Adapting to New Menu Options, Part 2

One of the reasons certain nonnative plants come to dominate the landscape is that they are released from the specialized insects and pathogens that keep them in check in their native range. So I’m always interested to see what insects are able to make use of these plants. I’ve often noticed little nibbles like this on Japanese barberry (Berberis thunbergii)…

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…but had never actually spotted anything feeding on the leaves until one autumn day when I stopped to investigate some particularly fresh-looking nibbles. This “window feeding” (leaving one epidermis intact) turns out to be the work of early instars of an inchworm, which starts eating all the way through the leaves as it grows larger.

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The above photo was taken on September 28. Here is the same caterpillar on October 2…

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…and October 8:

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The caterpillar burrowed into soil not long after this, emerging as an adult in March after I took its jar out of the fridge.

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Its patterned wings became good camouflage when it came to rest on the fallen white pine in my yard:

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This native moth is known as the barberry geometer (Geometridae: Coryphista meadii), and barberries are its only known host plants. Its original host is presumed to be American barberry (Berberis canadensis), but this plant does not occur in the Northeast. So like Stigmella rhamnicola (but even more dramatically), this moth has expanded its range because of the introduction of a nonnative plant. Sorry to say the little nibbles made by the caterpillars aren’t enough to keep the barberry from taking over places where the deer are selectively eating all the other understory plants.

Last October I discovered that barberry geometers aren’t the only caterpillars feeding on leaves of Japanese barberry. Whereas the inchworms “window feed” from the lower leaf surface, I found these two caterpillars grazing in little patches on the upper leaf surfaces:

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These are the same two caterpillars a week later:

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Five days later, they looked much more similar to one another:

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Unfortunately I was unable to rear these to adults. I have found similar caterpillars feeding on Christmas fern (Polystichum acrostichoides) and wintergreen (Gaultheria procumbens) in the fall, so I think this is just a species that feeds on whatever is still green at the end of the year rather than any kind of barberry specialist. It’s some sort of owlet moth (Noctuidae); I had one of the wintergreen feeders emerge in June, but I haven’t figured out what it is yet and I’m nowhere near there in my photo sorting, so stay tuned…

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Moths From A Willow Leaf

Over the past few days, a break in fieldwork has given me a chance to start catching up on going through my photos from this year—I’m exactly five months behind at the moment. On March 19 I finally got to the end of a story that started with the Berkshire BioBlitz on June 19 of  the previous year. I had collected this leaf of silky willow (Salix sericea) in the hope of finding out which of the nine willow-feeding species of Phyllonorycter (Gracillariidae) was responsible for the leaf mine on it, since as far as I know there is no way to distinguish among the mines of these species (though a few can be ruled out based on geography).

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In my hurry to document as many species as possible in a short time, I didn’t investigate what had made the webbing and leaf fold next  to the mine. This nearly proved disastrous, since it turned out a caterpillar was hiding in the leaf flap, and by the next day it had consumed much of the leaf. Fortunately, it didn’t eat too much of the mined portion.

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I moved the caterpillar to a separate vial with fresh willow leaves. I don’t usually collect non-micro moth caterpillars to rear, but when they show up on leaves I’ve already collected I figure I might as well.

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On June 28, the adult leafminer emerged, revealing itself to be Phyllonorycter salicifoliella, and leaving its pupal skin protruding from the mine.

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The externally feeding caterpillar developed much more slowly, and I had to keep collecting new willow leaves to feed it. Here it is on July 6 with an old willow leaf that has gone yellow:

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On July 29 it was looking pretty different:

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The last photo I took of it was on August 4…

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…but I noted in my journal that it turned pink and burrowed into soil on August 12. As it happens, I had tried to rear this same species eleven years earlier (from a caterpillar feeding on black cherry), so I have a photo to illustrate just how pink it was when ready to pupate:

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Nothing stirred in the jar for the rest of the year. I put it in the fridge over the winter, taking it back out on March 1. Almost three weeks later, an adult confused woodgrain (Noctuidae: Morrisonia confusa) appeared in the jar. Having no further use for it, I let it out to see if it could find others of its kind. Here it is resting on a fallen white pine at the edge of my yard:

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Giving Wasps Their Due

I often see pie charts like this one suggesting that about a quarter of all insect species are beetles. Suspiciously, other sources (e.g. here) say beetles represent about a quarter of all animal species, and Wikipedia goes so far as to say beetles constitute “almost 25% of all known life-forms”! (they may have meant to say “animal life forms”, but a pie chart here does show beetles representing about 20% of all organisms). The first statement is probably closest to the truth, but I suspect that it is based on numbers of described (named) species, and I can’t help but wonder if this figure is a result of a disproportionate number of people studying beetles. I’ve lost track of how many undescribed moths, flies, and wasps I’ve found in just the past few years; the moths and flies belong to groups that are studied by just one or two people in North America, and the situation is even worse for parasitoid wasps and gall wasps: for most of the wasps I’ve reared, I can’t even find a specialist to examine them and determine whether they match any described species.

According to Wikipedia, there are about 400,000 described beetle (Coleoptera) species in the world, with estimates of total species ranging from 850,000 to 4 million. There are about 180,000 described moths and butterflies (Lepidoptera), over 150,000 wasps, ants, bees, and sawflies (Hymenoptera), and about 150,000 flies (Diptera). I haven’t seen an estimated total number for Lepidoptera, but there are said to be one to three million Hymenoptera species and about a million Diptera species.

It seems like every insect has a set of more or less host-specific wasps that parasitize it. Even parasitoid wasps have wasps that parasitize them. So it’s hard for me to believe that there aren’t more wasp species than anything else. I looked around for estimated numbers for the major parasitoid groups, and found that the family Braconidae has roughly 12,000 described species, with 50,000 being a “probably highly conservative” estimate of the total number in the world. The related family  Ichneumonidae has 24,281 described species  and “probably includes more than 100,000 species” total. According to BugGuide.net, in North America it is estimated that there are 3000 species in this family alone remaining to be discovered and/or named, the same as the estimate for the entire beetle order (there are currently about 25,000 named beetle species and 5000 ichneumonid species in North America). The superfamily Chalcidoidea has about 22,000 described species, with an estimated total of more than 500,000 according to BugGuide.

With a thousand or so described species, the subfamily Platygastrinae (Platygastridae) is a relatively small group of wasps. (The wasps themselves are small, at a millimeter or so long.) As far as is known, all of them parasitize gall midges (Cecidomyiidae). As of 2014, there were 6203 known species of gall midges, with the number of species still unknown being “inestimable”.* It seems to me that the true number of platygastrines is therefore likewise inestimable, and I suspect that Peter Neerup Buhl’s report on the eight sets of platygastrid specimens I sent him last year is representative of the current state of knowledge of this group. As I mentioned here, he identified one of the wasps as Metaclisis floridana and another as a Synopeasnot identifiable to species because it was a male, but possibly S. pubescens; neither M. floridana nor S. pubescens had ever been reared before. The remaining six were all species of Platygaster, a genus with about a hundred known species in North America. Of these, one keyed to P. baccharicola but its antennae were broken off and it couldn’t be identified with certainty; two were males and therefore unidentifiable; and the remaining three didn’t match any known species. Our paper describing them was published last month**, so I can now introduce them to you:

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Platygaster pruni is a parasitoid of Contarinia cerasiserotinae, which forms galls on black cherry (Prunus serotina) that look like this:

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The larvae of Contarinia cerasiserotinae pop out of their galls in late May, burrow into the ground, and emerge as adults the following spring.

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However, in my one attempt to rear this species (from galls collected in Leverett, Massachusetts), all that emerged the following spring were a male and female of Platygaster pruni and this as yet unidentified eulophid:

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The other two new species were reared from gall midges that are themselves likely undescribed. The host of the first has never been reared (adults are unknown), but its larvae are common in western Massachusetts. They form leaf spot galls on wild oats (Uvularia sessilifolia), which are sometimes already empty by late May…

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…but sometimes larvae are present as late as early July. The whitish larva is only visible on the lower leaf surface.

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The larvae of this species likewise exit their galls and burrow into the ground, presumably emerging as adults the following spring.

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The four females and two males of Platygaster uvulariae all emerged on May 18.

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The last species came from the same site on Nantucket where I found Megaselia nantucketensis and is likewise known from a single specimen. Its host midge develops in little blisters on grape leaves; the blisters project from both leaf surfaces and have a little central tuft on each side.

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The larvae exit their galls in the first half of August, once again burrowing into the ground and overwintering.

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I did manage to rear two adults of this species, but they were in poor shape when I discovered them on May 15.

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Ray Gagné confirmed that they belong to the genus Vitisiella, and probably a new species, but the genus needs to be revised before that can be said with certainty. A third midge emerged on June 5 and was in good shape when I found it, but it turned out to be an inquiline (not the gallmaker), Lasioptera vitis.

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The single female of Platygaster vitisiellae emerged the same day.

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Hard to estimate how many Platygaster species there might be when every specimen I rear is either unidentifiable or a new species!

* Shortly after I posted this, Jason Dombroskie alerted me to this brand new paper (published just three days ago), which used DNA barcoding to estimate insect diversity in Canada. The estimate for gall midges is 16,000 out of 94,000 species, which they extrapolate to 1.8 million cecidomyiid species out of ten million insect species worldwide. Addressing the very point I was making at the beginning of this post, they state that if their estimates are accurate, “the global species count for this fly family may exceed the combined total for all 142 beetle families.”

** Buhl, Peter N. and Charles S. Eiseman. 2016. Three new reared species of Platygaster (Hymenoptera, Platygastroidea: Platygastridae) from the United States. International Journal of Environmental Studies, DOI: 10.1080/00207233.2016.120132

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Introducing Brachys howdeni

The state flower of Massachusetts is mayflower, which no longer seems an appropriate name since it blooms in April these days (I even saw some flowers in March this year). Another name for this plant is trailing arbutus, and I’ve always liked its Latin name, Epigaea repens (Ericaceae), because both the genus and species describe it well: Epigaea means “on the earth,” and repens means “creeping.” Its evergreen leaves form mats on the ground in dry, sunny areas, and its fragrant, white to pink flowers are visited by all sorts of insects in the spring.

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Mining bee (Andrenidae: Andrena)

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Greater bee fly (Bombyliidae: Bombylius major)

On January 7, 2012, I went for a walk with some friends on a ridge that happens to be just up the hill from where Julia and I now live. That was the winter that I started working on my leafminer book, and a year earlier I would probably have passed off the brown edging on this trailing arbutus leaf as something unrelated to insect feeding:

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However, since I had leafminers on the brain, I picked the leaf and held it up to the sky, and sure enough, I could see frass and a larva inside (the larva is at the lower left in the photo below).

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I was far enough along in my book project that I was pretty sure no insect was known to mine Epigaea leaves, so I kept an eye out for more of these mines over the next few months and collected them from several different locations.

At the same time, I was systematically going through all the known groups of leaf-mining insects and tracking down literature on their natural history. One day I went to the UMass science library to peruse (among other things) some papers by Henry Hespenheide on leaf-mining beetles*. I think that was the first time I had come across his name, so that evening I was surprised to see it pop up in that annoying “ticker” column on my Facebook newsfeed, which I normally ignore. He was discussing Philly cheesesteaks with a mutual acquaintance. I suppose my eye was conditioned to spot a newly familiar name in my peripheral vision, in the same way I was beginning to spot leaf mines where I never would have noticed them before. Anyway, I was glad to know how to reach Henry if any beetle questions should arise.

On April 16, the first adult emerged from one of the trailing arbutus leaf mines, and it was a buprestid (jewel beetle) in the genus Brachys.

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Brachys species are mostly associated with oaks, although some have been reared from leaf mines on various other trees. Since leaf-mining buprestids are among Henry’s specialties, I showed him this photo to see what he thought of it. He replied: “I know about this species and have a couple of specimens that Henry Howden reared decades ago. I would be happy to describe it if you have a nice series, say 5-10 specimens. . . Brachys is a VERY difficult genus with many cryptic species, but this one seems distinct enough to describe without revising the genus.”

Since then, I’ve provided Henry with around 20 specimens reared from trailing arbutus. I also collected some aborted mines and reared two different parasitoid wasps (Eulophidae) from them: three females of the purple-headed Neochrysocharis diastatae

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…and a tiny male of the genus Pnigalio, which is itself in serious need of revision:

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Over the past few years Henry has studied my Brachys adults from trailing arbutus and various other hosts, plus countless specimens others have reared or collected all over North America. Last year he described three new species from New Mexico, Texas, and Mexico**, and last month our paper describing the trailing arbutus miner was published***. Henry named it Brachys howdeni, since Henry Howden was the first to rear this species. It is very similar to B. aerosus, and the number of other species hiding under that name is yet to be determined. Stay tuned…

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* Hespenheide, Henry A. 1992. A review of the genus Tachygonus (Coleoptera: Curculionidae) north of Mexico. Proceedings of the Entomological Society of Washington 94(1):1-11.

* Hespenheide, Henry A. 2003. A reconsideration of Pachyschelus schwarzi Kerremans and a review of American Pachyschelus north of México (Coleoptera: Buprestidae). The Coleopterists Bulletin 57(4):459-468.

** Hespenheide, Henry A. 2015. Striking new species of Brachys Dejean, 1833 (Coleoptera: Buprestidae) from New Mexico, Texas, and Mexico. The Coleopterists Bulletin 69(2):221-224.

*** Hespenheide, Henry A. and Charles S. Eiseman. 2016. A new species of Brachys Dejean, 1833 (Coleoptera: Buprestidae) from the eastern United States using an unusual host. The Coleopterists Bulletin 70(2):335-340.

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Life and Death on Mt. Greylock

This past weekend I attended my third Berkshire BioBlitz. At my first one in 2011, I barely left the parking lot at the summit of Mt. Greylock and photographed 166 different species of insects and arachnids. This time, Julia and I actually made it into the woods, taking about four hours to complete the 0.7-mile Rounds Rock trail loop. Our list this time was not as impressive, mainly because we kept encountering things that were more interesting than just tallying as many species as we could. One of the first things Julia spotted was a leafminer that I’m confident is new to science, so we had to spend a while dealing with that. Not long after we resumed our slow walk, she flipped over an oak leaf to get a better look at a sawfly larva that was munching the edge, and she called me over to see an ichneumon wasp that was stalking up to it on the underside of the leaf. The wasp had almost reached the larva when I got there:

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The larva continued to munch away obliviously as the wasp inserted her ovipositor just behind its head.

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When she was done, she backed away but continued to watch the larva for a while, approaching it again and closely inspecting it, then wandered off.

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You may have noticed the yellow object stuck to the larva’s thorax.

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My best guess is that this is the egg of another ichneumon wasp in the subfamily Tryphoninae. These wasps have stalked eggs that they attach externally to their host larvae. There is a photo of an egg on a preserved specimen here; I’ve never seen a photo of a fresh one. With larvae of two different ichneumon wasps feeding on its innards, it’s probably safe to say this sawfly isn’t going to make it to adulthood.

We plodded along for about 40 minutes until Julia spotted something else that compelled me to spend 15 minutes sitting and photographing it: a treehopper emerging from its last nymphal skin. I got to it just as it was getting its abdomen free, but was able to watch the lengthening of its wings and pronotum.

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We didn’t hang around to see it become fully colored. There were a few adult treehoppers like the one below in the immediate vicinity, but the molting one seemed to be a different species.

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We had made it just a few steps down the trail when a colony of aphids caught my eye.

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There are always other insects to be found in association with aphids, and before long I noticed this syrphid fly larva resting on a stem:

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Then I turned around and saw a wasp with its ovipositor stuck in another syrphid larva on the underside of a leaf.

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As with the icheumonid parasitizing the sawfly larva, this wasp was poking the syrphid larva’s “neck.” I’m thinking the wasp is a figitid, maybe in the subfamily Aspiceratinae. Both of the observations of this rarely seen wasp group on BugGuide show females ovipositing in syrphid larvae. This one wandered off for a bit, then returned to the larva.

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She climbed onto the larva again, this time ovipositing into its abdomen.

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Meanwhile, Julia saw the first syrphid larva grab an aphid, and I turned around to see it holding its prey out from the stem, devouring it.

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I’m not sure what that yellow blob is on the aphid’s antenna. I initially thought it was a mite, but it doesn’t seem to be. Anyway, we pulled ourselves away from these two dramatic scenes and made our way to the top of Rounds Rock, an open area with lots of lowbush blueberry. I stopped to investigate what appeared to be a tentiform mine on a blueberry leaf, but it it turned out to be a midge gall… with a torymid wasp ovipositing in it.

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Here’s a shot of the underside of one of these galls—a simple fold at the edge of the leaf.

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There are a number of midge galls like this on various plants, but it appears that none have been recorded previously on blueberry. I’ll see what I can do about rearing some.

A lot to see on a short trail loop—and in fact we hadn’t quite covered half of the 0.7 miles at that point.

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Connecticut BioBlitz

A month ago Dave Wagner got in touch with me and Julia, saying we were needed at the 2016 Connecticut State BioBlitz on June 3 and 4. He wanted to make sure leafminers were properly represented in this attempt to break the record for the most species ever recorded in a single 24-hour BioBlitz. The area to be surveyed was a five-mile radius around a school in East Hartford, an area that is mostly pavement but includes a few fragments of floodplain and upland forest. We took the challenge, and this past weekend we helped document a total of 2765 species, breaking the previous record of 2519 that was set in 2001. It was hard not to just start writing down every plant and bird we encountered, but we knew other people had these covered, so we focused on leafminers, gallmakers, and other poorly known insect groups.  We ended up contributing 133 species to the tally; our gall wasps, sawflies, and leaf-mining moths represented small fractions of the 197 Hymenoptera and 442 Lepidoptera recorded, but our gall midges and leaf-mining flies accounted for about a fifth of the 183 Diptera. Here are a few of the things we saw along the way:

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Friday afternoon we explored the woods by the Mt. St. Benedict Cemetery north of Hartford. We encountered many different kinds of sawfly larvae, most of which I couldn’t hope to identify, but I recognized the spiky oak feeder above as a species of Periclista, and a Google search for “black locust sawfly” quickly identified the one below as Nematus tibialis (both Tenthredinidae). We also found four different leafminers (two moths, one beetle, and one fly) on black locust.

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I was also able to recognize these black cherry feeders as Sterictiphora (Argidae), based on their distinctive feeding pattern, which I have written about before.

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The woods around the cemetery also yielded a walkingstick nymph (Diapheromeridae: Diapheromera femorata)…

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…a distinctively marked long-horned beetle (Cerambycidae: Strangalepta abbreviata)…

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…an unusual abundance of buffalo treehoppers (Membracidae: Ceresa)…

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…signs of leafcutter bees (Megachilidae: Megachile)…

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…a leaf mine of the apple flea weevil (Curculionidae: Orchestes pallicornis) on shadbush (Rosaceae: Amelanchier)…

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…and this tortricid, which seems to be Cydia nigricana—a new state record, but not a particularly exciting one, since it’s an introduced species.

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On the way back to the cemetery parking lot, we spotted some mines of Parectopa plantaginisella on fleabane (Erigeron), as well as some superficially similar blotch mines on the same plants which backlighting revealed to be the work of Nemorimyza posticata (Agromyzidae). This fly has not been recorded from Erigeron before, but the intricate feeding lines produced by the larvae are characteristic. I have reared adults from several other genera in the aster family that were not previously known to be hosts of N. posticata, so there is no reason to doubt that it uses fleabane as well.

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Back at the school, all the naturalists gathered in a big room and pored over their finds. Assorted galls, cocoons, and the like were sent my way, including these beautiful cocoons of Hypera rumicis (Curculionidae) on curly dock (Rumex acetosella).

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I took these home with me, and one of the weevils emerged yesterday:

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On Saturday morning we walked around the school for a little while, at one point finding leaf mines of seven different genera of agromyzid mines within a one-meter radius. There were also some lovely (albeit nonnative) snails…

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…and some equally attractive (but likewise nonnative) fourteen-spotted lady beetles (Coccinellidae: Propylea quatuordecimpunctata).

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From there we headed to the nearby Hockanum River Linear Trail, where one of the first things we found was a new host record for the leaf-mining beetle Mantura floridana (Chrysomelidae). I’m trying to rear them to confirm the record. In the first photo below you can see three of the bright yellow larvae clustered together inside a leaf; the second photo is a good example of the finely squiggly frass that is sometimes produced by leaf-mining flea beetle larvae.

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A little further down the trail we found mines of a Bucculatrix angustata larva (Bucculatricidae) in leaves of an aster (Symphyotrichum).

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The larva first made a long, narrow mine along the margin of the lower leaf, then moved to the upper leaf and made a broader mine. The larva can be seen clearly in this backlit shot of the same mine:

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Also along that trail, I found this caterpillar mummified by a species of Aleiodes (Braconidae). I probably could have identified the wasp to species using this guide to eastern caterpillar mummies, but never got around to it.

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Right at the end of the trail, Julia spotted bark mines of the elusive Viburnum Marmara. Another new state record, and this one for a native species—but of course the species doesn’t have a name yet.

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On our way back to the car, I managed decent shots (both with slightly blurry faces) of two striking little insects: Mathildana newmanella (Oecophoridae)…

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…and Metriorrhynchomiris dislocatus (Miridae).

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Not a bad day’s work.

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No, Not Leafcutter Bees

Okay, there have been six votes for leafcutter bees being the culprit for yesterday’s mystery sign. Dave Almquist was the only one to qualify his guess: “They look almost like partial cuts from leafcutter bees, assuming that the leaves are small enough for that.” I would agree with that: they do look almost like partial cuts from leafcutter bees (Megachilidae: Megachile), but there are a couple of subtle clues suggesting something else. (They are, in fact, a bit larger than the cuts a leafcutter bee would make, but I realize I didn’t provide anything for scale.)

Here are a few examples of interrupted leafcutter bee cuts, along with completed cuts where the bee removed a circular or oblong piece from the leaf.

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Notice how the pieces are cut all along the leaf margins, in no way focused on the midrib or the major veins. Also notice that the cutouts are made by cutting in one direction: none of the interrupted cuts shows any evidence of the bee starting an opposing cut, and the complete cuts never have a disjointed part in the middle where two cuts didn’t quite line up. In some cases the asymmetry of the cut makes it clear which way the bee was moving (for instance, clockwise in the one in the lower right corner of the second photo). Each of the following photos has another clear example of one of these lopsided, spiral cuts:

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Now take another look at the leaf I found yesterday:

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See how there are two counterclockwise cuts that start at the leaf margin and end at the midrib or a major vein? Each of these has a paired cut that was supposed to meet the first one from the other side. The one associated with the midrib is very short and easy to overlook. The other one made it all the way to the intended vein, but missed the first cut terribly; the insect tried to correct this, but missed again.

On a neighboring leaf of the same oak sapling, the insect got it right:

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This one wasn’t so symmetrical, but it also worked out okay:

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In this second example, it is especially obvious that the leaf cutout was left dangling from the major vein in the middle. What did the insect do while the cutout was dangling like this? Five years ago I posted photos of this insect putting the finishing touches on its creation. When I saw these cut oak leaves yesterday, I knew the finished products should be lying on the ground directly below, but none were in evidence. Apparently they rolled a little when they fell; I picked one dead leaf up off the ground and found this under it:

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This little packet was 8 mm long. Congratulations to Laura Hughes for being the only person to recognize the work of a distracted leaf-rolling weevil.

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