Sunday, 6 December 2015

Columbine Graveyards

Serpentine Columbine (Aquilegia eximia)
Photo: David A. Hofmann (Creative Commons)

The oak savannah and Chamise (Adenostoma fasciculatum) chaparral of California's North Coast Ranges, are interrupted here and there by a unique and altogether surprising floral community: plants that grow on poison. In this case, the poison is serpentine, rocks that are so rich in magnesium and iron that they, and their associated soils, are toxic to most plants. Most, but not all.

There are some plants that can grow on serpentine deposits and many of those are rare and endemic, not to mention highly adapted, making serpentine flora one of a most intriguing element in California's generous biological endowment. Among the most exquisite serpentine plants is Serpentine Columbine (Aquilegia eximia), which displays large red and yellow flowers to attract the attention of pollinators. In addition to pollinators, Serpentine Columbine attracts great many other insects, but for a totally different reason.

Plants attract animals to help them with all kinds of tasks; the two most obvious, of course, are pollination and seed dispersal. Pollinators are attracted by scents and visually stimulating flowers. Take the elaborate deceptions of the Fly Orchid (Ophyrys insectifera), which wafts bee pheromone-like scents from its bee-shaped flowers. Real bees come not in search of pollen or nectar, as they might at a more conventional flower, but instead they come to mate with the lookalike blossom, in the process getting coated in pollen. The bees pollinate the next orchid they visit in another misguided hope for sex.


Some plants are entirely dependent on animals for dispersing their seeds. In the Rocky Mountains, Clark's Nutcrackers (Nucifraga columbiana) are the near-exclusive disperser of Whitebark Pine (Pinus albicaulis) seeds, transporting them great distances and planting them in suitable habitats.

Examples of animal pollinators and seed dispersers are virtually limitless, but there are lots of other reasons for plants to attract animals. Sometimes it's to eat them. Venus's Flytrap (Dionaea muscipula), is probably the most dramatic of the so-called carnivorous plants, capturing and later digesting insects and even small frogs between snap tap-like leaves. Even the ubiquitous Field Thistle (Cirsium discolor) may capture insects using sticky secretions on its flowers; those same secretions act as digestive enzymes, digesting stuck insects and providing an unusual food source, at least among thistles species.

Other plants feed off animals in more passive ways. The impressive Queen of the Andes (Puya raimondii) offers paramo birds a safe haven among its hooked leaves in exchange for the highly nutritious droppings the birds leave behind. As an added benefit to the plant, a bird occassionally gets hooked among the thorny leaves, dying and providing an even richer source of nutrients for the giant bromeliad.

Some plants attract animals to help them battle damaging herbivores. Azteca ants are provided with living spaces in Ceropia trees and are fed from extrafloral necataries, in exchange for doing battle against caterpillars and other damaging herbivores. Providing shelter and nectar is a pretty conventional way of attracting helpful predators, but there are a few plants, including the Serpentine Columbine, that attract protective insects in a completely different way, and that's by essentially becoming arthropod graveyards.

Serpentine Columbine stems are covered in glandular hair-like trichomes, making them very sticky. So sticky, that they trap insects by the dozens. These trapped insects in turn attract predatory arthropods. The predators come to dispatch trapped living insects or to feed on the corpses of those that have already died. The
predators are called upon to primarily combat caterpillars of the Darker Spotted Straw Moth (Heliothis phloxiphaga), which feed on the leaves, buds and even the flowers. The columbine's glandular hairs seem to be of little use in combating this caterpillar, so the plants rally Checker-rimmed Bugs (Pselliopus spinicollis), other true bugs (Order Hemiptera) and even the occasional crab spider (Mecaphesa spp), to help stavse off assault. Between meals of caterpillar, these predators feast on the stuck insects. This buffet style call to arms seems to be effective, columbines with more stuck insects (thus more helpful predators) usually experience greater reproductive success.

The insects that Serpentine Columbines capture aren't, for the most part, pollinators or herbivores that accidentally become stuck, but instead appear to be actively attracted by the plants through some kind of chemical signal. What exactly that signal is remains unclear, but it appears that Serpentine Columbine is the only plant so far known to actively attract insects in this way. Yet another remarkable find from the fascinating serpentine deposits of northern California!