The Beetle and the Bee

At the end of February, I finally found a beetle I had been looking for here, for years. Perhaps I had not been scouting for it early enough, but we barely had a winter this year, and I suspect that most years it would have been evident in March. The key was finding an aggregation of its host organism.

Tricrania sanguinipennis is a blister beetle in the family Meloidae. The adult insects measure 9-15 millimeters, but are bright enough, and active enough, that they are easily seen. Their wings, hidden under their red, leathery elytra, are almost vestigial, so they are flightless. Instead they crawl, but rather rapidly, over the surface of the soil, periodically digging where they detect the possible presence of a host.

Kansas is about as far west as this beetle is found, though there are records well north into Saskatchewan, Canada. It occurs over the eastern U.S. to extreme northern Florida, and up into adjacent southern Canada.

The limiting factor is that Tricrania sanguinipennis is a parasitoid of solitary bees that nest in dense aggregations, namely cellophane bees in the genus Colletes.

We have the Unequal Cellophane Bee, Colletes inaequalis, nesting in our yard and/or the adjacent neighbor’s lawn, depending on the year, but those locations are apparently blister beetle-free. It took finding a small aggregation of the bees in a forested park to locate the beetles.

An Unequal Cellophane Bee lurks just inside the entrance to her burrow.

The bees nest in burrows, ideally in sandy soil. The vertical, subterranean tunnels branch into several individual cells, each one an “apartment” for a single bee larva. The grub feeds on a nearly liquid loaf of pollen and nectar. The walls of its room are coated in a type of natural plastic manufactured in glands in its mother’s abdomen, which essentially waterproofs the chamber, and retards mold and fungus.

Back to the beetles. What the female beetles lack in mobility, they more than make up for in fecundity. Each lady can produce hundreds of eggs, over a thousand in some documented instances. How, then, do the beetles gain entry into these tunnels? The answer is that they do not. The larvae do.

The larval stage is unusually lengthy, progressing through six instars. An instar is the interval between molts. It is also strange in that it includes hypermetamorphosis. In this case that means the larval form changes radically in both appearance and behavior from one molt to the next.

Several male Unequal Cellophane Bees waiting for females to emerge.

The first instar larva that emerges from the egg is a sleek, streamlined, highly mobile bee-seeking missile called a triungulin. At the time they are active, it is almost exclusively male Colletes bees that are active, buzzing about and frequently landing to investigate a potential site where a female could emerge. It is at these brief moments when a blister beetle triungulin scampers aboard, affixing itself to the hairs on the underside of the bee’s abdomen.

Oops! An overeager male mistakes another male for a female.

When the male bee at last is able to mate, the triungulin transfers to the female bee. In at least a few instances, the triungulin may attach directly to a female bee that it encounters on the ground. She will eventually, and unwittingly, ferry that parasite, and probably several others, to her new nest burrow. Once inside, it disembarks and infiltrates one of the subterranean nest cells.

The beetle larva usually consumes the egg of the host immediately, but not always. The bulk of its diet will be the honey and pollen left by the bee for its offspring in the cell. It may be a frequent occurrence that more than one beetle larva invades a single bee cell. In that event, cannibalism of the competition resolves the conflict.

The meal of the host egg or larva is usually enough nutrition to trigger the beetle larva’s molt to the next instar. This results in a shocking change from that sleek, active larva into the insect equivalent of a couch potato. The second instar is, shall we say….rotund, and boat-shaped. It commences feeding on the pollen and nectar stores in the cell of the now missing bee offspring. The insect retains this form for the remainder of its larval life. The first three instars shuck their old exoskeleton completely, but the fourth and fifth instars retain each molt in its entirety. Think of it as an object inside a balloon (inside another balloon by the fifth instar). Those larvae actually shrink in size to fit inside the shed “skins.” When molting into the pupa stage, the sixth molt is again broken during shedding, and compacted at the rear of the pupa, which is still inside those other exoskeleton balloons.

It takes until late summer or early fall for the life cycle to complete, the adult beetle remaining encased in its final one or two larval exuviae, where it overwinters, still inside the cell in the host’s nest burrow.

You would think that the bee species hosting this diabolical beetle would be decimated by it, but such is not the case. Each spring there are plenty of the adult bees. What is more of a threat is the potential disconnect between the bees and their nectar plants. Colletes inaequalis visits flowering trees almost exclusively, especially Eastern Redbud, and maple trees. As the phenology of the blooming cycles becomes increasingly unpredictable thanks to climate change, the appearance of the flowers may cease to always coincide with the emergence of the bees that pollinate them.

Sources: Messinger Carril, Olivia, and Joseph S. Wilson. 2021. Common Bees of Eastern North America. Princeton, NJ: Princeton University Press. 286 pp.
Parker, J.B. and Adam G. Boving. 1925. “The Blister Beetle Tricrania sanguinipennis - Biology, Descriptions of Different Stages, and Systematic Relationship,” Proc U.S. Nat. Mus. 64(2491): 1-40. This is a wonderfully exhaustive article, with illustrations of all larval instars.

Fly Day Friday: Rat-tailed Maggots

I recently started working part-time at Songbird Supplies, LLC, located inside Summerland Gardens here in Colorado Springs. One of the perks has been encountering insects I seldom see. Julie, the proprietor of Summerland Gardens, called me over the other day to help her identify a "primordial soup" bug in one of the water gardens she is cultivating. I was surprised and delighted to find the critters in question were "rat-tailed maggots," the larvae of flower flies in the tribes Eristalini and Sericomyiini (subfamily Eristalinae, family Syrphidae). I had seen them maybe one other time previously.

The conundrum faced by folks with water container gardens is this: do I put in "mosquito dunks" to kill the mosquito larvae, or do I refrain so that the syrphid larvae can grow up into pollinator-capable adult flies? There is no right or wrong answer. Syrphid flies are definitely not endangered species, so I might deploy the dunks.

Note the leg-like "knobs" along the length of the body of the rat-tailed maggot on the left

The rat-tailed maggots are named for the posterior spiracles, which are fused into an extensible breathing siphon that connects them to the water surface at all times. Meanwhile, at the other end, they are filtering bacteria and other microbes from organic matter in the water. Larvae of some species exist in rather dense material like saturated manure, which barely qualifies as aquatic, but is too wet for other dung-eating fly larvae. The low oxygen levels in this kind of substrate are no problem because the maggot's spiracles are always connected to the surface.

The length of the "snorkel" varies greatly depending on the genus of flies. Eristalis larvae have very long tubes, as shown in the images here; Chrysogaster falls at the other end of the spectrum, with quite a short siphon.

Different genera occupy different aquatic niches. Mallota and Mylopeta, for example, live in the water that collects in tree holes. Sericomyia larvae live mostly in bog mat pools. Chrysogaster thrive along the edges of ponds and among emergent vegetation. The most commonly-encountered species are probably in the general Eristalis, Eristalinus, Helophilus, and Palpada, since they are most likely to be seen in urban, suburban, and rural areas. They live in putrid and organic-rich standing water. I know they are sometimes found while cleaning rain gutters.

Once the larva reaches maturity, it seeks dry land on which to pupate. Teh pupal capsule is hard, and resembles a tiny mouse in shape.

Adult Drone Fly, Eristalis tenax, Colorado

The "Drone Fly," Eristalis tenax, is the most common adult version of a rat-tailed maggot that one is likely to see. It is a stunning mimic of the honey bee. Likely introduced ages ago from Europe, it is now an established and widespread species across the entire North American continent. It is also a fairly respectable pollinator, visiting wildflowers, and the blossoms of orchard trees and landscape plants.

Here is an interesting piece of trivia: It is speculated that it was rat-tailed maggots that gave rise to the biblical account of "bees" spontaneously appearing from the rotting carcasses of animals. Certainly, the fluid decomposing material in a corpse would be a perfectly acceptable habitat for Drone Fly larvae.

Adult male Eristalis tenax, Arizona

Sources: Carr, John F. 2013. "Rat-tailed Maggots," Bugguide.net.
McAlpine, J.F. (editor). 1987. Manual of Nearctic Diptera Volume 2. Ottawa: Agriculture Canada. Monograph No. 28. pp. 675-1332.
Merritt, Richard W. and Kenneth W. Cummins. 1978. An Introduction to the Aquatic Insects of North America. Dubuque, Iowa: Kendall/Hunt Publishing Company. 441 pp.
Pfiester, Margaret, and Phillip E. Kaufman. 2009. "Rat-tailed Maggot," Featured Creatures. University of Florida Department of Entomology and Nematology.

Giant Silkmoths

We were fortunate to have hosted three species of giant silkmoths (family Saturniidae) in Butterfly Magic this year. They are spectacular insects, but so unique and short-lived that one cannot generalize to moths as a whole from their life cycle and behavior.

Estimates differ as to how many species are in the Saturniidae, but somewhere between 1,300 and 1,500 is fairly accurate. Gene sequencing could result in more or fewer species than currently recognized. Giant Silkmoths are found the world over in forest habitats, but are especially diverse in the neotropics (Mexico, Central and South America). You are probably already familiar with some U.S. species: the Luna Moth, Cecropia Moth, Polyphemus Moth, Royal Walnut Moth, Imperial Moth, and Cynthia Moth. Not all the silkmoths are giants. Buck moths (genus Hemileuca) are much smaller, and fly during the day.

The adult moths do not feed. They have vestigial mouthparts at best and fuel their flight on fat reserves accumulated as caterpillars. Consequently, the caterpillars of giant silkmoths are large and heavy at maturity. The “hickory horned devil,” larva of the Royal Walnut Moth, approaches the size of a frankfurter. It looks menacing but is harmless.

Meanwhile, the larvae of other silkmoths are studded in venomous spines. Beware the caterpillars of buck moths (I can attest from personal experience!), the io moth, and especially the South American Lonomia (see also more recent articles in medical journals).

The Forbes Moth, Rothschildia lebeau forbesi, is frequently mistaken for an Atlas Moth by our visitors. Both species have “windows” in their wings that are devoid of scales. The Forbes Moth ranges from the Lower Rio Grande Valley through much of eastern Mexico and south to Brazil. Relatives of this moth figure prominently in the culture of indigenous peoples. Bushels of cocoons are harvested to make rattles worn on the ankles during ceremonial dances.

The African Moon Moth, Argema mimosa, occurs over most of sub-Saharan Africa. Their cocoons are distinctive: compact, and made of dense silk with numerous small perforations. The adult moths are spectacular: a wingspan averaging 125 mm, yellowish green in color with long, streaming “tails” on the hindwings. What purpose those tails serve is open to speculation, but perhaps it further camouflages the insect when it is at rest among foliage.

Last but not least is the Atlas Moth, Attacus atlas, a real giant with a wingspan of 240 mm (compare that to the Forbes Moth, only 90-100 mm from wingtip to wingtip). The Atlas Moth is widely distributed, from India to Hong Kong, tropical Asia in general, plus Taiwan and Indonesia. The front wings have a conspicuous lobe with markings that suggest the head of a serpent, but whether this is real mimicry or not is debatable.

The adult females of all the silkmoths rarely stray far from their cocoons after emerging. They invest most of their energy in egg production, so instead of exerting calories in flight, they simply sit and emit a species-specific sexual attractant called a pheromone. This scent, while usually imperceptible to us, is potent when it comes to drawing males. The males, with broad wings and streamlined bodies, use their sensitive antennae to home in on the pheromones, following the scent trail up to a mile or more to find its source. Females, once mated, will then finally fly to appropriate host trees and deposit their eggs. Even unmated females will dump their eggs in great quantities, as is often seen in our greenhouse. Both genders live only a few days, assuming they can avoid predation by bats.

Breeding silkmoths in captivity is becoming quite the hobby and cottage industry. My friend Liz Day has a web page devoted to helpful hints for rearing common eastern U.S. species. Considering the threats faced by these magnificent moths, from suburban sprawl and light pollution to introduced parasites, our domestic species could use some help.

I would like to conclude by recommending Wings of Paradise by John Cody, published by University of North Carolina Press, 1996. Cody renders giant silkmoths in stunning watercolor paintings. He features all three species in the greenhouse, plus scores more. The art is complemented with anecdotes and factual information.