In a glass terrarium in Bratislava, a female harvestman with a male on top of her rotates downward, puts her mouthparts on his penis, and by doing so brings the encounter to an early end. Twenty three of the thirty five females in the experiment did it at least once, and the ones who did it in the first pairing tended to do it again in the second, which makes it a strategy rather than an accident.
Scientific American covered the finding on Monday. The study is sound and the reporting is fair, but the emphasis lands on the novelty. What reads as a curiosity is the visible edge of a sexual arms race that somebody has now measured, on both sides, in units of force. What follows is the study, one correction to the coverage, and the considerably stranger literature sitting a single citation away.
Both papers come from Pavol Prokop's group at Comenius University Bratislava. The first, in Scientific Reports in 2024, worked with Phalangium opilio, a harvestman common across North America and Europe and probably resident on your porch. The team collected roughly 230 juveniles, raised them to adulthood in isolation so every female started as a virgin, and then ran twenty minute pairings in glass terrariums.
The behavior showed up in 31 of 70 trials. Females that did it in the first pairing were significantly more likely to do it in the second (Fisher exact, p = 0.007), so this is a strategy some individuals run rather than a random twitch. Sixty eight percent of the episodes came after copulation. Prokop's framing is that the female assumes a position that puts her mouthparts on the penis and her genital opening out of reach, which is a fairly efficient way to end a conversation.
The quote"We already knew that some animals perform fellatio, like bats and bonobos. What we didn't know, and haven't seen in previous studies, is that females might use it to get out of engaging in intercourse."
Forty four percent of trials included the behavior. The proportion of individual females who did it at least once was 23 of 35, or 66 percent.
What the coverage saysScientific American writes "about 44 percent of the females licked the males' penises." Small slip, wrong denominator, and it happens to understate the finding. Two thirds of the females in that room had this move available to them.
Females showed resistance in 91 percent of trials (only 7 of 78 passed without it). Males bit females on the legs and body as a matter of routine, pursued up to six copulations in a single twenty minute window, and killed and partially ate four females. Two more lost limbs. Prokop, on the males who eat the females who refuse them, offers "it's a very pragmatic decision," which is one word for it.
The caveat the coverage skipsThis is a small glass box with no exit. Anita Aisenberg, the Uruguayan evolutionary biologist quoted in the piece, suggests the aggression may be spillover from the propensity of males to fight each other. A confinement effect is the obvious competing explanation and the paper cannot rule it out. Sexual conflict in P. opilio is real. The 5 percent cannibalism rate is a number from a terrarium.
The second study, published in Ethology in May, used Leiobunum limbatum. Males of this species carry a pair of sacs on the penis filled with a nonsperm secretion, and females lick them routinely. Females stayed receptive across sequential matings with different partners, with no drop off. Females in better body condition mated more often. Larger males got fewer copulations, which the authors read as an advantage to small size under scramble competition. Their conclusion is that coercion here "seems to be lower than in other species, particularly those lacking nuptial feeding."
Why it mattersSame order, same basic anatomy, opposite social contract. That contrast is the actual finding, and it is where the interesting literature starts.
One more result the coverage left out, because it complicates the story. Females that performed fellatio produced fewer eggs. Prokop's group reads the behavior as a cost of sexual conflict rather than a benefit the female extracts, and they explicitly reject the nuptial feeding explanation for this species. So the tidy version, in which the female trades a favor for a meal and walks away ahead, is not what the data show. She is paying to get out of something worse.
Here is the thread that runs under all of it, and it belongs mostly to Mercedes Burns at UMBC, who is an author on essentially every paper worth reading in this field.
Harvestmen in the eastern North American Leiobunum group come in two anatomical arrangements. Sacculate males have those gift sacs on the penis and open with a face to face embrace, handing the secretion directly to the female's mouth. Nonsacculate males have lost the sacs (though they kept the glands that fill them) and have instead evolved femoral hooks on the pedipalps that clamp the female's front legs and hold her still.
Burns, Hedin and Shultz mapped this across 29 species in 2013. The ancestral condition is gift giving males and unarmored females. Then, on at least four separate branches, the same thing happens in the same order. Males lose the gift sacs and gain the clamps. Females gain a sclerotized pregenital barrier, a plate of hardened cuticle that mechanically obstructs forced entry. The losses and the gains co-occur. Persuasion is replaced by leverage, and leverage is answered with armor.
Two years later the same group went and measured the forces. They dissected muscle cross sections, pennation angles, lever mechanics and cuticle mass across 28 species, and computed how hard a male can protract his penis and how hard a female can hold her operculum shut. The two scale together, R² = 0.482, p < 0.0001. This is a sexual arms race with units.
The 2015 paper also delivers the finding that ruins the clean version. The species do not fall into two camps. They distribute along a continuum, and the analysis failed to partition them by strategy at all. So the romantics and the brutes are the two ends of a gradient, and most harvestmen live in the middle, negotiating.
What is in the gift. Kahn and colleagues ran gas chromatography and mass spectrometry on the accessory glands and penial sacs of five Leiobunum species in 2018. They found eleven free amino acids, with alanine, threonine, isoleucine, glutamine and serine making up 91 percent of the total. The interesting part is the comparison. Gift giving species deliver a higher proportion of essential amino acids than their clamping relatives (0.392 against 0.330, a relative difference of about 19 percent). Chemistry tracked mating behavior rather than phylogeny, meaning it evolved convergently.
Put plainly, the males who have to persuade serve a better meal. Two caveats worth keeping. The study measured free amino acids only, not proteins or lipids or sugars, and "essential" was defined off the vertebrate list because nobody has worked out which amino acids are essential to a harvestman.
The myth is durable enough to have its own dedicated debunking pages at two universities. The claim is that daddy longlegs carry the most powerful venom in the world but have fangs too short to break human skin. Rod Crawford at the Burke Museum calls it "a full-fledged Urban Legend, with no basis in fact whatever," and the sentence does a lot of work, because the myth is wrong twice in three different animals.
Three unrelated creatures answer to the name. Harvestmen (Opiliones) have no venom glands, no fangs, no silk and one apparent body segment. They tear up solid food and eat it, which no spider does. Cellar spiders (Pholcidae) are actual spiders and do have venom. Crane flies (Tipulidae) are insects, and many adults have no functional mouthparts at all.
The pholcid is the only candidate that could in principle be dangerous, so somebody checked. Zobel-Thropp and colleagues sequenced and assayed the venom of Physocyclus mexicanus in 2019 and found it made mostly of metalloproteinases and peptide neurotoxins, with an effective paralytic dose in crickets of 3.9 micrograms per gram. That is unremarkable among spiders. Rick Vetter at UC Riverside is on record that he is aware of no publication showing any toxic effect of pholcid venom in humans.
My favorite detail in the whole file is that the 2019 venom proteomics paper, in a peer reviewed journal, cites MythBusters. The show tested it in 2004, verdict Busted, and the spider did bite Adam Savage. He reported a mild burning sensation that went away.
- Their sperm has no tail. Harvestman spermatozoa are aflagellate and immobile, which means whatever movement happens inside the female tract is probably being managed by the female. Machado and Burns call this one of the order's genuinely unique traits, and it gets a fraction of the attention the penis does.
- The anatomy is at least 400 million years old. Devonian harvestmen from the Rhynie chert in Scotland preserve a penis and an ovipositor, and the animals look startlingly modern. A 99 million year old specimen in Burmese amber, Halitherses grimaldii, is preserved with the organ fully everted. The popular coverage called it a 99 million year erection. There is no erectile tissue involved, but I understand the temptation.
- Precopulatory cannibalism has never been recorded in the order. Not once, in any suborder. Female spiders eat males before mating often enough that male spiders evolved elaborate vibratory courtship largely as a safe conduct pass. Harvestmen skipped the display because they never needed the pass.
- They cannot regrow a lost leg. They compensate instead. Prionostemma harvestmen that lose two or three legs take an immediate hit to speed and acceleration, then recover full performance in about two days by changing gait and pressing their sensory legs into walking duty. That is the fastest locomotor recovery recorded in any animal that sheds limbs.
- Losing a leg as a juvenile can determine which kind of adult male you become. In the New Zealand species Forsteropsalis pureora, males come in three morphs. Juveniles with at least one autotomy scar were 45 times more likely to mature into the small sneaker morph. One bad encounter with a predator in adolescence and the whole adult reproductive strategy changes. The study is correlational and the authors are careful about it.
- You can read their phylogeny off the smell. With no venom to work with, harvestmen went in for defensive chemistry. Naphthoquinones are ancestral, benzoquinones evolved later and independently at least twice, and one Costa Rican species does not synthesize its compound at all. It sequesters myrmicacin from the leafcutter ants it eats.
The headline treats this as a novelty item. What the literature actually contains is a conflict between the sexes that has been running for a very long time and that somebody has now measured in units of force, on both sides, and found to be evenly matched.
The honest part is the ending. Machado and Burns, reviewing the whole field in 2024, note that most of what we believe about how harvestman genitalia interact "can be regarded as educated guesses or hypotheses that need to be tested," and that exactly one paternity study has ever been done in the order. Roughly 6,700 described species, four hundred million years of anatomy, an arms race with a regression coefficient, and one paternity test.
Sources
The article: Christa Lesté-Lasserre, "Daddy long leg sex is weirder than you might imagine," edited by Andrea Thompson, Scientific American, 3 August 2026. scientificamerican.com. The page headline and the metadata title are not the same. Search engines and social cards get "Daddy Long Legs Perform Fellatio and Other Wild Facts about How These Arachnids Mate," which is also the URL slug. Readers get the milder one.
Study one: Prokop P, Litavský J, Provazník Z. "Female Phalangium opilio use fellatio to compensate sexual avoidance." Scientific Reports 14:25586, 2024. doi.org/10.1038/s41598-024-77209-9
Study two: Prokop P, Litavský J, Purkart A, Balcerčík J. "Mating Status Does Not Alter Sexual Receptivity in Females of the Harvestman Leiobunum limbatum." Ethology 132(8), 2026. doi.org/10.1111/eth.70080
The arms race: Burns M, Hedin M, Shultz JW. "Comparative analyses of reproductive structures in harvestmen reveal multiple transitions from courtship to precopulatory antagonism." PLOS ONE 8(6):e66767, 2013. doi.org/10.1371/journal.pone.0066767
The measured forces: Burns M, Shultz JW. "Biomechanical diversity of mating structures among harvestmen species is consistent with a spectrum of precopulatory strategies." PLOS ONE 10(9):e0137181, 2015. doi.org/10.1371/journal.pone.0137181
Gift chemistry: Kahn PC, Cao DD, Burns M, Boyer SL. "Nuptial gift chemistry reveals convergent evolution correlated with antagonism in mating systems of harvestmen." Ecology and Evolution 8(14):7103, 2018. doi.org/10.1002/ece3.4232
The field review: Machado G, Burns M. "Reproductive biology of harvestmen (Arachnida: Opiliones): a review of a rapidly evolving research field." Current Zoology 70(1):115, 2024. doi.org/10.1093/cz/zoac102
Venom myth: Rod Crawford, Burke Museum Spider Myths. burkemuseum.org · UC Riverside Spider Research. spiders.ucr.edu · Rick Vetter interview, UCR Entomology, 2022. entomology.ucr.edu
Pholcid venom assay: Zobel-Thropp PA et al. "Not so dangerous after all? Venom composition and potency of the pholcid (daddy long-leg) spider Physocyclus mexicanus." Frontiers in Ecology and Evolution 7:256, 2019. doi.org/10.3389/fevo.2019.00256
Devonian fossils: Dunlop JA, Anderson LI, Kerp H, Hass H. "A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts." Nature 425:916, 2003. doi.org/10.1038/425916a
Amber specimen: Dunlop JA, Selden PA, Giribet G. "Penis morphology in a Burmese amber harvestman." The Science of Nature 103:11, 2016. doi.org/10.1007/s00114-016-1337-4
Leg loss and recovery: Escalante I, Badger MA, Elias DO. "Rapid recovery of locomotor performance after leg loss in harvestmen." Scientific Reports 10:13747, 2020. doi.org/10.1038/s41598-020-70557-2
Juvenile leg loss and male morph: Powell EC, Painting CJ, Machado G, Holwell GI. Behavioral Ecology 34:613, 2023. doi.org/10.1093/beheco/arad029
Defensive chemistry: Raspotnig G et al. "Chemosystematics in the Opiliones." Frontiers in Ecology and Evolution 5:139, 2017. doi.org/10.3389/fevo.2017.00139 · Raspotnig G et al. Chemoecology 32:139, 2022. doi.org/10.1007/s00049-022-00373-9