Geitodoris, BERGH, 1891

Valdés, Ángel, 2002, A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia), Zoological Journal of the Linnean Society 136 (4), pp. 535-636 : 599

GENUS GEITODORIS BERGH, 1891

Geitodoris Bergh, 1891: 130 . Type species: Doris complanata Verrill, 1880 , by monotypy.

Carryodoris Vayssière, 1919: 67 . Type species: Carryodoris joubini Vayssière, 1919 , by original designation.

Verrillia Ortea & Ballesteros, 1981: 341 . Type species Geitodoris bonosi Ortea & Ballesteros, 1981 , by monotypy.

Diagnosis

Dorsum covered with simple tubercles, stiffened by integumentary spicules, which occasionally protrude from the dorsal surface in an irregular fashion. Head with two conical oral tentacles. Anterior border of the foot grooved and notched. Labial armature armed with jaw elements. Radula composed of hamate teeth, occasionally denticulate. Outermost lateral teeth multidenticulate. Reproductive system with a flattened, granular prostate, having two well differentiated regions. Penis and vagina devoid of hooks. There is a peduculate accessory gland, in some species armed with several copulatory hard structures.

Remarks

Bergh (1891) introduced the genus Geitodoris based on Doris complanata Verrill, 1880 , type species by monotypy, with a very short Latin description. According to Bergh (1891) Geitodoris is characterized by having the labium of the anterior border of the foot notched in the middle, the inner lateral teeth strong, hamate and outermost slender, multidenticulate, and by lacking a differentiated prostate. This diagnosis was based on Verrill’s (1880) original description of Doris complanata , rather than on newly examined specimens. Bergh (1894) completed the description of Geitodoris with anatomical studies based on one of Verrill’s original specimens.

Eliot (1906b) considered Geitodoris to be similar to Rostanga and also closely related to some archidorids, such as Archidoris stellifera Vayssière, 1904 . In contrast, Odhner (1926) speculated that Geitodoris is closely related to Discodoris , and is distinguished from it by the peculiar form of the outer radular teeth. He also noted other diagnostic characteristics of Geitodoris : the unarmed penis, absence of prostate and stomach, presence of jaws, anterior border of the foot notched and finger-like oral tentacles.

Vayssière (1919) described the genus Carryodoris for the new species Carryodoris joubini Vayssière, 1919 . In his description he did not mention Geitodoris or refer to the papers by Bergh (1891), Eliot (1906b) or Odhner (1926). Carryodoris was characterized by the presence of jaws with small rodlets and a radula with spatula-shaped outermost lateral teeth. Other features of this genus are the anterior border of the foot notched, perfoliate rhinophores and tripinnate branchial leaves.

Schmekel (1973) described a new species of Carryodoris from the Mediterranean, and considered this genus to be distinct from Geitodoris . She based her conclusion on two major differences between these two taxa, the absence of a differentiated prostate in Geitodoris , which is present in Carryodoris , and the absence of denticles on the outermost lateral teeth of Geitodoris , also present in Carryodoris . She also transferred Geitodoris ohshimai Baba, 1926 to this genus.

Ortea & Ballesteros (1981) regarded Carryodoris as a subgenus of Geitodoris . According to these authors, the name Geitodoris should be used for G. complanata and other species with smooth lateral teeth and lacking a differentiated prostate and Carryodoris for species with denticulate lateral teeth an a differentiated prostate. In addition, Ortea & Ballesteros (1981) described the new subgenus Verrillia for Geitodoris bonosi , which has smooth lateral teeth and a differentiated prostate. Other authors ( Perrone, 1984; Cervera, García-Gómez & García, 1985; Miller, 1996) followed this classification including three different subgenera, in subsequent papers. Martínez, Ortea & Ballesteros et al. (1996) considered that the presence of denticles on the lateral teeth of Geitodoris ‘should be considered as a specific character rather than a generic one’, but at the same time continued using the same classification.

An anatomical study of G. complanata , the type species of Geitodoris , shows that this species has a well developed prostate and therefore there are virtually no differences between Geitodoris and Verrillia . I agree with most of the authors mentioned above in that the presence of denticles in some mid-lateral teeth should not on its own be used to separate the two genera, and thus I regard Carryodoris as a synonym of Geitodoris .