Marsh Marigold Menagerie

In the spring of 2013, I wrote about the four-toed salamander surveys I was conducting at the time, which involved crawling around in swamps all over northwestern Massachusetts. On June 1, the last day of the survey, Julia tagged along, and one of the swamps we visited had a large population of marsh marigold (Ranunculaceae: Caltha palustris). The plants were probably all fruiting by that point, but this is what they would have looked like a month or so earlier:

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We noticed that many of the plants had brown markings that appeared to be leaf mines, radiating from the point where the petiole attached.

dsc_0407 A little investigating revealed that the brown streaks continued down the petiole, and in some cases into the stem.

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Julia split open a few of the petioles and found elongate, black-headed larvae feeding inside.

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I recognized them as being some sort of “lower” fly (Nematocera), which among the leafminers I knew of would indicate some sort of midge (Chironomidae). No other leaf-mining fly larvae have well-defined head capsules like this.

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We stuffed a bunch of the mined leaves and petioles into a bag to see what would come out. The first adult appeared six days later.

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It was not a midge, but some sort of dark-winged fungus gnat (Sciaridae). I hadn’t seen any references to leaf-mining sciarids in North America, but a little online search revealed that a few are known in Europe. As luck would have it, no one in the US studies sciarids, but I found someone who was willing to take a crack at identifying my specimens, so I sent them off once they had finished emerging. In the meantime, a number of other insects had appeared in the bag.

In addition to the thirteen sciarids, there were eight of these frit flies (Chloropidae)…

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…twelve of these scuttle flies (Phoridae)…

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…three of these weird little ichneumon wasps…

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…and this fruit fly (Drosophilidae).

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All of these insects had apparently emerged from the tunnels excavated by the sciarid larvae in the plant tissue. Their larvae were parasitoids or predators of the sciarid larvae, or else they were scavengers, feeding on the dead plant tissue or possibly the waste products of the sciarid larvae. There had also been some beetle larvae feeding on the surface of the leaves:

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Curious to see what they turned out to be, I hadn’t picked them off when we were collecting the mined leaves. After two weeks, they crawled away from the leaves and pupated…

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…and I ended up with five of these adult beetles:

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The beetles proved to be the easiest to identify. They were Prasocuris boreella (Chrysomelidae), which had been collected once before from marsh marigold, but apparently no one had ever found the larvae and reared them before.

I sent the other insects to various specialists, but I hadn’t heard back from anyone yet by the next spring, and the person with the sciarids didn’t respond when I inquired about them. So when I encountered more of the mines while doing wetland plant work in June 2014, I collected them to see if I could rear some more. I collected marsh marigold samples from two sites, and succeeded in rearing another seven or eight sciarids from each. I also ended up with another fruit fly, eleven more frit flies (five from one site and six from the other), and this braconid wasp:

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I asked Ray Gagné, the gall midge specialist, if he knew of anyone who studies dark-winged fungus gnats, and he put me in touch with Frank Menzel, a German, who referred me to Kai Heller, another German, who was happy to help. So off to Germany some of the new flies went, and before long Kai had identified them as Zygoneura calthae, a species that had been described from Finland in 1960 and was not known to occur in North America. The collection data for the type specimen included “Larven und Puppen im Blattstiel von Caltha palustris“: larvae and pupae in the petiole of marsh marigold. Nothing seems to have been published about the species since then.

I wanted to write something up for publication about the discovery of this species in North America, but first I needed to round up identifications of all the associated insects, which took another year or so.

Terry Wheeler and Julia Mlynarek reported back that my three sets of frit flies all represented the same undescribed species of Elachiptera. Julia is working on a revision of this genus, so she will give this fly a name sooner or later. My impression is that it is not a species either Terry or Julia had seen before, but maybe some will turn up in museums by the time the revision is complete. Most Elachiptera species seem to have larvae that feed on decomposing plant tissue damaged by other insects. This one seems to have a special relationship with marsh marigold, if not Zygoneura.

Brian Brown and Emily Hartop identified the scuttle flies as Megaselia limburgensis. Interestingly, this species has been reared in Europe in association with an agromyzid fly that bores in marsh marigold petioles. Hering* stated that “strange mine off-shoots” from the petiole were constructed in the leaf blade, and he wasn’t sure whether these were made by the agromyzid or the scuttle fly. These sound suspiciously like the leaf mines of Zygoneura, but apparently he didn’t find any associated sciarids.

The ichneumon wasps now reside in the British Museum of Natural History, where Gavin Broad identified them to the genus Neurateles. Two described species are known to occur in North America, but they more likely belong to one of several undescribed species. Neurateles is in the subfamily Orthocentrinae, which seems to be mostly made up of fungus gnat parasitoids.

The fruit fly, Rick Lapoint informed me, was Scaptomyza pallida, a very common, cosmopolitan species that feeds on decomposing plant tissue.

Joseph Fortier identified the braconid wasp as a species of Ephedrus (Aphidiinae), indicating that it must have emerged from some unseen, mummified aphid on the plant surface rather than having anything to do with the sciarid mines.

Having collected all of this information, I was now ready to write my paper. This past February, I wrote to Kai Heller with a quick question regarding where Zygoneura calthae had been found besides the original collection site in Finland. When he responded, he mentioned that DNA barcoding had shown my specimens “to be different genetically from the European ones by more than 6%. Additonally they look a bit different. It may be possible, that two different species or subspecies are present.”

I thanked him for this bit of information, and remarked that it agreed with my suspicion that this is a native species rather than a recent introduction (which should closely match a European population). He wrote back and said that he had rechecked the DNA results, and in fact the difference was nearly 10%: unquestionably a new species. So I put off submitting the paper for a little longer, as Kai wrote up a description of the species and Björn Rulik wrote up the DNA analysis. I decided to name it Zygoneura calthella, which seemed fitting for a species that’s pretty much the same as Z. calthae but a little smaller. There are also some subtle differences in the male genitalia between the two.

Our paper** describing Zygoneura calthella, along with a review of insects known to feed on marsh marigold, was published at the beginning of November.

* Hering, Erich Martin. 1951. Biology of the Leaf Miners. Junk, the Hague, Netherlands.

** Eiseman, Charles S., Kai Heller, and Björn Rulik. 2016. A new leaf-mining dark-winged fungus gnat (Diptera: Sciaridae), with notes on other insect associates of marsh marigold (Ranunculaceae: Caltha palustris). Proceedings of the Entomological Society of Washington 118(4):519-532.

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Quality Control

On June 22, I was doing botanical work in a shrub swamp in Southbridge, Massachusetts when I found this 5.5-mm larva on the underside of a highbush blueberry (Ericaceae: Vaccinium corymbosum) leaf:

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I find little tents like this from time to time on various trees and shrubs, and I’ve always assumed they’re the work of Machimia tentoriferella (Depressariidae), since that’s what it says on page 219 of Tracks & Sign of Insects. You can’t trust everything you read though, even if it’s something you wrote yourself, so I decided to take this opportunity to verify my assumption. I stuck the leaf in one of the little vials I carry in my backpack everywhere I go.

By July 3, the larva had grown to 8.5 mm and had spun its fourth tent on the same leaf. It had done some nibbling at the edges of the leaf in addition to its characteristic skeletonizing at either end of the tents.

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Five days later, it  had grown another millimeter and was eating large holes in the leaf.

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On July 25, it had reached 11.5 mm and was still working on the original leaf, which had turned completely brown at this point.

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By August 24, feeding on a fresh leaf, it had reached 16 mm and had developed a couple of yellow longitudinal stripes—hence the name “gold-striped leaftier.”

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That was the last time I photographed it, but it was still feeding into early September. I never imagined that rearing a little micro-moth would take all summer. On September 28, the adult emerged, confirming that it was what I had assumed it to be.

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It had pupated inside a tightly folded leaf. At no point was it actually a leaftier, which is a term reserved for larvae that use silk to fasten two or more leaves together.

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The HOSTS Database records Machimia tentoriferella from maples (Acer spp.), birches (Betula spp.), hickory (Carya), walnuts (Juglans spp.), chestnut (Castanea dentata), beech (Fagus grandifolia), oaks (Quercus spp.), balsam poplar (Populus balsamifera), basswood (Tilia americana), elms (Ulmus spp.), apple (Malus), cherry/plum (Prunus spp.), mountain-ash (Sorbus), true ashes (Fraxinus spp.), lilac (Syringa vulgaris), buttonbush (Cephalanthus occidentalis), dogwoods (Cornus spp.), and viburnum. If that is a complete list of published host records (which it probably isn’t), this rearing from blueberry is the first recorded instance of M. tentoriferella using a host in the heath family (Ericaceae).

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Life in a Monkeyflower Stem

I’m slowly making my way through all the photos I took this year, and I just finished August 6. That was the day of Julia’s family’s annual BioBlitz at Deep Woods Farm, their land in southeastern Ohio. A funny thing happened as Julia and I were heading back for dinner after the day’s exploration. We were walking past a little patch of monkeyflower (Phrymaceae: Mimulus ringens) that had popped up in an opening in the forest created when some trees blew down a few years ago…

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…when something made me say, “if I were a fly, I’d mine the stems of monkeyflower.” There is no fly known to mine in the stems of monkeyflower (or in the leaves, for that matter), but just for the heck of it I stopped and took a look. It took only a moment to spot the first mine:

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Since I was pretty sure this was something unknown to science, we of course had to delay dinner a little bit and spend a few minutes scouring the plants for larvae and pupae. We found several; here is the puparium of a larva that did its last bit of feeding in the midrib of a leaf:

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At the far left end you can see its little black spiracular horns (breathing structures) poking out through the leaf epidermis. I’ll tell you more about these flies some other time, but the reason I’m bringing this up now is just to explain how we came to be closely examining monkeyflower stems, when Julia noticed that some of them were weakened toward the base. She broke one at its weak point and discovered a caterpillar inside:

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The caterpillar had been busily converting the inside of the stem to mushy poop.

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I figured that while we were collecting monkeyflower stems, we might as well see if we could rear this caterpillar.

The next day, the caterpillar had popped out of its stem, and it seemed uninterested in trying to enter another stem.

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I had never tried to rear a stem-boring caterpillar before, but last summer when we visited MJ Hatfield in Iowa, she showed us how she was rearing all sorts of Papaipema (Noctuidae) caterpillars by offering them chunks of carrot to bore into. I wasn’t sure if this was a Papaipema, but I didn’t know what else to try, so I put it in a jar of soil with a carrot. It happily got to work turning the carrot to orange mush.

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Weeks passed, and the carrot turned brown and shriveled down to the soil. When I put all my bugs into the fridge to overwinter at the beginning of November, there was no sign of an adult moth in the jar, and I was skeptical that the caterpillar had survived. Since I was running out of space in the fridge, I left that jar on a shelf in my office along with a few others that I considered to be lost causes.

When I came to the photos of the caterpillar just now, I decided to go and have a look in the jar, and when I unscrewed the lid this moth was waiting just below it, perched on a section of monkeyflower stem I had put in there along with the carrot:

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I know next to nothing about noctuids, but I just browsed through the Papaipema pictures on BugGuide.net, and this seems like a good match for P. cataphracta, the so-called “burdock borer.” The guide page states that “larvae bore in rhizomes and stems of aster, burdock, corn, cottonwood, iris, lily, sunflower, thistle, tomato, and other plants.” So it sounds like this species will bore into just about anything with a stem. A couple of other species look pretty similar to this, but they are apparently specialists on the aster family; there are around 50 species in this genus, and my impression is that most of them are host-specific.

BugGuide also says that adults of Papaipema cataphracta fly from August to October, but there is one example of an adult that was found out and about in New Jersey on November 7 under natural conditions. I couldn’t say when in the past three weeks this monkeyflower one emerged; it’s perfectly alive, but it seems pretty sleepy and uninterested in flying at this point.

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Acorn Weevils

It’s another mast year for red oak (Quercus rubra) in my neck of the woods, which means that for the past couple of weeks, large nuts have been raining down from the treetops, making it a little treacherous to go for a walk in the forest. Last November, once the acorns had all fallen, Julia and I collected a few gallons of them to process and use for food. (We’ve made tasty acorn bread and acorn cake, but so far our favorite is acorn falafel.)

Anyone who has collected acorns is familiar with the whitish grubs that turn the insides to dust, eventually exiting through round holes like this one:

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The grubs are immature weevils in the genus Curculio (Curculionidae).

After collecting our acorns, we dried them by placing them on trays by the wood stove, and I think this arrested the development of most of the larvae that were feeding inside them. However, on December 7 I noticed two mature larvae squirming around on one of the trays.

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Just for the heck of it, I dropped both larvae into a jar of soil, and after they burrowed down I put them in the little refrigerator in the basement where all my other bugs pass the winter. I took the jar out on March 1 along with everything else, but it wasn’t until June 14 that an adult finally appeared.

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When I was satisfied with the photos I’d taken of it indoors, I brought it out to a red oak sapling in the yard to try for some more natural shots.

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It soon wandered onto a young stem, and as I continued to photograph it I realized that it was feeding by chewing a tiny hole into the stem.

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In the last photo above, it has finished feeding and the hole is visible just to the left of the tip of its proboscis.

Added 10/25/16: Here’s a great video Laura and David Hughes got last year of a larva chewing its way out of an acorn. Amazing to see it squeeze its blobby body through a hole that is just barely big enough to fit its head capsule.

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Mailbag

People regularly send me photos of mystery objects and other bug-related phenomena to see if I recognize them. Often I’m able to respond with a precise (or approximate) identification, and that’s the end of it. Other times I’m intrigued by the mystery but don’t have anything too useful to say about it. My last post featured an unusual case where I was able to collect the larva in question and raise it to an adult. More typically when a mystery piques my interest, I offer to post the mystery photo on my blog to see if any of my readers recognize it. Lately, a pattern has developed where the photographer says sure, go ahead and post it, but then I never quite get around to it… until now! Tonight I’m going through my old emails and rescuing these photos from the depths of my inbox. [Note: It seems that there is nothing I can do to convince WordPress to make proper spacing between the blockquotes and images. It all looks very nice while I’m editing it, but it’s all messed up when I preview/publish it. Sorry about that.]

First up is a pupa found by Scott Smyers on Mount Wachusett in eastern Massachusetts on July 11:

I found this interesting pupa under a snake board in a field at Wachusett on Monday.  I looked through your book and couldn’t find anything like it.  Any ideas?

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It’s definitely a lepidopteran pupa, but I don’t remember ever seeing one with bold longitudinal stripes like this. [Edit: Vazrick Nazari recognized it right away as the chrysalis of a common wood nymph, which is a butterfly I know well. That’s one mystery down!]

This next one was sent to me by Erin Banks Rusby around the same time (July 13):

I work for a non-profit in the Palo Alto, CA area and saw the following on a plant in Menlo Park, CA. I can’t tell you what plant it was on, unfortunately, but I can tell you that there were other markings like this one on other leaves of the plant. Is it an insect or bug that made it and do you have any idea what it could have been?
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My response at the time was:

Interesting pattern!  It’s not immediately recognizable to me… it appears to be something on the leaf surface, but I can’t quite tell from the photo whether it’s something clear and shiny or something white and fuzzy.  If the former, all I can think of is some kind of slug/snail, but that seems unlikely given how convoluted the pattern is.  If the latter, I’ve seen planthopper nymphs and related insects (e.g. psyllids) leave behind something vaguely similar, but again, not such an intricate pattern.  I’d be interested to hear more details or see other examples.

She consulted with her organization’s botanist/naturalist, who suggested it was a leaf mine of some sort. If it were a mine, it would pretty much have to be from a moth in the family Gracillariidae, since very shallow mines resembling snail trails are characteristic of that family.  But it would be very unusual for a gracillariid mine to be so tightly contorted, and there are a few outlying dots in the periphery of this pattern that make me think it is in fact some kind of whitish substance that has been deposited on the leaf surface. Unfortunately we don’t know what the host plant is, but maybe someone will recognize it from the photo above.

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The next one was sent to me a year ago (October 14, 2015) by Piet Tutelaers:
Since 2011 our insect working group of the KNNV Eindhoven in The Netherlands is puzzled with these strange cocoons. We have asked help from people from EIS (European Invertebrate Survey) in Leiden also in The Netherlands. They thought it would probably be cocoons from Asterodiaspis variolosa (Ratzeburg) [the golden oak scale; family Asterolecaniidae] but that turned out to be false. We have tried to breed the cocoons to see what animal comes out but that has failed so far (see photo 1
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and photo 2).
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The left cocoon in photo 1 and the one in photo 2 is the unknown one and the smaller right cocoon in photo 1 is from Asterodiaspis variolosa . The cocoons are 4mm long and 1.5-2mm wide. The small white cover is the place where the cocoon is opened and left by the inhabitant.

…The cocoons can be found in forests (plantations) of Douglas fir, a fir that is not native to Europe but is cultivated for its wood. Here we find the cocoons underneath fallen dead branches in the neighbourhood of Douglas fir. There should be some free air between the place of the cocoon and the soil.

I have opened one cocoon in order to see what is inside it and have made a photo with my stereo microscope. This photo gives me the strong impression that we have to do with an insect (see attached photo).

Because the Douglas fir is native in your country this insect is possibly known in the USA? So I hope you can help me in solving this puzzle.

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My response:

Looking at the first two photos, I was going to guess an egg sac of a spider such as Agroeca, but your photo of the opened one makes it clear that it is a cocoon, presumably of a moth.  Would it be possible to remove the object (pupa?) from the cocoon and get clearer photos of it?  Unfortunately I have never seen something quite like this before.  It may or may not be something related to the Douglas-fir; I live on the other side of the country from where Douglas-fir is native, and have only spent a little time exploring the Pacific Northwest.
Piet was going to try and get sharper photos of the pupa, but never got back to me with those, which is my excuse for taking so long to get around to posting these photos. (It occurs to me now to wonder if the object inside is an embryo rather than a pupa—it sort of reminds me of the egg coverings made by certain ground beetles, but I don’t know if any of them are capable of making a silken lid like this.)
I have no such excuse for not doing anything until now with the photos below that John van der Linden sent me in August 2015. Julia and I had met John a few weeks earlier when we stayed with MJ Hatfield in northeastern Iowa, but John’s photos are of things he found on tree trunks in India.
1. The top image shows a flattened case that has a two-lobed shape. This case was structured like a turtle shell. The visible part is larger in diameter and faces out (the “carapace”). The invisible part is smaller in diameter and touches the bark surface (the “plastron”). The larva hid sandwiched between these two layers.
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This well-camouflaged object is shown in the inset and in the upper left corner of the larger photo. I asked John if it was a portable case, and he replied that from what he recalled, both it and the next one were very loosely attached to bark with just a few strands of silk “…as if the larvae were camping out for a while…but could also sever the strands and move if they wanted?”
2. What I called the “cobra case” because it reminded me for some reason of the hood of a cobra. Fascinating architecture (see diagram, next picture). Yes, the case actually had two rows of holes built into it, along its margins!!
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3. Diagram of the cobra case.
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4. Cobra case stripped down to just the tube part of the case, with larva partly emerged.
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cobra-case-larva-45. Another view of the cobra case larva. The head was distinctly flattened into a blunt disc with the approximate inner diameter of the tube part of the case, so that when the larva rested inside the tube its flattened head acted just like a manhole cover, sealing off the tube.

My comment on the “cobra case” was:

The larva looks like something gelechioid to me, like a Coleophora… somehow the construction of the case is reminding me of Batrachedridae, which includes the octagonal casebearer as well as a species (pictured in my book) that makes a vaguely similar expanding frass shelter on palmetto leaves [here is an example on BugGuide; both of these moths are now placed in the family Pterolonchidae].

As a bonus, I’ll leave you with a mystery that got solved before I got around to posting it here. On June 17 of this year, Valerie Bugh wrote:

I have just found a second example of a mystery I can’t solve. I found the first about 5 years ago in FL. The second was found today in Austin, TX. Size is just under 10 mm if I remember correctly. They seem sort of waxy, and the “tail” is oriented down. I thought about some sort of diptera puparium, or maybe a wasp or bee nest. Anyway, I was wondering if you’ve seen this and know what it is.

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My response:

The shape reminds me of some spider egg sacs, but the material doesn’t seem to make sense for that… The closest thing I can think of is a resin bee nest, but the ones I’ve seen always are packed with pebbles and never have any kind of tail like that. . . I would strongly suggest that you put it in a container and give it a few months to see if something comes out, and if not, then cut it open to see if you can make sense of its contents.

In August Valerie wrote back with an update. The insect turned out to be one she knew as an adult—see her photo here.

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A Hidden Masterpiece

Last December, Jennifer Kleinrichert sent me this photo of a polka-dotted larva (prepupa) surrounded by an elaborate silken structure, wondering if I knew what it was:

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My response:

I’m afraid I don’t know this one… I’ve seen a photo of something similar once before; see the third one on this page. It would be interesting to know if this larva ends up spinning a similar dense, white cocoon. The polka-dots on your larva remind me a little bit of some pyraloid moths… can you tell me anything else about the context?  Looks like it was under a plywood board?  If you don’t mind, I’d love to post this as a mystery on my blog.

 
She replied that she and her husband had been changing some pictures in a bulletin board, and upon opening the door found the larva in between the two frames. In the photo below, the larva’s creation is visible on the horizontal board directly below the photo of a viceroy.

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I decided to hold off on posting the mystery here, because in January Julia and I happened to be visiting her parents in Ohio, just a half-hour drive from the bulletin board, and I figured I might as well take a crack at raising the larva to an adult. Jennifer and her husband Steve met us at the bulletin board and removed the panel for us. The larva was still in the same position within its gauzy double fence, which was about 6 cm across.

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We gently removed the larva and put it in a sandwich box with a  folded-up paper towel.

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I kept it in the refrigerator until March 1. After that, I took a peek at it every day, and although I didn’t actually see it move much, it seemed to be in a different position every time I looked. I expected it to pupate soon after it was exposed to warmer temperatures, but it wasn’t until nearly two months later, on April 23, when I found the larva suspended in some webbing in a fold of the paper towel where I hadn’t seen it venture before.

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Another week passed, and finally, on May 1, it shed its skin and pupated.

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I didn’t want to disturb it too much then, but I was able to get a better shot of the larval and pupal exuviae on May 27, when the adult moth at last emerged.

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My guess that it was some sort of pyraloid moth turned out to be correct. Specifically, it was a dogbane saucrobotys (Crambidae: Saucrobotys futilalis). I don’t think I had seen an adult before (at least, I had never been moved to photograph one), but the larvae were conspicuous in the dogbane patch by our mailbox last September.
 
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I never would have guessed that the larvae leave their messy feeding webs to spin such intricate structures for hibernating and pupating.

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Bean Borer

Last August, Julia noticed a few holes in some string beans in the garden.

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One of holes had droppings pouring out of it, and we could see someone fuzzy inside.

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I opened this tunnel up for a better look at its inhabitant.

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A quick look in a field guide told me what this was going to turn into, and I generally focus my rearing efforts on unknowns, but I decided to make an exception. After munching on beans for another week, it shed its fuzzy skin to reveal a fuzzy brown chrysalis.

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In the spring, it emerged as a gray hairstreak (Lycaenidae: Strymon melinus) as expected. It obligingly posed on a dandelion before fluttering away.

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We’re more than happy to sacrifice a few string beans to have these around.

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