Climate-Related, Decadal-Scale Assemblage Changes of Seagrass-Associated fishes in the Northern Gulf of Mexico
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Global Change Biology (2010) 16, 48–59, doi: 10.1111/j.1365-2486.2009.01889.x Climate-related, decadal-scale assemblage changes of seagrass-associated fishes in the northern Gulf of Mexico F. JOEL FODRIE*w , KENNETH L. HECK, JR *w ,SEANP.POWERS*w ,WILLIAMM. GRAHAM*w and K E L LY L . R O B I N S O N *w *Department of Marine Sciences, University of South Alabama, Mobile AL 36688, USA, wDauphin Island Sea Lab, Dauphin Island, AL 36528, USA Abstract Global temperatures are rising, and are expected to produce a poleward shift in the distribution of many organisms. We quantified changes in fish assemblages within seagrass meadows of the northern Gulf of Mexico (GOM) between the 1970s and 2006–2007, and observed changes consistent with this forecast. During 2006–2007 we sampled seagrass meadows using the same gears and methods previously employed by R. J. Livingston in coastal waters of northwest Florida throughout the 1970s. Comparisons between datasets revealed numerous additions to the fish fauna during 2006–2007 that were completely absent in the 1970s, including: Lutjanus synagris (lane snapper), Epinephelus morio (red grouper), Chaetodon ocellatus (spotfin butterflyfish), Mycteroperca sp (grouper, non gag), Centropristis philadelphica (rock sea bass), Fistularia tabacaria (bluespotted cornetfish), Ocyurus chrysurus (yellowtail snapper), Thalassoma bifasciatum (bluehead wrasse), Abu- defduf saxatilis (sergeant major), Acanthuridae spp. (surgeonfishes) and Sparisoma viride (stoplight parrotfish). Several other species showed large increases in abundance during the interval between 1979 and 2006, including Mycteroperca microlepis (gag grouper, up 200 Â ), Lutjanus griseus (gray snapper, up 105 Â ), and Nicholsina usta (emerald parrotfish, up 22 Â ). All of these are tropical or subtropical species that now make up a greater percentage of seagrass-associated fish assemblages in the northern GOM than in the past. Additionally, we observed regional increases in air and sea surface temperatures (43 1C) during the 30 years that separate Livingston’s samples and ours that correlate with northern shifts in the distribution of warm-water fishes. Documenting these range shifts is a critical first step in investigating the consequences of global warming for endemic marine communities and fishery production in the northern GOM. Keywords: climate change, community composition, geographic range shifts, ichthyofauna, seagrass, species distributions, temperature records Received 29 December 2008 and accepted 19 January 2009 ot et al., 2003); and poleward shifts in the range/ Introduction distribution of species (Perry et al., 2005). Over the Recent increases in global temperatures are expected to short-term, range expansions may increase local biodi- drive concurrent changes in the composition and ecol- versity as poleward-advancing species outpace pole- ogy of terrestrial and marine communities worldwide ward-retreating species (Hickling et al., 2006). Over (McCarty, 2001). The climate-related mechanisms be- longer timescales, however, poleward range expansions hind these community transitions may include altered may have consequences similar to those now being primary production resulting from increased CO2 or observed with nonnative species invasions, modifying photostress (Richardson & Schoeman, 2004); phenolo- local dynamics of competition, predation, herbivory gical changes in flowering, nesting or migration events and parasitism (C. Sorte, personal communication). (Ro- There is a growing literature documenting cases of range or distribution shifts involving taxa as diverse as Correspondence: F. Joel Fodrie, Mobile AL 36688, tel. 1 251 861 fishes (Perry et al., 2005), planktonic forams (Field et al., 7551, e-mail: [email protected] 2006) and butterflies (Parmesan et al., 1999). While these 48 r 2009 Blackwell Publishing Ltd TROPICAL FISH IN THE NORTHERN GULF OF MEXICO 49 cases can be correlated with regional warming, both representation of tropical or subtropical species that Sagarin et al. (1999) and Schiel et al. (2004) have noted would indicate a northern shift in the distribution of difficulties in directly linking current warming to fishes during this time; and (2) for species that have the dynamics and consequences of range shifts. For exited (those collected only by Livingston) or entered instance, many expectations regarding the rate and (those collected only by us) the fish community over the consequences of range shifts are drawn from ecosystem last 30 years, are the average northern range limits of models, known biotic responses to temperature, biolo- species in each of these groups equal? Additionally, we gical responses to decadal or interglacial climate cycles, explored climate conditions that might have affected and point counts separated over several decades. Un- seagrass fauna in the northern GOM from 1971 to 2007 fortunately, high-resolution historical data from closely using daily minimum air and sea surface temperature monitored reference sites that could be used to more (SST) data from the National Oceanographic and Atmo- rigorously examine the relationship between tempera- spheric Administration National Climate Data Center ture and species’ distributions are largely absent. Thus, (NCDC), and National Data Buoy Center (NDBC), we ususally rely on opportunistic studies that correctly respectively. Using these temperature data we asked identify and explore range shifts to increase our pre- if, and to what degree, seasonal temperature thresholds dictive capacity for the effects of the potential continua- have been lifted for tropical or subtropical species due tion or acceleration of the 0.5 1C global temperature to increases in either duration (using air temperature) or increase observed over the last century (McCarty, 2001). intensity (using SST) of warm conditions? In the dis- Despite reports on many taxa, documentation of cussion that follows, we examine the potential effects range shifts within some groups or ecosystems remain newcomers may have on endemic estuarine food-webs, relatively sparse. For instance, seagrass meadows pro- and subsequently, the offshore fishery production that vide numerous ecosystem services and may be biologi- originates from seagrass nurseries. We also consider a cal sentinels of local and climatological perturbations number of alternative hypotheses that may explain the (Orth et al., 2006). Temperature, CO2 concentration, changes we observed, including short-term population sea-level rise, and precipitation all impact seagrasses fluctuations/cycles, changes in regional physical ocea- (some positively, some negatively) making them valu- nography that may have altered larval transport, indir- bale indicators of climate change. However, the lack of ect species interactions, and anthropogenic habitat long-term records for seagrass distributions, combined alteration or degradation. with the confounding effects of other anthropogenic stressors, make any robust assessments of range shifts Materials and methods difficult (Orth et al., 2006). Conversely, coastal sea- grasses have long been identified as valuable nurseries The GOM is bisected (north–south) by the Tropic of for fishes and crustaceans (Heck et al., 2003), and there Cancer, with temperate environments/habitats in the is a comparatively rich literature on the distribution and northern GOM and tropical environments/habitats in characteristics of seagrass-associated fish communities the southern GOM (becoming the Caribbean Sea). We (e.g., Pearson, 1929; Livingston, 1982). Temperature expected the interface between two climate zones to affects many vital rates in the life history of fishes provide an excellent location for examining climate- (i.e., growth, reproduction; Pauly, 1994), and thus their related range shifts of fishes. Additionally, the GOM is geographic distribution. For instance, Kimball et al. bounded by North America, and marine species/sub- (2004) concluded that the northern range limit of invad- populations that might otherwise retreat northward in ing Pterois volitans/miles (lionfish) would be Cape response to global warming lack a latitudinal refuge. Hatteras, NC, due to the temperature at which mass Therefore, the consequences of climate change in this mortality occurs (o10.0 1C). Therefore, fishes can be region may be especially dramatic. valuable indicators of climate perturbations. We analyzed changes over the last 30 years in Here, we report on potential northern range shifts ichthyofaunal communities within seagrass meadows of fishes in to the northern Gulf of Mexico (GOM), many of the northern GOM by comparing survey data of which only use seagrass habitat as juveniles. Our obtained during 1971–1979 by Livingston (1985) to data analyses of climate-related changes in seagrass fish from our own surveys in 2006–2007. Livingston pro- communities were based on comparisons between vides the best reference point for comparisons between trawl surveys conducted during 1971–1979 (Livingston, current fish communities and those from several 1985) and 2006–2007 within seagrass meadows of the decades ago within northern GOM seagrass meadows, northern GOM. We asked two climate-related questions but there are notable differences in the survey regimes. regarding differences in these assemblages: (1) has there Livingston’s collections were spatially restricted and been an increase over the last three decades in the temporally expansive, while our surveys covered a r 2009 Blackwell Publishing Ltd, Global Change Biology, 16, 48–59 50 F. J. FODRIE et al.