A Thousand Acres of Silphiums

Review

After observing Vespula vulgaris foraging for the diaspores of Trillium ovatum in Oregon, Jules (1996) coined the term vespicochory to describe the dispersal of seed by Vespid wasps, specifically members of the Vespinae – the Vespidae subfamily that is comprised of yellowjackets and hornets. Few other descriptions have been recorded on observations of Vespinae acting as secondary seed dispersers in myrmecochorous plants. In controlled experiments, V. maculifrons has been reported to disperse T. cuneatum, T. undulatum and T. catesbaei in North Carolina and South Carolina (Zettler et al., 2001) and T. discolor in South Carolina (Bale et al., 2003). To my knowledge, this is the first recorded observation of the dispersal of T. recurvatum by Vespid wasps.

Before I begin describing my observations, I will first review and discuss the potential implications of Vespinae dispersal as an alternative to myrmecochory in trilliums. In most descriptions, myrmecochory has been described as a mutualist symbiose, meaning that both plant and ant species benefit from the relationship. Ants benefit by gaining the lipid and protein rich eliasome of the diaspore to feed their young while the plants benefit by having their seed dispersed from the parent plant and gain potential benefits in overwintering and germination environments. Similarly, for vespicochory to be considered as an important seed dispersal syndrome, we should consider the benefits to both sides and compare the role these wasps play to that of their ant cousins.

Yellowjackets, in addition to seizing seed directly from ants, removed more seeds (40%) from index cards than did each of three ant species observed (8 – 28%) (Bale et al., 2003). Zettler et al. (2001) measured dispersal distances by V. maculifrons and found an average distance of 1.4 m compared to a mean of 0.98 m in global cases of myrmecochory (Gómez and Espadaler, 1998). This difference in dispersal distance alone is significant; however, 53% of the seeds removed by V. maculifrons in this study were moved beyond 20 m – the furthest extent of their measuring capabilities, and were unrecovered, indicating a much higher than calculated average dispersal distance.

In addition to dispersal distance, another important thing when considering the benefits to the plant in a particular dispersal syndrome, is what is done with the diaspore once removed. The mandibles of Vespids are considerably larger and assumedly much more powerful than those of the ant species involved in dispersing trillium seed throughout their range. It is therefore a possibility that the seed could face catastrophic damage from the foraging wasp. Of the original seed recovered by Zettler et al. (2001), 95.7% had the eliasomes removed. Of these, 17% of the seeds had scarification near where the eliasome was attached but no seeds showed visible signs of embryo damage. The ultimate use of the eliasome was unknown in Zettler et al. (2001); however, Jules (1996) observed yellowjackets taking diaspores directly into their nest where, presumably, they were fed to developing young. Vespids typically nest underground and waste (i.e. seed portions of diaspores) are deposited below the nest. As mentioned by Zettler, “…we do not know how seed burial in these nests might affect seed germination and seedling emergence.” In cases where eliasomes are removed and seed are dropped randomly on the ground, it would be interesting to know how these seed fare in comparison to those buried within the nest. Further study is needed to determine the fate of seeds moved by these wasps when compared to myrmecochory.

Observations

The following observations and accompanying photographs were conducted at August G. Beckemeier Conservation Area in St. Louis County, Missouri. On August 5, 2021 at ~ 18:00 hrs., I collected a ripe fruit of T. recurvatum and placed it with about 25% of the seed exposed within 20 cm of a nest of Formica pallidefulva ants. My goal was to observe and photograph the ants carrying away the diaspores. The ants found the fruit within minutes and quickly began moving the loosely separated diaspores. After approximately 10 minutes the first V. maculifrons found the fruit and quickly left with a diaspore. It returned alone five times with gaps ranging between approximately one and three minutes. After the fifth visit, two to four wasps were at the scene at any given time, each working to free seeds from the fruit until all seeds were removed. I found that the wasps were able to pull the diaspores free from the fruit capsule matrix much easier than the ants. The ants tried, at times, to defend the fruit and the wasps did give them a wide berth. When two or more wasps were on the fruit at one time, however, the ants were ineffectual in their defense.

I was not setup to make accurate counts or to try and make seed dispersal distance measurements. It appeared the wasps moved at least 75% of the seed while F. pallidefulva moved the remaining into their nest. I believe this discrepancy was primarily due to the ability of the wasps to excise the diaspores from the fruit capsule matrix more quickly and easily than the ants. I watched one wasp perched on a short sapling approximately 1.5 m from the fruit. It removed the eliasome, letting the seed fall to the leaf litter below and then left carrying the eliasome with it.

Photography Details

These images were taken using a full-sized sensor digital camera and a 180mm macro lens with a 1.4x teleconverter and 30 mm extension tube stacked between the lens and camera body. This combination of equipment provides quite a long focusing distance, ensuring the photographer does not disturb the subjects. An off-the-body external speedlight “flash” was used at varying levels of power to obtain the extra light needed. Most of these images were taken at f/16, 1/100 sec. and ISO-640 and were taken handheld while using a fallen log for additional support.

Conclusion

This was an anecdotal observation of a single occurrence of vespicochory. This is a subject that warrants further investigation. Could vespicochory be just as or even more important in the dispersal and emergence of some “myrmecochorous” plants as myrmecochory? It would be interesting to know more about the frequency and dynamics of this unique seed dispersal mechanism.

References