ABSTRACT
Relationships between genome size and environmental variables suggest that DNA content might be adaptive and of evolutionary importance in plants. The genus Larrea provides an interesting system to test this hypothesis, since it shows both intra- and interspecific variation in genome size. Larrea has an amphitropical distribution in North and South American deserts, where it is most speciose. Larrea tridentata in North America shows a gradient of increasing autopolyploidy; while three of the four studied South American species are diploids, Larrea divaricata, Larrea nitida, Larrea ameghinoi, and the fourth is an allopolyploid, Larrea cuneifolia. We downloaded available focal species' georeferenced records from seven data reservoirs. We used these records to extract biologically relevant environmental variables from WorldClim at 30 arc seconds scale, to have a broad characterization of the variable climatic conditions of both regions, and a climatic envelope for each species. We estimated relative DNA content index and relative monoploid genome values, by flow cytometry, of four most abundant Larrea species throughout their respective ranges. Then we winnow the bioclimatic dataset down to uncorrelated variables and sampled locales, to analyse the degree of association between both intra- and interspecific relative DNA content and climatic variables that are functionally relevant in arid environments using Pearson correlations, general linear and mixed effects models. Within the genus Larrea, relative DNA content increases with rising temperature and decreases with rising precipitation. At the intraspecific level, all four species show relative DNA content variation across climatic conditions. Larrea is a genus that shows genome size variation correlated with climate. Our results are also consistent with the hypothesis that extreme environmental pressures may have facilitated repeated whole genome duplication events in North America, while in South America, reticulate evolution, as allopolyploidization, and speciation might have been climate-dependent since the Oligocene.
ABSTRACT
Flower-dwelling predators make flowers dangerous foraging sites for pollinators, potentially affecting their anti-predator behaviour. Moreover, predation vulnerability often varies among pollinators' body sizes with interspecific comparisons showing that smaller species are more vulnerable than larger ones. However, how intraspecific body size variation influences pollinator behaviour under predation risk is still unknown, especially under natural conditions. We hypothesized that bumblebee workers of different sizes will exhibit different foraging strategies under predation risk. We predict that (a) small workers should more often exhibit anti-predator behaviours than larger workers. We also hypothesized that the anti-predator behaviour should be influenced by predator size and reward availability; therefore, we expect (b) higher avoidance behaviour towards larger predator sizes and (c) more and longer visits to inflorescences with high nectar availability. Finally, we expect that (d) nectar availability should overcome the anti-predator behaviour in less vulnerable, large, workers. We recorded flower visitation, time spent and rejection behaviours of different sizes of Bombus terrestris (Apidae) workers (large, medium and small) to inflorescences of Alstroemeria aurea (Alstroemeriaceae) with different treatments of artificial spiders (small and large) and nectar availability (with, without). Anti-predator and foraging behaviour of bumblebees was affected by the size of the worker, the presence of artificial spiders and nectar availability. Large and medium size bumblebees strongly reduced flower visitation and time spent in the presence of artificial spiders, consistently avoiding flowers with spiders, regardless of spider size or nectar availability. Instead, small bumblebees seldom modified their behaviour when facing artificial spiders, only increasing their avoidance or decreasing their foraging time in nectarless flowers hosting large artificial spiders. This pattern of larger workers being more sensitive to predation risk than smaller ones at the intraspecific level in B. terrestris is contrary to the expected and acknowledged trend based on previous interspecific comparisons, but partially consistent with predictions of models of optimal foraging theory. Intraspecific behavioural variability was uncovered only when nectar was available, whereas artificial predator size rarely modified bumblebee anti-predator and foraging behaviour. Therefore, our findings suggest that the trade-off between maximizing resource intake and minimizing predation risk strongly varies across bumblebee worker body sizes.
Resumen Los depredadores que cazan sobre flores hacen que éstas sean sitios peligrosos de forrajeo para los polinizadores, pudiendo afectar su comportamiento. En general, la vulnerabilidad a la depredación varía con el tamaño del polinizador, siendo en comparaciones interespecíficas las especies más pequeñas las más vulnerables. Sin embargo, aún se desconoce cómo la variación intraespecífica del tamaño corporal influye en el comportamiento del polinizador bajo riesgo de depredación, especialmente en condiciones naturales.. Esperamos que abejorros de distinto tamaño exhiban diferentes estrategias de alimentación ante el riesgo de depredación. Predecimos que (i) obreras pequeñas manifiesten con mayor frecuencia comportamientos anti depredadores que obreras más grandes. Conjuntamente, esperamos (ii) una mayor evasión hacia tamaños de depredadores más grandes, y (iii) más visitas y mayor permanencia en inflorescencias con néctar. Finalmente, esperamos que (iv) la disponibilidad de néctar prevalezca a la manifestación del comportamiento anti depredador en obreras grandes, supuestamente menos vulnerables. Registramos a campo el número de visitas, tiempo de permanencia y el comportamiento de evasión de diferentes tamaños de obreras de Bombus terrestris (Apidae) (grandes, medianas y pequeñas) a inflorescencias de Alstroemeria aurea (Alstroemeriaceae) con arañas artificiales (pequeñas, grandes) y distinta disponibilidad de néctar (con, sin). El comportamiento de los abejorros se vio afectado por su tamaño corporal, la presencia de arañas artificiales y la disponibilidad de néctar. Abejorros grandes y medianos disminuyeron considerablemente las visitas y el tiempo de permanencia en inflorescencias, evadiendo consistentemente las flores que albergaron arañas artificiales, independientemente del tamaño del modelo o la cantidad de néctar. En cambio, abejorros pequeños rara vez modificaron su comportamiento, aumentando la evasión y disminuyendo el tiempo de permanencia solo en flores sin néctar y con arañas artificiales grandes. El patrón encontrado a nivel intraespecífico en B. terrestris resultó ser opuesto al observado en comparaciones interespecíficas, pero parcialmente consistente con modelos de teoría de forrajeo óptimo. Esta variabilidad intraespecífica se manifestó solo cuando había néctar disponible, mientras el tamaño del depredador apenas modificó el comportamiento. Nuestros hallazgos sugieren que el balance entre maximizar la ingesta de recursos y minimizar el riesgo de depredación varía fuertemente con el tamaño corporal de los abejorros.
