Caliphyllidae Tiberi, 1881

Moreno, Karina, Medrano, Sabrina, Gosliner, Terrence M., Wilson, Nerida G., Krug, Patrick J. & Valdés, Ángel, 2025, Phylogenetic systematics of the genus Cyerce Bergh, 1870 (Mollusca: Heterobranchia: Sacoglossa: Caliphyllidae) from the Pacific and Indian oceans with descriptions of nine new species, Zoological Journal of the Linnean Society 204 (1) : -

Family Caliphyllidae Tiberi, 1881

Genus Cyerce Bergh, 1870

Cyerce Bergh, 1870 : pl. 1, figs 9, 10; 1871: 98. Type species: Cyerce elegans Bergh, 1870 , by subsequent designation by Swennen (1961).

Lobiancoia Trinchese, 1881: 116 . Type species: Lobiancoia cristallina Trinchese, 1881 [= Cyerce cristallina ( Trinchese, 1881) ], by monotypy.

Diagnosis

Foot divided by transverse groove. Body oval, wider anteriorly, tapering posteriorly. Pericardium elevated, oval, papillate, or smooth. Anal papillae antero-medially situated between right rhinophore and pericardium. Head with bifurcated rhinophores, and ventral, enrolled oral tentacles shorter than rhinophores. Eyespots at base of rhinophores. Several dorsal leaf-like cerata along both sides of body, lacking tubules of the digestive gland; cerata able to autotomize. Cerata morphology variable, from bold to light coloration. Pharyngeal pouch attached to buccal bulb, variable in size. Radula with single row of teeth, constituting a descending and ascending limb, ascus at the base of descending limb. Ascus with several used teeth of variable sizes. Radular teeth morphology variable from elongate and slim to short and moderately wide. Teeth with two rows of denticles along either side. Penis with stylet at opening.

Remarks

As indicated by Yonow and Jensen (2018), Bergh (1870) introduced the genus name Cyerce in the legend of a plate ( Bergh 1870: pl. 1) and later provided a full description ( Bergh 1871). Two new species were originally included in Cyerce : Cyerce nigra Bergh 1870 and Cyerce elegans Bergh 1870 , both collected from Palau. Bergh (1870; 1871) did not specify the type species, but Swennen (1961) indicated that C. elegans is the type, and with this act C. elegans became the type by subsequent designation. Bergh (1871) described the genus Cyerce as including specimens with a dorsal anus, armed penis, short ‘crop’ [pharyngeal pouch], elongated radular teeth, a foot with a transverse groove, and cerata without ‘tubules’ [branches] of the digestive gland. A few years earlier, Pease (1866) had described the new species Lobifera nigricans ( Pease, 1866) from the Pacific Islands, which resembles the specimen of Cyerce nigra described by Bergh (1870; 1871) but displays some differences. The genus name Lobifera , also described by Pease (1866), is an unnecessary replacement name for Polybranchia Pease, 1860 (see: Swennen 1961 and, Medrano et al. 2019). Although the description of Lobifera nigricans by Pease (1866) was short and lacking details of the internal anatomy, external morphological traits such as the presence of a transverse groove across the foot and the presence of fan-like cerata suggests this species belongs to Cyerce . Comparisons of the external morphology of Cyerce nigricans Pease 1866 and Cyerce nigra Bergh 1870 are further elaborated under the remarks of Cyerce nigricans Pease 1866 (see: below).

Trinchese (1881) introduced the genus Lobiancoia for the new species, Lobiancoia cristallina collected from the Mediterranean Sea. Trinchese (1881) described Lobiancoia cristallina as having bifid rhinophores, head angled with ‘auriform tentacles’, a transverse groove dividing the foot, compressed dorsal papillae along both sides of the animal without ‘lobes of the liver’ or ‘cnidosacs’, and an anal papilla located to the right and front of the pericardial hump. Pelseneer (1892) reported for the first time the genus Cyerce from the Mediterranean Sea with the description of the new species Cyerce jheringi Pelseneer, 1892 . Pruvot-Fol (1954) regarded Lobiancoia as a valid genus, different from Cyerce which she considered a synonym of Lobifera . Portmann (1958) studied a specimen from the Mediterranean Sea he identified as L. cristallina and suggested that C. jheringi and L. cristallina belonged to the same species, effectively synonymizing Lobiancoia with Cyerce . However, Portmann (1958) maintained this species in the genus Lobifera , following the taxonomic revision by Pruvot-Fol (1954). Swennen (1961) clarified the taxonomic position of Lobifera as a synonym of Polybranchia and corroborated the synonymy of Lobiancoia with Cyerce , which is widely accepted in modern literature. Moreover, the monophyly of Cyerce has been supported by morphological ( Jensen 1996) and some molecular ( Christa et al. 2015) studies. Other molecular studies that included the closely related genus Sohgenia (which lacks a transverse foot groove) did not resolve whether Cyerce was monophyletic ( Krug et al. 2015). Here, Cyerce Bergh 1870 is regarded as a valid name for species of Sacoglossa with a transverse groove ventrally along the foot, anal papillae antero-medially between the right rhinophore and pericardium, bifurcated rhinophores, enrolled oral tentacles, cerata without tubules of the digestive gland, and a penis with a penial stylet.

Clade 1