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A new species of cryptobranch dorid nudibranch is described from the Western Indian Ocean. Doris ananas sp. nov. has previously been recorded from South Africa to Tanzania. The studied specimens were from the sub-tropical waters of southern Mozambique, from 10 to 40m deep and are frequently associated with yellow sponges. The new species is characterized by having several conical simple small tubercles surrounding each large tubercle. The integument is yellow and the top of the tubercles are pigmented in brown or black. This species distinguishes from other described taxa on both external and internal characteristics explored herein, mainly in the reproductive system, with the presence of two bags with internal spines annexed to the vagina. This conformation is described for the first time for cryptobranchs.
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Journal of ConChology (2016), Vol.42, no.4 205
A NEW LARGE AND COMMON SPECIES OF DORIS
(GASTROPODA, NUDIBRANCHIA) FROM THE WESTERN
INDIAN OCEAN
PatriCia oristanio V. lima13, yara tibiriçá2 & luiz riCardo l. simone3
1Instituto de Biociências da Universidade de São Paulo, Cx. Postal 42494: CEP04299–970 São Paulo, SP, Brazil
2University of Cádiz & SeaLife Research, Zavora Marine Lab., Inharrime, 1012, Praia de Zavora, Mozambique
3Museu de Zoologia da Universidade de São Paulo, Cx. Postal 42494: CEP04299–970 São Paulo, SP, Brazil
Abstract A new species of cryptobranch dorid nudibranch is described from the Western Indian Ocean. Doris ananas sp.
nov. has previously been recorded from South Africa to Tanzania. The studied specimens were from the sub- tropical waters of
southern Mozambique, from 10 to 40m deep and are frequently associated with yellow sponges. The new species is character-
ized by having several conical simple small tubercles surrounding each large tubercle. The integument is yellow and the top
of the tubercles are pigmented in brown or black. This species distinguishes from other described taxa on both external and
internal characteristics explored herein, mainly in the reproductive system, with the presence of two bags with internal spines
annexed to the vagina. This conformation is described for the first time for cryptobranchs.
Key words Doris, Dorid, Nudibranchs, Mozambique, Western Indian Ocean
IntroductIon
The nudibranchs of the Western Indian Ocean
are relatively poorly understood. Many species,
even common and large ones, still remain unde-
scribed or misidentified. The majority of nudi-
branch research in the region has been conducted
in Tanzania (Eliot, 1902, 1903, 1904a–c, 1905), and
South Africa (Barnard, 1927; Bergh, 1907; Fahey
& Gosliner, 1999; Gosliner, 1987, 1994; MacNae,
1971), leaving a gap in Mozambique despite a
few general marine invertebrate assessments
(MacNae & Kalk, 1958, 1962; Martens, 1879).
The genus Doris is characterized by the dor-
sum covered by simple, rounded tubercles, stiff-
ened by integumentary spicules, which do not
protrude from the dorsal surface. A head with
two lateral prolongations. An anterior border of
the foot grooved but not notched. A labial cuticle
lacking rodlets. A radula composed of simple,
hammate teeth, and outermost teeth being sim-
ple or denticulate. A reproductive system with
tubular, granular and simple prostate; penis and
vagina devoid of hooks; and vestibular or acces-
sory glands absent (Valdés, 2002).
The genus was described based on Doris ver-
rucosa Linnaeus, 1758 from the Atlantic Ocean,
the type species. Presently, it consists of 44 spe-
cies of which 20 are from the Indo- Pacific region,
six being unidentified species (Coleman, 2008;
Bouchet & Gofas, 2014). Samples of the here
introduced species, Doris ananas sp. nov., had
been found in the Western Indian Ocean, par-
ticularly South Africa, Tanzania (Gosliner, 1987;
Gosliner et al., 2011) and Mozambique (pre-
sent study). Additional records in Madagascar,
Mayotte and Seychelles Islands are found on- line
(www.seaslugs.free.fr). The description includes
anatomical information, which has been used
in an ongoing wider comparative, phylogenetic
study on the doridaceans.
MaterIal and Methods
The examined material was hand- picked dur-
ing SCUBA dives in Zavora and Ponta do Ouro,
Southern Mozambique. The material was first
stored at the A.C.C.M. – Zavora Marine Lab.
and is currently deposited at the Museum of
Kwazulu- Natal, Museu de Zoologia de São Paulo
and Museu de História Natural de Maputo.
After collection, all specimens were individu-
ally photographed and notes were taken with all
the data being entered in a database. The speci-
mens were either relaxed in magnesium chloride
7% solution or by freezing. Shortly after this,
they were transferred to formalin 4% (3 speci-
mens), ethanol 70% (1 specimen) or ethanol 96%
(5 specimens). Their dissections were performed
under a stereomicroscope using standard tech-
niques, with the specimens immersed in fixative.
Contact author : patylima84@gmail.com
PoV lima, y tibiriça & lrl simone
206
Digital photos of each step of the dissection were
obtained, as well as drawings aided by a camera
lucida. The radula was removed and placed in
10% sodium hydroxide in order to isolate it from
the soft tissue. A scanning electron microscope
(SEM) was employed to view details of the rad-
ula in the Laboratório de Microscopia Eletrônica
de Museu de Zoologia da Universidade de São
Paulo.
The following abbreviations are used in the
figures: am: ampulla; au: auricle; at: aortic trunk;
bc: bursa copulatrix; bg: blood gland; bm: buccal
mass; bs: buccal sphincter; cb: buccal commis-
sure; ce: cerebral ganglia; cg: connective buccal
ganglia; cp: pedal commissure; cu: caecum; dd:
duct of digestive gland; dg: digestive gland; es:
esophagus; ey: eye; fg: female gland; ft: foot; gb:
buccal ganglia; gc: gill circle; gf: gill filament; gg:
gastroesophageal ganglia; gp: pedal ganglia; gr:
rhinophoral ganglia; hd: hermaphrodite duct; in:
intestine; mo: mouth; m2 – m10: odontophore
muscles; mt: oral tube muscle; oc: odontophore
cartilage; od: odontophore; ot: oral tube; ov: ovi-
duct; pa: papilla; pc: pericardium; pe: penis; pl:
pleural ganglia; pr: prostate; ra: radula; rc: renal
chamber; ri: rhinophore; rm: retractor muscle gill;
rp: reproductive system; rs: radular sac; rv: renal
vesicle; sg: salivary gland. st: stomach; sn: nerv-
ous system; sr: seminal receptacle; ud: uterine
duct; va: vagina; vd: vas deferent; ve: ventricle;
vp: vaginal pouches; vv: auricoventricular valve.
Institutional Abbreviations
KZN – Museum of Kwazulu- Natal, South Africa.
MHN – Museu de História Natural de Maputo,
Mozambique.
MZSP – Museu de Zoologia da Universidade de
São Paulo, São Paulo.
systeMatIcs
Family Doridae Rafinesque, 1815
Doris Linnaeus, 1758
Type species Doris verrucosa Linnaeus, 1758
Doris ananas sp. nov.
(Figs 1–7)
Holotype MOZAMBIQUE, Inhambane, In -
harrime, Zavora Beach, 24°31'S 35°12'E (Y. Tibiriça
col., 06/v/2010, 30m depth), MZSP 111010.
Paratypes MOZAMBIQUE, Inhambane, In -
harrime, Zavora Beach (Y. Tibiriçá col.), MZSP
109879, 1 ex. (10/iii/2011, 12m depth); MZSP
109880, 1 ex. (10/iii/2011, 12m depth); MZSP
109882, 1 ex. (14/iii/2012, 18m depth); MZSP
109884, 1 ex. (10/viii/2012, 17m depth); MZSP
109885, 1 ex. (15/viii/2012, 17m depth); MZSP
109887, 1 ex. (06/iv/2010, 14m depth); NMSA
L9730/T4025, 1 ex. (14/III/2011, 16m depth);
NMSA L9731/T4026, 1 ex. (09/VIII/2011, 18m
depth); NMSA L9732/T4027, 1 ex. (17/III/2013).
Description
External morphology (Figs 1A–D; 3A; D; F)
Size of fixed animal 16- 31- 50mm. Color uniform
yellow with black tubercles. Body oval, elon-
gated, dorsum covered by rounded tubercles,
located mainly in median dorsal region; tubercles
decreasing in size towards edge. Rhinophores
Figure 1 Doris ananas live specimen. A. Dorsal-
slightly right view. B. Ventral view. C. Detail ante-
rior region, dorsal- slightly right view. D. Detail of
extended gill circle, dorsal view.
a new Doris from the western indian oCean 207
with 20–25 transverse lamellae, yellowish base,
2/3 black apex; rhinophoral sheaths with small
tubercles, two larger tubercles on left and right
sides. Gill composed of 6–7 tripinnate gill (Fig.
3F), occupying ~6% of haemocoel volume, yel-
lowish base and black apex, arranged in circu-
lar fashion surrounding anus; branchial sheaths
with small tubercles. Mouth opening in anterior
ventral region between anterior region of notum
and foot. Lateral extensions pore- like, on each
side of mouth (Fig. 3A). Anterior border of foot
grooved, but not notched. Foot yellow with some
small black dots.
Haemocoel organs (Figs 3B–C) Pericardium and
gill circle in posterior half of visceral mass occu-
pying ~15% of haemocoel volume. Buccal mass
located anteriorly, occupying ~25% of haemocoel
volume. Nervous system dorsal to buccal mass,
covered by blood gland, occupying ~5% of
haemocoel volume. Reproductive system on
right side of animal, occupying ~10% of haemo-
coel volume. Stomach on left side, intestine
with small curve at anterior portion, immersed
in haemocoel, both occupying ~15% of haemo-
coel. Digestive gland/gonad occupying ~30% of
haemocoel volume.
Circulatory and excretory systems (Fig. 3B; E–F)
Pericardial cavity dorsal and posterior to diges-
tive gland, anterior to gill circle. Gill retractor
muscle originating from base of gill circle, run-
ning longitudinally up to 1/4 of foot level, insert-
ing into dorsal surface of foot. Auricle funnel-
like (wider anteriorly) with thin walls. Ventricle
slightly taller than wide, with thick muscular
walls. Aortic trunk branched into posterior artery
irrigating stomach, caecum and digestive gland;
anterior artery irrigating reproductive system,
buccal mass, odontophore and nervous system
inserting on blood gland. Blood gland (bg) with
posterior portion three times larger than ante-
rior portion (Fig. 3B). Medial sinus connected to
afferent branchial ring, irrigating entire digestive
gland. Renal vesicle located on right dorsal side
of pericardium, near base of auricle, connected
to inner surface of pericardium (Fig. 3B); renal
Figure 2 Doris ananas, radula in SEM. A. Panoramic
view (MZSP109884). Scale: 300µm. B. Higher magni-
fication in central region (MZSP109880). Scale: 20µm.
C. Outer lateral teeth (MZSP109884). Scale: 20µm. D.
Detail of lateral teeth (MZSP109885). Scale: 30µm.
Figure 3 Anatomical details of Doris ananas. A.
Whole ventral view. Scale: 2mm. B. Dorsal view of
extracted visceral mass. Scale: 1mm. C, Same, ventral
view D. Rhinophore, dorsal view. Scale: 1mm E. Detail
of renal vesicle with the connection with pericardium.
Scale: 1mm. F. Gill circle, dorsal view. Scales: 1mm.
PoV lima, y tibiriça & lrl simone
208
chamber elliptical, color light yellow, with lon-
gitudinal folds, same size as ventricle. Renal
chamber extending from dorsal to medial sinus,
previously connected to renal vesicle, extending
posteriorly to center of gill circle and opening in
nephrostome papilla, next to base of anal papilla.
Digestive system (Figs 4A- B; 5A–D) Oral tube
composed of outer lip, with pleats lengthwise;
inner lip with transverse fold. Main oral tube
muscle, mt, three pairs of retractor muscles of
buccal mass, originating on oral tube, running
dorsally and ventrally to oral tube, inserting on
body side, about six times as wide and twice as
long as m10. Odontophore oval, connected to
oral tube by pair of ventral protractor muscles
(m10); thin longitudinal, dorsal and ventrolat-
eral protractors of oral sphincter, originating in
anterior region of odontophore, inserted in pos-
terior region of integument, close to oral tube
(Fig. 4A). Oral sphincter surrounding chitinous
part of oral tube. Odontophore muscles: m2, pair
of strong retractor muscles of buccal mass, twice
as long as wide, origin on anterior dorsal odon-
tophore, running laterally to m4 and inserted
ventrally on dorsal portion of foot; m4, main
pair of dorsal tensor muscles, strong and broad,
1/3 as long as wide, almost completely cover-
ing cartilage of odontophore, inserted on ventral
portion of subradular membrane; m5, pair of
dorsal auxiliary tensor muscles, twice as long
as wide, originating mostly in posterior region
of odontophore cartilages, covering ~1/3 of
posterior cavity of odontophore, as long as, and
with ~1/3 of m4 width, inserting on ventral side
of subradular membrane, around radular sac;
m6, unpaired horizontal muscle, with transverse
fibers connecting to median surface of left and
right odontophore cartilages, about same length
and half as wide as m4, posterior and anterior
portion about same width as m4 (Fig. 5D); m7,
pair of thin and short muscles, running parallel
to dorsal portion, originated in posterior part of
m6 and inserting on radular sac (Fig. 5B). Pair
of odontophore cartilages elliptical. Subradular
membrane thin, strong, translucent. Radular sac
~1/3 of odontophore length (Fig. 5A). Radular
teeth (Figs 2A–D): rachidian teeth absent; for-
mula 58 × 50.0.50 (in 25mm long specimen). Each
lateral tooth with broad base, tapering towards
apex, hook- shaped, with single terminal cusp;
outermost teeth thinner than internal teeth, inner
base width about half of lateral teeth width, apex
also hook- shaped, with single terminal cusp.
Pair of salivary glands long, tubular (Fig. 4A);
sponge- like, duct inserting in anterior region
of esophagus, extending posteriorly to anterior
region of digestive gland. Esophagus connected
with odontophore, making fold up to nerve ring,
running longitudinally until its connection with
stomach (Fig. 4A). Stomach oval (Fig. 4B), with
folds at entire inner surface; longitudinal pleats
thicker posteriorly, close to anterior region of
intestine. Intestine with longitudinal folds along
Figure 4 Doris ananas details of digestive system. A.
Foregut, dorsal view, some adjacent structures also
shown as in situ. B. Midgut as in situ, dorsal view.
Scales: 1mm.
Figure 5 Doris ananas odontophore anatomy. A.
Whole dorsal view, esophagus removed. B. Whole
ventral view, sphincter removed. C. Dorsal view,
radula removed, each cartilage slightly deflected. D.
Same, m4 and m5 deflected downwards to expose
odontophore cartilage. Scales: 1mm.
a new Doris from the western indian oCean 209
its entire length, diameter about half esophagus
diameter, but more uniform. Caecum as elon-
gated sac, located ventrally to stomach, opening
in anterior portion of stomach, close to esophageal
insertion, ~2/3 the length and ~1/3 the width of
stomach; typically containing dark brown sub-
stance (Fig. 4B). Common opening for esopha-
gus, stomach and caecum located on digestive
gland. Digestive gland dark beige, cone- shaped,
being largest organ of visceral mass, occupying
~30% of its volume; anterior portion about twice
wider than posterior portion, inner face of gland
sponge- like, bearing distinct main duct and vari-
ous secondary ducts. Anus opening into anal
papilla at center of gill circle, ~1/4 of gill fila-
ment length (Fig. 3F).
Genital system (Fig. 6A–E) Located between
buccal mass and digestive gland, mainly on
right- dorsal side. Gonad immersed into diges-
tive gland, difficult to distinguish between them.
Hermaphrodite duct thin, long. Ampulla located
on female gland, elongated and tubular. Prostate
tubular, granular, 2/3 of length of ampulla, nar-
rowing in vas deferens, with about half of length
of prostate, expanding up to penis (Fig. 6A).
Penis’ muscle absent. Penis muscular, cylindrical
and elongated, about half of length of prostate.
Female gland well- developed, rounded, occu-
pying ~20% of reproductive system volume,
divided into mucus gland (~2/3 of female gland,
color beige), and albumen gland (~1/3 of anterior
most region, dilated, irregularly shaped, color
dark brown) (Fig. 6E). Oviduct occupying ~1/4
of female gland volume. Uterine duct thin, rela-
tively short, length ~1/6 of vagina length, located
at base of vagina, inserted in female gland near
oviduct. Seminal receptacle rounded, as large as
bursa copulatrix, length ~1/3 of vagina length,
connected to vagina near uterine duct through
short stalk. Bursa copulatrix rounded, length
~1/3 of vagina length, connected to vagina after
seminal receptacle, also through short stalk.
Vagina cylindrical, elongated, approximately
~twice wider than penis, followed ventrally by
prostate and located parallel to penis in genital
opening. Pair of pyriform vaginal pouches ~1/2
of bursa copulatrix, one on each side of vaginal
opening, internally with white mass and chi-
tinous spine (Fig. 6C). Gonopore on right side,
anterior quarter of length of animal from head,
located between foot and notum.
Central nervous system (Fig. 3B; 4A; 7A–C)
Located dorsally to odontophore, mostly cov-
ered by blood gland. Pair of cerebral and pleural
ganglia fused with one another. Pedal ganglia
fused with cerebral and pleural ventrally, but
not fused among themselves. Pedal commissure
simple, surrounding esophagus and salivary
glands, same length as fused ganglia (cerebral,
pleural and pedal) (Fig. 7A–B). Buccal ganglia
short, located ventrally to odontophore, between
Figure 6 Doris ananas reproductive system. A. Dorsal
whole view, most structures uncoiled. B. Detail of penis
view. C. Detail of vaginal pouches, dorsal view, dor-
sal portion of walls artificially removed to show inner
spines. D. Female gland, dorsal view, uterine duct and
oviduct present. E. Same, ventral view, ampulla and
prostate connections present. Scales: 1mm.
Figure 7 Doris ananas central nervous system. A.
Dorsal view. Scale: 1mm. B. Ventral view. Scale: 1mm.
C. Detail of buccal and gastroesophageal ganglia, ven-
tral view. Scales: 0.5mm.
PoV lima, y tibiriça & lrl simone
210
radular sac and anterior portion of esophagus,
connected to cerebral ganglia through long
and slender connective tissue, united to gastro-
esophageal ganglia by short connective tissue.
Gastro- esophageal ganglia length about ~1/3
of buccal ganglia length, circular (Fig. 7C).
Rhinophoral (olfactory) ganglia bulb- shaped,
connected to anterior portion of cerebral ganglia.
Eyes dorsal, located on cerebral ganglia (Fig. 7A).
Statocysts small and iridescent, located ventrally
to pedal ganglia.
Etymology The name refers to the external simi-
larity with a pineapple, one of the most common
fruits in Mozambique. Ananas is the pineapple
genus and it comes from the Tupi word nanas for
the fruit.
Ecology Subtropical rocky reef, from 15 to 33
meters deep; throughout the whole year, with
higher abundance in June, when the water tem-
perature starts to drop from an average of 26°C
to 24°C. Specimens are usually seen crawling
on the reef, once, a specimen has been observed
embeded in an unidentified yellow sponge with
its gills retracted. Despite its large size, when
embeded in the sponge, the species became
well camouflaged and could be easily missed
by divers. They are relatively common but have
never been seen in great numbers, usually only
one or two individuals at a time.
Distribution Confirmed from Kenya to South
Africa, with possible records in Madagascar,
Mayotte, Tanzania, Indonesia, Queensland and
Marshall Islands (internet sources: http://www.
nudipixel.net/species/doris_sp/; http://seaslugs.
free.fr/; Gosliner, 1987; Gosliner et al., 2011;
present study).
dIscussIon
The external and internal anatomy of the spe-
cies described here is typical of the genus Doris
(Valdés, 2002), with the exception of the two
vaginal pouches, each one with a spine, located
very close to the vaginal opening (Figs 6A, C).
These vaginal pouches have never been recorded
for the genus before.
Because of its originality, the terminology of
vaginal pouches appears to be adequate. In the
literature, there are two other denominations for
genital accessory structures: accessory gland and
vestibular gland (Valdés et al., 2010). Doris ananas
has the vaginal pouches positioned in the vagi-
nal opening, and may have distinct functions
different from those in the literature. Other spe-
cies from different families may also have geni-
tal accessory structures. For example, Geitodoris
pusae (Marcus, 1955) has a single pouch around
the vagina without a spine, considered a vestibu-
lar gland (Marcus, 1955; Alvim & Pimenta, 2014);
and, Platydoris dierythros Fahey & Valdés, 2003
has an accessory gland connected to the penial
side (Fahey & Valdés, 2003).
Two genera of Dorididae have similar struc-
ture in the atrium with the vagina: Goslineria and
Pharodoris, both from the West- Pacific deep water.
Pharodoris has two large glands containing a long,
bifid, rigid spine, however it has a characteristic
elevated branchial sheath (Valdés, 2001) not pre-
sent in Doris ananas. The only species belong to
the genus Goslineria, Goslineria callosa has several
large sacs in the atrium, each containing a long,
simple and flexible spine (Valdés, 2001). A nota-
ble difference between Doris and Goslineria is the
prostate, which is tubular in Doris and flatted in
Goslineria.
Despite having armed appendices in the vagina
in common, D. ananas does not appear to belong
to the genera Goslineria or Pharadoris because of
the quantity of further similarities with D. verru-
cosa and several differences from species of those
genera (summarized in Table 1).
We decided to compare the new species with
the type species Doris verrucosa, because its
recent review provides more complete infor-
mation of the anatomy (Lima & Simone, 2015).
Furthermore, we compare the new species with
the common Indo- Pacific species: Doris granulosa
(Pease, 1860) and Doris immonda Risbec, 1928,
based on literature data (Valdés, 2002).
D. ananas differes from D. granulosa and D.
immonda in lacking lateral prolongations, with
only one pore on each side of the mouth (Fig.
3A). It is interesting to note that D. verrucosa has
the lateral prolongations on the mouth, which
are well developed and with triangle- shaped and
lateral groove on each one, while D. granulosa
and D. immonda present blunt prolongation.
The penis’ muscles of D. ananas are absent,
like in other species of Doris, such as D. granu-
losa and D. immonda (Marcus, 1955; Valdés, 2002;
Camacho- García & Gosliner, 2008), further con-
firming the generic statement.
a new Doris from the western indian oCean 211
The blood gland of D. ananas seems to be
divided with the posterior portion three time
greater than the anterior one (Fig. 3B), as with
D. immonda, while with D. verrucosa the blood
gland is undivided and covers the whole nerv-
ous system. Normally, the species of the fam-
ily Discodorididae have a divided blood gland
(Dayrat, 2010).
Another striking difference of D. ananas is
the presence of a papilla in the nephrostome
(Fig. 3F), which does not appear in D. verrucosa,
D. granulosa and D. immonda. The renal vesicle of
D. ananas is as large as ventricle (Fig. 3B), while
the ratio of vesicle/ventricle is normally ¼ the
size of ventricle in D. verrucosa. The gill filaments
of D. ananas are tripinnate (Fig. 3F), as well as
D. granulosa and D. immonda, and not unipinnate
as in D. verrucosa.
Despite of the geographic distance between
D. ananas with the typo species D. verrucosa, as
well as, anatomic differences with other known
doridids, mainly in the genital organs, a con-
servative approach has been applied here in con-
sidering it in the genus Doris. However, a more
complete taxonomic and molecular revision of
the group might prove different.
acknowledgeMents
We thank The Rufford Foundation for the finan-
cial support to conduct field research. We also
give thanks to Lara Guimarães (Laboratório
de Microscopia Eletrônica do Museu de
Zoologia da USP – MZSP) for helping with the
SEM examination. This work was supported
by Conselho Nacional de Desenvolvimento
Table 1 Some comparative anatomical data among the type species of the genus Doris, Goslineria and
Pharodoris with Doris ananas.
Doris ananas Doris verrucosa Goslineria callosa Pharodoris diaphora
Perfoliate rhinophores 20–25 lamellae 13 lamellae 22 lamellae 30 lamellae
Branchial leaves 6–7 tripinate 15–17 unipinate 7 tripinate 5 bipinate
Branchial sheath Bearing small
tubercles
Eight elongated
tubercles
Somewhat elevated
bearing numerous
tubercles
Very elevated
Oral tentacles Absent Two triangular
prolongation on
each side of the
buccal area
Absent Two triangular
prolongation on each
side of the buccal area
Notum/foot Dorsum margin
wider than foot
Dorsum margin
wider than foot
Dorsum margin as
wide as foot
Dorsum margin
narrower than foot
Radular formula
(animal size)
34 × 43.0.43 (21mm
long)
32 × 40.0.40 (9mm
long)
39 × 42.0.42 (21mm
long)
34 × 39.0.39 (16mm
long)
Blood gland Divided (two) undivided undivided Divided (two)
Color of rhinophores
and gill
Different color
(black) of the
dorsum (yellow);
Same color as the
dorsum
Same color as the
dorsum
Same color as the
dorsum
Outermost lateral teeth Denticles absent Denticles absent Small denticles Small denticles
Prostate shape Tubular and
glandular
Tubular and
glandular
Flattened Tubular
Vaginal pouches Two vaginal
pouches with
spine
Absent Several large sacs,
each containing a
long, simple, flexible
spine
Two large glands, each
containing a long,
bifid, rigid spine
Papilla of nephrostome Present Absent Absent Absent
Pedal commissure Simple Simple Simple With a visible triple
division in connection
with the left pedal
ganglia
Optical ganglia Not pedunculated Not pedunculated Pedunculated Pedunculated
PoV lima, y tibiriça & lrl simone
212
Científico e Tecnológico (CNPq) proc. 159446/
2012–0.
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... With nearly 2500 km of coast, spanning both subtropical and tropical areas, which support complex environments, Mozambique provides an ideal location to study the marine diversity of the western Indian Ocean (Pereira et al. 2014). In spite of this, there are only about 50 species of Nudipleura recorded in the literature from Mozambique (Martens 1879;Macnae & Kalk 1958;Gosliner et al. , 2015Pola et al. 2008;King & Fraser 2014;Lima et al. 2016;Matsuda & Gosliner 2017) The documentation of Mozambican Nudipleura fauna is relevant with regards to wider questions about biogeography and phylogenetics, as these questions can only be answered based on robust species assessment (Briggs & Bowen 2012). Moreover, knowledge of the biodiversity is the first step towards the discovery of products for the benefit of society (Fautin 1988;Flam 1994;Avila et al. 2000;Fusetani 2000; Benkendorff 2010). ...
... Western Indian Ocean. Kenya , South Africa (Gosliner 1987; and Mozambique (Lima et al. 2016). ...
... This species was recently described based on material from Mozambique collected during this study. See Lima et al. (2016) for material examined and full morphological details. ...
Article
Full-text available
This paper provides the first comprehensive annotated and illustrated inventory of Nudipleura from Mozambique. A total of 267 species are recorded, including 61 putative new species, documented over a period of seven years from several localities along the coast. At least 20 species need further investigation through molecular and taxomic analysis. Of the 186 confirmed described species, 118 are new records for the Mozambican fauna. Sampling was carried in tidal reefs and depths up to 60m on the subtropical and tropical coast of Mozambique. The most representative families were Chromodorididae (69 species), Discodorididae (30 species), Facelinidae (23 species) and Phyllididae (16 species). Nevertheless, a vast area of Mozambique remains unexplored, thus it is likely that the species documented here represent only a fraction of the true Nudipleura diversity of the country.
Article
Full-text available
Limacization is the evolutive process that transform a snail into a slug. The modifications and implications of this process are exposed and discussed herein, emphasizing the modification of the visceral mass that migrates to the head-foot haemocoel; the pallial structures also have similar modification or disappear. The limacization is divided in 2 levels: degree 1) with remains of visceral mass and pallial cavity in a dorsal hump; degree 2) lacking any vestige of them, being secondarily bilaterally symmetrical. Despite several gastropod branches suffered the limacization process, the degree 2 is only reached by the Systellommatophora and part of the Nudibranchia (doridaceans). Some related issues are also discussed, such as the position of the mantle in slugs, if the slugs have detorsion (they do not have), the absence of slugs in some main taxa, such as Caenogastropoda and the archaeogastropod grade, and the main branches that have representatives with limacization.
Article
Full-text available
The study of a large collection of cryptobranch dorid nudibranchs from deep waters in New Caledonia and the Philippines revealed the presence of Austrodoris kerguelenensis (Bergh, 1884); 18 new species belonging to the genera Cadlina, Austrodoris, Geitodoris, Discodoris, Peltodoris, Paradoris, Diaulula, Rostanga, Sclerodoris, Baptodoris and Dendrodoris; and two previously undescribed genera, Goslineria and Pharodoris. The anatomy of all these species, including the digestive, reproductive, and nervous system, are studied in detail. All these species are clearly distinguishable from other members of their genera. Most of the species have a pale, simple background coloration, and two of them lack eyes. Both characteristics seem to be adaptations to living in deep waters. Other deep-water Atlantic and Pacific species of dorid nudibranchs have similar adaptations. The two new genera are characterized by the presence of large copulatory spines, numerous flexible spines in Goslineria, and two solid, bifid spines in Pharodoris. No other cryptobranch dorid genera previously described have similar copulatory spines. Some of the species here described belong to genera previously reported from cold or temperate waters, such as Austrodoris, Cadlina and Diaulula. Most of the species belong to genera that are widespread in either cold, temperate or tropical waters (Rostanga, Paradoris, Geitodoris and Baptodoris), and only two belong to exclusively tropical genera (Sclerodoris and Dendrodoris). Vicariant events and vertical dispersal could explain the processes of speciation and the origin of these deep-water species.
Article
The phylogenetic relationships of the cryptobranch dorids are studied based on morphological characters of species belonging to all previously described genera. The phylogenetic hypothesis supports the cryptobranch dorids as a monophyletic group. There are two major clades within the Cryptobranchia: the radula-less dorids (Porostomata), and the radula-bearing dorids (Labiostomata new taxon). Labiostomata consists of those taxa sharing a more recent common ancestor with Actinocyclus than with Mandelia, and includes several monophyletic groups: Actinocyclidae, Chromodorididae, Dorididae and Discodorididae. The traditional group Phanerobranchia is probably paraphyletic. The new classification proposed for the Cryptobranchia addresses concepts of phylogenetic nomenclature, but is in accordance with the rules of the International Code of Zoological Nomenclature. The following genera of cryptobranch dorids are regarded as valid: Doris Linnaeus, 1758, Asteronotus Ehrenberg, 1831, Atagema J. E. Gray, 1850, Jorunna Bergh, 1876, Discodoris Bergh, 1877, Platydoris Bergh, 1877, Thordisa Bergh, 1877, Diaulula Bergh, 1878, Aldisa Bergh, 1878, Rostanga Bergh, 1879, Aphelodoris Bergh, 1879, Halgerda Bergh, 1880, Peltodoris Bergh, 1880, Hoplodoris Bergh, 1880, Paradoris Bergh, 1884, Baptodoris Bergh, 1884, Geitodoris Bergh, 1891, Gargamella Bergh, 1894, Alloiodoris Bergh, 1904, Sclerodoris Eliot, 1904, Otinodoris White, 1948, Taringa Er. Marcus, 1955 , Sebadoris Er. Marcus & Ev. Marcus, 1960, Conualevia Collier & Farmer, 1964, Thorybopus Bouchet, 1977, Goslineria Valdés, 2001, Pharodoris Valdés, 2001, Nophodoris Valdés & Gosliner, 2001. Several genera previously considered as valid are here regarded as synonyms of other names: Doridigitata d’Orbigny, 1839, Doriopsis Pease, 1860, Staurodoris Bergh, 1878, Fracassa Bergh, 1878, Archidoris Bergh, 1878, Anoplodoris Fischer, 1883, Etidoris Ihering, 1886, Phialodoris Bergh, 1889, Montereina MacFarland, 1905, Ctenodoris Eliot, 1907, Carryodoris Vayssière, 1919, Austrodoris Odhner, 1926, Guyonia Risbec, 1928, Erythrodoris Pruvot-Fol, 1933, Neodoris Baba, 1938, Siraius Er. Marcus, 1955, Tayuva Ev. Marcus & Er. Marcus, 1967, Nuvuca Ev. Marcus & Er. Marcus, 1967, Doriorbis Kay & Young, 1969, Pupsikus Er. Marcus & Ev. Marcus, 1970, Percunas Ev. Marcus, 1970, Verrillia Ortea & Ballesteros, 1981 . The genera Artachaea Bergh, 1882, Carminodoris Bergh, 1889 and Homoiodoris Bergh, 1882 have been poorly described and no type material is known to exist. They are regarded as incertae sedis until more material becomes available. The genus names Xenodoris Odhner in Franc, 1968 and Cryptodoris Ostergaard, 1950 are unavailable within the meaning of the Code. Hexabranchus Ehrenberg, 1831 is not a cryptobranch dorid, as suggested by other authors, because of the lack of a retractile gill. Other nomenclatural and taxonomic problems are discussed, and several type species, neotypes and lectotypes are selected. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 136, 535−636.
Nudibranch of Southern Africa. A guide to Opisthobranch Molluscs of Southern Africa. Sea Challengers
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GOSLINER TM 1987 Nudibranch of Southern Africa. A guide to Opisthobranch Molluscs of Southern Africa. Sea Challengers, Jeff Hamann & California Academy of Science, California.
The fauna and flora of sand flats at Inhaca Island
MACNAE W & KALK M 1962 The fauna and flora of sand flats at Inhaca Island, Moçambique. Journal of Animal Ecology 31: 93-128.
South African nudibranch Mollusca, with descriptions of new species, and a note on some specimens from Tristan d'Acunha
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BARNARD KH 1927 South African nudibranch Mollusca, with descriptions of new species, and a note on some specimens from Tristan d'Acunha. Annals of the South African Museum 25: 171-215.
A new species of Platydoris (Mollusca: Nudibranchia) from North-Western Australia
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FAHEY S AND VALDÉS A 2003. A new species of Platydoris (Mollusca: Nudibranchia) from North-Western Australia. In Wells FE, Walker DI & Jones DS (eds) The Marine Flora and Fauna of Dampier, Western Australia. Western Australian Museum, Perth: 395-404.
CNPq) proc. 159446/ 2012–0. references ALVIM & PIMENTA AD 2014 Taxonomic review of the family Discodorididae (Mollusca: Gastropoda: Nudibranchia) from Brazil, with descriptions of two new species
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Científico e Tecnológico (CNPq) proc. 159446/ 2012–0. references ALVIM & PIMENTA AD 2014 Taxonomic review of the family Discodorididae (Mollusca: Gastropoda: Nudibranchia) from Brazil, with descriptions of two new species. Zootaxa 3745 (2): 152–198.
Indo-Pacific Nudibrancchs and Sea Slugs. A field guide to the World's most diverse fauna. Sea Challengers Natural History Books & California Academia of Sciences
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GOSLINER TM BEHRENS DW & VALDÉS A 2011 Indo-Pacific Nudibrancchs and Sea Slugs. A field guide to the World's most diverse fauna. Sea Challengers Natural History Books & California Academia of Sciences, California.
Boletim da Faculdade de Filosofia, Ciencias e Letras
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MARCUS ER 1955 Opisthobranchia from Brazil. Boletim da Faculdade de Filosofia, Ciencias e Letras. Universidade de São Paulo, Zoologia 20: 89–261. (Pls. 1–30).