Is Love Vine Parasitizing Wasps?

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No, that's not dodder (Cuscuta sp.), its love vine (Cassytha filiformis), a member of the same family as the avacados in your kitchen (Lauraceae). It is a pantropical parasite that makes its living stealing water and nutrients from other plants. To do so, it punctures their vascular tissues with specialized structures called "haustoria." Amazingly, a recent observation made in Florida suggests that this botanical parasite may also be taking advantage of other parasites, specifically gall wasps.

Gall wasps are also plant parasites. They lay their eggs in developing plant tissues and the larvae release compounds that coax the plant to form nutrient-rich galls packed full of starchy goodness. Essentially you can think of galls as edible nursery chambers for the wasp larvae. While looking for galls on sand live oak (Quercus geminata) growing in southern Florida, Dr. Scott Egan and his colleagues noticed something odd. A small vine seemed to be attaching itself to the galls.

Love vine draping a host plant. 

Love vine draping a host plant. 

The vine in question was none other than love vine and they were attached to galls growing on the underside of the oak leaves. What is strange is that, of all of the places that love vine likes to attach itself to its host (new stems, buds, petioles, and on the top and edge of leaves), the only time this vine showed any "interest" in the underside of oak leaves was when galls were present. Obviously this required further investigation.

The team discovered that at least two different species of gall wasps were being parasitized by love vine - one that produces small, spherical galls on the underside of oak leaves and one that forms large, multi-chambered galls on oak stems. Upon measuring the infected and uninfected galls, the team discovered that there were significant differences that could have real ecological significance.

(A)  Cassytha filiformis  vine attaching haustoria to a leaf gall induced by the wasp  Belonocnema treatae  on the underside of their host plant,  Quercus geminata . (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria. (C) Box plots of leaf gall diameter for unparasitized galls (control) and galls that have been parasitized by  C. filiformis . (D) Proportion of  B. treatae  leaf galls that contained a dead ‘mummified’ adult for unparasitized galls (control) and galls that have been parasitized by the vine  C. filiformis .  [SOURCE]

(A) Cassytha filiformis vine attaching haustoria to a leaf gall induced by the wasp Belonocnema treatae on the underside of their host plant, Quercus geminata. (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria. (C) Box plots of leaf gall diameter for unparasitized galls (control) and galls that have been parasitized by C. filiformis. (D) Proportion of B. treatae leaf galls that contained a dead ‘mummified’ adult for unparasitized galls (control) and galls that have been parasitized by the vine C. filiformis. [SOURCE]

For the spherical gall wasp, infected galls tended to be much larger, however, the team feels that this may actually be due to the fact that the vine "prefers" larger galls. Astonishingly, larvae living in the infected galls were 45% less likely to survive. For the multi-chambered gall wasp, infection by love vine was associated with a 13.5% decrease in overall gall size. They suggest this is evidence that love vine is having net negative impacts on these parasitic wasps.

Subsequent investigation revealed that these wasps were not alone. In total, the team found love vine attacking the galls of at least two other wasps and one species of gall-making fly (though no data were reported for these cases). To be sure that love vine was in fact parasitizing these galls, they needed to have a closer look at what the vine was actually doing.

Figure S2. (A)  Cassytha filiformis  vine attaching haustoria to a leaf gall induced by the wasp  Callirhytis quercusbatatoides  on the stem of their host plant,  Quercus geminata . (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria on  C. quercusbatatoides . (C) Exemplar of parasitic vine wrapping tightly around the stem directly proximate to a gall induced by the wasp  Disholcaspis quercusvirens  on  Q. geminata . (D) Field site where love vine,  C. filiformis , is attacking the sand live oak,  Q. geminata , and many of the gall forming wasps on the same host plant.  [SOURCE]   

Figure S2. (A) Cassytha filiformis vine attaching haustoria to a leaf gall induced by the wasp Callirhytis quercusbatatoides on the stem of their host plant, Quercus geminata. (B) Labeled graphic of insect gall, parasitic vine, and vine haustoria on C. quercusbatatoides. (C) Exemplar of parasitic vine wrapping tightly around the stem directly proximate to a gall induced by the wasp Disholcaspis quercusvirens on Q. geminata. (D) Field site where love vine, C. filiformis, is attacking the sand live oak, Q. geminata, and many of the gall forming wasps on the same host plant. [SOURCE]
 

Dissection of the galls revealed that the haustoria were plugged into the gall itself, not the wasp larvae. However, because the larvae simply cannot develop without the nutrients and protection provided by the gall, Eagan and his colleagues conclude that these do indeed represent a case of a parasite being parasitized by another parasite.

At this point, the next question that must be asked is how common is this in love vine or, for that matter, across all other parasitic plants that utilize haustoria. Considering that parasites of parasites are nothing new in the biosphere, it is a safe bet that this will not be the last time this phenomenon is discovered.

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

Further Reading: [1]

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]