Novel adaptation to hawkmoth pollinators in Clarkia reduces efficiency,not attraction of diurnal visitors

Background and Aims

Plant populations experiencing divergent pollination environments may be under selection to modify floral traits in ways that increase both attractiveness to and efficiency of novel pollinators. These changes may come at the cost of reducing overall effectiveness of other pollinators. The goal of this study was to examine differences in attractiveness and efficiency between Clarkia concinna and C. breweri, sister species of annual plants with parapatric distributions.

Methods

An assessment was made as to whether observed differences in visitors between natural populations are driven by differences in floral traits or differences in the local pollination environment. Differences in floral attractiveness were quantified by setting out arrays of both species in the geographical range of each species and exposing both species to nocturnal hawkmoths (Hyles lineata) in flight cages. Differences in visitor efficiency were estimated by measuring stigma–visitor contact frequency and pollen loads for diurnal visitors, and pollen deposition on stigmas for hawkmoths.

Key Results

The composition of visitors to arrayed plants was similar between plant species at any particular site, but highly divergent among sites, and reflected differences in visitors to natural populations. Diurnal insects visited both species, but were more common at C. concinna populations. Hummingbirds and hawkmoths were only observed visiting within the range of C. breweri. Despite attracting similar species when artificially presented together, C. concinna and C. breweri showed large differences in pollinator efficiency. All visitors except hawkmoths pollinated C. concinna more efficiently.

Conclusions

Differences in the available pollinator community may play a larger role than differences in floral traits in determining visitors to natural populations of C. concinna and C. breweri. However, floral traits mediate differences in pollinator efficiency. Increased effectiveness of the novel hawkmoth pollinator on C. breweri comes at relatively little cost in attractiveness to other visitors, but at large cost in their efficiency as pollinators.     

Responses of Cyperus brevifolius (Rottb.) Hassk. and Cyperus kyllingia Endl. to varying soil water availability

Different responses of Cyperus brevifolius and Cyperus kyllingia to varying soil water regimes were examined to explain their successful existence in a diverse range of habitats throughout the year in Indonesia. Thirty 43-day sprouts of each species were grown in three soil conditions, namely drought, field capacity, and flooding under greenhouse conditions. Plant height, leaf length, tiller number, and flower number were measured twice a week, from 45 to 98 days after sowing (DAS), while the other 12 traits were recorded at the end of the observation time. Ten out of the twelve traits were substantially influenced by the soil water content. Both species exhibited their best growth, production, and reproduction under field capacity conditions, and these traits were greatly subdued under drought conditions. Under drought conditions, both species manifested reduced growth and leaf expansion; however, stomatal aperture and frequency did not exhibit strong response to the soil water content. C. brevifolius showed a significantly greater biomass production and reproductive traits in field capacity and flooded conditions, but under drought conditions, growth was greatly hampered and only vegetative propagation occurred. On the other hand, C. kyllingia showed a higher tolerance to drought conditions, indicated by both a higher biomass and a higher number of flowers. The results obtained suggest that survival ability when faced with drought conditions was more apparent in C. kyllingia than in C. brevifolius, but where there was sufficient soil water, C. brevifolius was more prolific. This could be the explanation for the dominance of C. brevifolius in flooded areas and during the rainy season, and the occurrence of C. kyllingia in a wider range of habitats throughout the year.     

Effects of moderate high-temperature stress on photosynthesis in three saplings of the constructive tree species of subtropical forest

During the exposition to moderate high-temperature stress, photosynthetic rates and fluorescence of chlorophyll a were measured with a photosynthetic measurement system (Li-Cor 6400) and leaf chamber fluorometer (Li-Cor6400 LCF), respectively, in leaves of saplings, sun-adapted species (Schima superba), shade-adapted species (Cryptocarya concinna), and in mesophytic plant (Castanopsis hystrix) (42°C). The results showed that moderate high-temperature stress led to a decrease in F_v/F_m, namely the primary photochemical quantum efficiency, indicating that moderate high-temperature stress causes a partial inhibition of PSII. It also showed that such an effect was more severe in the shade-adapted plant C. concinna than in the sun-adapted species S. superba. However, except for the sun-grown leaves of C. concinna, the moderate high-temperature stress increased the photosynthetic rate of leaves at high light intensity. Simultaneously, less photoinhibition was found to occur under high-light intensity, suggesting that the capacity of resistant-photoinhibition was stimulated by moderate high-temperature stress. The quantum yield of PSII (ϕ_PSII) decreased in the sun-grown leaves of S. superba and C. hystrix but did not show any significant change in leaves of the shade plant C. concinna and shade-grown leaves of sun plant S. superba or the mesophytic plant C. hystrix because they already had a very low ϕ_PSII under this condition. Moderate high-temperature stress led to a decrease in ϕ_PSII/ϕ_CO2 ratios, an estimate of the quantum requirement for CO₂ assimilation, in the sun plant S. superba and the mesophytic plant C. hystrix because they were associated with the dissipation of a lower fraction of excitation energy. However, no significant changes were found in shade plant C. concinna and in shade-grown leaves of the other examined plants. The effect of moderate high-temperature (42°C) on photosynthesis depends on species and leaf type (sun and shade leaves) in the saplings of subtropical broad-leaved trees.     

The physiological light response of two tree species across a hydrologic gradient in Brazilian savanna (Cerrado)

Tropical savanna ecosystems are extremely diverse and important for global carbon storage. In the state of Mato Grosso, tropical savanna (locally known as the Cerrado), turns from well-drained, upland areas into seasonally flooded areas within the Pantanal; however, the Cerrado and the Pantanal share many common tree species, such as Vochysia divergens, a flood-adapted tree native to the Amazon Basin, and Curatella americana, a tree, adapted native to the welldrained the Cerrado. We measured the photosynthetic light response of these species in the the Cerrado and the Pantanal over a 1-year period to determine how these species physiologically adjust to these hydrologically distinct habitats. We hypothesized that neither species would experience a significant decline in maximum, light-saturated photosynthetic rate (P _max) in their naturalized habitat. Physiological performance of each species was generally higher in the habitat that they were adapted to; however, our data indicated that both species have broad tolerance for seasonal variations in hydrology, allowing them to tolerate seasonal drought during the dry season in the Cerrado, and seasonal flooding during the wet season in the Pantanal. In V. divergens, flexible water-use efficiency, higher specific leaf area (SLA), and a greater ability to adjust mass-based P _max (P _max,m) to variations in leaf N and P concentration appeared to be key traits for withstanding prolonged drought in the Cerrado. In C. americana, increases in SLA and higher nutrient-use efficiency appeared to be important in maintaining high rates of P _max,m in the seasonally flooded Pantanal. Flexibility in physiology and resource-use efficiency may allow these species to survive and persist in habitats with broadly differing hydrology.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

Floral volatiles fromClarkia breweri andC. concinna (Onagraceae): Recent evolution of floral scent and moth pollination

Clarkia breweri (Onagraceae) is the only species known in its genus to produce strong floral fragrance and to be pollinated by moths. We used gas chromatography-mass spectrometry (GC-MS) to identify 12 abundant compounds in the floral headspace from two inbred lines ofC. breweri. These volatiles are derived from two biochemical pathways, one producing acyclic monoterpenes and their oxides, the other leading from phenylalanine to benzoate and its derivatives. Linalool and linalool oxide (pyran form) were the most abundant monoterpenoids, while linalool oxide (furan form) was present at lower concentrations. Of the aromatic compounds detected, benzyl acetate was most abundant, whereas benzyl benzoate, eugenol, methyl salicylate, and vanillin were present as minor constituents in all floral samples. The two inbredC. breweri lines differed for the presence of the additional benzenoid compounds isoeugenol, methyleugenol, methylisoeugenol, and veratraldehyde. We also analyzed floral headspace fromC. concinna, the likely progenitor ofC. breweri, whose flowers are odorless to the human nose. Ten volatiles (mostly terpenoids) were detected at low concentrations, but only when headspace was collected from 20 or more flowers at a time. Trans--ocimene was the most abundant floral compound identified from this species. Our data are consistent with the hypothesized recent evolution of floral scent production and moth pollination inC. breweri.     

Summer Precipitation Predicts Spatial Distributions of Semiaquatic Mammals

Climate change is predicted to increase the frequency of droughts and intensity of seasonal precipitation in many regions. Semiaquatic mammals should be vulnerable to this increased variability in precipitation, especially in human-modified landscapes where dispersal to suitable habitat or temporary refugia may be limited. Using six years of presence-absence data (2007–2012) spanning years of record-breaking drought and flood conditions, we evaluated regional occupancy dynamics of American mink (Neovison vison) and muskrats (Ondatra zibethicus) in a highly altered agroecosystem in Illinois, USA. We used noninvasive sign surveys and a multiseason occupancy modeling approach to estimate annual occupancy rates for both species and related these rates to summer precipitation. We also tracked radiomarked individuals to assess mortality risk for both species when moving in terrestrial areas. Annual model-averaged estimates of occupancy for mink and muskrat were correlated positively to summer precipitation. Mink and muskrats were widespread during a year (2008) with above-average precipitation. However, estimates of site occupancy declined substantially for mink (0.56) and especially muskrats (0.09) during the severe drought of 2012. Mink are generalist predators that probably use terrestrial habitat during droughts. However, mink had substantially greater risk of mortality away from streams. In comparison, muskrats are more restricted to aquatic habitats and likely suffered high mortality during the drought. Our patterns are striking, but a more mechanistic understanding is needed of how semiaquatic species in human-modified ecosystems will respond ecologically in situ to extreme weather events predicted by climate-change models.     

Photosynthetic response of four fern species from different habitats to drought stress: relationship between morpho-anatomical and physiological traits

Ferns flourish in many habitats, from epiphytic to terrestrial and from sunny to shady, and such varied conditions require contrasting photosynthetic strategies to cope with drought. Four species of temperate ferns from different habitats were subjected to drought by withholding irrigation in order to investigate their photosynthetic responses. Lepisorus thunbergianus (epiphytic) had low stomatal density and showed high water-use efficiency (WUE) retaining photosynthetic activity with low relative frond water content under drought stress, which suggested their high adaptation to drought. On the other hand, low WUE with low light-saturated photosynthetic rate in Adiantum pedatum (terrestrial, shady environment) was associated with much lower photosynthesis than in the other species under drought stress, suggesting lower adaptation to drought-prone habitats. Morphological stomatal traits such as stomatal density and photosynthetic response to drought in ferns involved species-specific adaptation to survive and grow in their natural habitats with different levels of drought.     

Does human-induced habitat transformation modify pollinator-mediated selection? A case study in Viola portalesia (Violaceae)

Pollinator-mediated selection is one of the most important factors driving adaptation in flowering plants. However, as ecological conditions change through habitat loss and fragmentation, the interactions among species may evolve in new and unexpected directions. Human-induced environmental variation is likely to affect selection regimes, but as yet no empirical examples have been reported. In the study reported here, we examined the influence of human-induced habitat transformation on the composition of pollinator assemblages and, hence, pollinator-mediated selection on the flower phenotype of Viola portalesia (Violaceae). Our results indicate that pollinator assemblages differed substantially in terms of species composition and visitation rate between nearby native and transformed habitats. Similarly, the insect species that contributed most to visitation rates differed between plant populations. While the magnitude and sign of pollinator-mediated selection on flower length and width did not differ between sites, selection for flower number lost significance in the transformed habitat, and a significant pattern of disruptive selection for flower shape, undetected in the native habitat, was present in the transformed one. Overall, the results of this study suggest that human-induced habitat change may not only modify the species composition of pollinator assemblages, relaxing the selection process on some flower characters, but they may also create new opportunities for fitness-trait covariation not present in pristine conditions.     

Effects of three flower field types on bumblebees and their pollen diets

Agri-environment schemes, like flower fields, have been implemented in the EU to counteract the dramatic decline of farmland biodiversity. Farmers in Lower Saxony, Germany, may receive payments for three flower field types: annual, perennial (five years old), and mixed flower fields composed of yearly alternating annual and biannual parts. We assessed the effectiveness of these flower field types in providing bumblebee foraging habitat compared to control cereal fields. We sampled bumblebees with transect walks and assessed the richness of exploited pollen plants using DNA meta-barcoding and direct observations.All flower field types enhanced bumblebee abundance and species richness compared to control fields but attracted mostly three generalist species. Although we expected highest benefits from the more heterogeneous mixed flower fields, abundance was highest in annual, only intermediate in mixed, and lowest in perennial flower fields. Bumblebee species richness did not differ between flower field types.Overall, the proportion of sown plants in pollen loads was surprisingly low (< 50%). Bombus pascuorum, but not B. terrestris agg., exploited 10% of the sown plant species in perennial, 36% in annual and 45% in mixed flower fields, respectively. Compared to direct observations, pollen samples revealed 4.5 times more visited plant species and thus assessed floral resource use more reliably. Plant species richness in pollen loads decreased with local flowering plant species richness and increased with proportion of annual crops in the landscape, potentially due to the exploitation of more diverse and scattered resources, including flowering crops, in homogenized landscapes to fulfil dietary requirements.Our results indicate that under the current management, both annual and mixed flower fields provide the most attractive food resources, while perennial flower fields offered the poorest foraging habitats. Conclusively, flower fields seem important but resources from the surrounding landscape are still needed to sustain bumblebees in agricultural landscapes.     

Agave turneri (Agavaceae), a new species from northeastern Baja California, Mexico

Agave turneri, a new species of Agave from the Sierras Cucapá and El Mayor in northeastern Baja California, Mexico, is a medium-sized species that does not produce offsets, has a relatively short and narrow panicle, and has a distinctive flower structure. The closest relatives to this new species are Agave moranii, which occurs approximately 200 km to the south of the type locality, and A. deserti var. simplex, which occurs in Arizona and California. This new species is a narrow endemic restricted to specific granodiorite and tonalite habitats in a hyperarid environment. Agave turneri appears to be a critically endangered owing to its habitat preference for specific types of granite in the Sierra Cucapá, threats due to prolonged drought and global change, and its close proximity to the Mexicali metropolitan area.     

Comparative Population Dynamics of Two Closely Related Species Differing in Ploidy Level

Background

Many studies compare the population dynamics of single species within multiple habitat types, while much less is known about the differences in population dynamics in closely related species in the same habitat. Additionally, comparisons of the effect of habitat types and species are largely missing.

Methodology and Principal Findings

We estimated the importance of the habitat type and species for population dynamics of plants. Specifically, we compared the dynamics of two closely related species, the allotetraploid species Anthericum liliago and the diploid species Anthericum ramosum, occurring in the same habitat type. We also compared the dynamics of A. ramosum in two contrasting habitats. We examined three populations per species and habitat type. The results showed that single life history traits as well as the mean population dynamics of A. liliago and A. ramosum from the same habitat type were more similar than the population dynamics of A. ramosum from the two contrasting habitats.

Conclusions

Our findings suggest that when transferring knowledge regarding population dynamics between populations, we need to take habitat conditions into account, as these conditions appear to be more important than the species involved (ploidy level). However, the two species differ significantly in their overall population growth rates, indicating that the ploidy level has an effect on species performance. In contrast to what has been suggested by previous studies, we observed a higher population growth rate in the diploid species. This is in agreement with the wider range of habitats occupied by the diploid species.     

Space Use Variation in Co-Occurring Sister Species: Response to Environmental Variation or Competition?

Coexistence often involves niche differentiation either as the result of environmental divergence, or in response to competition. Disentangling the causes of such divergence requires that environmental variation across space is taken into account, which is rarely done in empirical studies. We address the role of environmental variation versus competition in coexistence between two rodent species: Rhabdomys bechuanae (bechuanae) and Rhabdomys dilectus dilectus (dilectus) comparing their habitat preference and home range (HR) size in areas with similar climates, where their distributions abut (allopatry) or overlap (sympatry). Using Outlying Mean Index analyses, we test whether habitat characteristics of the species deviate significantly from a random sample of available habitats. In allopatry, results suggest habitat selection: dilectus preferring grasslands with little bare soil while bechuanae occurring in open shrublands. In sympatry, shrubland type habitats dominate and differences are less marked, yet dilectus selects habitats with more cover than bechuanae. Interestingly, bechuanae shows larger HRs than dilectus, and both species display larger HRs in sympatry. Further, HR overlaps between species are lower than expected. We discuss our results in light of data on the phylogeography of the genus and propose that evolution in allopatry resulted in adaptation leading to different habitat preferences, even at their distribution margins, a divergence expected to facilitate coexistence. However, since sympatry occurs in sites where environmental characteristics do not allow complete species separation, competition may explain reduced inter-species overlap and character displacement in HR size. This study reveals that both environmental variation and competition may shape species coexistence.     

Thermal plasticity of mitochondria: A latitudinal comparison between Southern Ocean molluscs

Mitochondrial volume density (Vv_(mt,f)), cristae surface density (Sv_(im,mt)), cristae surface area (Sv_(im,f)) and citrate synthase (CS) activity were analysed as indicators of thermal acclimation in foot muscle of the limpet, Nacella concinna, and the clam, Laternula elliptica, collected from 4 locations within the Southern Ocean, South Georgia (54°S, N. concinna only), Signy (60°S), Jubany (L. elliptica only ? 62°S) and Rothera (67°S). Animals were acclimated to 0.0 °C whilst a sub-set of N. concinna (South Georgia, Signy and Rothera) and L. elliptica (Rothera) were acclimated to 3.0 °C. At 0.0 °C N. concinna had higher Vv_(mt,f), Sv_(im,mt), Sv_(im,f) and muscle fibre specific CS activity than L. elliptica which correlated with the more active life style of N. concinna. However, mitochondrial density was very low, 1–2% in both species, suggesting that low temperature compensation of mitochondrial density is not a universal evolutionary response of Antarctic marine ectotherms. Both Sv_(im,mt) and Sv_(im,f) were reduced by warm acclimation of N. concinna. South Georgia N. concinna maintained muscle fibre specific CS activity after acclimation, in contrast to N. concinna from Rothera and Signy and L. elliptica from Rothera, indicating that they have the physiological plasticity to respond to their warmer, more variable thermal environment.     

Diet of the mara (Dolichotis patagonum), food availability and effects of an extended drought in Northern Patagonia (Mendoza,Argentina)

The mara is a near threatened Caviomorph, endemic to Argentina. Studies on feeding ecology allow assessment of the dietary adaptability of maras to habitat changes. The mara diet and food availability on two sampling sites, belonging to distinct landscapes of Northern Patagonia, were estimated using microhistological analysis and point-quadrat transects, over four seasons, and besides during an extended drought. Significant differences were detected by Kruskall-Wallis ANOVA, feeding selection by the χ² test, and dietary preferences by Bailey's confidence interval. Grasses dominated food availability, with Panicum and Poa as major species, followed by shrubs and scarce forbs. Plant cover and forbs increased in spring and summer. The drought caused a strong decrease in plant cover and proportion of grasses. Maras ate all grass species, most forbs and several shrubs. Grasses dominated the diet, with Poa and Panicum being the major species, supplemented by the shrubs Lycium and Prosopis. Maras ate more grasses and forbs in spring and summer, and shrubs in autumn and winter. More shrubs and forbs, and less grasses, were eaten during the drought. Plant categories were used selectively only in autumn and winter, and in the drought period, with preference for shrubs and avoidance of grasses. Bromus, Poa, Plantago and Prosopis were preferred, and Panicum avoided. The mara qualified as a grazer but shifted to a mixed feeder during the drought. Dolichotis patagonum shared habitats with several big and medium-sized herbivores and showed the highest dietary similarities with plain vizcachas, brown hares and horses. Protective measures for natural habitats are needed, given that increasing impacts on food resources and habitat quality could be threats to the survival of maras and other wild vertebrates in Northern Patagonia.     

Do clonal growth form and habitat origin affect resource-induced plasticity in Tibetan alpine herbs?

To investigate how growth form and habitat origin affect phenotypic plasticity to resource supply in the Tibetan alpine herbs, the phalanx-type species Stipa capillacea and the guerilla-type species Carex montis-everestii were sampled from two different habitats (alpine steppe and alpine scrubland) and grown under three levels of light intensity and two levels of nutrient supply. Interspecific differences in light-induced plasticity were detected only in number of ramets, specific leaf area and leaf sheath length. Plasticity in plant biomass, number of ramets and rhizome length in response to light intensity differed between the two habitats. Stipa plants were more plastic than Carex plants in number of ramets and specific leaf area in response to light intensity. Carex plants from the alpine scrubland expressed greater light-induced plasticity in plant biomass and number of ramets than those from the alpine steppe, and Stipa plants showed less interhabitat differences in plasticity, which may be closely related to their contrasting growth forms. Clonal growth form and habitat origin affected nutrient-induced plasticity in none of the measured traits. It may be the guerilla growth form that makes Carex plants more efficiently adapted to highly heterogeneous light conditions in scrubland, and less habitat-dependent plasticity contributes to success of the phalanx-type Stipa plants in alpine habitats. The results are discussed in the context of foraging for heterogeneously distributed essential resources and adaptation to habitat origin.     

Temperature relations of photosynthetic response in populations of Verbascum thapsus L.

Summary Seedlings representative of Verbascum thapsus L. populations from thermally diverse habitats were grown under uniform, controlled conditions. The plants were used to obtain temperature response curves for net photosynthesis over a range of 15–40°C. In general, all experimental plants exhibited similar rates of net photosynthesis at 20, 25, 30, and 35°C. Plants representative of cool habitat populations (high-latidude and high-altitude) had greatest rates of net photosynthesis at the lower temperatures and much lower rates at 40°C. Plants representative of warm habitat populations (low-latitude and low-altitude) exhibited rates of net photosynthesis at 40°C which were nearly twice those of plants representative of cool habitat populations. Carbon dioxide transfer resistances are discussed with reference to plant control of photosynthesis at different temperatures. Patterns of photosynthesis and resistance response among plants representative of different habitats suggest ecotypic variation has occurred only to a very limited extent. Therefore, the patterns exhibited by experimental plants suggest that Verbascum thapsus' success in a number of diverse sites is related to the ability of all members of the species to photosynthesize over a broad range of temperatures.     

Effect of simulated macroalgae on the fish assemblage associated with a temperate reef system

Increased habitat complexity is supposed to promote increased diversity, abundance and biomass. This study tested the effect of the macroalgal cover on temperate reef fishes by mimicking macroalgae on artificial reefs in NW Sicily (Mediterranean Sea). Macroalgal cover affected reef fishes in different ways and independently of intrinsic temporal trends. The fish assemblages of manipulated and control artificial reef units differed in the relative abundances of the associated species, but little in species composition. In line with studies in seagrass habitats, fishes were most abundant in reefs covered by artificial macroalgae. Three species (Boops boops, Serranus scriba and Symphodus ocellatus) exhibited consistently greater abundance on vegetated reef units than on control reef units. The total number of species and the abundance of three particular species (S. scriba, S. ocellatus and Thalassoma pavo) displayed temporal trends which were independent on short and large temporal scales. Only fish total biomass and one species (Spicara flexuosa) displayed strong effects of interaction among the experimental factors. Mechanisms to explain these findings are discussed from observational evidence on habitat use and interactions among multiple species. This study highlights that manipulative experiments involving repeated sampling of fish in artificial habitats appear to be a valid approach to study fish-habitat relationships in fluctuating environments. It is also concluded that macroalgae mimics may serve as a tool for restoring lost marine vegetated habitats when current human-induced conditions prevent the recovery of pristine macroalgal stands.     

Thermal Reaction Norms and the Scale of Temperature Variation: Latitudinal Vulnerability of Intertidal Nacellid Limpets to Climate Change

The thermal reaction norms of 4 closely related intertidal Nacellid limpets, Antarctic (Nacella concinna), New Zealand (Cellana ornata), Australia (C. tramoserica) and Singapore (C. radiata), were compared across environments with different temperature magnitude, variability and predictability, to test their relative vulnerability to different scales of climate warming. Lethal limits were measured alongside a newly developed metric of “duration tenacity”, which was tested at different temperatures to calculate the thermal reaction norm of limpet adductor muscle fatigue. Except in C. tramoserica which had a wide optimum range with two break points, duration tenacity did not follow a typical aerobic capacity curve but was best described by a single break point at an optimum temperature. Thermal reaction norms were shifted to warmer temperatures in warmer environments; the optimum temperature for tenacity (T_opt) increased from 1.0°C (N. concinna) to 14.3°C (C. ornata) to 18.0°C (an average for the optimum range of C. tramoserica) to 27.6°C (C. radiata). The temperature limits for duration tenacity of the 4 species were most consistently correlated with both maximum sea surface temperature and summer maximum in situ habitat logger temperature. Tropical C. radiata, which lives in the least variable and most predictable environment, generally had the lowest warming tolerance and thermal safety margin (WT and TSM; respectively the thermal buffer of CT_max and T_opt over habitat temperature). However, the two temperate species, C. ornata and C. tramoserica, which live in a variable and seasonally unpredictable microhabitat, had the lowest TSM relative to in situ logger temperature. N. concinna which lives in the most variable, but seasonally predictable microhabitat, generally had the highest TSMs. Intertidal animals live at the highly variable interface between terrestrial and marine biomes and even small changes in the magnitude and predictability of their environment could markedly influence their future distributions.