Unknown stem galls on Common Mugwort
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This is a story about an unknown gall former, and the story is still only partly understood.
Last summer (August 2020) in Central Park and on Randall's Island, I found six stem galls on the very common and extremely invasive weed Common Mugwort, Artemisia vulgaris. The galls look like this one:
https://www.inaturalist.org/observations/79560187
@steven-cyclist was with me several times when I found them, and he probably also photographed one or two of them.
I had never noticed these galls on mugwort before 2020, so I photographed them all, but I had no luck working out what the gall-making organism was that had caused the galls to form.
Fortunately iNatter @jeffmollenhauer is a lot more of a thorough and persistent naturalist than I am. Jeff collected some of these galls in New Jersey. Jeff then carefully cut one gall open, and photographed the larva that was inside the gall. It appeared to be a beetle larva:
https://www.inaturalist.org/observations/65600322
Jeff also carefully saved a few of the intact galls to see what hatched out in the next spring or summer.
Sure enough, this spring, 2021, beetles started hatching out, like this one:
https://www.inaturalist.org/observations/79554086
It is a Tumbling Flower Beetle, in the genus Mordellistena.
However, it seems that the beetle is almost certainly not the original gall former, but instead is an inquiline parasite. Mordellistena is a stem borer and a gall borer, but it does not create galls.
The Mordellistena larva apparently takes over the gall, a gall which is probably initially occupied by, and created by, a Eurosta fly larva. Eurosta are gall flies, and they are the causal agent of some similar galls on Golden Rod species. For example this large gall, which was caused by the Goldenrod Gall Fly, Eurosta solidaginis:
https://www.inaturalist.org/observations/10167994
After an initial gall is formed on Common Mugwort, then presumably a mother Mordellistena beetle lays an egg on the outer surface of the gall. That egg hatches and the young beetle larva chews its way into the gall. The beetle larva starts out by eating the soft inner tissues of the gall, and then probably it often goes on to eat the little Eurosta larva too! Talk about eating you out of house and home!
For more info on this type of interaction, read the results section in this paper (especially under the Differentiation of Mordellistina convicta section). https://academic.oup.com/icb/article/41/4/928/2046532
Perhaps the Mordellistena species in our case might also be Mordellistena convicta? On BugGuide that species has been reported from inside galls on Common Ragweed, Ambrosia artemisifolia, so maybe it could also bore into galls on Common Mugwort, a plant which is somewhat closely related?
https://bugguide.net/node/view/170446
I asked @borisb, a beetle expert, what he thought about this beetle, and Boris commented, "What you have resembles those WE [in Europe] have in ragwort (Mordellistena weisei-group), though ours do not produce galls." I am assuming that Boris means that this group of Mordellistena beetles in Europe bore into the stems of Ragwort plants (does Boris mean the plant Jacobaea vulgaris?), but that they do not cause galls to form. I explained to Boris that we assume that the Mordellistena beetles here do not actually cause the galls in the Common Mugwort, but instead they take advantage of galls already created by a gall-forming fly.
And incidentally, here is a 2008 paper about differentiating the larvae of that European group of Mordellistena beetles:
https://agris.fao.org/agris-search/search.do?recordID=BY2008000753
@megachile made an entry in the excellent gallformers.org database (a great new resource from @megachile and @jeffdc) for this still currently unknown stem gall:
https://www.gallformers.org/gall/1592
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@susanhewitt Thanks for the great summary on what we have learned to date on the Common Mugwort galls! I would still love to figure out which Mordellistina species this is and it would be super interesting to discover which insect is the original gall former before the Mordellistina larvae bores into the gall. I'm happy to have helped uncover some of the story, but would love to learn the rest of it. The galls were abundant on Common Mugwort in southern New Jersey in the fall of 2020.
I agree. This is a fascinating story and still very incompletely understood.
Maybe we can try to find some of these galls in a very early stage (perhaps now) and cut the galls open to try to find what we assume may be the fly larva that causes the gall to form in the first place.
@srall -- you might like to read about this.
I had the same thought. I'll check on the Mugworts tomorrow and see if there are any galls forming yet. :-)
Great! Please do let me know if you find any.
Enjoying following along with this story! I do feel like these galls suddenly popped up all over last year, but maybe it is just an increasing number of eyes on the lookout for such things.
That's a good point. I don't know if maybe I was being more carefully observant last year, on the lookout for galls, or whether there was indeed a sudden increase in the number of these galls last summer.
I must say that I suspect the latter.
Neat, thanks for tagging me. I've seen these many times.
@Thanks @srall -- have you. seen good numbers of them every year, or have they been increasing in recent years?
I've only been looking for them for two summers (before this one), so I can't really say if they are trending. Generally, in the fall, if I look at any large stand of mugwort I will find some.
I think, in general, with galls, mines, diseases, etc., people simply didn't photograph them, didn't notice them before, so they look explosive. For instance, I never knew I could ID oak shothole miners until @susanhewitt started posting them, but my oaks have had holes in them for a decade at least.
I imagine you are right in your opinion about these phenomena, although I do remember reading about the oak shot hole leaf miner that there was big outbreak a few years ago in the northeast.
Images of an adult Mordellista sp. reared from one of these galls:
https://www.inaturalist.org/observations/79554086
I just became aware of these galls about a month ago, and I was wondering if you all had any luck finding any of them last spring. It seems that this late in the season, all that's in the galls are the mordellid larvae, but based on the lack of updates- were the galls absent from the mugwort in the spring? I wonder if they might be a later season gall. In any case, ever since I first learned of them I've been seeing these stem swellings everywhere, they seem to be quite abundant. (I've seen them here in PA, as well as in CT and NJ when I've traveled.)
Yes, I could not find any of these galls in the spring this year, although I looked. I don't know when they first started appearing this year. I suppose it must be in the early summer? Or late summer?
That might make sense. I'll keep an eye out for them more this next season, too. Hopefully I'll be able to catch when they start appearing then.
I will also try more carefully to search month by month in an attempt to find out when it is that they start appearing.
I assume the mugwort needs to get fairly tall before these galls start forming.
Unfortunately I was not able to search for the galls until late summer this year due vacation and work schedule in the spring and early summer. Hoping to do better next year. Planning to rear some more galls this winter to see if I get more Mordellistena though.
Hoping to find some earlier ones this year.
@susanhewitt:
Susan, when I said "resembles those we [in Europe] have in ragwort", what I thought of was common mugwort.
There is a record of a species of uncertain identity (19th century!) from a tall Senecio, however.
I now tried the key of Liljeblad (1945) on your individual - without reaching a conclusion. But we can be sure it neither is M. convicta (legs dark, elytral vestiture stripy, tibial spurs yellow), nor M. aspersa (legs dark, tibial ridges of about same length, not reaching near opposite side of tibia), which also has been found in galls of Eurosta flies.
See the observation for more.
Thanks @borisb!
Very interested in this thread. We have been collecting and rearing these galls as part of the Stone Barns Center Gall Project. We have seen mostly emergences of the same Mordellistena beetle (see https://www.inaturalist.org/observations/149353922 and https://www.inaturalist.org/observations/149356112 for gall images). Interestingly, I had one Braconid wasp (I think from subfamily Doryctinae, but that is yet to be confirmed) emerge from this gall (see https://www.inaturalist.org/observations/149358703). Unfortunately the wasp emerged while I was away and my team missed it during gall checks (it is pretty tiny), so the specimen was badly desiccated by time I returned. Both this wasp and the Mordellistena beetles will be sent to the University of Guelph for DNA barcoding (likely in March 2023) and I will update if we receive any additional taxonomic information from the barcoding. We have collected many more samples for rearing this year, from a variety of locations, times, and maturity. While many Doryctinae are idiobiont ectoparasitoids of wood-boring beetle larvae and thus possibly this individual parasitized one of the Mordellistena larva, some members of the subfamily have been shown to be gall formers (https://pubmed.ncbi.nlm.nih.gov/34563693/).
@stonebarnsecology : Barcoding is an excellent idea - the common European it resembles is in the database, we are lucky:
https://www.barcoding-zsm.de/tax/taxon/318140
In your specimen photographed, we can see the pubescence on elytra is in two colours:
Mainly golden, dark along suture. Just like in Mordellistena bicoloripilosa, and nothing else seen that would differ.
Morphological evidence in this genus is inferiour to genetics . . .
@stonebarnsecology thanks for posting additional information about the gall here! If you need any samples from New Jersey, please let me know. I would be happy to collect and send some of the galls to you if it would help your study. The galls are abundant on Artemisia vulgaris in New Jersey.
@borisb @jmole Thanks for the responses. I will keep you updated as the data comes in and as we get new emergences from the collected galls this spring/summer. Our current gall emergence specimens are scheduled to be sent to the University of Guelph for Sanger sequencing along with a bunch of samples from targeted pollinator collection, I'm just slowly working my way through data cross-checks with our ArcGIS system and making sure the photography for each specimen is up to par before we send the specimens out. Hopefully I will be finished in the next couple of weeks, and then it is in Guelph's hands.
If helpful, here are some additional images of another Mordellistena specimen that emerged from a different group of galls on a nearby patch of mugwort in the same field (https://www.inaturalist.org/observations/149376416).
@borisb I think this observation https://www.inaturalist.org/observations/151166094 has a better view of the color of the pubescence you mentioned in your post above.
@stonebarnsecology : Right - this about the median in range of variation (few golden hair in shoulder region, to narrowly dark along suture, and tips)
I created an iNaturalist project for tracking the observations of this unknown mugwort gall: https://www.inaturalist.org/projects/unknown-stem-galls-on-common-mugwort
Also I haven't seen it mentioned yet, but there does seem to be a mugwort stem gall midge (Lasioptera artemisiae) in the old world: https://www.inaturalist.org/observations/37844399
Boston record of the beetle:
https://www.inaturalist.org/observations/165939064
Thanks Boris!
@susanhewitt What an interesting mystery, Susan! I too have seen similar galls on mugwort.
came across this observation, and it looks a bit different than what i've seen of these galls, possibly a different mugwort species also? and the larva doesn't look like the usual mordellid: https://www.inaturalist.org/observations/54002485
@cecildomyiidae That looks an awful lot like a fly puparium (or larva) from what I can see...
These stem galls are quite noticeable wherever Common Mugwort grows & I’ve seen them for some time without bothering to pay special attention to them. What I am still unclear about is, is there any evidence that these galls are the result of infestation by anything other than a species of Mordellidae? Every gall I have opened has contained one larva. apparently a beetle & apparently a species of Mordellidae.
https://www.inaturalist.org/observations/232963335
Has anybody asked Rob Naczi about these beetles? He works at the New York Botanical Garden but studies mordellids on the side.
Thanks for that tip, Charley. I've just written to him. Here is his BugGuide profile: https://bugguide.net/user/view/143385
It's odd that there is no mention of any mordellid larva associated with Artemisia vulgaris in the paper by Ford & Jackman (1996).
Ford, E. J. & Jackman, J. A. 1996. New larval host plant associations of tumbling flower beetles (Coleoptera: Mordellidae)
in North America. The Coleopterists Bulletin 50(4): 361-368. https://www.jstor.org/stable/4009273
These stem galls of Artemisia vulgaris and the consistent association of a mordellid species (one or more) can hardly be new! Also, if Artemisia vulgaris is introduced in North America from Europe surely there should be many records of these stem galls from Europe, but so far (with admittedly casual searching) I have not been able to find any mention of stem galls of this type in Artemisia vulgaris in Europe.
Hi all, I heard back from Rob Naczi and have his permission to share with the iNaturalist community what he wrote about the Mordellidae associated with these Common Mugwort stem galls. Here is what Rob wrote:
Thank you very much for making me aware of the interest in the galls, plant, fly, and mordellid. I am very pleased to see such a lively discussion and so many naturalists involved. This is wonderful!
First, please realize that to provide a determination of the mordellid, I would need to examine physical specimen(s). Many, many mordellids cannot be determined definitely from photographs.
Now, more to the point, I have received a few specimens of mordellids reared from Common Mugwort in New York state, and these are Mordellistena aspersa. Please realize that the taxonomy of this species is incompletely resolved; what is called M. aspersa may be a complex of species. Also, this name has not been typified. So, the true identify of Mordellistena aspersa is unclear at this time. Further, it’s possible that more than one mordellid species inhabits galls on Common Mugwort, though the specimens I have seen belong to one species.
https://bugguide.net/node/view/2288007 maybe not same.
wow, cool!
We found many stems of Artemisia vulgaris to have galls apparently similar to these in Providence, Rhode Island, on August 17. Cut them open and found a larva that could be a beetle. Here's an observation: https://www.inaturalist.org/observations/236789734
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