What the shell tell in Aglajidae: A new genus for Aglaja felis (OPISTHOBRANCHIA: CEPHALASPIDEA)

              Rev. Acad. Canar. Cienc., Vol. XXVI, 83-1 19 (diciembre de 2014)
WHAT THE SHELL TELLS IN AGLAJIDAE:
ANEW GENUS FOR Aglaja felis (OPISTHOBRANCHIA: CEPHALASPIDEA)
Ortea, J .,
2 J *Caballer, M., 4 Moro, L. & 5 Espinosa, J.
1 Departamento BOS, Oviedo University, Spain. E-mail: jortea@uniovi.es
2 Museum National d'Histoire Naturelle, 55 rue de Buffon, 75005 Paris, France
1 Centro de Oceanologia y Estudios Antarticos. IVIC. Ctra. Panamericana Km 1 1, Miranda, Venezuela
E-mail: manuelcaballergutierrez@hotmail.com
4 Servicio de Biodiversidad, Gobiemo de Canarias, S/C de Tenerife, Canary Islands, Spain
E-mail: leopoldo.moroabad@gobiemodecanarias.org
Tnstituto de Oceanologia, Playa, La Habana, Cuba
* Corresponding author: mcaballergutieiTez@mnhn.fr
ABSTRACT
In the most recent molecular phylogeny of the Aglajidae, the Caribbean species
Aglaja felis Marcus & Marcus, 1970 was transferred to the Japanese genus Nakamigawaia
Kuroda & Habe in Habe, 1961 without justification or support. The assumed N. felis was
used as the only representative of the genus in the phylogeny, which did not include the
type species: Nakamigawaia spiralis Kuroda & Habe in Habe, 1961. The material deter-mined
as A. felis came from Bahamas, Papua New Guinea and Philippines and the conclu-sion
was that they were a complex of species; the Indo-Pacific ones possibly new and all
of them belonging to the genus Nakamigawaia. Nonetheless, a simple comparison of shells
of the type species of Aglaja Renier, 1 88 1 and Nakamigawaia with A. felis, shows remark-able
and supra-specific differences. Thus, in absence of N. spiralis, Nakamigawaia would
not be represented in the phylogeny of the Aglajidae, but a distinct taxa, distributed in the
Caribbean and the Indo-Pacific. Migaya Ortea, Caballer & Espinosa new genus, is proposed
to relocate A. felis. This genus is characterized by bearing sensorial bristles on the head, by
its internal shell and supported by the molecular phylogeny. In the same phylogeny, the
genus Chelidonura A. Adams, 1850 was fragmented in several independent clades, one of
them, characterized by the shell bearing a crest in the protoconch. Two new species of this
clade are described in this paper.
The shell can be very helpful to assign the right names in the Aglajidae and using it is
recommended even in papers where the molecular techniques are applied. A visual analysis
of the usefulness of the shell within the group is performed using the established phylogeny
as a frame for comparison.
Key words: Mollusca, Gastropoda, systematics, Migaya, new genus, new species,
Caribbean Sea.
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RESUMEN
En la filogenia molecular mas reciente realizada sobre la familia Aglajidae, la especie
caribena Aglajafelis Marcus & Marcus, 1970 fue transferida al genero japones Nakamigawaia
Kuroda & Habe en Habe, 1961 sin ninguna argumentacion o soporte. La asumida como N. felis
fue usada como el linico representante del genero en la filogenia, que no incluyo la especie
tipo:
Nakamigawaia spiralis Kuroda & Habe en Habe, 1961 . Dicho material provenia de Ba-hamas,
Papua Nueva Guinea y Filipinas y la conclusion del trabajo fue que se trataba de un
complejo de especies; las Indo-Pacificas posiblemente nuevas y todas ellas pertenecientes al
genero Nakamigawaia. Sin embargo, una simple comparacion de las conchas de las especies
tipo de Aglaja Renier, 1881 y Nakamigawaia con la de A. felis, deja en evidencia grandes di-ferencias
que las situan en generos distintos. Por ello, en ausencia de N. spiralis, Nakamiga-waia
no estaria representada en la filogenia de Aglajidae sino un taxon distinto distribuido en
el Caribe y en el Indo-Pacifico. Se propone Migaya Ortea, Caballer & Espinosa nuevo genero
para reubicar A. felis. Este genero se caracteriza por presentar quetas sensoriales en la cabeza
y por su concha interna, estando respaldada por los resultados de la ultima filogenia molecu-lar
publicada. Adicionalmente, en dicha filogenia el genero Chelidonura A. Adams, 1850 es
fragmentado en varios clados independientes, uno de los cuales se caracteriza por su concha
interna, la cual presenta una cresta en la protoconcha. En este trabajo se describen dos espe-cies
nuevas de dicho clado.
La concha es un caracter muy util para asignar nombres en aglajidos y usarlo es reco-mendable
incluso en articulos en los que se aplican tecnicas de biologia molecular. Por ello,
se aborda un analisis visual de la utilidad de la concha dentro de la Familia, usando la filoge-nia
establecida como marco de comparacion.
Palabras clave: Mollusca, Gastropoda, sistematica, Migaya, nuevo genero, nuevas
especies, mar Caribe.
1. INTRODUCTION
Aglaja felis Marcus & Marcus, 1970, is a little black-coloured sea slug originally de-scribed
from the Caribbean Sea. Prior to 2014, Nakamigawaia Kuroda & Habe in Habe, 1961
was considered monotypic and endemic to the south east coast of Japan. In the latest molecu-lar
phylogeny of the Aglajidae Pilsbry, 1895 owed to CAMAC'HO-GARCIA, ORNELAS-GATDULA,
GOSLINER, & VALDES (2014), these authors transferred A. felis to the genus
Nakamigawaia with no explanation or support. ‘W. felis” was used as the only nominal rep-resentative
ofNakamigawaia and was recorded from Bahamas, Papua New Guinea and Philip-pines
(CAMACHO-GARCIA et al. 2014. Accordingly, the genus “Nakamigawaia”
represented by A. felis, resulted in a complex of 3 clades: one including specimens from Ba-hamas,
other including specimens from the Indo-Pacific (possibly composed of two distinct
species) and the last including two specimens determined as Melanochlamys sp. 1 . Nakami-gawaia
spiralis Kuroda & Habe in Habe, 1961, described from Japan, is the type species of
the genus Nakamigawaia, but it was not included in the phylogeny of the family (CAMA-C'HO-
GARCIA et al. 2014). A simple comparison of A. felis with N. spiralis (not performed
by CAMAC'HO-GARCIA et al., 2014), reveals that they can’t belong to the same genus. Mi-gaya
Ortea, Caballer & Espinosa, new genus, is described in this paper to relocate Aglaja felis.
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In the same phylogeny, CAMACHO-GARCIA et al. (2014) found several different
clades formerly belonging to the genus Chelidonura A. Adams, 1 850, two of them in the
Caribbean. The existence of these clades is consistent with the proposal of two different lin-eages
of Chelidonura in the Caribbean proposed by ORTEA, ESPINOSA, CABALLER,
MORO & BACALLADO (2012). One of this clades, includes the type species of the genus,
Chelidonura hirundinina (Quoy & Gaimard, 1833), the other is distinguished by bearing a
crest in the protoconch of the shell, and includes C. pusilla Ortea, Moro & Espinosa, new
species, and C. quadrata Ortea, Caballer & Espinosa, new species, described in this paper.
The external anatomy and coloration are not typically valid characters to distinguish
species in the Aglajidae (ORNELAS-GATDULA, DUPONT & VALDES, 2011; ORNELAS-GATDULA&
VALDES, 2012; ORNELAS-GATDULA, CAMACHO-GARCIA, SCHRODL,
PADULA, HOOKER, GOSLINER & VALDES, 2012). Thus, to warrant the correct use of the
names (even in molecular papers), the characters of the species used must be consistent with
their original descriptions. Select the correct ones depending on the family is a heavy matter.
However, the shell has been proposed to be a valid systematic character to distinguish taxa and
to assign the correct names (ORTEA et al., 2012). The usefulness of the shell to distinguish
genus within the Aglajidae, is visually analyzed in the frame of the phylogeny performed by
CAMACHO-GARCIA et al. (2014).
2.
MATERIAL AND METHODS
The specimens ofA.felis were captured by hand from the intertidal to 6 m depth, from
1981 to 2013, in several localities covering the whole extension of the Caribbean Sea: Cuba,
Mexico, Costa Rica, Venezuela and Guadeloupe. The specimens of C. pusilla Ortea, Moro &
Espinosa, new species, and C. quadrata Ortea, Caballer & Espinosa, new species, were cap-tured
by hand between 1 and 6 m depth, in sandy bottoms in Cuba. They were photographed
alive (when possible) and data were taken on behavior, anatomy and coloration. Afterwards,
the animals were preserved in ethanol 96 %. To compare them with other Aglajidae, diagrams
and photos were made of the internal shell using an Olympus SZ16 stereomicroscope and
Nikon cameras.
For the comparison of the shells within the different clades in the Aglajidae (Figure 6):
1.
- The molecular phylogeny by CAMACHO-GARCIA et al. (2014) was assumed as
the correct frame.
2.
- The identity of the species given by CAMACHO-GARCIA et al. (2014) was as-sumed
to be true in the most of the cases (see Table 1).
3.
- Data on the species were obtained from supplementary material, original descrip-tions
or recent papers (Table 1 ).
4.
- No differences in the shell type where found between the members of the same
clade, thus, only one shell, the typical one, is showed in Figure 6.
5.
- When no data was available on the shell of one species, it was considered to have
the same kind than the rest of the clade.
6. When no data and no specific name were given by CAMACHO-GARCIA et al.
(2014) the specimen was considered to have the same kind of shell than the rest of
the clade on which it was included. If these specimens belonged to a separate clade
they were marked with a red question mark and each case was treated differently.
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7.
- Each terminal clade with different kind of shell was identified with a single genus.
When incongruences were found between clades that share species that were pre-viously
considered to belong to the same genus, the type species determined the true
generic identity of the clade.
8.
- Five clades were found lacking a formal generic name (after the descriptions in
this paper): Gen 1-5. No data at all was available for the Gen 1
.
9.
- The shells of the three genus not considered by CAMACHO-GARCIA et al. (2014)
are shown in Figure 6 in separated and unrelated lines.
A formal reconstruction of ancestral characters in the Aglajidae could not be performed
due to the lack of data on the real specimens used by CAMACHO-GARCIA et al. (2014).
Abbreviations:
IES, Instituto de Ecologia y Sistematica, Havana, Cuba.
JC, Jesus Ortea’s collections, Noreha, Asturias, Spain.
JEC, Jose Espinosa’s collections, Havana, Cuba.
LC, Leopoldo Moro’s collections, La Laguna, Tenerife, Canary Islands, Spain.
MC, Manuel Caballer collections, Boo de Pielagos, Cantabria, Spain.
MNHN, Museum National d’Histoire Naturelle, 55 rue de Buffon, 75005 Paris, France.
3. SYSTEMATICS
Order Cephalaspidea Fischer, 1883
Family Aglajidae Pilsbry, 1895
Genus Migaya Ortea, Caballer & Espinosa new genus
Type species:. Aglaja felis Marcus & Marcus, 1970: Studies on the Fauna of Curasao and other
Caribbean island 33: 9-10, figs. 1-3. Type locality: Majimo Reef, La Parguera, Puerto Rico.
Diagnosis: (Figure 6, B.2) Head bearing sensorial bristles. Internal shell simple, with a wing
in the upper edge. Teleoconch vestigial, reduced to an arched and narrow plate attached to the
protoconch (“wing”), which crosses the visceral region. Protococh is smooth, strong and
proportionally big, on the right side of the animal. See Migaya felis new comb, for further
description.
Etimology: Migaya, in the Cantabrian-Asturian language means crumb, small, fragmented,
breadcrumb, referring to the size of the shell of Migaya felis new comb., whose protoconch
detaches as a crumb.
Remarks: In the most recent approach to the classification of the Aglajidae, CAMACHO-GARCIA
et al. (2014) transferred A. felis, one of the most common species in the Caribbean,
to the genus Nakamigawaia in absence of data to support that nomenclatural act or any dis-cussion.
This authors clustered their specimens from Bahamas (A. felis), the Indo-Pacific
“felis” and the
“Melanochlamys spp” used by ANTHES, SCHULENBURG & MICHIELS
(2008), under the name N. felis, may be because they found external resemblances with N. spi-ralis?.
They didn’t provide any data on the specimens.
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In 1992 Gosliner determined some specimens from Hawaii as N. felis (PITTMAN &
FIENE, 2014). Photos of these specimens are available online and, as it can be observed, their
external appearance and their shells are really similar to the shell of “felis” from the Caribbean.
But the external anatomy and coloration are not typically valid characters to distinguish
species in the Aglajidae (ORNELAS-GATDULA, DUPONT & VALDES, 201 1; ORNELAS-GATDULA&
VALDES, 2012; ORNELAS-GATDULA, CAMACHO-GARCIA, SCHRODL,
PADULA, HOOKER, GOSLINER & VALDES, 20 1 2; VALDES, ORNELAS-GATDULA &
DUPONT, 2013). In consequence, it must have been the shell the reason why Gosliner
(PI TTMAN & FIENE, 2014) considered some Aglajidae from Hawaii conspecifics with A.
felis, a species from the Caribbean. The motivation to transfer this species to a genus endemic
to the south coast of Japan (Nakamigawaia ), with only one species to that date (N. spiralis),
with a radically different kind of shell (Figures 1 and 6) is unknown. Any support was given
to this act, but we have to consider (in absence of other explanation) that CAMACHO-GAR-CIA
et al. (2014) followed him.
ANTHES et al. (2008) described a “sickle-shaped shell” and draw a sketch (ANTHES
et al., 2008: fig. 2) from their “temporary Melanochlamys sp.” (used by CAMACHO-GAR-CIA
et al., 2014). This kind of shell would match with that of A. felis (Figure 1 C-E; RED-FERN,
2013), but is really different from the shell of N. spiralis (Figure 1 I-J).
N. spiralis has been studied in Japan by several authors in the last 50 years (HABE,
1961; HABE 1975; BABA, 1985; HABE 2001; SASAKI, 2008), so, we can asume that if
they illustrate always the same kind of shell with very little variations, this must be the shape
it has. In addition, BABA (1985) studied the species in detail, so as SASAKI (2008); a spec-imen
with a sickle-shaped shell as the one beared by A. felis would have been detected and il-lustrated
or described.
CAMACHO-GARCIA et al. (2014) used four specimens of A. felis from Bahamas,
which on their phylogeny appeared as a sister group with the two specimens of
“Melanochlamys spp” from Australia (ANTHES et al., 2008). This group was the sister
group of the clade formed by the specimens of “A. felis” from Philippines and Papua New
Guinea.
The facts are that, in their phylogeny, CAMACHO-GARCIA et al. (2014) used sev-eral
specimens of three different species from two different Oceans, with shared a “sickle-shaped
shell” that match with the shell of A. felis, whose morphology is very constant (own
data; REDFERN, 2013) and is shared by specimens from Hawaii determined by one of the au-thors
(Gosliner: PITTMAN & FIENE, 2014). This type of shell is radically different from
that of N. spiralis, from Japan, which is characterized by housing a lot of endemic fauna
(OKUTANI, 2000).
The conclusion is that CAMACHO-GARCIA etal. (2014) used three species from the
same genus, but they didn’t belong to Nakamigawaia. Thus, Nakamigawaia wouldn’t be rep-resented
in their phylogeny and the clade of “V. felis” (formerly A. felis), very distant from
Ag/aja (Figure 6: A. 2. 1.2. part) would lack a formal name, which is proposed in this paper as
Migava Ortea, Caballer & Espinosa new genus. Accordingly, the position of Nakamigawaia
within the Aglajidae remains to be clarified.
Migaya felis new comb., type species of the genus Migava Ortea, Caballer & Es-pinosa
new genus, has been included in the genus Ag/aja Renier, 1881. But, the internal shell
of the type species of Ag/aja is composed by a reduced protoconch, from which it emerges
a dextral haliotiform shell with remarkable growth striae (Figure 1 H). In the outer edge it
bears a prominence where the concavity changes to form a wing-shaped callus. In the upper
87
edge, the shell forms a wide sinistral spiral: the big wing, in whose edge there is an almost-transparent
membrane. This kind of shell is clearly different from the one in Migaya felis
new comb.
BABA (1985) made a detailed anatomical revision of Nakamigawaia spiralis, type
species of the genus Nakamigawaia, and gave complementary data to its original description,
which was based solely on the internal shell (Figures 1 I—J): solid, very calcified on its pos-terior
region, membranous in the anterior region, rolled in spiral, with three whorls, a longi-tudinal
groove and lacking a visible protoconch in the apex of the spire. This shell, whose
calcified region is illustrated by SASAKI (2008: Figure 13 E), is quite different from that
found in Migayafelis new comb. (Figures 1 C-E). Additionally, an external characteristic that
distinguishes Migaya felis new comb, from Nakamigawaia spiralis is the presence of senso-rial
bristles in the head or at the sides of the mouth (common in Aglajidae), absent in the
Japanese species.
As it can be seen in Figure 6, the shell of Migaya Ortea, Caballer & Espinosa new
genus, is unique within the Aglajidae.
Migaya felis (Marcus & Marcus, 1970) comb. nov.
(Figure 1)
Material examined: More than 100 specimens 2-6 mm long alive, collected in sandy bottoms up to 6
m depth in: Quiebrahacha, La Habana, Jibacoa, Cayo Coco, Maria La Gorda, Golfo de Batabano, Cien-fuegos,
Jardines de la Reina (Cuba) (JC, JEC); Puerto Morelos (Mexico) (JC); Manzanillo (Costa Rica)
(JC); Mochima, La Tortuga (Venezuela) (MC) and Guadeloupe (MNHN).
Supplementary material: Aglaja tricolorata Renier, 1807, 1 specimen, 40 mm long alive, collected at
night in a sandy-muddy bottom at 2 m depth, 8/04/2008, Arrecife, Lanzarote, Canary Islands, Spain
(LC).
Description: Body usually black, rarely with white heather, very flexible and flattened, four
times as long as wide (Figure 1 A-B). Cephalic shield occupies approximately the anterior 2/3
of the animal, with the anterior end bearing a narrow hyaline visor with maroon points. An-terior
lobes black, with 5-6 white sensorial filaments (Figures 1 F-G), of which the innermost
and the outermost are smaller. Posterior lobes rounded; equal in shape and proportions at rest,
but the left lobe is up to twice as big as the right one when the animal moves. Internal shell
amber-colored, with the protoconch in the right lobe and an arched plate, 450 pm long (in 4
mm long specimens), crossing the left one (Figures 1 C-E). REDFERN (2013: 727B & D) il-lustrates
similar shells, 500 pm long, from Bahamas. Shell plate absent. Teleoconch reduced
to a wing with fine oblique striae in the upper edge united to the larval shell. Protoconch rolled
one and a half whorls, extended forward as a soft visor, with a dorsal apophysis strengthen-ing
the union with the shell plate. There is no anterior membranose plate in the shell. Proto-conch
splits up easily when dry.
Habitat: Inhabits sandy bottoms, from the shore to 6 m depth. Feeds on cypridacid ostracods
(at least), a kind of prey not reported anteriorly in these animals.
88
Genus Chelidonura A. Adams, 1850
Type species: Bulla hirundinina Quoy & Gaimard, 1833
Shell: (Figure 6, A.2. 1.1) Thick, oval, scoop-shaped, with remarkable growth lines and a wing
in the upper edge, that ends in a small spike and do not surpass the protoconch. Teleoconch
long, rounded, with a solid edge. Protoconch lacking spines or a crest, strong and propor-tionally
big, separated from the shell by a callus.
Remarks: Head with sensorial bristles.
Clade “’Crested” Chelidonura
Diagnosis: (Figure 6, A. 2. 3) Scoop-shaped, with remarkable growth lines and the shoulder of
the teleoconch forming a “wing” proportionally big. Sometimes calcified. Protoconch with a
remarkable and strong crest. The callus sometimes covers the protoconch entirely and part of
the “wing”.
Species included: C. mariagordae Ortea, Espinosa & Moro, 2004, C. africana Pruvot-Fol,
1953, C. juancarlosi Ortea & Espinosa, 1998 and C. berolina Marcus & Marcus, 1970 ??.
Remarks: ORTEA et al. (2012) based on the morphology of the internal shell, pointed out the
existence of two different evolutive lines within the genus Chelidonura in the Atlantic; one
containing at least C. hirundinina, C. cubana Ortea & Martinez, 1997 and C. hummelincki
(Marcus & Marcus, 1970) and the other one, grouping at least C. africana, C. mariagoradae
(C. normani) and C. juancarlosi. CAMACHO-GARCIA et al. (2014), based on the analysis
of molecular data, found three monophyletic clades:
- A.2. 1 . 1 (= Chelidonura sensu stricto). including C. hirundinina among many others.
- A.2. 2 (part) (= Gen 3 in Figure 6), including several species from the Pacific.
- A. 2. 3 (=”Crested” Chelidonura ), including species from the Atlantic: C. mariagor-dae
(as C. normani Omelas-Gatdula, Dupont & Valdes, 2011) + C. africana + C.
berolina (?).
Thus, the “Crested” Chelidonura species are separated from other clades formerly in-cluded
in Chelidonura by molecular evidence (CAMACHO-GARCIA et al., 2014), and dis-tinguished
from them ( Chelidonura s.s. and Gen 3) by the general shape of the shell, by the
crest that bears in the protoconch and by the “wing” in the shoulder of the teleoconch.
Aglaja hummelincki, described from Puerto Rico, was redescribed by THOMPSON
(1977) based in specimens from Jamaica. He gave a good account on its external anatomy and
coloration. ORTEA, MORO & ESPINOSA (2007) considered C. hummelincki synonymous
to C. berolina. Later, ORNELAS-GATDULA etal. (2011), under the principle of first reviser
(?), transferred again C. hummelincki to the synonymy. They gave no data on the shell of C.
berolina, which remains unknown. So, the latter species is tentatively considered within this
clade, assuming that the name given by CAMACHO-GARCIA et al. (2014) is correct (see
Material and methods). The synonymy of C. hummelincki needs to be clarified with specimens
from the type locality.
89
Chelidonura pusilla Ortea, Moro & Espinosa new species
(Figures 2-3)
Holotvpe: 2 mm long alive, deposited in IES (39-101). Type locality: Reparto Nautico, Havana, Cuba.
Material examined: 7 specimens, 2-6 mm long alive, collected in sandy bottoms up to 6 m depth in the
type locality, july 1, 1999.
Etimoiogy: pusilla , in Latin: dwarf, because of the small size of this species.
Description: Body flattened, black, with the anterior end of the head and the posterior end ot
the cephalic shield somewhat discolored. One specimen with some disperse white spots.
Cephalic shield of the same length that the posterior end of the body, with 2 small triangular
lobes. Internal shell white, calcified, with marked growth striae, filling the whole posterior half
of the body (Figure 2 C). Shoulder wide. Small lobe of the shell extended over the posterior
region of the body. Protoconch big, globose, with a crest, partially embedded in and rein-forced
by the teleoconch, which do not close the aperture.
Remarks: This is a gregarious species that was initially misidentified and mixed Runcina
prieta Ortea, Moro & Espinosa, 2007. The calcified internal shell of Chelidonura pusilla
Ortea, Moro & Espinosa, new species, is well developed and very complex considering the
size (1-1.2 mm) (Figures 2 B and 3). Together with the small size of the body, the propor-tions
and the shape of the shell distinguish C. pusilla from all the other members of the
genus.
ORTEA et al. (2012) show the most of the shells of the species included in this clade,
one of them strongly calcified (C. juancarlosi).
Chelidonura quadrata Ortea, Caballer & Espinosa new species
(Figures 4-5)
Holotvpe: 3.5 mm long alive, deposited in IES. Type locality: Playa Pilar, Cayo Guillermo, Cuba, sandy
bottom, 3 m depth.
Etimoiogy: Quadrata, in Latin squared, by the quadrangular appearance of the shell.
Description: Body dark greyish brown, with some disperse opaque white spots and notice-able
brilliant blue patches on the edge of the parapodia, on the sides of the front edge of the
head and on the posterior lobes of the body. Posterior edge of the cephalic shield white, some-what
depressed in the middle, with the same proportions than the posterior region of the body.
Posterior region of the body topped in two lobes; the left one sharper and much bigger than
the right one (Figures 4 A).
Internal shell amber, slightly calcified, fragile, quadrangular (Figures 4 C and 5 C),
occupying 1/3 of the posterior region of the body (Figure 4 D). Protoconch with a crest (Fig-ures
4 E and 5 C-E) that is prolonged ventrally (Figure 5 D) and partially covers the inner side.
Teleoconch shows mild growth lines and a wide shoulder (or “wing”) (Figure 5 F), laid on the
posterior region of the body.
90
Remarks: Based on its external characters this species is somewhat related to Aglaja unsa
Marcus & Marcus, 1969, from Brazil, but the shell ofA unsa has a wrinkled shoulder (MAR-CUS,
1970: pt. 1, tig. 2). These wrinkles are present even in the protoconch, which lacks the
crest present in Chelidonura quadrata Ortea, Caballer & Espinosa, new species. In addition,
the length ot the cephalic shield in A. unsa is twice as big as the posterior region of the body,
ditferent from that of C. quadrata.
Within the species of the clade, C. quadrata shall be compared with these that bear a
non-calcified shell. The internal shell of C. africana bears a protococh with a crest prolonged
ventrally, but the external morphology and coloration of the animal is different and the shell
is stronger, bigger, with different shape and proportions. C. mariagordae bears certain exter-nal
resemblance, but lacks the ventral prolongation of the crest and the proportions of the
shell and the shoulder (“wing”) are quite different.
Internal shells of the remaining genus/clades within the Aglajidae
This comparison is based on the best information available (Table 1).
Odontoglaja Rudman, 1978
Type species: Odontoglaja guamensis Rudman, 1978
Shell: (Figure 6) Oval, flattened, calcified, thick and strongly calcified, with a broad, elongate
and curved wing in the upper edge. Teleoconch rounded, concave-flattened, at least twice as
long as the protoconch, usually reinforced on the edge. Protoconch big, usually covered by the
callus, which strongly joins it with the teleoconch, lacking spines or other processes.
Remarks: It has a radula.
Melanochlamys Cheeseman, 1881
Type species: Melanochlamys cylindrica Cheeseman, 1881
Shell: (Figure 6, A. 1 ) Rolled, opencoiled, inflated, rectangular with rounded edges, strongly cal-cified,
with a broad, short and curved wing in the upper edge. Teleoconch remarkedly concave,
long and wide, with an extremely thin and fragile margin, remarkable growth striae and a spiral
central groove, reinforced in the early whorl. Margin usually shows an indentation. Protoconch
proportionally small, lacking spines or other processes. The callus does not join the protoconch
with the teleoconch, so, because of the rolled shape of the shell, it looks like being separated.
Remarks: It lacks a radula. It shows certain resemblance with Gen 4, that may be solved when
new specimens and better information on the shell anatomy of its species are available.
Aglaja Renier, 1807 (Figure 6)
Type species: Aglaja tricolorata Renier, 1807 (Figure 1 H)
Shell: (Figure 6, A.2. 1 .2: part) Opencoiled, rolled, inflated, slightly calcified, whitish, with a
broad, elongate and curved wing in the upper edge. Teleoconch concave, very short, practi-cally
reduced to the “wing”, prolongued in a very short and inconspicuous anterior membra-nose
plate, with a change of concavity and some growth striae. Protoconch vestigial, continued
in a dextral haliotiform shell with remarkable growth striae (Figure 1 H).
Remarks: It lacks a radula.
91
Navanax Pilsbry, 1895 (Figure 6)
Type species: Strategus inermis Cooper, 1862
Shell: (Figure 6, A. 2. 1.2: part) Rolled, flattened, elongated, partially calcified, with a narrow,
long and curved calcified wing in the upper edge. Teleoconch membranose, translucend, non
calcified, fragile and easy to split, long (see the two stages in Figure 6), with a slight groove
at the end of the last whorl. Protoconch vestigial, white, calcified, covered by the callus.
Remarks: It lacks a radula. It shows certain resemblance with Gen 2 and Gen 5, that may be
solved if new specimens and better information on the shell anatomy of the species is avail-able.
Philinopsis Pease, 1860
Type species: Philinopsis speciosa Pease, 1860
Shell: (Figure 6, B.l: part) Rolled, moderatflatened, strongly calcified, rounded to quasi-tri-angular
with rounded edges, covered by some kind of vitreous bamish, with a broad, long
and curved wing in the upper edge. Teleoconch concave, elongated and narrow, with growth
striae and a spiral slight central groove. Margin becomes progressively thinner from the early
whorl, but it doesn’t look like fragile. Margin do not shows an indentation or it is very slight.
Protoconch small, lacking spines or other processes, covered by a callus. The callus does not
join the protoconch with the teleoconch, it looks like being separated.
Remarks: It lacks a radula. Head lacking sensorial bristles (ORTEA et al., 2007). This shell
is certainly very similar to that of Melanochlamys. It shows certain resemblance with Gen 4,
that may be solved when new specimens and better information on the shell anatomy of its
species are available.
Spinoaglaja Ortea, Moro & Espinosa, 2007
Type species: Chelidonura petra Marcus, 1976.
Shell: (Figure 6, B.l
:
part) Oval, scoop-shaped, completely calcified, with growth lines and
a broad, long and curved wing in the upper edge that widely surpass the protoconch. Teleo-conch
long, rounded, with a solid edge. Protoconch vestigial, covered by a callus with two
spines pointing backwards into the left lobe of the mantle.
Remarks: It lacks a radula. Head lacking sensorial bristles. Caudal lobes equal (ORTEA et
al., 2007). Includes 4 species (ORTEA et al. 2012; ORTEA et al. 2013). Only “
Spinoaglaja
petra” (see Table I and Discussion), was considered in the last phylogeny of the family by CA-MACHO-
GARCIA et al. (2014), who, despite of the differences in the shells, considered
Spinoaglaja synonymous to Philinopsis given that they were close in the phylogeny. Indeed,
these two genera are monophyletic only if other two terminal clades are included in the group
(Gen 4 and Gen 5) (CAMACHO-GARCIA etal., 2014). These two clades bear a different kind
of shell than the rest. Given the separation of the 4 clades based on molecular evidences and
the existence of a synapomorphy (as a different shell is) we propose the revalidation of the
genus Spinoaglaja.
92
Nakamigawaia Kuroda & Habe, 1961
Type species: Nakamigawaia spiralis Kuroda & Habe, 1961 (Figure 1 I-J)
Shell: (Figure 6) Planispirally opencoiled, solid, very calcified on its posterior region, mem-branous
in the anterior region, with three whorls and a longitudinal groove. Teleoconch plate
absent. Protoconch absent or vestigial, prolongued with a wing in the upper edge. That wing
shows oblique growth striae and gets wider as it grows and rolls up.
Remarks: It lacks a radula. Head lacking sensorial bristles.
Pseudophiline Habe, 1976
Type species: Pseudophiline hayashii Habe, 1976
Shell: (Figure 6) Quasi-spherical, scoop-shaped, completely calcified, and a broad, short and
rounded wing in the upper edge. Teleoconch very wide, rounded, lacking a groove or an in-dentation,
with a solid edge. Protoconch vestigial, covered by a callus, smooth.
Remarks: It has a radula (KITAO & HABE, 1982). The systematic position of this genus re-mains
to be confirmed.
Noalda Iredale, 1936
Type species: Hydatina exigua Hedley, 1912
Shell: (Figure 6) Bulloid, thin, truncate above, partly external, white with a yellow spiral band
with wide reddish-brown edges (BURN & THOMPSON, 1998). Teleoconch rounded, not
opencoiled, with slight growth striae.
Remarks: It lacks a radula. The systematic position of this genus remains to be confirmed.
4. DISCUSSION
As discussed, the phylogeny tackled by CAMACHO-GARCIA et cil. (2014) incurs in
several assumptions that lead to a divergent approach in the relative position of some genera
within the Aglajidae. This work also includes some Caribbean species, whose taxonomy is dis-cussed
herein, that could affect their conclusions:
1.
- Chelidonura normcini, from Bahamas, synonymized with the Cuban species C.
mariagordae Ortea, Moro & Espinosa, 2004 by ORTEA et al. (2012), because of
their identical internal shell, protoconch and coloration.
2.
- Philinopsis petra (Marcus, 1976) described from Brazil and included originally in
the genus Chelidonura. The distribution of this species reaches Guadeloupe
(ORTEA, ESPINOSA, CABALLER & BUSKE, 2012) but not Bahamas (the ori-gin
of the specimens used by CAMACHO-GARCIA et al., 2014), where there are
two different species transferred to the genus Spinoaglaja by ORTEA, MORO &
BACALLADO (2013) (see Table 1).
3.
- Philinopsis pusa (Marcus & Marcus, 1966) was originally described as a member
ofAglaja, it bears a calcified and squamous internal shell and it has never been re-
93
captured. The shell of this species is very different from the fragile and smooth
shell of the species which inhabits Bahamas, P. bagciensis Ortea, Moro & Espinosa,
2007, whose color variations have been studied VALDES etal. (2013) based solely
in specimens from Stocking Island (Bahamas).
The shell has been the base for the taxonomy of mollusks (PONDER & LINDBERG,
1992; FURUHASHI et al. 2009) since LINNAEUS (1758) published the 10th edition of Sys-tema
Naturae and the zoological nomenclature started in 1758 (ICZN, 1999: article 3), but
with the evolution of malacology as a science, new and diverse characters have been consid-ered
to establish the classification in different hierarchical levels (BOUCHET & ROCROI,
2005). The advent of the molecular biology applied to taxonomy has brought many benefits
to the classification of Gastropods (i.e. GRANDE et al. 2008, WILLIAMS et al. 2010,
BOUCHET et al. 2011), but the shell, when exists, remains as a good character to distinguish
taxa in mollusks.
In aglajids, the shell has been considered to have taxonomic value by some authors
(RUDMAN, 1974, ORNELAS-GATDULA et al. 2011), refused by others (GOSLINER,
1980) or proposed as a main anatomic character for the separation of genus and species
(ORTEA et al. 2012). The delicate internal shells in the Aglajidae are difficult to extract (in-tact)
and are easily damaged by fixation in formalin or bouin (very common in the pre-mol-ecular
period). This may be the reason why they were abandoned as a useful character
(ORTEA etal. 2012).
If the shell is really a valid character to separate genera within the family Aglajidae,
each genus should have a typical kind of shell. A visual and gros comparison of the shells of
the different clades found by CAMACHO-GARCIA et al. (2014) is shown in Figure 6. It is
based in several assumptions that may contain some degree of error (see Materials and meth-ods),
the main of them is the specific determination given by CAMACHO-GARCIA et al.
(2014) to their specimens. Anyway, no species was found with a different kind of shell than
the remaining in the shame clade (see Material and methods: point 4). Consistently, the most
of the previously established genera, and the one described in this paper, have a typical shell
that allows alone to distinguish it from the rest of the members of the family. Only
Melanochlamys and Philinopsis show shells that are difficult to distinguish between them,
but we assume that other sinapomorphies must exist in their anatomy. The same is applicable
to the unnamed clades (Gen 1?, Gen 2-5) with shells resembling those of other groups (Gen
2-Gen 5 -lNavanax\ Melanochlamys-Philinopsis-GQn 4; Chelidonura-?Gen 3). However, for
each monophyletic clade (Figure 6) we observe terminal nodes with different kinds of shells.
Contemporary papers such as the one owed to GOSLINER (201 1 : Figures 2A-B), show
the typical calcified shoulders and the smooth protoconchs in the shells of Philinopsis falci-phallus
Gosliner, 201 1 (Gen 2) and Philinopsis coronata Gosliner, 201 1 (Gen 5), indeed mem-bers
of different clades that we are not able to distinguish based only of their shells. In the other
hand, this paper also shows the shells of Chelidonura mandroroa Gosliner, 201 1 (Gen 3) and
Chelidonura alisonae Gosliner, 201 1 (true Chelidonura), that we are now able to distinguish;
the first one with an extended wing and the second one with this structure reduced, not sur-passing
the protoconch.
CAMACHO et al. (2014) discussed the possibility of a polyphyletic origin for the
genus Chelidonura. On the contrary, we think that there are still many genera to be described
in the family Aglajidae, with sinapomorphies that remain to be observed. This would be con-sistent
with the idea of a Chelidonura-Odontoaglaja shell-type for an ancestral aglajid, with
94
only a certain number ot paths to evolve from this shape. In such case Noalda and Pseudophi-line
would not be members of the family.
In synthesis, with the information available, the shell alone is not enough to distin-guish
all the different genus/clades within the Aglajidae, but is a good character to identify the
members ot the most ot the described genera. Knowing that all the members of the same clade
share a typical shell, better accounts on the characters of the specimens/species/genus are nec-essary
to find new synapomorphies that will allow, together with the shell, to distinguish all
the genera in the tamily. Some of these characters have been already identified: the radula
and the sensorial wristles. Further studies including Nakamigawaia, Nocdda and Pseudophi-line
are advisable to complete the knowledge on the Aglajidae.
5. ACKNOWLEDGMENTS
To our collegue Andrea Zamora (University of Bergen), for the cession of the images
of some shells (see Table 1 ).
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Figure 1.- Migaya felis (Marcus & Marcus, 1970) new comb, from the Caribbean (A - G). A. Dorsal
view of the animal. B. Lateral view. C. Right and left side of the internal shell. D - E. Right side of the
shell. F. Scheme of the sensorial bristles on the head. G. Sensorial bristles on the head. H. Internal shell
of Aglaja tricolorata from the Canaries. Nakamigawaia spiralis Kuroda & Habe in Habe, 1961 from
Japan (I - J). I. Scheme of the internal shell adapted from SASAKI (2008). J, Iconotype (KURODA &
HABE IN HABE, 1961).
JU 250 pm
500 pm
99
Figure 2.- Chelidonura pusilla Ortea, Moro & Espinosa new species, from Cuba: A. Shell in the fixed
specimen. B. Shell in dorsal view. C. Shell in the posterior end of the specimen.
Figure 3.- Chelidonura pusilla Ortea, Moro & Espinosa new species, from Cuba, Shell: A. In dorsal
view. B. In ventral view after the extraction. C. In dorso-lateral view. D. In dorso-lateral view.
100
Figure 4.- Chelidonura quadrata Ortea, Caballer & Espinosa new species: A. Scheme of the Animal in
dorsal view. B. Fixed specimen in dorsal view. C-E. Shell in dorsal view during the process of extraction.
101
A
Figure 5.- Chelidonura quadrata Ortea, Cabal ler & Espinosa new species: A. Lateral view of the ani-mal.
B. Snout of the animal relaxed and in motion. C. Shell in dorsal view. D. Shell in ventral view. C.
Shell in lateral view. C. Shell in lateral view.
102
- Radula |
Aglaja ocellgera
A « . - Navanax: > Navanax
• C. . T . c.
| oolyafrhos * Navanax aengmalicus +
I Navanax gemmatus
Gen 1?: Chehdonura sp ?
Philinopsis: -*• Philinopsis
cyanea * Philinopsis bagaenas (as Philinopsis pusa )
*
Philinopsis dapicta * 'Philinopsis speciosa-gardinen'
Spinoaglaja: Spmoagla/a aea (as P petra)
Philinopsis onentahs
Gen 4: Philinopsis gardmen * Phitnopas pilsbryi
Philinopsis reticulata * Philinopsis Imeolata
Gen 5: Philinopsis coronata * Philinopsis sp
I
Aglejidae sp 1 A^aiidae sp 2 ?
Migaya n. gen. : n comb
Migaya sp Pacific 'Melanochlamys sp
* Anthes ec al (2008)
I
Melanochlamys:
Melanochlamys forrame * Melanochlamys diomedea -
Melanochlamys sp. 1
Chelldonura:
Chetidonura livida * Chelldonura vanans 7
pA.2.1.1
Gen 2: Philinopsis faiciphailus *
Ag/aja regiscorona
A.2.2
A.2.3
|
Chetdonura electra * Chelldonura cubana *
Chetidonura atisonae * Chelldonura amoena *
Chehdonura flavolobala * Chehdonura punctata
* Chehdonura amoena * Chehdonura inornate
Gen 3: Chehdonura mandroroa * Chehdonura inornate *
Chehdonura sp * Chehdonura sandrana *
Chehdonura tsurungensis * Chehdonura pallida
"Crested” Chelldonura: Chehdonura managordae
(as Chehdonura nomnani) + Chehdonura afncana *
Chehdonura berohna 77
^ a .
f *3
Nakam/gawaia:
Pseudophillne:
Noalda
:
jr
Figure 6.- Comparison of the shells within the different clades in the Aglajidae based on the molecular
phylogeny of the family (CAMACHO et al., 2014).
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111
Name
Name
given
by
Camacho-Garcia
et
al.
(2014)
Procedence
(Camacho-Garcia
et
al.,
2014)
Origin
ot
data
Aglaja
tricolorata
Renier,1807
Aglaja
tricolorata
Renier,1807
Italy:
Giglio
Canary
islands,
this
paper,
Pruvot-Fol
(1954),
Martinez,
Ballesteros,
Avila,
Dantart
&
Cimino
(1993)
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