Urban ecosystems cover less than 3% of the Earth's land surface, yet more than half of the human population lives in urban areas. The process of urbanization stresses biodiversity and other ecosystem functions within and far beyond the city. To understand the mechanisms underlying observed changes in biodiversity patterns, several observational and experimental studies were performed in the metropolitan area of Phoenix, Arizona, and the surrounding Sonoran Desert. The first study was comprised of seven years of arthropod monitoring using pitfall traps in common urban land-use types. This study revealed differences in community structure, diversity and abundance over time and between urban and wildland habitats. Urban habitats with high productivity had higher abundances of arthropods, but lower diversity compared to wildland habitats. Arthropod abundance in less-productive urban habitats was positively correlated with precipitation, but abundance in high-productivity urban habitats was completely decoupled from annual fluctuations in precipitation. This study showed the buffering capacity and the habitat heterogeneity of urban areas. To test the mechanisms controlling community diversity and structure in urban areas, a major field experiment was initiated. Productivity of the native shrub Encelia farinosa and bird predation of associated arthropods were manipulated to test whether bottom-up or top-down forces were more important in urban habitats compared to wildland habitats. Abundance, richness and similarity were monitored, revealing clear differences between urban and wildland habitats. An unusually cold and dry first season had a negative effect on plant growth and arthropod abundance. Plants in urban habitats were relatively unaffected by the low temperature. An increase in arthropod abundance with water availability indicated bottom-up forces in wildland habitats, whereas results from bird exclusions suggested that bird predation may not be as prominent in cities as previously thought. In contrast to the pitfall study, arthropod abundance was lower in urban habitats. A second field experiment testing the sheltering effect of urban structures demonstrated that reduced wind speed is an important factor facilitating plant growth in urban areas. A mathematical model incorporating wind, water and temperature demonstrated that urban habitats may be more robust than wildland habitats, supporting the empirical results.
The reproductive ground plan hypothesis of social evolution suggests that reproductive controls of a solitary ancestor have been co-opted during social evolution, facilitating the division of labor among social insect workers. Despite substantial empirical support, the generality of this hypothesis is not universally accepted. Thus, we investigated the prediction of particular genes with pleiotropic effects on ovarian traits and social behavior in worker honey bees as a stringent test of the reproductive ground plan hypothesis. We complemented these tests with a comprehensive genome scan for additional quantitative trait loci (QTL) to gain a better understanding of the genetic architecture of the ovary size of honey bee workers, a morphological trait that is significant for understanding social insect caste evolution and general insect biology.
Results
Back-crossing hybrid European x Africanized honey bee queens to the Africanized parent colony generated two study populations with extraordinarily large worker ovaries. Despite the transgressive ovary phenotypes, several previously mapped QTL for social foraging behavior demonstrated ovary size effects, confirming the prediction of pleiotropic genetic effects on reproductive traits and social behavior. One major QTL for ovary size was detected in each backcross, along with several smaller effects and two QTL for ovary asymmetry. One of the main ovary size QTL coincided with a major QTL for ovary activation, explaining 3/4 of the phenotypic variance, although no simple positive correlation between ovary size and activation was observed.
Conclusions
Our results provide strong support for the reproductive ground plan hypothesis of evolution in study populations that are independent of the genetic stocks that originally led to the formulation of this hypothesis. As predicted, worker ovary size is genetically linked to multiple correlated traits of the complex division of labor in worker honey bees, known as the pollen hoarding syndrome. The genetic architecture of worker ovary size presumably consists of a combination of trait-specific loci and general regulators that affect the whole behavioral syndrome and may even play a role in caste determination. Several promising candidate genes in the QTL intervals await further study to clarify their potential role in social insect evolution and the regulation of insect fertility in general.