Southern Beeches and Biogeography

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If you spend any time learning about paleontology, you are bound to come across at least one reference to the southern beeches (genus Nothofagus). This remarkable and ecologically important group of trees can be found growing throughout the Southern Hemisphere at high latitudes in South America, Australia, New Zealand, New Guinea, and New Caledonia. Not only are they prominent players in the forests in which they grow, their fossil history has provided scientists with invaluable data on plate tectonics and biogeography.

Southern beeches may not be the tallest trees in any forest but that doesn’t mean they aren’t impressive. Numbering around 37 species, southern beeches have conquered a range of climate zones from temperate to tropical. Those living in lowland tropical forests tend to be evergreen, holding onto their leaves throughout the year whereas those living in temperate or montane habitats have evolved a deciduous habit. Some species of southern beech are also known for their longevity, with individuals estimated to be in excess of 500 years in age.

Nothofagus alpina

Nothofagus alpina

Anyone familiar with the true beeches (genus Fagus) will quickly recognize many similarities among these genera. From their toothy leaves to their triangular nuts, these trees are strikingly similar in appearance. Indeed, for much of their botanical history, southern beeches were included in the beech family (Fagaceae). However, recent molecular work has revealed that the southern beeches are genetically distinct enough to warrant their own family - Nothofagaceae.

The beech-like fruits of Nothofagus obliqua var. macrocarpa

The beech-like fruits of Nothofagus obliqua var. macrocarpa

As mentioned, the southern beeches, both extant and extinct, have been important players in our understanding of plate tectonics. Their modern day distribution throughout the Southern Hemisphere seems to hint at a more concentrated distribution at some point in the past. All of the continents and islands on which they are found today were once part of the supercontinent of Gondwana, which has led many to suggest that the southern beech family arose before Gondwana broke apart during the Jurassic, with ancestors of extant species riding the southern land masses to their modern day positions. Indeed, the paleo record seems to support this quite well.

Fall colors of Nothofagus cunninghamii.

Fall colors of Nothofagus cunninghamii.

The southern beeches have an impressive fossil record that dates back some 80 million years to the late Cretaceous. Their fossils have been found throughout many of the Southern Hemisphere continents including the now-frozen Antarctica. It would seem that the modern distribution of these trees is the result of plate tectonics rather than the movement of seeds across oceans. This is bolstered by lines of evidence such as seed dispersal. Southern beech nuts are fairly large and do not show any adaptations for long distance dispersal, leading many to suggest that they simply cannot ocean hop without serious help from other forms of life.

Nothofagus fusca

Nothofagus fusca

However, life is rarely so simple. Recent molecular work suggests that continental drift can’t explain the distribution of every southern beech species. By studying trees growing in New Zealand and comparing them to those growing in Australia and Tasmania, scientists have discovered that these lineages are far too young to have originated before the breakup of Gondwana. As such, the southern beeches of Austrialasia more likely got to their current distributions via long distance dispersal events. Exactly what allowed their seeds to cross the Tasman Sea is up for debate, but certainly not impossible given the expanse of time available for rare events to occur. Regardless of where anyone stands on this recent evidence, it nonetheless suggests that the biogeographic history of the southern beech family isn’t as clear cut as once thought.

Nothofagus fusca

Nothofagus fusca

Unfortunately, while southern beeches hold a prominent place in the minds of naturalists, the same cannot be said for the rest of the world. Little care has been given to their scientific and ecological importance and massive quantities of these trees are logged each and every year. Today it is estimated that 30% of all southern beech species are threatened with extinction. Luckily, large portions of the remaining populations for these trees are growing on protected lands. Also, because of their scientific importance, numerous southern beeches can be found growing in botanical collections and their seeds are well represented in seed banks. Still, southern beeches and the forests they comprise are worthy of respect and protection.

Photo Credits: [1] [2] [3] [4] [5] [6]

Further Reading: [1] [2] [3]

Tropical Oaks - Lessons in Biogeography from a Giant Acorn

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Seeing the nut of Quercus insignis in person for the first time was a peculiar experience. I didn't know acorns came that big! What was even stranger was encountering this species in the tropics. I thought that in leaving my temperate home behind, I had left trees such as oaks behind as well. Thus, picking up this gigantic acorn was a challenge to my ignorance of tropical forest diversity. What it did for me was ignite a fury of questions regarding the biogeography of the genus Quercus.

Quercus insignis is native from Mexico to Panama. It is a member of the white oak grouping and, despite having one of the largest acorns of any oak species, relatively little is known about this species. What we do know is that it is in trouble. It is considered critically endangered in Mexico and near threatened in Guatemala and Panama with a remaining stronghold in Nicaragua. Habitat loss and changing environmental conditions seem to be at the core of its disappearance.

One big question was looming over me. What was an oak doing this far south? Call it a northern bias but I have always associated oaks with more temperate climes. I needed to get over this. My investigation lead me to some very interesting work done on the family to which oaks belong - Fagaceae. Based on some incredible paleontological and genetic detective work, we now know that Fagaceae originated in Asia. The first fossil evidence of a member of this family dates back some 100 million years, during the early part of the Cretaceous.

At this time, the continents of Asia, Europe, and North America were still connected. Some 60 million years ago, the genus Quercus diverged from Castanea. They were also starting to radiate across the Northern Hemisphere. The first fossil evidence of oaks in North America comes from Paleogene deposits dated to 55 to 50 million years before present. This is when the oaks really started to hit their stride.

Between 22 and 3 million years ago the genus Quercus underwent numerous speciation events. The new terrain of North America must have presented countless opportunities for oaks because they quickly became the most specious genus of all the Fagaceae. This radiation was particularly fruitful in what would become the U.S. and Mexico. Of the roughly 220 species that exist in this region today, 160 occur in Mexico, and of those, 90 species are endemic.

This brings us to the tropics. Evergreen and semi-evergreen oaks have done quite well in this region. However, their astounding diversity quickly drops once you hit the isthmus of Panama. South America is home to only one species of oak. What happened that limited the oaks reign south of the equator?

To put it simply, geology happened. For much of the Earth's history, North and South America shared no connection. Though the exact time frame is debated, tectonic forces joined the two continents some 4.5 million years ago. The Great American Interchange had begun. The two continents were able to freely exchange flora and fauna like never before. The migrations are thought to have been a bit lopsided. Tropical flora and fauna did not do as well farther north but temperate flora and fauna seemed to find warmer climes more favorable. As such, South America gained disproportionately more biodiversity as a result.

This pattern did not hold true for everything though. For the oaks, only one species (Quercus humboldtii) made it through. As such, the genus remains a dominant fixture of the Northern Hemisphere. Sadly, much of this diversity is at serious risk of being lost forever. Like the magnificent Quercus insignis, many of the world's oaks are on the decline. Disease, habitat loss, and countless other issues plague this genus. A 100 million year old journey is quickly being undone in less than two centuries. The hand of man is time and again proving to be a force unrivaled in the biological world.

Leaf Credit: http://www.oaknames.org/

Further Reading:
http://www.sciencedirect.com/science/article/pii/S0378112713006580