A wasp has its day

If you’ve followed this blog for any length of time, you know that a lot of my attempts to rear insect larvae to adults end up producing parasitoid wasps. You have probably also heard me lament from time to time how hard it is to get a specialist to examine these wasps and tell me what they arenot because no one is interested, but because there are so many wasps in the world, and so few people that study them, that those who do are perpetually overwhelmed and need to focus on whatever little slice of the unending diversity they’ve chosen to focus on at the moment if they’re ever going to get anything done. I guess that goes for pretty much all entomologists, actually.

So it was a happy day in May 2016 when Dave Wagner directed Scott Shaw and Eduardo Shimbori to me for specimens they might use in their revision of the New World braconid wasps of the tribe Adeliini. These wasps specialize in parasitizing the larvae of tiny leaf-mining moths in the family Nepticulidae. The wasp larva waits until its host has finished feeding and spun its cocoon before finally devouring it; some time later, the adult wasp chews its way out of the moth cocoon. Although there are a few records of these wasps from other hosts, they seem to all be bogus records resulting from nepticulid larvae wandering into the leaf mines of other insects before spinning their cocoons.

I sent Scott and Eduardo the few Adelius specimens I had in my possession, and José Fernández-Triana kindly located some others that I knew were among the heaps of braconids I had deposited in the Canadian National Collection, and he passed them along. When Eduardo sent me the newly published paper* yesterday, I learned that the latter specimens were the most interesting ones: they constituted the entire type series of a new species, Adelius floridensis Shimbori & Shaw.

I reared the wasps from leaf mines of Fomoria hypericella that Julia and I collected on two different species of St. Johnswort at St. Sebastian River Preserve State Park in Florida back in March 2013. Here’s a larva of F. hypericella munching away in a leaf of roundpond St. Johnswort (Hypericum cistifolium):

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The finished mine of this species is more blotchy. Unlike virtually all other nepticulids, Fomoria hypericella spins its cocoon within the mine rather then popping out of the leaf and wandering for a bit before getting ready to pupate. The cocoon is visible in the lower right corner of the photo below (there is also a mine of another larva just getting started in the upper left corner):

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Before spinning its cocoon, the larva cuts a crescent-shaped exit slit in the leaf, just as larvae of other nepticulids do before exiting. This gives the pupa a place to pop out when the adult (which has no chewing mouthparts) is ready to emerge.

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The 2-mm adult moth may not have an exciting wing pattern, but you have to love the flashy orange pompom on its head.

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The wasp, Adelius floridensis, is a zippy little thing, just over 1.5 mm long (not counting the antennae), but I was able to get a few decent shots of one as it ran around:

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This female’s ovipositor is visible in the second photowell suited to poking through the leaf epidermis to insert its egg in a nepticulid larva.

There are now 19 described species of Adelius in the Americas (up from three), with just five of these known from the US and Canada. The other one I’ve met is A. coloradensis, which is widespread in the western US and seems to have an affinity for Stigmella species mining leaves of buckthorns (Rhamnaceae).

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This particular one was reared from an as yet unnamed species of Stigmella that mines leaves of cascara buckthorn (Frangula purshiana). Here are three of the moth cocoons, two with moth pupal exuviae poking out and one with an exit hole chewed by the wasp:

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Besides Adelius floridensis, the other newly described North American species is A. canadensis, which is known from just two specimens caught in fir-pine-aspen forest in Alberta in 1988. Adelius nigripectus is known from Indiana and Kansas and has been reared from an unidentified poplar leafminer. The final species, A. fasciipennis, was reared from Zimmermannia phleophaga in 1913. The moth was described at the same time and hasn’t been seen since. Its larvae mined in the inner bark of American chestnut and the species is believed to have gone extinct (perhaps along with its parasitoid) after the chestnut blight made larger chestnut trees scarce. It would be a tricky moth to find if it is still around though; there is no external evidence of the mine until the larva chews a tiny exit hole in the bark and drops to the ground. This was reported to happen between April and June, so if there are American chestnuts near you, it’s worth keeping an eye out, starting right about now!

* Shimbori, Eduardo M., Marco A. Bortoni, Scott R. Shaw, Carolina Da S. Souza-Gessner, Paula De C. M. Cerântola, and Angélica M. Penteado-Dias. 2019. Revision of the New World genera Adelius Haliday and Paradelius de Saeger (Hymenoptera: Braconidae: Cheloninae: Adeliini). Zootaxa 4571(2): 151–200.

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Origami Weevils

I always get excited when I encounter the work of leaf-rolling weevils (Attelabidae), even though they are by no means uncommon. I just find it fascinating that these insects have learned to fold leaves into neat little cylindrical packets for their larvae to live inside, without the use of silk or any other adhesive. The female starts by making two cuts in a leaf that meet at the midrib, as shown here:

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She then folds, rolls, and tucks the severed portion of the leaf, laying a single egg inside this packet, with the end result looking like this:

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The species shown above is Synolabus nigripes, which apparently feeds only on sumac (in this case winged sumac, Rhus copallinum). I’ve only encountered it once or twice; what I usually see are rolls on oak (or occasionally chestnut) made by Attelabus bipustulatus (shown here) or perhaps other species.

Last August when I was teaching my Tracks & Sign of Insects… workshop at the Eagle Hill Institute in Maine, we came across some weevil leaf rolls on speckled alder (Alnus incana), which was a first for me.

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We took a couple of them back to the lab to examine, and one of the participants unraveled one to reveal the egg inside:

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I kept the other one in a vial to see if an adult would eventually emerge. Eight months later—yesterday—it did!

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I opened the roll up to see what had been going on in there all this time, and not surprisingly, beneath the outer layer of leaf, everything inside had been converted to weevil poop.

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It wasn’t just a disorganized wad of poop though. When it had had its fill of the leaf tissue, the larva had packed all of its frass against the remaining outer leaf layer, creating a smooth-walled, roomy cell in which to pupate.

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At one end of the cell, there were a few differently textured, yellowish fecal pellets, which I assume were meconium deposited by the newly emerged adult weevil.

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Naturally, before taking the above photos I finished my portrait session with the weevil.

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It turns out this is a species (Himatolabus pubescens) that I had met once before, six years earlier and 3000 miles away, nibbling on an oak leaf at Madera Canyon. According to Art Evans’ Beetles of Eastern North America, adults of this species have been found on hazelnut (Corylus) and rosemallow (Hibiscus) as well as alder and oak.

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Some more new flies

Nope, attention-grabbing titles just aren’t my forte.

Anyway, you may recall that last fall Owen Lonsdale and I published a paper describing 30 new species of agromyzid flies, two of which I found right along my driveway, and the rest of which I summarized here. That paper also gave new information on nearly 100 other species that already had names, gleaned from my first five years of rearing leaf-mining insects. During those five years, several other people around the US got inspired to start collecting leafminers too, and ended up contributing a number of specimens that were used in that paper. The #1 contributor was Tracy Feldman of North Carolina, and in recognition of his efforts, I set aside all of his rearing records of species that I hadn’t reared myself in other states (with the exception of Cerodontha feldmani) to be included in a separate paper dedicated to North Carolina Agromyzidae. That paper has just been published*, and in it Owen, Tracy, and I describe another nine new species.

One of these was a species Tracy first found in the spring of 2015 on a plant I’d never heard of, Carphephorus bellidifolius (Asteraceae), which apparently has the common name of “sandywoods chaffhead.” The leaf mines were partially on the upper surface…

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…but the larvae often did most of their feeding on the lower surface, switching sides when they were almost mature. This is the underside of the same leaf that is shown above:

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Naturally, all the mines Tracy collected that year turned out to be parasitized by eulophid wasps, so all we got were these:

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He tried again the following May, with the same result. But then in late June, he found some more larvae, and these survived to form puparia…

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…and emerged as adult flies within a few weeks:

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When Owen examined the specimens, he informed us that this was a new species, so we named it Liriomyza carphephori—I always like to name species after their host plants if that name isn’t already taken.

Tracy and I were both surprised when Owen told us that the fly Tracy found mining leaves of Spanish needles (Bidens bipinnata) was the same species. Not only was the plant not very closely related (belonging to a different tribe of the aster family), but the mine seemed very different: on Bidens it was entirely on the upper surface and always started at the tip of the leaf, where it made a very contorted track before eventually meandering elsewhere in the blade.

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I had collected similar mines in Vermont and in my front yard in Massachusetts on Bidens frondosa (“devil’s beggar-ticks”) in October 2016. The larvae were mining in leaves that had already turned completely red.

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The adults that emerged the following spring proved to be the same species, so L. carphephori ended up being the one species in the North Carolina paper with paratypes from other states. The paratypes also included some older specimens of mine that we had mentioned in the first paper but hadn’t been able to put a name on yet because I had only reared females. One of these females had been reared from Bidens frondosa back in July 2014, and we discussed it as “Liriomyza sp. 3.” Two others had been discussed as “Liriomyza sp. 5,” and they were reared from Mikania scandens (climbing hempvine).

Mikania scandens, as it happens, is in the same tribe as Carphephorus (Eupatorieae) but has pointed leaves like Bidens. Interestingly enough, on Mikania some of the mines look like the ones on Bidens

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…and some are like the ones on Carphephorus: switching between the two leaf surfaces and not crammed into the tip of the leaf.

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When I found these mines while doing botanical fieldwork back in August 2013, I was standing knee-deep in water. Some of them were already empty, and the larvae had glued their puparia to the undersides of the leavesa useful adaptation to this habitat since they probably would have drowned if they had dropped from the leaves as Liriomyza larvae normally do.

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Incidentally, one of those two females from Mikania emerged within a few weeks and the other not until the following spring. It’s always hard to know when to give up and throw away old puparia, and every once in a while I’m rewarded for hanging onto them until the next year.

One final note on Liriomyza carphephori: I’ve found similar leaf mines in Ohio on white snakeroot (Ageratina altissima), another member of the tribe Eupatorieae, and I predict these larvae will turn out to be the same species.

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* Eiseman, Charles S., Owen Lonsdale, and Tracy S. Feldman. 2019. Nine new species of Agromyzidae from North Carolina, USA, with new host and distribution records for additional species. Zootaxa 4571(3): 301–333.

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Parasite of a parasite of a parasite

I’m slowly making my way through the photos I took last summer, and I just got to the conclusion of an interesting series that started with this leaf I was given on June 22:

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The leaf is from hairy honeysuckle (Lonicera hirsuta), a species that is listed as Endangered here in Massachusetts. I was given the leaf by a botanist at the state’s Natural Heritage & Endangered Species Program with the hope that I could determine what the leafminer was. No leafminer has been reared from this particular honeysuckle species, but based on what miners are known from other honeysuckles, I can say that the larva was a fly in the genus Aulagromyza (Agromyzidae). My best guess is that it is none of the six described Aulagromyza species that are known to feed on honeysuckle in North America. Long, linear, greenish mines have been reported from a similar plant, limber honeysuckle (L. dioica), in Ontario, and these were tentatively associated with an undescribed Aulagromyza species that is known from a single female reared from “Lonicera columbiana” (a plant that doesn’t exist) in Montana. So I’m thinking it’s either that or… something else.

But that’s all beside the point. The point is, a healthy mature Aulagromyza larva would pop out of its leaf and form a puparium like this…

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…from which an adult fly would emerge:

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(This fly is Aulagromyza luteoscutellata, which makes much shorter, wider mines that are common on introduced Asian bush honeysuckles.)

But if you look at the leaf mine photo above, you’ll see that there are a couple of objects toward the end of the mine. Here’s a backlit detail of the relevant portion:

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The object at right is the remains of the fly larva, which was devoured by a parasitoid braconid wasp in the genus Colastes (or a related genus in the subfamily Exothecinae). We know this because the object at left is the wasp’s cocoon.

Normally, the braconid larva would pupate within the cocoon and then the adult would chew a hole at one end and emerge:

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On July 5, an adult wasp did chew its way out of the cocoon in the hairy honeysuckle leaf…

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…but it was a eulophid wasp, which as a larva had devoured the braconid larva within its cocoon.

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I probably rear a lot more hyperparasitoids than I realize, so it’s always nice to see a clear case like this.

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In Search of the Lost Sawfly

Greetings to everyone who subscribed to this blog after my recent post about the inhabitants of a clump of sod in my front lawn! This one will take us a little farther afield. As most of you know, I’ve been working on a guide to North American leafminers for the past seven years or so, and in order to make its coverage as complete as possible, I periodically go on road trips with my wife Julia in search of species that can’t be found in our own yard.

Insect species known or presumed to mine leaves in North America include around 1200 moths, 800 flies, 200 beetles, and 40 sawflies. There are numerous species in the first three groups that are known only from caught adults, so no one knows what types of mines they make and on what plants. The situation is better with the sawflies: when I started this project, the hosts were known for all but two species in the main leaf-mining group, the tribe Fenusini. One of these was Metallus ochreus, which Dave Smith had described in 1988 from adults caught in Virginia and Maryland. This turned out to be a common species feeding on dewberry here in New England, including in my own backyard, as discussed here. The remaining enigma was Prolatus artus, which Dave had described back in 1967 from a single female he had caught in southwestern Oregon in 1962.

When Julia and I took our first trip to the western US in the fall of 2012, we found sawfly larvae mining huckleberry leaves in western Washington. I thought maybe we had discovered the host of Prolatus artus, but no, it turned out to be a new species, which Dave and I named Scolioneura vaccinii.

When Julia and I took our second trip to the western US in the summer of 2015, we didn’t go as far as the Pacific Coast states, but in Colorado Julia spotted some sawfly larvae mining leaves of a wild rose. I thought maybe these could be Prolatus artus, but no, they turned out to be just another new species, which Dave and I named Fenusa julia.

In the late winter of 2017 we took a short trip to the Southwest, and we met up with our friend Eric LoPresti outside of Phoenix, Arizona. Here we found some larvae mining leaves of a plant called “blue dicks” (Asparagaceae: Dichelostemma capitatum).

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To my great surprise, these turned out to be sawfly larvae. There were no previous records of any sawfly mining leaves of a monocot (unless you count Eutomostethus luteiventris, which bores in stems of rushes and sometimes extends its tunnels into the bracts). For all I knew this could be a weird species in some group of sawflies whose larvae normally do something other than mine leaves.

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My #1 priority on this trip was to follow up on a discovery Eric had made the previous year in southern California, of sawfly larvae mining leaves of a plant called fiddleneck (Boraginaceae: Amsinckia intermedia).

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I suspect these larvae belong to the genus Kerita, which is in the smaller, lesser known tribe of leaf-mining sawflies (Pseudodineurini). The eastern species K. fidala feeds on another boraginaceous plant, Virginia bluebells (Mertensia virginica), and there are two western species whose hosts are unknown. When we got to California we found the larvae in great abundance everywhere we encountered fiddleneck.

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Unlike the Fenusini, which have to complete their development in whatever leaf they start out in, larvae of Pseudodineurini are able to establish new mines in fresh leaves. So we were able to supply the larvae we’d collected with a constant supply of fresh fiddleneck leaves, and a few dozen reached maturity, burrowing into the jars of soil I provided them to wait until it was time to pupate and then emerge as adults.

Disappointingly, nothing ever came of any of those fiddleneck larvae. A year later I sifted through the soil in one of the jars and could find no evidence that any insects had ever been there. However, two of the nine larvae from the blue dicks leaves did emerge as adults ten months after we collected them.

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I passed them along to Dave Smith, and I can only imagine his surprise when he matched them to a species he had collected just once, a thousand miles away and 55 years earlier: Prolatus artus. Our short paper documenting this was just published:

Eiseman, Charles S. and David R. Smith. 2019. Prolatus artus Smith (Hymenoptera: Tenthredinidae), a leafminer of Dichelostemma capitatum (Benth.) Alph. Wood (Asparagaceae). Proceedings of the Entomological Society of Washington 121(1):115-118.

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Life in a Cubic Foot of My Lawn

Late last March, I noticed some neatly cut goldenrod stems in my lawn. Or I should say, what used to be my lawn; five years earlier, it was a “proper” lawn (shown here), but it is now more of a meadow, interspersed with young fruit trees, berry bushes, strawberry and asparagus patches, vegetable gardens, and so forth. Anyway, you can see three of the cut stems in this photo:

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And a closer look at one here:

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I felt like I should know who was leaving this distinctive sign, but I was drawing a blank, and I didn’t find the answer in my book. So I split one of the stems open and found it filled with frass, with a tunnel leading to the base and into the roots.

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Since the tunnel was plugged with frass at the top of what remained of the stem, I guessed that the larva (or maybe pupa) of the insect responsible was overwintering in the roots. So toward the end of April, I dug up a chunk of my lawn including a few of these cut stems and put it in a clear garbage bag in the front hallway, where I could watch to see what emerged.

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The above photo was taken about a month later, the day the bugs in question finally started to emerge. But a whole bunch of things appeared in the bag before then:

A couple of young sac spiders (Clubionidae: Clubiona)…

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…a case-bearing moth larva (Coleophoridae)presumably a species of the Coleophora duplicis complex. I often see these larvae in the fall, feeding in goldenrod seedheads while wearing portable cases cleverly disguised with pieces of the flowers. If I’m right that this is one of those, the flower bits have mostly worn away, leaving the bare silken core of the case…

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…a nymph of a leafhopper in the genus Agallia (Cicadellidae)…

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…a sheet-web weaver (Linyphiidae: Linyphiinae). This is the group that includes the “filmy dome spider” and “bowl and doily spider”, but I’m not sure exactly who this one is…

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Zenodosus sanguineus (Cleridae), a “checkered beetle” that is normally found under bark, where it feeds on bark beetles…

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…lots of these fancy little humpbacked springtails (Entomobryidae: Lepidocyrtus paradoxus), which are a common sight in my yard…

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…also a good number of these woodlice: Philoscia muscorum (Philosciidae), a species introduced from Europe…

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…a handsomely antlered little parasitoid wasp (Eulophidae: Eulophini), possibly one of the species I regularly rear from leafminers…

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…several of these weevils in the genus Tychius (Curculionidae), presumably one of the European species that feeds on clover seeds…

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…a clear-winged little delphacid planthopper…

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…a so-called “pleasing fungus beetle,” Cryptophilus obliteratus (Erotylidae)—according to BugGuide this is an Asian species that has been in Europe since 1982 and hasn’t yet been officially documented in North America (guess I should have saved the specimen; oh well)…

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…a tiny tube-tailed thrips (Phlaeothripidae)…

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…the strawberry bud weevil (Curculionidae: Anthonomus signatus), or a close relative…

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…a great many dark-winged fungus gnats (Sciaridae), probably of several species, but I’ll just show two examples here…

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…a couple of these platygastrid wasps, which are parasitoids of gall midges…

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…at least three of these little red cobweb spiders (Theridiidae: Thymoites unimaculatus)…

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…a dance fly (Empididae)…

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…an agromyzid fly in the genus Melanagromyza—a stem or root borer or possibly a flower feeder, very likely from the goldenrod. Given the probable host association, it would have been worth saving this one to be identified to species, had it not been an unidentifiable female…

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…the wee six-legged larva of a whirligig mite (Anystidae)…

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…plenty of “cornfield ants” (Formicidae: Lasius neoniger)…

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…some adult Agallia leafhoppers—maybe the same species as the nymph that appeared earlier…

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…a tachinid fly in the genus Graphogaster—a parasitoid of what most people call “microlepidoptera” but which to me are medium-sized moths (Depressariidae, Gelechiidae, Scythrididae, Tortricidae)…

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…several shiny green soldier flies (Stratiomyidae: Beridinae)…

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…the stilt bug Berytinus minor (Berytidae), a clover-sucking bug from Europe…

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…some manner of moth caterpillar—your guess is as good as mine…

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…a dark-banded owlet (Erebidae: Phalaenophana pyramusalis) that lost some of its scales in the bag—this is one of the “litter moths,” whose caterpillars feed on dead leaves…

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…several of these Asian broad-nosed weevils (Curculionidae: Myosides seriehispidus)…

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Asagena americana (Theridiidae), another cobweb spider…

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…a good old-fashioned non-Lyme-disease-carrying dog tick (Ixodidae: Dermacentor variabilis)—one of our chickens eagerly eats these out of my hand, and that was the fate of this one…

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Tetramesa (Eurytomidae), a wasp that forms galls in grass stems…

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…a figitid wasp, which Matt Buffington tells me is in the tribe Ganaspini (Eucoilinae)—either Ganaspis (a parasitoid of fruit flies, Drosophilidae) or Hexacola (a parasitoid of shore flies, Ephydridae)…

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…a nymph of a leafhopper in the genus Aphrodes—probably a grass feeder…

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…a sweet little jumping spider (Salticidae)…

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…a wolf spider in the genus Trochosa (Lycosidae)…

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…and the rust fly Psila collaris (Psilidae), who I long ago photographed slurping bird poop.

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Now, if you’ve stuck with me through that whole month’s worth of incidental bugs, you, like me, get to be rewarded with a look at what was responsible for those neatly cut goldenrod stems that prompted me to dig up that little chunk of my lawn in the first place:

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A plume moth! Three of them emerged, in fact—one for each of the cut stems. I consulted the plume moth specialist, Debbie Matthews, and she told me they belonged to one of two Hellinsia species (Pterophoridae). Examination of genitalia is needed to distinguish the adults, but she told me that if I had a male of H. glenni I could recognize it without dissection by teasing out an appendage from near the tip of its abdomen and seeing that it had a goose-head shape at the end.

And lo, tucked in there among its fur was the goose head!

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The other species in question, Hellinsia kellicottii, is also a goldenrod borer. Does its larva produce these same neatly cut stumps? I’m not sure. The answer may or may not be in the literature somewhere, but for now I’m happy to be able to say approximately who is responsible when I encounter this distinctive feeding sign.

Edit: Oh yeah, I now remember that I asked Debbie that question, and she said: “Not always. You can look for piles of frass coming out of a hole about an inch from ground level in the fall. Probably not easy to find after overwintering.” Which I guess means, yes, sometimes. For what it’s worth, John van der Linden investigated the same stumps in Iowa the previous spring and also came up with H. glenni. I had seen his post on BugGuide six months before I noticed the stumps in my own yard, and that was what I was not quite managing to remember when I had the feeling that I should already know the answer to this riddle.

 

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How Many New Species?

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) 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 is 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.

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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.

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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.

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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.

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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.

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So that’s six in the first category. I 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. For the 43 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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.”

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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).

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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.

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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.

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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:

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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).

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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).

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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.”

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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.

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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…

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