Today we have a special contribution by Athayde Tonhasca Júnior. The photos are not his, but he’s writteen an illustrated mini-essay on a case of parasitism. Athayde’s words are indented, and you can enlarge the photos by clicking on them. You don’t often see life cycles as complicated as this one!
If you are getting low on wildlife contributions, your readers may enjoy -(or be horrified by, depending on one’s take) an introduction to stylopids.
Stylopized, emasculated and zombified: the risks of visiting a flower
‘Bizarre’ and ‘weird’ are overused adjectives for describing many characters and events of the natural world. Life is way too complex and varied to conform to familiar patterns, so the out-of-the ordinary is all around us, even though we not always see it. But when the discussion turns to stylopids, it’s difficult to avoid talking about the bizarre and the weird.
Stylopids (or stylops) are small, seldom seen and poorly known insects with about 700 described species, 10 to 16 of them in Britain. The true figures are likely to be much higher. They are parasites of other insects such as bees, wasps, plant hoppers and leaf hoppers. From a distance, male stylopids can be mistaken for flies, but their ruffled wings give them away and explain the name of this group of insects: the order Strepsiptera, from the Greek strephein (to twist) and pteron (wing). Twisted-wing insects is another common name for them.
Males have branched antennae and their eyes are berry-like structures comprising dozens of image-forming eyelets. This unusual array inspired the development of new cameras of reduced size and sharp images, which are handy for smartphones. Males cannot feed because their mouthparts are not developed. But never mind going hungry; they don’t live for more than a few hours. Their only objective in life is to use their fancy eyes to find a female and mate.
Females look nothing like the males. In fact, they don’t look like your ordinary insect at all because they don’t have wings, antennae, legs, mouthparts or eyes: they are neotenic, i.e., they retain their larval features. An adult female does not need a fully formed body since she never leaves her host: she will develop and die semi-buried in another insect. ‘Semi-buried’ because the tip of her cephalothorax (the head and the thorax fused together) protrudes from the host’s abdomen.

Through this exposed area, the female releases a pheromone to attract males. Once suitors finds her, they face an anatomical challenge. Only parts of her head/thorax are exposed, which doesn’t bode well for conventional insect romance. But this setback is nothing compared to the facts that she doesn’t have genitalia, and her eggs float in the haemolymph (‘blood’). So a male has only one course of action: the disturbingly sounding traumatic or hypodermic insemination. He pierces the female’s cuticle with his penis and injects his sperm into her haemolymph. Watch it at this link.

The deed done, males soon die. The fertilised eggs hatch inside the female, giving birth to thousands of tiny planidia (singular planidium, from the Greek planis, meaning ‘wanderer’). These are a type of larva that don’t look at all like larvae. They have well developed legs, are quite nimble, and they’re phoretic, that is, they use another organism to be transported to a new location. The planidia feed on mum’s innards and eventually crawl out of her body to disperse and start looking for a host of their own.

A wandering planidium climbs a flower to wait for an unsuspecting visitor. When a bee or wasp lands, the planidium somehow hitchhikes a ride to their nest. We are not sure how it does this: it could hide in the pollen, or possibly be swallowed with the nectar sucked up by the flower visitor, then released when the host regurgitates nectar inside the nest. Chances are it will end up in the wrong nest, so most planidia are done for. But the staggering fecundity of female stylopids compensates bad odds: they can dish out 750,000 planidia, so a few are likely to find a suitable host.
Once inside the right nest, the planidium burrows into a host’s egg or larva and transforms into a traditional legless, grub-like larva. It is followed by other larval stages, pupation, and finally adulthood – by then the host has also become an adult. If it’s a male stylopid, it squeezes out of the host and flies away, usually leaving a big gap behind and killing the host. If it’s a female, it will park itself in the host’s abdomen.

JAC: I added this video of a male emerging:
The story above is a peep at stylopids’ life histories, as there is considerable variation depending on the species and type of host. And if all this sounds outlandish, there is more to come:
Stylopization (parasitism by stylopids) causes all sorts of physical and behavioural eccentricities in the host, all for the parasite’s advantage. Host infertility is one example. Reproduction involves mating, nest building, nest provision, etc., all of which are risky and energy-consuming, and therefore not beneficial for a parasite. Stylopids solve this problem by disabling the host’s reproductive organs, functionally castrating them. Some stylopized bees have reduced scopae (pollen-carrying structures) and seldom if ever carry pollen: there’s no point, as they don’t have a brood to provide for. Contrary to what happens to most parasitized insects, stylopization often lengthens host development; they live longer so that stylopids have more time to mature. Some stylopized bees are led to stand still on a grass or flower stems with their head downwards. Such a zombie state greatly facilitates stylopids’ mating business.
In Britain, furrow bees (genera Halictus and Lasioglossum), yellow-face bees (genus Hylaeus) and especially mining bees (genus Andrena) are victims of stylopids, but we have little information about their interactions and no idea about consequences of parasitism.
Stylopids are odd and enigmatic, but they are also one of the most complex and intriguing groups of animals. They are evolutionary marvels that have puzzled and awed generations of entomologists and naturalists, and more surprises will be revealed from future research. It seems quite fitting, then, for the august Royal Entomological Society to have adopted a stylopid (Stylops kirbii) as a representative of the organisation:

Athayde, this is utterly fascinating, thank you.
Thanks, Janet.
Well that made my day! Thank you very much for sharing this very weird stuff.
I’ve seen many Stylopidized wasps with females, but have yet to see a male. I believe they were placed in the Coleoptera (beetles) for a time, but are now in their own insect order.
You are right, they have been placed with beetles. They are still considered a phylogenetic headache, I think.
Thank you, Athayde! It has been nearly 50 years since I was shown Strepsipteran females on pinned bee specimens in the university collection. Never since,.
Interesting, as you likely know, there is one primitive genus, Mengenilla, with free living females…
The relatedness to beetles is still possible, but at most as sister-group [so Coleoptera+Strepsiptera]. Even more bizarre was the suggestion that they arose from flies [Diptera] or fly-ancestor — requiring a switch [of wings and halteres] between the second and third thoracic segments
. . . why am I suddenly thinking about politicians and religious leaders?
Seriously, amazing life cycle, and thank you.
You are welcome!
Thanks!
Please tell me that stylopids have their own parasites. They would definitely have to have an even weirder life cycle.
So, naturalists observe, a flea
Hath smaller fleas that on him prey;
And these have smaller still to bite ’em;
And so proceed ad infinitum.
Thus every poet, in his kind,
Is bit by him that comes behind.
Jonathan Swift, “On Poetry, A Rhapsody”, 1733
(I don’t know…)
Nature is endlessly fascinating – thank you for this wonderful account! As you say, the males look superficially like true flies but the amongst other differences the relative positions of the halteres and the wings are reversed.
Indeed, Jonathan. We can’t never become bored with the natural world.
I know that name. These being insects, that in itself is a good predictor of a gory story.
Oh, them. Yeah, I remember them being discussed, possibly here, as one of the more challenging cases for insect immune systems. Or maybe they came up in a discussion on SF stories taking inspiration from real life.
Delightful creatures. Mother Nature doing her thing on Mrs Grundy. Are textbooks about these banned in Tennessee? Yet?
Strepsipterans are awesome! Traumatic insemination occurs in bedbugs and their relatives, among other insects, so isn’t unique to them, though — all of the information on the immune implications comes from work on bedbugs, I think. The females have an organ that buffers some of the effects of piercing the exoskeleton, I believe . . .
Nature is astounding. Thanks for sharing.
You are very welcome.
Thank you for this! My daughter told me about these a few years ago. I can not wait until my kids are done with school, and I can show them that this and (especially) the video. The writing style and the subjects and photographs of the latest posts on insects are so good. I loved this one.
Thank you for the kind words, Desirée. Perhaps your daughter is a budding entomologist?
Incredible creatures, I wonder what god was thinking when he made them!
Obviously, he was thinking of ways to make bees suffer. Pretty devious, he be. Just ask Job.
Evolutionary marvel indeed. Thanks for your time in writing this essay, very much appreciated.
Thankyou for a fascinating (and strangely horrifying) article. After ‘flu and covid and tapeworms, you’ve found something new that I’m now terrified is going to make the leap across to humans….
🙂