Behaviour broadens thermal safety margins on artificial coastal defences in the tropics.

Tropical species are predicted to be among the most vulnerable to climate change as they often live close to their upper limits to thermal tolerance and in many cases, behavioural thermoregulation is required to persist in the thermal extremes of tropical latitudes. In concert with warming temperatures, near-shore species are faced with the additional threat of shoreline hardening, leading to a reduction in microhabitats that can provide thermal refuges. This situation is exemplified in Singapore, which lies almost on the equator and so experiences year-round hot temperatures, and much of its coastline is now seawall. To investigate the thermal ecology of a common intertidal gastropod, Nerita undata, on these artificial structures, we measured thermal conditions on two seawalls, the temperatures of habitats occupied by the snail, and compared these with the snail's thermal tolerance by measuring heart rate and behavioural thermoregulation (as preferred temperature, Tpref). At one of the two seawalls (Tanjong Rimau), temperatures experienced by N. undata exceeded all measures of thermal tolerance in the sun, while at the other (Palawan Beach), they did not. Temperatures in habitats occupied by the snails on the seawalls were similar to their measured Tpref in the laboratory and were lower than all measures of thermal tolerance. Behavioural thermoregulation by the snails, therefore, significantly increased the thermal safety margins of N. undata on the relatively homogenous seawalls in Singapore, and at one of the two seawalls were necessary to allow snails to survive. Accordingly, to facilitate motile species to maintain broad thermal safety margins through behavioural regulation, the provision of additional refuges from thermal stress is recommended on artificial coastal defences such as seawalls.

Why are 'suboptimal' temperatures preferred in a tropical intertidal ectotherm?

In thermally extreme environments, it is challenging for organisms to maximize performance due to risks associated with stochastic variation in temperature and, subsequently, over evolutionary time minimizing the exposure to risk can serve as one of the mechanisms that result in organisms preferring suboptimal temperatures. We tested this hypothesis in a slow-moving intertidal snail on tropical rocky shores, where temperature variability increases with time from 30 min to 20 hr when recorded at 30 min intervals (due to short-term environmental autocorrelation where temperatures closer in time are more similar as compared to temperatures over a long period of time). Failure to accommodate temporal variation in thermal stress by selecting cool habitats can result in mortality. Thermal performance curves for different traits (heart rate and locomotion) were measured and compared to the snail's thermal preferences in both the field and laboratory. Predicted performances of the snails were simulated based on thermal performance curves for different traits over multiple time-scales and simulated carryover effects. A strong mismatch was found between physiological and behavioural thermal maxima of the snails (physiological thermal maximum being higher by ~7°C), but the snails avoided these maxima and sought temperatures 7-14°C cooler. Such a risk-averse strategy can be explained by their predicted performances where the snails should make decisions about preferred temperatures based on time periods ≥5 hr to avoid underestimating the temporal variation in body temperature. In extreme and stochastic environments, where the temporal variation in environmental conditions can lead to substantial divergence between instantaneous and time-averaged thermal performances, 'cooler is better' and 'suboptimal' body temperatures are preferred as they provide sufficient buffer to reduce mortality risk from heat stress.

Annual temperature variation as a time machine to understand the effects of long-term climate change on a poleward range shift.

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long-term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.

Rapid range shift in an introduced tropical marine invertebrate.

The barnacle Megabalanus coccopoma is native to shorelines from Baja California to Peru and has been introduced to a number of other locations including the Atlantic US SE coast, where it was first recorded in 2006. In 2009, the range of M. coccopoma in the SE US extended from Ft. Pierce, FL north to Cape Hatteras, NC with seasonal populations found as far north as Kitty Hawk, NC. During the exceptionally cold winter of 2009/2010, the range of M. coccopoma shifted dramatically due to the dieback of all monitored populations north of Florida. We examined body size, distribution, and density of M. coccopoma during the summers of 2010, 2011, and 2012 to describe the extent of the range retraction and the rate of range re-expansion. In 2010, recruits were found as far north as Tybee Island, Ga, but no established populations were found north of Florida. In 2011 recruits were found at Rodanthe, NC but established populations were still limited to Florida. By 2012 populations were established in Rodanthe, NC, slightly north of its previously known range limit. Estimated rates of range re-expansion were 255.8 km/yr in 2010 and 794.1 km/yr in 2011. Rates of re-expansion to the north in 2010 and 2011 were faster than have previously been reported for any marine species, and are one of the few rates published for any tropical marine invertebrate.