Hoplodoris, BERGH, 1880

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 : 591

GENUS HOPLODORIS BERGH, 1880

Hoplodoris Bergh, 1880: 51 . Type species: Hoplodoris desmoparypha Bergh, 1880 , by monotypy.

Diagnosis

Dorsum covered with simple, large and rounded tubercles, stiffened by integumentary spicules. Head with two conical oral tentacles. Anterior border of the foot grooved and notched. Labial armature armed with jaw elements. Radula composed of simple, hamate teeth, occasionally denticulate. Reproductive system with a flattened, granular prostate, having two well differentiated regions. Penis armed with hooks. Vagina devoid of armature. One or two large and pedunculated accessory glands armed with copulatory spines.

Remarks

Bergh (1880) described the genus Hoplodoris based on Hoplodoris desmoparypha Bergh, 1880 , the type species by monotypy. The genus Hoplodoris is characterized by having the dorsum covered with long tubercles, the anterior border of the foot grooved and notched, presence of jaws, hamate radular teeth, large prostate, penis armed with hooks and presence of an accessory gland with a spine. The type species of Hoplodoris has not been collected since, and there is no information on its external morphology. Unfortunately, the type material of this species collected from Palau could not be located at ZMUC and is presumed lost. The information for this genus used in the phylogenetic analysis has been obtained from Hoplodoris novaezelandiae ( Bergh, 1904) .

Thompson (1975) regarded Carminodoris as a junior synonym of Hoplodoris based on his description of Hoplodoris nodulosa Angas, 1864 from Australia. However, the original description of Carminodoris ( Bergh, 1889) , based on Carminodoris mauritiana , states that this genus is characterized by having the anterior border of the foot grooved and notched, the dorsum covered with small tubercles, presence of jaws, hamate lateral teeth, denticulate outermost lateral teeth, large prostate and penis armed with hooks. As mentioned below, it is probable that Carminodoris , which lacks accessory glands and has small dorsal tubercles, is a synonym of Discodoris , but this point needs confirmation.

Other genera having accessory glands with spines are Asteronotus Ehrenberg, 1831 ; Jorunna Bergh, 1876 and Paradoris Bergh, 1884 . According to the results of the phylogenetic analysis (see below), all these taxa belong to different clades, and it is very likely that they acquired the copulatory spines independently.

Several species have been added to the genus Hoplodoris since its original description. Burn (1969) transferred Doris nodulosa Angas, 1864 , Miller (1991) transferred Homoiodoris novaezelandiae Bergh, 1904 and Gosliner & Behrens (1998) described the new species Hoplodoris estrelyado Gosliner & Behrens, 1998 . Some Indo-Pacific species previously assigned to the genus Carminodoris Bergh, 1889 , should also probably be transferred to Hoplodoris (see Gosliner & Behrens, 1998). Gosliner & Behrens (1998) described some variation within Hoplodoris to accommodate species with one or two accessory glands armed or unarmed with spines. All the species included in Hoplodoris are characterized by having rounded dorsal tubercles, except for Hoplodoris desmoparypha , which has elongate tubercles.