A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia) Author Valdés, Ángel text Zoological Journal of the Linnean Society 2002 2002-12-31 136 4 535 636 https://academic.oup.com/zoolinnean/article-lookup/doi/10.1046/j.1096-3642.2002.00039.x journal article 10.1046/j.1096-3642.2002.00039.x 8acc9095-eaff-47d7-b3da-91b6c2fb636e 0024-4082 4634200 PARADORIS INDECORA ( BERGH, 1881 ) ( FIGS 34A , 35 , 36 ) Discodoris indecora Bergh, 1881: 108–112 , pl. J, figs 26–33, pl. K, figs 11–19. Paradoris granulata Bergh, 1884a: 686–691 , pl. 76, figs 10–24. Paradoris granulata var. Bergh, 1884a: 691–693 , pl. 77, figs 25–32. Type material Discodoris indecora Bergh . The original type material, collected from Trieste , Italy , is lost. Paradoris granulata Bergh, 1884 . LECTOTYPE (here selected): Trieste , Italy , April–May 1979 – 80, 24 mm preserved length, leg. Graeffe ( ZMUC GAS-2120 ); PARALEC TOTYPES: Trieste, Italy , April–May 1979 –80, six specimens, 8–24 mm preserved length, leg. Graeffe ( ZMUC ). Two other specimens labelled as Paradoris granulata var. belong to the same species. They were probably collected from Trieste, Italy , April–May 1979 –80, 11– 19 mm preserved length, leg. Graeffe ( ZMUC GAS- 2121). Additional material Cabo de Palos , Murcia , Spain , 4 August 1984 , one specimen, 14 mm preserved length, leg. J. Templado ( MNCN 15.05 /18231) . External morphology The general colour of the living animals is uniformly pale grey with a pale brown tinge in the centre of the dorsum ( Fig. 34A ). There are several dark brown spots on the tips of the larger tubercles, also associated with groups of small opaque white dots. The rhinophores are dark brown with the apex and some spots opaque white. The gill is pale grey with the apices of the leaves bright yellow and dark brown. The whole dorsum is covered with small, rounded tubercles ( Fig. 35E ). The largest tubercles occur in two lines running from the rhinophores to the gill. The rhinophoral and branchial sheaths have tubercles no different from those on the rest of the dorsum. There are eight tripinnate branchial leaves, forming a circle. The anal papilla is situated in the centre of the branchial circle of leaves. The rhinophores are elongate, having 14 lamellae in a 24-mm preserved length specimen. Ventrally there are two long and conical oral tentacles ( Fig. 36E ). The tentacles are grooved longitudinally. The anterior border of the foot is grooved and notched. Anatomy The posterior end of the glandular portion of the oral tube has six strong retractor muscles ( Fig. 36C ) which attach to the body wall. The oval, muscular buccal bulb has two additional muscles attached; two long and thin salivary glands connect with it at each side of the oesophageal junction. The buccal bulb is half the length of the glandular portion of the oral tube. The labial cuticle has two areas covered with a number of simple rodlets ( Fig. 35D ). The radular formula is 20 ¥ 22.0. 22 in a 24-mm long specimen. Rachidian teeth are absent. The lateral teeth are narrow and elongate, having a single cusp and lacking denticles ( Fig. 35A ). The teeth from the middle portion of the half-row are larger than those closer to the medial portion of the radula, and they have a short and strong cusp compared to the base ( Fig. 35B ). The outermost teeth are smaller and also lack denticles ( Fig. 35C ). Some of them completely lack a cusp. The oesophagus is long and connects directly to the stomach ( Fig. 36A ). Figure 34. Living animals. A, Paradoris indecora , Southern Spain, photo by D. Moreno. B, Otinodoris sp. (CASIZ 073238), photo by T. M. Gosliner. C, Sebadoris nubilosa , Seychelles, photo by T. M. Gosliner. D, Conualevia marcusi , La Paz Bay, Baja California Sur. The ampulla is very long and convoluted ( Fig. 36C ). It branches into a short oviduct and the prostate. The oviduct enters the female gland mass near to its centre. The prostate is tubular and has two portions distinguishable by their colour and texture ( Fig. 36B ). It connects with a long duct that narrows and expands again into the large ejaculatory portion of the deferent duct. The muscular deferent duct opens into a short common atrium with the vagina. Connected to the atrium there us a large, ramified accessory gland and two muscular sacs each containing a rigid spine. The vagina is long and thin. At its proximal end it joins the bursa copulatrix. From the bursa copulatrix leads another duct connecting to the uterine duct and the seminal receptacle. The bursa copulatrix is oval in shape, about four times as large as the seminal receptacle ( Fig. 36B ). In the central nervous system ( Fig. 36D ) the cerebral and pleural ganglia are fused and distinct from the pedal ganglia. There are three cerebral nerves leading from each cerebral ganglion and three pleural nerves leading from each pleural ganglion. There is a separate abdominal ganglion on the right side of the visceral loop. The buccal ganglia are near to the rest of the central nervous system, joined to the cerebral ganglia by two relatively short nerves. Gastrooesophageal, rhinophoral and optical ganglia are present. The pedal ganglia are clearly separated, having two nerves each one. The pedal and parapedal commissures are enveloped together with the visceral loop. Figure 35. Paradoris indecora (ZMUC) , SEM images of the radula, jaws and dorsal tubercles. A, inner lateral teeth; scale bar = 60 Mm. B, mid-lateral teeth; scale bar = 75 Mm. C, outer lateral teeth; scale bar = 60 Mm. D, jaw elements; scale bar = 30 Mm. E, dorsal tubercles; scale bar = 250 Mm. Figure 36. Paradoris indecora (MNCN 15.05/18231). A, general view of the anatomy; scale bar = 1 mm. B, reproductive system; scale bar = 1 mm. C, lateral view of the buccal bulb; scale bar = 1 mm. D, central nervous system; scale bar = 1 mm. E, ventral view of the mouth area; scale bar = 2 mm. The circulatory system ( Fig. 36A ) consists of a large heart and a two blood glands situated in front of and behind the central nervous system. Remarks Ortea (1995) revised the Atlantic species of Paradoris and concluded that Paradoris granulata Bergh, 1884 , the type species of the genus, is a junior synonym of Discodoris indecora Bergh, 1881 . The material from the Mediterranean examined here is identical to Ortea’s (1995) redescription of this species. He also described more new species from the area that appear to be distinct in several anatomical details.