BULLETIN OF MARINE SCIENCE, 81(2): 283–296, 2007 IS CLIMATE CHANGE CONTRIBUTING TO RANGE REDUCTIONS AND LOCALIZED EXTINCTIONS IN NORTHERN (HALIOTIS KAMTSCHATKANA) AND FLAT (HALIOTIS WALALLENSIS) ABALONES? Laura Rogers-Bennett ABSTRACT Abalone abundance surveys from the 1970s were repeated 30 yrs later follow- ing a period of increased sea surface temperatures along the Pacific coast of the United States. Northern abalone, Haliotis kamtschatkana (Jonas, 1845) once abun- dant enough to support commercial fishing in Washington and Canada, are now extremely rare in the southern portion of their range in southern and central Cali- fornia. They have also declined 10 fold in northern California in the absence of hu- man fishing pressure. In Washington, northern abalone are in decline and exhibit recruitment failure despite closure of the fishery. Flat abalone, Haliotis walallensis (Stearns, 1899) no longer occur in southern California, and in central California have declined from 32% to 8% of the total number of abalones, Haliotis spp., inside a marine reserve. The distribution of flat abalone appears to have contracted over time such that they are now only common in southern Oregon where they are sub- ject to a new commercial fishery. Given these range reductions, the long-term per- sistence of flat abalone and northern abalone (locally) is a concern in light of threats from ocean warming, sea otter predation, and the flat abalone fishery in Oregon. The likelihood of future ocean warming poses challenges for abalone restoration, suggesting that improved monitoring and protection will be critical, especially in the northern portions of their distributions. Range shifts towards the poles have been documented for a number of species in meta-analyses and these shifts are consistent with predictions of global warming (Walther et al., 2002; Parmesan and Yohe, 2003; Root et al., 2003). A growing num- ber of range shifts and contractions are documented from marine systems in which southern species replace northern species (Barry et al., 1995; Southward et al., 1995; Field et al., 2006). Northward shifts in species ranges (Zacherl et al., 2003) may not necessarily have negative impacts on abundance unless population shifts are unsuc- cessful due to biotic or abiotic factors limiting expansion. For example, black aba- lone, Haliotis cracherodii (Leach, 1814), populations could expand into the central California coast in response to oceanic warming, however, disease, sea otters, and poaching are acting synergistically to limit abundances in this region, north of Point Conception (Haaker et al., 1992; Raimondi et al., 2002; Harley and Rogers-Bennett, 2004). Concern is rising over the fate of abalone populations on the Pacific coast of the United States (see Transboundary Abalone Recovery Group, Vancouver, March 2007), sparking interest in the potential impacts of oceanic warming for restoration planning (Hobday and Tegner, 2002; Vilchis et al., 2005). Abalone populations have declined dramatically precipitating the closure of multiple abalone fisheries (Karpov et al., 2000; Rogers-Bennett et al., 2002), as well as the listing of pink, Haliotis corru- gata (Wood, 1828), green, Haliotis fulgens (Philippi, 1845), black, and northern, Hali- otis kamtschatkana (Jonas, 1845) abalone as species of concern <www.nmfs.noaa. Bulletin of Marine Science 283 © 2007 Rosenstiel School of Marine and Atmospheric Science of the University of Miami 284 BULLETIN OF MARINE SCIENCE, VOL. 81, NO. 2, 2007 gov/pr/species/concern/>, and white abalone, Haliotis sorenseni (Bartsch, 1940), as endangered (Federal Register 66: 103, June 2001). Recent work has focused on the po- tential impact of ocean warming on abalone populations (Tegner et al., 2001; Hobday and Tegner, 2002; Vilchis et al., 2005) and its implications for successful restoration. Determining the impacts of ocean warming is complicated by confounding factors such as fishing. One way to discern the impacts of ocean warming, as distinct from fishery influences, is to examine population trends in unfished species (Moser et al., 2000), such as flat abalone Haliotis walallensis (Stearns, 1898), and populations in- side marine reserves (Roberts and Hawkins, 1999). The distribution of the northern subspecies, Haliotis kamtschatkana kamtschat- kana (Jonas 1845), extends from southeast Alaska to Point Conception where the subspecies morphology converges with the southern threaded subspecies, Haliotis kamtschatkana assimilis, (Dall, 1878) which extends from Point Conception south into northern Baja California, Mexico (Geiger, 1999). Northern abalone and the for- mer threaded abalone were determined to be subspecies of H. kamtschatkana be- cause of little divergence between the two with respect to sperm lysin (Geiger, 1998). The threaded subspecies was taken in the commercial fishery (> 21,000 abalone) in southern California from 1971 to 1980 (Rogers-Bennett et al., 2002), but since then has remained scarce. All northern abalone fisheries are now closed along the Pacific coast including southeast Alaska (closed in 1995), British Columbia (closed in 1990), and Washington State (closed in 1994). Workshops have been convened to exam- ine prospects for rebuilding northern abalone stocks in British Columbia (Campbell, 2000) and Washington, (J. Gaydos and B. Peabody, SeaDoc Society, pers. comm.). Flat abalone, H. walallensis, have never been considered an abundant species (Mc- Millen and Phillips, 1974; Owen, 2006). The traditional range of this species is quite narrow, extending from Newport, Oregon to Baja California, Mexico, with the type specimen from Gualala, California (Geiger, 1999; Owen, 2006). Flat abalone were never targeted by fisheries until 2001 when a commercial fishery was opened in Or- egon [there is also a small recreational red abalone fishery in the state (S. Groth, Oregon Department of Fish and Wildlife, pers. comm.)]. Here, I examine northern and flat abalone along the Pacific coast of the United States, comparing recent data with historic records from 30 yrs ago to test whether there has been a shift in the southern range as would be predicted with ocean warm- ing. I focus on northern abalone which is a species of concern, has a broad geograph- ic distribution, and was once the basis of important commercial and recreational fisheries. Flat abalone has a narrow geographic distribution and has been unfished throughout its range (except recently in Oregon). The modern distribution of north- ern and flat abalone is assessed in relation to historic accounts of their distribution, abundances in marine reserves, and changes in water temperatures. Potential causes for reductions in range are discussed as are the implications of these range shifts for restoration. Methods Southern California.—In southern California, abalone populations are surveyed by the Kelp Forest Monitoring Program (KFMP), part of the Channel Islands National Parks Service. This program quantifies abalone density along transects once per year at multiple sites around the northern Channel Islands. Two band transects each 10 × 3 m are placed on ROGERS-BENNETT: ABALONE AND CLIMATE CHANGE 285 either side of a fixed 100 m line for a total search area of 60 m2. Detailed descriptions of their methods can be found elsewhere <www. nature.nps.gov>. Abalone numbers are also monitored annually within artificial habitats called Abalone Re- cruitment Modules (ARMs). Each module is composed of twenty ½ cinder blocks enclosed in a wire mesh cage stacked to leave an open core. The ½ cinder blocks are stacked in five layers of four bricks with a surface area of 5 m2 (see Davis, 1995). Once a year these “habitats” are disassembled and the numbers of juvenile and adult abalone inside are recorded. Between 80–100 of these habitats are sampled each year at three islands: Anacapa, Santa Cruz, and Santa Rosa Island. Central California.—In central California, we conducted abalone density estimates at the Hopkins State Marine Reserve (no fishing) in central California using the same methods as surveys in the 1970–1980s (Lowry and Pearse, 1973; Hines and Pearse, 1982). A fixed cable (270 m long) with meter demarcations still exists in the subtidal having remained since the earlier surveys. This cable allowed us to examine the same subtidal area (approximately 90 × 90 m) inside this small reserve as in the past. Dive teams started at positions 10 m apart start- ing at the 60 m cable mark and ending at the 150 m cable mark. Abalone were marked with yellow lumber crayon to prevent double counting. We estimate that our dive team searched approximately 800 m2 of crevice habitat during 15 hrs and 13 min of search time over two consecutive days each year. Density is often difficult to quantify within the range of sea otters, Enhydra lutris (Lin- naeus, 1758), in which abalone occupy cryptic habitats. In this study, abalone density was determined within crevice habitat (about 10% of the total area) as had been done previously (Hines and Pearse, 1982). Earlier surveys of flat abalone recorded their numbers as a propor- tion of the total number of abalone (flat and red abalone) rather then as density (m–2). Thus, we report our results using a similar metric to compare with historic records. Northern California.—California Department of Fish and Game divers conducted timed swim surveys searching for northern, flat, and red abalone in 1975 at popular abalone fishing sites in northern California. Included in this survey were sites at two State Parks: Van Damme in Mendocino County and Fort Ross in Sonoma County. During each survey, two-person dive teams conducted 20 min swims enumerating the species and size of the abalone they observed. Similarly, California Department of Fish and Game assessed subtidal benthic communities dur- ing a series of surveys funded by the Pacific Gas and Electric Company in preparation for a pro- posed nuclear power plant at Point Arena, Mendocino County (Gotshall et al., 1974). Subtidal surveys of abalone populations were included in the ecological surveys from 1971 and 1972. This survey used a combination of 30 × 2 m2 transects along with 30 m2 arcs.