Date of Award


Degree Name

MS in Biological Sciences


Biological Sciences


Matt Ritter


Our understanding of the natural world is constantly evolving and strengthening as more observations are made and experiments are performed. For example, we understand that tree stems grow toward the light (positive phototropism; Darwin 1880, Loehle 1986, Christie et al. 2013) and against gravity (negative gravitropism; Knight 1806, Hashiguchi et al. 2013). We also know that plants respond to mechanical stimulus and perturbation (thigmotropism; Braam 2005). Genes and their resulting proteins have been described to uncover some of the mechanisms for these environmental responses, but relatively speaking, we have just scratched the surface (Wyatt et al. 2013). While the discovery of the molecular mechanisms responsible for these behaviors is certainly dependent on the ever-improving lab technology available, every molecular discovery is dependent on a macroscopic observation.

In this manuscript I present the two novel macroscopic observations I made on members of Araucaria in the urban forest. The first describes a hemisphere-dependent lean in A. columnaris, and the second provides genetic and morphological evidence that hybrids exist between A. columnaris and A. heterophylla.

Araucaria columnaris (J.R. Forst.) Hooker, or the Cook Pine is a conifer with a narrow native range that has been cultivated worldwide and grows unlike any other tree known. The initial observation we made was that trees in California and Hawaii lean south, and trees in California lean to a greater extent than trees in Hawaii. Measuring 250 trees in 16 regions worldwide, however, produced statistically significant evidence for a hemisphere dependent directional leaning pattern. Trees in the northern hemisphere lean south, and trees in the southern hemisphere lean north. Additionally, the lean becomes more pronounced at greater distances from the equator.

We also gathered morphological and genetic evidence in the California urban forest that A. columnaris and A. heterophylla (Salisb.) Franco are hybridizing. Many individuals have intermediate characteristics of both species, which originally led me to believe that hybrids exist in cultivation. After analyzing several individuals with microsatellite genetic markers, I have enough evidence to conclude that hybrids between A. columnaris and A. heterophylla exist. This is an important observation mainly for municipalities and arborists interested in properly identifying trees in the urban forest. Knowing the proper identity of trees is imperative to informing decisions about their protection or removal.

As we continue to ask questions about the inner workings of nature we will continue to gain a better appreciation for what we still do not know. The evidence provided in this manuscript better informs our future questions about a leaning pattern in A. columnaris and about the history of the cultivation of Araucaria.

Award received:

Publishing in Journal "Ecology"