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Tropical ectotherms are particularly vulnerable to global warming because their physiologies are assumed to be adapted to narrow temperature ranges. This study explores three mechanisms potentially constraining thermal adaptation to global warming in tropical insects: (a) Trade-offs in genotypic performance at different temperatures (the jack-of-all-trades hypothesis), (b) positive genetic covariance in performance, with some genotypes performing better than others at viable temperatures (the 'winner' and 'loser' genotypes hypothesis), or (c) limited genetic variation as the potential result of relaxed selection and the loss of genes associated with responses to extreme temperatures (the gene decay hypothesis). We estimated changes in growth and survival rates at multiple temperatures for three tropical rain forest insect herbivores (Cephaloleia rolled-leaf beetles, Chrysomelidae). We reared 2,746 individuals in a full sibling experimental design, at temperatures known to be experienced by this genus of beetles in nature (i.e. 10-35°C). Significant genetic covariance was positive for 16 traits, supporting the 'winner' and 'loser' genotypes hypothesis. Only two traits displayed negative cross-temperature performance correlations. We detected a substantial contribution of genetic variance in traits associated with size and mass (0%-44%), but low heritability in plastic traits such as development time (0%-6%) or survival (0%-4%). Lowland insect populations will most likely decline if current temperatures increase between 2 and 5°C. It is concerning that local adaption is already lagging behind current temperatures. The consequences of maintaining the current global warming trajectory would be devastating for tropical insects. However, if humans can limit or slow warming, many tropical ectotherms might persist in their current locations and potentially adapt to warmer temperatures.

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AbstractTropical mountains might protect species from global warming by facilitating biotic migrations upslope. Current predictions of tropical biotic responses to global warming are based on correlations between species elevational distributions and temperatures. Because biotic attritions, range shifts, and mountaintop extinctions result from complex demographic processes, predictive models must be based on mechanistic associations between temperature and fitness. Our study combines long-term temperature records with experimental demography to determine the contribution of local adaptation to organismal resilience in a warming world. On the Barva volcano in Costa Rica, Cephaloleia belti (Coleoptera: Chrysomelidae) displays high-elevation (960-2,100 m asl) and low-elevation (50-960 m asl) mitochondrial haplotypes. We reared haplotype cohorts at temperatures prevalent along the elevational gradient (i.e., 10°-30°C). Based on ambient temperatures recorded every half hour for 4 years, we projected average instantaneous population growth rates ([Formula: see text]) at current and future temperatures (i.e., +1° to 6°C) for each beetle haplotype. Haplotypes are adapted to local temperatures, but with a temperature increase beyond 2°C, both haplotypes will face lower-elevation demographic attritions and extinctions. Upper distribution limits serve as potential elevational refugia from global warming. This study shows how species resilience to global warming emerges from complex fitness responses of locally adapted phenotypes facing novel environments.

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Determining responses of organisms to changing temperatures is a research priority, as global warming threatens populations and ecosystems worldwide. Upper thermal limits are frequently measured as the critical thermal maximum (CTmax), a quick bioassay where organisms are exposed to increasing temperatures until individuals are not able to perform basic motor activities such as walking or flying. A more informative approach to understand organism responses to global warming is to evaluate how vital rates, such as growth or survival, change with temperatures. The main objectives of this study are: (1) to determine if factors affecting insect vital rates such as diet quality, developmental temperatures or acclimation also affect CTmax and (2) to determine if vital rates of different life stages (i.e., insect larvae or adults) display different responses to temperature changes. If different life stages have particular thermal requirements, this may indicate different susceptibility to global warming. This study focuses on Cephaloleia belti (Coleoptera: Chrysomelidae), a tropical insect currently expanding its diet to an exotic host plant. We determined how high and low-quality diets (i.e., native vs novel host), as well as exposure temperatures affect CTmax of adult beetles. We also estimated larval and adult survival when feeding on high and low quality host plants, when exposed to temperatures typical of the elevational distribution of this species, or when exposed to projected temperatures in 100 years. We did not detect an effect of diet quality or acclimation on CTmax. However, larvae and adults had different thermal requirements. CTmax is not affected by previous diet or acclimation as an adult. We propose that to understand processes involved in the adaptation and persistence of ectotherm populations in a warming world, studies must explore responses beyond CTmax, and focus on the response of vital rates to changing temperatures.

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Geographic isolation is the first step in insect herbivore diet specialization. Such specialization is postulated to increase insect fitness, but may simultaneously reduce insect ability to colonize novel hosts. During the Paleocene-Eocene, plants from the order Zingiberales became isolated either in the Paleotropics or in the Neotropics. During the Cretaceous, rolled-leaf beetles diversified in the Neotropics concurrently with Neotropical Zingiberales. Using a community of Costa Rican rolled-leaf beetles and their Zingiberales host plants as study system, we explored if previous geographic isolation precludes insects to expand their diets to exotic hosts. We recorded interactions between rolled-leaf beetles and native Zingiberales by combining DNA barcodes and field records for 7450 beetles feeding on 3202 host plants. To determine phylogenetic patterns of diet expansions, we set 20 field plots including five exotic Zingiberales, recording beetles feeding on these exotic hosts. In the laboratory, using both native and exotic host plants, we reared a subset of insect species that had expanded their diets to the exotic plants. The original plant-herbivore community comprised 24 beetle species feeding on 35 native hosts, representing 103 plant-herbivore interactions. After exotic host plant introduction, 20% of the beetle species expanded their diets to exotic Zingiberales. Insects only established on exotic hosts that belong to the same plant family as their native hosts. Laboratory experiments show that beetles are able to complete development on these novel hosts. In conclusion, rolled-leaf beetles are pre-adapted to expand their diets to novel host plants even after millions of years of geographic isolation.

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The critical thermal maximum (CTmax), the temperature at which motor control is lost in animals, has the potential to determine if species will tolerate global warming. For insects, tolerance to high temperatures decreases with latitude, suggesting that similar patterns may exist along elevational gradients as well. This study explored how CTmax varies among species and populations of a group of diverse tropical insect herbivores, the rolled-leaf beetles, across both broad and narrow elevational gradients. Data from 6,948 field observations and 8,700 museum specimens were used to map the elevational distributions of rolled-leaf beetles on two mountains in Costa Rica. CTmax was determined for 1,252 individual beetles representing all populations across the gradients. Initial morphological identifications suggested a total of 26 species with populations at different elevations displaying contrasting upper thermal limits. However, compared with morphological identifications, DNA barcodes (cytochrome oxidase I) revealed significant cryptic species diversity. DNA barcodes identified 42 species and haplotypes across 11 species complexes. These 42 species displayed much narrower elevational distributions and values of CTmax than the 26 morphologically defined species. In general, species found at middle elevations and on mountaintops are less tolerant to high temperatures than species restricted to lowland habitats. Species with broad elevational distributions display high CTmax throughout their ranges. We found no significant phylogenetic signal in CTmax, geography, or elevational range. The narrow variance in CTmax values for most rolled-leaf beetles, especially high-elevation species, suggests that the risk of extinction of insects may be substantial under some projected rates of global warming.

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The species of the Neotropical genus Cephaloleia Chevrolat, 1836 are revised. We present a key to the known larvae of Cephaloleia (8 species), a key to the 95 species known to occur in Mexico, Central America and the West Indies, and a key to the 138 species known to occur in South America. All identification keys were translated to Spanish. Descriptions for the 214 known species of Cephaloleia as well as illustrations for 212 species are presented. The following species are removed from Cephaloleia: C. bipartita Pic, 1926c is transferred to Hybosispa Weise, 1910; C. minasensis Pic, 1931 and C. viridis Pic, 1931 are transferred to Stenispa Baly, 1858. The following species are described as new: C. abdita sp. n. from Brazil; C. amba sp. n. from Colombia, Ecuador, and Peru; C. angustacollis sp. n. from Ecuador; C. brevis sp. n. from French Guiana; C. calathae sp. n. from Costa Rica; C. chica sp. n. from Peru; C. conforma sp. n. from Costa Rica; C. crenulata sp. n. from Ecuador; C. gemma sp. n. from Bolivia and Brazil; C. horvitzae sp. n. from French Guiana; C. interrupta sp. n. from Costa Rica; C. kressi sp. n. from Costa Rica; C. lenticula sp. n. from Ecuador, French Guiana, Peru, and Suriname; C. nana sp. n. from Ecuador; C. ochra sp. n. from Ecuador; C. stainesi sp. n. from Costa Rica; and C. susanae sp. n. from Brazil and Ecuador. Cephaloleia simoni Pic, 1934 is treated as Incertae sedis. The larvae of C. erichsonii Baly, 1858 and C. puncticollis Baly, 1885 are described and illustrated.

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A total of 301 adult hispine beetles of the genera Cephaloleia and Chelobasis were found in rolled leaves of plants of 17 species of Zingiberales (families Costaceae, Heliconiaceae, Maranthaceae, Musaceae, and Zingiberaceae) during a field study at La Gamba, Golfito region, Costa Rica. Of these beetles, Cephaloleia belti was recorded from 12 potential host plant species, C. distincta from 7, C. dilaticollis from 5, C., Chelobasis bicolor, C. championi, and C. histrionica from 3, Chelobasis perplexa and C. instabilis from 2, whereas C. trivittata from only one. Of the plant species, Heliconia latispatha had 7 beetle species in its leaf rolls, Calathea lutea had 5, H. imbricata and H. rostrata had 4, H. stricta and Musa paradisiaca had 3, H. wagneriana had 2, while on H. vaginalis, H. danielsiana, H. densiflora, H. longiflora, Calathea crotalifera, C. platystachya, Goeppertia lasiophylla, Alpinia purpurata, Costus pulverulentus and Costus barbatus, H. densiflora, H. vaginalis, and H. danielsana only hispines of one species were found. Cephaloleia belti occurred together with beetles of six other hispine species, whereas Cephaloleia trivittata never shared a leaf roll with another hispine species. The remaining beetle species aggregated with one to four other hispines. Adults of C. belti and C. championi were frequently seen, occasionally also with C. dilaticollis, C. histrionica, and Chelobasis perplexa, to co-occur with the carabid Calophaena ligata in the same leaf roll without any sign of interspecific aggression. A comparison of host choices and the phylogeny of the hispines and of their host plants revealed no signs that beetles used species level phylogenetic relationships within the Zingiberales to select food plants. Obviously, within this plant order, rolled-leaf hispines choose their plant hosts in a nearly opportunistic manner. Seemingly, they use differences among plants at higher taxonomic levels but within the Zingiberales, the availability of young - rolled - leaves might be the actual decisive factor.