The Dispersal of Ancient and Modern Apples by Humans and Other Megafauna

Crop domestication often involves selection for larger fruits. In some crops, humans took plant species with relatively small fruits and, over many generations of artificial selection, developed a plant with much larger fruits. Consider giant pumpkins as an extreme example. Yet in the case of apples, relatively large fruits already existed in the wild. Producing larger apples happened quickly and, perhaps even, unconsciously. Apples were practically primed for domestication, and as Robert Spengler explains in a paper published last year in Frontiers in Plant Science, looking back in time at the origins of the apple genus, Malus, can help us understand how the apple we know and love today came to be.

Apples are members of the rose family (Rosaceae), a plant family that today consists of nearly 5000 species. According to the fossil record, plants in the rose family were found in large numbers across North America as early as the Eocene (56 – 33.9 million years ago). They were present in Eurasia at this time as well, but Spengler notes, “there is a much clearer fossil record for Rosaceae fruits and seeds in Europe and Asia during the Miocene and Pliocene (20 – 2.6 million years ago).” Around 14 million years ago, larger fruits and tree-form growth habits evolved in Rosaceae subfamilies, giving rise to the genera Malus and Pyrus (apples and pears). Small, Rosaceae fruits were typically dispersed by birds, but as Sprengler writes, “it seems likely that the large fruits [in Malus and Pyrus] were a response to faunal dispersers of the late Miocene through the Pliocene of Eurasia.” Larger animals were being recruited for seed dispersal in a changing landscape.

Glacial advances and retreats during the Pleistocene (2.6 million – 11,700 years ago) brought even more changes. Plants with effective, long distance seed dispersal were favored because they were able to move into glacial refugium during glacial advances. Even today, these glacial refugium are considered genetic hot spots for Malus, and could be useful for future apple breeding. As the Pleistocene came to a close, many megafauna were going extinct. This continued into the Holocene. Large-fruited apple species lost their primary seed dispersers, and their ranges became even more contracted.

Humans have had an extensive relationship with apples, which began long before domestication. Foraging for apples was common, and seeds were certainly spread that way (perhaps even intentionally). Favorable growing conditions were also created when forests were cleared and old fields were left fallow. Apple trees are early successional species that easily colonize open landscapes, gaps in forests, and forest edges, so human activity that would have created such conditions “could have greatly promoted the spread and success of wild Malus spp. trees during the Holocene.”

The earliest evidence we have of apple domestication (in which “people were intentionally breeding and directing reproduction”) occurred around 3000 years ago in the Tian Shan Mountains of Kazakhstan, where Malus sieversii – a species that is now facing extinction – was being cultivated. This species was later brought into contact with other apple species, a few of which were also being cultivated, including M. orientalis, M. sylvestris, and M. baccata. These species easily hybridized, giving us the modern, domesticated apple, M. domestica. As Spengler writes, “the driving force of apple domestication appears to have been the trans-Eurasian crop exchange, or the movement of plants along the Silk Road.” Continued cultivation and further hybridization among M. domestica cultivars over the past 2000 years has resulted in thousands of different apple varieties.

The unique thing about domesticated apples is that their traits are not fixed in the same way that traits of other domesticated crops are. Growing an apple from seed will result in a very different apple than the apple from which the seed came. Apple traits instead have to be maintained through cloning, which is accomplished mainly through cuttings and grafting. Apples hybridize with other apple species so readily that most apple trees found in the wild are hybrids between wild and cultivated populations.

Spengler considers the study of apple domestication to be “an important critique of plant domestication studies broadly, illustrating that there is not a one-size fits-all model for plant domestication.” The “key” for understanding apple domestication “rests in figuring out the evolutionary driver for large fruits in the wild – seed dispersal through megafaunal mammals – and the process of evolution for these large fruits – hybridization.” He notes that “domestication studies often ignore evolutionary processes leading up to human cultivation,” which, in the case of apples, involves “hybridization events in the wild” that led to the evolution of large fruits “selected for through the success in recruiting large megafaunal mammals as seed disperses.” Many of those mammals went extinct, but humans eventually assumed the role, selecting and propagating “large-fruiting hybrids through cloning and grafting – creating our modern apple.”

Excerpt from Fruit from the Sands by Robert N. Spengler:

Indeed, the relationship between apples and people is close and complex, spanning at least five millennia. The story of the apple begins along the Silk Road… In recent years genetic studies have resolved much of the debate over these origins. Nevertheless, the ancestry of the apple is highly complex. Cloning, inbreeding, and reproduction between species have created a genealogy that looks more like a spider’s web than a family tree. To growers, the beauty of the apple lies not in its rosy skin but in its genetic variability and plasticity, its ability to cross with other species of Malus and other distant lines of M. domestica, and the ease with which it can be grafted onto different rootstocks and cloned.

See Also: Science Daily – Exploring the Origins of the Apple

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Interested in learning more about how plants get around. Check out the first issue of our new zine Dispersal Stories.

Seed Dispersal via Caching – The Story of Antelope Bitterbrush

Generally speaking, individual plants produce an enormous amount of seeds. This may seem like a huge waste of resources, but the reality is that while each seed has the potential to grow into an adult plant that will one day produce seeds of its own, relatively few may achieve this. Some seeds will be eaten before they get a chance to germinate. Others germinate and soon die from lack of water, disease, or herbivory. Those that make it past the seedling stage continue to face similar pressures. Reaching adulthood, then, is a remarkable achievement.

Antelope bitterbrush is a shrub that produces hundreds of seeds per individual. Each seed is about the size of an apple seed. Some seeds may be eaten right away. Others fall to the ground and are ignored. But a large number are collected by rodents and either stored in burrows (larder hoarding) or in shallow depressions in the soil (scatter hoarding). It is through caching that antelope bitterbrush seeds are best dispersed. When rodents fail to return to caches during the winter, the seeds are free to sprout in the spring. Some of the seedlings will dry out and others will be eaten, but a few will survive, making the effort to produce all those seeds worth it in the end.

Fruits forming on antelope bitterbrush (Purshia tridentata)

Antelope bitterbrush (Purshia tridentata) is in the rose family and is often simply referred to as bitterbrush. It occurs in grasslands, shrub steppes, and dry woodlands throughout large sections of western North America. It is a deciduous shrub that generally reaches between three and nine feet tall but can grow up to twelve feet. It has wedge-shaped leaves that are green on top, grayish on bottom, and three-lobed. Flowers are yellow, strongly fragrant, and similar in appearance to others in the rose family. Flowering occurs mid-spring to early summer. Fruits are achenes – single seeds surrounded by papery or leathery coverings. The covering must rot away or be removed by animals before the seed can germinate.

Bitterbrush is an important species for wildlife. It is browsed by mule deer, pronghorn antelope, bighorn sheep, and other ungulates, including livestock. It provides cover for birds, rodents, reptiles, and ungulates. Its seeds are collected by harvester ants and rodents, its foliage is consumed by tent caterpillars and other insects, and its flowers are visited by a suite of pollinators. For all that it offers to the animal kingdom, it also relies on it for pollination and seed dispersal. The flowers of bitterbrush are self-incompatible, and if it wasn’t for ants and rodents, the heavy seeds – left to rely on wind and gravity – would have trouble getting any further than just a few feet from the parent plant.

Antelope bitterbrush (Purshia tridentata) in full bloom – photo credit: wikimedia commons

In a study published in The American Naturalist (February 1993), Stephen Vander Wall reported that yellow pine chipmunks were the primary dispersal agents of bitterbrush seeds in his Sierra Nevada study area. The optimal depth for seedling establishment was between 10-30 millimeters. Seeds that are cached too near the surface risk being pushed out of the ground during freeze and thaw cycles where they can desiccate upon germination. Cached bitterbrush seeds benefit when there are several seeds per cache because, as Vander Wall notes, “clumps of seedlings are better able to push through the soil and can establish from greater depths than single seedlings.”

Another study by Vander Wall, published in Ecology (October 1994), reiterated the importance of seed caching by yellow pine chipmunks in the establishment of bitterbrush seedlings. Seed caches, which consisted of anywhere from two to over a hundred seeds, were located as far as 25 meters from the parent plant. Cached seeds are occasionally moved to another location, but Vander Wall found that even these secondary caches produce seedlings. Of course, not all of the seedlings that sprout grow to maturity. Vander Wall states, “attrition over the years gradually reduces the number of seedlings within clumps.” Yet, more than half of the mature shrubs he observed in his study consisted of two or more individuals, leading him to conclude that “they arose from rodent caches.”

A study published in the Journal of Range Management (January 1996) looked at the herbivory of bitterbrush seedlings by rodents. In the introduction the authors discuss how “rodents [may] not only benefit from antelope bitterbrush seed caches as a future seed source, but also benefit from the sprouting of their caches as they return to graze the cotyledons of germinating seeds.”  In this study, Ord’s kangaroo rats, deer mice, and Great Basin pocket mice were all observed consuming bitterbrush seedlings, preferring them even when millet was offered as an alternative. The two species of mice also dug up seedlings, possibly searching for ungerminated seeds. Despite seed dispersal via caching, an overabundance of rodents can result in few bitterbrush seedlings reaching maturity.

A cluster of antelope bitterbrush seedlings that has been browsed. “Succulent, young seedlings are thought to be important in the diets of rodents during early spring because of the nutrients and water they contain.” — Vander Wall (1994)

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Photos of antelope bitterbrush seedling clusters were taken at Idaho Botanical Garden, where numerous clusters are presently on display along the pathways of the native plant gardens and the adjoining natural areas.