> Habitat & Distribution

Populations of Fontigens benfieldi inhabit streams draining caves of the Wabash Spring Basin, a tributary of the New River in Giles County, VA.  Two populations have been documented: one in the main stream of the Sugar Run Cave System and a second in the main stream of Dulaney's Cave.  The snails are found grazing on and under rocks and cobbles.  A broad-brush review of the zoogeography of North American cavesnails has been offered by Hershler & Holsinger (1990), and a more detailed review for the genus Fontigens by Gladstone and colleagues (2021).  FWGNA incidence unranked.

> Ecology & Life History

Fontigens benfieldi is apparently stygobiontic, entirely unpigmented and blind, known only from limestone caves.  The waters of such caves are (of course) rich in minerals, but poor in organics.  Presumably the entire ecosystem is based on fine organic matter raining down from the lighted world above.

Weck (2022) reported approximately one year to maturity (3 mm) in laboratory populations of the cave-dwelling Fontigens antroecetes, constant low levels of reproduction (25 - 80 eggs/pair/yr), and a lengthy hatch time of 70 - 80 days.

> Taxonomy & Systematics

This species was among three newly described by Dillon and colleagues in an appendix to Volume 5 of the FWGNA series (Dillon et al. 2023).  It is a member of the "orolibas group" of Hershler et al. (1990), bearing a tripartite penis with a tubular distal lobe and a bulbous proximal lobe.  It may be distinguished from all other species of Fontigens by its unique ovoid or fusoid shell and a minimum CO1 sequence divergence of 9.2% from all previously -sequenced species and 9.1% from Fontigens davisi, with which it co-occurs, from which it appears to be reproductively isolated.

A survey of CO1 sequence diversity across 13 populations of 9 Fontigens species was offered by Liu and colleagues (2021).  See my essay of 9Aug22 from the link below for a review.

Taylor (1966) suggested that Fontigens comprises a distinct hydrobiid subfamily, the Fontigentinae, which Hershler et al. (1990) synonymized under the Emmericiinae of Brusina (1870).  Wilke et al. (2013) did not confirm a close association between Fontigens (represented by a single sample) and the European genus Emmericia, however, tentatively returning the Fontigentinae to subfamilial status.  Despite this evidence, self-appointed experts insisted on placing Fontigens in the Emmericiidae for several years (Bouchet et al. 2017), until Gladstone & Whelan (2022) split the genus to its own separate family, the Fontigentidae. 

> Supplementary Resources

• Montage of Fontigens benfieldi shells, from Dillon et al. (2023).
  

> Essays

• Earlier versions of this website, online until August of 2016, adopted the large, broadly-inclusive concept of the Hydrobiidae (sl) following Kabat & Hershler (1993).  More recently the FWGNA project has shifted to the Wilke et al. (2013) classification system, distinguishing a much smaller Hydrobiidae (ss) and elevating many hydrobioid taxa previously ranked as subfamilies to the full family level.  For more details, see The Classification of the Hydrobioids.
• See my essay of 9Aug22, Startled by Fontigens, sort-of, I suppose for a review of the paper by Liu et al. (2021) documenting unusually high levels of intraspecific mtDNA sequence divergence.
• I shared the backstory behind the discovery and description of Fontigens hershleri, F. benfieldi, and F. davisi in my essay of 9Jan24, Three New Fontigens From Virginia. Additional photos showing shell variation are also available.

> References

Bouchet, P., J. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nutzel, P. Parkhaev, M. Schrodl, and E. Strong (2017) Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia, 61: 1–526.
Dillon, R.T., Jr., T.E. Malabad, W.D. Orndorff & H-P. Liu (2023) Three new Fontigens (Caenogastropoda: Fontigentidae) from caves in the Appalachian Ridge and Valley Province, Virginia. Pp. 283 - 306 in Dillon, R.T., Jr. et al. The Freshwater Gastropods of North America Volume V: Ohio, Cumberland, and Tennessee River Systems. FWGNA Press, Charleston. [pdf]
Gladstone, N.S., E. Pieper, S. Keenan, A. Paterson, M. Slay, K. Dooley, A. Engel, and M. Niemiller (2021)  Discovery of the Blue Ridge springsnail, Fontigens orolibas Hubricht 1957 (Gastropoda: Emmericiidae) in East Tennessee and its conservation implications.  Freshwater Mollusk Biology and Conservation 24: 34 - 42.
Gladstone, N. S. and N. Whelan (2022) Pushing barcodes to their limits: phylogenetic placeament of Fontigens Pilsbry, 1933 (Caenogasatropoda: Littorinimorpha: Truncatelloidea) and elevation of Fontigentidae Taylor, 1966.  Journal of Molluscan Studies 88: eyab038.
Hershler, R. H. & J. R. Holsinger (1990)  Zoogeography of North American hydrobiid cavesnails.  Stygologia 5: 5-16.
Hershler, R., J.R. Holsinger & L. Hubricht (1990) A revision of the North American freshwater snail genus Fontigens (Prosobranchia: Hydrobiidae). Smithsonian Contributions to Zoology 509:1-49,  
Hubricht, L. (1963)  New species of Hydrobiidae.  Nautilus 76: 138 - 140.
Hubricht, L. (1976)  The genus Fontigens from Appalachian caves (Hydrobiidae: Mesogastropoda).  Nautilus 90:86 - 88.
Kabat, A.R., and R. Hershler (1993) The prosobranch snail family Hydrobiidae (Gastropoda: Rissooidea): review of classification and supraspecific taxa. Smithsonian Contributions to Zoology 547:1-94. 
Liu, H-P., L. Schroeder, A. Berry, and R. T. Dillon, Jr. (2021)  High levels of mitochondrial DNA sequence divergence among isolated populations of Fontigens (Truncatelloidea: Emmericiidae) in eastern USA.  Journal of Molluscan Studies 87:  eyab026. 
Taylor, D.W. (1966) Summary of North American Blancan nonmarine molluscs.   Malacologia  4: 1 - 172.
Weck, R.G. (2022)  Life history observations of the Illinois state endangered Enigmatic Cavesnail, Fontigens antroecetes (Hubricht, 1940) made under simulated cave conditions.  Subterranean Biology 43: 185 - 198. 
Wilke T., Haase M., Hershler R., Liu H-P., Misof B., Ponder W. (2013)  Pushing short DNA fragments to the limit: Phylogenetic relationships of “hydrobioid” gastropods (Caenogastropoda: Rissooidea).  Molecular Phylogenetics and Evolution 66: 715 – 736.