Field of Science

Biantidae: The Importance of Titillators

An unidentified member of Biantinae, photographed in Singapore by WJ.

Long-time readers of this site will be familiar with my rants about the influence of early 20th-century arachnologist Carl-Friedrich Roewer on Opiliones taxonomy (just enter 'Roewer' into the search box near top right on this page). One of Roewer's larger errors (but an understandable one in context) involved the species he included in the family Phalangodidae. As Roewer had it, this was an almost cosmopolitan family, with representatives on all continents. However, as research has progressed, it has become clear that Roewer's Phalangodidae was a polyphyletic assemblage of a number of different lineages of relatively generic-looking Laniatores (short-legged harvestmen). Firm distinction of some of these lineages (now recognised as separate families) often requires examination of the male genitalia, something Roewer never did.

Specimen of Lacurbs, from A. B. Kury.

The Biantidae are one of these families of ex-phalangodids. They are a pantropical family, with representatives in South America, Africa and Asia. Biantids have eyes well-separated on the carapace instead of on a common central eye-mound, and they bear a strong external resemblance to another family of Laniatores that I've covered before, the South American Stygnidae. Distinguishing biantids from stygnids depends on two features: the presence of a process of the tarsi of the third and fourth legs in Stygnidae, and the presence of a ventral sclerotised plate on the penis of the Stygnidae versus no plate and dorsal processes called titillators (named for an obvious possible function) on the penis of Biantidae. However, despite the strong external similarity, stygnids and biantids are not closely related: a recent molecular phylogenetic analysis of Laniatores places them in separate superfamilies, with stygnids in the Gonyleptoidea and biantids in the Samooidea (Sharma & Giribet 2011).

Representative biantid penes, from Kury & Pérez González (2007). Left: Caribbiantes (Stenostygninae), dorsolateral view; right: Biantes sherpa (Biantinae), lateral and dorsal views. Ti = titillators; Co = conductors.

Biantidae are divided between four subfamilies (Kury & Pérez González 2007). The African genera Lacurbs and Zairebiantes are more divergent than the other two subfamilies: Lacurbs has the scutum (dorsal shield) of the opisthosoma (the rear part of the body) widest in the middle and narrowing towards the front and back, while other biantids are more or less straight-sided, and has the tibiae and metatarsi of the hind legs armed and swollen. Zairebiantes has its eyes placed closer together and further forward than other biantids, and Pinto-da-Rocha (1995) suggested that its classification as a biantid may require re-evaluation. The other two subfamilies, the South American Stenostygninae and the African and Asian Biantinae, contain the great majority of biantids and share the presence of dense scopulae (pads of hairs) on the third and fourth tarsi. Apart from their distribution, the latter two subfamilies are distinguished by genital morphology: in Stenostygninae, the titillators are rigid and sit forward to cover the capsula interna of the penis, while in Biantinae they are soft and fold back and out so that they don't cover the capsula interna.


Kury, A. B., & Pérez González, A. 2007. Biantidae Thorell, 1889. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds) pp. 176-179. Harvard University Press: Cambridge (Massachusetts).

Pinto-da-Rocha, R. 1995. Redescription of Stenostygnus pusio Simon and synonymy of Caribbiantinae with Stenostygninae (Opiliones: Laniatores, Biantidae). Journal of Arachnology 23 (3): 194-198.

Sharma, P. P., & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106-142.

1 comment:

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