Adventures in Taxonomy

Apparently today is “Taxonomist Appreciation Day,” so I suppose it’s an appropriate time to write a little something about my latest paper, which I discovered had been published right after I clicked “publish” on my previous blog post. I never intended to become involved in taxonomy at all, but as a naturalist, it gets really hard to make sense of the world when so many of the species I’m studying don’t have names. If I meet a species that no one has gone to the trouble to describe and name, how can I tell how it relates to anything anyone has found before? Once there are names to which to attach observations, I can start to understand patterns in the habits and host plants (or host insects) of different species, and how these differ from species to species. Somebody needs to do the hard work of deciding what the limits are between similar species, and illustrating (in words and pictures) what the differences are. As summarized here, I’ve now had the pleasure of collaborating with various taxonomists to give names to over 60 species of insects whose natural history I’ve written about. This latest paper* doesn’t describe any new species, but it makes several taxonomic changes involving the names of leaf-mining moths in the family Gracillariidae.

It all started in March of 2017, when Julia and I stopped in Arizona on our way home from checking out the “super bloom” in southern California. Although nothing had leafed out yet in the canyons we visited, we found a number of different active mines in the leaves of evergreen trees and shrubs, and we had good success in rearing adults from most of them. These included a Cameraria species on Arizona madrone (Ericaceae: Arbutus arizonica)…


…and another Cameraria on silverleaf oak (Fagaceae: Quercus hypoleucoides):

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When we got home and I started trying to identify all the leaf mines we had found using the keys I’d written, I found that most of them were new to science. These two species were among the few that had names (or at least, that had names that could be linked to leaf mines that had been described in the literature). To my surprise, they were Phyllonorycter arizonella and P. cretaceella. But they both were clearly Cameraria, based both on the leaf mines (flat, on the upper surface, with a longitudinal crease along one margin when complete) and the adult wing pattern (the black edging was on the outer edge of the white markings). I checked the original descriptions; both were species that Annette Braun had described in 1925 from specimens she reared in Arizona, and no one had published anything about them since. She described both species in the genus Lithocolletis, which is now considered a junior synonym of Phyllonorycter but in 1925 also included all the species that are now placed in Cameraria. The genus Cameraria was actually described in 1902—it was named for the Kentucky lepidopterist V. T. Chambers (“camera” being Latin for “chamber”)—but it was decades before the name was adopted for North American species, and most species of “Lithocolletis” didn’t receive their currently recognized names until Don Davis transferred them to Cameraria and Phyllonorycter in the 1983 Check List of the Lepidoptera of America North of Mexico.

Phyllonorycter species mostly make leaf mines on the undersides of leaves, and rather than remaining flat they usually become strongly wrinkled and “tentiform.” I decided to review the leaf mine descriptions of all the other North American Cameraria and Phyllonorycter species to see if any others were on the “wrong” side of the leaf. I found a number of other examples of Phyllonorycter species that sometimes or always make upper-surface mines, but based on features of the larvae and adults (larvae becoming round in cross section in later instars, rather than remaining flat; adult forewings with black edging on the inner margins of the white markings, rather on the outer margins), they appeared to be proper Phyllonorycter species. However, among the Cameraria species there were two that formed underside tentiform mines, and a little investigation suggested that they actually belonged in Phyllonorycter. One was C. leucothorax, which mines leaves of tanoak (Notholithocarpus densiflorus) and canyon live oak (Quercus chrysolepis) in Oregon and California. I’ve never reared it, but this is probably its mine:


The other was Cameraria affinis, an eastern species that mines leaves of honeysuckles and snowberries (Caprifoliaceae: Lonicera and Symphoricarpos).


I wrote to Don Davis about this, and he agreed that all four species were misplaced and should be transferred as I suggested, changing their names to Cameraria arizonella, C. cretaceella, Phyllonorycter affinis, and P. leucothorax. I thought I would just write a short note to set things right, but he felt that something more detailed was warranted, including illustrations of the male and female genitalia of each species, which he offered to provide. He had never seen specimens of C. arizonella (the six that Julia and I reared may be the only ones beyond Annette Braun’s original five, which are in Philadelphia now), so I sent him a male and female of that species, along with whichever sex he was missing for P. affinis. He had reared his own series of C. cretaceella in New Mexico in 1989, and fortunately he already had what he needed for P. leucothorax as well.

When I submitted the manuscript to Zootaxa, I suggested Terry Harrison as a possible reviewer. This turned out to be a very good idea, because when Terry was looking over the manuscript and illustrations, he was struck by the resemblance of Phyllonorycter affinis—in both wing pattern and genitalia—to the species he calls P. mariaeella. As it happens, last October when I visited Harvard University with Julia to give a slideshow about leafminers to the Cambridge Entomological Club, we showed up early and spent a few hours photographing V. T. Chambers’ type specimens that are housed at the Museum of Comparative Zoology. Unfortunately there is no type specimen for P. mariaeella—whatever existed may have been devoured by dermestid beetles, as happened to many of Chambers’ specimens while he was still alivebut filed under that name there are three specimens reared by Mary Murtfeldt in Kirkwood, Missouri. She was the namesake of P. mariaeella, and the species was described from specimens she reared at that same location, so these were as close to type specimens of P. mariaeella as could be hoped for.

Here’s one she reared from snowberry:


And one of two she reared from honeysuckle:


Annette Braun, in her 1908 revision of the genus “Lithocolletis“, had distinguished affinis and mariaeella based on the forewing fasciae (white markings) being “nearly straight”
in the former and “distinctly bent outward near the middle” in the latter. Terry dissected a series of Illinois specimens showing a full range of variation in these markings and found that they all had matching genitalia. David Lees at the Natural History Museum in London was kind enough to send photos of one of the affinis type specimens and its genitalia, and these too matched Terry’s specimens from Illinois as well as Mary Murtfeldt’s specimens from Missouri, and these in turn matched Chambers’ description of mariaeella. Since Chambers described mariaeella in 1875 and affinis was described in 1876 (by Frey & Boll, a couple of German lepidopterists studying material from Texas), mariaeella has priority, and Don and I have designated affinis as a synonym. One less leaf-mining moth to worry about!

The other reviewer I suggested for this manuscript, Steven Whitebread, noticed another possible synonymy when reviewing our illustrations. This is one of the type specimens of Phyllonorycter gemmea, another Frey & Boll species, but one collected in Massachusetts rather than Texas (and that, I suppose, is why part of its type series is deposited in Cambridge, MA rather than in London):


It was allegedly reared from black locust (Fabaceae: Robinia pseudoacacia), but Frey & Boll had some mix-ups with the leaves they had collected and they weren’t sure what type of mine it had come from. They described three different types of mines they had found on black locust, which clearly correspond with three other gracillariids: Chrysaster ostensackenella, Macrosaccus robiniella, and Parectopa robiniella. Braun (1908) incorrectly cited Frey & Boll as having said that Phyllonorycter gemmea makes an upper-surface mine, which is how this species ended up getting mentioned in this “wrong side of the leaf” paper. Chambers was convinced that this moth hadn’t come from black locust at all, and I’m inclined to agree. Steven suggested that it might be synonymous with P. diversella, a leafminer of huckleberry (Ericaceae: Gaylussacia baccata) that Annette Braun described decades later. It does seem plausible that if Frey & Boll had a bunch of mixed-up leaves, they could have mistaken a huckleberry leaf for a leaflet of black locust. But as I considered this possibility, I found two other species that Braun describedP. martiella from birch, and P. viburnella from arrowwoodthat also have a similar wing pattern, and the only way to sort this out will be to dissect the specimen in the above photo (which is the only specimen in the type series that still has an abdomen) and compare its genitalia with those of specimens reared from huckleberry, birch, and arrowwood. A bit beyond the scope of this paperone of many taxonomic issues that still need to be worked out involving species that already have names, let alone the countless ones that remain to be described.

And so, a big thanks to all taxonomistspast, present, and futurefor your ongoing work to help the rest of us make sense of the complexity of nature!

* Eiseman, Charles S. and Donald R. Davis. 2020. Wrong side of the leaf: assigning some Lithocolletinae species (Lepidoptera: Gracillariidae) to their proper genera. Zootaxa 4751(2): 201–237.

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Mallow Munchers

Three years ago, Don and Mignon Davis described a new species of moth, Telamoptilia hibiscivora (Gracillariidae), whose larvae mine leaves of swamp rosemallow (Malvaceae: Hibiscus moscheutos) along the Atlantic Coast*. Specimens were known from Maryland, Louisiana, Texas, and adjacent Mexico, dating back to 1932—sometimes these things take a little time. Swamp rosemallow isn’t a plant I normally encounter in my wanderings, but nonetheless I added T. hibiscivora to my mental list of species to watch for—and, of course, to my complete guide to North American leafminers.

Last spring, when Kelly Omand asked me about some beetles that were emerging from swamp rosemallow seeds she had collected on Nantucket, I immediately made checking those plants a priority for my next visit to the island. In September Julia and I followed Kelly’s directions to the plants and found them in bloom:


More importantly, we found a thriving population of Telamoptilia hibiscivora, with multiple larvae often mining a single leaf.


The above photos show the upper and lower surface of the same three mines. Each mine begins as a linear track on the underside of the leaf, then becomes a greenish, interparenchymal blotch: leaving some green tissue adjacent to both the upper and lower epidermis, with the result that the mines are less distinct than leaf mines typically are. But the older larva continues to nibble away at the cells in the blotch, so that it develops more distinct whitish or brown patches. Here’s a backlit leaf with at least three larvae visible inside:


As noted by Davis & Davis, the mature larva exits its mine “and forms an elliptical, white to brownish white cocoon usually on the leaf surface.”


A detail they didn’t mention is that the finished cocoon is decorated with a few “frothy bubbles” as in certain other gracillariid moths (see this post for more examples).


The original description noted that larvae pupate only when subjected to cold temperatures, and that the pupal stage overwinters, so once everybody had finished mining and spun their cocoons, into the fridge they went. I emptied out the bug fridge a little less than a month ago, and yesterday the first adult Telamoptilia hibiscivora emerged:


And just like that, the species’ known distribution was extended by 500 miles or so.

Thanks to the “Leafminers of North America” project I created on iNaturalist, I recently learned of the existence of similar mines on saltmarsh mallow (Kosteletzkya pentacarpos) in Everglades National Park. Another new species of Telamoptilia, or a new host record for T. hibiscivora? Someone down there will have to investigate.

As for the beetles that live in swamp rosemallow seeds, we got to meet those too. Here’s a portion of a seed capsule with at least two beetles visible:


After spending its whole larval life eating out the inside of a single seed, and pupating in the resulting chamber, the adult weevil emerges by chewing a neat circular lid at one end of the seed.


This is one of three Althaeus species in the US north of Mexico, all of which develop inside mallow seeds; I’m not sure exactly which one. They belong to the subfamily Bruchinae (Chrysomelidae), which are known as “bean weevils,” but that name doesn’t work for them since they don’t feed on beans (not to mention that they aren’t really weevils). Kelly calls them “Gonzos,” which seems reasonable to me.

* Davis, Donald R. and Mignon M. Davis. 2017. First report of the genus Telamoptilia from the western hemisphere with descriptions of two new species (Gracillariidae). Journal of the Lepidopterists’ Society 71(4)261-273.

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Ten Years Later

book_cover_awardSince we have ten fingers and ten toes, I suppose it’s worth mentioning that it was ten years ago today that Tracks & Sign of Insects and Other Invertebrates was first published. People are often surprised when I tell them that Noah and I didn’t know much about the subject matter when we started writing it, and that it didn’t take years and years to finish. We set out to write it precisely because we didn’t know: we wanted a book that would solve those nagging natural history mysteries, like what are those shiny red discs we always find on the undersides of rocks, and what makes those feathery patterns in the algal film on birch bark, and what’s the deal with the flattened, dead flies we keep seeing stuck to the undersides of leaves?


Mark Elbroch had suggested we give ourselves a year to put this book together, but given the vastness of the topic and how little we knew about it, I decided on a year and a half. In the fall of 2007 I began scouring the literature in earnest; throughout 2008 we took thousands of photos, traveling throughout the US to photograph things that couldn’t be found at home in New England; and the next winter was spent compiling and writing. I ended up getting a one-month extension on the deadline for submitting the manuscript—I think from March 1 to April 1, 2009—and even so, I remember we were still working on finishing the introduction the day before the new deadline (including Noah running out into the woods behind his house to take the pictures of white pine trees that appear on pages vi and ix).

It was a good thing we had a firm deadline, because we could easily have spent the rest of our lives adding and revising. As it was, I did keep slipping new tidbits in for a while longer until the editor cut me off. I still remember the last thing I tried to slip in that didn’t make the cut; it was my discovery that a beetle named Chalepus walshii was responsible for those distinctive white rectangles I kept seeing on Canada bluejoint and other marsh grasses while conducting botanical fieldwork:


The answers to other riddles have remained elusive. In May 2011, I wrote here about a Bucculatrix cocoon Noah and I found in Tennessee that was curiously marked with little figure-eights.

As you can read in that post, I speculated that these markings might have been the work of a parasitoid wasp. This seems far-fetched to me, but I still don’t have a better explanation. Annette Braun never mentioned any such markings in her 1963 revision of the genus Bucculatrix, and I have never found another cocoon like this among the hundreds or thousands of Bucculatrix cocoons I’ve encountered over the past decade.

Or rather, I had never until October 10 of last year, when I went for a run down the road from my house and stopped to inspect a heart-leaved aster (Symphyotrichum cordifolium). John van der Linden had told me about a fly he found in Iowa that sneakily mines the petioles of heart-leaved asters, hardly leaving any discernible evidence besides its puparium; I wanted to see if I could find it in Massachusetts and rear adults to learn its identity. Curled in one of the petioles of the aster I stopped to examine was a Bucculatrix cocoon covered with little “8”s.


Were these “8”s inscribed in the surface or just a result of the Bucculatrix larva using two different colors of silk to spin its cocoon? I couldn’t tell for sure—the latter seemed almost plausible with this new cocoon, but looking back at the one from Tennessee, the “8”s are just too rounded and irregular; I think it has to have been done from outside. Naturally I collected the cocoon to see what would emerge.

Two weeks later I was doing some fieldwork in Northampton and I spotted two more cocoons decorated with little “8”s, this time attached to pine needles.


The same sprig of pine needles also had a similar cocoon without the “8”s:


Both the aster and the pine were directly below red oak trees, and I believe oak was the actual larval host of the Bucculatrix larvae that spun these cocoons. Many species of Bucculatrix start out life mining in oak leaves, then complete their development feeding on the surface of these leaves. When mature, they drop down on strands of silk and may wander some distance before finding a suitable place to spin a cocoon. There is a Bucculatrix cocoon just outside my second-floor bathroom window, 100 feet from any tree. Maybe it’s one of the aster-feeding species and didn’t have to walk that far, or maybe it blew there on a very windy day. The point is, one shouldn’t assume that the plant a cocoon is found on is the caterpillar’s actual food plant; my hope would be that the moth that emerges from the cocoon would be a species whose larval host is already known. If the “8”s indicate parasitism, though, this will make it very tricky to learn the moth’s exact identity—unless a moth emerges from that one unmarked cocoon and can be assumed to be the same species as the others.

Well, a few days ago this wingless wasp emerged from the cocoon on the aster petiole:


I believe it is an ichneumonid in the genus Gelis. Whereas many parasitoid wasps that emerge from their hosts’ cocoons arise from eggs that are laid when the host is still a larva or even an egg, Gelis species oviposit in their hosts’ cocoons. So I suppose if it’s really a wasp scribbling little “8”s on these Bucculatrix cocoons, Gelis would be a good candidate for the culprit. If the two cocoons on the pine needles have the same fate, that will be good supporting evidence. I’m still open to other explanations, but have none to offer at the moment.


This is the (now somewhat moldy) cocoon with the exit hole chewed by the emerging adult wasp. The “8”s are fainter now but still discernible.

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Introducing Grapholita thermopsidis

Almost every year since we bought our house, Julia and I have spent a few weeks on a road trip in search of exciting new leafminers. At this point we’ve visited nearly every US state and two Canadian provinces. This year, we’re looking forward to sticking around the homestead for the entire growing season for a change, but our past collecting trips continue to bear fruit. Today, for instance, a moth species we discovered in Colorado five years ago finally has a name.

In July 2015, our travels were centered around a visit to our friends in Colorado, Sally Waterhouse and Denny Radabaugh, who were Julia’s biology professors in college and who officiated at our wedding. This was the trip on which we found the rose leaf-mining sawfly Fenusa julia. Exactly one week earlier, we were poking around Sally and Denny’s yard and checking out the unusual (to us) plants in it when we spotted some mines on Thermopsis (Fabaceae), a plant known as goldenbanner or false lupine.


From above, the mines were visible as only slight discoloration, and they were only conspicuous because of the way they caused the leaflets to buckle.


The mines were perfectly distinct when viewed from below.


“Underside tentiform mines” like these are characteristic of certain moths in the family Gracillariidae, and on a legume like Thermopsis the expected genus would be Macrosaccus. However, upon collecting some of these mines, it soon became apparent that the larvae inside were expelling their frass:


A few gracillariids expel frass from their mines, but none of these are species that make tentiform mines, so this was something weird. At this point my leafminer book manuscript was far enough along that I knew the only documented leafminer on Thermopsis was Parectopa thermopsella (Gracillariidae), described in 1875 by V. T. Chambers, who collected the species in Colorado. No specimens of Parectopa thermopsella are known, but there is nothing in Chambers’ description of the leaf mine or the adult that indicates how it would be distinguished from the moth now known as Micrurapteryx occulta, which Annette Braun described in 1922 as Parectopa occulta and mines leaves of a variety of legumes. We found mines of “Parectopa thermopsella” during that trip to Colorado. They are flat, upper-surface mines, with a more or less lobed or “digitate” shape. Each lobe represents a different feeding excursion; in between bouts of feeding, the larva returns to a point along the midrib where it expels its frass through a hole in the lower epidermis.


In the lower-surface view above, you can see the initial linear track the larva makes on the lower leaf surface before switching to the upper surface to make a blotch mine. Unfortunately we were unable to rear any adults of “Parectopa thermopsella,” but they would look something like this Micrurapteryx we reared from a similar mine on Astragalus during the same trip to Colorado:


Anyway, ten days after we collected the mystery mines on Thermopsis, some larvae had abandoned their mines and were continuing to feed on the leaves externally, within shelters made by rolling or tying the leaves with silk.


A few days after that, parasitoid wasps started emerging from some of the mines. These little ~1 mm beauties are a female and male Zagrammosoma mirum (Eulophidae):


There were also a few braconid wasps, a little shy of 3 mm long:


So far, what José Fernández-Triana has been able to tell me about them is that they are in the genus Dolichogenidea. Their DNA barcode matches that of some specimens from Canada, but the species has not yet been identified (if it in fact has a name).

Fortunately, some of the moth larvae escaped parasitism and pupated in early August.


It was clear by now that this moth was not a gracillariid at all, but I wasn’t quite sure what it was. It wasn’t until the following spring that the adults finally emerged:


I knew they belonged to the family Tortricidae, which are commonly known as the “leafroller moths.” I sent them to Jason Dombroskie at Cornell University, who confirmed his impression from my photos that they belonged to the genus Grapholita.  He suspected they represented an undescribed species, but he didn’t have time to investigate further.

Last year, when I heard that moth aficionado Kyhl Austin had become Jason’s grad student and was focusing on Tortricidae, I seized the opportunity to ask Kyhl if he could take a look at these specimens. He was happy to, and within a few weeks he had confirmed that they belonged to a new species and described them in intricate detail. He also dissected and identified all of the other tortricids I’d sent to Cornell, several of which had not been reared before or were not known to start out life as leafminers. Today our paper* was published, in which the new species is named Grapholita thermopsidis and the natural history of six other leaf-mining tortricids is discussed.

At ~5 mm long (wingspan ~1 cm), this is the largest insect species I’ve had the pleasure of naming. It is the third moth species I’ve coauthored, and it occurs to me that each moth has taken five years after its initial discovery to actually get described (although with Marmara viburnella  what took so long was successfully rearing adults, and the species description was published the following year). Many more new moths await, so hopefully I can pick up the pace a bit!

* Eiseman, Charles S., Kyhl A. Austin, Julia A. Blyth, and Tracy S. Feldman. 2020. New records of leaf-mining Tortricidae (Lepidoptera) in North America, with the description of a new species of Grapholita. Zootaxa 4748(3): 514–530.

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Backyard Bowerbird

This morning I heard some bluebirds twittering outside the window, so Julia and I took the cue to go out and clean out the nest boxes. The bluebirds’ nest from last year was a dense mass of pine needles and other miscellaneous vegetation, including a single tendril from one of the nearby grapevines (visible below the center in the photo below).


The tree swallow nest on the other side of our front yard vegetable garden was lined with feathers, as is typical for this species. There were some fancy spotted ones that came from the guinea hens that live at a neighbor’s house down the hill and through the woods, several hundred feet away. The ones at the upper edge in the photo below look like barred owl feathers.


Last year was the first time we had a house wren use any of our boxes. House wrens are notorious for filling up multiple boxes with twigs and not necessarily using any of them to nest in. This was the case with both of our backyard boxes last year. Here’s the one from the northwest corner, which in the past has been occupied by chickadees:


When we opened the box at the southwest corner, we were surprised to find the twigs decorated with numerous spider egg sacs:


Now, when I’ve heard people talk about house wren nesting behavior before, I’ve gotten the impression that the wrens just fill up all the boxes because they’re jerks (or, say, trying to crowd out the competition), but this collection of little silken decorations reminded me of the male bowerbirds in Australia and New Guinea, which build elaborate structures in order to attract mates. I checked the Peterson guide to bird nests, and sure enough, it is the male house wren that arrives first and “builds dummy nests of twigs in all or most of available nest sites. Female may or may not accept prechosen site; may or may not accept male’s incomplete twig nest.” If she does accept the twig nest, she adds her own cup of finer material in which the eggs are laid. Nothing in there about the male adding decorations to the twigs, but clearly the one in our yard shared my appreciation for invertebrate architecture. The nest included at least thirteen egg sacs of the common house spider (Theridiidae: Parasteatoda tepidariorum); seven bright yellow egg sacs of pirate spiders (Mimetidae: Mimetus); four tufted egg sacs that I believe belong to cobweb spiders in the genus Euryopis (Theridiidae); four smooth, shining, disc-shaped egg sacs of a hunting (as opposed to web-spinning) spider (Gnaphosidae or maybe Corinnidae); four cocoon bundles of wasps in the genus Cotesia (Braconidae), some of which had a yellow tint suggestive of C. glomerata, a parasitoid of cabbage white caterpillars; three loose, white, spherical egg sacs that I believe belong to the cobweb spider Steatoda triangulosa; three big, fluffy egg sacs that I suspect are from some kind of orbweaver (Araneidae); two cocoons of a spider-parasitizing wasp in the Polysphincta genus group (Ichenumonidae); and one tent caterpillar cocoon (Lasiocampidae: Malacosoma).


Left to right: cocoon of a spider-parasitizing ichneumon wasp; egg sacs of pirate spider and common house spider.


Left, bundle of braconid wasp cocoons; center, egg sac probably of a cobweb spider in the genus Euryopis; lower right, more house spider egg sacs.


Two more pirate spider egg sacs; an egg sac of Steatoda triangularis (bottom right), and in the background, a tent caterpillar cocoon and egg sacs of three other spider species.


A clear view of the tent caterpillar cocoon.


Three hunting spider egg sacs (Gnaphosidae or maybe Corinnidae).

Shortly after we cleared out that box, a couple of chickadees came to check it out. Maybe they’ll beat the wren to it this year.


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Leafminers of North America, Season 2

Leafminer season never really ends, of course, but what I’m announcing in this post is that I’ve just sent out the first monthly installment of the fully revised and updated second edition of Leafminers of North America, my nearly 2000-page e-book about these fascinating insects that I’ve spent much of the past decade studying. Actually, I sent it out last weekend, but I got so wrapped up in working on finishing some papers about leaf-mining beetles that I didn’t get around to telling anyone except those who already have the first edition. So for the rest of you, here is a modified version of the message I sent out.


A white oak leaf with mines of three different leaf-mining moths. Clockwise from upper left: Stigmella (Nepticulidae), Cameraria, and Phyllonorycter (Gracillariidae).

It’s been two years since I sent out the first monthly installment of Leafminers of North America. As you may know, I just completed the first edition last June, but “complete” is a moving target when there is so much left to learn. In the past two years, at least 50 new species of North American leafminers have been described, and I now have notes in every single chapter of the book about text and photos to be added or modified. So I’m going to start cycling through the chapters again, and the updated introduction has already been completed. It includes some new illustrations and some added text, based on feedback from readers as well as my experience teaching a week-long leafminer workshop last summer. I’m going to try to stick to the same schedule as before to begin with, and I should be able to pick up the pace a bit once I get past the Lepidoptera and Diptera chapters (many new adult images to add there, along with taxonomic updates to be made). As I go through the chapters, I am also planning on compiling a spreadsheet of leafminer mysteries that need investigating, including fields for location and season. I will provide this as an Excel file, so that people interested in helping to solve these mysteries can reorganize it however they like.

As before, there are monthly and annual subscription options or you can just purchase the whole thing at once. Anyone who does the latter will receive the full first edition right away and the second edition as it becomes available, at no extra cost. See this page for details. And anyone who doesn’t really want a 2000-page book on leafminers but values the work I’m doing is welcome to make a contribution, either using the button at the bottom of that page or the “make a donation” button at the top of the right sidebar on this one. As you’ll see below, the scope of my bug-related pursuits doesn’t leave much space in my life for gainful employment.


The larva of a Periclista sawfly (Tenthredinidae) face to face with an adult of the leaf-mining beetle Brachys aerosus (Buprestidae) on a scrub oak leaf.

A few other updates while I’m at it: I have a paper in press documenting some new rearing records for (mostly non-leafmining) sawflies. I’m still excited to start working on a hostplant-based guide to North American sawfly larvae, but so far this winter I’ve been focusing my attention on finishing up a number of leafminer-related publications:

  • I coauthored the sections on Bucculatricidae, Gracillariidae, Heliozelidae, Nepticulidae, and Tischeriidae for the new checklist of North American Lepidoptera that will be published later this year
  • A paper on leaf-mining Tortricidae, including the description of a new species, is now in press
  • A paper on some lithocolletine Gracillariidae, including correcting the generic assignments of several species, was just resubmitted today
  • Today I also resubmitted a second paper on leaf-mining muscoid flies
  • A fourth paper on Agromyzidae is nearly ready to submit (describing thirteen new species, with new hosts etc. for a number of others)
  • A large paper on leaf-mining Chrysomelidae and a smaller paper on leaf-mining Buprestidae are nearly ready to submit
  • I’m halfway through writing a large paper on eulophid wasps reared from leafminers, but I plan to write a smaller paper on leaf-feeding weevils before I finish that
  • A number of publications on various moth groups are in progress
  • I recently completed a report on a leafminer survey of Black Rock Forest in New York (as mentioned in my last blog post), and will soon be writing reports on surveys conducted last year on Nantucket, in various midwestern prairies, and in Michigan’s Huron Mountains

If you’re an iNaturalist user, check out the Leafminers of North America project I created and please add your leaf mine observations there. The umbrella project that keeps track of that one along with several smaller regional leafminer projects now has over 6000 observations in it, and this has allowed me to glean a number of new host and distribution records as well as learn about some leaf mines that no one has documented before.

And finally, this summer I will be teaching several workshops in the northeastern US on invertebrate tracks & sign (including, but not limited to, leafminers), ranging in length from a few hours to a full week. For details, see my schedule.

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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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