The evolutionary ecology of nut dispersal

Abstract

A variety of nut-producing plants have mutualistic seed-dispersal interactions with animals (rodents and corvids) that scatter hoard their nuts in the soil. The goals of this review are to summarize the widespread horticultural, botanical, and ecological literature pertaining to nut dispersal inJuglans, Carya, Quercus, Fagus, Castanae, Castanopsis, Lithocarpus, Corylus, Aesculus, andPrunus; to examine the evolutionary histories of these mutualistic interactions; and to identify the traits of nut-bearing plants and nut-dispersing rodents and jays that influence the success of the mutualism. These interactions appear to have originated as early as the Paleocene, about 60 million years ago. Most nuts appear to have evolved from ancestors with wind-dispersed seeds, but the ancestral form of dispersal in almonds (Prunus spp.) was by frugivorous animals that ingested fruit.

Nut-producing species have evolved a number of traits that facilitate nut dispersal by certain rodents and corvids while serving to exclude other animals that act as parasites of the mutualism. Nuts are nutritious food sources, often with high levels of lipids or proteins and a caloric value ranging from 5.7 to 153.5 kJ per propagule, 10–1000 times greater than most wind-dispersed seeds. These traits make nuts highly attractive food items for dispersers and nut predators. The course of nut development tends to reduce losses of nuts to insects, microbes, and nondispersing animals, but despite these measures predispersal and postdispersal nut mortality is generally high. Chemical defenses (e.g., tannins) in the cotyledons or the husk surrounding the nut discourage some nut predators. Masting of nuts (periodic, synchronous production of large nut crops) appears to reduce losses to insects and to increase the number of nuts dispersed by animals, and it may increase cross-pollination. Scatter hoarding by rodents and corvids removes nuts from other sources of nut predation, moves nuts away from source trees where density-dependent mortality is high (sometimes to habitats or microhabitats that favor seedling establishment), and buries nuts in the soil (which reduces rates of predation and helps to maintain nut viability). The large nutrient reserves of nuts not only attract animal dispersers but also permit seedlings to establish a large photosynthetic surface or extensive root system, making them especially competitive in low-light environments (e.g., deciduous forest) and semi-arid environments (e.g., dry mountains, Mediterranean climates). The most important postestablishment causes of seedling failure are drought, insufficient light, browsing by vertebrate herbivores, and competition with forbs and grasses. Because of the nutritional qualities of nuts and the synchronous production of large nut crops by a species throughout a region, nut trees can have pervasive impacts on other members of ecological communities. Nut-bearing trees have undergone dramatic changes in distribution during the last 16,000 years, following the glacial retreat from northern North America and Europe, and the current dispersers of nuts (i.e., squirrels, jays, and their relatives) appear to have been responsible for these movements.