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This paper discusses the systematics of the aeolid genus Baeolidia Bergh, 1888. To date, this monophyletic genus is the most diverse within Aeolidiidae with sixteen valid species. Excluding Baeolidia cryoporos Bouchet, 1977, the genus is restricted to the Indo-Pacific and Eastern Pacific. Species of Baeolidia show a huge intrageneric variability in several morphological characters. Only oral glands, if present, may distinguish Baeolidia from other aeolidiids genera. Aeolidiella occidentalis Bergh, 1875, Aeolidiella faustina Bergh, 1900 and Spurilla orientalis Bergh, 1905 are transferred to Baeolidia but they are considered nomina dubia. Five new species, Baeolidia rieae sp. nov., Baeolidia variabilis sp. nov., Baeolidia lunaris sp. nov., Baeolidia gracilis sp. nov. and Baeolidia scottjohnsoni sp. nov. are described.
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Accepted by M. Schroedl: 28 Mar. 2014; published: 28 May 2014
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http://dx.doi.org/10.11646/zootaxa.3802.4.5
http://zoobank.org/urn:lsid:zoobank.org:pub:4095CA09-8EA4-4941-8286-32E95F0206AE
Review of Baeolidia, the largest genus of Aeolidiidae (Mollusca: Nudibranchia),
with the description of five new species
LEILA CARMONA
1,4
, MARTA POLA
2
, TERRENCE M. GOSLINER
3
& JUAN LUCAS CERVERA
1
1
Departamento de Biología, Facultad de Ciencias del Mar y Ambientales, Campus de Excelencia Internacional del Mar (CEI·MAR,)
Universidad de Cádiz. Polígono Río San Pedro, s/n, Ap.40. 11510 Puerto Real (Cádiz), SPAIN.
2
Laboratorio de Biología Marina, Departamento de Biología, Edificio de Biología, Campus de Excelencia Internacional UAM+CSIC,
Universidad Autónoma de Madrid, C/ Darwin, 2, 28049 Madrid, SPAIN.
3
Department of Invertebrate Zoology, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco,
CA 94118, USA.
4
Corresponding author. E-mail: leila.carmona@uca.es; phone: +34 956016012
Abstract
This paper discusses the systematics of the aeolid genus Baeolidia Bergh, 1888. To date, this monophyletic genus is the
most diverse within Aeolidiidae with sixteen valid species. Excluding Baeolidia cryoporos Bouchet, 1977, the genus is
restricted to the Indo-Pacific and Eastern Pacific. Species of Baeolidia show a huge intrageneric variability in several mor-
phological characters. Only oral glands, if present, may distinguish Baeolidia from other aeolidiids genera. Aeolidiella
occidentalis Bergh, 1875, Aeolidiella faustina Bergh, 1900 and Spurilla orientalis Bergh, 1905 are transferred to Baeoli-
dia but they are considered nomina dubia. Five new species, Baeolidia rieae sp. nov., Baeolidia variabilis sp. nov., Bae-
olidia lunaris sp. nov., Baeolidia gracilis sp. nov. and Baeolidia scottjohnsoni sp. nov. are described.
Key words: Cladobranchia, molluscan diversity, morphology, new species, systematics
Introduction
The aeolidiid genus Baeolidia was introduced by Bergh (1888) based on a single preserved specimen of Baeolidia
moebii from Mauritius. The original description has contradictory information about the cerata arrangement, which
has been considered as one of the most important diagnostic characters in Aeolidiidae. Bergh (1888) originally
described the cerata in rows and then in arches (page 779 of that contribution). This lack of clarity has produced a
great deal of confusion in the literature about the morphological characteristics of this genus. While some authors
characterized Baeolidia by having the cerata in arches (Miller 2001) others ascribed or transferred species that have
rows instead of arches to this genus (Gosliner 1985). Additionally, Baeolidia has been considered as a junior
synonym of Spurilla Bergh, 1864 (Rudman 1982), whereas some authors that have considered Baeolidia as a valid
genus rejected the validity of Limenandra (Gosliner 1980, 1985; Valdés et al. 2006; Gosliner et al. 2008). Some
species have been assigned to Baeolidia: Baeolidia moebii Bergh, 1888; Baeolidia major Eliot, 1903; Baeolidia
japonica Baba, 1933; Baeolidia fusiformis Baba, 1949; Baeolidia benteva Er. Marcus, 1958; Baeolidia cryoporos
Bouchet, 1977 and Baeolidia palythoae Gosliner, 1985. Furthermore, Limenandra nodosa Haefelfinger & Stamm,
1958; Aeolidiopsis harrietae Rudman, 1982 and Spurilla australis Rudman, 1982 were also transferred to
Baeolidia because of their ceratal arrangement or/and the ornamentation of the rhinophores (Gosliner1985; Miller
2001).
The first comprehensive study on Aeolidiidae (Carmona et al. 2013) rendered Baeolidia as a monophyletic
genus once Spurilla salaamica Rudman, 1982 and Aeolidiopsis ransoni Rudman, 1982 were transferred to
Baeolidia. The latter contribution also showed that not only the ceratal arrangement but also the rhinophorial
ornamentation and the position of the anus lacked of any phylogenetic significance within Baeolidia. Additionally,
Carmona et al. (2013) pointed out the existence of, at least, three undescribed species (Fig. 1). The molecular
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phylogeny of Aeolidiidae (Carmona et al. 2013) also showed that Baeolidia was closely related with Spurilla and
Berghia Trinchese, 1877, although the relationships among these three genera were not resolved. Recently, the
morphological characteristics of Spurilla, Berghia, and Limenandra Haefelfinger & Stamn, 1958 have been re-
defined (Carmona et al. 2014a, b, c, respectively). While the species of Spurilla have a broad body, cerata curved
inwards and perfoliate rhinophores, Berghia species usually are slender, with papillate rhinophores. Nevertheless
the systematics of the genus Baeolidia have not been studied. Therefore, the main objectives of the present study
are: 1) Undertake a general revision of Baeolidia (revising species of all names available, when possible); 2)
Describe five new species; 3) Compare the morphological characters of the genus Baeolidia with Spurilla, Berghia
and Limenandra.
FIGURE 1. Phylogeny of Baeolidia, based on Carmona et al. (2013). Names on right side of vertical bars refer to revised
classification.
Materials and methods
Literature review. A comprehensive review of the literature was conducted to determine the valid names for the
species recognized in the present study. After the description of the type species, all available names for Baeolidia
species are organized and discussed in this paper according to the year of publication. In the synonymy lists,
references to the original description of the valid name and all synomyms (basionyms and primary synonyms) as
well as the first proposed change of binomen (secondary synonyms) are included, but subsequent references are
not.
Morphology. Whenever possible, two or more specimens of each species were examined anatomically.
Specimens were dissected by dorsal incision. Their internal features were examined and drawn through a dissecting
microscope with the aid of a camera lucida. Special attention was paid to the morphology of the reproductive
system and oral and salivary glands. The buccal mass was removed and dissolved in 10% sodium hydroxide until
the radula was isolated from the surrounding tissue. The radula was then rinsed in water, dried, and mounted for
examination by scanning electron microscopy (SEM).
The specimens examined are deposited in the California Academy of Sciences, CASIZ (San Francisco, USA)
and the Museo Nacional de Ciencias Naturales, MNCN (Madrid, Spain), being named in agreement to the
classification of Aeolidiidae proposed by Carmona et al. (2013).
Nomenclatural acts. This published work and the nomenclatural acts it contains have been registered in
ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be
resolved and the associated information viewed through any standard web browser by appending the LSID to the
prefix "http://zoobank.org/". The LSID for this publication is: LSID urn:lsid:zoobank.org:pub:1DB70C57-A6EB-
491D-A0FD-0B1652E5B61E
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Results
Systematics
Nudibranchia Cuvier, 1817
Cladobranchia Willan & Morton, 1984
Family Aeolidiidae Gray, 1827
Genus Baeolidia Bergh, 1888
Diagnosis of the genus Baeolidia based on Bergh (1888). Forma corporis minus depressa quam in Aeolidiis
propriis; rhinophoria annulate moriformia, tentacula digitiformia; papillae dorsales non numerosae, compressae,
sat breves; margo anterior podarii profunde sulcatus, angulis fortiter productis.- Penis inermis. Mandibulae
applanatae: margo masticatorius sublaevis. Radula uniseriata, dentibus simpliciter pectinatis.
Body low. Rhinophores perfoliate. Oral tentacles digitiform. Cerata scarce, compressed, short. Anterior margin
grooved. Penis unarmed. Jaws flattened, masticatory border almost smooth. Radula uniseriate, teeth with simple
denticulation (translated from Bergh, 1888)
Baeolidia moebii Bergh, 1888
(Figs. 2A–B, 3A, 4A–B, 5, 6A)
Baeolidia moebii Bergh, 1888: 778, pl. LXXIX, Figs. 10–16, pl. LXXX, Figs. 1–4.
Baeolidia major Eliot, 1903: 252.
Baeolidia major amakusana Baba, 1937: 335.
Berghia major (Eliot, 1903): Edmunds 1969, 467, Fig. 10.
Spurilla major (Eliot, 1903): Rudman 1982, 169, Figs. 18–19.
Type locality. Grand Bay, Rodrigues Island, Mauritius.
Type material. To our knowledge no type material remains. We designate the specimen MNCN 15.05/63447
from Mozambique as the neotype in order to avoid confusion with similar species.
Material examined. Neotype: MNCN 15.05/63447, one specimen, dissected, 18 mm in length preserved,
Mozambique, Zavora Bay, collected by Yara Tibiriça, January 2012. Other material: CASIZ 177602, one
specimen, dissected, 10 mm in length preserved, Philippines, Luzon Island, Batangas Bay, Calumpan Peninsula,
collected by Terrence M. Gosliner, 16 April 2008; CASIZ 157017, two specimens, dissected, 25 and 24 mm in
length preserved, Philippines, Luzon Island, Batangas Province, Balayan Bay, Seafari Beach, collected by Ángel
Valdés and Mary James Adams, 12 May 2001; CASIZ 186211, one specimen, dissected, 10 mm in length
preserved, Philippines, Luzon, Batangas Province, Calumpan Peninsula, Batangas Bay, collected by Alicia
Hermosillo, 04 May 2011; CASIZ 180327, one specimen, dissected, 6 mm in length preserved, Hawaii, Maui,
Airport Beach, collected by Cory Pittman, 04 October 2008; CASIZ 076055, one specimen, dissected, 16 mm in
length preserved, Mexico, Baja California, Gulf of California, San Jose Island, Punta Colorado, collected by
Antonio J. Ferreira 15 July 1971.
Geographical distribution. Originally described from Mauritius (Bergh 1888), it is also known from the
Seychelles (Gosliner et al. 2008), Tanzania (Eliot 1903; Edmunds 1970), Mozambique (present study), the
Reunion Island (Bidgrain 2013), Australia (Cobb & Mullins 2013), Indonesia (Gosliner et al. 2008), the
Philippines (Gosliner et al. 2008), Japan (Baba 1933b, 1937, 1955; Ono 1999, 2004; Nakano 2004) and the
Marshall Islands (present study). This species is also found in the Eastern Pacific: Hawaii (Kay 1979; Gosliner
1980), tropical Pacific North America (Gosliner et al. 2008) and Mexico (Farmer 1966). Recently, it has also been
reported in Turkey (Turk & Furlan 2011).
External morphology (Figs. 2A–B, 3A): Body large, broad, tapering gradually towards posterior end of foot.
Foot corners tentaculiform. Body colour variable, commonly greyish white or brownish green (Fig. 2A). Some
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specimens reddish (Fig. 2B). Background colour brownish green, overlaid with reticulate ochre pattern. Dorsum
usually with bright white patches. Whitish or yellow ring on anteriormost edge of head, continuing towards oral
tentacles and forming a distinct light band; both, ring and band, may be interrupted. Rhinophores, oral tentacles and
foot corners same as body colour. Rhinophores shorter than oral tentacles. Rhinophores studded with minute knobs
(Fig. 3A); apex white. Oral tentacles with white tips.
FIGURE 2. Photographs of the living animals. A, B, Baeolidia moebii: (A) specimen from the Philippines, photo by Terrence
M. Gosliner, CASIZ 177602; (B) specimen from Mozambique, photo by Yara Tibiriça, MNCN 15.05/63447; (C) Baeolidia
macleayi, specimen from Australia, photo by Terrence M. Gosliner, CASIZ 71923; (D-E) Baeolidia japonica: (D) specimen
from Japan, photo by Akira Kawahara, CASIZ 1814520; (E) specimen from Marshall Islands, photo by Scott Johnson, CASIZ
186795; (F) Baeolidia ransoni, specimen from the Philippines, photo by Terrence M. Gosliner, CASIZ 186208.
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FIGURE 3. Different papillate rhinophores in Baeolidia: (A) from Rudman (1982); (B) from Gosliner (1985). Scale bars: A,
3mm; B, 1 mm; C, 1.2 mm; D, 1.5 mm.
Cerata length variable. Cerata flattened, almost leaf-like. Branches of digestive gland brownish-green, shining
through translucent body wall. Whitish patches on outer side of cerata, probably with bright yellow marks. Cerata
usually with bright yellow subapical band and just below it a purplish area. Apex translucent white. Cerata
composed of up to nine arches, leaving a distinct gap between pre and post-pericardial groups. Each arch with 4–15
cerata, decreasing in size towards foot. Anus cleioproctic, within second right arch. Genital opening placed among
cerata of anteriormost group on right side.
Anatomy. Masticatory border of jaws smooth (Fig. 4A). Radular formulae 17 x 0.1.0 (MNCN 15.05/63447, 18
mm), 19 x 0.1.0 (CASIZ 076055, 16 mm), 20 x 0.1.0 (CASIZ 177602, 10 mm) and 22 x 0.1.0 (CASIZ 180327, 6
mm). Radular teeth bent upwards with up to 100 elongate, fine and acutely pointed denticles from side to side,
without central notch or central cusp (Fig. 4B). Teeth progressively smaller towards posterior region of radula. Oral
glands fragile, small and spongy, lying dorso-laterally to buccal bulb (Fig. 5). Oral gland morphology variable,
from rounded to elongate. Salivary glands large.
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FIGURE 4. Scanning electron photographs. (A–B) Baeolidia moebii (CASIZ 180327): (A) Detailed view of the masticatory
border; (B) Radular teeth; (C–D) Baeolidia macleayi (CASIZ 195247): (C) Detailed view of the masticatory border; (D)
Radular teeth; (E–F) Baeolidia japonica (CASIZ 181357); (E) Detailed view of the masticatory border; (F) Radular teeth. Scale
bars: A, 10 µm; B, 30 µm; C-D, 10 µm; E, 100 µm; F, 150 µm.
Reproductive system diaulic (Fig. 6A). Preampullary duct widening into elongate ampulla. Postampullary duct
dividing into oviduct and vas deferens. Vas deferens elongate, moderately wide, penetrating into wider proximal
portion of penial sac with unarmed penial papilla. Receptaculum seminis ovoid, short stalk connecting to short
oviduct, before latter forms female glands. Vagina ventral to penis.
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Remarks. This species has not been studied under the name of Baeolidia moebii since its original description
(Bergh 1888). When Eliot (1903) described B. major, he pointed out that the latter could be merely a full-grown
individual of B. moebii since the main difference between both species was the size. Nevertheless, Eliot (1903)
finally described B. major as a valid species, owing to the ambiguities in the original description of B. moebii.
Taking into consideration Eliot’s statements together with the proximity of both type localities (Mauritius for B.
moebii and Zanzibar for B. major), Carmona et al. (2013) considered B. major as a junior synonym of B. moebii.
Neither Bergh (1888) nor Eliot (1903) mentioned the presence of oral glands in B. moebii. Only Rudman (1982)
pointed out the existence of salivary glands in this species. Two of our specimens possessed oral glands, and
salivary glands were found twice. This variability could be an artefact of the state of preservation of these
specimens. An alternative hypothesis could be an intraspecific variation for these two characters, which has also
been observed in Limenandra nodosa (Carmona et al. 2014c).
FIGURE 5. Oral glands of Baeolidia. Scale bar: 0.5 mm.
Despite the large variability in colouration, our specimens identified as Baeolidia moebii match with the
information given by Bergh (1888) and Eliot (1903) as well as with more recent descriptions (Edmunds 1969;
Gosliner 1980; Rudman 1982). Gosliner (1985) stated the presence of a penial gland in the reproductive system of
B. moebii (identified as Berghia major). The only aeolidiid where this condition has been described is Limenandra
nodosa (Schmekel 1970), although Carmona et al. (2014c) did not find any traces of a penial gland. Since none of
the specimens here examined exhibited this structure, the existence of a penial gland for B. moebii is not confirmed
(Fig. 6A).
Finally, excluding Baeolidia australis (see B. australis remarks), B. moebii colouration is clearly
distinguishable from the remaining species of this genus. In terms of internal anatomy, the radular morphology of
Baeolidia moebii is quite particular since teeth are bent and do not have any central notch or denticle. So far, only
B. australis and B. lunaris sp. nov. have similar radular teeth. Baeolidia moebii and B. lunaris sp. nov. differ in size
(B. moebii is larger than B. lunaris sp. nov.), and rhinophorial ornamentation (B. lunaris sp. nov. has elongate
papillae instead of minute knobs and colouration). Additionally, the ampulla of B. moebii is larger than the ampulla
found in B. lunaris sp. nov. Differences between B. moebii and B. australis are recorded in B. australis remarks.
This species was studied molecularly by Carmona et al. (2013) (Fig. 1).
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FIGURE 6. Reproductive system. (A) Baeolidia moebii, specimen from Mozambique (MNCN 15.05/63447); (B) Baeolidia
macleayi, specimen from Australia (CASIZ 195247); (C) Baeolidia japonica, specimen from Marshall Islands (CASIZ
181357); (D) Baeolidia ransoni, specimen from the Philippines (CASIZ 186209). Scale bars: A–D, 0.5 mm. Abbreviations:
am, ampulla; fgm, female gland mass; ps, penial sac; rs, receptaculum seminis; va, vagina; vd, vas deferens.
Baeolidia macleayi (Angas, 1864)
(Figs. 2C, 3B, 4C–D, 6B)
Aeolis macleayi Angas, 1864: 65, pl. VI, Fig. 4.
Coryphella macleayi (Angas, 1864): Bergh 1878, 16.
Spurilla macleayi (Angas, 1864): Burn 1969, 96.
Type locality. Port Jackson, New South Wales.
Type material. To our knowledge no type material remains. We designate the specimen CASIZ 071923 (6
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mm) from Long Reef, North of Sydney, New South Wales, Australia, as the neotype in order to avoid confusion
with similar species.
Material examined. Neotype: CASIZ 195247, one specimen, dissected, 6 mm in length preserved, Australia,
New South Wales, North of Sydney, Long Reef, collected by Terrence M. Gosliner, 17 September 1989. Other
material: CASIZ 071923, two specimens, dissected, 8 and 6 mm in length preserved, Australia, New South Wales,
North of Sydney, Long Reef, collected by Terrence M. Gosliner, 17 September 1989.
Geographical distribution. Originally described from Port Jackson, Australia (Angas 1864), this species is
also known from New Zealand (Burn 1962, 1969; Coleman 2001; Miller 2001; Burn 2006; Rudman 2007a).
External morphology (Figs. 2C, 3B): Body fairly short, broad, tapering close to posterior end of foot. Foot
corners tentaculiform. Body colour translucent. Some specimens with white and lemon-yellow speckling over
dorsum and head. Bright orange mark on pericardium with different intensity; it continues through head and
posterior end of foot. Rhinophores ribbed (Fig. 3B), shorter than oral tentacles. Proximal two-thirds reddish or
brownish. Distal part of rhinophores with lemon-yellow pigment. Oral tentacles translucent, may have white and
lemon-yellow spots.
Cerata elongate and slender, or moderately short and bulb-shaped. Cerata translucent, being visible the
brownish or bluish ramifications of digestive gland. Cnidosacs white. Cerata with bright lemon-yellow ring
subapically. White spots over ceratal surface might form a second ring below the yellow one. Cerata composed of
four arches and three rows, leaving a distinct gap between pre and post-pericardial groups. Each group with 3–10
cerata, decreasing in size towards foot. Anus cleioproctic, below second right row. Genital aperture below first
right arch.
Anatomy. Masticatory process smooth (Fig. 4C). Radular formula 16 x 0.1.0 (CASIZ 195247, 6 mm). Radular
teeth bilobed with 30–40 elongate and acutely pointed denticles on either side of central cusp (Fig. 4D). Central
cusp prominent, elongate. Oral glands spongy, moderately large, delicate, with uniform diameter throughout most
of length. Oral glands dorso-laterally to buccal bulb. Salivary glands absent.
Reproductive system diaulic (Fig. 6B). Preampullary duct widening into moderately short ampulla.
Postampullary duct dividing into oviduct and vas deferens. Vas deferens inserting into wider proximal portion of
penial sac with unarmed penial papilla. Receptaculum seminis heart-shaped, short stalk connecting to short
oviduct, before latter forms female glands. Vagina ventral to penis.
Remarks. We transfer Aeolis macleayi to Baeolidia because of the type of oral glands, which are exclusive of
this genus in Aeolidiidae.
Excluding Baeolidia faustina (see B. faustina remarks), the colouration and the shape of the cerata distinguish
B. macleayi from the remaining Baeolidia species. Regarding the radular morphology, the teeth found in B.
macleayi are similar to B. rieae sp. nov., B. variabilis sp. nov. and B. gracilis sp. nov. Nevertheless, the central
cusp of B. macleayi is larger than the minute central cusp found in the species mentioned above. Moreover, the
shape of the receptaculum seminis is quite different between B. macleayi and B. rieae sp. nov., B. variabilis sp.
nov. and B. gracilis sp. nov.: in B. macleayi it is heart-shaped; in B. rieae sp. nov. and B. variabilis sp. nov. it is
bean-like, and the receptaculum seminis of B. gracilis sp. nov. is ovoid. In addition, it is worth mentioning that B.
macleayi is a native species from Australia and New Zealand, whereas B. rieae sp. nov. was found in Japan, B.
gracilis sp. nov. in the Philippines and
B. variabilis sp. nov. is from the Philippines, Papua New Guinea and the
Marshall Islands.
The oral glands of B. macleayi are first described and reported here.
Baeolidia occidentalis (Bergh, 1874)
Aeolidiella occidentalis Bergh, 1874: 397, pl. VIII, Figs. 9–19.
Spurilla occidentalis (Bergh, 1874): Labbé 1930, 623.
Type locality. The Antilles, not very far from Saint Thomas.
Type material. According to Bergh (1874), the material was deposited in the Museum of Copenhagen. The
registration number of the holotype of Baeolidia occidentalis is ZMUC-GAS 2042.
Geographical distribution. So far, this species is only known from the Antilles (Bergh, 1874).
External and internal morphology. Not completely described by Bergh (1874). No specimens available for
the present study.
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Remarks. Bergh (1874) described this species, Aeolidiella occidentalis, based on three specimens. Since they
were already preserved and in a bad state of conservation, Bergh (1874) did not describe the colouration of the
living animal. The radular teeth, the morphology of the cerata and the oral glands described by Bergh (1874) seem
to associate this “species” with the genus Baeolidia. Although Bergh’s original description does not allow
identifying this “species”, B. occidentalis have been reported in some contributions about Sargassum and its
ecological aspects (e.g. Adams 1960; Thiel & Gutow 2005). However, none of these contributions provided a
detailed description of this “species” or of the specimen identified as B. occidentalis. The lack of any singular
morphological feature in this species leads us to conclude that Baeolidia occidentalis should be treated as nomen
dubium.
Baeolidia faustina (Bergh, 1900)
Aeolidiella faustina Bergh, 1900: 235, pl. 20, Figs. 39–40.
Spurilla faustina (Bergh, 1900): Miller 2001, 641, Figs. 4–5.
Type locality. Tasmania.
Type material. To our knowledge no type material remains.
Geographical distribution. To date, only known from Tasmania (Bergh 1900), New Zealand (Miller 2001),
and Australia (Burn 1962, 1969, 2006).
External and internal morphology. Not properly described by Bergh (1900). No specimens available for the
present study.
Remarks. Based on the original description and the figure of the radular tooth (Bergh 1900), it is very likely
that this species belongs to Baeolidia. Nevertheless, Bergh (1900) did not provide any distinctive feature, which
has blurred the identity and validity of this species. Since Bergh (1900), only Burn (1962) and Miller (2001) have
studied this species but both authors had some difficulties separating B. faustina from B. macleayi. Burn (1962)
distinguished both species mainly by the position of the anus and the external colouration. Seven years later, Burn
(1969) transferred all the specimens previously identified as B. faustina to B. macleayi. Finally, Burn (2006)
rendered B. faustina as junior synonym of B. macleayi. Hence, since this name cannot be assigned to specimens,
we conclude that Baeolidia faustina should be considered as nomen dubium.
Baeolidia orientalis (Bergh, 1905)
Spurilla orientalis Bergh, 1905: 223, pl. XIX, Figs. 29–31.
Type locality. Tual Reef, Kei Island, Indonesia (Station 258 of the Siboga Expedition).
Type material. The material was deposited in the National Museum of Natural History, Leiden. The
registration number of the syntype of Baeolidia orientalis is ZMA.MOLL.139366
Geographical distribution. To date, this species is only known from the Indo-Pacific (Bergh 1905).
External and internal morphology. Not properly described by Bergh (1905). No specimens available for the
present study.
Remarks. Since Bergh (1905), new specimens of Spurilla orientalis have not been discovered again.
However, its name can be found in the literature as part of the discussion of other Spurilla species (e.g. Er. Marcus
1961; García-Gómez 2002). Based on the ceratal insertion (in arches and rows), the figure of the radular tooth
(Bergh 1905) and the geographical distribution of Spurilla and Baeolidia (Carmona et al. 2013), this “species”
should be ascribed to Baeolidia. However, the lack of any singular morphological characters in the original
description of Baeolidia orientalis leads us to render this species as nomen dubium.
Baeolidia japonica Baba, 1933
(Figs. 2D–E, 3C, 4E–F, 6C).
Baeolidia japonica Baba, 1933: 282, Fig. 8.
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Spurilla japonica (Baba, 1933): Rudman 1982, 193.
Berghia japonica (Baba, 1933): Gosliner 1985, 261.
Type locality. Tomioka Bay, Japan.
Type material. To our knowledge no type material remains. We designate the specimen CASIZ 1814520 from
Ose, Japan, as the neotype in order to avoid confusion with similar species.
Material examined. Neotype: CASIZ 1814520, one specimen, dissected, 2 mm in length preserved,
immature, Japan, Ose, collected by Akira Kawahara, 10 February 2010. Other material: CASIZ 186795, one
specimen, dissected, 8 mm in length preserved, Marshall Is., collected by Scott Johnson, 12 September 2011;
CASIZ 181357, one specimen, dissected, 5 mm in length preserved, Marshall Is., collected by Scott Johnson, 27
December 2008.
Geographical distribution. Originally described from Tomioka Bay, Japan (Baba 1933) it is also known from
Shimoda (Nakano 2004), the Ryukyu Is. (Ono 2004), Indonesia (Gosliner et al. 2008), the Marshall Is. (present
study), Philippines (Gosliner et al. 2008), and Papua New Guinea (Gosliner et al. 2008).
External morphology (Figs. 2D–E, 3C): Body short, broad, tapering close to posterior end of foot. Foot
corners tentaculiform. Body colour brown or translucent white. Bright white spots over dorsum, forming
continuous patch from pericardial area to posterior end of foot. Tiny white spots form white mark over head.
Rhinophores, oral tentacles and foot corners translucent white. Rhinophores shorter than oral tentacles.
Rhinophores full of elongate, white papillae (Fig. 3C); apex white. Oral tentacles with white spots. Cerata
moderately short, flattened, almost leaf-like. Cerata usually brownish with white spots, or translucent covered by
white pigment; density of white spots varies; some specimens with bluish band on anterior side; a minute white
ring on apex. Cerata in two arches followed by rows, leaving a distinct gap between pre and post-pericardial
groups. Each group with 4–11 cerata, decreasing in size towards foot. Anus cleioproctic, within right second arch.
Gonopore among cerata of anteriormost arch on right.
Anatomy. Masticatory borders smooth (Fig. 4E). Radular formula 33 x 0.1.0 (CASIZ 181357, 5 mm). Radular
teeth slightly bilobed with 27–36 elongate, acutely pointed denticles on either side of the inconspicuous central
notch (Fig. 4F). Teeth progressively smaller in posterior region of radula. Oral glands small, fragile, spongy. Oral
glands dorso-laterally to buccal bulb. Salivary glands long.
Reproductive system diaulic (Fig. 6C). Preampullary duct widening into moderately short ampulla. Postampullary
duct dividing into oviduct and vas deferens. Vas deferens entering into wider proximal portion of penial sac with
unarmed penial papilla. Receptaculum seminis huge, rounded, short stalk connecting to long oviduct, before latter
forms female glands. Vagina ventral to penis.
Remarks. When Baba (1933a) described Baeolidia japonica, he did not mention anything about colouration.
It was sixteen years later (Baba 1949) when the first colour figure of B. japonica was published. In that
contribution, Baba depicted this species as “ground-colour dark brown, the whole back sprinkled with opaque
white spots”. This description clearly does not match with our specimens, which are mainly white. Nevertheless, in
the same area where Baba described B. japonica, both colour types (brownish and whitish) cohabit (Rudman 2002;
Nakano 2004; Ono 2004). Since one of our specimens is from the type locality of B. japonica, we conclude that
both colour types are conspecific.
Excluding Baeolidia scottjohnsoni sp. nov., the main internal differences between B. japonica and the
remaining species of the genus is the shape and the size of the receptaculum seminis. This structure is bigger and
more rounded in B. japonica than in the remaining Baeolidia species. Due to this particular receptaculum seminis,
it cannot be confounded with the remaining species of the genus. Its distinction from B. scottjohnsoni sp. nov. will
be discussed in the remarks section for that species.
Carmona et al. (2013) studied this species from a molecular approach (Fig. 1).
Baeolidia ransoni (Pruvot-Fol, 1956)
(Figs. 2F, 6D, 7A–B)
Aeolidiopsis ransoni Pruvot-Fol, 1956: 228, Fig. 1.
Type locality. Kaukura Atoll, French Polynesia.
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FIGURE 7. Scanning electron photographs. (A–B) Baeolidia ransoni (CASIZ 065417); (A) Detailed view of the masticatory
border; (B) Radular teeth; (C–D) Baeolidia salaamica (CASIZ 184524); (C) Detailed view of the masticatory border; (D)
Radular teeth. Scale bars: A, 50 µm; B, 100 µm; C, 50 µm; D, 150 µm.
Type material. To our knowledge no type material exists. We designate the specimen CASIZ 065417 from
North of Madang, Papua New Guinea, as the neotype in order to avoid confusion with similar species.
Material examined. Neotype: CASIZ 065417, one specimen, dissected, 3 mm in length preserved, Papua
New Guinea, North coast, North of Madang, collected by Richard C. Willan, 31 January 1988. Other material:
CASIZ 065300, one specimen, dissected, 10 mm in length preserved, Papua New Guinea, North coast, North of
Madang, collected by Richard C. Willan, 31 January 1988; CASIZ 186209, one specimen, dissected, 7 mm in
length preserved, Philippines, Luzon, Batangas Province, Calumpan Peninsula, Balayan Bay, collected by Terrence
M. Gosliner, 28 April 2011; CASIZ 186208, one specimen, Philippines, Luzon, Batangas Province, Calumpan
Peninsula, Balayan Bay, collected by Terrence M. Gosliner, 28 April 2011.
Geographical distribution. First described for Kaukura Atoll, French Polynesia (Pruvot-Fol 1956), it is also
known from the Indo-Pacific (Australia, Papua New Guinea and the Philippines) (Gosliner et al. 2008), Japan
(Nakano 2004; Ono 2004) and Hawaii (Pittman & Fiene 2012a).
External morphology (Fig. 2F): Body broad, large, tapering close to posterior end of foot. Foot corners
rounded. Body colour translucent with small light ochre and medium bright white spots all over. Density of spots
varies. Rhinophores smooth, translucent with ochre pigment. Apex white. Oral tentacles short, slender, tapering
near apices. Oral tentacles translucent with small light ochre and medium bright white spots. Cerata flattened,
almost leaf-like. Cerata large, closely oppressed to the body surface rather than erect. Cerata translucent with small
light ochre and medium bright white spots. Tips translucent. Cerata up to twenty-three rows. Each row contains
1–4 cerata, decreasing in size towards the foot. Anus acleioproctic, dorsally to notal brim. Gonopore located among
cerata of anteriormost group on right.
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Anatomy. Masticatory edge smooth (Fig. 7A). Radular formulae 11 x 0.1.0 (CASIZ 186209, 7 mm) and 13 x
0.1.0 (CASIZ 065417, 3 mm). Radular teeth pectinate, short with 18–46 elongate, acutely pointed denticles from
side to side, without a notch or central cusp (Fig. 7B). Oral glands absent. Salivary glands present.
Reproductive system diaulic (Fig. 6D). Preampullary duct widening into wide ampulla. Postampullary duct
dividing into oviduct and vas deferens. Vas deferens elongate, moderately thin, penetrating into wider proximal
portion of penial sac, with unarmed penial papilla. Receptaculum seminis rounded, short stalk connecting to short
oviduct, before latter forms female glands. Vagina ventral to penis.
Remarks. The other two Palythoa feeders, Baeolidia harrietae (Rudman, 1982) and B. palythoae Gosliner,
1985, were transferred and ascribed to Baeolidia respectively by Gosliner (1985). However, Aeolidiopsis ransoni
was retained as the sole species of Aeolidiopsis because of its acleioproctic anus dorsal to the notal brim (Pruvot-
Fol 1956; Gosliner 1985; Miller 2001). Carmona et al. (2013) showed that, from a molecular point of view, the
position of the anus and the rhinophoral ornamentation are not significant phylogenetically in Baeolidia. Hence,
Aeolidiopsis ransoni was transferred to Baeolidia.
Only B. harrietae and B. palythoae have some resemblance with B. ransoni in their colouration, the reticulate
pattern of the digestive gland and the cerata lying out the body. However, the smooth rhinophores of the latter
species distinguish B. ransoni from B. harrietae and B. palythoae. The teeth of B. ransoni are a diagnostic
character for this species (Fig. 7B). Teeth of B. ransoni lack a central cusp or central notch, are quite arched,
pectinate, and have elongate and acutely pointed denticles from side to side. So far, this radular morphology has not
been found in any other Baeolidia species.
This species was included in the molecular study by Carmona et al. (2013) (Fig. 1).
Baeolidia dela (Er. Marcus & Ev. Marcus, 1960)
Berghia dela Er. Marcus & Ev. Marcus, 1960: 924, Figs. 83–86.
Type locality. Addu-Atoll, Maldives Islands.
Type material. To our knowledge no type material exists.
Geographical distribution. So far, this species is only known from the Maldives Islands (Er. Marcus & Ev.
Marcus 1960).
External and internal morphology. Described and figured by Er. Marcus & Ev. Marcus (1960). No
specimens available for the present study.
Remarks. Ernest & Eveline Marcus (1960) described Berghia dela, based on a single specimen. Since this
specimen was already preserved, Er. Marcus & Ev. Marcus (1960) could not provide any information about the
colouration of the living animal. However, they described and drew the papillate rhinophores, the jaws, the radular
tooth and the penis (Er. Marcus & Ev. Marcus 1960, figs. 84–86). The cerata were depicted as cylindrical and were
arranged in arches plus rows. Recently, Carmona et al. (2014b) stated the main characteristics of the aeolidiid
genus Berghia. One of the most important is the papillate morphology of the rhinophores, which can be easily
distinguished from the rhinophorial ornamentation of Baeolidia (see general Discussion). Additionally, based on
these authors’ investigations, Berghia is restricted to the Atlantic and Mediterranean. Therefore, according to these
statements, Berghia dela should be transferred to Baeolidia. Although Er. Marcus & Ev. Marcus (1960) did not
give any information about colouration, which is usually critical to recognize and separate species within
Aeolidiidae (personal observation), the cylindrical cerata could be the distinctive feature that may help to identify
this species if it is ever reported again.
Baeolidia cryoporos Bouchet, 1977
Baeolidia cryoporos Bouchet, 1977: 60, Figs. 26–27.
Type locality. Atlantic Ocean (47º 33 N, 08º 34 W).
Type material. According to Bouchet (1977), the type material was deposited in the Muséum National
d’Historie Naturelle, Paris.
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Geographical distribution. Deep waters, 2110 m, of the Atlantic Ocean (47º 33 N, 08º 34 W) (Bouchet 1977).
External and internal morphology. Described in detail and figured by Bouchet (1977). No specimens
available for the present study.
Remarks. Since Bouchet’s description (1977), this species has not been found again. Therefore, the only
available information stems from the original description. This deep waters aeolidiid has fusiform cerata in rows
and papillate rhinophores. Based on these morphological characters Bouchet (1977) placed this particular member
of Aeolidiidae in Baeolidia. This author also described the reproductive system, which has an extremely large vas
deferens (see figure 27 of Bouchet’s contribution), and figured the jaws and one radular tooth. More information
about the radula and the oral glands, sometimes absent, would be essential to confirm the allocation of Baeolidia
cryoporos to this genus. Additionally, this species is the only Baeolidia species from the Atlantic and is the only
deep-water aeolidiid together with Aeolidia herculea Bergh, 1894. The occurrence of this genus in the Atlantic
Ocean would need further comparatives studies.
Baeolidia harrietae (Rudman, 1982)
(Fig. 8A)
Aeolidiopsis harrietae Rudman, 1982: 157, Figs. 3C, 5D–F, 7–9.
Baeolidia harrietae (Rudman, 1982): Gosliner 1985, 260, Table 1.
Type locality. Lizard Island, Australia.
Type material. The holotype is deposited in the Australian Museum, Sydney (C124689).
Geographical distribution. Originally reported for the Great Barrier Reef of Australia (Rudman 1982), it is
also known from Papua New Guinea (Gosliner et al. 2008), the Philippines (Gosliner et al. 2008) and the Ryukyu
Is., Japan (Ono 2004).
External and internal morphology. Figured and described in detail by Rudman (1982). No specimens
available for the present study.
Remarks. When Rudman (1982) first described Aeolidiopsis harrietae, he pointed out the difficulties of
discerning the systematic position of this species within Aeolidiidae. The author finally attributed this species to
Aeolidiopsis based on the diet similarities with Aeolidiopsis ransoni. Gosliner (1985) considered that Aeolidiopsis
harrietae shares some morphological aspects with Baeolidia palythoae, such as the papillate rhinophores, and
therefore transferred A. harrietae to Baeolidia. The molecular phylogeny conducted by Carmona et al. (2013)
rejected the validity of the genus Aeolidiopsis, since Aeolidiopsis ransoni clustered within the Baeolidia clade.
Thus, we agree with Gosliners (1985) statements and confirm its placement within Baeolidia.
Baeolidia ransoni, B. harrietae and B. palythoae are very similar from a morphological point of view.
However, while B. harrietae has papillate rhinophores, B. ransoni has smooth ones. B. palythoae has two pairs of
oral glands, but Rudman (1982) could not find any traces of oral glands in B. harrietae.
Baeolidia australis (Rudman, 1982)
(Fig. 8B)
Spurilla australis Rudman, 1982, 164, Figs. 12–16.
Berghia australis (Rudman, 1982): Gosliner 1985, 261.
Baeolidia australis (Rudman, 1982): Miller 2001, 633.
Type locality. Long Reef, Australia.
Type material. According to Rudman (1982), the holotype is deposited the Australian Museum, Sydney
(C92613).
Geographical distribution. Only reported from New Zealand and Australia (Morton & Miller 1968 as
Baeolidia major; Powell 1976, 1979; Willan & Morton 1984).
External and internal morphology. Described in detail and figured by Rudman (1982). No specimens
available for the present study.
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FIGURE 8. Photographs of the living animals. (A) Baeolidia harrietae, specimen from Papua New Guinea, photo by Terrence
M. Gosliner; (B) Baeolidia australis, specimen from Australia, photo by Lean and David Atkinson; (C) Baeolidia salaamica,
specimen from the Philippines, photo by Terrence M. Gosliner, CASIZ 177599; (D) Baeolidia salaamica, specimen from the
Philippines, photo by Terrence M. Gosliner, CASIZ 177397; (E) Baeolidia chaka, specimen from South Africa (6 mm), photo
by Terrence M. Gosliner; (F) Baeolidia palythoae, specimen from South Africa (8 mm), photo by Terrence M. Gosliner.
Remarks. We transfer Spurilla australis to Baeolidia based on the fact that its rhinophoral papillae (small
knobs) are one of the types of Baeolidias (Fig. 3) and because its morphological similarities with the type species
of this genus (Baeolidia moebii). Regarding the characteristic of the radula, the teeth of B. moebii seem to be wider
and have shorter denticles than those of B. australis. However, due to the intraspecific variability of this structure
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in Aeolidiidae (personal observation), further material of this species would be essential to test the validity of this
character. In terms of colouration, B. australis has a thin orange sub-apical band above a broad blue band on each
cerata, which it is never present in B. moebii. B. australis looks bluish whereas B. moebii is more yellowish or
brownish. The body of B. australis also seems to have a brown and white reticulate pattern that does not appear to
be present in B. moebii. Recently Rudman (2007b) questioned the validity of B. australis, but until this species
could be examined from a morphological and/or molecular point of view we retain B. australis as a distinct species.
Baeolidia salaamica (Rudman, 1982)
(Figs. 3D, 7C–D, 8C–D, 9A)
Spurilla salaamica Rudman, 1982: 173, Figs. 21–23.
Berghia salaamica (Rudman, 1982): Gosliner 1985, 261.
Type locality. Dar es Salaam, Tanzania.
Type material. According to Rudman (1982), the material was deposited in the Australian Museum, Sydney
(C124695).
Material examined. CASIZ 184524, one specimen, dissected, 6 mm in length preserved, Japan, Manadaru,
collected by Rie Nakano, 19 April 2006; CASIZ 177397, one specimen, dissected, 3 mm in length preserved,
Philippines, Luzon Island, Batangas Province, Balayan Bay, Anilao, Matotonggil, collected by Terrence M.
Gosliner, 18 March 2008; CASIZ 177599, one specimen, 2 mm in length preserved, Philippines, Luzon, Batangas
Province, Calumpan Peninsula, collected by Terrence M. Gosliner, 16 April 2008.
Geographical distribution. Originally described from Dar es Salaam (Tanzania) (Rudman 1982), this species
is also known from Papua New Guinea (Gosliner et al. 2008), the Philippines (Gosliner et al. 2008), Japan (Ono
1999, 2004; Nakano 2004), Korea (Koh 2006) and Hawaii (Pittman & Fiene 2012b).
External morphology (Fig. 3D, 8C–D): Body short, broad, tapering close towards posterior end of foot. Foot
corners tentaculiform. Body colour translucent with white marks scattered over dorsum. White ring, which may
have opaque white pigmentation in centre, on head. White diamond-shape mark behind rhinophores. Rhinophores,
oral tentacles and foot corners translucent with white marks. Rhinophores shorter than oral tentacles. Rhinophores
studded of minute white knobs (Fig. 3D). Apex white. Oral tentacles with white tips.
Cerata moderately long, flattened, almost leaf-like. Branches of digestive gland shining through translucent
body wall. Cerata may have white pigmentation (as speckles or covering outer side of cerata) on them. Apex
translucent white. Cerata forming two arches plus three rows. Each arch or row contains 2–11 cerata, decreasing in
size towards foot. Anus cleioproctic located within second right arch. Genital opening among cerata of
anteriormost group on right side.
Anatomy. Masticatory border smooth (Fig. 7C). Radular formulae 13 x 0.1.0 (CASIZ 177397, 3mm) and 19 x
0.1.0 (CASIZ 184524, 6 mm). Radular teeth slightly bi-arched with up to 25 moderately broad, acutely pointed
denticles on either side of minute central cusp (Fig. 7D). Teeth progressively smaller towards posterior region of
radula. Oral glands small, relatively elongate, spongy. Oral glands dorso-laterally to buccal bulb. Salivary glands
absent.
Reproductive system diaulic (Fig. 9A). Preampullary duct widening into ampulla Postampullary duct dividing
into oviduct and vas deferens. Vas deferens elongate, entering into wider proximal portion of penial sac with
unarmed penial papilla. Receptaculum seminis bean-shaped, connecting to oviduct, before latter forms female
glands. Vagina ventral to penis.
Remarks. Since Rudman (1982) did not consider Berghia and Baeolidia as valid genera he described this
species as Spurilla. Gosliner (1985) transferred Spurilla salaamica to Berghia based on its papillate rhinophores
and cerata in arches. The molecular phylogeny conducted by Carmona et al. (2013) showed that Spurilla/Berghia
salaamica clustered within the Baeolidia clade, breaking the monophyly of Baeolidia. Thus, this species was
transferred to Baeolidia, as the most parsimonious alternative. Baeolidia salaamica also shares some
morphological features with other members of this genus such as the knob-like papillae of the rhinophores (similar
to those found in B. moebii), the leaf-like cerata and the cerata arrangement in arches and rows (also in e.g. B.
japonica). Therefore, we can confirm that B. salaamica belongs to this genus.
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FIGURE 9. Reproductive system. (A) Baeolidia salaamica, specimen from Japan (CASIZ 184525); (B) Baeolidia rieae sp.
nov., specimen from Japan (CASIZ 184525); (C) Baeolidia variabilis sp. nov., specimen from the Philippines (CASIZ
177715); (D) Baeolidia lunaris sp. nov., specimen from Tanzania (CASIZ 099221). Scale bars: A–D, 0.5 mm. Abbreviations:
am, ampulla; fgm, female gland mass; ps, penial sac; rs, receptaculum seminis; va, vagina; vd, vas deferens.
Concerning the original description, only one of our specimens from the Philippines (Fig. 8D) matches
completely with Rudman’s (1982) diagnosis. Figure 8C shows that the other specimen from the Philippines has
somewhat large white patches over the outer side of the cerata. Moreover, none of the specimens here examined
bear a denticulate masticatory border (Fig. 7C) as Rudman (1982) stated in the original description. Based on our
results in previous studies (Carmona et al. 2014a, b, c) we conclude that the latter morphological feature is not
significant in Aeolidiidae.
The translucent body, the white ring on the head, the white diamond-shape mark just behind the rhinophores,
and mainly the kind of rhinophoral papillae, which is exclusive of Baeolidia but being only present in B. moebii
and B. australis, allow us to distinguish B. salaamica from other species of this genus. Concerning the anatomical
characters, it seems that B. salaamica has the wider radular teeth denticles of Baeolidia (Fig. 7D). Carmona et al.
(2013) studied this species from a molecular approach (Fig. 1).
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Baeolidia chaka (Gosliner, 1985)
(Fig. 8E)
Berghia chaka Gosliner, 1985, 245, Figs. 11–14.
Type locality. Jesser Point, Sodwana Bay National Park, South Africa.
Type material. The type material was deposited in the South African Museum, Cape Town (Gosliner 1985).
The registration number of the holotype of Baeolidia chaka is SAM-A35634.
Geographical distribution. So far, only known from South Africa (Gosliner 1985).
External and internal morphology. Described in detail and figured by Gosliner (1985). No specimens
available for the present study.
Remarks. Berghia chaka was described originally from South Africa (Gosliner 1985). The original
description was based on two specimens, both deposited in the South African Museum, Cape Town (SAM). The
only available information stems from the original description since this species has not been found again. The
body and teeth shape, the flattened cerata and the oral glands described and figured by Gosliner (1985), lead us to
transfer Berghia chaka to Baeolidia.
Gosliner (1985) considered the anteriorly rounded foot as the diagnostic character of B. chaka. This species
also has some cerata noticeably longer than the rest, which is not very common in Baeolidia. In external
appearance, B. chaka most closely resembles B. japonica. However, the latter species possesses tentacular foot
corners, a bluish band on the anterior side and cerata uniform in size.
Baeolidia palythoae Gosliner, 1985
(Fig. 8F)
Baeolidia palythoae Gosliner, 1985, 237, Figs. 4–10.
Type locality. Umgazana, South Africa.
Type material. The holotype is deposited in the South African Museum, Cape Town (SAM-A35640).
Geographical distribution. To date, only known from South Africa (Gosliner 1985).
External and internal morphology. Described in detail and figured by Gosliner (1985). No specimens
available for the present study.
Remarks. Baeolidia palythoae was originally described from South Africa by Gosliner (1985). So far, the
only available information about this species is the original description. Gosliner (1985) provided a complete
description of this species and figured the rhinophores, the ventral and lateral view of the species, the cerata, the
oral glands, the jaws, the central nervous and the reproductive system. This author also presented scanning electron
micrographs of the jaws and their masticatory border as well as the radular teeth. The most distinctive features of
this species are the second pair of oral glands, which are ovoid in shape and empty on the ventral side of the head,
as well as the small tubercles on the masticatory border. B. palythoae closely resembles B. harrietae in external
appearance but both species differ in several features such as the number of cerata per row (4–9 in B. harrietae and
3–5 in B. paylothoae), the position of the gonopore, the number of oral glands and the ornamentation of the
masticatory border (Gosliner 1985).
Baeolidia rieae sp. nov.
LSID urn:lsid:zoobank.org:act:35F88299-66D8-4726-8796-410D907D86F8
(Figs. 9B, 10A, 11A–B).
Baeolidia sp. B: Carmona et al. 2013, 6.
Material examined. Holotype: CASIZ 184525, one specimen, dissected, 3 mm in length preserved, Japan,
Amami-Ohoshima Island, collected by Rie Nakano, 06 March 2010.
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FIGURE 10. Photographs of the living animals. (A) Baeolidia rieae sp. nov., specimen from Japan, photo by Jum Imamoto,
CASIZ 184525; (B–C) Baeolidia variabilis sp. nov.: (B) specimen from Marshall Islands, photo by Scott Johnson, CASIZ
187741; (C) specimen from the Philippines, photo by Terrence M. Gosliner, CASIZ 177715.
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FIGURE 11. Scanning electron photographs. A, B, Baeolidia rieae sp. nov. (CASIZ 184525); (A) Detailed view of the jaw;
(B) Radular teeth; (C–D) Baeolidia variabilis sp. nov. (CASIZ 177715); (C) Detailed view of the masticatory border; (D)
Radular teeth; (E-F) Baeolidia lunaris sp. nov. (CASIZ 099221); (E) Detailed view of the masticatory border; (F) Radular
teeth. Scale bars: A, 30 µm; B, 100 µm; C, 2 µm; D, 10 µm; E, 10 µm; F, 20 µm.
Type locality and habitat. The Amami-Ohoshima Island, Japan. Found in 7 m of water close to green
zoanthids.
Geographical distribution. So far, only known from Japan (present study).
Etymology. This species is named after Rie Nakano who kindly provided us material from Japan, including
this new species.
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External morphology (Fig. 10A): Body short, broad, tapering close towards posterior end of foot. Foot
corners rounded. Body colour translucent. Dark ochre and pearly white pigmentation over notum. Rhinophores
same colour as ground body colour with inconspicuous papillae. Apex pearly white. Cerata large, flattened, almost
leaf-like. Cerata light ochre with pearly white, dark ochre and iridescent green pigmentation. Cnidosacs white.
Cerata in one arch followed by three rows, leaving a distinct gap between pre and post-pericardial groups. Each
arch with 2–5 cerata, decreasing in size towards the foot. Anus cleioproctic, behind first right row. Gonopore
behind the anteriormost row on right.
Anatomy. Ornamentation of masticatory border could not be determined (Fig. 11A). Radular formula 20 x
0.1.0 (CASIZ 184525, 3 mm). Radular teeth bilobed with 31–39 elongate, acutely pointed denticles on either side
of minute central cusp (Fig. 11B). Teeth progressively smaller towards posterior region of radula. Oral glands
small, spongy, dorso-laterally to buccal bulb. Salivary glands absent.
Reproductive system diaulic (Fig. 9B). Preampullary duct widening into moderately long ampulla. Postampullary
duct dividing into oviduct and vas deferens. Vas deferens short, penetrating into wider proximal portion of the
penial sac with unarmed penial papilla. Receptaculum seminis bean-shaped, short stalk connecting to short oviduct,
before latter forms female glands. Vagina ventral to penis.
Remarks. Baeolidia rieae sp. nov. is clearly distinguishable from other members of the genus by its
conspicuous iridescent pigmentation. Only Baeolidia lunaris sp. nov. and Baeolidia gracilis sp. nov. (see below)
have this kind of pigment. Nevertheless, in B. lunaris sp. nov., the cerata have a completely different colouration
and a very characteristic white spot on their bases. In addition, the body colour of B. gracilis sp. nov. is much
darker than in B. rieae sp. nov.. B. gracilis also has larger cerata with an opaque white line on their inner margin
(see below), which is absent in B. rieae sp. nov.. Anatomically, these three species are similar but some differences
could be found. While the radular teeth of B. lunaris sp. nov. are pectinate, B. rieae sp. nov. and B. gracilis sp. nov.
have bilobed teeth. In addition, the teeth of B. rieae sp. nov. show a minute central cusp, but B. gracilis sp. nov.
only has a central notch. Baeolidia rieae sp. nov. and B. gracilis sp. nov. also differ in number of oral glands.
Regarding the receptaculum seminis, there are differences in shape. While the receptaculum seminis of B. rieae sp.
nov. is bean-like, in B. lunaris sp. nov.
and B. gracilis sp. nov. it is rounded and ovoid respectively. Moreover, the
length of the vas deferens also varies among these species, having an intermediate length in B. rieae sp. nov..
This new species was included in the molecular study by Carmona et al. (2013) (Fig. 1).
Baeolidia variabilis sp. nov.
LSID urn:lsid:zoobank.org:act:B6B01F57-6AD8-49AD-B95D-B7F7EE91C9DC
(Figs. 3C, 9C, 10B–C, 11C–D)
Berghia sp. 1: Gosliner et al. 2008, 404.
Berghia sp. 4: Gosliner et al. 2008, 405.
Baeolidia sp. A: Carmona et al. 2013, 6.
Material examined. Holotype: CASIZ 103739, one specimen, dissected, 4 mm in length preserved, Philippines,
Mindoro, Medio Island, collected by Terrence M. Gosliner, 28 February 1995; Paratype: CASIZ 190618, one
specimen, dissected, 3 mm in length preserved, Papua New Guinea, North coast collected by Terrence M. Gosliner,
29 January 1988. Other material: CASIZ 177715, one specimen, dissected, 4 mm in length preserved, Philippines,
Luzon, Batangas Province, Calumpan Peninsula, collected by Terrence M. Gosliner, 21 April 2008; CASIZ
177716, one specimen, dissected, 3 mm in length preserved, Philippines, Luzon, Batangas Province, Calumpan
Peninsula, collected by Terrence M. Gosliner, 21 April 2008; CASIZ 186210, one specimen, dissected, 2 mm in
length preserved, Philippines, Luzon, Batangas Province, Maricaban Island, collected by Terrence M. Gosliner, 21
May 2011; CASIZ 187741, two specimens, dissected, 11 and 10 mm in length alive, Marshall Island, Kwajalein
Atoll, collected by Scott Johnson, 24 July 2011.
Type locality and habitat. Medio Island, Mindoro, Philippines. Found in shallow water reefs under coral
rubble, where it likely feeds on small sea anemones.
Geographical distribution. So far, only known from the Philippines, the Marshall Islands and Papua New
Guinea (present study).
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Etymology. In Latin the word “variabilis” means variable. The specific name refers to the two colour types of
this species.
External morphology (Figs. 3C, 10B–C): Body short, broad, tapering close to posterior end of foot. Foot
corners, short and rounded. Body colour translucent with two different colour patterns. First one with opaque white
patches on head (its density varies) and notum (Fig. 10B). Light ochre reticulate on both sides of body. Second one
with small light ochre spots all over body (Fig. 10C). Rhinophores approximately equal in length to oral tentacles.
Rhinophores densely covered by elongate papillae (Fig. 3C). Rhinophores translucent with ochre pigment and
white apex. Oral tentacles short, slender, tapering near apices. Oral tentacles same colour as ground colour and
white tips.
Cerata flattened, almost leaf-like. In first colour pattern, cerata ochre with white pigmentation on posterior side
of basal portion. Some cerata noticeably longer than others. In second colour pattern, cerata recurved inwardly,
with some papillae or bulbs. Cerata completely white, usually having bright yellow pigment on posterior side of
their distal area. Cerata in two or three arches followed by a couple of rows. Cerata groups leaving a distinct gap
between pre and post-pericardial groups. Each group with 1–5 cerata, decreasing in size towards foot. Anus
cleioproctic, below second right arch. Genital aperture among cerata of anteriormost group on right.
Anatomy. Masticatory process smooth (Fig. 11C). Radular formulae 8 x 0.1.0 (CASIZ 103739, 4 mm), 14 x
0.1.0 (CASIZ 177715, 4mm). Radular teeth bilobed with 47–52 elongate and acutely pointed denticles on either
side of minute central cusp (Fig. 11D). Teeth progressively smaller towards posterior region of radula. Oral glands
small, moderately elongate, fragile, spongy, laying dorso-laterally to buccal bulb. Ventral gland present. Salivary
glands absent.
Reproductive system diaulic (Fig. 9C). Preampullary duct widening into conspicuous ampulla. Postampullary
duct dividing into oviduct and vas deferens. Vas deferens short, entering into wider proximal portion of penial sac
with unarmed penial papilla. Receptaculum seminis bean-shaped receptaculum seminis, short stalk connecting to
long oviduct, before latter forms female glands. Vagina ventral to penis.
Remarks. This particular species highlights how variable the colouration can be within the same species.
Gosliner et al. (2008) identified both colour patterns as separate species but the molecular analyses undertaken by
Carmona et al. (2013) showed that they are conspecific. Further studies would help to elucidate if these colour
patterns depend on habitat, prey and/or an ontogenetic process.
Both colour patterns differ consistently from the remaining Baeolidia species but B. japonica, B. salaamica, B.
gracilis sp. nov. and B. scottjohnsoni sp. nov. (see below) are somewhat similar in external appearance. However,
B. salaamica is completely translucent, including the cerata, while the body colour of B. variabilis sp. nov. is
opaque white or covered by small ochre flecks. B. scottjohnsoni sp. nov. and B. japonica have a characteristic blue
band at the anterior side of the cerata, which is not found in B. variabilis sp. nov.. Internally, differences in the size
of the receptaculum seminis between B. variabilis sp. nov. and B. scottjohsoni sp. nov. are also obvious (Fig. 8C
and 14B respectively). None of these species has a ventral oral gland except B. gracilis sp. nov. (see below) but B.
variabilis sp. nov.
lacks any traces of the iridescent pigment found on B. gracilis sp. nov..
Carmona et al. (2013) included this new species in their molecular phylogeny (Fig. 1).
Baeolidia lunaris sp. nov.
LSID urn:lsid:zoobank.org:act:249BBCD0-ED70-4513-8954-BB8566ECFC0E
(Figs. 3C, 9D, 11 E–F, 12A)
Baeolidia sp. 1: Gosliner et al. 2008, 405.
Material examined. Holotype: CASIZ 099221, one specimen, dissected, 10 mm in length alive, Tanzania, Mtwara
Region, Mana Huanja Island, collected by Terrence M. Gosliner, 01 November 1994.
Type locality and habitat. Mtwara Region, Mana Huanja Island, Tanzania. Found in shallow water feeding on
zoanthids.
Geographical distribution. So far, only known from Tanzania (present study).
Etymology. The specific name refers to the white patch present at the base of the cerata, similar to the moon
(from latin: lunaris, pertaining to the moon).
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FIGURE 12. Photographs of the living animals. (A) Baeolidia lunaris sp. nov., specimen from Tanzania, photo by Terrence M.
Gosliner, CASIZ 099221; (B) Baeolidia gracilis sp. nov., specimen from the Philippines, photo by Terrence M. Gosliner,
CASIZ 083766; (D) Baeolidia scottjohnsoni sp. nov., specimen from Marshall Islands, photo by Scott Johnson, CASIZ
184503.
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External morphology (Figs. 3C, 12A): Body short, broad, tapering close to posterior end of foot. Foot corners
rounded. Body colour translucent brownish. Fine iridescent opaque white spots over body. Rhinophores
approximately equal in length to oral tentacles. Rhinophores densely covered by elongate papillae (Fig. 3C).
Rhinophores translucent brownish with small white spots. Apex white. Oral tentacles translucent brownish short,
slender, tapering near apices. Cerata moderately long, flattened, leaf-like. Cerata same colour as ground body
colour, but a little bit darker. White spot on anterior side of cerata, close to base. Cerata recurved inwardly. Apex
white. Cerata in one arch and four rows, leaving a distinct gap between pre and post-pericardial groups. Each group
with 2–5 cerata, decreasing in size towards foot. Anus cleioproctic, below first right row. Gonopore housed among
cerata of anteriormost right group.
Anatomy. Masticatory process smooth (Fig. 11E). Radular formula 21 x 0.1.0 (CASIZ 099221, 10 mm).
Radular teeth blended with 52–71 elongate, acutely pointed denticles (Fig. 11F). Central cusp absent. Teeth
progressively smaller towards posterior radular region. Oral glands small, ovoid, delicate. Salivary glands long.
Reproductive system diaulic (Fig. 9D). Preampullary duct widening into short ampulla. Postampullary duct
dividing into oviduct and vas deferens. Vas deferens entering into wider proximal portion of penial sac with
unarmed penial papilla. Receptaculum seminis rounded, short stalk connecting to short oviduct, before latter forms
female glands. Vagina ventral to penis.
Remarks. Three Baeolidia species are known from Tanzania. Only B. moebii (Eliot, 1903; Edmunds, 1970),
B. salaamica (Rudman, 1982) and B. lunaris sp. nov. (present study) have been reported in that country. In terms of
colouration, B. lunaris sp. nov. differs from B. moebii and B. salaamica in several features. B. lunaris sp. nov.
lacks the whitish or yellow ring usually present in B. moebii. The latter species also has a bright yellow subapical
band, which does not appear in B. lunaris sp. nov.. B. salaamica lacks any traces of iridescent pigmentation, being
mainly translucent white or brown. Moreover, both species, B. salaamica and B. lunaris sp. nov. differ in the shape
of the radular teeth (see Figs. 7C–D and 11E–F respectively). The radular teeth of B. salaamica are bilobed while
B. lunaris sp. nov. has pectinate teeth. However, teeth of B. moebii and B. lunaris sp. nov. are very similar (Figs.
4A–B and 11E–F respectively). In terms of reproductive system, B. lunaris sp. nov. is the only of these three
species with rounded receptaculum seminis. The main difference among these species from Tanzania is the
morphology of the papillae of their rhinophores. While B. moebii and B. salaamica have rounded and short papilla,
B. lunaris’ sp. nov. are clearly longer.
Regarding the remaining members of this genus, B. lunaris sp. nov. is similar to B. rieae sp. nov. and B.
gracilis
sp. nov. (see below) in that the three species have iridescent pigment. Neither B. rieae sp. nov. nor B.
gracilis sp. nov. have a white spot at the base of the anterior side of the cerata. Additionally, the shape of the
radular teeth of B. gracilis sp. nov. and B. rieae sp. nov. is different of B. lunaris’ sp. nov.. The former species
have a radular teeth more bilobed and less bended than the radular teeth of B. lunaris sp. nov. (see Figs. 11A–B,
11E–F and 13A–B respectively). Because of the lack of material properly conserved for a molecular study, we
could not test the validity of this species from a molecular perspective. However, we consider that the
morphological and colour characteristics presented here support the validity of Baeolidia lunaris sp. nov..
Baeolidia gracilis sp. nov.
LSID urn:lsid:zoobank.org:act:A50C22AD-2E31-4D77-841F-7DEF78087E20
(Figs. 12B, 13A–B, 14A)
Berghia sp. 2: Gosliner et al. 2008, 404.
Material examined. Holotype: CASIZ 083766, one specimen, dissected, 9 mm in length preserved, Philippines,
Luzon, Batangas Province, Calumpan Peninsula, Maricaban Strait, collected by Terrence M. Gosliner, 22 February
1992.
Type locality and habitat. Arthur's Rock (South West side of Calumpan Peninsula), Maricaban Strait,
Calumpan Peninsula, Batangas Province, Luzon, Philippines. Found under coral rubble in 10 m of water.
Geographical distribution. Thus far, known only from the Philippines (present study).
Etymology. The specific name refers to the slender body of this species.
External morphology (Figs. 12B): Body moderately elongate, tapering close to translucent posterior end of
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foot. Foot corners rounded. Head and pericardium dark greyish blue. Posterior part of notum and cerata ochre.
Iridescent greyish blue and bright ochre pigment over pericardium. Reticulate brown pattern close to foot edges.
Rhinophores approximately equal in length to oral tentacles. Rhinophores dark greyish blue with only few
moderately short papillae, mainly in posterior part. Oral tentacles short, slender, tapering near apices. Oral tentacles
with same colour as rhinophores.
FIGURE 13. Scanning electron photographs. (A–B) Baeolidia gracilis sp. nov. (CASIZ 083766); (A) Detailed view of the
masticatory border; (B) Radular teeth; (C–D) Baeolidia scottjohnsoni sp. nov. (CASIZ 184503); (C) Detailed view of the
masticatory border; (D) Radular teeth. Scale bars: A, 3 µm; B, 10 µm; C, 50 µm; D, 100 µm.
Cerata long, flattened, pod-bean-like. Cerata with greenish ochre net, converging on translucent apex. Inner
margin with opaque white line. Cerata in three arches followed by five rows, leaving a distinct gap between pre-
pericardial group and post-pericardial groups. Each group with 1–9 cerata, decreasing in size towards foot. Anus
cleioproctic, below second right arch. Gonopore among cerata of anteriormost group on right.
Anatomy. Masticatory process smooth (Fig. 13A). Radular formula 9 x 0.1.0 (CASIZ 083766, 9 mm). Radular
teeth bilobed with 48–63 elongate, acutely pointed denticles on either side of notch (Fig. 13B). Two pairs of oral
glands. Oral glands delicate. First pair elongate, laying dorso-laterally to buccal bulb. Second pair smaller,
rounded, emptying on ventral side of head. Salivary glands absent.
Reproductive system diaulic (Fig. 14A). Preampullary duct widening into short ampulla. Postampullary duct
dividing into oviduct and vas deferens. Vas deferens inserting into wider proximal portion of penial sac, with
unarmed penial papilla. Receptaculum seminis moderately large, ovoid, short stalk connecting to short oviduct,
before latter forms female glands. Vagina ventral to penis.
Remarks. While most of the Baeolidia species have a stout body, Baeolidia gracilis sp. nov. is much more
slender than the others members of this genus (see e.g. Fig. 12). The cerata of this species are also particular. They
are long, flattened and pod-bean-like rather than being leaf-like. This morphological features and its colour pattern
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characterize this species. Regarding colouration, B. gracilis sp. nov. most closely resembles B. rieae sp. nov. and
B. lunaris sp. nov.. However, B. gracilis sp. nov. has a white line on the inner margin of the cerata that is not
present in B. rieae. sp. nov. Moreover, B. rieae sp. nov. lacks the second pair of oral glands of B. gracilis sp. nov..
The latter species does not have the white spot close to the base of cerata of B. lunaris sp. nov.. Both species also
differ in the radular morphology (see Fig. 11E–F for B. lunaris sp. nov. and 13A–B for B. gracilis sp. nov.). In
terms of internal anatomy, in addition to this new species, only B. variabilis sp. nov. and B. palythoae have two
pairs of oral glands. B. variabilis sp. nov. and B. gracilis sp. nov. have consistent differences in colouration, while
B. palythoae has papillae over the masticatory border, which is smooth in B. gracilis sp. nov..
FIGURE 14. Reproductive system. (A) Baeolidia gracilis sp. nov., specimen from the Philippines (CASIZ 083766); (B)
Baeolidia scottjohnsoni sp. nov., specimen from Marshall Islands (CASIZ 184503). Scale bars: A, 0.5 mm; B, 1 mm.
Abbreviations: am, ampulla; fgm, female gland mass; ps, penial sac; rs, receptaculum seminis; v, vagina; vd, vas deferens.
Baeolidia scottjohnsoni sp. nov.
LSID urn:lsid:zoobank.org:act:EDE11CBD-6EE9-43D1-9970-158C1FEB2E1A
(Figs. 3C, 12C, 13C–D, 14C)
Baeolidia sp. C: Carmona et al. 2013, 6.
Material examined. Holotype: CASIZ 184503, one specimen, dissected, 12 mm in length alive, Marshall Islands,
Kwajalein Atoll, Enewetak Pinnacle, collected by Scott Joshnson, 11 October 2010. Paratype: CASIZ 180341, one
specimen, dissected, 12 mm in length alive, Hawaii, Maui, Wahikuli, collected by Pauline Fiene, 30 November
2003.
Type locality and habitat. Enewetak Pinnacle, Kwajalein Atoll, Marshall Islands. Found in 10 m of water
over Caulerpa.
Geographical distribution. So far, only known from the Marshall Islands and the Hawaiian Islands (present
study).
Etymology. This species is dedicated to Scott Johnson who kindly provided us with abundant material from
the Marshall Islands, including specimens of this new species.
External morphology (Figs. 3C, 12C): Body short, broad, tapering close to posterior end of foot. Foot corners
tentaculiform. Body brownish. Bright white spots over head, dorsum and foot. White spots over notum form almost
a continuous patch from pericardial area to posterior end of foot, excluding ceratal insertions. Iridescent blue
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pigmentation over white spots, more evident on those parts of dorsum where groups of cerata from both sides are
closer as well as close to ceratal insertion. Rhinophores, oral tentacles and foot corners brownish with bright white
spots. Rhinophores approximately same length of oral tentacles. Rhinophores with elongated papillae (Fig. 3C).
Distal papillae whiter. Apex white. Oral tentacles with white spots. Cerata moderately short, flattened, leaf-like.
Cerata slightly recurved inwards. Cerata brown with white spots on posterior side. Double subapical band on
anterior side of cerata. Upper half of cerata white; second half iridescent blue that may reach their base. Orange or
light ochre patch may be found on distal part of posterior side of cerata, being overlaid by bright ochre reticulate.
Latter converges on apex. Apex with minute white ring. Cerata in two arches and two rows, leaving a distinct gap
between pre and post-pericardial groups. Each group with 2–4 cerata, decreasing in size towards foot. Cleioproctic
anus within second right arch. Genital aperture placed within anteriormost arch on right.
Anatomy. Masticatory process smooth (Fig. 13C). Radular formula 14 x 0.1.0 (CASIZ 184503, 12 mm).
Radular teeth slightly bilobed with 25 moderately broad and acutely pointed denticles on each side of the small and
elongate central cusp (Fig. 13D). Oral glands small, delicate (spongy), elongate. Salivary glands absent.
Reproductive system diaulic (Fig. 14B). Preampullary duct widening into ampulla. Postampullary duct dividing
into oviduct and vas deferens. Vas deferens moderately long vas deferens inserting into wider proximal portion of
penial sac with unarmed penial papilla. Receptaculum seminis huge, folded, short stalk connecting to short oviduct,
before latter forms female glands. Vagina ventral to penis.
Remarks. Excluding Baeolidia japonica, B. scottjohnsoni sp. nov. is easily distinguishable from other
members of the genus due to its colouration and the shape and size of its receptaculum seminis. However, the
brownish colour pattern of B. japonica has some resemblances with B. scottjohnsoni sp. nov.. Both species have
brownish body colour, with white spots all over the notum and a blue band on the posterior side of the cerata.
Furthermore, both species have a huge receptaculum seminis (Figs. 6C and 14B). Nevertheless, the molecular
phylogenetic analysis conducted by Carmona et al. (2013) (see Fig. 1) clearly illuminated the presence of distinct
cryptic species that are separated by some morphological differences. The blue pigment over the notum of B.
scottjohnsoni sp. nov., differences in the radular morphology (denticles of B. scottjohnsoni sp. nov. are broader
than those of B. japonica), and the folded receptaculum seminis of B. scottjohnsoni sp. nov. distinguish these
species.
Carmona et al. (2013) studied this species from a molecular approach (Fig. 1).
Discussion
Genus Baeolidia
Recently, the monophyly of Baeolidia was recovered in the first molecular phylogenetic study of Aeolidiidae
(Carmona et al. 2013). So far, Baeolidia is the largest genus within Aeolidiidae with sixteen valid species. Some
species traditionally ascribed to Baeolidia such as Baeolidia fusiformis, Baeolidia benteva and Baeolidia nodosa
belong to other aeolidiid genera but not to Baeolidia (Carmona et al. 2013; Carmona et al. 2014b,c). Moreover,
Baeolidia macleayi, Baeolidia dela and Baeolidia chaka are transferred to this genus for the first time. Figure 1
shows the phylogenetic hypothesis for Baeolidia presented by Carmona et al. (2013.) but showing the final names
of the new species on the right side of vertical bars.
Except for the Atlantic species Baeolidia cryoporos, this genus is entirely Indo-Pacific with just a few
representatives in the Eastern Pacific. Only B. moebii, B. ransoni, B. harrietae and B. salaamica present
populations in the Indo-Pacific as well as in the Eastern Pacific.
The inclusion of Baeolidia ransoni and Baeolidia macleayi, with smooth and ribbed rhinophores respectively,
and the existence of different cerata arrangements within this genus require modifying the diagnosis of the genus
Baeolidia. All the traditional morphological characters (e.g. the reproductive system and radula) and new
morphological traits, such as the oral glands and the rhinophoral papillae, have been deeply studied here. At a
generic level, the oral glands, the different types of rhinophoral papillae, the radular morphology and the leaf-
shaped cerata were shown to be informative. However, there are species that lack some of these characters. In fact,
this genus has the greatest diversity of morphological patterns within Aeolidiidae (see revised diagnosis), which
makes it very difficult to determine new synapomorphies at this level based on anatomical data. This situation has
also been demonstrated with other heterobranch groups such as Tamb ja and Roboastra (Pola et al. 2007, 2008) as
well as within Aeolidiidae with Berghia (Carmona et al. 2014b).
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Some morphological traits also have intraspecific variation. The ornamentation of the masticatory border (e.g.
Baeolidia salaamica) and the presence or absence of oral glands (e.g. Baeolidia moebii) may vary within the same
species. Although the intraspecific varibilty of the precence/absence of the oral glands has not been reported in the
literature, this variation has also been observed in Limenandra nodosa (Carmona et al., 2014c). Thus, in order to
separate species, the colouration of the living animal can be considered as the main character in Baeolidia together
with the ornamentation of the rhinophores (including the different types of papillae) and, in some cases, the radular
morphology (Table 1).
FIGURE 15. Schematic representation of differences of body shape among the genera Spurilla (A), drawing of Ascensión
Cespedosa; Berghia (B), drawing of Ascensión Cespedosa; Limenandra (C), taken from Haefelfinger & Stamm (1958); and
Baeolidia (D), taken from Rudman (1982).
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FIGURE 16. Comparison of the radular morphology among the genera Spurilla (A), Berghia (B), Limenandra (C) and
Baeolidia (D).
Baeolidia vs Spurilla, Berghia and Limenandra
The aeolidid genera Spurilla, Berghia and Baeolidia were considered to be closely related. Indeed, Odhner
(1939) placed Spurilla and Berghia in a new family called Spurillidae, including Baeolidia ten years later and
overlooking Limenandra (Odhner in Franc 1968). In the manner of Odhner (in Franc 1968), many authors regarded
Limenandra as a junior synonym of Baeolidia. Recently, the molecular phylogeny of Aeolidiidae (Carmona et al.
2013) rejected the validity of Spurillidae sensu Odhner and the synonymy of Limenandra with Baeolidia.
However, the morphological differences among these genera have not been well defined.
Despite difficulties in finding synapomorphies for some genera within Aeolidiidae, it is possible to
differentiate Baeolidia, Spurilla, Berghia and Limenandra based on some features. Regarding body size and
morphology, we could divide these genera in two groups: Spurilla and Baeolidia, and Berghia and Limenandra
(Fig. 15). The body of the members of the first group is quite broad although species of Baeolidia are usually
smaller and wider than species of Spurilla. On the other hand, Berghia and Limenandra species are both slender but
Limenandra species are more cylindrical. The rhinophorial ornamentation is also very useful to separate these
genera. Although Berghia, Baeolidia and Limenandra were supposed to have papillate rhinophores, we could find
consistent differences in the morphology of the papillae. While the rhinophorial papillae of Berghia are usually
asymmetrical, being more or less rounded on posterior side, and elongate and perpendicular to rhinophores on their
lateral sides (see figure 2 of Carmona et al. 2014b), in Limenandra they are bifurcated (see figure 4 of Carmona et
al., 2014c). In Baeolidia, we find more variability in the morphology of these papillae and in the ornamentation of
the rhinophores in general (Fig. 2). However, due to the particular rhinophores together with other morphological
features, Baeolidia cannot be confounded with Spurilla, Limenandra, and Berghia. Although the arrangement of
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the cerata is not appropriate to separate genera in Aeolidiidae (Carmona et al. 2013), the morphology of the cerata
may give some information. Our research suggests that all the Berghia species have cylindrical cerata, with a round
apex and uniform diameter throughout most of their length (Carmona et al. 2014b); the cerata of Limenandra may
have papillae over their surface (Carmona et al. 2014c); Spurilla has cerata that are recurved inwards (Carmona et
al. 2014a), which are also present in some species of Baeolidia. However, the latter usually have leaf-shaped
cerata, being broad at their base and narrowing towards the apices.
Regarding the radular morphology (Fig. 16), only Limenandra is easily recognizable from the remaining
genera (Fig. 16C). The radular teeth of Spurilla, Berghia and Baeolidia are very similar, corresponding with the
radular teeth represented in the Figure 17. Although there is quite a lot of intrageneric variability, Figure 16 depicts
the archetypical radular teeth for each genus.
FIGURE 17. Most common radular teeth morphology in Aeolidiidae.
In terms of biogeography, so far Berghia is the only genus restricted to one ocean basin (Atlantic-
Mediterranean) (Fig. 18A) although the occurrence of otherwise Indo-Pacific Baeolidia in the Atlantic Ocean with
B. cryoporos would need further comparatives studies (Fig. 18B).
In order to facilitate the identification of these four genera, here we present a small key based on external
characters and distribution.
1 Rhinophores perfoliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spurilla
- Rhinophores ribbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baeolidia macleayi
- Rhinophores smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baeolidia ransoni
- Rhinophores papillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Notum with colouration pattern of concentric circles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limenandra
- Notum without colouration pattern of concentric circles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Cerata cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Cerata non-cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 Indo-Pacific distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baeolidia dela
- Atlantic-Mediterranean distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Berghia
5 Remaining species of Baeolidia.
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FIGURE 18. Map of the range of the different species of Spurilla, Berghia, Limenandra, and Baeolidia. A. Berghia and
Limenandra. B. Spurilla and Baeolidia. Abbreviation: Isolated records of Baeolidia cryporos (star) and Spurilla braziliana
(square).
Revised diagnosis of Baeolidia
Body mainly short and broad. Foot corners tentaculiform or rounded. Excluding B. ransoni and B. macleayi,
rhinophores are papillate. Rhinophorial papillae may be elongate or short and rounded. Cerata arranged in arches,
rows or both, arches and rows. Cerata dorso-ventrally flattened and normally leaf-like. Radular teeth pectinate,
usually bilobed and with a central cusp. Masticatory jaw edge usually smooth. Oral glands, when present, fragile
and spongy.
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Acknowledgments
We are deeply grateful to all individuals who helped to collect and provided specimens and images for this study,
including C. Pittman, R. Nakano, S. Johnson and Y. Tibiriça. This work was supported by several research grants
(CGL2006-05182/BOS and CGL2010-17187, Spanish Ministry of Economy and Competitiveness (includes the
early Ministry of Sciences and Innovation) to J. L. Cervera; California Academy of Sciences, National Science
Foundation (DEB 0329054 PEET) to T. M. Gosliner; and a Research Grant of the Malacological Society of
London to L. Carmona. This is CEI·MAR journal publication 56.
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... The aeolidiid B. moebii is native to the Indo-Pacific realm, being described from Mauritius (Bergh 1888) and subsequently recorded from additional localities (Tanzania, Mozambique, Reunion Island, Seychelles, Indonesia, east Australia, the Philippines, South Korea, Japan, the Marshall Islands, Hawaii, California, and west Mexico: Koh 2008;Carmona et al. 2014b and references therein). Recently, B. moebii has been also recorded from the northern and central Red Sea from Egypt, Israel (Eilat, Gulf of Aqaba), and Saudi Arabia (Jeddah) (Yonow 2000(Yonow , 2008, from where it presumably entered the Mediterranean Sea through Lessepsian migration (Paz-Sedano et al. 2019;Çinar et al. 2021). ...
... Further confirming this statement, all Mediterranean records of the species come from the Levant Sea (Figure 1 As for A. lurana, B. moebii also suffered from various nomenclatural and taxonomic issues in the past, and was misidentified with other taxa (Bergh 1888;Gosliner 1979Gosliner , 1985Miller 2001). However, the review of the genus carried out by Carmona et al. (2014b), coupled with the integrative taxonomic analysis carried out on the Cypriot specimen (Paz-Sedano et al. 2019), allowed clarification of the above-mentioned mismatches. In particular, the taxa Baeolidia major Eliot, 1903 and Baeolidia major amakusana Baba, 1937 were considered as junior synonyms of B. moebii (see Carmona et al. 2014b), whereas the Red Sea photographic records of Baeolidia australis (Rudman, 1982) were confuted as they were based on juvenile B. moebii, thus suggesting that only B. moebii occurs in the Red Sea and that it is the only Baeolidia species that has entered the Mediterranean Sea since at least 2007 (Paz-Sedano et al. 2019). ...
... However, the review of the genus carried out by Carmona et al. (2014b), coupled with the integrative taxonomic analysis carried out on the Cypriot specimen (Paz-Sedano et al. 2019), allowed clarification of the above-mentioned mismatches. In particular, the taxa Baeolidia major Eliot, 1903 and Baeolidia major amakusana Baba, 1937 were considered as junior synonyms of B. moebii (see Carmona et al. 2014b), whereas the Red Sea photographic records of Baeolidia australis (Rudman, 1982) were confuted as they were based on juvenile B. moebii, thus suggesting that only B. moebii occurs in the Red Sea and that it is the only Baeolidia species that has entered the Mediterranean Sea since at least 2007 (Paz-Sedano et al. 2019). Despite the resolution of these taxonomic issues, little is known about the ecology and reproduction of B. moebii, both in its native and invaded ranges, apart from the facts that it feeds on different genera of anemones (Gosliner 1979;McDonald and Nybakken 1997) and hosts symbiotic zooxanthellae (Rudman 1982). ...
Article
Full-text available
The nudibranch Baeolidia moebii Bergh, 1888 is a sea slug widely distributed in the Indo-Pacific, including the Red Sea, from where it presumably entered the Mediterranean Sea through Lessepsian migration. Although the first record of this species in the invaded area dates back to at least 2007, it is only known so far from the Mediterranean basin based on three scattered records of single individuals, all found in the Levant Sea (Turkey and Cyprus). The present study reports the presence of B. moebii in Kalloni Gulf, a semi-enclosed shallow embayment located in Lesvos Island, a record that widens its known distribution to Greece and the Aegean Sea. Field observations of more than 150 individuals encountered in a single sampling event also allowed observation of intraspecific morphological variation and several specimens laying egg masses, thus first reporting of a spawning aggregation and providing concrete confirmation of its establishment success in the entire basin. Monitoring of such spawning events and aggregations may contribute to a better understanding of the population dynamics of the species. The discontinuous known distribution of B. moebii in the eastern Mediterranean can be attributed to false absence of the species due to its cryptic behaviour, its under-sampled habitat, or to a truly very patchy distribution because of secondary dispersion by currents or vessels, with establishment only in areas of particularly favourable conditions, such as the Kalloni Gulf.
... nia de Palythoa caribaeorum observada en Sao Vicente con aeólidos (1) y sus puestas (2). A la hora de ubicar nuestra especie en Aeolidiopsis Pruvot-Fol, 1964(según Ortea, Caballer & Espinosa, 2003 o en Baeolidia, Bergh, 1888, era obligado tener en cuenta la publicación de Rudman (1982) sobre el primero y la revisión del género Baeolidia, que hacen Carmona, Pola, Gosliner. & Cervera (2014), autores que lo definen como un género Indopacífico, en el que incluyen Baeolidia cryoporos Bouchet, 1977, de aguas profundas del Atlántico Nordeste. Sobre Baeolidia benteva Marcus, 1958, de Ubatuba (Brasil), matizan que es de otro género y no hacen ninguna referencia a B. quoyi Pruvot-Fol, 1934, de localidad tipo desconocida. En García ...
... En la revisión de Baeolidia de Carmona et al. (2014) no se hace referencia a una especie del género registrada por Pruvot-Fol (1953) en las vecinas costas de Senegal como: Baeolidia möbii Bergh, 1888, syn. ¿Baeolidia quoyi Pruvot-Fol, 1934. ...
... Fig. 19, sin localidad tipo; fue discutida en Ortea, Caballer & Espinosa (2003) comparando sus figuras originales con Burnaia helicochorda (Miller, 1987) sugiriendo la posibilidad de que fueran especies sinónimas por la forma de las placas raquídeas de la rádula y por el borde masticador de las mandíbulas, muy largo y espiculoso en ambas especies. Dicho trabajo de Ortea et al. (2003) es ignorado por Carmona et al. (2014) en la revisión de Baeolidia al igual que lo hicieron más tarde en la de Burnaia Miller, 2001 y su reubicación en Facelinidae (Carmona, Pola, Gosliner & Cervera, 2015), un ejemplo más de su desconocimiento de la literatura relacionada. (2010) estudian los simbiontes de zoantarios en las islas de Cabo Verde, recogiendo la presencia de zooxantelas del género Symbiodinium en los tejidos de P. caribaeorum, género con una coloración anaranjada y de forma esférica (D, tomado de Wikipedia © Allisonmlewis). ...
Article
Full-text available
A unique aeolidacean from Cape Verde Islands, whose diet on zoantarios (Cnidaria: Anthozoa) relates it to Baeolidia palythoae Gosliner, 1985, of the coast of South Africa, is described as a new species of Aeolidiopsis Pruvot-Fol, 1956, without Atlantic representatives, at the same time a new genus and a new family are proposed to relocate it in the taxonomic discussion. Data on its external and internal anatomy are provided and it is discussed with other species that feed on Zoantarios in the Indo-Pacific.
... No additional data are available for the specimen from Turkey but in Cyprus, the leaves were covered by hydrozoans such as Aglaophenia harpago (Schenck, 1965) and Sertularia sp., and by bryozoans such as Electra posidoniae Gautier, 1954 and Calpensia nobilis (Esper, 1796). (Baba, 1933) CASIZ184520 Japan JQ997058 Baeolidia japonica (Baba, 1933) CASIZ 181357 Philippines JQ997057 Baeolidia moebii (Bergh, 1888) CASIZ186211 Philippines JQ997061 Baeolidia moebii (Bergh, 1888) CASIZ177602 Philippines HQ616770 Baeolidia moebii (Bergh, 1888) CASIZ 180327 Hawaii JQ997060 Baeolidia moebii (Bergh, 1888) MNCN 15.05/54987 Hawaii HQ616771 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51948 Marshall Island JX087550 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51949 Marshall Island JX087551 Baeolidia moebii (Bergh, 1888) a SZN-MOL0020 Cyprus MK922511 Baeolidia ransoni (Pruvot-Fol, 1956) CASIZ 186209 Philippines JQ997043 Baeolidia rieae (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184525 Japan JQ997046 Baeolidia salaamica (Rudman, 1982) CASIZ 177599 Philippines JQ997062 Baeolidia salaamica (Rudman, 1982) CASIZ 177397 Philippines JQ997047 Baeolidia salaamica (Rudman, 1982) CASIZ 180330 Philippines JQ997048 Baeolidia scottjohnsoni (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184503 Marshall Islands JQ997045 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.1 Marshall Islands JQ997055 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.2 Marshall Islands JQ997056 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 186210 Philippines JQ997054 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 177716 Philippines JQ997051 a The Baeolidia specimen sequenced during the present study. ...
... No additional data are available for the specimen from Turkey but in Cyprus, the leaves were covered by hydrozoans such as Aglaophenia harpago (Schenck, 1965) and Sertularia sp., and by bryozoans such as Electra posidoniae Gautier, 1954 and Calpensia nobilis (Esper, 1796). (Baba, 1933) CASIZ184520 Japan JQ997058 Baeolidia japonica (Baba, 1933) CASIZ 181357 Philippines JQ997057 Baeolidia moebii (Bergh, 1888) CASIZ186211 Philippines JQ997061 Baeolidia moebii (Bergh, 1888) CASIZ177602 Philippines HQ616770 Baeolidia moebii (Bergh, 1888) CASIZ 180327 Hawaii JQ997060 Baeolidia moebii (Bergh, 1888) MNCN 15.05/54987 Hawaii HQ616771 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51948 Marshall Island JX087550 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51949 Marshall Island JX087551 Baeolidia moebii (Bergh, 1888) a SZN-MOL0020 Cyprus MK922511 Baeolidia ransoni (Pruvot-Fol, 1956) CASIZ 186209 Philippines JQ997043 Baeolidia rieae (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184525 Japan JQ997046 Baeolidia salaamica (Rudman, 1982) CASIZ 177599 Philippines JQ997062 Baeolidia salaamica (Rudman, 1982) CASIZ 177397 Philippines JQ997047 Baeolidia salaamica (Rudman, 1982) CASIZ 180330 Philippines JQ997048 Baeolidia scottjohnsoni (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184503 Marshall Islands JQ997045 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.1 Marshall Islands JQ997055 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.2 Marshall Islands JQ997056 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 186210 Philippines JQ997054 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 177716 Philippines JQ997051 a The Baeolidia specimen sequenced during the present study. ...
... No additional data are available for the specimen from Turkey but in Cyprus, the leaves were covered by hydrozoans such as Aglaophenia harpago (Schenck, 1965) and Sertularia sp., and by bryozoans such as Electra posidoniae Gautier, 1954 and Calpensia nobilis (Esper, 1796). (Baba, 1933) CASIZ184520 Japan JQ997058 Baeolidia japonica (Baba, 1933) CASIZ 181357 Philippines JQ997057 Baeolidia moebii (Bergh, 1888) CASIZ186211 Philippines JQ997061 Baeolidia moebii (Bergh, 1888) CASIZ177602 Philippines HQ616770 Baeolidia moebii (Bergh, 1888) CASIZ 180327 Hawaii JQ997060 Baeolidia moebii (Bergh, 1888) MNCN 15.05/54987 Hawaii HQ616771 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51948 Marshall Island JX087550 Baeolidia moebii (Bergh, 1888) MNCN/ADN: 51949 Marshall Island JX087551 Baeolidia moebii (Bergh, 1888) a SZN-MOL0020 Cyprus MK922511 Baeolidia ransoni (Pruvot-Fol, 1956) CASIZ 186209 Philippines JQ997043 Baeolidia rieae (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184525 Japan JQ997046 Baeolidia salaamica (Rudman, 1982) CASIZ 177599 Philippines JQ997062 Baeolidia salaamica (Rudman, 1982) CASIZ 177397 Philippines JQ997047 Baeolidia salaamica (Rudman, 1982) CASIZ 180330 Philippines JQ997048 Baeolidia scottjohnsoni (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 184503 Marshall Islands JQ997045 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.1 Marshall Islands JQ997055 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 187741.2 Marshall Islands JQ997056 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 186210 Philippines JQ997054 Baeolidia variabilis (Carmona, Pola, Gosliner & Cervera, 2014) CASIZ 177716 Philippines JQ997051 a The Baeolidia specimen sequenced during the present study. ...
... The nudibranchs that feed on zoantharians (zoaeating nudibranchs) have been known to the aquarium community for several decades and are considered a pest to Zoanthus species in tropical marine aquariums. Several individuals of the West Pacific aeolid nudibranch Baeolidia rieae Carmona, Pola, Gosliner & Cervera, 2014 were found four times feeding on specimens of Zoanthus Lamarck, 1801, a commercial zoantharian genus, in the aquarium store Urban Natura (Naturalis Insignia SL) and once again in a private aquarium, both in Barcelona (Spain). In this study we provide new morphological data about this species and a first description of its spawn. ...
... Finally, Bayesian Inference and Maximum Likelihood analyses were performed for a dataset concatenating the COI, 16S and H3 Baeolidia sequences extracted from Genbank (Supplementary Material). Carmona, Pola, Gosliner & Cervera, 2014 Type locality and habitat Japan, Amami-Ohoshima Island found in 7 m depth of water close to green zoantharians. ...
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Baeolidia rieae Carmona, Pola, Gosliner & Cervera, 2014 was described based on a single specimen from the subtropical Amami-Oshima Island (SW Japan). In this study, we recorded B. rieae feeding on a zoantharian, Zoanthus sp., in an aquarium store in Barcelona (Spain). The morphological and molecular match from the original description corroborated its identification. We discuss the possible origin of the B. rieae in aquarium stores, its resistance to ordinary disinfection processes, the harm it may cause to commercial coral species, and how it might develop into an invasive species if it continues to be found in aquarium stores around the world without treating the problem properly. Finally, we describe for the first time the spawn of B. rieae and its eggs features, and provide new morphological and biological data. Résumé : Risque associé au nudibranche Baeolidia rueae (Mollusca : Gastropoda : Heterobranchia) via le commerce international d'aquariophilie. Baeolidia rieae Carmona, Pola, Gosliner & Cervera, 2014 a été décrite avec un seul spécimen de l'île subtropicale d'Amami-Oshima (sud-ouest du Japon). Dans cette étude, nous signalons B. rieae se nourrissant du zoanthaire Zoanthus sp. dans un magasin d'aquarium à Barcelone (Espagne). La correspondance morphologique et moléculaire de la description originale a corroboré son identification. Nous discutons de l'origine du B. rieae dans les aquariums, de sa résistance aux processus de désinfection ordinaires, des dommages qu'il cause aux espèces de corail commerciales et de la façon dont il pourrait se développer en une espèce envahissante s'il continue à être trouvé dans les magasins d'aquarium du monde entier sans traiter correctement le problème. Enfin, nous décrivons pour la première fois la ponte de B. rieae et les caractéristiques de ses oeufs, et fournissons de nouvelles données morphologiques.
... Distribution. Across the Pacific including Japan (Ono 2004), French Polynesia (Pruvot-Fol 1956), Hawaii (Carmona et al. 2014a), the Philippines, Australia, and Papua New Guinea ). Here representing a first record for Thai waters. ...
... Distribution. Across the Indo-Pacific including Tanzania (Rudman 1982), Japan (Ono 2004), Korea (Koh 2006), Hawaii (Carmona et al. 2014a), Philippines, and Papua New Guinea ). Here representing a first record for Thai waters. ...
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Improved access to field survey infrastructure throughout South-East Asia has allowed for a greater intensity of biodiversity surveys than ever before. The rocky bottoms and coral reef habitats across the region have been shown to support some of the highest sea slug biodiversity on the planet, with ever increasing records. During the past ten years, intensive SCUBA surveys have been carried out at Koh Tao, in the Gulf of Thailand, which have yielded remarkable findings in sea slug biology and ecology. In this work a brief history of sea slug biodiversity research from Thailand is covered and a complete inventory of sea slugs from Koh Tao, Gulf of Thailand is provided. This inventory is based on surveys from 2012 to 2020, with previously unreported findings since 2016. Habitat specificity and species-specific ecology are reported where available with a focused comparison of coral reef habitats and deeper soft-sediment habitats. The findings contribute 90 new species records for Thai waters (92 for the Gulf of Thailand) and report a remarkable consistency in the proportional diversity found to be exclusive to one habitat type or another. Additionally, taxonomic remarks are provided for species documented from Koh Tao that have not been discussed in past literature from Thailand, and a summary of previous records in the Indo-West Pacific is given.
... Zoanthids are identified morphologically based on the types of polyp (immersae/ liberae/ intermediae) (Reimer et al., 2006), colors of polyps, tentacles and oral slits, number of tentacles, size of colonies and oral discs, and sediment encrustation inside tissue or not. The collected zoanthid and associated species were identified morphologically with the aid of online available literatures (Esper 1805;Haddon and Shackleton, 1891;Carlgren, 1900;Pax and Muller, 1957;Reimer et al., 2011;Gul, 2013;Khushali, 2014;Koupaei et al., 2014;Reimer et al., 2014;Nasir et al., 2018, andLinnaeus, 1758;Rudy et al., 1983;Bosch et al., 1995;Budd, 2007;Apte et al., 2010;Bano et al., 2011;Afsar et al., 2012;Carmona et al., 2014a, 2014b, Agustiadi and Luthfi, 2017Ghani et al., 2017;Ghani et al., 2018aGhani et al., , 2018bKazmi et al., 2018;Ghani et al., 2019). ...
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One intertidal (Buleji) and two diving sites (Mubarak village and Churna Island (2 stations)) were visited for the collection of zoanthids (with morphotypes) (Zoanthus sansibaricus (6), Z. vietnamensis (2), Palythoa tuberculosa and P. mutuki (2) from Buleji), while Z. sansibaricus (morph-4) and P. tuberculosa from both diving sites. Moreover, 29 intertidal (26 Molluscans and 1 Annelida, Arthropoda and Echinodermata) and 1 underwater Porite (Tubipora musica) associates were identified from Buleji and diving station-I of Churna Island, respectively, in order to study their ecology. All zoanthids (including morphotypes) and porite were reared in aquaria to determine their growth potentials in artificial settings. During culture, the recorded parameters ranged between 35-36.5‰ salinity, 7.3-7.7 pH, 25.4-31.0˚C temperature, 0.4-0.3ppm NO2-, 0.12-0.08ppm NO2-N, 500-545mg/l Ca, 0.23-0.25mg/l NH3 and 7-10mg/l DO2. The Z. sansibaricus (morph-1c) and Z. vietnamensis (morph-1), P. tuberculosa(c), and P. mutuki (morph-1c & 2a) lived up to 21 weeks, showing 41, 51, 18, and 32 & 66 average growth rate or average polyp growth percentage (APG%) respectively, while Z. sansibaricus (morph-4a), P. mutuki (morph-2b) and T. musica were sustained up to 19, 6 and 4 weeks with APG% of 0,-92 and-93, respectively.
... In 1888, Bergh first collected and described species from Mauritius Island belonging to both lineages; Cladobranchia and Anthobranchia (Bergh, 1888). Bergh introduced the genus Baeolidia in 1888, based on the description of a single specimen, Baeolidia moebii which eventually contained contradictory information and thus, led to morphological confusion (Carmona et al., 2014a). ...
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Nudibranchs are considered one of the most diverse groups of opisthobranchs. Their history in Mauritius dates from 1832, with first records appeared in expedition reports and systematic works. Recent review of their biodiversity in Mauritius identified 23 species. The present study provides a list of nudibranch species using data from both systematic works and internet records as a means of maintaining an inventory from Mauritius. Some 105 species belonging to 20 families (excluding undescribed taxa and those recorded as cf.) have been identified in Mauritius. Most species have been collected in the north-west part of the island which is dominated by hotels and not subjected to easterlies which could be one reason explaining their abundance. Providing a list of nudibranchs species is important, to be able to design better ways of conserving them in the future, if the need arises. With a wide maritime zone and considered as a striking biodiversity hotspot, further species might be discovered from both Mauritius and Rodrigues.
... Although the Brazilian specimens of Okenia studied here closely fit the holotype and the topotype of O. polycerelloides analyzed, it is important to consider the possibility that they are part of a species complex. Recent molecular studies have shown that sea slug species that are morphologically similar and were previously considered as only a single taxon are in fact a complex of different species with more restricted distributions than first thought (e.g., Carmona et al. 2011Carmona et al. , 2014aCarmona et al. , 2014bCarmona et al. , 2014cCarmona et al. , 2014d. In order to test that, we tried to obtain DNA sequences of O. polycerelloides from Brazil and the Canary Islands, but we failed in amplifying the topotype. ...
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Okenia polycerelloides (Ortea & Bouchet, 1983) is a small goniodoridid nudibranch originally described from the Canary Islands. Its taxonomic history has been problematic since its original description, a situation that worsened after this taxon was synonymized with Okenia zoobotryon (Smallwood, 1910), one of the most problematic Okenia Menke, 1830 species. Because of their external similarity, it has been difficult to determine the differences between the two taxa without a meticulous anatomical study. Thus, we present herein the first complete anatomical study of O. polycerelloides, based on specimens from the type locality and from Brazil (the latter previously identified as O. zoobotryon). A topotype of O. zoobotryon was also analyzed. Additionally, we also performed a preliminary molecular analysis on these species. O. polycerelloides can be distinguished externally from O. zoobotryon by the absence of integumentary spicules and the presence of a posterodorsal papilla, and internally by characteristics of the digestive (shape of the salivary glands, length of the typhlosole) and reproductive systems (diameter and length of the penial sac; cilia on the penis; and shapes of the seminal receptacle, bursa copulatrix, and ampulla). Molecular analysis revealed a high genetic divergence in both COI and H3 genes between O. polycerelloides and O. zoobotryon, which support both species as distinct. Therefore, we present a redescription and propose to restore the species status of O. polycerelloides, and consider the records of O. zoobotryon in the South Atlantic Ocean as attributable to O. polycerelloides, until new evidence may suggest otherwise.
... Although the Brazilian specimens of Okenia studied here closely fit the holotype and the topotype of O. polycerelloides analyzed, it is important to consider the possibility that they are part of a species complex. Recent molecular studies have shown that sea slug species that are morphologically similar and were previously considered as only a single taxon are in fact a complex of different species with more restricted distributions than first thought (e.g., Carmona et al. 2011Carmona et al. , 2014aCarmona et al. , 2014bCarmona et al. , 2014cCarmona et al. , 2014d. In order to test that, we tried to obtain DNA sequences of O. polycerelloides from Brazil and the Canary Islands, but we failed in amplifying the topotype. ...
Article
Okenia polycerelloides (Ortea & Bouchet, 1983) is a small goniodoridid nudibranch originally described from the Canary Islands. Its taxonomic history has been problematic since its original description, a situation that worsened after this taxon was synonymized with Okenia zoobotryon (Smallwood, 1910), one of the most problematic Okenia Menke, 1830 species. Because of their external similarity, it has been difficult to determine the differences between the two taxa without a meticulous anatomical study. Thus, we present herein the first complete anatomical study of O. polycerelloides, based on specimens from the type locality and from Brazil (the latter previously identified as O. zoobotryon). A topotype of O. zoobotryon was also analyzed. Additionally, we also performed a preliminary molecular analysis on these species. O. polycerelloides can be distinguished externally from O. zoobotryon by the absence of integumentary spicules and the presence of a posterodorsal papilla, and internally by characteristics of the digestive (shape of the salivary glands, length of the typhlosole) and reproductive systems (diameter and length of the penial sac; cilia on the penis; and shapes of the seminal receptacle, bursa copulatrix, and ampulla). Molecular analysis revealed a high genetic divergence in both COI and H3 genes between O. polycerelloides and O. zoobotryon, which support both species as distinct. Therefore, we present a redescription and propose to restore the species status of O. polycerelloides, and consider the records of O. zoobotryon in the South Atlantic Ocean as attributable to O. polycerelloides, until new evidence may suggest otherwise.
... Only B. alba has a widespread distribution, whereas the remaining species of the genus are endemics with restricted ranges. Other aeolidiid genera such as Anteaeolidiella (Carmona et al., 2014a) and Spurilla (Carmona et al., 2014b) have the same kind of distribution patterns. These genera show endemics in peripheral areas such as South Africa and Mozambique, suggesting that examination of supposed B. alba from those localities, which could not be studied here, are necessary in order to confirm its identity. ...