A Peculiar Patch of Pussytoes

This week (on account of tomorrow is the deadline) I got around to writing up the results of the two-day survey of leaf-mining moths that Julia and I conducted at Black Rock Forest at the end of August. I showed the list to Jason Dombroskie, who is compiling a list of the Lepidoptera of New York, and he informed me that 17 of the species we found have never been documented in New York before—not counting the four that don’t have names yet. And since he is keeping track of which species have been found in each of New York’s seven ecoregions, he was able to determine that another 27 species we found are new for the Lower New England / Northern Piedmont ecoregion. Not bad for a two-day survey!

Near the end of the second day—shortly after finding the previously unknown immature stages of a Bucculatrix species (more on that later, maybe)—we walked past a patch of pussytoes (Asteraceae: Antennaria ?neglecta) and I did a double-take: “That’s not what pussytoes normally looks like, is it?”


Nearly every rosette bore a dense cluster of leaves, extra densely covered with woolly white hairs.


These reminded me of some midge galls that occur on willows and goldenrods, but I’d never heard of such a thing on pussytoes. I opened one up and sure enough, it had a central cell with something inside…


…and since I recognized the “something” inside as the pupa of a parasitoid wasp, we filled several vials with galls, with the hope that at least one would contain a midge that wasn’t parasitized.

Over the next two weeks, nine eurytomid wasps emerged…


…along with that single midge I had hoped for:


In these last two shots, she’s showing off her incredible ovipositor, no doubt specially designed for piercing pussytoes plants.


Naturally, I sent her off to Ray Gagné at the Smithsonian. He had never found these galls before either, in over fifty years of studying gall midges, but he reported that the specimen matches the description of Asphondylia antennariae, which was described in 1889 from specimens reared from similar galls on plantain-leaved pussytoes (Antennaria plantaginifolia) collected in Milwaukee. In the key to Antennaria galls in Ray’s 1989 The Plant-feeding Gall Midges of North America, the galls of this species are distinguished from the similar galls of Rhopalomyia antennariae by the latter’s having recurved tips on some of the leaves that form the gall. Asphondylia antennariae galls are not supposed to have leaves that are recurved at the tips, but this turns out not to be a consistent difference, based on the galls I found.


The descriptions in Ray’s species accounts provide a more reliable distinction: the Rhopalomyia galls are polythalamous (multi-celled), whereas the Asphondylia galls are monothalamous, each containing a single larva in a central cell—as I found when I tore open the first gall.

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Swallowtail Surprise

I’ve made it up to the end of August in my photo sorting, which means I’m now going through all the pictures I took over the two days Julia and I spent surveying for leaf-mining moths at Black Rock Forest in New York (the results of which, incidentally, you can peruse here if you like). You may recall that on the first day of the survey we bumped into a bald-faced hornet devouring a yellowjacket. One of the highlights of the second day was getting to witness the defense display of a young tiger swallowtail caterpillar. It was just sitting there on a black cherry leaf, doing a very good job of mimicking a bird dropping…


…when suddenly—maybe because it was startled by my camera’s flash—it decided to stick out its fake snake tongue!


That thing is called an osmeterium, and you can read all about it on Wikipedia. All swallowtail caterpillars have one, and the effect is particularly striking with older larvae that also have big fake eyes on their backs:


…Although maybe it’s even more surprising to have a forked tongue pop out of a piece of bird poop than out of something that looks like a little cartoon snake.

From above, this caterpillar seems to be sporting a goofy grin:


That yellow-orange “birthmark” under its right “eye” is troubling; I’m not sure what might be behind that. Anyway, the caterpillar’s real head, of course, is down below:


And if you look really close, you can see that rather than having a single eye on each side of its head, it has a cluster of five ocelli. This feature is common to all butterfly and moth caterpillars (except for a few tiny leafminers), and it’s a good thing to look for when trying to decide if you’re looking at a caterpillar or a sawfly larva (which would only have one ocellus).


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Another year, another 20 new species

2020 is off to a good start! This afternoon I gave my “Native Plants as Insect Habitat” slideshow in the Berkshires, and on the way home stopped at Noah’s house because his not-quite-six-year-old son wanted to show me the bobcat tracks they had been following all week. We got to the tracks just as it was getting too dark to see, but we followed them by flashlight for a little while, and before too long met the tracks of a yearling bear—not a common sight on January 1. So we got to trail the bear for a bit as it stopped to sniff and follow tracks of other animals, including the bobcat, a red fox, and a coyote.

Anyway, a year has gone by since I wrote my “How Many New Species?” post, so I’m re-posting it here with the 20 additional species described in 2019. Species are listed in the order in which they were described, so if you already read the previous post, you can just go to each list and skip down to the species with “2019” after the authors’ names.


I am often asked how many new insect species I have found (or “discovered”). I’m never quite sure how to answer this. I’ve certainly reared dozens of undescribed species of moths, for instance, that are now sitting in my office or in various museums, waiting to be named. The number for parasitoid wasps is probably even higher. But simply having “found” new species doesn’t count for much if they haven’t been properly documented and named. Also, to me the credit for “discovering” a new species mostly belongs to the taxonomist who does the hard work of comparing it with all the similar species in the world to demonstrate that it is really something new. Of all the species I have coauthored, Marmara viburnella is the only one I felt certain was undescribed (because I went to the trouble of reviewing the larval biology and adult morphology of all the previously described species in that genus) before I passed it along to a taxonomist.

With that said, for my final post of the year, I thought it would be fun to put together a list of all the species that have been either described in papers I coauthored or described based (at least in part) on specimens I collected. (This was partly inspired by my realization that I never got around to writing anything here about most of the 30 fly species I recently described with Owen Lonsdale.) With any luck, this list will continue to grow. What limits the number of new species I’m able to help describe is a shortage not of “new” species to name, but of time that my collaborators and I have available to devote to this task. So the take-home message from this post should not be “Wow, look at all the new species Charley has found!”, but rather, “Wow, we have so much left to learn about our natural surroundings, and we need to support more funding for taxonomy!”

For species I’ve written about before, you can click on the name to see the relevant post.

First, the species I did not coauthor (of these, Orchestomerus eisemani and Adelius floridensis are the only species for which one of my specimens was designated as the holotype):

1. Celticecis cornuata Gagné, 2013 – A hackberry gall midge I found in Kentucky while traveling with Noah to check out the periodical cicadas in Nashville and Sam Droege’s bees in Maryland.

2. Orchestomerus eisemani Yoshitake & Anderson, 2015 – A leafminer of Virginia creeper I found at work one day in Plymouth County, Massachusetts. This seems to be its northern range limit; although I’ve since found it in Nashville and at several sites in Ohio, I have found it just one other time in Massachusetts, at Lake Chaubunagungamaug.


3. Brachys howdeni Hespenheide (in Hespenheide & Eiseman, 2016) – I first found this trailing arbutus leafminer while hiking along the ridge just above the house where I now live. I see the mines in just about every sizable patch of the host plant I encounter.


4. Liriomyza limopsis Lonsdale, 2017 – Owen had already given this species a name based on Canadian specimens collected as adults, but no host plant was known until I reared it from white wood aster (Eurybia divaricata) and whorled aster (Oclemena acuminata) in New York and Massachusetts.


5. Liriomyza pilicornis Lonsdale, 2017 – Similar story, except that Graham Griffiths was the first to rear this species, 45 years before Julia and I found it mining leaves of bastard toadflax (Comandra umbellata) in Massachusetts.


6. Liriomyza pistilla Lonsdale, 2017 – Ditto, except in this case the host is cow-wheat (Orobanchaceae: Melampyrum lineare) and Griffiths reared it 40 years before me. I find the leaf mines pretty regularly.


7. Adelius floridensis Shimbori & Shaw, 2019 – This braconid wasp species is known only from a few specimens I reared in 2013 from the St. Johnswort leafminer Fomoria hypericella (Nepticulidae) in Florida.


So that’s seven in the first category. I mostly just happened to give specimens to the right taxonomists at the right time: Ray Gagné was finishing up a revision of all the gall midge species on hackberries; Henry Hespenheide was (and is) in the midst of revising the genus Brachys, and Owen Lonsdale was finishing up a revision of the Canadian species of Liriomyza. In the case of Orchestomerus eisemani, Bob Anderson was inspired to revise that genus after having initially identified the weevils I had reared as O. wickhami Dietz, then discovered his error after I had published a note documenting their natural history. Adelius floridensis got to be described because Dave Wagner directed Scott Shaw and Eduardo Shimbori to me when they asked him if he had any specimens they might use in their revision of the New World braconid wasps of the tribe Adeliini. For the 62 species listed below that I have coauthored, I’m extremely grateful to the taxonomists who took time away from whatever other projects they were working on to help me put names to my natural history observations.

1. Scolioneura vaccinii Smith & Eiseman (in Smith et al. 2015) – A sawfly that mines leaves of huckleberries (Vaccinium spp.), which Julia and I found in western Washington on our first cross-country trip in search of leafminers (though we only were able to rear parasitoids, and the type specimen was reared from a larva Noah and his wife Sydne collected the following year).

2. Megaselia nantucketensis Eiseman & Hartop, 2015 – A scuttle fly that emerged from a midge gall on black oak, collected on Nantucket during the gall and leaf mine survey I’ve been conducting there since 2011.

3. Megaselia risoria Hartop, Wong & Eiseman, 2016 – The naming of this species was a byproduct of my having reared specimens of M. globipyga from a dead tussock moth caterpillar I found at work.

4. Platygaster pruni Buhl & Eiseman, 2016 – A platygastrid wasp that emerged from a midge gall on black cherry, which I collected at work one day in western Massachusetts.

5. Platygaster uvulariae Buhl & Eiseman, 2016 – A platygastrid wasp that emerged from a midge gall on wild oats (Uvularia sessilifolia)—again collected at work in western MA. No one has yet been able to rear the midge that causes this gall.

6. Platygaster vitisiellae Buhl & Eiseman, 2016 – A platygastrid wasp that emerged from a midge gall on wild grape, collected as part of the Nantucket survey. The midge species is probably undescribed (but I was able to rear some adults, which are sitting in the Smithsonian waiting for someone to decide to revise the genus Vitisiella).

7. Zygoneura calthella Eiseman, Heller & Rulik, 2016 – A dark-winged fungus gnat that feeds in leaves and petioles of marsh marigold. Julia and I first found it while surveying for four-toed salamanders in western Massachusetts.

8. Fenusa julia Smith & Eiseman, 2017 – A sawfly that mines leaves of wild rose, which Julia spotted in Colorado on another leafminer-hunting road trip.


9. Marmara viburnella Eiseman & Davis (in Eiseman et al. 2017) – Another product of the Nantucket survey. The larva of this moth begins as a leafminer, then disappears down the petiole and spends most of its life feeding in the stem. Julia and I reared it from arrowwood, but mines have also been found on various other viburnums.

10. Platygaster tephrosiae Buhl & Eiseman, 2017 – Another one from Nantucket; I reared the two known specimens from midge galls that happened to be on some goat’s rue leaves that Kelly Omand collected for me to feed some leaf-tying caterpillars. I failed to rear the caterpillars, and I haven’t been able to rear the midge yet either.

11. Platygaster vaccinii Buhl & Eiseman, 2017 – The single known specimen emerged from a gall on lowbush blueberry that I collected at the 2016 Berkshire BioBlitz on Mt. Greylock—caused by another midge that has never been reared.

12. Macrosaccus coursetiae Eiseman & Davis, 2017 – Another one Julia and I collected on our first cross-country trip; this one from Arizona, mining leaves of a shrub called rosary babybonnets (Coursetia glandulosa).

13. Phytosciara greylockensis Eiseman, Heller & Rulik, 2018 – Another one from the Mt. Greylock BioBlitz; a dark-winged fungus gnat that mines leaves of bluebead lily (Clintonia borealis).

14. Agromyza fission Eiseman & Lonsdale, 2018 – Owen had already named this species based on a specimen collected in the DC area in 1914, but the type specimen is one Julia and I collected at MJ Hatfield’s “Red Oak Prairie” in eastern Iowa on the way home from Colorado. One of the paratypes came from a larva we collected the next day on Marcie and Mike O’Connor’s land in Wisconsin, and Mike Palmer provided two from Oklahoma. The larvae mine leaves of hackberry.


15. Agromyza soka Eiseman & Lonsdale, 2018 – This is another one that Owen had already named based on a 1914 specimen from the DC area, but as with A. fission its host was unknown. It turns out to be responsible for leaf mines on black locust that since 1982 have been attributed to Phytoliriomyza robiniae (Valley), adults of which were repeatedly associated with black locust but never actually reared. Some paratypes came from specimens Julia and I reared from larvae we collected at the 2016 Connecticut BioBlitz, and the rest came from larvae Tracy Feldman found mining both black locust and wisteria in North Carolina.


16. Melanagromyza palmeri Eiseman & Lonsdale, 2018 – The only known specimen is one that Mike Palmer reared from a sunflower stem (or possibly the roots) in Oklahoma. Stem feeding members of this genus are borers rather than miners, meaning that they don’t form any externally visible trails. So rearing them takes special dedication and/or luck.

17. Ophiomyia euthamiae Eiseman & Lonsdale, 2018 – This species mines leaves of grass-leaved goldenrod (Euthamia graminifolia), mostly on the lower surface. I first noticed mines like this on Nantucket, but those were possibly made by O. maura; the whole type series of O. euthamiae came from my yard.


18. Ophiomyia mimuli Eiseman & Lonsdale, 2018 – This species mines in stems of monkeyflower. I first found it at a bioblitz on Julia’s family’s land in southern Ohio, and some paratypes came from the swampy woods right behind our house in Massachusetts.


19. Ophiomyia parda Eiseman & Lonsdale, 2018 – Another species whose holotype I collected in my yard. It is a common leafminer of asters (Symphyotrichum spp.) and seems to be responsible for all of the mines previously attributed to O. quinta.

20. Calycomyza artemisivora Eiseman & Lonsdale, 2018 – This species is known only from two specimens I reared from leaf mines on Artemisia ludoviciana that Mike Palmer collected in Oklahoma.


21. Calycomyza avira Eiseman & Lonsdale, 2018 – Another one that Owen had already named before I reared it; there are several specimens at the Smithsonian from Connecticut, New York, and West Virginia, dating back to 1929. The larvae mine leaves of beggar-ticks (Bidens spp.). I reared some from mines I collected at work, and Tracy Feldman provided some from North Carolina.


22. Calycomyza eupatoriphaga Eiseman & Lonsdale, 2018 – This belongs to the same species complex as C. artemisivora. It has been reared from three plants in the tribe Eupatorieae: I found it on white snakeroot (Ageratina altissima) in Massachusetts and on blue mistflower (Conoclinium coelestinum) in Tennessee, and Mike Palmer found it on late boneset (Eupatorium serotinum) in Oklahoma. In teasing apart the members of this complex, Owen found a specimen that was collected in Ontario in 1947, which he included as a paratype. The holotype is from the woods right behind our house.


23. Calycomyza vogelmanni Eiseman & Lonsdale, 2018 – I reared the only known specimen from a leaf mine on thin-leaved sunflower (Helianthus decapetalus), which I collected near Burlington, Vermont, where I went to grad school. I named the species after Hub Vogelmann, who founded my graduate program (the Field Naturalist program). He had retired long before I attended, but he was very enthusiastic about my first book when it was published, and he was supportive as I got started on my leafminer book project.


24. Cerodontha edithae Eiseman & Lonsdale, 2018 – This species is an iris leafminer, the only known specimen of which Julia and I reared as part of our Nantucket surveys. I named it after Edith Andrews, who died in 2015, a day after her 100th birthday. She lived on Nantucket for most of her life and was an enthusiastic naturalist to the end. Birds were her main passion, but not long after Julia gave her a copy of my book, Julia went to visit her and found her and her daughter Ginger on their hands and knees in her driveway, getting a closer look at some wasp burrows. When I first met her, I was amazed at the memory of this nearly 100-year-old woman as she quoted from various parts of my book.


25. Cerodontha feldmani Eiseman & Lonsdale, 2018 – Another species known from a single specimen; in this case one I reared from a sedge leaf mine that Tracy Feldman collected in North Carolina. Tracy has been intensively collecting leafminers in North Carolina and elsewhere for the past few years and has found numerous new state records, new host records, and new species.


26. Liriomyza ivorcutleri Eiseman & Lonsdale, 2018 – I reared the single known specimen from a leaf mine on cup plant (Silphium perfoliatum) that Julia and I found in Iowa. When Owen told me this yellow fly was a new species, I couldn’t resist naming it after Ivor Cutler, the Scottish recording artist responsible for “Yellow Fly,” along with other classics like “I Believe in Bugs.”


27. Liriomyza valerianivora Eiseman & Lonsdale, 2018 – I found the leaf mines of this species in a scrappy wetland in north-central Massachusetts where I was conducting botanical fieldwork with Sally Shaw. I was lucky she was with me, because I never would have recognized the basal leaves of garden valerian, which are totally different from those on mature plants.


28. Phytomyza actaeivora Eiseman & Lonsdale, 2018 – I tried for several years to rear adults from leaf mines on red baneberry (which I’ve found in Vermont and Ohio) before finally succeeding with some mines I found on white baneberry in my neighbors’ woods. I also found mines of what is probably the same species on black cohosh (these are all Actaea species) at Jason Dombroskie’s house in Ithaca, NY, but these were all parasitized like the ones on red baneberry.


29. Phytomyza aesculi Eiseman & Lonsdale, 2018 – I first became aware of this species because of photos of buckeye leaf mines that several different people posted to BugGuide.net. One spring when Julia was visiting her parents in Ohio (the Buckeye State), she managed to collect a bunch of larvae, from which I reared the type series. This species is active only in spring, with a pupal diapause lasting nearly a year. The author of this article was grateful when I let him know that his mystery “buckeye leafmining fly” now has a name.


30. Phytomyza confusa Eiseman & Lonsdale, 2018 – I named this fly “confusa” because everything about it was confusing. I found the leaf mines at the base of a tree in the middle of a lawn in a big park in Iowa. I tentatively identified the plant as Virginia waterleaf (Hydrophyllum virginianum), but it looked a little weird to me (not to mention that Virginia waterleaf is normally a forest species). Iowa botanist John Pearson suggested that it might be a buttercup such as Ranunculus fascicularis. When Owen initially determined the flies as belonging to a new species in the Phyomyza aquilegiae group, this seemed to fit, since all members of this group feed on plants in the buttercup family as far as is known. I showed my photo of the plant to several other botanists, and they all shared my initial impression that it was Virginia waterleaf, but most were also willing to believe it was Ranunculus fascicularis, and one even examined some herbarium specimens of that species that she said matched in every respect. But Owen later determined P. confusa to be closely related to another new species that I reared from two species of waterleaf (see below), and decided both flies probably are better placed in the P. obscura group, all species of which feed on plants in the mint and borage families (waterleafs are in the latter). This species was also confusing because the leaf mines were hard to characterize—some began with a distinct linear portion and some did not, and by the time the adults emerged the leaves were so crumpled and degraded that I couldn’t decide whether the puparia were formed inside or outside the mines.


31. Phytomyza doellingeriae Eiseman & Lonsdale, 2018 – While working in Maine in July 2013, I collected leafminers from flat-topped aster (Doellingeria umbellata) that Owen determined to be a new species near P. solidaginivora Spencer based on the genitalia. Both of the adults I reared were underdeveloped (as shown here), so when I returned in August I collected some more. I reared some good specimens this time, but Owen determined them to be a different new species, which I named P. doellingeriae. Meanwhile, he decided the first flies were close enough to P. solidaginivora to call them that for now. Incidentally, Spencer (1969) reared P. solidaginivora from a plant in Alberta that he thought was goldenrod (Solidago; hence the name), but Graham Griffiths examined his pressed leaf mines and didn’t think they looked like any goldenrod that occurs in Alberta. Spencer’s drawing of the leaf looks exactly like a flat-topped aster leaf, so that fly is probably misnamed.


32. Phytomyza erigeronis Eiseman & Lonsdale, 2018 – I first found this leafminer of daisy fleabane (Erigeron) in my front yard. Some of the paratypes came from the 2016 Connecticut BioBlitz.


33. Phytomyza hatfieldae Eiseman & Lonsdale, 2018 – When Julia and I stayed with MJ Hatfield in northeastern Iowa on the way home from Colorado, we spent a little time exploring the woods on the bluff next to her house with MJ and John van der Linden. Leaf mines that we collected there on sweet cicely (Osmorhiza claytonii) yielded the holotype of this new species. The paratypes also included a number of specimens Graham Griffiths had reared from various Osmorhiza species in the 1970s, plus a few that Ed Stansbury reared in Washington just in time to be included in the paper.


34. Phytomyza hydrophyllivora Eiseman & Lonsdale, 2018 – This species is common on broad-leaved waterleaf (Hydrophyllum canadense) in Ohio, and I collected the mines several times from the woods by Julia’s parents’ house before I finally got a few adult flies instead of parasitoid wasps. I later reared one from the same host in Tennessee (during our brief trip to see the solar eclipse last year), and one from a mine I found on Virginia waterleaf while conducting a rare plant survey in the Berkshires.


35. Phytomyza palmeri Eiseman & Lonsdale, 2018 – This is another species (like Melanagromyza palmeri) known only from Mike Palmer‘s yard in Oklahoma, and although he gave me a number of leaf mines, only he has been able to rear adults. The larvae mine leaves of coralberry (Symphoricarpos orbiculatus).


36. Phytomyza palustris Eiseman & Lonsdale, 2018 – I found this leafminer of swamp saxifrage while conducting botanical fieldwork in the Berkshires. I check this plant for mines every time I see it, but as far as I can tell the range of this species is limited to one square meter in the town of Washington, MA.


37. Phytomyza sempervirentis Eiseman & Lonsdale, 2018 – Julia and I first found this species when we visited Cane Creek Canyon in northern Alabama on our way home from Florida in spring 2013. The larvae form mines on coral honeysuckle (Lonicera sempervirens) very similar to those formed by the closely related P. nigrilineata (Griffiths) on limber honeysuckle (L. dioica) in Alberta. I found more (including the holotype) three years later at the Montague Plains in western MA. Tracy Feldman also provided a bunch of specimens from North Carolina, and Mike Palmer reared a few from orange honeysuckle (L. ciliosa) in Oregon.


38. Phytomyza tarnwoodensis Eiseman & Lonsdale, 2018 – I reared the only known specimens of this species from leaf mines on bush honeysuckle (Diervilla lonicera) I collected in my parents’ yard in western MA. “Tarnwood” is the name my parents gave to their property many years ago, and when I was little this sign that my mother painted used to be on a post at the edge of our yard by the road:



39. Phytomyza tigris Eiseman & Lonsdale, 2018 – The larvae of this species mine leaves of foamflower (Tiarella cordifolia). The leaf mines are very common, but it took me many tries (always getting only parasitoid wasps) until I finally managed to rear a few adults—in my neighbors’ woods right near where I finally found unparasitized puparia of P. actaeivora. The name Phytomyza tiarellae was already taken, so I named this one “tigris” after the tiger stripes on its puparium (going with the “big cat” theme Owen had started with Ophiomyia parda).


40. Phytomyza triangularidis Eiseman & Lonsdale, 2018 – This is another one Julia and I found on our first cross-country trip, this time in northern Idaho. The larvae mine leaves of arrowleaf ragwort (Senecio triangularis).


41. Phytomyza vancouveriella Eiseman & Lonsdale, 2018 – Although Julia and I found a few leaf mines of this species on the Olympic Peninsula on that same trip, the only known specimens are a few that Mike Palmer reared in Oregon five years later. The host is Vancouveria hexandra, whose common names include “white inside-out flower.”


42. Phytomyza verbenae Eiseman & Lonsdale, 2018 – One last species (for now) from that first road trip; Julia and I found the mines on western vervain (Verbena lasiostachys) in California.


43. Phytomyza ziziae Eiseman & Lonsdale, 2018 – I reared the holotype and some of the paratypes from leaf mines on golden Alexanders (Zizia aurea) I collected while conducting botanical fieldwork in western MA. Another came from the same Berkshire BioBlitz that produced the only known specimens of Phytosciara greylockensis and Platygaster vaccinii. There are also a few specimens that Graham Griffiths reared from heart-leaved golden Alexanders (Zizia aptera) in Alberta in 1973. It’s a bit curious how many agromyzid species are known only from Alberta and Massachusetts…


44. Agromyza arundinariae Eiseman, Lonsdale & Feldman, 2019 – This and the next eight species were described in a paper dedicated to agromyzid flies that Tracy Feldman collected in North Carolina—I helped with rearing and described the leaf mines, and Owen Lonsdale described the adult flies. Agromyza arundinariae is one of three new species Tracy found on the native bamboo Arundinaria tecta.


45. Agromyza indistincta Eiseman, Lonsdale & Feldman, 2019 – I gave this one the name “indistincta” because there was nothing distinctive about it; it’s the fourth species to be reared from seemingly identical mines on grasses in the genus Dichanthelium, and the adult is pretty darn similar to two of the other three.


46. Calycomyza chrysopsidis Eiseman, Lonsdale & Feldman, 2019 – A leafminer of Maryland goldenaster (Asteraceae: Chrysopsis mariana).


47. Cerodontha enigma Eiseman, Lonsdale & Feldman, 2019 – This one is a similar situation to Agromyza distincta; it is a leafminer of Dichanthelium and is an enigma because it is known from a single specimen Tracy collected in his yard. The adult is very similar to Cerodontha angulata, which is the species I’ve reared every time I’ve collected similar mines on Dichanthelium.


48. Cerodontha arundinariella Eiseman, Lonsdale & Feldman, 2019 – The second species reared from leaf mines on Arundinaria tecta.


49. Cerodontha saintandrewsensis Eiseman, Lonsdale & Feldman, 2019 – The third species reared from leaf mines on Arundinaria tecta, named for St. Andrews University, where Tracy works and does much of his collecting (and the only known locality for this species).


50. Liriomyza carphephori Eiseman, Lonsdale & Feldman, 2019 – Tracy first found this species mining leaves of sandywoods chaffhead (Asteraceae: Carphephorus bellidifolius), and I had already decided to name at after this plant when Owen determined that it was the same species Tracy and I had reared from beggarticks (Bidens spp.) in North Carolina, Vermont, and my own front yard in Massachusetts. It’s still a good name for it; several other Liriomyza species mine leaves of Bidens, but this is so far the only leafminer of any kind to be found on Carphephorus.


51. Liriomyza polygalivora Eiseman, Lonsdale & Feldman, 2019 – A leafminer of whorled milkwort (Polygalaceae: Polygala verticillata).


52. Liriomyza triodanidis Eiseman, Lonsdale & Feldman, 2019 – A leaf and stem miner of small Venus’ looking-glass (Campanulaceae: Triodanis biflora).


53. Agromyza princei Eiseman & Lonsdale, 2019 – I reared this species from a leaf mine on black raspberry (Rosaceae: Rubus occidentalis) that Julia and I collected in the parking lot of a cemetery in Hartford at the 2016 Connecticut BioBlitz. It is known from a single specimen, which emerged as an adult a year after I collected the larva. When Owen told me he needed a name for this new species, “Raspberry Beret” popped into my head, so I named it after Prince, who had died shortly before we found the leaf mine.


54. Melanagromyza vanderlindeni Eiseman & Lonsdale, 2019 – This species is named for John van der Linden, who reared it from dead stems of Joe-Pye weed (Asteraceae: Eutrochium) he collected in Iowa. John has an incredible knack for finding stem-feeding insects that leave little or no external evidence. He has written about some of his natural history discoveries on his blog, and many more can be found on his BugGuide page.

55. Ophiomyia antennariae Eiseman & Lonsdale, 2019 – Julia and I found this leafminer of plantain-leaved pussytoes (Asteraceae: Antennaria plantaginifolia) in beautiful Cane Creek Canyon in northern Alabama in the spring of 2013, shortly after getting in a car wreck in Mobile. The mines are much like those from which Mike Palmer and I have reared O. coniceps in Oklahoma and New England, and Owen and I almost described this new species in our first (2018) paper, but I temporarily convinced him that we should consider the Alabama specimens to be O. coniceps. However, before that paper went to press, Owen reasserted his original position—though he considers one female from Cane Creek Canyon to be a better match for O. coniceps—so we removed the remaining specimens from that paper and dealt with them in the 2019 paper. As it happens, Julia and I had separated out what we thought might be two different mine types on plantain-leaved pussytoes, and that female was the only one that emerged from mines of the type that O. coniceps makes in New England. But Mike has reared O. coniceps from both mine types in Oklahoma, so evidently the difference isn’t entirely consistent.


56. Ophiomyia osmorhizae Eiseman & Lonsdale, 2019 – Another John van der Linden discovery from Iowa; this one is a stem miner of sweet cicely (Apiaceae: Osmorhiza).

57. Calycomyza smallanthi Eiseman & Lonsdale, 2019 – In August 2017, Julia and I met up with Noah’s family in Nashville to see the full solar eclipse, and we found this species mining leaves of hairy leafcup (Asteraceae: Smallanthus uvedalius) just around the corner from Noah’s mother’s house.


58. Liriomyza euphorbiella Eiseman & Lonsdale, 2019 – Mike Palmer reared this one from fire-on-the-mountain (Euphorbiaceae: Euphorbia cyathophora) in Oklahoma.

59. Liriomyza garryae Eiseman & Lonsdale, 2019 – Julia and I found this species mining leaves of silktassel (Garryaceae: Garrya spp.) in Arizona and Texas on the way home from checking out the “super bloom” in southern California in 2017.


60. Liriomyza phloxiphaga Eiseman & Lonsdale, 2019 – I reared the only known specimen of this species in 2017 from a leaf mine on phlox (Polemoniaceae: Phlox paniculata) in my mother’s garden in Massachusetts.


61. Phytomyza nemophilae Eiseman & Lonsdale, 2019 – Mike Palmer reared this species from leaf mines on Nemophila parviflora (Hydrophyllaceae) in Oregon.

62. Phytomyza salviarum Eiseman & Lonsdale, 2019 – Julia and I found this species mining leaves of several different sage species (Lamiaceae: Salvia) on Ann and Bruce Hendrickson’s ranch in Texas in 2017, the same day we collected the holotype of Liriomyza garryae.


That’s the current tally—more coming soon!

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Cup Plant Creature

At the end of June, Julia and I visited Marci and Jim Hess and explored some of their remnant and restored prairie in Lafayette County, Wisconsin. We were of course focusing on collecting leafminers, and I spotted something unusual on a cup plant leaf that I decided to collect even though it turned out not to be a leaf mine. Cup plant (Asteraceae: Silphium perfoliatum) gets its name from the way opposing petioles are fused at the base, forming a “cup” that catches rainwater:


These brown patches at the edge of one of the leaves looked like they could be mines of a casebearer moth larva (Coleophoridae: Coleophora)…


…but flipping the leaf over revealed that they were actually “window feeding,” the work of a caterpillar feeding on the lower leaf surface under the protection of some webbing it had spun.


The white object to the left looked to me like the cocoon of a metalmark moth (Choreutidae). I’m only familiar with a few species, and wasn’t sure if one was known to feed on cup plant.


On July 7, my hunch proved to be correct when the gorgeous adult moth emerged:


Knowing the host plant, I was quickly able to identify it as Tebenna silphiella, which has been given the common name of “Rosinweed Moth” because apparently rosinweed (Silphium integrifolium) is (was) its only known host. The Latin name, which allows for the possibility of it eating other Silphium species, turns out to be more appropriate.

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Life Lurking in Lousewort

Not long after we started to let our lawn go wild, a couple of blue-eyed grass plants (Sisyrinchium montanum) popped up in one area. Blue-eyed grass owes its name to its grass-like leaves; it is actually in the iris family (Iridaceae).


In the past year or so the plants have really started to multiply; here’s a shot of one patch from this past June:


I’ve been keeping a close eye on this part of the yard to see what else shows up, and this spring I was excited to see some Pedicularis canadensis (Orobanchaceae)—known as wood betony or Canadian lousewort—peeking out among the Queen Anne’s lace, plantain, and dandelion.


The latter three, of course, are all weedy European species, though not without value to native insects (in addition to all being edible for humans). The yard is full of adults and leaf-mining larvae of plantain flea beetles every spring; black swallowtail caterpillars devour Queen Anne’s lace as readily as any native member of the parsley family; and a variety of bees, flies, and other nectar feeders visit dandelion flowers (and later, goldfinches and indigo buntings come to eat the seeds). Pedicularis canadensis, on the other hand, is a native plant supporting a suite of specialized, little-known insect herbivores. It’s also interesting because it is a hemiparasite, stealing nutrients from neighboring plants through its roots.


I wondered how it had gotten there, and as I thought about it more, I wasn’t entirely sure I hadn’t sprinkled some seeds there myself. I have a favorite conservation area I try to visit every spring, which includes a meadow that is full of all sorts of unusual wildflowers, including the largest and densest population of Pedicularis canadensis I’ve ever seen. On May 6, 2012, I photographed this ~2 mm fly on one of the Pedicularis flowers there:


At that point I was just beginning to focus my attention on leafminers, and I didn’t recognize this as a Phytomyza, a member of the most diverse genus of “leafminer flies” (Agromyzidae). When I posted photos of this fly on BugGuide.net, Owen Lonsdale identified the genus and added that of the three species known to feed on Pedicularis, this fly most resembles Phytomyza pedicularicaulis, but it doesn’t perfectly match any of them. It turns out that none of these Pedicularis-feeding Phytomyzas are leafminers, and none have been found in New England. Phytomyza pedicularicaulis (known only from Quebec) is a stem borer of Pedicularis canadensis; larvae of Phytomyza tenella (known from Colorado, Quebec, Labrador, and Europe) feed in the seed capsules; and those of Phytomyza pedicularidis (again known only from Quebec) also feed in seed capsules but of Pedicularis groenlandica. So if I could find these flies again, I would at minimum be adding a new geographic record for one of these species and maybe they would turn out to represent a fourth, previously unknown species.

This occurred to me when I visited the conservation area on June 7, 2018, but the flowers had gone by at that point, so I decided to open a few seed capsules and see if there were any larvae inside. I found some right away, so I filled a few vials with seed capsules to try and rear them to adults. I didn’t get around to taking any photos until June 18, when several larvae had emerged and formed puparia:


These were 4 mm long, and at the time I thought “gosh, these are the biggest agromyzid puparia I’ve ever seen!” Of course, the fly I was after was only 2 mm long, so these were clearly something else, but I was distracted right then by this gorgeous 2 mm pteromalid wasp that had also just emerged from one of the seed capsules:


From the photo, Roger Burks identified this as belonging to the genus Lyrcus. Species of Lyrcus have been reared mostly from various weevils and gall midges, as well as a few other insects—the common theme evidently being larvae that feed concealed in plant tissues. I found the hole where this wasp had chewed its way out of a seed capsule,


but I couldn’t find any identifiable host remains inside. The next day, a few tiny gall midge larvae (Cecidomyiidae) appeared in one of the vials,


and three days later, a 1.5 mm adult…


The latter, at least, was a Clinodiplosis. These appear fairly regularly in my rearing vials and it’s usually not quite clear what they’re up to. Larvae of most species feed on fungi; many feed in galls induced by other insects, and some are apparently plant feeders.

On July 6, two of these 1.5 mm frit flies (Chloropidae: Oscinellinae) appeared.


These are another mystery; I get frit flies in my rearing vials from time to time as well, and this family includes herbivores, scavengers, parasites, and predators, so again it isn’t obvious what the larvae were doing in the Pedicularis (especially since the vials contained the peduncles/stems as well as the fruits).

On July 10, out popped a 5.5 mm adult of Endothenia hebesana (Tortricidae), leaving its pupal exuviae protruding from the seed capsule in which the caterpillar had fed.


This moth has been reared from seeds of a variety of plants, but the recorded hosts show a decided preference for plants in the order Lamiales.

Nothing further emerged before I put all my rearing containers in the fridge for the winter. But then, beginning a few weeks after I took them out, half a dozen of these ~4 mm flies emerged:


These were, of course, the adults of the larvae that I had seen feeding inside the seed capsules and that had emerged and formed the 4 mm puparia. I immediately recognized them as scathophagids, having become familiar with several leaf-mining species, and I should have recognized the puparia too. What threw me off is that virtually all scathophagids are associated with monocots—100% of the leafminers are (hosts include orchids, true and false Solomon’s seal, Canada mayflower, and false hellebore); as I discussed here, species of Cordilura are stem borers in sedges; and so on. I learned today that these Pedicularis flies are Gimnomera cerea; as far as is known, the larvae develop exclusively in fruits of P. canadensis, whereas the related G. incisurata feeds in fruits of several Penstemon species, which are also hemiparasites. When Brad Sinclair identified the flies, I realized that I already had a scan of a paper (Neff 1968*) that goes into great detail about their natural history. At the end of the paper, Neff speculates that when these hemiparasitic plants parasitize grasses, sedges, and other monocots, they take up organic compounds that are attractive to monocot-feeding insects, and that this may have facilitated the evolutionary shift in host plants.

And finally, just after these flies emerged, two pteromalid wasps emerged from two of the remaining puparia.


I have yet to find anyone who has both the expertise and the time to identify my pteromalid specimens, but luckily Roger Burks can recognize many of the genera from photos, and he says this one is a Lamprotatus. Species in this genus are mostly parasitoids of muscoid flies (which include scathophagids, but mainly anthomyiids), but this is the first record of one (or apparently of any parasitoid) from Gimnomera.

Now the question is, how long will it take all of these bugs to find the Pedicularis that just showed up in my yard?

* Neff, S. E. 1968. Observations on the immature stages of Gimnomera cerea and G. incisurata (Diptera: Anthomyiidae, Scatophaginae). The Canadian Entomologist 100: 74-83.


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I’ve been making sporadic progress in sorting through my backlog of photos, and I’m now exactly seven months behind. April 13 saw the conclusion of a story that began with my collecting this scrub oak (Quercus ilicifolia) leaf in early October of 2018:


The brown, wrinkled patch is a leaf mine of a moth in the genus Cameraria (Gracillariidae), but which of the 30 oak-feeding Cameraria species is it? I periodically collect oak Cameraria mines to see if I can figure out ways to distinguish the different species.

I removed this leaf from winter refrigeration on March 1, and on March 10 I discovered this little caterpillar spinning a cocoon in the bottom of the rearing vial:


This was clearly not a Cameraria larva, which would be flattened and legless and would pupate inside the leaf, from which the adult would eventually emerge. It was a larva of one of the bigger micro-moths, probably something in the superfamily Gelechioidea (or maybe a tortricid). It had evidently been inside the mine when I collected the leaf and had spent the winter there.

The next day it was still working on its cocoon; in the photo below it’s lying on its back and adding silk to the ceiling:


On March 20 the larva abandoned its cocoon and disappeared into the crumpled-up piece of toilet paper I had stuffed into the vial for humidity control. At this point I knew my chances of learning the identity of this interloper were not good; a happy, healthy caterpillar does not leave its carefully constructed cocoon behind. On April 13 this ichneumon wasp emerged from a cocoon spun next to the caterpillar’s remains inside the wad of paper.


It turns out I was a bit hasty in saying at the beginning of this post that April 13 was the end of this story. Nearly two months later, on June 10, the adult Cameraria emerged from the same mine:


I tried running it through the key I made here to the known eastern oak-feeding Cameraria species, and I think C. bethunella may be the best match. Unfortunately this is a species complex that needs more work, and I’ve reared much smaller moths with a similar wing pattern from completely different mines. So no name for this moth just yet…

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Foiled Again

Now that I’ve put most of my bugs in the fridge for the winter, I can get back to catching up on sorting through my backlog of photos, which goes back to September of last year. I just reached the conclusion of a little story that started with this leaf mine on an oxeye (Asteraceae: Heliopsis helianthoides) growing along my driveway:


I see these mines in my yard regularly, but they are always empty so I have been unable to rear the larvae out to determine what species they are. On September 1 last year, I finally found this one mine with a larva inside. I spotted it just in time: the moment I picked it, the larva began to exit through a slit it had cut in the lower epidermis:


It jumped a few millimeters once it was all the way out, which would have propelled it to the ground if I hadn’t been holding the leaf upside-down. Within a few hours, its skin had hardened and contracted to form a puparium:


But alas, 24 days later, what emerged was not an adult fly but a figitid wasp, which had been living as a larva inside the fly larva all along:


Matt Buffington tells me (based on looking at the above photo) that the wasp is a Gronotoma species. This genus is in need of revision, and when I met him last December he told me that he probably won’t get around to that until after he retires.

There’s a good chance that the fly was Liriomyza arctii, the only species that has been reared from mines like this on Heliopsis (by yours truly, in Wisconsin). The species was named by Kenneth Spencer in 1969, who reared it from Arctium (burdock) in Canada. The fact that this apparently native fly was only known to feed on a nonnative plant was puzzling, until I reared it from Heliopsis and three other native genera of Asteraceae over the past few years. I’ve also repeated the rearing from burdock in my own yard, which is why I’m reasonably sure L. arctii is what is mining the oxeye and wingstem in my yard (probably also the Jerusalem artichokes and other sunflowers, but Helianthus is not yet a confirmed host for this fly). Here’s one I reared from nodding beggarticks (Bidens cernua), also not far from my house:


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