify the particular blocks MICROREFUGIA AND PROTECTED Bolly Wilderness areas and redwood parks of old-growth forest and bottomland sites AREAS encompass portions of likely mesorefu- that have the highest potential to act as gia. In general, Wilderness areas largely microrefugia. The nature of the landscape protect higher elevations, not the middle and the mosaic of late-seral forests can have Representation and Existing and lower slopes where most of the mi- a major influence on the efficacy of mi- Protected Areas crorefugia are likely to occur. Proposed crorefugia. For example, even a relatively expanded reserve networks would repre- small old-growth forest fragment situated We also intersected remaining late-seral sent all of the provisional mesorefugia, if in a steep, north-facing canyon that experi- forests with north-facing slopes (N, NE, implemented, including those at lower and ences shade most of the time will likely NW, Figure 3ab) and areas of relatively middle elevations (e.g., contrast Figures function well as a long-term refuge for high precipitation with microrefugia char- 2 vs. 3b). We also propose three priority mesophilic species. acteristics (see Appendix for methods). Us- mesorefugia corridors to link: (1) Siskiyou ing ecoregion-scale data on forest cover and Crest–Kalmiopsis, (2) Kalmiopsis–Sis- In sum, several microrefugia deserve topography, it was challenging to identify kiyou Mountains, and (3) Trinity/Scott Bar immediate conservation attention, includ- the small river valleys and bottomlands that River–Siskiyou Crest (numbers correspond ing: southern bend of the Klamath River, consistently pool cooler air and may func- to Figure 3a). California; lower slopes of the Klamath tion as additional microrefugia. More local- River from around China Point eastwards scale data and on-the-ground surveys are to Hamburg, California; northern slope required to identify potential bottomland PRIORITIZING MICROREFUGIA of the Scott Bar Mountains and along the refugia. For similar reasons, we also did not lower Scott River in California; old-growth attempt to identify potential microrefugia We used our microrefugia site features to fragments close to the coast in Oregon for the vulnerable serpentine flora. identify a set of provisional areas outside and in the foothills behind the redwood extant protected areas that warrant im- belt in northwestern California; north- Based on this analysis, the current protected mediate conservation attention. For the facing slopes of the Middle Smith River, area network under-represents most of the portion of the ecoregion outside of for- California; larger old-growth pockets to important microrefugia for the KSE (Figure mal protected areas, 22 highest-priority the west of the Kalomiopsis Wilderness, 3a). For instance, only 16% of remaining microrefugia and 40 high-priority areas southwest Oregon; southeastern watersheds old-growth forest occurs within strictly containing late-seral forest and other key of the Siskiyou Mountains (e.g., Dillon protected areas (Table 1). Some important
Volume 32 (1), 2012 Natural Areas Journal 69 as are not identified. The mature forest shown is north-fac- rity and white ovals are high priority gaps in protected areas r to locations described in the text) of formal protected areas in e Klamath-Siskiyou Ecoregion, southwest Oregon and northern Figure Figure 3. (A) Provisional microrefugia (highest priority and priority) and proposed mesorefugia corridors outside (numbers refe the Klamath-Siskiyou Ecoregion, southwest Oregon and northern California. High-quality microrefugia inside formal protected are ing Circles only. were drawn based on visual inspection of the mapped old forest polygons. (B) Some priority microrefugia in th California, which are not encompassed in the proposed expanded reserve network (Noss et al. 1999). Black ovals are highest prio coverages. Circles were drawn by visual inspection of old-growth forest concentrations.
70 Natural Areas Journal Volume 32 (1), 2012 Table 1. Area and percentage of north-facing late-seral and old-growth forest (collectively LSOG) in the Klamath-Siskiyou Ecoregion (KSE), southwest Oregon and northern California, by climate change vulnerability zones, extant protected areas, and proposed conservation areas. Data for old-growth forests for some portions of the ecoregion (southwestern coastal, southeastern border) were unavailable; thus, coastal old-growth forest area, inside and outside of protected areas, is underestimated and the drier zone old-growth forest area to a smaller extent.
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Volume 32 (1), 2012 Natural Areas Journal 71 Representation and Proposed assemblages in order to refine the location ACNKOWLEDGMENTS Protected Areas of mesorefugia and better design represen- tative networks of microrefugia. Potential We greatly appreciate the contributions to Prior reserve designs proposed for the refugia for the endemic serpentine flora this analysis by the following individuals: KSE include the Phase 1 reserves and need to be identified and prioritized. Such J. Cokendolpher, L. Farley, J.A. Johnstone, Representation Zones proposed by Noss areas are likely to be mesic serpentine sites J.M Ledford, R.A. Progar, J. Sawyer, et al. (1999), the Siskiyou Crest National that remain relatively moist even under a and W. Peterman. We thank KS Wild for Monument (KS Wild 2010) and Siskiyou changing climate, due to terrain position permission to use their ecoregion map as Wild Rivers (Sisikyou Project 2010) pro- and other biophysical features (e.g., seeps a basemap, S. Harrison for reviewing an posed by conservation groups, and Scenario and bogs). The sites and their surround- earlier draft, K.E. Iron, and an anonymous 3 Plan “interacting current and near-future ing buffer habitats need to be identified reviewer. This study was sponsored by a habitat” of Carroll et al. (2010). These and prioritized using a similar approach grant to the Geos Institute from The 444-S reserves, if implemented, would protect as for the old-growth forest microrefugia. Foundation. The opinions expressed are a large proportion of the critical micro- Identifying and protecting microrefugia those of the authors and do not necessarily refugia within high-precipitation zones complements ongoing modeling of range reflect the views of 444-S Foundation. and mesorefugia (i.e., 47% the remaining shifts for vulnerable species and natural old-growth forest and 22% of north-facing communities (e.g., Pearson and Dawson old-growth; Table 1). If all the proposed 2003; Loarie et al. 2008; Carroll et al. David Olson is a Conservation Biologist reserve expansions were implemented, 2010; Damschen et al. 2010; Harrison et with Conservation Earth Consulting. then 70% of remaining old-growth forest al. 2010), studies of climate sensitivity of and 33% of north-facing old-growth for- species, analyses of how a changing climate Dominick A. DellaSala is Chief Scientist est would be protected. We identified only will affect wide-ranging species, and as- and President of the Geos Institute, Ash- five gaps of highest priority and six high sessing the cost and cost-effectiveness of land, Oregon, and current President of the priority microrefugia that were not fully alternative conservation actions. Society for Conservation Biology, North contained within the proposed protected America Section. area network (Figure 3b). All of the highest CONCLUSION priority gaps are critically important sites James R. Strittholt is the Executive Direc- and should receive immediate conservation Large natural landscapes and wilderness, tor and Chief Scientist of the Conservation attention. Collectively, these gaps contain the foundation of reserve designs, remains Biology Institute, Corvallis, Oregon. important coastal and intact old growth the mainstay of conservation efforts in this areas, local pockets of species endemism, and many other localities and is especially Reed F. Noss is the Davis-Shine Professor and transitional areas; and they may pro- important in a changing climate. Without vide additional mesorefugia corridors. at the Department of Biology, University of large natural landscapes in relatively good Central Florida, Orlando, and the Presi- condition, many of the remaining pockets dent and Chief Scientist for the Florida NEXT STEPS of old-growth forest may not persist or Institute for Conservation Science. function well as microrefugia. However, Additional and more finely resolved prior- for ensuring a robust reserve design that is Jamie Kass is a GIS specialist and con- ity setting of microrefugia is warranted in responsive to climate change, it is prudent sultant. the near future. GIS-based spatial analyses to secure priority old-growth forest micro- supported by field evaluation of candidate refugia as swiftly as possible while the Thomas F. Allnutt is a Conservation Spe- microrefugia can assess their species more time-consuming and uncertain task cialists with the Department of Environ- assemblages, landscape context, terrain of conserving larger landscapes continues. mental Science, Policy and Management, position, habitat condition, defensibility, Waiting decades for formal “gazettement” University of California, Berkeley. and complementarity with other candidate of large protected areas without securing sites. These evaluations can be augmented microrefugia now may allow continued Marni E. Koopman is a Climate Change by additional analyses of past and future degradation of these critical refuges. Our refugia based on species distributions and Scientist with the Geos Institute, Ashland, recommended approach is somewhat novel biophysical predictors of climatic and Oregon. vegetation stability and identification of for most conservation advocacy, where areas predicted to experience wildfires securing larger priority landscapes pro- within historic ranges of frequency and posed in comprehensive strategies is often LITERATURE CITED intensity. In addition, targeted surveys acted upon first, but the rapidly warming of old-growth forest invertebrates and landscape may require a diversification of Ackerly, D.D., S.R. Loarie, W.K. Cornwell, S.B. Weiss, H. Hamilton, R. Branciforte, non-vascular plants (e.g., fungi, lichens, tactics. As Voltaire cautioned, we should and N.J.B. Kraft. 2010. The geography of bryophytes) are needed to improve our not let the perfect be the enemy of the climate change: implications for conserva- understanding of the distribution of distinct good. tion biogeography. Diversity and Distribu-
72 Natural Areas Journal Volume 32 (1), 2012 tions: (doi:10.1111/j.1472-642.2010.00654. D.M. Meko. 1998. North–South precipita- servation in old-growth Northwest forests. x) 1-12. tion patterns in Western North America on American Zoologist 33:578-587. interannual-to-decadal timescales. Journal Alexander, J., N. Seavy, C. Ralph, and B. Hogo- Loarie S.R., B.E. Carter, K. Hayhoe, S. Mc- of Climate 11:3095-3111. boom. 2006. Vegetation and topographical Mahon, R. Moe, C.A. Knight, and D.D. correlates of fire severity from two fires in Dobrowski, S.Z. 2010. A climatic basis for Ackerly. 2008. Climate change and the the Klamath-Siskiyou Region of Oregon and microrefugia: the influence of terrain on cli- future of California’s endemic flora. PLOS California. International Journal of Wildland mate. Global Change Biology: (doi:10.1111/ 1(3):e2502, 1-10. Fire 15:237-245. j.1365-2486.2010.02263.x). Loarie, S.R., P.B. Duffy, H. Hamilton, G.P. As- Ashcroft, M.B. 2010. Identifying refugia from Frest, T.J., and E.J. Johannes. 1993. Mollusc ner, C.B. Field, and D.D. Ackerly. 2009. The climate change. Journal of Biogeography Species of Special Concern within the velocity of climate change. Nature 462:1052- 37:1407-1413. range of the Northern Spotted Owl with 1057. (doi:10.1038/nature08649). an addendum addressing new management Briggs, T.S. 1969. A new Holarctic Family of Mead, L.S., D.R. Clayton, R.S. Nauman, D.H. options proposed in June, 1993. Final report Laniatori Phalangids. Pan-Pacific Entomolo- Olson, and M.E. Pfrender. 2005. Newly to Forest Ecosystem Management Working gist 45:35-50. discovered populations of salamanders Group, U.S. Department of Agriculture, Briggs, T.S. 1971a. Relict harvestmen from the from Siskiyou County, California represent Forest Service, Deixis Consultants, Seattle, a species distinct from Plethodon stormi. Pacific Northwest. Pan-Pacific Entomologist Wash. 47:165-178. Herpetologica 61:158-177. Gardner, M.R., and R.M. Shelley. 1989. New Briggs. T.S. 1971b. The harvestman of family Moilanen A., B.A. Wintle, J. Elith, and M. records, species, and genera of Caseyid Burgman. 2006. Uncertainty analysis for Triaenonychidae in North America (Oplion- millipedes from the Pacific Coast of es). Occasional Papers of the California regional-scale reserve selection. Conserva- North America (Diplopoda: Chordeuma- tion Biology 20:1688-1697. Academy of Sciences 90:1-43. tida: Caseyidae). Pan-Pacific Entomologist Bury, R.B. 1973. Western Plethodon: system- 65:177-268. Mote, P., and E.P. Salathé. 2009. Future climate atics and biogeographic relationships of in the Pacific Northwest. Climatic Change Harrison, S., H.D. Safford, J.B. Grace, J.H. 102:29-50. the Elongatus group. HISS News Journal Viers, and K.F. Davies. 2006. Regional 1:56-57. and local species richness in an insular en- Murphy, J.M., D.M.H. Sexton, D.N. Barnett, Carroll, C., J.R. Dunk, and A. Moilanen. 2010. vironment, serpentine plants in California. G.S. Jones, M.J. Webb, M. Collins, and Optimizing resiliency of reserve networks to Ecological Monographs 76:41-56. D.A. Stainforth. 2004. Quantification of modelling uncertainties in a large ensemble climate change: multispecies conservation Harrison, S., E.I. Damschen, and J.B. Grace. planning in the Pacific Northwest. Global of climate change simulations. Nature 2010. Ecological contingency in the 430:768-772. Change Biology 16:891-904. effects of climatic warming on forest Chen, J.S., S.C. Saunders, T.R. Crow, R.J. herb communities. PNAS: doi:10.1073/ Niwa, C.G., and R.W. Peck. 2002. Influence of Naiman, K.D. Brosofske, G.D. Mroz, B.L. pnas.1006823107. prescribed fire on carabid beetle (Carabidae) and spider (Araneae) assemblages in forest Brookshire, and J.F. Franklin. 1999. Micro- Jiang, H., J.R. Strittholt, P.A. Frost, and N.C. climate in forest ecosystems and landscape litter in southwestern Oregon. Environmen- Slossera. 2004. The classification of late tal Entomology 31:785-796. ecology. BioScience 49:288-297. seral forests in the Pacific Northwest, USA Cokendolpher, J.C., R.W. Peck, and C.G. Niwa. using Landsat ETM+ imagery. Remote Sens- Noss, R.F. 2001. Beyond Kyoto: forest man- 2005. Mygalomorph spiders from southwest- ing of Environment 91:320-331. agement in a time of rapid forest change. ern Oregon, USA with descriptions of four Conservation Biology 15:578-590. new species. Zootaxa 1058:1-34. Johnstone, J.A., and T.E. Dawson. 2010. Cli- matic context and ecological implications of Noss, R.F., J.R. Strittholt, K. Vance-Borland, C. Coleman, R.G., and A.R. Kruckeberg. 1999. summer fog decline in the coast redwood Carroll, and P. Frost. 1999. A conservation Geology and plant life of the Klamath- region. PNAS 10:4533-4538. plan for the Klamath-Siskiyou ecoregion. Siskiyou Mountain region. Natural Areas Natural Areas Journal 19:392-411. Journal 19:320-340. Koopman, M.E., R.S. Nauman, B.R. Barr, Olson, D.M. 1992. The northern spotted owl Daly, C., R.P. Neilson, and D.L. Phillips. 1994. S.J. Vynne, and G.R. Hamilton. 2009. Projected future conditions in the Klamath conservation strategy: implications for Pa- A statistical-topographic model for mapping cific Northwest invertebrates and associated climatological precipitation over mountain- Basin of Southern Oregon and Northern California. National Center for Conserva- ecosystem processes. Final Report prepared ous terrain. Journal Applied Meteorology for the Northern Spotted Owl EIS Team, 40- 3:140-158. tion and Science Policy, Climate Change Leadership Initiative, MAPPS Team of U.S. 04HI-2-1650, The Xerces Society and U.S. Damschen, E.I., S. Harrison, and J.B. Grace. Department of Agriculture, Forest Service, Department of Agriculture, Forest Service, 2010. Climate change-effects on an endemic- Ashland, Ore. Portland, Ore. rich edaphic flora: resurveying Robert H. Kruckeberg, A.R. 1984. 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Volume 32 (1), 2012 Natural Areas Journal 73 W.W. Wettengel, Y. Kura, P. Hedao, and biological integrity in the Klamath-Siskiyou Taylor, A.H., and C.N. Skinner. 2003. Spatial K. Kassem. 2001. Terrestrial ecoregions ecoregion using multiple spatial scales. and temporal patterns of historic fire regimes of the world: a new map of life on Earth. WWF & Conservation Biology Institute, and forest structure as a reference for res- BioScience 51:933-938. Ashland, Ore. toration of fire in the Klamath Mountains. Olson, D.M., M. O’Connell, R. Rayburn, Y-C. Staus, N.L., J. R. Strittholt, and D.A. DellaSala. Ecological Applications 13:704-719. Fang, and J. Burger. 2009. Managing for 2010. Evaluating areas of high conserva- Tecklin, D, D.A. DellaSala, F. Luebert, and P. climate change within protected area land- tion value in western Oregon with a deci- Pliscoff. 2011. Valdivia temperate rainforests scapes. Natural Areas Journal 29:501-506. sion-support model. Conservation Biology of Chile and Argentina. Pp. 132-153 in D.A. Pearson, R.G., and T.P. Dawson. 2003. Pre- 24:711-720. DellaSala, ed., Temperate and Boreal Rain- dicting the impacts of climate change on Staus, N.L., J.R. Strittholt, D.A. DellaSala, forests of the World: Ecology and Conserva- the distribution of species: are bioclimate and R. Robinson. 2002. Rate and pattern tion. Island Press, Washington, D.C. envelope models useful? Global Ecology of forest disturbance in the Klamath-Sis- Vance-Borland, K. 1999. Physical habitat clas- and Biogeography 12:361-371. kiyou ecoregion, U.S.A. Landscape Ecology sification for conservation planning in the Rull, V. 2009. Microrefugia. Journal of Bioge- 17:455-470. Klamath Mountains region. M.S. thesis, ography 36:481-484. Stebbins, G.L., and J. Major. 1965. Endemism Oregon State University, Corvallis. Rull, V. 2010. On microrefugia and cryptic and speciation in the California flora. Eco- Vicente, F. 2010. Micro-invertebrates conser- refugia. Journal of Biogeography 37:1623- logical Monographs 35:1-35. vation: forgotten biodiversity. Biodiversity 1625. Steele, C.A., and A. Storfer. 2006. Coales- Conservation: (doi:10.1007/s10531-010- 9898-6). Salathé, E.P. Jr, R. Steed, C.F. Mass, and P.H. cent-based hypothesis testing supports Zahn. 2008. A high-resolution climate model multiple Pleistocene refugia in the Pacific Wagner, D.H. 1997. Klamath-Siskiyou region, for the US Pacific Northwest: mesoscale Northwest for the Pacific giant salamander California and Oregon, USA. Pp. 74-76 in feedbacks and local responses to climate (Dicamptodon tenebrosus). Molecular Ecol- S.D. Davis, V.H. Heywood, O. Herrera-Mac- bryde, J. Villa-Lobos, and A.C. Hamilton, change. Journal of Climate 21:5708-5726. ogy 15:2477-2487. eds., Centres for Plant Diversity. Volume 3: Sawyer, J.O. 2007. Why are the Klamath Moun- Strittholt, J.R., and D.A. DellaSala. 2001. Im- The Americas. World Wildlife Fund for Na- tains and adjacent north coast floristically portance of roadless areas in biodiversity ture and IUCN (World Conservation Union), diverse? Fremontia 35:3-11. conservation in forested ecosystems: case Information Press, Oxford, U.K. study of the Klamath–Siskiyou ecoregion Whittaker, R.H. 1960. Vegetation of the Sis- Siskiyou Project. 2010. Siskiyou Wild Rivers of the United States. Conservation Biology kiyou Mountains, Oregon and California. Protection and Community Enhancement 15:1742-1754. Campaign. Available online
Appendix.
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74 Natural Areas Journal Volume 32 (1), 2012