(NUDIBRANCHIA: DISCODORIDIDAE) WITH A MORPHOLOGICAL PHYLOGENETIC ANALYSIS

The genus Jorunna is characterized by a dorsum covered with caryophyllidia, a prostate with two sections, a penis usually unarmed but occasionally armed with hooks, a copulatory spine, the presence of an accessory gland and a labial cuticle smooth or armed with jaw elements. The examination of 216 non-type specimens, 30 types, and a review of the literature show that there are 16 valid species of the genus Jorunna : J. tomentosa (Cuvier, 1804); J. funebris (Kelaart, 1859); J. pantherina Angas, 1864; J. rubescens (Bergh, 1876); J. labialis (Eliot, 1908); J. parva (Baba, 1938); J. spazzola (Marcus, 1955); J. hartleyi (Burn, 1958); J. alisonae Marcus, 1976; J. lemchei (Marcus, 1976); J. evansi (Eliot, 1906); J. pardus Behrens & Henderson, 1981; J. ramicola Miller, 1996 and J. onubensis Cervera, Garcı ´ a-Go´mez & Garcı´a, 1986. In addition, two new species from the Eastern Paciﬁc are described: J. osae n. sp. and J. tempisquensis n. sp. We propose two new combinations: Jorunna parva and J. evansi . New records for the genus Jorunna are provided from Italy, Algeria, Seychelles, Madagascar, Thailand, Marshall Islands, New Caledonia, Iˆle de la Re´union, Sudan, Papua New Guinea, Indonesia, Panama, Costa Rica, Bahamas, and Southern Mexico. We present the ﬁrst preliminary phylogenetic analysis of this cryptobranch dorid genus, based on morphological anatomical data, and discuss the biogeography and evolution of several characters in this group. The phylogeny supports the hypo-thesis that the genus Jorunna is a monophyletic group and shows that Kentrodoris is nested within it.


INTRODUCTION
The genus Jorunna Bergh, 1876 was previously composed of 19 species described from the Atlantic, Eastern Pacific, and the Indo-Pacific. Valde´s (2002) proposed that this genus belongs to the family Discodorididae Bergh, 1891. The first reference to species of this genus was the description of Doris tomentosa Cuvier, 1804 from La Rochelle, France. Since then, additional species have been described by Kelaart (1859), Angas (1864), Bergh (1876Bergh ( , 1878Bergh ( , 1881, Eliot (1906Eliot ( , 1908, Baba (1938), Er. Marcus (1955), Ev. , Risbec (1956), Burn (1958), , Cervera, Garcı´a-Go´mez & Garcı´a (1986) and Miller (1996). Kay & Young (1969) considered all specimens worldwide to represent a single species, Jorunna tomentosa, and this idea was supported by Edmunds (1971). Bergh (1876) established the genera Jorunna and Kentrodoris. He based Kentrodoris on three species, with Kentrodoris rubescens Bergh, 1876 from the tropical Indo-Pacific as its type. The other two species were Kentrodoris gigas Bergh, 1876 and Kentrodoris annuligera Bergh, 1876. All were characterized by a penis armed with a long spine (which we show is associated with the accessory gland rather than the penis), a large accessory gland which opens near the vagina, and a lack of jaws and rachidian teeth. Bergh (1876) based Jorunna on Doris johnstoni Alder & Hancock, 1845, which is considered a synonym of Jorunna tomentosa (Cuvier, 1804) (Fischer, 1869). The main features of the genus include a dorsum covered with caryophyllidia, low rhinophoral and branchial sheaths, a massive prostate with two sections, an unarmed vagina, a penis occasionally with hooks, an accessory gland with a spine, a labial cuticle (either smooth or with jaw elements), inner and lateral hamate radular teeth, and outermost teeth either smooth or denticulate (Valde´s & Gosliner, 2001). According to Bergh (1876), the main difference between Jorunna and Kentrodoris is that the spine of Kentrodoris is situated on the penis, whereas it is in the accessory gland in Jorunna. Er. Marcus (1955) described the genus Awuka, based on Awuka spazzola Marcus, 1955, with the same features as Jorunna and Kentrodoris. According to this description, the presence of jaws and one single denticle on the innermost radular teeth are the distinctive features of Awuka. Marcus also described a genital spine situated on the penis. Later, Ev.  regarded the genus Awuka as a synonym of Jorunna. Marcus (1976) also found that the spine described by Bergh (1876) and Er. Marcus (1955) on the penis of Kentrodoris, was actually situated in the large accessory gland of both Kentrodoris and Jorunna. However, she concluded that Kentrodoris was different from Jorunna because of the 'soft notum, the high gills, the acinous prostate, the loculated spermatocyst and the sheathed male organ' of Kentrodoris. She designated K. rubescens, the type species of the genus, as the only member of Kentrodoris and regarded the other two species, originally included by Bergh (1876) in Kentrodoris (K. gigas and K. annuligera), as members of Jorunna.
The genus Jorunna was reviewed by Ev.  who demonstrated significant differences in the anatomy of species from different regions, described five new species and reinstated several others. She characterized Jorunna as having stiffened notal tubercles or caryophyllidia, simple lateral radular teeth, marginal radular teeth (sometimes with irregular denticles), a smooth prostatic epithelium, an unsheathed male duct ending with a spineless papilla, or even without a papilla, and an accessory gland with a spine.
The phylogenetic relationships of the genera Kentrodoris and Jorunna were studied by Valde´s & Gosliner (2001), supporting the idea that the morphological differences between the genera are just autapomorphies of Kentrodoris rubescens. They therefore regarded Kentrodoris and Jorunna as synonyms, and the name Jorunna was selected as having precedence over Kentrodoris (ICZN, 1999: Art. 24).
The aims of the present study are: (1) to provide a comprehensive systematic revision of the genus Jorunna; (2) to test whether Jorunna is a monophyletic group and to present the first phylogenetic analysis of the genus; (3) to discuss the biogeography and evolution of several characters in the context of the phylogeny. No phylogenetic studies based on cladistic criteria have previously been published for Jorunna.

MATERIAL AND METHODS
Living animals, non-type material and type material were studied from the collections of several museums and institutions: Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco (CASIZ); Muse´um National d'Histoire Naturelle, Paris (MNHN); Zoologisk Museum, Copenhagen (ZMC); National Museum of Natural History, Washington DC (USNM); Natural History Museum, London (BMNH); and University of Costa Rica (UCR) (which now houses the collections of the Instituto Nacional de Biodiversidad, INBio).
In total, 246 specimens were examined, including 30 types. Eighty-eight of these specimens were dissected. For most of the specimens that were dissected we observed the external coloration and morphology (such as dorsum-caryophyllidia, gills, lamellae, oral tentacles, foot, rhinophores, mantle glands etc.), and the internal anatomy (labial cuticle, radula and reproductive system). For the rest of the specimens that were not dissected, we observed the external coloration and morphology. Specimens were dissected by dorsal incision. Their reproductive systems were examined and drawn under a dissecting microscope with camera lucida. At least two specimens of each species were examined anatomically. The buccal bulb was extracted from the specimen by making two dorsal longitudinal incisions from behind the rhinophores to the gill. The jaw and radula were dissected and isolated from each other. Radulae were placed in a 10% solution of NaOH to remove tissue. Jaws and radulae were examined by scanning electron microscope. Mantle samples were taken from each specimen, placed in 100% alcohol and critical-point dried. The accessory gland was dissected from the reproductive system, stained using acid fuchsin, dehydrated in a series of ethanol, cleared in xylol and mounted on a slide. The copulatory spine was drawn using a microscope with camera lucida.
Jorunna has one of the most difficult reproductive systems to draw among dorids. We tried to represent the different structures in our drawings exactly as we saw them. In most of the cases, however, we had to produce up to three drawings (with the exception of J. parva and J. ramicola) to represent these structures. For example, in most of the cases the first thing we see is the accessory gland (covering at least 25-70% of the reproductive system in mature individuals), part of the bursa copulatrix, seminal receptacle and sometimes part of the prostate and ampulla. Second, we had to move or remove the accessory gland to see the details of the seminal receptacle, bursa copulatrix and ducts. Third, in most cases after moving or removing the bursa copulatrix and seminal receptacle we were able to see the connections between the ampulla and prostate.
Photographs of living animals for most of the species were taken by the collectors and used to record features of living animals. External morphological data were obtained from the original descriptions, and from redescriptions when specimens or type material were not available. The names of the species have been treated in chronological order as they were originally described.

SYSTEMATIC DESCRIPTIONS Part A: The genus Jorunna (including
Kentrodoris and new taxa) External morphology: Body oval, convex (Fig. 1A, B). Caryophyllidia elongate, c. 150 mm in length, homogeneously distributed around the branchial, rhinophoral sheaths and notum, with short spicules and ciliated tubercles (Fig. 2E). Eleven bipinnate gills and rhinophores with 14 lamellae in 16 mm preserved length specimen (CASIZ 073939). Ten bipinnate gills and 13 lamellae in 15 mm preserved length specimen (CASIZ 086912). Eleven bipinnate gills and 14 lamellae in 13 mm preserved length specimen (CASIZ 072598). Gills imbricate, perpendicular relative to body. Background colour of living animal yellow-cream to grey (Fig. 1A, B). Three to six small, light brown spots distributed randomly over dorsum forming two vertical lines relative to rhinophores. Light spots composed of aggregation of minute spots. In specimens with yellow-cream background, rhinophores and gill also yellow-cream. Apices of rhinophores same colour, but with some dark brown pigmentation on lamellae. Branchial leaves also with some dark brown pigmentation on some branches at their apices. Specimens with a greyish background with rhinophores and gills of same colour, although apices of rhinophores yellow to cream. Small, white, sphaerical glandular structures surrounding mantle edge. Anal papilla with seven divisions, speckled with dark pigment. Foot and hyponotum same colour as notum without spots. Cream oral tentacles, conical in shape. Posterior end of foot visible dorsally when animal moves.
Ampulla very long, divided into the short oviduct and prostate (Fig. 3A, C). Ampulla entering female glands near upper part of mass. Prostate very large, granular, narrowing into short, coiled deferent duct, expanding again into muscular, wide ejaculatory portion. Deferent duct opening into common atrium with vagina (Fig. 3B). Penial hooks absent. Long, convoluted accessory gland connecting to atrium (Fig. 3A). Accessory gland with spine about 920 mm long. Spine straight, pointed (Fig. 3D). Vagina long, tubular, wider than deferent duct. Uterine duct short, thin, opening in distal part of female gland mass. Seminal receptacle oval, at least one-fifth as large as rounded bursa copulatrix (Fig. 3B).
Distribution: From 658N on the Norwegian coast to Faeroes, all coasts of Britain, France, Portugal, Morocco , and South Africa (Gosliner, 1987), Italy and Algeria (present study).
Remarks: Cuvier (1804) incompletely described Doris tomentosa as a species with a white to grey semi-translucent background colour, a curved body, a mantle larger than the foot, and a woolly dorsal texture. Johnston (1838) described Doris obvelata as having a uniform yellowish-white background colour, usually dashed with a few dark spots, a broad margin, caryophyllidia on the dorsum, and bipinnate gills.
A more complete description of Doris johnstoni Alder & Hancock, 1845, was later published by Alder & Hancock (1845-1855, including an anatomical description. Fischer (1869) synonymized Doris johnstoni with Doris tomentosa for the first time.
In her revision of the genera Kentrodoris and Jorunna, Marcus (1976) divided Jorunna tomentosa, into four different species: J. malcomi, J. alisonae, J. lemchei and J. luisae, based on differences in the radular and jaw morphology and the reproductive system. Thompson & Brown (1984) synonymized J. lemchei with J. tomentosa, noting that these two are indistinguishable in habits, external morphology and internal anatomy. These authors also included J. luisae as a possible synonym of J. tomentosa. Later, Valde´s & Gosliner (2001) indicated that these three species (J. luisae, J. lemchei and J. tomentosa) have sufficient differences to justify their separation as different species. According to these authors, J. lemchei is clearly different from J. tomentosa because of the presence of large hooks on the penis and the absence of denticulation on the outermost lateral teeth. Jorunna luisae differs from this last species in the presence of jaws and denticulated innermost lateral teeth. However, in our study we have synonymized J. luisae with J. spazzola (Marcus, 1955).
In a specimen from Plymouth, England, Marcus (1976) described the outer lateral teeth from the right side of the radula as having denticles, whereas some of the teeth on the left side do not have denticles. The specimen studied by Cervera, Garcı´a & Garcı´a (1986) from Spain lacks denticles on the outermost teeth. In the present study, we examined several specimens of J. tomentosa from different localities and we found that the specimens from Spain (CASIZ 115215) and England (CASIZ 079486) show intra-specific variation, whereas specimens from South Africa (CASIZ 073939), Norway (MNHN, no catalogue number), and Quiberon, France (MNHN, no catalogue number) do not have any denticles on the outermost lateral teeth.   alisonae Marcus, 1976 from Hawaii can be distinguished from J. tomentosa by the presence of a labial cuticle with jaw elements, their radular morphology and differences in the reproductive system. Jorunna onubensis has a non-prostatic deferent duct with two regions, which is absent in J. tomentosa. Jorunna alisonae has only one small denticle on the outermost teeth, whereas J. tomentosa has more than one denticle on two or more of the outermost teeth, in some cases. Additionally, while the background colour of J. tomentosa is yellow-cream to grey with small light spots randomly distributed, J. labialis has a white to very dark grey background colour with light brown spots and J. alisonae has a pale grey background colour with two vertical lines of black spots and opaque white spots around the mantle margin.

Kentrodoris gigas
External morphology: Body rigid, finely tomentose. Body profile high, dorsum densely covered with caryophyllidia, about 220 mm in length (Fig. 4D). Rhinophoral and branchial leaves low. Six tripinnate branchial leaves and 14 lamellae on rhinophores in a 20 mm preserved length specimen (CASIZ 120962), 6 tripinnate gills, 15 lamellae in 11 mm preserved length specimen (CASIZ 106445), 6 tripinnate gills, 20 lamellae in 15 mm preserved length specimen (CASIZ 074203). Branchial leaves large in size when compared to size of animals, varying in length within a single gill circle. Posterior prolongation of foot visible dorsally when animal moves.  Background colour of dorsum and foot white to yellowcream. Notum with several dark brown to black rings of different sizes, larger in middle portion of middle body (Fig. 1C). Dark rings not completely covered with pigment, instead each ring composed of several minute dark brown to black spots homogeneously distributed inside. Border of notum and hyponotum scattered with similar dark spots all around margins. No rings present on sole of foot; however a couple of dark brown dots at each side of notched upper lip in some specimens. Rhinophores white with black clubs. Branchial leaves white, rachises of first and second pinnae of each leaf with black line. White mantle glands present around margin of notum.
Wide, tubular ampulla convoluted in middle region, branching into the short oviduct and prostate (Fig. 5C). Oviduct enters closely adjacent to female gland mass. Female gland about the same size as accessory gland. Short vas deferens separates from ampulla and opens into the longer, widened portion of glandular prostate. Prostate consists of two distinct glandular sections, well differentiated. Deferent duct short, undulate. Long uterine duct emerges from female gland mass, joining pyriform seminal receptacle at its base (Fig. 5B). Duct connecting seminal receptacle and oval bursa copulatrix, long, coiled. Bursa copulatrix smaller than seminal receptacle in some specimens, whereas in others, twice as large as seminal receptacle (Fig. 5B). Vaginal duct emerging from base of bursa copulatrix long, thin, smooth. Muscular deferent duct opens into common atrium with vagina. No penial hooks. Large accessory gland connected to atrium (Fig. 5A). Accessory gland consisting of a muscular portion, with a curved, pointed spine about 717 mm long (Fig. 5D, E).
Remarks: Kelaart (1859) described Doris funebris as a dorid with an ivory-white mantle ornamented with jet-black spotted circles and half-rings or imperfect annular spotted figures. Alder & Hancock's (1864) redescription is more complete, including an illustration of the animal. Bergh (1876) introduced two species from the Philippines. Kentrodoris annuligera, described as having a bright yellowish mantle, with black rings and rhinophores and a black gill, for which he included anatomical descriptions and illustrations of the species and its radular teeth, and Jorunna gigas, which was characterized by caryophyllidia, black rhinophores, oral tentacles with black tips, a black-brown foot, curved body, an accessory gland with a spine, a labial cuticle smooth, six tripinnate branchial leaves, a dorsum with blackish marks, and hamate teeth without denticles. According to , the description of these two species is very similar. Their radular morphology is very similar, and they both have dark blotches or rings on the dorsum, and oral tentacles with black tips.
Considering the similarity between these two descriptions, and the fact that both species come from the same locality (Aibukit, Philippines), we synonymize them. Eliot (1906) described Kentrodoris maculosa as having a yellow background with black ring-like spots, black rhinophores with white bases and apices, a yellow gill with black rachises, and oral tentacles with black apices. This species was synonymized by  with J. funebris.
The original description of Discodoris wetleyi Allan, 1932 includes two figures (Allan, 1932: pl. 4, figs 1, 2) in which it is possible to recognize a colour pattern very similar to that of J. funebris. Discodoris wetleyi was described as having a notum with minute granulations, a prolongation of the foot that is visible when the animal is in motion, a yellowish cream background colour with irregular sized patches of dark brown composed of minute dots. The rhinophoral club is black with a white base. Branchial leaves are yellow-cream with dark brown tips and have numerous small dark spots along the marginal end of the foot. Kenny (1970) synonymized this species with J. funebris and, after studying the original descriptions, we agree with this conclusion.
According to  the differences between J. zania and J. funebris are the size and shape of the penial papilla, which is large and bulbiform in J. funebris and smaller in J. zania. In addition,  found that in J. funebris the atrium and the copulatory spine are longer, whereas in J. zania the atrium is partly everted and the copulatory spine smaller.  also mentioned that both species have black spots on their notched upper lip and that J. zania sometimes has a few needle-like serrations in one or two of the outermost teeth, a feature that was not reported by Edmunds (1971) nor was found in the material studied here. The examination of several specimens of J. funebris, and the review of the original description and subsequent redescriptions, confirms that the external and internal features of J. zania fits with those of J. funebris. The presence of a smooth labial cuticle, a pointed accessory spine, the simple hamate shape of the teeth lacking denticulations, the external coloration of the body, including the presence of black spots on the notum and the presence of black spots on their notched upper lip, are characteristics of J. funebris. The presence of an everted atrium and the size and shape of the penial papilla could be due to preservation or intra-specific variation. For these reasons, we do not consider the features mentioned by  enough to justify the existence of two different species, and therefore synonymize J. zania with J. funebris.
Jorunna pantherina (Angas, 1864) ( Table 2) redescription of the same species by Miller (1996) in the radular and external morphology and details of the reproductive system. According to these authors, J. pantherina is characterized by having a purplish brown or dark grey to pale brown or greenish background colour with small dark blotches, bipinnate branchial leaves, an accessory gland with a spine and hamate outermost and midlateral teeth, lacking denticles. However, Thompson (1975) and  reported the presence of jaw elements in their specimens, a feature that is not present in the description by Miller (1996).  illustrated the jaw elements in her description. Miller (1996) mentioned that there are some differences between specimens from New Zealand and Australia; the former can be very dark in colour, with a smooth labial cuticle and without autotomization of the mantle rim. Unfortunately, due to the absence of material, we are unable to clarify the presence of jaw elements to confirm the identity of this species. The presence of acutely pointed lateral teeth and jaws in J. hartleyi Burn, 1958, andJ. pantherina (Angas, 1864), the verification that J. hartleyi posseses an accesory gland spine and that both species are found in Australia, strongly suggest that they are possible synonyms.
Jorunna rubescens (Bergh, 1876) (Figs 1E, 6, 7, Table 2) External morphology: Body elongate, elevated (Fig. 1E). Caryophyllidia about 176 mm in length, densely distributed covering entire dorsum, with long, ciliated base, spicules projecting over tip and elongate tubercle (Fig. 6E). Rhinophoral and branchial sheaths highly elevated. Seven tripinnate branchial leaves and 23 lamellae on rhinophores in 35 mm preserved length specimen (CASIZ 109797), 6 tripinnate gills, 23 lamellae in 42 mm preserved length specimen (CASIZ 068673), 7 tripinnate gills, 25 lamellae in 41 mm preserved length specimen (CASIZ 073074). Background colour cream greyish to yellow, with a distinctive pattern on dorsum in living animals that can also be seen in preserved specimens. Pattern consisting of several oval yellow spots surrounded by a cream ring, distributed all over dorsum. Several horizontal black stripes along body (Fig. 1E). Anterior part of head black with some yellow dots. Rhinophoral sheaths black. Base of rhinophores black or cream yellowish, club black-spotted with some cream white spots. Branchial sheaths black-spotted with some oval yellow spots at base becoming bigger at tip. Base of branchial leaves white, rachises dark brown. Upper lip of anterior border of foot speckled with black. Sole of foot white-creamish with some black stripes also present on laterals of foot. Posterior end of foot dorsally visible when the animal moves. Foot cream greyish to yellow with some dark black spots on edges. Ventrally, anterior border of foot grooved, notched. Oral tentacles conical, white to yellow.
Ampulla long and convoluted (Fig. 7B), branching into short oviduct and prostate. Flattened prostate short, wide with a loop in middle portion (Fig. 7C). Prostate narrowing into long, convoluted deferent duct, expanding into ejaculatory portion. Penis unarmed. Highly differentiated, pear-shaped accessory gland ( Fig. 7A) with spine about 3.7 mm in length (Fig. 6D). Vagina long, almost straight. At proximal end, branching into duct that connects to a lobate seminal receptacle and uterine duct (Fig. 7B). Uterine duct short, wide opening near the upper part of female gland mass. Seminal receptacle one-third as large as oval bursa copulatrix.
Distribution: Western Pacific -Japan, China Sea, Marshall Islands, Malaysia, Thailand, Philippines, Indonesia, Papua New Guinea and Australia ; Western Indian Ocean -Aldabra Atoll, Mauritius, Sri Lanka, Iˆle de la Re´union (present study).
Remarks: According to Valde´s & Gosliner (2001), Doris venosa Quoy & Gaimard, 1832 is a likely senior synonym of J. rubescens (Bergh, 1876). Doris venosa was described as being a dorid with a white background and broken irregular reddish lines resembling 'veins' on the dorsum, and which also has yellow along the edges with some orange spots; the mantle has blue dark spots and the animal has an elevated body. Although the description resembles that of J. rubescens, we regard Doris venosa as a possible synonym, following Valde´s & Gosliner (2001), due to the uncertainty in establishing the generic position of the species and lack of type material.
Remarks: This species was originally placed in the genus Rostanga by Eliot (1906), presumably because of the presence of a prostate, unipinnate branchial leaves and caryophyllidia. According to Rudman & Avern (1989), the narrow radula (with twenty or less teeth in a half row) is a characteristic of the genus Jorunna rather than Rostanga. Rudman & Avern (1989) also mentioned that J. spazzola (Marcus, 1955), described from Brazil, is similar to this species in external coloration and radular morphology. However, Marcus (1955) described an accessory gland with a spine in J. spazzola, a characteristic that was not found by Eliot (1906) in J. evansi.
The present study of one of the paralectotypes (9 mm) of Rostanga evansi showed that an accessory gland with a curved copulatory spine is present (Fig. 25B). In addition, although Eliot (1906) mentioned the lack of oral tentacles in the specimens studied from Cape Verde, the 4 mm lectotype specimen showed oral tentacles (Fig. 25C).
From the descriptions of J. evansi and J. spazzola, as well as the examination of a new specimen from Costa Rica, we concluded that these two species share a similar external morphology and anatomy. The only difference we can see is the absence of a denticle on the innermost teeth in J. evansi that is sometimes present in J. spazzola. Whether these two species are synonyms or not remains uncertain in the absence on new material for comparison from Cape Verde.
Jorunna labialis (Eliot, 1908)   length ( Fig. 8E) with long conical base, long spicules, short and rounded, ciliated tubercle. Spicules higher than tubercles. Rhinophoral and branchial sheaths low, regular. Eleven bipinnate branchial leaves forming a circle. Rhinophores wide, with eight lamellae in 10 mm preserved length specimen (CASIZ 081804). Eleven gills and 11 lamellae in 10 mm preserved length specimen (CASIZ 081804). Lamellae arranged almost vertically. According to Eliot (1908), the colour of living specimens of J. labialis may vary from white to very dark dull grey with light brown spots. The general colour of preserved animals studied here is uniformly cream yellowish without spots. Rhinophores and gills same colour as mantle. Anterior border of foot notched and grooved. Oral tentacles conical. Mantle margin slightly wider than foot. Ventral colour same as dorsum in preserved animals.
Ampulla elongate, tubular, with a loop in middle portion, branching into a short oviduct and prostate (Fig. 9C). Oviduct enters female gland near distal part of mass. Prostate flattened, folded, granular ( Fig. 9C), connecting to a long duct leading into ejaculatory portion of deferent duct. Deferent duct thinner in its proximal part and opening into a common atrium with vagina. No penial hooks. Accessory gland, amorphous, convoluted, connecting to the atrium (Fig. 9A), with a muscular portion, a curved spine about 600 mm long (Fig. 9D), a wide duct, and glandular region connected to it. Vagina long, with some loops, thinner than deferent duct, connecting to bursa copulatrix at its distal end. Another duct, connecting to both the uterine duct and seminal receptacle leading from bursa copulatrix. Bursa copulatrix oval, about three times larger than oval seminal receptacle (Fig. 9B).
Remarks: Eliot (1908) described Kentrodoris labialis from the Red Sea as a dorid with caryophyllidia, jaw elements, bipinnate gills, hamate outermost teeth without denticles, a curved spine and a white to dark dull grey mantle with some light brown spots. Edmunds (1971) redescribed J. tomentosa (Cuvier, 1804) from Tanzania as having caryophyllidia, a labial cuticle with jaw elements, a convoluted prostate, outermost teeth with no basal plate, a lanceolate cusp, and a pale grey-green to grey background colour speckled with dark or purple-brown spots and white dots along the edges. Edmunds considered that J. tomentosa was a cosmopolitan species extending from Europe to the Marshall and Hawaiian Islands, following Kay &Young (1969).
Later, Marcus (1976) described J. malcomi, based on the specimens identified by Edmunds (1971) as J. tomentosa from Tanzania. According to  J. malcomi is present in both the Atlantic Ocean (Ghana) and the Indian Ocean (Tanzania and Madagascar). Although the drawings of the reproductive systems provided by  show no significant differences, the presence of this species in both areas seems very unlikely from a biogeographical perspective. The identification of Atlantic specimens remains doubtful until more specimens from both localities are studied.
Comparing the original descriptions of J. malcomi  from Tanzania and Ghana and of J. labialis (Eliot, 1908) from Suakin and Suez, we found that both species share the presence of caryophyllidia, a yellowish brown to pale grey-green background colour with rows of dark brown spots along the sides speckled with white marks, a labial cuticle with jaw elements, an accessory gland with a spine, midlateral and outermost teeth hamate and lacking denticles, and bipinnate gills. In addition, the reproductive system of these two species is very similar (Eliot, 1908;Edmunds, 1971;. Our material from Port Sudan fits these descriptions and, therefore, J. malcomi and J. labialis are synonyms, the latter being the older name. Jorunna labialis differs from J. tomentosa by the presence of a labial cuticle with jaw elements, the shape of the outermost teeth and the absence of denticles. There are also some differences in the reproductive system such as the absence of a large penial papilla in J. labialis. Finally J. labialis is white to very dark grey with light spots, while J. tomentosa is cream yellowish to grey with light spots that form two vertical lines. Jorunna labialis is distinguished from other species found in the Red Sea (J. funebris) and Indian Ocean (J. rubescens) by the external and radular morphology. Jorunna funebris and J. rubescens both have the mantle covered with dark rings or stripes, whereas these are not present in J. labialis. In addition, J. rubescens and J. funebris lack jaw elements.
External morphology: Body elongate. Dorsum with long caryophyllidia, about 250 mm in length (Fig. 10D). Caryophyllidia with long cylindrical base, long spicules, and long ciliated tubercle. Caryophyllidia of different shapes and sizes, the larger ones sparsely distributed between the smaller ones. Larger caryophyllidia leaf-like in shape, homogeneously distributed, the rest generally smaller, elongated. Rhinophoral and branchial sheaths low. Seven short bipinnate branchial leaves and rhinophores with 15 lamellae in 8 mm preserved length specimen (CASIZ 115730). Six bipinnate gills and 14 lamellae in 6 mm preserved length specimen (CASIZ 74251). Background colour of living animal dark yellow-orange to dark brown (Fig. 1F, G). Most specimens with a dark yellow background colour becoming lighter in areas close to rhinophores and gill. Other specimens with a dark brown colour in central part of dorsum, becoming dark yellow towards edges of mantle (forming a V-shape in area behind rhinophores). Light yellow colour following, becoming darker again towards front part of rhinophores (Fig. 1G). Higher caryophyllidia dark brown in colour, resembling dark spots in mantle. In some specimens, base of rhinophore rachis light yellow with dark brown clubs, base of gill light yellow with dark brown tips. Other specimens with rhinophores with dark brown rachises containing light yellow clubs and light yellow gills with dark brown rachises. Foot with a dark brown line on middle part of visible portion, with dark brown spots along edges clearly visibly when the animal moves. Large white mantle glands around mantle.
Anterior border of foot notched, grooved. Oral tentacles elongate. Margin of hyponotum with dark brown spots. Upper lip yellowish with two brown spots on each side of lip in some specimens.
Ampulla long, curved, branching into a short oviduct and prostate (Fig. 11B). Oviduct entering female glands close to distal part of mass. Prostate flattened, glandular, folded in middle region, connecting to a short folded duct expanding into short ejaculatory portion of deferent duct. Muscular deferent duct opening into a common atrium with vagina. No penial hooks. Large, convoluted accessory gland connecting to atrium, with a muscular portion, a pointed spine about 477 mm long (Fig. 11C), a long, thin folded duct and a glandular, convoluted proximal area. Vagina short, wider in its proximal region connecting to oval bursa copulatrix. Another thin duct leading from bursa copulatrix, connecting to long uterine duct and pear-shaped seminal receptacle. Bursa copulatrix larger than seminal receptacle (Fig. 11A).
Distribution: Tanzania, Seychelles ( present study) to Kii Peninsula (Baba, 1938) and Okinawa in Japan, Philippines and Papua New Guinea ( present study).
Remarks: According to Baba (1938), Thordisa parva is characterized by having digitiform oral tentacles, a smooth labial cuticle, some black spots on the dorsal tubercles and on the edges of the hyponotum, black rhinophoral tips and branchial leaves, most lateral teeth hamate without denticles and pectinate outermost teeth. From this and our own material of Thordisa parva, we conclude that this species should be transferred to the genus Jorunna, because of the presence of caryophyllidia and a large accessory gland with a long spine. In Thordisa, there are simple elongate tubercles and the accessory gland is much smaller with short spines (Chan & Gosliner, 2007).
Jorunna parva differs from other species of Jorunna from the Indo-Pacific in its external coloration and radular morphology. Jorunna parva is the only species of the genus that has dark tubercles resembling dark spots on the mantle; a visible foot with a dark brown line on the middle part of the dorsal side with dark brown spots on the edges is also a characteristic of the species. Although the radular morphology of J. parva is similar to that of J. ramicola, there are some differences. The labial cuticle of J. parva is smooth, whereas there are jaw elements in the other species. Also, there are no denticles on the innermost teeth of J. parva, whereas they are present in J. ramicola.
The reproductive system of the holotype of J. luisae was studied. Ampulla thick and convoluted, divided into short oviduct and prostate (Fig. 13A). Ampulla entering small female gland near upper part of mass. Prostate large, granular and folded into itself, expanding into short ejaculatory portion. Deferent duct opening into common atrium with vagina. Penial hooks absent. Convoluted accessory gland connecting to atrium, with a spine visible through the tissue (Fig. 13B). Vagina wider than defferent duct, tubular and coiled. Duct that connects bursa copulatrix with seminal receptacle long and coiled. Seminal receptacle absent.
The reproductive systems of the specimens from Costa Rica and the Bahamas were not fully mature. Ampulla particularly wide in both specimens examined (Fig. 13D), entering small female gland near distal part of mass. Prostate granular, flattened, connecting to deferent duct, other portion connecting to female gland. Deferent duct long and coiled, expanding again into tubular ejaculatory portion. Deferent duct opening into a common atrium with vagina. No penial hooks.
Small accessory gland connecting to atrium (Fig. 13D). No spine was found in the inmature specimens. Vagina long, tubular, connecting to oval bursa copulatrix at its distal end. Another convoluted duct, connecting to oval seminal receptacle and uterine duct, leading from bursa copulatrix. Bursa copulatrix bigger than seminal receptacle (Fig. 13D).
Distribution: Mediterranean, and in Western Atlantic from Brazil, Mexico, Costa Rica and the Bahamas ( present study).
Remarks: The holotype of J. luisae has been dissected. The coloration of the preserved animal is unknown, the specimen being stained with purple ink. There are caryophyllidia all over the dorsum. The radula and jaws (buccal bulb) of the holotype are missing. The reproductive system remains in the body, all the organs are present but the receptaculum seminis (Fig. 13A). The penis is everted and the spine is visible through the tissue (Fig. 13B). Other internal organs such as stomach, digestive gland and intestine are missing. Twelve lamellae in rhinophores. No branchial leaves present. Oral tentacles present.
Er. Marcus (1955) described Awuka spazzola from Brazil. Later, Ev.  redescribed the same species and placed it in the genus Jorunna. Ev. Marcus was aware of the error in the original description, where the accessory spine was interpreted as situated in the penis instead of in the accessory gland, as noted by Valde´s & Gosliner (2001).
Jorunna spazzola differs from most other members of the genus by the presence of jaws. The innermost lateral teeth are with or without a denticle and the outermost radular teeth are denticulate. However, J. luisae  from the Mediterranean is another species that shares these features. According to the original descriptions (Er. Marcus, 1955;Ev. Marcus, 1976), they both have caryophyllidia, a labial cuticle with jaw elements, unarmed penis, an accessory gland with a spine, the innermost radular teeth with a small denticle, outermost teeth with irregular denticles and uni to tripinnate leaves. According to the redescription of J. luisae by Schmekel & Portmann (1992) based on several specimens from Naples, this species has innermost teeth lacking a denticle and outermost teeth pectinate (2-4 denticles). In the specimen we examined from Costa Rica, we also found the innermost teeth lacking denticles but the outermost tooth pectinate. The presence or absence of denticles in the innermost teeth seems to be variable within the species.
The drawings of the reproductive systems in the original descriptions of these two species are very similar. No major differences can be identified other than the absence of loops in some of the ducts in J. spazzola, which could be due to the manipulation of the organs during dissection.  stated that ".... the differences between J. spazzola and J. luisae are insignificant....", however she preferred to keep the species from the Mediterranean separate from the Brazilian one. These differences are a larger number of lamellae on the rhinophores and teeth, shorter jaw elements, and a smaller female gland mass in J. luisae.
The similarity between these two species does not allow us to justify their separation of the species, so we regard them as synonyms. See also remarks on J. evansi from Cape Verde Islands.

Remarks:
The generic position of this species from southern Australia has recently been clarified by Rudman & Avern (1989). According to these authors, Rostanga hartleyi is most appropriately placed in the genus Jorunna, because of the radular morphology and the anatomy of the reproductive system. Jorunna hartleyi is characterized by having a pale pink mantle with large brown patches encircled with white or dark purple spots, caryophyllidia, an accessory gland without a spine, a smooth labial cuticle, an unarmed penis, elongate outermost teeth lacking denticles, and hamate midlateral and innermost teeth lacking denticles.
In contradiction to what these authors found in the accessory gland, we examined the holotype of R. hartleyi and found that the accessory gland is partially extruded, so that we could confirm the presence of a long spine of about 198 mm (Fig. 25A).
Comparing the original descriptions of J. hartleyi (Burn, 1958) and of J. pantherina (Angas, 1864), we found that these species share the presence of pointed lateral teeth and the presence of jaws, and both have been found in Australia. This suggests that they are possible synonyms, remains uncertain in the absence of new material for comparison. Jorunna alisonae Marcus, 1976 (Figs 1I -K, 14, 15,   External morphology: Body oval. Dorsum covered with caryophyllidia about 133 mm long (Fig. 14F). Caryophyllidia elongate, with long spicules. Twelve tripinnate branchial leaves and 10 lamellae in rhinophores in both specimens of 9 and 10 mm preserved length (CASIZ 031866). Oral tentacles conical. Anterior portion of foot notched, grooved.
Background colour pale grey, with numerous grey dots of different sizes on mantle margin (Fig. 1I-K). Two lines of spots nearly forming two vertical lines arranged from rhinophores to gills. Viscera visible in the centre of dorsum. Rhinophores cream-brownish with a pale grey-brownish club. Gill grey to dirty brown with some cream glandular spots. Ventral part of animal, the same colour as dorsum. Posterior end of foot visible dorsally when animal moves.
Ampulla long, thin with a loop in distal portion, dividing into short oviduct and prostate (Fig. 15B). Ampulla entering female gland mass near upper middle part of mass. Prostate large, flattened, glandular, convoluted in middle section, narrowing into short, coiled deferent duct, and expanding again into muscular, narrow ejaculatory portion. Penis unarmed. Amorphous accessory gland with a distinct stylet sac containing a spine about 1.15 mm in length (Fig. 15A). Copulatory spine straight, pointed (Fig. 15D). Deferent duct opening into a common atrium with vagina. Vagina long, tubular (Fig. 15C). Uterine duct short, thin, opening in upper middle part of female gland mass. Convoluted duct from oval bursa copulatrix leading to seminal receptacle. Seminal receptacle slightly oval, almost as large as bursa copulatrix (Fig. 15C).
Distribution: Only from Hawaii , Marshall Islands and Indonesia ( present study).
Remarks: The holotype has not been dissected; it is well preserved and of creamish colour. There are caryophillydia all over the dorsum. There are 15 bipinnate branchial leaves. We could only count about six lamellae in the rhinophores, which are strongly retracted into their rhinophoral sheaths. The penial papilla is partially everted. Jorunna alisonae (Marcus, 1976) can be differentiated from J. tomentosa, a similar species from the northeastern Atlantic, Mediterranean and South Africa, by the presence of jaw elements, a single denticle on the outermost teeth that is present in some specimens, and tripinnate branchial leaves.
According to , the presence of a single denticle close to the tip on the outermost lateral teeth in J. alisonae may be a malformation. Among the several specimens examined in this study from Hawaii and the Marshall Islands, we found a single denticle present in only one. We conclude that its presence is not a malformation, but a variable feature of this species.  ( Distribution: Only from western Ireland .

Jorunna lemchei
Remarks: The holotype has been dissected and largely destroyed. The coloration of the preserved animal is creamish. The radula and jaws (buccal bulb) of the holotype are missing. The oral tube is present. Most of the internal organs of the holotype are also missing (reproductive system, stomach, intestine); however, part of the digestive gland is present. There are 11-12 bipinnate branchial leaves and about 15 lamellae in the rhinophores. Oral tentacles present.
Although the external coloration of this species is very similar to that of J. tomentosa, J. lemchei is clearly different from this and all other congeners owing to the presence of cuticular spines on the penial papilla and male atrium . Confirmation of this feature in other specimens from the type locality would be highly desirable.

Jorunna pardus Behrens & Henderson, 1981
(Figs 1L, 16, 17, Table 3) External morphology: Body shape oval, elongate (Fig. 1L). Centre of dorsum elevated. Dorsum densely covered with caryophyllidia, about 170 mm, homogeneously distributed on mantle. Border of mantle undulate. Rhinophoral sheath moderately elevated. Eight tripinnate gills and 14 lamellae in a Y. E. CAMACHO-GARCI´A AND T. M. GOSLINER 35 mm preserved length specimen (CASIZ 072359), 9 gills, 16 lamellae in 29 mm preserved length specimen (CASIZ 072420). Anal papilla located in centre of branchial leaves. Anterior border of foot grooved, notched. Oral tentacles long, digitiform with a bifid tip present in some specimens. Background colour of body yellow-cream. Entire dorsum speckled with numerous dark brown spots of different sizes composing of minute dark brown dots. Caryophyllidia present within these brown spots (Fig. 16E) also spotted dark brown, the rest yellow-cream. Base of branchial sheaths and rachis yellow-cream with dark brown tips. Rhinophores dark brown with base and rachis yellow-cream. In some specimens, the yellow-cream sole of foot and notum are lightly speckled with small dark brown spots. Anterior border of foot with large dark brown dots on upper lip. Foot dorsally visible when the animal moves. Big dark brown spots around margin of foot.
Ampulla long, wider in middle part (Fig. 17B), entering female gland near distal part of mass. Prostate flattened, granular, divided into two different portions, as in other members of genus. Thicker portion of prostate connecting to deferent duct, other portion connecting to female gland, next to opening of ampulla. Deferent duct short, expanding again into short, narrow ejaculatory portion. Deferent duct opening into a common atrium with vagina. No penial hooks (Fig. 16F). Large, convoluted accessory gland connecting to atrium (Fig. 17A), consisting of a muscular portion with a pointed spine 950 mm long (Fig. 17D), a long and thin folded duct and a granular convoluted area connected to it. Vagina long, wide, connecting to oval bursa copulatrix at its distal end (Fig. 17C). Another convoluted duct, connecting to oval seminal receptacle and uterine duct, leading from bursa copulatrix. Bursa copulatrix about the same size as seminal receptacle (Fig. 17C).
Distribution: Southern California  to Sacramento Reef, Baja California (Behrens & Gatewood, 1986). It is found on rocks in the intertidal and subtidal, from 5 to 18 m depth.

Remarks:
The holotype and the paratypes are in good condition and not dissected. All of the specimens have caryophyllidia on the notum. The background colour of the specimens is cream yellowish with dark black spots all over the dorsum and dark gills and rhinophores. There are dark spots around the margin of the foot in most of the specimens.
The holotype has an extruded buccal bulb covered with dark spots. The penis in the holotype and in the 32-mm paratype specimen (CASIZ 15863) is extruded and a pointed spine is visible outside. An extruded buccal bulb with dark spots is also found in the 28-mm paratype specimen (CASIZ 15863). The 30-mm paratype specimen (CASIZ 15863) has a partially extruded penis, the mantle is damaged on the right side. The margin of the foot has a few spots around the mantle margin. Both tips of the oral tentacles are bifurcate. One of the oral tentacles of the paratype specimen of 28 mm preserved length is bifurcate as well.
Behrens & Henderson (1981) described J. pardus as a species that has caryophyllidia, leopard-like spots on the notum, a smooth labial cuticle, an accessory gland with a spine, a marginal foot with dark spots, and outermost teeth with denticles. This description is consistent with the characteristics of the specimens we studied.
Based on two of the paratypes, we observed that the oral tentacles are bifurcate or with four prolongations. We also found considerable variability in the presence or absence of denticles in the outermost teeth. This was also found in specimens of J. tomentosa and J. alisonae. Intra-specific and/or ontogenetic radular variation may explain these findings.
Jorunna pardus can be distinguished from J. funebris by its external features such as the absence of rings on the notum, the presence of dark spots on the edges of the foot and by differences in the reproductive system. Jorunna parva, an Indo-Pacific species, differs from J. pardus by the presence of higher brown caryophyllidia, dark rhinophoral clubs, mantle glands, bipinnate brachial leaves and the presence of five outermost teeth with up to five pectinate denticles.  External morphology: For a complete description of the species see Cervera et al. (1986).
Remarks: Examination of the type material confirms the original description (Cervera et al., 1986). This species is the only member of the genus known to have a non-prostatic deferent duct composed of two regions: a long convoluted proximal part, and a distal part that is short, thin, and slightly sinuous. The study of the reproductive system of the holotype verified the presence of this unique feature. According to Cervera, Garcı´a & Garcı´a (1986) J. tomentosa differs from J. onubensis by the presence of 5 -6 dark spots on both sides of the body, the absence of the non-prostatic deferent duct and labial cuticle with jaw elements. Jorunna onubenis differs from J. lemchei by the presence of a labial cuticle with jaw elements and the absence of spines on the penial papilla.
Jorunna onubensis can also be distinguished from J. tomentosa by the presence of jaw elements and the absence of outermost teeth with denticles. Although the external coloration of J. onubensis is similar to that of J. labialis, it differs by the length of the deferent duct. In J. labialis, the deferent duct is short with some small loops, whereas in J. onubensis the deferent duct is extremely long and convoluted. Also, the radula formula differs and J. labialis does not have a long nonprostatic deferent duct (see Table 4).
Background of living animals pale grey to light brown (Fig. 1M, N). Dorsum with a number of light grey to light brown patches of different sizes. Patches composed of an aggregate of minute dark brown spots more densely arranged near centre of notum. Minute dark spots homogeneously distributed over dorsum. Rhinophores pale grey speckled with dark brown and white on club. Branchial leaves dark grey. White spots around margin. Branchial and rhinophoral sheaths with small white glandular structures. Anterior border of foot notched and grooved. Oral tentacles conical. Hyponotum and notum same colour as dorsum.
Ampulla long, thick, tubular, convoluted in its distal part (character consistent in all specimens examined; Fig. 20B). Ampulla enters female gland in middle portion. Prostate flattened, granular, convoluted, divided into two different portions clearly distinguishable by their different texture and coloration. Deferent duct short, expanding into wide ejaculatory portion, opening into a common atrium with vagina. No penial hooks. Small accessory gland connected to atrium, consisting of a muscular portion with a pointed spine 2.2 mm long (Fig. 20C).
Penis entirely covered by cilia (Fig. 19G). Vagina short, with a loop in middle portion, connecting to large oval bursa copulatrix at its distal end. Another duct, connecting to rounded seminal receptacle and uterine duct, leading from bursa copulatrix. Seminal receptacle slightly smaller than bursa copulatrix (Fig. 20A).
Remarks: Jorunna ramicola is clearly distinguishable from other members of the genus by having a brown mantle with irregular brown to pale grey patches, light grey to light brown spots, denticulate innermost radular teeth, long and denticulate outermost teeth, and a labial cuticle with jaw elements. According to Miller (1996), J. ramicola also differs from other Indo-Pacific species by the presence of jaw elements that are rounded at the base and flat on top with up to 12 denticles. In the specimens studied here, the SEM photographs show that the jaw elements do not have as many regular denticles as those described by Miller (1996); however, other features studied match those that he described.
Jorunna ramicola can be distinguished from other Indo-Pacific species by the external morphology, radular morphology and anatomy. Jorunna rubescens, J. funebris and J. parva do not have jaw elements, whereas in J. ramicola they are well developed.
The dorsum of these three species has stripes (J. rubescens), rings (J. funebris) or elevated dark tubercles resembling dark spots in the mantle (J. parva). In contrast, J. ramicola has a brown to light grey dorsal colour with some irregular brown patches.
Jorunna hartleyi from Australia differs from J. ramicola by the absence of denticulate outermost teeth, as well as by the external coloration. Jorunna hartleyi has a pale pink background with large brown patches encircled with white papillae.
Two other Indo-Pacific species, J. pantherina from Australia and New Zealand and J. alisonae from Hawaii and Marshall Islands, differ from J. ramicola by the radular morphology and external coloration. In these species, the outermost teeth can be hamate and smooth (J. pantherina), smooth, or a single denticle may be present near the tip (J. alisonae); also, the innermost teeth in both species lack denticles. In addition, the dorsal colour of J. pantherina is purplish brown to dark grey or black with patches and opaque white spots, whereas J. alisonae is pale grey with two vertical lines of spots. Etymology: Jorunna osae is named after the Osa Conservation Area in Costa Rica.
External morphology: Body oval, elongate (Fig. 1O). Dorsum with elongate caryophyllidia, about 410 mm long. Caryophyllidia with a conical base, long spicules, a small elongate ciliated tubercle (Fig. 21D). Rounded, ciliated areas covering base of caryophyllidia. Body pale cream to yellow. Dorsum with light yellow-brown patches or spots, darker in some specimens. Patches homogeneously arranged on mantle, composing of minute dark spots. Some white glandular structures along mantle edge, larger posteriorly, visible only in living animals. Rhinophoral sheaths low, pale cream or yellow, speckled brown. Gill sheath pale cream or yellow speckled with brown spots. Rhinophores conical, pale cream or yellow with white apex, club lightly speckled brown. Gill pale, cream to yellow. Twelve erect bi-tripinnate branchial leaves and 10 lamellae in rhinophores in a 5 mm preserved length specimen (UCR-INBI01496535), imbricated in living animals, 12 bipinnate gills, 10 lamellae in a 6 mm preserved length specimen (CASIZ 162254), six tripinnate gills, 10 lamellae in a 3.5 mm preserved length specimen (UCR-INB03792183). Oral tentacles conical. No spots present on sole of foot or ventral part of notum. Innermost teeth hamate small, lacking denticles (Fig. 21A). Lateral teeth hamate, with a single cusp, lacking denticles (Fig. 21B). Teeth increase in size gradually towards medial portion of half-row. Outermost teeth small, elongate, lacking denticles (Fig. 21C).
Ampulla thick, long, convoluted in middle portion (Fig. 22C), dividing into a short oviduct entering female glands near their nidamental opening. Prostate flattened, granular, convoluted, dividing into two different portions clearly distinguishable by different texture and coloration. Deferent duct short, expanding into narrow ejaculatory portion.
Convoluted accessory gland (Fig. 22A) containing copulatory spine 600 mm in length. Spine curved, pointed (Fig. 22D). Vagina short, wide, with a curve in middle portion, connecting to small, oval bursa copulatrix at its proximal end. Another convoluted duct, connecting to oval seminal receptacle and uterine duct, leading from bursa copulatrix. Bursa copulatrix about same size as seminal receptacle (Fig. 22B).
Distribution: Pacific Coast of Costa Rica from Playa Gallardo to Playa Avellanas. It occurs in intertidal areas, under rocks to 13 m depth.
Remarks: This species differs from J. tempisquensis, another species found in the tropical American Pacific, by the presence of mantle glands on the margin, and the erect gill with imbricate branchial branches, and the external and radular morphology. Jorunna osae has a pale cream to brown background colour with light brown spots, whereas J. tempisquensis varies from white creamish to dark black purplish with dark brown spots on the dorsum and has a spreading gill. There are jaw elements present in J. tempisquensis, but these are absent in J. osae. Although J. osae has small and elongate outermost teeth, very similar to those in J. tempisquensis, the innermost teeth have a shorter base than those of J. tempisquensis.
Jorunna osae can also be distinguished from J. pardus from California and J. spazzola from Brazil and the Mediterranean by external and radular morphology. Jorunna pardus has a cream yellowish background colour with large dark brown spots, dark brown rhinophores with cream yellowish bases, and cream yellowish branchial leaves with dark brown tips. Jorunna spazzola has a light grey or white yellow to orange background with brown spots on each side of the notum lined up to form a row, and grey rhinophores, whereas in J. osae the background colour is pale cream to yellow with light brown-yellowish patches and the rhinophores are pale cream to yellow with white apices. In addition, J. spazzola has outermost teeth with up to four irregular denticles and innermost teeth sometimes with a small denticle, whereas in J. osae the outermost and innermost teeth lack denticles. Jorunna pardus also differs from J. osae by the presence of denticles in the outermost teeth.
Jorunna osae also differs from other similar species in the Mediterranean and Western Atlantic by its internal anatomy. For example, J. tomentosa has outermost teeth with denticles, whereas in J. osae the outermost are smooth. Jorunna lemchei and J. onubensis are distinct from J. osae; the former has an armed penis and the latter a very long non-prostatic duct. Jorunna osae lacks these features.
Jorunna tempisquensis new species (Figs 1P,23,24,  External morphology: Body oval, wide. Dorsum with caryophyllidia, about 200 mm in length (Fig. 23E). Caryophyllidia with a long, conical base, long spicules and a small, elongate, ciliated tubercle. Rhinophoral and branchial sheaths low. Eleven long, widely spreading, tripinnate branchial leaves and nine lamellae in rhinophores in 8 mm preserved length specimen (UCR-INBI01496480), 12 tripinnate gills, 11 lamellae in 14 mm preserved length specimen (CASIZ 167976), 10 tripinnate gills, 10 lamellae in 16 mm preserved length specimen (CASIZ 166890). Gills not imbricate, forming circle around anal papilla. Background colour of living animal light cream, light brown to dark purplish black. In darker specimens, centre of dorsum covering with large, light brown or black spots of different sizes. Dorsum with minute dark brown spots homogeneously arranged, covering tubercles and caryophyllidia. Rhinophores light cream to light brown, speckled with minute dark brown spots. Tips of rhinophores yellowish white. Base of branchial leaves dark brown to almost purple with light yellow tips and some minute brown spots.
Anterior border of foot notched, grooved with minute brown spots more densely arranged on edges of mantle. Oral tentacles conical. Small brown spots present on ventral side of head including tentacles. Foot same colour as dorsum, with some minute dark spots irregularly distributed. Posterior end of foot visible dorsally when the animal moves.
Ampulla short, convoluted, branching into a short oviduct and prostate (Fig. 24C). Oviduct enters female gland in centre of mass. Prostate flattened, folded, granular (Fig. 24C), connecting to a long, convoluted duct expanding into short ejaculatory portion of deferent duct. Deferent duct opening into common atrium with vagina. Vagina long, wide as deferent duct. Convoluted accessory gland (Fig. 24A), connecting to atrium with a distinct stylet sac containing a 550 mm long copulatory spine, almost entirely straight, pointed (Fig. 24D). Vagina connecting to oval bursa copulatrix at its proximal end. Another duct leading from bursa copulatrix to uterine duct and seminal receptacle. Bursa copulatrix slightly larger than oval seminal receptacle (Fig. 24B).
Distribution: Southern Mexico and Costa Rica. It occurs in intertidal areas, under rocks from 0 to 2 m depth. Remarks: Jorunna tempisquensis can be distinguished from J. osae, also from Costa Rica, by the external coloration, the presence of jaw elements, the absence of mantle glands, by having a spreading rather than erect gill and by differences in radular morphology such as the shape of the innermost teeth.
Jorunna tempisquensis also differs from J. pardus from California and J. spazzola from Brazil and the Mediterranean by the external and radular morphology. Jorunna pardus has cream yellowish background colour with large dark brown spots, dark brown rhinophores with cream yellowish bases and cream yellowish brachial leaves with dark brown tips. Jorunna spazzola has a light grey or white yellow to orange background with brown spots on each side of the notum forming a row, whereas J. tempisquensis has a cream white to dark purplish brown background with light brown spots on the dorsum. Jorunna pardus also differs from J. tempisquensis by the absence of jaw elements and by the presence of denticles on the outermost teeth. Additionally, Jorunna spazzola has outermost teeth with up to four irregular denticles and innermost teeth sometimes with a small denticle. Jorunna tempisquensis lacks denticles on the outermost and innermost teeth.

Part B. Misidentified and doubtful species of Jorunna and Kentrodoris
There are several species of dorids which have been erroneously classified as members of the genus Jorunna (K. pseudofusca, J. marchadi), some whose status within Jorunna remains uncertain (J. atypha, K. nigra, Jorunna sp., Kentrodoris (?) sp.) and others which were included by  as 'potential' members of the genus Jorunna (Platydoris inframaculata, Doris infranaevata), and which recently have been considered as members of Platydoris by Dorgan, et al. (2002) and Dayrat (2006). These are discussed in detail in this section.
Remarks: According to Bergh (1881), J. atypha has a greyish white to light yellowish-white background colour, white rhinophores with brown upper lamellae, white branchial sheaths, a yellowish foot, rhinophores with 20 lamellae, 11 bi-to tripinnate branchial leaves, short and flat caryophyllidia, broad oral tentacles, an accessory gland with a spine, hamate midlateral teeth and outermost teeth with 3-4 denticles. The presence of jaw elements in this species could not be confirmed by Bergh (1881) due to the bad preservation of the single specimen. Bergh (1881) also mentioned that the shape of the oral tentacles is quite different from other members of the genus Jorunna.
Due to the incomplete description provided by this author (also emphasized by , and his own hesitation about whether this species really belongs to the genus Jorunna, the generic position of this species remains unclear. Kentrodoris nigra Risbec, 1928 Kentrodoris nigra Risbec, 1928: 91, pl. 2, fig. 6 (New Caledonia, type probably lost, not at MNHN). Jorunna sp. Pruvot-Fol, 1953Jorunna sp. Pruvot-Fol, 1953 Remarks: Pruvot-Fol (1953) described Jorunna sp. from Dakar, Senegal. According to , the little information provided in the original description is insufficient to allow placement in this genus. There is no mention of the presence of an accessory gland with a spine, one of the principal features of Jorunna.
Remarks: Burn (1966) collected two specimens from Port Phillip Bay, southeastern Australia. According to Burn, the background colour of the animals is creamy-white with dark brown spots scattered over the back. Unfortunately, he did not provide any further description that would allow us to determine the generic position of the specimens. Distribution: Amboina, Sri Lanka, Philippines (Dorgan et al., 2002).

Platydoris inframaculata (Abraham, 1877)
Remarks:  included this species as a potential member of Jorunna. She was not certain whether this species belonged to Jorunna or Platydoris, although she agreed that " Farran's (1905) figures of teeth and elements of penial spines and vaginal plates certainly belong to a Platydoris". Dorgan et al. (2002) placed this species in the genus Platydoris based on the similarity of the external morphology of the original specimens, specifically the presence of large and small brown spots on the ventral side, with the preserved specimens they examined from the Philippines.
Remarks:  included this species as a potential member of Jorunna, stating that it was possibly synonymous with J. tomentosa.
Based on the examination of the external morphology of the holotype we confirm, with Dayrat (2006), that this species belongs to the genus Platydoris. The body is depressed, the mantle is wide with a crenulate border, and the oral tentacles are flattened and grooved.
The drawings of the radular morphology and copulatory spine made by Bergh (1878) show a very wide base and short copulatory spine instead of a slender and long spine that is characteristic of the genus Jorunna. Also, the wide base in most of the outermost teeth is different from the thin base and elongated shape of the outermost teeth of Jorunna. Since some of the features of J. maima resemble those present in the genus Sclerodoris, the identity of this species remains unclear.

Phylogenetic methods
The characters described below were placed in a data matrix (Table 1) using MacClade 4.0 (Maddison & Maddison, 2000).
Data were analysed by Phylogenetic Analysis Using Parsimony (PAUP) version 4.0 (Swofford, 2000) to find the most parsimonious phylogenetic tree, using an heuristic search. The stepwise addition option of Random Trees was used, with 100 replicates. The multistate characters were treated as unordered and unweighted. The percentage of missing data in the matrix was 4%. The strict consensus tree was used to trace character evolution using the character trace option in MacClade 4.0 (Maddison & Maddison, 2000). A Bremer support (or decay) analysis was performed using a heuristic search by PAUP to estimate branch support (Bremer, 1994).
Characters were polarized using the outgroup taxa Discodoris boholiensis Bergh, 1877 andDoris verrucosa (Linnaeus, 1758), which were used for rooting the tree. Diaulula sandiegensis (Cooper, 1863) was also included for comparison since, like Jorunna, it has caryophyllidia. This outgroup selection was made following Valde´s &Gosliner (2001) andValde´s (2002), who showed that these taxa are basal to the rest of the caryophyllidia-bearing dorids, and that Diaulula sandiegensis forms a polytomy with Jorunna.

Taxa
Sixteen members of Jorunna were selected to sample the morphology of the group and 35 characters were included in the data matrix (Table 1). Data were obtained from the examination of preserved and living specimens, from some types and, in a few cases, from the literature (see Systematic Descriptions). Poor original descriptions and lack of living animals for studies of J. lemchei, J. hartleyi and J. evansi added noise to the analysis. These species were excluded a posteriori from the analysis due to the lack of complete anatomical information.

Characters
The characters used are listed below. Thirty-five characters were coded as either binary or multistate. The character states are indicated with numbers: 0, presumed plesiomorphic condition that was tested by the phylogenetic analysis; 1-3, apomorphic states. All multi-state characters were treated as unordered. Characters 1-22 are from external morphology, 23-28 from radular morphology, and 29 -35 from the reproductive system.
(1) Oral tentacles: Some species of cryptobranch dorids, including Doris verrucosa, lack oral tentacles (0). In some other cryptobranch dorids such as Jorunna there is a pair of oral tentacles around the buccal area that are digitiform in shape (1).
(2) Prolongation of the posterior foot: In most Jorunna and some other dorids, the foot is relatively long and extends a short distance from the posterior end of the mantle (1) (1). On the contrary, other species as well as the outgroup taxa lack these spots (0).  (1) or singly (2). (23) Outer-lateral teeth shape: The shape of the outermost radular teeth is very different among species. In some Jorunna and the outgroup, the teeth are hamate (0), whereas in other species of Jorunna they have a plate-like shape (1). (24) Labial cuticle: Jaw elements are present in most of the cryptobranch dorids including some species of Jorunna (0), whereas in others the labial cuticle is smooth (1). (25) Shape of the jaw elements: In some species the jaw elements are well developed, with a rectangular shape (1), whereas in others the jaw elements are amorphous (0). (26) Outermost lateral teeth: Two character states can be recognized in the outer lateral teeth: they can be smooth (0) as in most species of Jorunna, or in a few species denticulate (1). (27) Innermost radular teeth with denticles: Two members of Jorunna have one to three denticles on the inner lateral tooth (1), whereas in other species and in the outgroup taxa the innermost tooth is smooth (0). (28) Basal extension of lateral teeth: The radula of most species of Jorunna has a swelling on the lateral teeth that can be situated basally (0), or in the middle of the teeth (1). (29) Shape of the copulatory spine tip: The shape of the copulatory spine can be curved with a little hook (1), or straight and lacking a hook (0).
(30) Stomach free: In Jorunna the stomach is not covered by the digestive gland (1), whereas in the outgroup the stomach is covered by the digestive gland (0). (31) Accessory gland: The accessory gland consists of a structure located next to the distal part of the female gland mass, which opens into the albumen gland. There is an accessory gland present in all species of Jorunna (1). Doris, Discodoris and Diaulula lack an accessory gland (0). (32) Copulatory spine: All species of Jorunna have a copulatory spine present in the accessory gland, which is used in the copulatory process (1). The outgroup taxa lack a copulatory spine (0). (33) Shape of accessory gland: The accessory gland can be globular in shape (0), or not globular (1). (34) Prostate portions: In Jorunna and Diaulula (as well as other genera such as Halgerda and Asteronotus), the prostate has two parts that are well differentiated in colour and texture (1), whereas in most dorids the prostate has a single part (0). (35) Relative size of bursa: The bursa colpulatrix can be the same size as the seminal receptacle (0), or much larger (1).

RESULTS AND DISCUSSION
The heuristic analysis of the data set produced five equally parsimonious trees with a length of 65 steps. The Consistency Index (CI ¼ 0.585) and the Retention Index (0.675) show that 42% of all characters exhibit homoplasy (reversal and/or parallelism). The strict consensus tree is shown in Figure 26A. The italic numbers indicate reversals or parallelism. All the branches but one were present in each of the equally parsimonious trees, i.e. 100% in the majority rule tree (Fig. 26B). The strict consensus tree shows that the genus Jorunna is a monophyletic group supported by four synapomorphies, with Diaulula sandiegensis being the sister group to Jorunna (Fig. 26A). The synapomorphies of the genus Jorunna are: prolongation of the posterior foot (2), stomach free (30), accessory gland (31) and copulatory spine (32). The synapomorphies that support the sister relationship of Diaulula sandiegensis and Jorunna are: oral tentacles (1), angle of the gills (3), anterior border of the foot (5), ciliated tubercle (6), shape of the