Ginkgoes and dodos are my two favorite icons of “ghost” mutualistic interactions. So, I had the two last posts on megafauna and extinct interactions dedicated to them.

There are only five living groups of seed plants, and ginkgo is one of them; just a single species. Ginkgoes (Ginkgo biloba, Ginkgoaceae) have fleshy “fruits” and their “seeds” were dispersed by animals including, most probably, from dinosaurs to Pleistocene megafauna, and to extant frugivores nowadays. The reason is that the ginkgo has survived on Earth for a really extended period of time, with the earliest fossils of ginkgo-like plants dated more than 200 million years ago. Among the many ginkgo-like tree species, only Ginkgo biloba has survived (up to five Ginkgo species are known as fossils).


Living ginkgo very nearly went extinct, in fact, as of two million years ago, existing in only a small area in eastern China, the Tian Mu Shan mountains in Guizhou Province. Ginkgoes have then survived just by human intervention, with an assisted dissemination for cultivation starting at least 1200 yr ago by Buddhist monks, and introduced to Europe just by 1730-1750.

From Engelbert Kaempfer in his Amoenitates, 1712: the first illustration of ginkgo by a Western botanist.

Most likely a combination of extreme dispersal limitation due to lack of efficient seed dispersers combined with large-scale climate shifts and habitat modification contributed to their nearly extinction in the wild. Contrary to other tree species, retractions to small refugia populations failed to recover the original range, especially in North America and Europe. As with other megafauna-dependent species, it resprouts vigorously from buds buried in its underground parts, and human use certainly rescued the ginkgoes, probably because of their nutritous “nut”. In Peter Crane’s words: “It is irrepressible; its capacity for self-preservation has helped it survive through millions of generations.”

We know very little about how seed dispersal works in living ginkgo. The fleshy “fruit” is really the mature, fertilized ovule with a a three-layered integument: a fleshy outer sarcotesta, a stony inner sclerotesta, and a thin endotesta. Its smelly, large seeds (20-30 mm x 16-24 mm) are one of its most well-known and distinctive features: the seed’s soft outer layer starts to break down after a few days on the ground and produces butyric acid, CH3(CH2)2COOH giving it the “interesting” odor. Germination improves after the fleshy seed coat has been removed by passing through the gut of an animal or being teared-off. In one of the potentially wild ginkgo populations in China it is documented that the seeds are eaten by a wild cat, and in Japan they are eaten by badgers. Yet, there were very few seedlings in this population, located in 1989 by Del Tredici, despite good fruiting. People harvested the nuts, which are very nutritious, as well as Pallas’s squirrels (Callosciurus erythraeus), which also may act as good dispersers by scatter-hoarding the seeds.

Yet who were the seed dispersers that mediated the range expansion of ginkgoes over continents and islands (Japan) before human-mediated propagation? As in other megafauna-dependent plants, most likely a combination of dispersal agents, including large and medium-sized mammals and, well before that, dinosaurs. As with other extant large-‘seeded’ Coniferopsida like Cephalotaxus and Torreya with very large seeds, scatter-hoarding animals like the extinct multituberculates (i.e., the ‘rodents’ of the Mesozoic; g. Ptilodus) would have played a role in active seed dispersal of ginkgoes by scatter-hoarding the seeds.

We can see ginkgoes as survivors with a long history of mutualistic interactions involving a diverse array of animals, whose actual diversity we can only speculate about, then replaced by extensive human use.

  • van Beek, T.A. (2003) Ginkgo biloba. CRC Press, NY.
  • Crane, P. (2013). Ginkgo. The tree that time forgot. Yale University Press, New Haven.
  • del Tredici, P. (1989) Ginkgos and multituberculates: evolutionary interactions in the Tertiary. Bio Systems, 22, 327–339.

Text: Pedro Jordano with excerpts from Del Tredici (1989) and Crane (2013). Illustrations: WikiMedia.

Updated dataset on fruit colors in Dryad

  I’ve updated our dataset on fruit colors that we used in the Journal of Evolutionary Biology paper. The full metadata is here, in the Dryad open-source repository. You can find this and other datasets also in the web page.

Megafauna fruits and seeds photos updated

I’m in the process of moving photo repositories to iCloud and have my previous photos of megafauna fruits and seeds moved to this gallery. You can access the photos here.

These photos were taken in the herbarium of Museu Goeldi (Belém, Pará, Brazil) in 2002. They include many of the species we discuss in our paper in PLoS One on megafauna fruits.

Fruit colors paper just published in Journal of Evolutionary Biology

Our study on the evolution of visual displays in plants appeared in the last issue of JEB. We are still working on this subject, now exploring how effective are visual displays (fruit colors) in attracting avian frugivores that might use color cues to build up complex and diverse fruit diets. We have evidence that strongly frugivorous birds might use color cues to guide foraging for fruit combinations that maximize the yield of important nutrients. Rather than consuming fruits at random, frugivorous warblers are quite selective not only on the fruit species they take, but in which combinations they consume the different fruit species.

Fruit colors

Fruits show an immense diversity of colors and displays. However, we are still far from a general theory for the evolution of fruit displays. The main elements of those displays do not only include color itself, but also characteristics of the fruit “design” (ow the fruit is built) like number of seeds, amount of pulp, size, etc., and the nutrients in the pulp (both macro- and micro-nutrients, as well as secondary compounds). All this adds an extraordinary complexity and diversity to the fruit displays. Together with Alfredo Valido and Martin Schaefer I’ve been exploring the evolutionary patterns of fruit traits for the Iberian Peninsula fleshy-fruited flora (ca. 120 species). We studied whether correlated trends between these elements of the display (design, nutrients, color) have been maintained through the phylogenetic diversification of the flora. We found some interesting patterns of covariation between sugar content, lipid content, and color that suggest predictable patterns of fruit evolution in relation to the main types of frugivores feeding on the fruits. Our results suggest that the evolution of fruit displays has been quite constrained by history, yet selection by frugivores might have contributed to marked and predictable covariation among color and nutrient contents. This is an interesting finding to understand the evolution of visual signals in plants, acting to attract diverse suites of animal frugivores that can act as legitimate dispersers of the seeds. Our work is now in press in Journal of Evolutionary Biology.

Fruit color brief course

Course on measurement of colors of flowers and fruits

Together with Alfredo Valido, I gave a short and introductory course on fruit colors and their measurement. We gave that course just prior to the ECOFLOR meeting in Valencia, one of the nicest meetings we have here, where a bunch of great people working on pollination biology get together.


Photos of megafauna fruits

Go here if you want a view of megafauna fruits photos . These were taken by Mauro Galetti and me at the Museo Goeldi Herbarium (Belem, Brazil) and correspond to species we analyzed in our ongoing project on megafauna fruits.