Zoo1. J . Linn. Soc., 54: 185-212.With 15 figurns
April 1974
A comparison of CheZidonuru. Nuuanaz and
AgZuja with other genera of the Aglajidae
(Opisthobranchia: Gastropoda)
W . B RUDMAN
Department of Zoology. University of Dar es Salaam.
P.O. Box 35064. Dar es Salaam. Tanzania
Accepted for publication August 1973
An account of the anatomy of Chelidonura inomata Baba. C. elecrra R u d m a . “Navanax”
inermis (Cooper) and Aglaja ocelligera Bergh . Navanax Pilsbry is shown to be ajunior synonym
of Aglaja Renier . A detailed comparative study of the anatomy. and in particular. the external
body form. diaphragm. labial glands. alimentary canal. reproductive system. penis structure and
shell. show that the described species of the Aglajidae can be divided into four distinct genera:
Aglaja Renier Chelidonura A . Adams. Philinopsis Pease and Melanochlamys Cheeseman . All
described species. with available anatomical information. are listed; 10 species in Aglaju
17 species in Chelidonura. 12 species in Philinopsis. and 8 species in Melanochlamys .
Twenty-three other species are listed. but insufficient published information is available to
determine their correct generic position .
.
.
CONTENTS
Introduction
. . . . . . . .
Section I . Chelidonura A. Adarns
. .
External features and mantle cavity
Body cavity and alimentary canal
. . .
Reproductivesystem
Nervoussystem
. . . . .
Discussion of Chelidonura
. .
Section 2 . Navanax Pilsbry
. . . .
External features and mantle cavity
Body cavity and alimentary canal
Reproductivesystem
. . .
Nervoussystem
. . . . .
DiscussionofNavanax
. . .
Section 3 Aglqa Renier
. . . .
Section 4 . The genera of the Aglajidae
Aglaja
. . . . . . . .
Chelidonura
. . . . . .
Philinopsis
. . . . . .
Melanochlamys
. . . . .
Features of taxonomic value
.
Achecklist of species
. . .
Material
. . . . . . . . .
References
. . . . . . . .
Abbreviations used in figures
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W. B. RUDMAN
186
INTRODUCTION
This is the third of three papers on the genera of the Aglajidae. As I have
shown in the two earlier papers (Rudman, 1972a,e), most species of this
family have been placed in either Chelidonura A. Adams, 1850 or Aglaja
Renier, 1807. Chelidonura. although easily characterized externally by the long
posterior lobes, has been interpreted by some authors as a group into which all
species with posterior processes could be placed. A mound of sensory bristles
on each side of the mouth has also been considered characteristic of
Chelidonura. but I have found that these peculiar bristles are present in species
of all the genera and is a family, rather than a generic feature.
The result has been the arbitrary definition, by many authors, of the genera
Chelidonura and Aglaja. Published anatomical information on both genera is
very limited and this series of papers is the first attempt at a comparative
anatomical study of the genera of the Aglajidae. In this paper I have included
the results of investigations into the anatomy of Chelidonura inornata Baba,
C. electra Rudman, “Navanax inermis (Cooper) and Aglaja ocelligera Bergh.
As in the previous two papers I will present detailed accounts of the anatomy
of the species studied, and compare this in each case, with published
information on the relevant genus. I have also included in this paper a general
comparative account of the four genera of the Aglajidae.
”
SECTION 1
Chelidonura A. Adams, 1850
The followhg description is based on a study of Chelidonura inornata Baba,
1949 and C. electra Rudman, 1970. The anatomy of the two species is very
similar and any differences will be indicated.
External features and mantle cavity
The body is elongate; the headshield triangular, with the wide anterior end
formed into a rounded lobe at each comer (Fig. 1). The headshield tapers
backwards into a rounded point. The posterior triangular region is a loose flap
lying over the midline of the body. The rounded anterior end of the posterior
shield just reaches the posterior limit of the headshield. Two long tapering
lobes, the left usually longer than the right, extend from the posterior end of
the posterior shield. These lobes are characteristic of the genus and both the
generic name “Chelidonura” meaning dove-tail, and the name of the type
species “hirundinina“, alluding to the swallow, refer to these extensions of the
posterior shield. The parapodia, in both these species, are large, folding over
and partially covering the back of the animal (Rudman, 1970). In C. electra the
anterior end of the head is quite simple, the mouth opening centrally and with
a mound on each side bearing long, retractile, sensory bristles. Below the
mouth is a deep pouch into which a large labial gland opens. The two anterior
corners of the head of C. inornata are, however, unlike C. electra, rolled and
form a funnel on each side, directing water down over the yellow-brown
Hancock’s organs (Fig. 2). These funnels are mobile and in life are projected
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
187
Figure 1. Chelidonuru electra: semi-diagrammatic view of anatomy.
either outwards or upwards. The mounds bearing sensory bristles are much
larger in C.inornata. The eyes are quite close together and are visible through
the dorsal surface of the head.
The posterior shield in the specimens studied: is much fleshier than in the
other genera of the Aglajidae and a thick layer of muscle, interspersed with
blood spaces, lies over the shell cavity. As in Melanochlamys and Philinopsis,
there is a thin ciliated duct running from the shell cavity and opening in the left
posterior quarter of the dorsum. Because of the thick layer above the shell
cavity in C. electra, the duct is very long. At the base of the V, formed by the
lobes of the posterior shield, sits the “fossette glandulaire”. This small gland,
W. B. RUDMAN
188
H .O.
L.GL.V .
-
L
lorn
Figure 2. Chelidonura inornata: left view of head.
also found in Melanochlamys, is described in a separate study of the related
genus Philine (see Rudman, 1972g).
The general organization of the mantle cavity is similar to that of other
aglajid genera. It is, however, relatively small and the ciliated raphae run back
postero-ventrally, the upper raphe ending on the edge of the right lobe of the
posterior shield, and the lower raphe on the edge of the left. These lobes fold
under the shield to meet in the ventral midline. The aglajid “yellow gland” was
not found in C. electra. but a small one was observed in C. inornata.
Body cavity and alimentary canal
A significant feature which distinguishes Chelidonura from Philinopsis and
Melanochlamys is the absence in Chelidonura of a diaphragm separating the
body cavity into two parts (Fig. 14). Loss of this has been accompanied by an
elongation of the body and can be noticed externally in the distinct separation
of the headshield region and the posterior shield. In general, the arrangement of
the viscera is as in the other genera.
The muscular, unarmed, buccal bulb is relatively small in Chelidonura, being
approximately one-fifth of the body length (excluding the long posterior
lobes), whereas it is half the body length in the other genera. From the buccal
bulb the oesophagus quickly widens into the crop. The stomach lies under the
digestive gland and above the mucous gland of the reproductive system. From
the stomach the intestine curves through the digestive gland and opens into the
mantle cavity (Fig. 1).
Reproductive system
The penis of both species has been described before (Rudman, 1970). I t
consists of a small sac containing a wedge-shaped papilla. The sides of the
CHELIDONURA AND OTHER ACLAJID OPISTHOBRANCHS
189
papilla fold under to form a ventral channel which acts as an excurrent sperm
groove.
Serial sections (Fig. 3) of the reproductive system of C. inornutu and
dissections of C. electru show that in both species the reproductive system is
the same. The ovotestis lies diagonally over the digestive gland and from the
ventral surface arises the narrow spermoviduct which quickly enlarges to form
the folded ampullar region which is packed with endogenous sperm. Near the
ALE. G L .
.GL
A
,
1
IOmm
Figure 3. Chelidonura inornuto: A, section of genital gland mass showing opening of oviduct
into albumen gland and capsule gland into common genital chamber; B. section showing sperm
duct opening into common genital chamber and albumen gland opening into capsule gland;
C,view of dissected reproductive system.
190
W. B. RUDMAN
.N.
Figure -1. Chelidonura efectra: left anterior ganglia viewed from outside
genital gland mass the spermoviduct narrows and coils around the albumencapsule gland complex before splitting, one short branch, the sperm duct,
opening into the common genital chamber, the other, the oviduct, opening into
the albumen gland. At h e point of branching is a small caecum which probably
acts as a fertilization chamber (Fig. 3). The histology of the capsule and
albumen glands is the same as described for Mefaizochfamys (see Rudman,
1972a), though the glands differ in form. Unlike Melarzocklarnys where the
tubular albumen gland opens into the tubular capsule gland, in Chelidonura the
two glands, although communicating with one another, are sac-like. As in
Melanocklamys, only the capsule gland opens directly into the common genital
chamber. The mucous gland, consisting of only one arm, lies on the floor of the
body cavity under the digestive gland. A short duct from the common genital
chamber runs to the exogenous sperm sac, a small spherical bulb lying alongside
the albumen gland-capsule gland complex. From just inside the genital opening,
a thin duct runs to the gametolytic sac lying completely enclosed in the
posterior part of the digestive gland (Fig. 15). In both Mefanochlamys and
Pkilinopsis, the gametolytic sac lies outside the digestive gland and at the right
anterior edge of it.
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
191
Nervous system
The following description of the nervous system is based on Chelidonura
electra. As the nervous system is fully illustrated in Figs 4, 5 , 6 and 7, only a
brief description and explanatory notes will be given here.
The cerebral ganglia are closely linked by a short commissure. The pedal
ganglia, close below the respective cerebral ganglia, are joined by a long pedal
commissure. The pleural ganglia are quite distinct, and on the right side, the
supra-oesophageal ganglion is joined to the pleural. As in Melanochlumys, a
long cord from the supra-oesophageal ganglion runs back to innervate the gill
and upper raphe. At the rear end of the body cavity sits the large visceral
ganglion, to which is attached the sub-oesophageal ganglion. A large cord from
the visceral ganglion gives rise to two major branches (Fig. 6B), one supplying
the small right lobe, the “floor” of the mantle cavity, and in particular, the
lower raphe, and the other supplying the large left lobe of the posterior shield.
Figure 4 shows the arrangement of the cerebral nerves. A huge cord gives rise
Figure 5. Chelidonura electra: left anterior ganglia folded out to show inner side,
W. B. RUDMAN
192
J 111111
Figure 6. Chelidonuru elecrm: A, pedal nerves of left anterior end; B, innervation of extensions
of posterior shield.
to branches running to the Hancock’s organs (H.), to the sensory bristles
(S.BR.) and to the anterior lip of the headshield (Cl). Some branches of the
labial nerve (L.N.)run to either side of the mouth, while others (Ll) innervate
the extrinsic musculature of the buccal bulb. A small nerve from the cerebral
ganglion (0T.N.) runs down to the otocyst sitting on the pedal ganglion. The
pedal nerves are homologous to those of Melunochlurnys and Philinopsis and, as
in Philinopsis, P3 and P4 arise from a common cord as do P5 and P6. In
Melunochlamys only P3 and P4 are joined.
Discussion of Chelidonura
Some information is available on the anatomy of C. hirundininu, the type
species of the genus (Bergh, 1900; Risbec, 1951) and some on C. punctufu
Eliot, 1903 (Marcus & Marcus, 1970b). Other published information on species
of Chelidonuru is usually limited to a description of the external body form.
Apart from C. electru and C. inomutu, I have investigated the anatomy of
C. philinopsis Eliot and C.sundrunu Rudman, both from Zanzibar, and
specimens provisionally identified as C. vuriuns Eliot, from Fiji. These species
showed little variation from the anatomical organization of the two described
here in detail (Rudman, 1973).
One feature in C. inomutu, the curling up of the anterior edges of the foot
into funnels, was not present in the other species and appears to be a specific
peculiarity. Another feature is the range of adult size within the genus, and of
individuals within a species. As I have noted elsewhere (Rudman, 1970), adult
specimens of C. inomutu from the Solomon Islands measured up to 65 mm in
length while fully mature specimens from Queensland were approximately
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
193
Figure 7. Chelidonura electra: showing major nerves of remainder of body-dotted sections
represent nerves passing under lateral muscle straps or lying at a shallow distance below body
wall.
one-third this length. A similar observation was made with C. electra. In th,e
small specimens the posterior shield is not fleshy, and as in Melunochlumys, the
shell is just under the dorsal surface. The posterior processes, in the small
specimens, consist only of the epithelial layers, while in the large specimens the
shell is hidden deep below the surface, as described previously, and the
posterior processes are thick and fleshy, and well supplied with blood spaces
and nerves.
The specimens of C. phihopsis and C. sandrana fall into the smaller size
category, and neither the posterior lobes nor the posterior shield are noticeably
fleshy. Another point of difference in external body form is that in these two
latter species the parapodia, although quite large, do not fold over one another
in the midline.
Although over 20 species have been assigned to this genus, my understanding
of it calls for the transfer of some of these to the Nuvanax-Agluju genus group
(which is discussed later). In the final section of this paper there are definitions
and species lists for each of the genera of the Aglajidae.
194
W. B. RUDMAN
SECTION 2
Navanax Pilsbry, 1895
Two species of this genus have been described, both from the west coast of
America: N. inermis (Cooper, 1862) from California and N. aenigmaticus
(Bergh, 1894) from the Bay of Panama (Pilsbry, 1896). A number of references
in the literature describe the colour and external form of N. inermis and also
the shape of the penis (Marcus, 1961; Marcus & Marcus, 1967; MacFarland,
1966), and the interesting work of Paine (1964) shows that it tracks its prey
which consists of Bulla, Haminoea and other opisthobranchs. Although
previous authors have considered Navanax to be a doubtful genus, they could
not be sure because no account of the anatomy existed. The following account
of the anatomy of N. inermis is based on specimens obtained from San Diego,
California, and shows that Navanax should be merged with Aglaja.
External features and mantle cavity
This species can grow up to 180 mm in length (Marcus, 1961). The largest
specimen examined in this study was only 35 mm long. The body is elongate,
the headshield extending half way down the body, the posterior end rounded.
At the anterior end, the edges can be rolled, forming funnels directing water
over the Hancock’s organs (Fig. 10B). The posterior shield begins under the
headshield, and at the posterior end is developed into a pair of thin triangular
lobes, which fold under to form the posterior border of the mantle opening
(Figs 8 and 11). The parapodia are large, folding over and partially enclosing
the body. On either side of the mouth are the mounds bearing the sensory
bristles, quite complex structures (Fig. 10B, S.BRM.), while below the mouth
is the sac into which the large labial gland opens (the cells staining blue in
Mallory and Heidenhain, and light pink in Weigert’s iron haematoxylin and van
Gieson).
The shell is large and flattened, as in Philinopsis, and the upper edge is
strongly calcified, while the rest is very thin. A duct from the shell cavity opens
in the left posterior quarter of the notum, and in the posterior midline, as in
Chelidonura, there is the “fossette glandulaire”. The mantle cavity is similar to
that of Chelidonura but differs in having a compound gill, as in Hydatina physis
(see Rudman, 1972c), rather than the simple plicate form found in Chelidonura, Philinopsis, and Melanoch lamys.
Body cavity and alimentary canal
The most significant difference in the body cavity is the position of the
diaphragm. In Melunochlumys and Philinopsis the diaphragm is vertical,
separating the cavity into two halves, while in Chelidonura it is absent. In
Navanax, however, it forms a sac, hanging from the roof of the body cavity and
enclosing the digestive gland, posterior gut and reproductive system.
The buccal bulb is large (Fig ‘8) and is attached to the body wall by four
pairs of extrinsic muscles. Two long lateral muscles on each side
(Fig. lOD,PR.M.) arise from the posterior end of the buccal bulb and run
CHELZDONURA AND OTHER AGLAJID OPISTHOBRANCHS
195
!
L.M.S
Figure 8. Aglqa inemis (Cooper): dorsal dissection to show organs of the body cavity.
forward through the nerve ring to the respective sides of the mouth. There are
also two pairs of short muscles;both pairs arise in the midline, one dorsally and
the other ventrally, at the anterior end of the buccal bulb. There is a short,
thin-walled oral tube enclosing a pair of large warty lips (Fig. 1OC). A series of
W. B. RUDMAN
196
N.C.
Figure 9 . Aglqa inennis (Cooper): section through head showing sensory bristles (-cut sagitally
just off centre).
colour slides showing the feeding behaviour of Nuvunax inermis were kindly
made available by Professor R. T. Pine. These show that, as in PhiZinopsis
tarongu, the buccal bulb is partially everted and used to envelop the prey
(Rudman, 1972f). The oesophageal crop is large and is attached to the body
wall by a pair of long muscles (Fig. 8). The yellow gland (Y.GL.) is extremely
large, lying alongside the digestive gland and opening into the mantle cavity
alongside the anus.
Reproductive system
Unfortunately, the only specimens available were immature. However, it was
possible to determine the structure of the penis and the basic plan of the
reproductive system.
The prostate has two arms which open at the base of the thin-walled penis
sac. A deep incurrent sperm groove runs from the penial opening on the right
side of the head down along the wall of the penis sac to the prostate gland. A
large muscular flap on the left side of this groove folds over it. The penial
papilla (Fig. 1OA) is bulb shaped with a deep excurrent sperm groove running
from the junction of the incurrent groove and the prostate opening, to the tip
of the papilla.
The genital gland mass is quite different from that of the other aglajid
genera. From the ampullar region of the spermoviduct the duct narrows and
coils around the capsule-albumen gland mass. A large thin-walled caecum opens
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
197
M
Figure 10. ig24a inemis (Cooper): A, penis structure; B, anterior view of head; C, anterior
region of buccal bulb-lateral view; D, buccal bulb-right lateral view.
off the spermoviduct in this region (EX.S.S.) and is possibly the exogenous
sperm sac. Because the capsule-albumen gland complex was undeveloped it was
difficult to interpret the rest of the pattern of ducts. It does appear, however,
that the spermoviduct splits, a branch opening into the gland complex, and the
other, the sperm duct, running forward alongside the mucous gland to the
vestibule at the genital opening. The mucous gland is a large coiled structure
and the albumen-capsulp gland opens into it, some distance from the genital
opening. The gametolytic sac is connected to the genital opening by a long
thin-walled duct.
W. B. RUDMAN
198
G.OF!
-
Figure 11. Aglqa inermis (Cooper): v e n d view of mantle cavity.
S.BR. N.
+
L.N.
O.N.
Figure 12. Agluja inem&(Cooper): showing anterior nerve complex
CHELIDONURA AND OTHER ACLAJID OPISTHOBRANCHS
199
Nervous system
The nervous system is illustrated in Figs 8 and 12, and is similar to that of
Chelidonuru. As in that genus, pedal nerves 3 and 4, and 5 and 6, arise from
single nerve cords.
Discussion of Navanax
A number of authors have doubted the validity of the genus Nuvanax, some
considering it to be a synonym of Chelidonuru and others of Agluju. These
comparisons were made with little understanding of the anatomy of the genera
within the Aglajidae. Bergh (1905) considers Navunux to be- a synonym of
Chelidonuru because both genera have backward extensions of the posterior
shield and well developed mounds on which the sensory bristles sit. Many
features of the anatomy however, show h a t these two genus groups are quite
distinct. Marcus (1961) considers that Nuvanax and Chelidonuru are distinct
because of the smooth penial papilla in Nuvanax which “is absent or warty in
Chelidonuru”. Although these two genera are distinct, Marcus’s argument is
invalid because, as I will show later, the species he ascribes ta Chelidonuru do
not belong there. However, Marcus & Marcus (1970b), on reconsideration,
agree with Bergh in considering Nuvanax to be a synonym of Chelidonuru. I
hope to show in the following sections that Nuvunux is not a synonym of
Chelidonuru but rather a synonym of Aglaju.
SECTION 3
Agluju Renier, 1807
The information available on species definitely belonging to Agluja is
scattered and patchy. Fortunately Vayssiere (1880) described the basic
anatomy of the type species A. tricolorutu (as Doridium membranaceum) and
Guiart (1901) has described the anatomy of the other species of Renier,
A . depictu. I have investigated the anatomy of specimens of A . ocelligeru.
These specimens, from San Juan Is, Puget Sound, ranged to 30 mm in length,
when alive. They were brownish in colour, with yellow specks, and were similar
in body form to Nuvunux inermis, although much smaller. A good illustration
of this species appears in MacFarland (1966; Plate 2, fig.4) although it was
misidentified as Melunochlamys dionzedea (Bergh). Dissections of A . ocelligeru
showed it to be very similar in structure to “Nuvanax” inermis with a large
buccal bulb and with the diaphragm surrounding the viscera. As in “Nuvanax”
inermis the exogenous sperm sac opens off the ovotestis duct, some distance
from the genital vestibule.
The penial apparatus is somewhat similar to that described for Agluja phocue
(Marcus, 1961). The penis sac is typically thin-walled and the prostate gland
extends from its base (Fig. 13). The prostate is a long, folded, glandular tube,
the part nearest the penis sac being tightly folded and enclosed in a thin-walled
envelope. Inside the penis sac, the seminal groove runs from the opening to the
base of the sac, where it meets the opening of the prostate gland. A common
groove runs up the base of the penis, which has small tubercles scattered over
200
W. B. RUDMAN
SD R
Figure 1 3 . Agtqa ocelligera (Bergh): penis StNCNTe. Lower illustration showing base of papilla
folded back to show seminal grooves.
it, and then runs along the conical penial papilla, which in preserved specimens
at least, has annular wrinkles (Fig. 13).
Only small anatomical points, such as the shape of the penis, the presence or
absence of a flagellum on the posterior shield, and of course the colour,
separate A. ocelligeru, A. depictu and A. tricolorutu from one another and from
“Nuvanax ” inemis. Two species described by Marcus & Marcus and attributed
by them to Chelidonuru, in fact belong to Agluja. The good anatomical
information provided by Marcus & Marcus for C. evelinue (1955, 1961,1970a)
and for C. phocue (1961) clearly place both species in the genus Agluju and give
more evidence of the taxonomic value of the features of the reproductive
system in this family. In both species the buccal bulb is large, unlike the
situation in Chelidonuru. and is partially eversible. Also the posterior processes
or “flagella” on the posterior shield are of a different nature to the long
elongate “tails” found in members of Chelidonuru.
One feature of the reproductive system does appear to be variable in the
genus Aglqu, or at least open to some doubt. Because of the immaturity of the
specimens available, I am unsure, in “Nuvanax”, whether the spermoviduct
branches, with one branch going to the female gland mass, the other to the
genital opening, or not. However, in Agluja evelinue, Marcus (1955) shows that
the spermoviduct does indeed split in that species. Similarly, Marcus (1961)
shows that this also occurs in Agluju phocue. However Guiart (1901) described
in Aglaja depictu, a condition in which the spermoviduct remains unbranched,
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
201
opening only into the genital vestibule. Vayssiere (1880) also showed this to be
the case in A . tricolorata. Because the branch between the spermoviduct and
the female gland mass is difficult to see without serial sections, it is obvious
that the two Mediterranean species, A . depicta and A . tricolorata, will need to
be reexamined before the position can be clarified. There is one major point
though, that distinguishes the reproductive system of this genus group from the
other three genera. In this genus alone, the exogenous sperm sac opens off the
spermoviduct whereas in the other three genera the exogenous sperm sac opens
independently into the genital vestibule (Fig. 15).
SECTION 4
The genera of the Aglajidae
In this section I will summarize the information presented in this paper and
the two preceding papers (Rudman, 1972a,e). First there are definitions of
each of the four genera. This will be followed by a discussion on the taxonomic
importance of parts of the anatomy and finally there are species lists for each
genus.
Aglaja Renier, 1807 (Figs 14 and 15)
Type species: AgZaja tricolorata Renier 1807 (Type by subsequent
designation-Suter, 1913)
Synonymy: Aglaja Renier, 1804 (non-binomial)
Doridium Meckel, 1809
Navanax Pilsbry, 1895 new synonym
Body elongate, ovate; posterior edge of headshield rounded, sometimes with
a median crest. Parapodia large, partially enclosing notum. Posterior shield with
or without processes developed from the posterior end. Shell flattened,
internal, slightly calcified. Diaphragm reduced to thin sac enclosing viscera.
Reproductive system: spermoviduct splits, sperm duct running to vestibule
and oviduct opening into albumen-capsule gland complex. (Information
concerning A . tricolorata and A. depicta suggests that in some species the
spermoviduct may not branch into the female gland mass-see preceding
discussion.) Exogenous sperm sac opening off the spermoviduct. Gametolytic
sac opening into vestibule, and mucous gland large, coiled and unbranched.
Alimentary canal: buccal bulb large, capable of partial eversion. Only one
labial gland present; it opens below the mouth and stains blue in Mallory and
Heidenhain.
Chelidonura A. Adams, 1850 (Figs 14 and 15)
Type species: Bulla hirundinina Quoy 8t Gaimard, 1832
(Type by original designation)
Synonymy: Hirundella Gray, 1850
Body elongate, narrow; headshield triangular, the posterior end tapering to a
rounded point. Parapodia large, folding over to enclose notum. Posterior shield
15
202
W. B. RUDhiAN
not eversible
labial gland
2
dia&agm
partially
eve;,
Ible
\
labial giand
type
1
shell cavity
not eversible
Figure 14. Diagrammatic transverse sections showing the general body plan in each of the four
genera of the Agiajidae.
distinct, beginning behind headshield, the posterior edge developed into two
large tapering lobes, the left usually longer than the right. Shell flattened,
internal and fairly strongly calcified. Diaphragm absent.
Reproductive system: spermoviduct divides, sperm duct running to the
vestibule at genital opening and oviduct opening into a sac-like albumen gland
which opens into capsule gland. Capsule gland and ducts from gametolytic sac
and exogenous sperm sac open separately into vestibule. Mucous gland is large
and not divided into two arms.
Alimentary canal: buccal bulb is small, one-fifth of body length, and
muscular,probably incapable of eversion. Only one labial gland; it opens below
the mouth, is large and stains blue in h4allory & Heidenhain.
Phifinopsis Pease, 1860 (Figs 14 and 15)
Type species: Philinopsis speciosa Pease, 1860
(Type designated here by position precedence)
Body elongate, ovate, posterior end of headshield rounded at each corner
with a raised median pointed crest. Parapodia large, partially covering notum;
posterior shield is without posterior lobes. Shell large and flat, internal, inner
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
C
-
/
exogenous
sperm sac
20 3
\
ampullar region
of spermoviduct
Figure 15. The reproductive systems typical of each of the four genera in the Aglajidae. (The
different stippling patterns show homologous organs in each genus.)
edge is strongly calcified and remainder is only thinly calcified. Diaphragm
vertical, separating anterior and posterior parts of body cavity.
Reproductive system: spermoviduct opens directly into the vestibule at
genital opening without branching. Albumen gland opening into capsule gland
which in turn opens into vestibule. Gametolytic sac and exogenous sperm sac
each open into vestibule through narrow ducts. Nucous gland is divided into a
long and short arm.
Alimentary canal: buccal bulb either large, muscular and capable of partial
eversion, or long, narrow and muscular. Labial glands of two types; one'staining
blue in Mallory and Heidenhain, the other not. Only one gland of the
blue-staining type, it opens below the mouth, while there are two of other
type, one opening above, the other below the mouth.
Melanochlamys Cheeseman, 1881 (Figs 14 and 15)
Type Species: Melanochlamys cylindrica Cheeseman, 188 1
(Type by monotypy)
Body usually cylindrical, posterior end of headshield rounded, with a small
indentation in the midline. Posterior shield simple, no processes extending from
204
W.
B. RUDMAN
the posterior end. Shell internal, strongly calcified and often of one and one
half whorls. Diaphragm is vertical, separating anterior and posterior parts of
body cavity.
Reproductive system: spermoviduct branches, testis duct opening into
vestibule at genital opening, while oviduct enters tubular albumen gland, which
in turn opens into tubular capsule gland. The latter opens into vestibule.
Separate ducts communicate with the exogenous sperm sac and gametolytic
sac. Large mucous gland is divided into a long and short arm.
Alimentary canal: buccal bulb large, muscular, rigid, incapable of eversion.
Labial glands of two types, one blue in Mallory and Heidenhain, the other not.
There are two glands of each type, one of each opening above, and one below
the mouth.
Features of taxonomic value
The family Aglajidae can be divided into four distinct genera, easily
recognizable on external body form and construction of the reproductive
system. Other features, such as the form of the shell, the diaphragm in the
body cavity, and the nervous system can be useful. An attempt to divide the
family on penial types (Marcus & Marcus, 1966) is as unsatisfactory as any
other classificatory system based on a single character. Separate discussions on
different parts of the anatomy in relation to their taxonomic importance now
follow.
1. External body form
Melunochlamys and Chelidonuru can be easily distinguished from one
another and from the other two genera. In MeZunochlumys the parapodia are
small, barely reaching onto the dorsal surface. The body is compressed and the
posterior end of the headshield is either straight or has a small median
indentation. In Chelidonuru, the lobed anterior end of the headshield and the
long posterior extensions to the posterior shield make this genus instantly
recognizable. It is somewhat more difficult to separate Aglaja and Philinopsis
from one another, although as a group they are quite distinct from
MelunochZumys and Chelidonuru. Unfortunately I am not familiar with the
body shape of living members of Agluju except from photographs and drawings.
One characterktic feature of Philinopsis is that the median point on the
posterior edge of the headshield stands raised in a crest when the animal is
burrowing. I cannot find any information on whether this occurs in Agluju.
One feature separating Agluju and Philinopsis is that in the latter genus, the
posterior edge of the posterior shield is blunt, with a median dorsal notch,
whereas in Aglqu there is considerable variation with A . tricolorutu having a
small thin “flagellum”, and A . inermis having quite long extensions somewhat
reminiscent of Chelidonuru.
2 . Diaphrugm (Fig. 14)
In more primitive opisthobranchs, such as the Acteonacea, (see Rudman,
1972b, c, d) and Hurninoeu (see Rudman, 1971), the organs of the body are
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
205
contained in two separate regions. The anterior gut, the ganglia of the nervous
system, and in Haminoea, the penis, are found in the anterior haemocoel in the
head-foot of the body. The other organs, the gonad, digestive gland and
reproductive system, occupy the visceral spiral or are associated with the
mantle cavity. In the Aglajidae, in which the visceral spiral is lost, all these
organs are found in the enlarged body cavity.
However, the phylogenetic origin of this body plan can be seen in the
presence of a thin fibrous wall, which in three of the genera, separate the
organs found in the head-foot haemocoel of more primitive forms, from those
organs originally associated with the mantle cavity and visceral spiral. In
Melanochlamys and Philinopsis, this diaphragm sits vertically, running from the
roof to the floor and separating the body into an anterior and posterior body
cavity. In Aglaja it forms a sac suspended from the roof of the body cavity and
encloses the organs normally associated with the visceral spiral. In Chelidonura
the diaphragm is absent.
3 . Labial glands (Fig. 14)
Although these glands are difficult characters to use, from a practical
taxonomic point of view, they are characteristic. The numbering of the two
gland types follows that used in this, and the earlier two papers of this series.
Glands of type 1 stain deep blue in Mallory & Heidenhain stain and light pink
in Weigert’s iron haematoxylin and van Gieson. Glands of type 2 stain only
very lightly in Mallory & Heidenhain and not at all in the second stain.
In Melanochlamys, two glands, one of each type, open above the mouth, and
a similar two glands open below the mouth. In Philinopsis, a gland of each type
opens below the mouth and a gland of type 2 opens above the mouth. In both
Aglaja and Chelidonura, there is only one gland present; it is large, of type 1,
and opens below the mouth.
4. Alimentary canal (Fig.14)
The most significant feature of the alimentary canal, distinguishing each
genus, is the relative size of the buccal bulb, and its mode of operation. In
Melanochlamys it is large, approximately half the body length, it is rigid and
non-eversible, and it is used as a suction pump. In Aglaja, and most species of
Philinopsis, the buccal bulb is large, flexible and partially eversible. As I
described in an earlier paper (Rudman, 1972e), three species of Philinopsis have
a long tubular pharynx. In Chelidonura the buccal bulb is relatively small, and
although no information is available, it is most probably unable to be everted.
5 . Reproductive system (Fig.15)
This is probably the most significant feature of the anatomy, from a
phylogenetic and taxonomic point of view. Even if there were no other parts of
the anatomy which differed from genus to genus, the distinctly different
structure of the reproductive system in each speciesgroup would alone warrant
serious consideration.
In Chelidonura, Melanochlamys and Philinopsis the exogenous sperm sac
206
W. B. RUDMAN
communicates with the genital vestibule by way of a separate duct. In Aglaja
the exogenous sperm sac opens into the hermaphrodite duct. The first three
genera can be separated from one another by the arrangement of the albumen
and capsule glands. In Melanochlumys the hermaphrodite duct branches, a
short sperm duct opening into the genital vestibule. The other branch, the
oviduct, opens into a long tubular albumen gland which in turn opens into a
similarly tubular capsule gland, which itself opens into the genital vestibule. In
Chelidonura the arrangement is somewhat similar, but the albumen and capsule
glands are sac-like rather than long and tubular. In Philinopsis the hermaphrodite duct does not branch and the capsule-albumen gland complex open
separately into the genital vestibule.
Another feature is that in Chelidonuru and Agluja the mucous gland consists
of only one arm while in the other two genera there is a small secondary arm to
the mucous gland.
6 . Penis structure
The penis was used by Marcus 8c Marcus (1966) in an attempt to clarify the
confusion existing over the generic classification of the group. Unfortunately
the use of a single character, which is itself variable within a genus, as defined
here, has only added to the confusion. In Melunochlamys, the structure of the
penis of those species which have been studied is generally constant, with the
exception of the rather specialized penis of M . diomedea (see Rudman, 1972a).
In Philinopsis there appears to be two subgroups, those with a large buccal bulb
having one type of penis and the three with a long narrow pharynx having a
different type (Rudman, 1972e). In Chelidonuru the penial papilla is usually
conical with a seminal groove running up its v.entral side, but there are
exceptions. Not enough information is available about species of Agluju to say
much about the penis structure. It would seem that it is more variable in this
genus than in the other three.
7. Shell
The internal shell can be used to distinguish Chelidonuru and Melunochlamys
from each other and from the Aglaja-Philinopsis group. In Melunochlumys the
shell is completely calcified, it is strong and partly coiled. In the other three
genera the shell is flattened and only lightly calcified (Rudman, 1972e, 1973).
In Agluju and Philinopsis the inner edge is fairly strongly calcified while in
Chelidonuru some species have a brownish tinge to the shell. Apart from the
strongly calcified shell in Melunochlumys this is not a good taxonomic
character either for differentiating species or genera. Also because the shell is
usually fragile, and internal, it is often broken or dissolved in preserved
specimens.
A checklist of species
The following lists, as far as I am able to determine, are complete for the
four genera of the Aglajidae. In each case the genus each species was originally
assigned to is placed in parenthesis after the original author’s name. The
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
207
numbers following each species name, correspond to the numbers 1-7 used in
the preceding section and indicate those characters which were available to me
in determining the correct generic position of each species.
Aglaja Renier, 1807
1. A . uenigmuficu (Bergh, 1894 as Navurchus); (1, 4, 6 , 7 , Bergh, 1894)
2. A . depictu Renier, 1807 (1-7, Guiart, 1901)
= Doridium coriuceum Meckel, 1809
= Acera curnosu Cuvier, 1810
= Eidofheu marmorufa Risso, 1826
3. A . evelinae evelinae (Marcus, 1955 as Chelidonura) (1-7, Marcus,
1955)
4. A . evelinae dicu (Marcus & Marcus, 1970a as Chelidonura); (1, 3, 7,
Marcus & Marcus, 1970a)
5 . A . felis Marcus & Marcus, 1970a; (1, 3, 4, 6, Marcus & Marcus,
1970a)
6. A . hummelincki Marcus & Marcus, 1970a; (1, 3, 4, 6, 7, Marcus &
Marcus, 1970a)
7. A . inermis (Cooper, 1862 as Navurchus); (1-7, Marcus, 1961; this
paper)
8. A . ocelligera (Bergh, 1894 as Doridium); (1-7, this paper)
9. A. phocue (Marcus, 1961 as Chelidonuru); (1,4-7, Marcus, 1961)
10. A. tricolorata Renier, 1807; (1,4, 5 , 7, Vayssiere, 1880)
= Doridium membrunuceum Meckel, 1809
Chelidonuru A. Adams, 1850
1. C. umoenu Bergh, 1905; (1,4-7, Bergh, 1905)
2. C berolina Marcus & Marcus, 1970a; (1,6, Marcus & Marcus, 1970a)
3. C.conformufa Burn, 1966; (1,7, Burn, 1966); may = C. fulvipunctuta
4. C. elecfra Rudman, 1970; (1-7, Rudman, 1970; this paper)
5 . C. elegans Bergh, 1900; (1,4-7, Bergh, 1900)
6. C. fulvipuncfufu Baba, 1938; (1, Baba, 1938; Allan, 1959)
7. C. hirundininu (Quoy & Gaimard, 1832 as Bullu); (1, 3, 4, 6, 7,
Risbec, 1951; Marcus & Marcus, 1970a)
= C. adumsi Angas, 1867
8. C. inornafa Baba, 1949; (1-7, Rudman, 1970; this paper)
9. C obscuru (Bergh, 1901 as Doridium);(1, Bergh, 1901)
10. C. pullidu Risbec, 1951; (1,4,7, Risbec, 1951)
11. C. philinopsis Eliot, 1903; (1-7, Rudman, 1973)
12. C. plebeia Bergh, 1900;(1,4, Bergh, 1900)
13. C. puncfufuEliot, 1903; (1, 3 , 6 , 7 , Marcus & Marcus, 1970b)
14. C. sabina Marcus & Marcus, 1970; (1, 3-7, Marcus & Marcus, 1970b)
15. C. sandruna Rudman, 1973; (1-7, Rudman, 1973)
16. C. fsurugensis Baba & Abe, 1959; (1, Baba & Abe, 1959)
17. C varians Eliot, 1903; (1-7, Rudman, 1973)
= C. velufina Bergh, 1905 in part (Plate 3, fig 5 ) .
208
W. 6. RUDMAN
Philinopsis Pease, 1860
Species marked with an asterisk have a long cylindrical pharynx, atypical of
the genus. Bergh (1908) suggested a new genus Boridium, in a footnote, for
A. velutinu Bergh in which he considered the buccal bulb was absent. As I have
shown, (Rudman, 1972e), A. velutinu (a junior synonym of P. gurdineri), has a
long elongate buccal bulb but in other features is characteristic of Philinopsis.
1. P. ceylonica (White, 1946 as Agluju); (1, 3-7, White, 1946)
2. P. cyanea (Martens, 1879 as Doridium); (1-7, Rudman, 1972e)
= Doridium nigrum Martens, 1879
= Doridium guttutum Martens, 1880
= Doridium murmorutum E. A. Smith, 1884
= Doridium cupense Bergh, 1907
= Agluju iwasai Hirase, 1936
*3. P. gurdineri (Eliot, 1903 as Doridium); (1-7, Rudman, 1973)
= Chelidonuru velutina Bergh, 1905 in part (Plate 3, fig 6)
= Agluju velutinu Bergh, 1908
= Agluju splendidu Risbec, Marcus, 1965: not Risbec, 1951
4. P. gigliolii (Tapparone-Canefri, 1874 as Agluja); (1-7, Rudman,
1972e)
*5. P. lineolutu (H. & A. Adams, 1854 as Agluju); (1, 3-7, Bergh, 1902;
White, 1945)
6. P. minor (Tchang-Si, 1934 as Aglaju); (1, 4-7, Tchang-Si, 1934)
*7. P. pilsbryi (Eliot, 1900 as Doridium); (1-7, Rudman, 1972e)
= Philinopsis nigru Pease, 1860 (nomen oblitum)
= Doridium alboventralis Bergh, 1897 (nomen oblitum)
= Agluja pilsbryi huwuiiensis Pilsbry, 1920
8. P. speciosu Pease, 1860;(1-7, Rudman, 1972e)
9. P. splendidu (Risbec, 1951 as Agluju); (1,4-7, Risbec, 1951)
10. P. turongu (Nlan, 1933 as Agluju); (1-7, Rudman, l968,1972e, f)
= Chelidonuru uureopunctutu Rudman, 1968
11. P. troubridgensis (Verco, 1909 as Aglaju); (1-7, Rudman, 1972e)
12. P. Virgo (Rudman, 1968 as Agluju); (1-7, Rudman, 1968,1972e)
Melunochlamys Cheeseman, 1881
1. M . cylindrica Cheeseman, 1881;(1-7, Rudman, 1972a, f)
2. M. diomedeu (Bergh, 1893 as Doridium); (1-7, Marcus, 1961;
Rudman, 1972a)
3 . M. ezoensis (Baba, 1957 as Agluju); (1, Baba, 1957)
4. M. henri (Burn, 1969 as Agluji); ( 1 , 4 , 6 , 7 , Burn, 1969)
5. M. lorrainue (Rudman, 1968 as Agluju); (1-7, Rudman, 1968; 1972a)
6. M. nunu (Steinberg & Jones, 1960 as Agluju); (1, 4, 7, Steinberg &
Jones, 1960)
7. M. queritor (Bum, 1958 as Agluju); (1-7, Rudman, 1972a)
8. M . seurati (Vayssiere, 1926 as Doridium); (1, 7, Vayssiere, 1926)
CHELIDONURA AND OTHER AGLAJID OPISTHOBRANCHS
209
Incertae sedis
Because insufficient information is available, the following species cannot be
assigned, with certainty, to any genus. They are listed with their original
generic names.
1. Doridium adellae Dall, 1894
2. Chelidonura africana Pruvot-Fol, 195 3
3. Aglaja baken MacFarland, 1924
4. Doridium berrieri Dieuzeide, 1935
5. Aglaja dubia O’Donoghue, 1929
6. Doridium gemmata Morch, 1863
7. Doridium laurentiana Watson, 1897
8. Posterobranchaea maculata d’Orbigny, 1837
9. Doridium maderensis Watson, 1897
10. Chelidonura mediterranea Swennen, 1961
11. Aglaja minuta Pruvot-Fol, 1953
12. Aglaja nuttalli Pilsbry, 1895
1 3 . Chelidonura nyanyana Edmunds, 1968
14. Doridium orbignyana Rochebrune, 1881
15. Aglaja orientalis Baba, 1949
16. Aglaja pelsunca Marcus & Marcus, 1966
17. Chelidonura perparvum Risbec, 1928
18. Doridium punctilucens Bergh, 1893a
19. Doridium purpureum Bergh, 1894
20. Aglaja pusa Marcus & Marcus, 1967
21. Doridium reticulatum Eliot, 1903 : may = P. cyanea
22. Aglaja sanguinea Allan, 1933
23. Aglaja taila Marcus & Marcus, 1966
MATERIAL
Specimens of Chelidonura inomata and C.electra (all approximately 65 mm
in length) were made available by Professor J. E. Morton from collections made
during the 1965 Royal Society Expedition to the Solomon Islands. They were
collected at Guadalcanal. Specimens of C.electra from Orpheus Is, Queensland,
and C. inomata from Clarence River Heads, New South Wales, (all approximately 20 mm in length), were made available by Mr R. Bum.
Specimens OT Aglaja inermis, (approximately 35 mm in length), were
collected by Mr G. McDonald at the San Diego Flood Control Channel, San
Diego, California. Specimens of Aglaja ocelligera (18-30 mm in length), were
collected by Dr T. E. Thompson, at San Juan Is, Puget Sound, in July, 1969.
I am very grateful to the above mentioned colleagues for making these
specimens available.
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ABBREVIATIONS USED IN FIGURES
A.
ALB.GL.
A.0V.T.D.
A.PR.M
RB.
B.C.
C.
CAP.GL.
C.C.
CER.
C.G.
C.G.C.
CR.
D.GL.
D.G.S.
DIG.GL.
EX.
EX.S.S.
FERT.C.
G.0P.
G.S.
H.
H.O.
IN.
INT.
L.
L.GL.
L.GL.1
L.GL.V
L.M.S.
L.N.
L.R.
L.W.
M.
M.F.
M.GL.
anus
albumen gland
ampullar region of ovotestis duct
anterior protractor muscle
buccal bulb
cerebro-buccal connective
cerebral nerve
capsule gland
cerebral commissure
cerebral ganglion
cerebral ganglion
common genital chamber
crop
digestive gland
duct to gametolytic sac
digestive gland
excurrent sperm groove
exogenous sperm sac
fertilisation chamber
genital opening
gametolytic sac
nerves to Hancock’s organ
Hancock’s organ
incurrent sperm groove
intestine
nerves to extrinsic musculature of
buccal bulb
labial gland
labial gland type 1
labial gland vestibule
lateral muscle straps
labial nerve
lower raphe
wart lips of buccal bulb
mouth
muscular flap over sperm groove
mucous gland
M P.C.
minor pedal commissure
nerves to sensory bristles
nerve cells
optic nerve
oral tube
otocyst nerve
oviduct
ovotestis
ovotestis duct
opening of yellow gland
pedal ganglion
P 1-8
pedal nerves
penial papilla
PAP.
pedal commissure
P.C.
penis
PEN.
pedal ganglion
P.G.
pleural ganglion
PL.G.
PL.SU9.G. pleuro-&boesophageaI connective
P.P.
penial papilla
PR.
prostate
protractor muscles of buccal bulb
PR.M.
S.BR.
sensory bristles
S.BR.N.
nerves to sensory bristles
S.9R.M.
mound bearing sensory bristles
S.GL.
salivary gland
S.GR.
seminal groove
SP.D.
sperm duct
ST.
stomach
SUB.G.
suboesophageal ganglion
SUP.G.
supraoesophageal ganglion
SUP.G.C. connective from supraoesophageal to
pallid ganglion
SUP.V.C. connective from supraoesophageal to
visceral ganglion
U.R.
upper raphe
V.G.
visceral ganglion
Y.GL.
yellow gland
N.
N.C.
O.N.
O.T.
0T.N.
0V.D.
0V.T.
0V.T.D.
0.Y.GL.
P.