Fossils Shine Light On the History of Gall-Making Wasps

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We can learn a lot about life on Earth from the fossil record. I am always amazed by the degree of scrutiny involved in collecting data from these preserved remains. Take, for instance, the case of gall-making wasp fossils found in western North America. A small collection of fossilized oak leaves is giving researchers insights into the evolutionary history of oaks and the gall-making wasps they host.

Oaks interact with a bewildering array of insects. Many of these are gall-making wasps in the family Cynipidae. Dozens of different wasp species can be found on a single oak tree. Female wasps lay their eggs inside developing oak tissues and the larvae release hormones and other chemicals that cause galls to form. Galls are essentially edible nursery chambers. Other than their bizarre shapes and colors, the compounds released by the wasp larvae reduce the chemical defenses of the oak and increase the relative nutrition of the tissues themselves. Often, these relationships are precise, with specific wasp species preferring specific oak species. But when did these relationships arise? Why are oaks so popular? What can fossil evidence tell us about this incredible relationship?

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Though scant, the little fossil evidence of oak galls can tell us a lot. For starters, we know that gall-making wasps whose larvae produce structures similar to that of the Cynipids have been around since at least the late Cretaceous, some 100 million years ago. However, it is hard to say for sure exactly who made these galls and exactly what taxonomic affinity the host plant belongs to. More conclusive Cynipid gall fossils appear again in the Eocene and continue to pop up in the fossil record throughout the Oligocene and well into the Miocene (33 - 23 million years ago).

Miocene aged fossils are where things get a little bit more conclusive. Fossil beds located in the western United States have turned up fossilized oak leaves complete with Cynipid galls. The similarity of these galls to those of some present day species is incredible. It demonstrates that these relationships arose early on and have continued to diversify ever since. What's more, thanks to the degree of preservation in these fossil beds, researchers are able to make some greater conclusions about why gall-making wasps and oaks seem to be so intertwined.

Holotype of Antronoides cyanomontanus galls on fossilized leaves of  Quercus simulata . 1) Impression of the abaxial surface of the leaf, showing the galls extending into the matrix. 2) Galls showing close association with secondary veins. 3) Gall showing the impression of rim-like base partially straddling the secondary vein. 4) Close-up of gall attached at margin extending down into the matrix. 5) Gall uncovered revealing spindle-shaped morphology.

Holotype of Antronoides cyanomontanus galls on fossilized leaves of Quercus simulata. 1) Impression of the abaxial surface of the leaf, showing the galls extending into the matrix. 2) Galls showing close association with secondary veins. 3) Gall showing the impression of rim-like base partially straddling the secondary vein. 4) Close-up of gall attached at margin extending down into the matrix. 5) Gall uncovered revealing spindle-shaped morphology.

1)  Xanthoteras clavuloides  galls on fossilized  Quercus lobata , showing gall attached to secondary vein. Specimen in California Academy of Sciences Entomology collection, San Francisco. 2) Two galls of attached to a secondary vein showing overlap of their bases. Specimen in California Academy of Sciences Entomology Collection, San Francisco. 3) Three galls collected from leaf of California  Quercus lobata  showing clavate shape and expanded, ring-like base. 4) Gall showing the annulate or ribbed aspect of the base, which is similar to bases of  Antronoides cyanomontanus  and  A. polygonalis . 5) Galls showing clavate shape, pilose and nonpilose surfaces, and bases.

1) Xanthoteras clavuloides galls on fossilized Quercus lobata, showing gall attached to secondary vein. Specimen in California Academy of Sciences Entomology collection, San Francisco. 2) Two galls of attached to a secondary vein showing overlap of their bases. Specimen in California Academy of Sciences Entomology Collection, San Francisco. 3) Three galls collected from leaf of California Quercus lobata showing clavate shape and expanded, ring-like base. 4) Gall showing the annulate or ribbed aspect of the base, which is similar to bases of Antronoides cyanomontanus and A. polygonalis. 5) Galls showing clavate shape, pilose and nonpilose surfaces, and bases.

Gall-making wasps seem to diversify at a much faster rate in xeric climates. The fossil records during this time show that mesic tree speciess were gradually being replaced by more xeric species like oaks. Wasps seem to prefer these drier environments and the thought is that such preferences have to do with disease and parasite loads.

Again, galls a large collections of nutrient-rich tissues that are low in defense compounds. Coupled with the juicy grub at their center, it stands to reason that galls make excellent sites of infection for fungi and other parasites. By living in drier habitats, it is believed that gall-making wasps are able to escape these environmental pressures that would otherwise plague them in wetter habitats. The fossil evidence appears to support this hypothesis and today we see similar patterns. White oaks are especially drought tolerant and its this group of oaks that host the highest diversity of gall-making wasps.

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

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

Red or White?

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Who doesn't love a nice oak tree? One cannot overstate their importance both ecologically and culturally. Although picking an oak tree out of a lineup is something many of us are capable of doing, identifying oaks to species can be a bit more challenging. This is further complicated by the fact that oaks often hybridize. Still, it is likely you have come across some useful tips and tricks for narrowing down your oak choices. One such trick is distinguishing between the red oaks and the white oaks. If you're anything like me, this is something you took for granted for a while. Is there anything biologically or ecologically meaningful to such a split?

In short, yes. However, a true appreciation of these groups requires a deeper look. To start with, oaks are members of the genus Quercus, which belongs in the family Fagaceae. Globally there are approximately 400 species of oak and each falls into one of three categories - the red oaks (section Lobatae), the white oaks (section Quercus), and the so-called "intermediate" oaks (section Protoblanus). For the sake of this article, I will only be focusing on the red and white groups as that is where most of the oak species reside. The intermediate oak group is made up of 5 species, all of which are native to the southwestern United States and northwestern Mexico.

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As is common with oak identification, reliable techniques for distinguishing between the two groups can be tricky. Probably the most reliable feature is located on the inner surface of the acorn cap. In white oaks, it is hairless or nearly so, whereas in red oaks, it is covered in tiny hairs. Another useful acorn feature is the length of time it takes them to germinate. White oak acorns mature in one season and germinate in the fall. As such, they contain lower levels of tannins. Red oak acorns (as well as those of the intermediate group) generally take at least two seasons to mature and therefore germinate the following spring. Because of this, red oak acorns have a much higher tannin content. For more information on why this is the case, read this article.

Tyloses in white oak xylem.

Less apparent than acorns is the difference in the wood of red and white oaks. The wood of white oaks contains tiny structures in their xylem tissues called tyloses. These are absent from the wood of red oaks. The function of tyloses are quite interesting. During extreme drought or in the case of some sort of infection, they cut off regions of the xylem to stop the spread of an embolism or whatever may be infecting the tree. As such, white oaks tend to be more rot and drought resistant. Fun fact, tyloses are the main reason why white oak is used for making wine and bourbon barrels as it keeps them from leaking their contents.

More apparent to the casual observer, however, is leaf shape. In general, the white oaks produce leaves that have rounded lobes, whereas the red oaks generally exhibit pointed lobes with a tiny bristle on their tips. At this point you may be asking where an unlobed species like shingle oak (Quercus imbricaria) fits in. Look at the tip of its leaf and you will see a small bristle, which means its a member of the red oak group. Similarly, the buds of these two groups often differ in their overall shape. White oak buds tend to be smaller and often have blunted tips whereas the buds of red oaks are generally larger and often pointed.

Tricky leaves of the shingle oak ( Quercus imbricaria ). Note the bristle tip!

Tricky leaves of the shingle oak (Quercus imbricaria). Note the bristle tip!

Despite this broad generalizations, exceptions abound. This is further complicated by the fact that many species will readily hybridize. Quercus is, after all, a massive genus. Regardless, oaks are wonderful species chock full of ecological and cultural value. Still, oak appreciation is something we all need more of in our lives. I encourage you to try some oak identification of your own. Get outside and see if you can use any of these tricks to help you identify some of the oaks in your neighborhood.

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

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