American Opisthobranch Mollusks
American Opisthobranch Mollusks
American Opisthobranch Mollusks
Frederick M. Bayer
Decem ber • 19 6 7
P u b l i s h e d by the U n i v e r s i t y of M i a m i
I n s t i t u t e of M a r in e S c ie n c e s
P reface
P re fa c e ......................................................................................................... v
vii
E x p la n a tio n o f P la t e 1
(See page 2)
A bst r ac t
I n t r o d u c t io n
E x p l a n a t io n of L e t t e r in g
Discussion.— Size, color, shell, and male copulatory organ are the principal
elements by which the species of Aglaja are distinguished. The color may
vary, e.g., in A. tricolorata (Pruvot-Fol, 1954: 51) and A. cyanea (Mac-
nae, 1962: 193); in other species it is constant and is preserved rather
well in alcohol (Bergh, 1893: 114, 126, note 1). The shell is easily broken,
and the outer rim (often of conchiolin), adheres to the wall of the shell
cavity and is hardly delimited. Spines on the inner rim of the shell,
as in A. pusa, have not been described in previous species. The male
copulatory organ of A. pusa is similar to that of A. ceylonica White
(1946: 167); this organ is not described for all species. Pruvot-Fol’s
figures of the “penis avec prostate de deux especes d’Aglaja” (1954: 50,
Fig. 8, g) are actually Bergh’s figures (1894: PI. 10, Figs. 13 and 11) of
Navanax inermis (Cooper, 1862) and N. aenigmaticus (Bergh, 1894);
Pilsbry (1895-1896: PI. 15, Figs. 89 and 86) had copied them with the
correct names.
With A. pusa we compare those species of Aglaja whose range is geo
graphically related to the coast of Florida today or was in the past.
19
West Indian species:
A. gemmata (Morch, 1863: 25). Yellow or dull fleshy with close longi
tudinal black lines. Shell not described. Unfigured. Possibly identical with
the following species.
A. punctilucens (Bergh, 1893: 131). Outer surface of parapodia and sole
with alternate yellowish and brown longitudinal lines. Head shield with
brilliant emerald-green dots, preserved in alcohol for 30 years. Shell pro
portionally smaller than that of pusa.
Species from the Atlantic Ocean and the Mediterranean Sea:
A. depicta Renier, 1807 (Bergh, 1893: 123). Cephalic shield and para
podia edged with an inner orange and an outer sky-blue stripe. Head
shield with two or four stripes of buff in front. Shell similar to that of
pusa in size and shape, but without spines. Penis tubular, not coiled;
generally paired prostates with common duct.
A. tricolorata Renier, 1807 (Bergh, 1893: 111). Edges of head shield and
parapodia frequently as in preceding species; sole velvety black (also in
depicta). Cephalic shield less than half body length. Left posterior mantle
lobe generally with flagelliform filament. Diameter of shell 6.5 mm. Male
organ as in depicta.
A. orbignyana (Rochebrune, 1881; Pilsbry, 1896: 239). Black violet with
yellowish longitudinal lines.
A. maderensis (Watson, 1897: 238). Total length 12 mm; in life, opaque
white with minute specks of crimson.
A. capensis (Bergh, 1907: 29). Greyish white with black marks, possibly
(material preserved in formalin) remains of original color. Shell does not
seem to project into right posterior mantle lobe. Male organ black in front.
A. seurati (Vayssiere, 1926: 125). Total length up to 10 mm; wax-white;
diameter of shell 3 mm.
A. dubia O’Donoghue, 1929: 11. Color as in capensis, but without black
marks. Head shield Vs of back. Shell very little produced on right side.
Left lobe with flagellum.
A. minuta Pruvot-Fol, 1953: 30. Length 5.2 mm preserved. Color as in
tricolorata, but sole brown.
American Pacific species:
A. purpurea (Bergh, 1893: 133; 1894: 209). Blackish purple alive. Right
posterior wing of shell “not long.” Male atrium black.
A. diomedea (Bergh, 1893: 133; 1894: 211). Diameter of shell 5 mm,
its left spur unusually large. Prostate paired (Marcus, 1961a: 6-7; in
PI. 1, Fig. 12, read 67 for 91).
A. ocelligera (Bergh, 1893: 133; 1894: 212). Left posterior lobe with
flagellum. Diameter of shell 4 mm; inner border broadly thickened,
especially on right, rather short wing. End of prostate bifurcate.
A. adellae (Dali, 1894: 73). Color plum with vermiculate golden yellow
spots. End of right wing of shell blunt.
20
A. bakeri MacFarland, 1924: 391. Back with irregular longitudinal lines,
light on dark ground. Head shield V3 of total length. Diameter of shell
4.7 mm; right wing long, narrow, and pointed.
A. nana Steinberg & Jones (1960: 73). Preserved 9.8 mm long. Translu
cent greyish white with irregular black flecks, small yellow brown dots and
larger opaque white spots. Shell without projecting process. Male organ
not described.
Remarks.— The absence of dark striations on the lighter ground color and
of “eye spots” (ocelli) of different color (Engel, 1936a: 59) distinguish
St. citrinus from St. longicauda (Quoy & Gaimard, 1824). Most of the
present living slugs were translucent yellow, as were the first slugs from the
same locality and Rang’s material. Some specimens, however, were more
reddish, approximately sienna. The radula is highly variable. Fine denticles
occur on the outermost lateral teeth in citrinus and sometimes in longi-
41
cauda; in citrinus they number up to eight. The length of the gut, which
we had found very much longer in longicauda from the Lesser Antilles
(Marcus, 1963: 14-15) than in citrinus, is not much longer in material
from Eniwetok, Marshall Islands. The penial spines of citrinus are smaller
and more numerous than those of longicauda, but this difference is of
degree, and, moreover, the armature of the penis becomes stronger with
age. As in our first material as well as in the present one, the secretion of
the mantle gland was purplish, not yellow as in Rang’s specimens. The
color of this secretion is not a constant specific character in Stylocheilus
(Marcus, 1962a: 18). The present animals have black contents of the gut
due to particles of pitch ingested together with the rasped algae.
One animal of the collection from Florida (Virginia Key, beach facing
Bear Cut, Biscayne Bay, J. Ewald, May 11, 1960) is enigmatic. It shows
remains of yellow color without striae or ocelli as citrinus; its seven outer
most radular teeth are smooth (hitherto only seen in longicauda), the gut
is short (citrinus), the penial spines are large and few (longicauda). We
mention this specimen in order to stress the difficulties in classifying a
deficient, possibly faded, example of Stylocheilus, but do not intend to
unite citrinus with longicauda.
46
The gill (Fig. 54) bears 12-15 pairs of pinnate leaflets, about 5 of which
are free from the branchial membrane. The anus (ar) lies over the end of
this membrane, and the prebranchial gland opens (nr) at the root of the
gill.
The elements of the jaw (Fig. 55 A) are high rodlets in the middle of
the mandible, low platelets near the borders. Their maximum height is
seven times their breadth, their free borders have 5-16 uniform denticles,
while Bergh figured 2-6 irregular ones. In the present material the radula
(Fig. 55 B) has 34 rows and 60-70 teeth per half-row; no fold or minute
tooth was seen on the rhachis. The second cusp is absent in the outermost
20 or fewer teeth; this reduction was mentioned also for m eckelii (Vays-
siere, 1885: 132). The connection of the basal region of the teeth with
the radular membrane is distended. In certain positions it appears like a
denticle. Hydrozoa, polychaetes, and crustaceans were found in the gut.
There is no flap on the collar-like membrane that surrounds the genital
apertures. The male atrium (Fig. 54, mo ) is everted in most of the present
specimens. The hermaphrodite duct (Fig. 55 C, ei) widens into a long,
winding ampulla (a) full of sperm. The exit of the ampulla is connected
with the gland mass (g) by a thin strand and is continued into a common
efferent and insemination duct ( d + xo) . Farther outward these ducts
separate. The efferent duct ( d) enters the prostate ( q) and then forms as
in P. capensis (Vayssiere, 1901a: Fig. 232) a long loop, whose ectal limb
is differentiated into a penial sheath (si) and a cuticularized penis papilla
( p ) . The papilla projects into the wrinkled male atrium ( am) .
The vagina (v) leaves the nidamental or outer oviduct (ov) as a wide
tube with folded epithelium and runs straight forwards to the thin-walled
bursa or spermatheca ( b) . The eggs in the spermatheca are evidently a
surplus of female germ cells which are digested in this organ. The insemi
nation duct ( xo) begins in the ectal third of the vagina, loops inwards and
forms a series of widenings which represent a spermatocyst (y). The
innermost of these widenings is joined with the insemination-efferent duct
by a narrow canal, halfway between prostate and ampulla. Evidently the
alien sperms rising through the insemination-efferent duct reach the eggs in
the glandular oviduct or gland mass. Studies of living material may reveal
whether the inner oviduct, the communication of the hermaphrodite duct
with the gland mass is wider in other phases of the reproductive cycle.
Discussion.— The most important difference between meckelii and occi-
dentalis, which leads us to separate them specifically, resides in the penial
sac. This is voluminous in m eckelii and lodges many circumvolutions of
the efferent duct. The sac of m eckelii is separated from the male atrium by
a closed septum. Less important differences are the lesser size of occi-
dentalis, its less numerous branchial leaflets, and its naked rhachis of the
radula. P. m eckelii has a longitudinal fold (Bergh, 1897: 18) or a median
47
tooth (Vayssiere, 1901a: 35) on the rhachis. The mandibular elements
stressed by Bergh (p. 30) as peculiar in occidentalis agree in the present
material better with the regularly dentate ones of m eckelii (Vayssiere:
PL 3) than with those of Bergh’s specimens of occidentalis (Bergh: PL 7,
Figs. 16, 17).
P. morosa (Bergh, 1892b: 28) from the western Atlantic Ocean has the
gill free for half of its length, hence is a different species.
The caudal spur figures in the diagnoses of the genus Pleurobranchaea
given by Bergh (1897: 4), Vayssiere (1901a: 22), and O’Donoghue
(1929b: 48). This is not correct, as it is absent in tarda, m aculata ,
generally called novaezealandiae (see Burn, 1958: 6), and japonica Thiele
(1925: 283). The spur is inconstant in capensis Vayssiere, 1900 (Bergh,
1907: 31, 32: absent; O’Donoghue, 1929b: 49: present), melanopus
Bergh (1907: 34: absent; O’Donoghue, 1929b: 51-52: present), and
probably in meckelii (Pruvot-Fol, 1954: 215).
photo; 63, preserved animal, showing sub-notal glands; 64, labial hooks; 65,
teeth from middle of half-row, in situ; 66, radular teeth.
54
The labial cuticle is covered with pointed hooks bearing secondary
cusps; the central hooks are 34 ^ high, the peripheral ones minute. The
formula of the radula is 60X27.0.27. The rhachis is naked. The innermost
lateral tooth is 41 ^ high, its base 24 ^ broad (Fig. 66), the edge has 2-6
inner and 4-5 outer denticles. The height of the second and third teeth
measures 42 ^ the width of the base 16 The cutting edge has one
pointed cusp and four outer denticles, the foremost of which is stronger
than the rest, so that a bicuspid aspect results. The same holds for the
further teeth, whose number of denticles generally decreases outwards.
Some teeth with only one strong cusp and several small denticles must be
defined as unicuspid. The marginal teeth are short, 24 ju,; their upper prong
is longer than the lower one and followed by about 4 weak denticles. The
lower prong is often cleft in any of the teeth.
Discussion. — The combination of blue with red is rare among the Chromo-
doridinae, while blue with yellow or orange is frequent. Therefore we also
compared the species with the latter colors, but did not find the same
pattern in any species with the peculiar radula of the present one. We even
pondered possible allocation to N oum ea Risbec (1928: 165), due to the
broad innermost lateral tooth whose form is similar to that of Noumea
flava Risbec (1928: 169; 1953: Fig. 47A). However, the different width
of the first and second lateral tooth is less pronounced in nyalya than in
Risbec’s species. Pruvot-Fol (1951a: 126, 139) questions the validity of
N oum ea , with exception of the type species. Even if Noumea is maintained
in the sense of Risbec (1953: 85-90) and Baba (1949: 54-55, 144) none
of its species agrees with nyalya, whose lateral teeth are quite peculiar.
Description .— The color of the living animals, from photos and collector’s
notes, is blue, hidden by yellow pigment on the notum. The area around
the mouth is blue, the anterior border of the foot yellow. The thin margin
of the notum (Bergh’s “Mantelgebrame” ) is brimmed with a fine white
line, closely accompanied by a pale red one. The inwards following yellow
color of the margin darkens gradually and ends on a bright yellow band,
visible also on the hyponotum, due to the thinness of the margin. On the
inner edge of the yellow band there is a row of about a dozen blackish
brown spots on either side, and next to them a row of irregular brilliant
blue blotches. The middle area of the back is stippled with yellow tubercles
and blue dots in the spaces of a yellow network. The same pattern occurs
on the sides of the foot, and further two rows of blue blotches as well as
one of violet spots on the limit towards the lavender-blue sole. The rhino
phores are lavender, the bases of the gills brick-red, their tips yellowish.
The preserved slugs are up to 60 mm long, 18 mm broad, 15 mm high.
The tentacles are small and folded. The rhinophores have about 30 leaves
of which 20 are complete. The anterior border of the foot is grooved, its
lateral angles projecting and pointed. The sole is narrow, 4 mm in width.
Thick whitish glands lie along the border of the notum. There are eleven
bipinnate gills, in one specimen unipinnate with several bifid tips. The anal
papilla completes the circle of the branchiae behind.
The labial elements (Figs. 73, 74) are unicuspid. The radular formula
is 83X 175.0.175. The rhachis is naked. The innermost and lateral teeth
(Fig. 75) are of the same type, bicuspid with a stronger upper and a
considerably smaller lower prong. In about ten marginal teeth there are
two blunt cusps of equal size and a weak denticulation on the lower edge.
One specimen had an aberrant innermost tooth with a single hook widely
distant from the apex and a second tooth with one cusp in similar position
and a lower much smaller one.
60
Discussion.— The teeth in the center of the half-row agree with White’s
Figure 16 A; but there the innermost teeth (Figs. 16 B, C) are a little
shorter. White had no record of the living animal and classified it as
Glossodoris aegialia (Bergh, 1905b: 70) from the Gulf of California. As
that species has two lateral rows of eye-spots bearing pupillae and a
similar row in the anterior half of the back, it can hardly be identical
with White’s species which is smooth. The elements of the labial plates
and the radula of the present species agree sufficiently with Bergh’s, but
both, as well as White’s species, are of the regular H ypselodoris- type, so
that the agreement is of little weight. The arrangement of the pupillae in
aegialia excludes a nearer relation to acriba.
Description .—According to the collector’s notes, the living slugs are “dirty
yellow with blackish spots” and “tan with greenish black spots.” The
preserved specimens are white, in one animal with yellow gills. The biggest
slug is 12 mm in length, 6 mm in breadth, 4 mm in height. The width of
the hyponotum is 2 mm, that of the foot 3 mm. The body is rigid. The
center of the notum is sculptured with low bosses of different sizes stiffened
by perpendicular spicules which do not project from them (Fig. 81). The
bosses on the sides of the notum are smaller and disappear near the edge.
There is a horizontal layer of larger spicules in the connective tissue under
the bosses, around which they lie radially. The underside is smooth, also
with horizontal spicules; a band of glands lies around the border of the
hyponotum.
Four lobes surround the pits of the rhinophores, whose clubs bear about
15 foliations. The tentacles are barrel-shaped with an outer furrow. There
are up to 11 unipinnate gills. The border of the branchial pocket is circular
with a lobed outline, or contracted to semilunar or reniform shape. The
anal papilla lies in the center of the plumes. Often some plumes are bitten
off; in one slug, only three remain. The foot is bilabiate and notched in
front, pointed behind; it nearly reaches the posterior border of the notum.
The labial cuticle is smooth, without markings. The radula (Fig. 82)
comprises 47 rows with 2.35.0.35.2 teeth. The inner teeth are strongly
64
F i g u r e s 80-84. Siraius kyolis, sp . n o v . 80, p r e s e r v e d s p e c im e n ; 81, sk in ;
82, ra d u la r te e th ; 83, d ia g r a m o f a lim e n ta r y tra ct; 84, d ia g r a m o f r e p r o d u c tiv e
org an s.
curved hooks; in the middle of the half-row the teeth become straighter;
the marginal teeth are pectinate with stronger denticles on inner and outer
edges. The teeth of the oldest rows are bigger than those of the younger
ones, suggesting that growth continues after the formation of the tooth in
the radular papillae. In the oldest row (compared with the 40th row), the
height of the teeth is: innermost tooth 44 ^ (30 ^ ); middle tooth 80 ^
(68 /x); 34th tooth 95 ^ (80 ^ ). The size of the outermost hook-shaped
and pectinate teeth decreases suddenly. The salivary glands are ribbon-like,
the left is longer than the right one. The stomach projects from the
intestinal gland; the right liver (r) has about half the breadth of the
stomach (so). One stomach contained part of a dorididan radula com
prising three rows of teeth.
The thick winding ampulla (a ) divides at its outlet into the spermatic
65
duct and the inner oviduct. The latter is quite short, merging into the
large female gland mass (g). The former becomes glandular immediately.
This bent prostatic section ( q ) is followed by a narrow and short efferent
duct ( d ), whose ectal half is contained in a muscular sheath; its tip
projects into the male atrium.
The vagina (v) begins wide, narrows gradually, and enters the sperma
theca ( b ) with a thin canal. The dark contents of the spermatheca shine
through the skin as a blackish spot. The spermatocyst ( y ) lies near the
spermatheca; all sperms are fastened with their heads to the fundus of the
spermatocyst. This seminal receptacle is connected with the insemination
duct ( xo ) by its own canal. The insemination duct opens into the gland
mass near the entrance of the inner oviduct.
Discussion .— Since the type-species of Siraius, S. ilo Marcus (1955: 135),
was described, 19 specimens of it have been seen, all greyish yellow without
dark spots. From the coast of Sao Paulo the range of ilo was extended to
Cabo Frio (1958b: 57). It has 22 branchiae and more teeth per half-row
than kyolis, but only 37 radular rows. In Siraius as in Austrodoris (Odhner,
1934: 269), the shape of the salivary glands cannot be maintained as a
generic character. In S. ilo they are sac-like, short and broad, in kyolis
ribbon-like, similar to those of Doris bovena Marcus (1955: Fig. 117, r).
They are still longer in Doris bicolor Bergh (1884b: 655), a small
Mediterranean species which probably belongs to Siraius. With respect to
the number of gills and of teeth per half-row, bicolor is near kyolis but it
has a maximum of 34 radular rows, up to six pectinate teeth, and a con
siderably longer right liver than kyolis.
The seminal receptacles, called seminal vesicles by Bergh, were called
vaginal in Siraius. If, however, those of Homoiodoris Bergh (1882: PI.
6, Fig. 16) were qualified as semiserial (Odhner, 1926: 54), the vesicles
of S. ilo and S. kyolis would be semiserial too. In the vaginal type (Odhner,
1926: 51), the two vesicles lie quite near to one another at the inner end
of the vagina, while in the semiserial and in the serial type the spermatocyst
is connected with the insemination or uterine duct. In the former type, the
insemination duct originates from the entrance of the vagina into the
spermatheca, as in Siraius; in the latter it leaves the spermatheca inde
pendent from the vagina, as in Peltodoris greeleyi (Fig. 98). The true
vaginal arrangement occurs, e.g., in Austrodoris Odhner (1926; 1934),
but it is rare in the doridids. In Doris, another genus with vaginal seminal
vesicles, these do not lie so near together (Marcus, 1955: Fig. 106, the
type-species) as in Austrodoris. In Neodoris Baba (1938: 13) the semi
serial and the serial types occur (Marcus, 1959a: 37 ff).
39. Anisodoris worki, spec. nov.
Figures 85-89
Material.— Florida: Biscayne Bay; under rocks, east end of County
66
(MacArthur) Causeway. R. C. Work, August 7, 1964. One specimen.
Holotype .— USNM 576289.
Description .— The color photo shows a slug 28.5 mm long, 16 mm broad.
This is evidently the natural size, as the preserved specimen is 20.5 mm
in length, 11.5 mm in breadth, and 6.5 mm in height. The breadth of the
hyponotum is 4 mm; the foot is 3.5 mm broad but rolled in.
The color alive is yellowish; the papillae, rhinophores, and gills are
darker yellow. Dark spots in the connective tissue are interrupted by the
papillae, which are lighter than the spots. The middle of the back is darker
yellow. The papillae are smaller on the margin of the notum. The hypo
notum and the foot show small black specks; there is a row of eight
larger, longish spots on either side of the inner border of the hyponotum
(Fig. 86). In the preserved animal there are small brown chromatophores
on the tips of the round papillae, in the rhinophores, and in the gills.
The notal sculpture is cauliflower-like (Fig. 86), as the interspaces
between the larger papillae (0.8 mm) are filled with small ones (0.15 mm).
The pits of the rhinophores and branchiae are surrounded by middle-sized
papillae. The spicules are often blunt and do not project above the surface.
The tentacles are rather long, flattened clubs without furrows; there are
about 25 foliations on the rhinophores, and six multipinnate gills with
warts on their outer sides. The anterior border of the foot is transversely
grooved, the upper lip notched in the middle.
The labial cuticle is smooth. The radula has 27 rows and 38 hamate
teeth per half-row (Fig. 87). The size of the innermost tooth is 40-47
from the second to the third tooth the size increases from 50 to 80
The teeth in the middle of the half-row measure 160 the outermost
tooth 45-52 [A, the penultimate and the antepenultimate teeth 85 and 100 ^
respectively. As the fourth tooth from the outer border is 150 ^ long, the
fourth from the middle 95 the pararhachidian teeth are smaller than the
outer ones. The stomach is not covered by the intestinal gland.
The ampulla (a) is long and winding, the female gland mass (g) large,
wrapping the seminal reservoirs. The male duct leaves the gland mass,
widens into a folded prostate ( q ), and runs looping ( d ) towards the
muscular, strong penis ( p ) lodged in the atrium. The vagina opens
together with the penis, the nidamental duct farther ventrally. The vagina
(v) is narrow; it enters the spermatheca ( b ). The short insemination duct
( x o ) leaves the spermatheca at this entrance. The spermatocyst (y) is
inserted on the duct which continues into the gland mass. The ectal section
of the vagina bears a flattened gland ( va ), connected with the vagina by a
short, muscular duct (vz). The wall of this duct (Fig. 89) contains a large
papilla with a spikelet and many smaller papillae, also with spikelets,
around the bigger one.
67
The species is named for the collector.
Discussion .— In his original diagnosis of Anisodoris, Bergh (1898b: 508)
included “tentacles as in Archidoris;,> that is: “low, fold-like” (Bergh,
1878: 616). In the type-species, Archidoris tuberculata, they bear an
68
external furrow. When Eliot (1910: 95) selected the type-species of
Anisodoris, A . punctuolata (d’Orbigny, 1837: 187), he did not consider
its pointed tentacles; in other species there are furrowed lobules (Marcus,
1959a: 48). Nor can “tentacles finger-shaped” (Pruvot-Fol, 1954: 239)
be used as a character of Anisodoris, and even less “vestibular gland
present” (ibid.). The smooth labial cuticle is not mentioned in Pruvot-Fol’s
diagnosis, though it is the only character that separates A nisodoris from
Discodoris with labial rodlets. Sometimes these are very short (MacFarland,
1909: 75) or absent (Marcus, 1955: 148) in Discodoris, possibly worn
or in regeneration. Often the notal papillae of A nisodoris are stronger than
the granules of the generally softer Discodoris, while the rigid notum of
Peltodoris becomes velvety by densely standing, uniform caryophyllidia.
Therewith the general aspect of most species of these genera is different. In
preserved specimens, however, the notum may be swollen or shrunken
and the sculpture modified. The characters of sculpture are difficult to
describe unequivocally in different languages, and, moreover, exposed to
subjective preferences, e.g., “pustules” (Burn, 1962c: 156) and “Nopp-
chen” (Bergh, 1905a: 98).
The species nearest to Anisodoris worki is Archidoris marmorata Bergh
(1881: 86) which belongs to the Discodoridinae due to its prostate
(Odhner, 1926: 64-65), and to A nisodoris due to its smooth labial cuticle
and notal sculpture. The Anisodoris m arm orata Bergh (1898b: 515) from
northern and middle Chile (Marcus, 1959a: 45, 88) must be re-named
as A. rudberghi, nom. nov.
Bergh’s Archidoris m arm orata came from Trieste. It differs from w orki
by the absence of spines in the duct of the vestibulo-vaginal gland (Bergh,
1881: 92), by the reddish brown back with white nodules and violet black
clubs of the rhinophores, by the size of the body (living 47, preserved
nearly 40 mm), and by eight gills.
Vayssiere’s A rchidoris marmorata Bergh (Vayssiere, 1901b: 16) from
Marseille cannot be the same species, because its labial cuticle bears
rodlets, nor can it be an Archidoris. It is not known whether it has a
prostate and a vestibular gland. Both features, labial rodlets and undes
cribed prostate and accessory gland, also make it difficult to define A rchi
doris stellifera Vayssiere (1903: 82) from the Gulf of Lions and Naples.
Certainly it is not an Anisodoris. Vayssiere (1904: 129; 1913: 314-315)
considers the radular teeth of his marmorata Bergh and stellifera as
specifically different.
Pruvot-Fol (1954: 240) unites marm orata Bergh, 1881, marmorata
Vayssiere, 1901, and stellifera Vayssiere, 1903, under the name A nisodoris
stellifera Vayssiere, 1904. Her figure 90 combines details from the publi
cations in 1881, 1901, and 1904, though only the reproductive organs of
marmorata Bergh (her figure 90, e) are known. It is possible that she had
material of marmorata Bergh, 1881, as she writes: “labial cuticle unarmed,
69
folded, sometimes pierced by some hairs, which do not constitute an
armature.”
Accessory vaginal glands are frequent in the Dorididae (Bergh, 1892a:
1086); if not combined with other structures, they are no more than
specific characters.
The vaginal appendage of Anisodoris fiindersi Burn (1962c: Fig. 11,
sp) is considered as a spermatheca by the author. The sculpture of the
notum of fiindersi differs by two rows of high tubercles from that of worki.
Also A. timorensis Bergh (1905a: 97), whose reproductive organs are
not known, differs by the tubercles of its back, 80-120 ^ in diameter,
from worki.
40. Anisodoris prea, spec. nov.
Figures 90-93
Material.—Florida: Biscayne Bay; Matheson Hammock; Thalassia and
sand flats. F. M. Bayer, February 21, 1948. Four specimens.
Syntypes .— USNM 576278.
Description .— The biggest animal is preserved 28 mm in length, 19 mm in
breadth, and 8 mm in height. The hyponotum is 4 mm broad. The smallest
specimen is 17 mm long, 10 mm broad. The color is a creamy white,
without traces of pigment; the body is stiff. The notum is covered with low,
flat knobs of different size, each big one surrounded by a number of small
ones. The middle of the back is nearly smooth, sharply delimited on either
side by a row of large tubercles which form a kind of crest. Towards the
sides the knobs are smaller. The spicules are partly dissolved, but sections
(Fig. 91) show that they were radially dispersed in the knobs and
projected slightly from them.
The pits of the rhinophores bear a bigger tubercle on either side. There
are about 30 rhinophoral leaflets containing spicules. The tentacles are
flat, broader than long, contracted. The border of the branchial pocket is
retracted; there are seven multipinnate gills. The foot is bilabiate, its upper
lip notched.
The labial cuticle is smooth. The radula is 2.3 mm in length, 2 mm in
width. Its formula is 25X29-30.0.29-30; the last 5 rows are still weak.
The rhachis is broad. All teeth (Fig. 92) are hook-shaped, the inner ones
bigger (0.13 mm) than the outer ones (50 ^ ). The maximum height in
the middle of the half-row is 0.2 mm. The stomach is free, covering the
spherical right liver. The salivary glands are long.
The ampulla ( a ) is winding, its outlet separates into the spermatic and
the oviduct. Where the latter enters the female gland mass (g), lies the
small albumen gland. The male duct passes immediately into the folded
prostate ( q ), which is much longer than wide. The following section of the
efferent duct ( d ) is coiled and ectally enclosed in a muscular sheath. A
70
short conical penial papilla projects into the atrium. The vagina (v) begins
wide, narrows farther inwards, forms a loop, and enters the spermatheca
( b ). As in the other species of the genus the entrance into and the outlet
from the spermatheca is established by a short single duct; the seminal
reservoirs are arranged semiserially. The insemination duct (xo ) or com
munication between the receptacles is angled in a similar way as in A.
nobilis (Marcus, 1961a: Fig. 57), and also its continuation, the insemi
nation duct of the mentioned figure, is quite short in the present species;
it goes out from the spermatocyst ( y ).
F i g u r e s 90-93. Anisodoris prea, sp. nov. 90, dorsal view of preserved slug; 91,
section of skin; 92, radular teeth; 93, diagram of reproductive organs.
77
2. Half-row of radula with a minimum of 71 teeth ............... evelinae
2. Half-row of radula with a maximum of 53 teeth ............................ 3
3. Rhachis of radula deeply grooved longitudinally; penis of adult
4 mm long, its hooks up to 27 p in h e ig h t................................. branneri
3. Rhachis of radula with longitudinal serpentine fold; penis of adult
11 mm long, its hooks up to 250 ^ in h e ig h t.......................... hedgpethi
The outlet of the ampulla (a) divides into the male and female ducts.
The spermatic duct immediately enters the massive prostate ( q ) whose
transparent ental part consists of small cells, the yellow ectal part of large
ones. Also the following tubular section is glandular; the next coiled stretch
of the efferent duct ( d ) is narrow, and its short ectal portion runs within
a muscular sheath. The strongly muscular male atrium lodges the small
penial papilla. Neither atrium nor papilla bears spines, as was determined
by the examination of sections.
The rather long vagina (v) leads to the spermatheca ( b ) which is
surrounded by the coils of the efferent duct. The insemination duct ( xo )
begins at the entrance of the vagina. The spermatocyst ( y ) has its own
canal; behind its connection with the insemination duct the latter merges
into the gland mass ( g ) near the ampulla and the quite short inner oviduct.
79
Within the gland mass the insemination duct forms a thin-walled, globular
vesicle, the fertilization chamber ( b e ) , filled with sperm, similar to that
of Thordisa diuda (Marcus, 1955: Fig. 140, r ).
Discussion — Of the Atlantic species of Discodoris, four have a spiny penis
by which they differ from phoca. All of these occur on the American coast.
They are: (1) branneri MacFarland (1909: 66) with a longitudinal row
of five or six larger black spots along either side of the notum; (2) evelinae
Marcus (1955: 143) has mamillate warts, not caryophyllidia; (3) hedg-
pethi Marcus (1959b:254) differs from evelinae by more radular rows,
fewer teeth per half-row, and longer and stronger spinous male organ; and
(4) mortenseni Marcus (1963: 30) is similar to phoca, but different by
minute penial spines, small size, in life up to 20 mm, a white foot, the
breadth of the radula little inferior (92.3 per cent) to its length, one
fourth of the outer teeth decreasing in size, and often the two outermost
teeth are dentate.
The remaining Atlantic species of D iscodoris have no penial spines, as
phoca, or the armature of the male organ is not described. American
species are: (5) alba White (1952: 113), in which the last third of the
half-row decreases in size, and whose black spots were only described for
the back; (6) m uta Bergh (1877b: 532), with the base of its innermost
tooth (PI. 62, Fig. 17) split, and without black spots on the sole; (7)
notha Bergh (1877b: 530) with a muscular and cuticularized cone beside
the aperture of the male atrium; (8) purcina of the present collection,
whose radula is broader than long; (9) pusae Marcus (1955: 147) with
8-12 marginal teeth developed as thin lamellae; and (10) voniheringi
MacFarland (1909: 73), whose six gills are bipinnate and whose labial
cuticle consists of three fields, a median and two lateral ones.
The Central and Eastern Atlantic and Mediterranean species of D isco
doris differ from phoca; (11) cavernae Starmiihlner (1955: 217) by two
ramified vestibular glands; (12) edwardsii Vayssiere (1902: 232) by
2-3 rudimentary innermost teeth; (13) erubescens Bergh (1884b: 662)
by its color, white with reddish hue alive and 12 bipinnate gills; (14)
indecora Bergh (1881: 108) by its small size (alive 21 mm) and color,
olive brown with whitish spots; (15) maculosa Bergh (1884b: 658) by
its rhachidian platelets; (16) rubens Vayssiere (1919: 65) by its 7-8
lamellate marginal teeth; and (17) tristis Bergh (1899: 11) by its tentacles
grooved on the outer side.
The Indo-Pacific representatives of Discodoris were only cursorily
compared with phoca, with exception of the two species from the American
Pacific coast, heathi MacFarland (1905: 39; 1906: 118) and aurila in the
present collection. The former— as well as D. fulva O’Donoghue (1924:
27), evidently a juvenile heathi— has 12-16 outermost teeth in form of
thin, concave plates. D. aurila has a very differently colored notum and
no melanophores on the underside.
80
45. Discodoris purcina, spec. nov.
Figures 102-104
Material.— Florida: Southwest Point, Biscayne Key. F. M. Bayer and E.
Deichmann, 1962. One specimen.
Holotype .— USNM 576281.
Description .— The length of the preserved animal is 25 mm, its breadth
18 mm, the height 7.5 mm. The foot is curved, 17 mm long, 4 mm broad.
The notum is transparent greyish with fine black granules, swollen, not
well preserved, probably bearing small papillae that had been stiffened by
spicules which are dissolved. The border of the notum is even, not
undulated. The hyponotum is broad, the underside colorless. A general
sulphureous hue may be due to preservation.
The tentacles are rather thick and pointed, not grooved. The rhinophores
are blackish owing to melanophores, and they have about 20 leaflets.
The anterior pedal border is bilabiate, the upper lip notched. The six
black bipinnate gills are of small size. The borders of the rhinophoral and
branchial pockets are smooth.
81
The labial cuticle bears two triangular fields of yellowish, stratified
rodlets, up to 50 ^ high and 5 ^ in diameter. The radula is colorless, very
short, broader (3.3 mm) than long (2.5 mm), and comprises 16 rows
and 43 teeth per half-row. All teeth are hook-shaped with smooth cusps.
The innermost tooth is 90 f± long, the six following teeth are also small.
Then the size of the teeth increases to 300 ^ and decreases in about six
of the outer teeth. The stomach is free, not enclosed in the intestinal gland.
The ampulla is fusiform; the spermiduct and the female duct separate
from one another in the female gland mass. The prostate is sausage-shaped.
The male atrium contains crystals, probably precipitates resulting from
the “hexamine” buffering of the formalin (letter of Prof. Bayer of May 12,
1964), which hinder the examination of the copulatory organ. The vagina
has no special gland; the insemination duct leaves the spermatheca to
gether with the entrance of the vagina into the latter. The insemination
duct is very long, and its entrance into and outlet from the spermatocyst
are united.
D iscussion.— The principal characters of D. purcina are the unicolored
notum with black gills, and the exceptionally short radula which is broader
than long. By its radula D. purcina comes near to only two of the nine
species of the genus that were recorded from West Atlantic warm waters,
D. alba White (1952: 113) from Dry Tortugas, and D. mortenseni
Marcus (1963: 30) from Curasao and Tobago. In both of them the
radula is a little longer than broad. The innermost tooth of alba is bluntly
hamate, and the teeth in the center of the half-row have rather short
square bases. D. mortenseni has an expansion near the base of the inner
most tooth, and also the basal expansions in the middle of the half-row
are more pronounced than in purcina. Outermost teeth with denticles
are frequent in mortenseni.
A small number of radular rows occurs also in the Californian D. heat hi
MacFarland, 1905. Moreover, its notum is similar to that of purcina in
color and sculpture. However, it differs from purcina by its 8-10 bran
chiae and the quite different shape of all radular teeth (Marcus, 1961a:
Fig. 63).
46. D iscodoris pusae Marcus, 1955
Figures 105-107
R eference.— Marcus, 1955: 147-150, Figs. 151-165.
M aterial. — 1. Florida: Biscayne Bay. Under rocks at east end of County
Causeway. R. C. Work, April 11, 1963. One specimen.— 2. Florida:
Biscayne Bay, off Matheson Hammock. March 6, 1965. Shrimp trawler.
Three specimens.
Further distribution .— Brazil, northern coast of the State of Sao Paulo,
shallow water.
82
Description .— As the specimens from the two mentioned localities are
different in size and some inner characters, they are described separately.
The animal from the first locality, accompanied by a color photo, has a
pinkish red notum, darker in the middle than on the sides. There are
nearly contiguous violet red blotches along the mid-line and more spaced
smaller ones on either side between the rhinophores and the branchiae.
Small dark and blunt knobs of irregular sizes and intervals are scattered
all over the notum, those near the margin are smallest. The tip, shaft,
and anterior side of the rhinophorial club are whitish; basal and posterior
parts of the rhinophores dark. The outer side of the gills is chalk-white,
their inner side violet with rare white stipples, probably secreted matter.
83
Also the red anal papilla is frosted with white. Long white spicules form
a horizontal lattice under the surface and around the border of the notum.
The length of the preserved animal measured over the back is 30 mm,
lineal 20 mm, the breadth 18 mm, the height 10 mm. The width of the
hyponotum is 7 mm, equal to that of the sole. The anterior border of the
foot is bilabiate, its upper lip notched. There are about 20 rhinophoral
foliations; the border of the pit is slightly lobate. The tentacles are flat,
with broad base and pointed tip. Of the six gills the anterior ones have
small, bipinnate plumes; the posterior ones are much bigger and tripinnate.
This character is probably due to a regeneration of lost gills. The border
of the branchial pocket is smooth. The spicules were dissolved.
The labial armature consists of two nearly triangular fields. The
stratified rodlets measure 60 /a in height, nearly 10 ^ in diameter. The
radula comprises 26 rows, each half-row contains 36-38 lateral and 15-18
marginal teeth. The rhachis is naked, the innermost tooth is often rudi
mentary. The marginal teeth are soft, blunt, and concave lamellae; their
edges bear minute irregular denticles, hardly discernible. The lateral teeth
are smooth hooks whose size increases outwards, only three laterals pre
ceding the marginals are smaller.
The ampulla (a) is long, winding. At its ectal end the male duct enters
the prostate ( q ) directly, and the inner oviduct enters the female gland mass
(g). The inner massive portion of the prostate consists of small cells.
The middle part is also massive, but composed of large cells; the outer
most section is tubular. The sheathed muscular, efferent duct ( d ) ends
with a short penial papilla.
Between the outer oviduct or nidamental duct (ov), that is the outlet
of the gland mass (g), and the penial papilla an uncommonly narrow
vagina (v) runs inwards and opens into the longish spermatheca ( b ).
Close to the vaginal entrance the insemination duct ( x o ) originates, so that
the arrangement of the seminal receptacles is serial. The considerable
width of the insemination duct is exceptional, and the same holds for the
position of the spermatocyst (y) quite near to the gland mass. A long
sausage-shaped gland ( vn ) with cuticular spines in its duct opens into the
female vestibule.
The specimens from the second locality were red alive; preserved they
are whitish with brown spots. Their length is 11 mm, the width 6 mm,
and the height 4 mm. The breadth of the sole is 2 mm. As far as the skin,
which is not well preserved, is recognizable, it resembles that of the original
description (Marcus, 1955: 147). The rhinophoral clubs have about 16
foliations on either side. The five to six gills are bipinnate, as frequently
those of small doridids of a species are, whose larger specimens have a
greater number of branchiae and more ramified pinnules.
The labial rodlets are 50 high and 8 ^ in diameter. The radula is
1.6 mm in length, 1.17 mm in width, hence half the size of that of the
84
specimen from the first locality. In Brazilian D. pusae the corresponding
measurements are 2.3 and 1.5 mm. The number of radular rows is 23,
that of the hooks 15, and that of the marginal lamellae 10. The innermost
lateral tooth bears one or two inner denticles, absent in the specimen
from the first locality. In the original material there is sometimes one den
ticle. The lamellae are smooth, as in the specimens from Brazil; while
they bear minute irregular denticles in the slug from the first locality.
This minor character observed in a single specimen cannot be considered
as taxonomically significant.
The reproductive organs combine characters of the animal from the
first locality with such from Sao Paulo. The section of the insemination
duct between spermatocyst and female gland mass is short as in the former,
and its width is the same as that of the vagina, as in the latter. Cuticular
spines in the duct of the vestibular gland are wanting as in the small
specimens of the original material (Marcus, 1955: 149). Both Floridian
materials have simple, not stratified, labial rodlets.
Remark .— The original material, of which we have about 20 specimens,
is united with the animal of April 1963 by the slugs of March 1965.
F i g u r e s 108-111. Discodoris aurila, sp. nov. 108, living animal, from color
photo; 109, labial armature and its elements; 110, radular teeth; 111, diagram
of reproductive organs.— F ig u r e 112. Platydoris angustipes Morch, living
animal, from color photo.
The ampulla ( a ) is long; its outlet divides immediately into male and
female ducts. The male duct enters the voluminous prostate ( q ) whose
nuclei are small in the inner, large in the outer part. Ectally to the
86
prostate the efferent duct ( d ) begins with a muscular bulb, narrows, and
projects into the vestibule with a minute papilla.
The female gland mass (g) is conspicuous, its nidamental duct ( o v )
is glandular to near its aperture. The vagina (v) courses inwards beside
the penial papilla. The vaginal entrance into the large spermatheca ( b )
and the origin of the insemination duct ( xo ) are neighboring but separated.
The insemination duct is long, looping; the duct of the small, silky
spermatocyst (y), enters it near its opening into the gland mass.
Discussion .— The range of most doridids is little known, because many
of them were described only once, and based upon very few, often a single
specimen. The two descriptions of the Antillean Discodoris notha, for
example (Bergh, 1877b: 530; 1904: 53), refer to a total of three specimens
collected in 1860 and 1899. A doridid of the almost unexplored opistho-
branch fauna of Panama requires comparison with many congeneric species,
so much the more, as the present collection reveals obvious relationships
between the Pacific side of the Canal Zone and the Caribbean. Therefore
we went through the descriptions of the 28 species from the Indo-Pacific
Ocean and the 18 from the Atlantic Ocean and the Mediterranean Sea.
The geographically least distant D. heathi Mac Farland (1905: 39; 1906:
118), whose southernmost record on the American Pacific coast is Laguna
Beach (33° 32' N ), is not related with aurila. Its 12-16 outermost radular
teeth are reduced to thin, concave plates. Though it is not considerably
bigger than aurila, it has 8-10 branchial plumes.
Among the Indo-Pacific species we found D. pallida Baba (1937: 305)
from Amakusa (32° 30' N) similar to aurila. It differs by its nine gills,
the radular formula 15 X 14-17.0.14-17, and the larger labial plates. Also
D. modesta Bergh (1877b: 534) from the Palau Islands, Western Caro
lines, resembles aurila. Its rhinophores have 50 leaflets, the ends of the
labial rodlets are thickened and often fringed; the radular formula is
29X48.0.48. The Atlantic D. voniheringi MacFarland (1909: 73) from
Alagoas, Brazil (9° S) has bipinnate gills, and the labial armature consists
of a median and two lateral plates.
90
entally the vagina narrows and enters the spermatheca ( b ). Beside this
entrance, but with an independent origin, the narrow insemination duct
( x o ) leaves the spermatheca. This duct forms two loops and is connected
with the large, sausage-shaped spermatocyst (y) by a short canal before
it enters the gland mass. The entrance of the insemination duct is far
distant from that of the inner oviduct.
Discussion. — Since the introduction of Thordisa Bergh (1877b: 540), the
diagnosis of that genus has been reduced too much (Bergh, 1892c: 132-
133). The characters of the type species must be considered, in order to
define Thordisa : notum villous, labial cuticle smooth, lateral radular teeth
without denticles, marginal teeth pectinate, penis unarmed. The dorsal
sculpture of densely set soft villous papillae should not be considered in a
restricted sense, and Th. diuda Marcus (1955: Fig. 136) with conical
tubercles, longer than broad and of different length, can be retained in
the genus. The tentacles, pointed conical knobs in the living animal (Fig.
134), round buttons in preserved specimens (Fig. 137, /), are difficult
to judge.
E tidoris Ihering (1886: 234) has sparsely set long tubercles mixed with
smaller ones, hence is not villous (Marcus, 1958a: 25). More important
than this difference from Thordisa is the glandular section of the efferent
duct in E. ladislavii, which contrasts with the massive prostate of Thordisa.
Species whose marginal radular teeth are not pectinate, e. g., Th. clan-
destina Bergh (1884a: 106) and Th. maculosa Bergh (1905a: 123), must
be excluded from Thordisa.
For species with denticles on the outer side of the lateral teeth we adopt
A porodoris Ihering (1886: 238). This character was stressed by Eliot
(1910: 106-107); it was not seen in the type species by Alder & Hancock
(1855: 42, D oris m illegrana) . Even Thiele (1931: 438) left the denti-
culation unmentioned. Thordisa dubia Bergh (1894: 178) is identical with
A porodoris millegrana. Bergh’s description of a voluminous prostate and
an unarmed penis completes Eliot’s. Further species of A porodoris are
Thordisa souriei Pruvot-Fol (1953: 66) and Thordisa hilaris Bergh, 1905,
variety Risbec (1953: 55), which we name A porodoris risbeci, nom. nov.
A porodoris rubra Bergh (1905a: 94), whose cusps of the lateral teeth
are smooth, does not belong to A porodoris. The cones in its efferent duct
suggest a Baptodoris, but the type-species of this genus has bipinnate gills
(Bergh, 1884b: 671), while rubra seems to have unipinnate ones.
Taringa Marcus (1955: 151) is separated from the preceding genera by
an armed penial papilla; the radula is as in Aporodoris. The three species
differ by their dorsal sculpture and the shape of the male organ. The
typical T. telopia, telopia disa, and T. armata Swennen (1961: 64) have
only caryophyllidia, while these are mixed with longer tubercles in aivica.
In the Atlantic and the Mediterranean species the penis is approximately
91
bell-shaped, in the Pacific one conical. The innermost tooth of the latter
has no inner denticles.
0.5 mm. The skin is leathery; the notum contains large spicules, up to 1
mm in length and 0.1 mm in diameter. The hyponotum is 2.5 mm broad,
reticulated, with dense white strands and transparent interspaces.
The color alive, from the photo, is light red. Two blackish bands, which
have a common origin between the rhinophores, end under the branchial
97
tuft, and leave the tubercles free. Minute white dots on the blunt tips of the
latter are cutaneous glands. Around the base of every tubercle, except the
quite marginal ones, there are about five black spots, evidently single
melanophores. There are two pairs of bigger white tubercles and many
small, dark red spots, the latter especially in the middle. The rhinophores
are dark red with small white tips, and white dots occur on the rim of the
rhinophoral pits. The black pigment is retained after preservation; the
white glandular secretion and the red color have disappeared. The absence
of black pigment in the inner organs contrasts with several other dendro-
doridids.
The tentacles are quite short, the rhinophores have around 18 leaflets on
either side; the sheaths of the rhinophores are high, with smooth rims and
small tubercles on the outer side. The anterior pedal border is bilabiate; the
sole is 7 mm in breadth. There are five multipinnate gills; the anal papilla
lies to the left, between the anterior and posterior plumes.
The central nervous system is smooth when examined from below.
A constriction shows between the pedal ganglia, and apposed to their
hinder surface lie the buccal ganglia, far from the oral vestibule. The latter
is gradually dilated into a pharynx, whose anterior part is invaginated into
the vestibule, as in our figure of D. areolata Bergh (Marcus, 1962b: Fig.
18). Two retractors insert at the point of this invagination.
The ampulla ( a ) is longish; the spermoviduct enters the female gland
mass ( g ) where the oviduct and the spermiduct separate. The latter
emerges from the mass as a short duct and forms the prostate ( q ), the
uppermost organ of the anterior genital mass. The prostate consists of
branched diverticula. The male duct ( d ) passes through it and is in its
farther ectal course sheathed by muscles. Before the efferent duct enters
the male atrium, it becomes spinous ( p ). The spines are numerous and
pointed, about 50 ^ in length.
Beside the male aperture lie the openings of the vagina (v) and the
oviduct. The terminal part of the vagina is widened and heavily ciliated.
The vagina runs to a spherical spermatheca (b). A long duct connects the
spermatheca and the spermatocyst ( y ). This duct leaves the spermatheca
at the entrance of the vagina, and the insemination duct ( xo ) starts from
the same point, coursing to the gland mass, which it enters near the
emerging male duct.
The egg ribbon is about 2 mm broad and forms a spiral of a little more
than three coils, 11 mm in diameter; unrolled it would be 5 cm long.
The diameter of the eggs is 0.14 mm, the length of the capsule 0.23 mm.
There are about 200 eggs per mm, hence a total of ca. 10,000 in the
present ribbon.
Discussion.— Doriopsilla albopunctata (Cooper, 1863: 58), known from
Monterey to Point Eugenia, 27° 51' N (Lance, 1961: 67), is related to
the new species by the white-tipped notal tubercles and the lobed prostate.
98
The latter character was described in Doriopsis fulva MacFarland (1905:
45; 1906: 131), now united with albopunctata (Steinberg, 1961: 57 ff.).
Steinberg describes the color variations of albopunctata as bright yellow,
orange, and warm brown. Cockerell & Eliot (1905: 41) found a deep
chestnut colored living animal of their D oriopsis reticulata which is synony-
mized with albopunctata too. In reticulata 30 rhinophoral leaflets were
indicated, but if this species is identical with fulva, which has 15-20 leaflets
on either side of the rhachis (Marcus, 1961a: 30), Cockerell & Eliot must
have summed them from both sides.
Red and pink tones and blackish bands are not mentioned in the descrip
tions referring to albopunctata and its synonyms. The branchiae are
yellowish white in living albopunctata , as the descriptions of fulva and
reticulata show, and the rhinophores are dark yellow.
The position of the buccal ganglia apposed to the pedal ganglia, and the
absence of a ptyaline gland (Marcus, 1962b: Fig. 18) distinguish D oriop-
silla Bergh, 1880, from D endrodoris Ehrenberg, 1831 (ibid.: Fig. 19).
The location of the anus to the left of the gills cannot be considered
as a generic character of D oriopsilla as compared with a median position
in Dendrodoris. The records referring to the type-species of Doriopsilla do
not agree in this respect, as Bergh (1880b: 319) found the anus symmet
rical between the two posterior gills of a total of four, Pruvot-Fol (1954:
337) and we (Marcus, 1962b: 472) to the left of five and four gills re
spectively. MacFarland described the anal papilla of Doriopsilla fulva (now
called albopunctata ) at the right of the center of the circle of branchiae
(1906: 131); we found the anus surrounded by the gills in the same species.
F i g u r e s 131-134. Tritonia (Candiella) bayeri, sp. nov. 131, living animal, from
color photo (locality 2 ), the white lines drawn black; 132, masticatory border
of jaw; 133, radular teeth; 134, diagram of reproductive organs.
103
lateral teeth occur (Vayssiere, 1912: 94-95; Baba, 1949: Fig. 103 B, b;
106 c), but the combination of a differentiated innermost lateral tooth with
a second one distinguished from the remaining teeth is a novelty.
Only four appendages of the veil, two outer tentacles and two inner
processes, as in T. bayeri, are rare, at least in adult tritoniids. During
growth these appendages as well as the branchial tufts increase in number
(Haefelfinger, 1963: 64; Tardy, 1963: 5). Whether they are short papillae,
pointed lobes, or finger-shaped, is systematically significant only when the
slugs are observed alive; otherwise, the lengths can be widely different. One
species with an undifferentiated innermost lateral tooth (Pruvot-Fol, 1937:
69) and with Duvaucelia-like openings of the intestine and reproductive
organs has four lobes. Four digitiform processes occur in Duvaucelia lineata
(Alder & Hancock, 1848: 191; 1851: Fam. 2, PI. 4 ), a true Duvaucelia.
With Odhner (1963: 51) we separate Duvaucelia and Tritonia as genera
by the position of the anus and the reproductive apertures. The species
classified correctly by Haefelfinger (1963) and Tardy (1963) show that
the distinction between Duvaucelia and Tritonia is sound: Haefelfinger’s
three species belong to Tritonia, and that of Tardy to Duvaucelia. It is
true that Tardy in his comparative notes does not assign the species called
Candiella villafranca by Vayssiere (1901b: 101) to the correct genus, as
it belongs to Tritonia. Furthermore, Duvaucelia insulae Baba (1955: 25,
50) is a Tritonia .
Tritonia bayeri belongs to the subgenus Candiella Gray, 1850, due to
its conic male copulatory organ and the entire rather than bilobed veil
with a small number of appendages.
There are no tritoniids from Western Atlantic warm waters which
deserve a special comparison with the new species. T. palmeri Cooper,
1862 (Cockerell & Eliot, 1905: 33), from the warm temperate area of
California, and the more northern T. festiva (Stearns, 1873) (Marcus,
1961a: 31) do not have the mentioned features of the rhinophores, the
second lateral tooth, and the veil characteristic of bayeri. Also the male
copulatory organ of / estiva (not known of palm eri ) is different.
113
corners, not around the big eyes. No color marks are retained in the
preserved specimens.
The pointed cerata of the preserved slugs measure up to 1.5 mm in
height. They are arranged in five groups: the three first are arches, the two
last simple, unarched. The cerata on the arches stand in more than one
row; their number is respectively 19, 10, 12, 2, 2. The genital opening
lies near the hind limb of the first arch, the renal pore in the interhepatic
space, the anus within the second arch.
The mandible is light yellow, its masticatory border bears around 30
conical, knobbed denticles. The radula comprises 23 teeth; the cusp of
the tooth is prominent, flanked by 4-6 slightly curved denticles. The
diverticula of the digestive gland in the cerata are nodular, the cnidosacs
indistinctly set off from the wall of the ceras.
The penis is everted in one specimen (Fig. 147); it is strong, cylindrical
and curved upward. It ends with a disc (Fig. 148) whose edge is beset
with 10 broad warts, each bearing a tiny spine. There are two further
warts on the surface of the disc, also with spines. Between them lies the
opening of the thick, winding male duct. There is no penial gland.
Discussion.— Edmunds’ useful survey of the penial characters of the
Facalaninae (1964: 26) leads to the Australian genus Austraeolis Burn
(1962b: 120) for the present species, especially if one adds “or warts”
to “minute fleshy filaments” in the original diagnosis. The type-species of
Austraeolis, A . ornata (Angas, 1864) differs from catina by the absence
of cuticular hooks on the six filaments of the penial glans, 9 rhinophoral
rings, about 20 denticles on the masticatory border of the jaw, and 20-22
rows of radular teeth. A . fucia Burn (1962b: 122), whose penial armature
is not known, has six rhinophoral annuli and only the two first arches of the
liver have distinct anterior and posterior limbs. Its radula contains 19 teeth.
149-150. Aeolidiella lurana, sp. nov. 149, dorsal view of living slug,
F ig u r e s
from a colored figure by Sergio Teixeira da Silva, orange pattern stippled;
150, youngest and oldest tooth.
116
rhinophores and A . ? takanosimensis Baba (Risbec, 1956: 31, 32)
with foliated ones cannot be included in Aeolidiella (Marcus, 1961a: 56).
The only Aeolidiella known from the West Atlantic warm waters is
A . occidentalis Bergh (1875b: 397) from St. Thomas. As only three very
badly preserved slugs have been examined, this species remains problem
atic. The radula and the location of the anus, somewhat farther in front
than in most true Aeolidiidae, disagree with the characters known for
Aeolidiella. The double arch of the tooth occurs in this genus and in
Spurilla (Marcus, 1955: Figs. 283, 284). In the latter the double arch
is chiefly brought about by the different length of the denticles, in A eo
lidiella also by widenings of the basal plate. The tooth of A . occidentalis
is rather of the Spurilla- type. When the smooth rhinophores of A e o
lidiella, in some species provided with a spiral groove (Vayssiere, 1888:
109, PL 1, Fig. 8b), are contracted in preserved animals, the circular or
oblique folds that Bergh described of occidentalis may appear. Bergh’s
ventral view of the fore end (1875b: PL 8, Fig. 18) agrees with that of
the present Aeolidiella in preserved condition, not with that of Spurilla
(Marcus, 1955: Fig. 278). The white lines of occidentalis (1875b: Pl.
8, Fig. 9) disagree with Spurilla, whose white dots or spots on the cerata,
head and back will hardly appear as lines in preserved material. In any
case, lurana differs widely from occidentalis.
The 11 valid species which result from Macnae’s list and our additions,
as well as the synonyms (Engel, 1925: 42) or near relatives (Pruvot-Fol,
1954: 429-431) of A . glauca (Alder & Hancock, 1845), were compared
with A . lurana. A more detailed comparison is needed for A . saldanhensis
Barnard (1927: 201) from Southwest Africa; A . takanosimensis Baba
(1930; 1949: 111, 183) from the Pacific coast of Japan from about 35° N
southwards; and A . multicolor Macnae (1954: 36) from False Bay,
South Africa.
Macnae compared his species with A . indica Bergh (1888: 781) from
Mauritius. As he did not see the figure of the radula (PL 78, Fig. 2),
he could not note its less developed wing-like expansions. Evidently,
indica is a distinct species closely allied to multicolor. Risbec (1956:
31) thought that m ulticolor and takanosimensis are one and the same
species. However, the three first groups of cerata of the posterior liver
stand in arches in m ulticolor, while these groups are many-branched in
takanosimensis. A slightly different distance between the nephroproct
and the anus in these species is taxonomically insignificant, as we know
from Spurilla and Berghia (Marcus, 1957a: Figs. 236, 245). As the three
first groups of cerata of the posterior liver in lurana agree with multicolor,
not with takanosimensis, our species approximates multicolor.
The insufficiently described A . saldanhensis differs from m ulticolor and
from lurana by its rounded, not prolonged, antero-lateral angles of
the foot.
117
The radulae of multicolor and lurana differ by the central denticle,
much broader than the adjacent lateral denticles in the former and only
slightly broader than these in the latter. The number of denticles is
different too, but this character is highly variable, at least in certain
species, e. g., A . japonica Eliot, 1913 (Baba, 1949: 183). The tail of
living m ulticolor (Macnae, 1954: PI. 2, Fig. 4), is 1/10 of the total
length, that of lurana 1/4. The foot of multicolor is narrower than the
back, that of lurana broader. The anterior right liver and its left partner
have 6-7 branches in multicolor, five in lurana.
The differences between the species of A eolidiella are slight, and the
same holds for those between m ulticolor and lurana. Nevertheless, as far
as one dares compare an ample material whose average length is 20 mm
(m ulticolor) with a single slug 6.5 mm long (lurana), the differences are
sufficient for a separation.
Z o o g e o g r a p h ic R em arks
Already more than fifty years ago Thiele (1910b: 109) called the
Malacofauna of the West Indies “on the whole well known,” mentioning
that nudibranchs and cephalopods were not considered. In 1906 (p. 132)
Eliot found the knowledge of the West Indian nudibranchs very imperfect.
We think that now Thiele’s opinion can be generalized for the opistho
branchs. The tropical West Atlantic species of the present collection
gathered between Florida and Cabo Orange (4° 46' N) can be grouped
according to their entire range or systematic affinities as follows:
1. Species only known from the West Indies: Oxynoe antillarum, Tri-
dachia crispata, Cyerce antillensis, Umbraculum plicatulum (perhaps
better in group 3 or even in 7), Pleurobranchaea occidentalis and P.
agassizii, H ypselodoris edenticulata, Discodoris hedgpethi, D. purcina,
A phelodoris antillensis, and Phyllidiopsis papilligera.
2. Species of the western Atlantic Ocean and the American Pacific
coast: Berthellina quadridens, Pleurobranchus areolatus, Phidiana lynceus
and, not in the present collection, Tritonia exsulans Bergh, 1894 (Marcus.
1961a: 32), occur on either side of the Isthmus of Panama and are
probably testimonies of the intercommunication between the western
Atlantic and the eastern Pacific Oceans during part of the Pliocene epoch
(Ekman, 1953: 30-38; Weisbord, 1962: 8). Collier & Farmer (1964:
380, 389) recorded Cadlina evelinae and D endrodoris krebsii from several
localities in the Gulf of California. The Tertiary connection of the tropical
opisthobranch fauna in the West Atlantic and East Pacific Oceans explains
the affinities of Onchidella floridana with O. steindachneri from the Gala
pagos Islands. Other cases were previously recorded (Marcus, 1961a: 57).
3. Species from the West Indies and the Bermudas: The original de
scriptions of the Bermudian species are so incomplete that only Elysia
papillosa can be allotted to this group. Elysia tuca seems to be related to
a species from the Bermudas, and Umbraculum bermudense is probably a
young U. plicatulum. Whether H ypselodoris edenticulata and H. acriba
are identical with Bermudian species, cannot be decided yet.
4. West Indian and Brazilian species: This most numerous group evi
dences the unity of the littoral opisthobranch fauna of the warm West
Atlantic Ocean. M icrom elo undata, H ydatina vesicaria (belongs also to
group 5), Haminoea elegans, Lobiger souverbiei (see also group 6),
Elysia evelinae, E. cauze scops (also group 6), Phyllaplysia engeli,
123
Berthella tupala, Pleurobranchaea hedgpethi hamva, Cadlina rumia,
Chrom odoris neona, Siraius kyolis (a new species with Brazilian affinity),
Peltodoris greeleyi, D iscodoris evelinae, D. phoca, D. pusae, Taringa telopia
disa, Platydoris angustipes, D endrodoris krebsii, Bornella calcarata, Phidi-
ana lynceus (belongs also to group 2), Cratena kaoruae, and Spurilla nea-
politana (belongs also to group 5).
5. West Indian species occurring also in the eastern Atlantic Ocean
including the Mediterranean Sea or related to species of this region:
H ydatina vesicaria (also group 4), H aminoea antillarum, Anisodoris w orki
(a new species related to the Adriatic A . m arm orata ), and Spurilla
neapolitana (also group 4).
6. West Indian species which also occur in the Indo-West Pacific Ocean
(the three first) or have systematic relationships with inhabitants of this
region (the three la st): Chelidonura hirundinina, Lobiger souverbiei,
Petalifera ramosa, Felimare bayeri, G odiva rubrolineata, and Austraeolis
catina. Elysia aff. cauze is reported from Hawaii.
7. Species of the West Indies and of most warm seas: Dolabrifera
dolabrifera, A plysia dactylom ela, Stylocheilus citrinus on flotsam, and
the two pelagic slugs Fiona pinnata and Glaucus atlanticus.
8. In all seas with exception of the polar regions: Lamellaria per-
spicua.
9. New West Indian species whose zoogeographic affinities cannot yet
be recognized: Aglaja pusa, Elysia duis, Chromodoris nyalya, A nisodoris
prea, and Tritonia bayeri.
10. Of the nine species collected on the Pacific side of Panama three,
which are also known from the tropical West Atlantic Ocean, were
mentioned in the preceding. Also two further species, Tridachiella diom edea
and Taringa aivica have Atlantic relationships. The former is the represen
tative of a monotypical genus, only known in the Bay of Panama and in
the likewise tropical Gulf of California, and is vicarious for the Caribbean
Tridachia crisp at a, also the only species of its genus. Taringa aivica
belongs to a well characterized genus which is recorded from Brazil,
Florida, and the eastern Mediterranean Sea.
The four remaining Pacific species do not evidence Atlantic relations.
Those of D oriopsilla janaina are distinctly Californian, as the most similar
species reaches its southern limit at 27° 51' N. Two further new Pacific
species belong to the circumtropical and circumsubtropical genera
Chrom odoris and D iscodoris. Their geographic affinities cannot be defined
positively; only the absence of Californian and Caribbean similarities can
be stated. It must be stressed that the opisthobranchs of the tropical
Pacific coast of Central and South America are very little known.
The most enigmatic range is that of Titiscania limacina. It is known
from the Pacific coast of Panama (present collection), the Gulf of Califor
124
nia (Dr. Peter E. Pickens, Tucson, Ariz., coll.), the West Pacific Ocean
(Marshall Islands), the Eastern (Philippines) and the Western Indian
Ocean (Sea of Mauritius). Even if the anatomical differences between ours
and the previously described material resulted in a specific separation, the
Indo-West Pacific and the East Pacific tropical forms would be closely
related. As long as the biology of T. limacina is not known, not even a
reasonable hypothesis can be ventured to explain how Titiscania has over
come the East Pacific barrier.
11. In the present paper, five species are treated that occur on a section
of the coast of southern middle Brazil, but are not recorded from the
West Indian region. The benthic opisthobranch fauna of the upper littoral
is reasonably well inventorized for the referred section, the coast of
Sao Paulo, which, however, comprises less than one tenth of the Brazilian
coast. Therefore a subdivision of the Brazilian opisthobranch fauna,
perhaps an inner tropical local fauna north of Cabo Frio and an outer
tropical one south of it, would be premature. A generalized qualification
of the Brazilian opisthobranch fauna as an impoverished West Indian fauna
with a number of endemic species is justified, because the relatively well
known area lies not far from the southern limit of the western Atlantic
warm-water fauna. This limit can be assumed to lie in northern St.
Catarina, hence farther south than indicated by Ekman (1953: 47).
The mangrove ends at about 27° S, not at Rio, and also large blocks of
stony corals, it is true, no reefs, occur south of Rio. The continental
shelf of St. Catarina is relatively narrow. Therefore the high temperature
and salinity of the Brazilian Current running along the edge of the shelf
reach the littoral of St. Catarina, which is little influenced by conti
nental inflows.
One of the five Brazilian species, Tritonia wellsi, can be expected to
occur also in the West Indian region, because it is recorded from North
Carolina, south of Cape Hatter as. In contrast with this warm-water species
Marionia cucullata is eurythermal; its range extends from the upper littoral
at 38° 50' S to that of Rio de Janeiro. A third zoogeographically in
teresting species of this Brazilian group is Aeolidiella lurana. A single
specimen of it was gathered on a place we frequently visited, 100 m in
front of the beach of Santos. The species is allied to a South African
one from False Bay, and is the first case of a southern transatlantic
relationship in the Brazilian opisthobranch fauna. In that from Florida
a parallel case, G odiva rubrolineata , is known.
R esu m o
Z U SA M M E N F A SSU N G
137
P art II
A bstr ac t
The paper contains a total of 47 species and subspecies collected by
Dr. Peter E. Pickens (University of Arizona, Tucson) in the intertidal
zone of the northeastern part of the Gulf of California. Of the 20 new
forms lnuda luarna, sp. n., and Tayuva ketos, sp. n., represent new genera
of the Dorididae. Several species recently collected on the Pacific coast of
Panama by Dr. Frederick M. Bayer occur also in the Gulf, among them
the Indo-West Pacific neritacean Titiscania limacina. About two thirds of
the present collection are warm-water opisthobranchs; relationships with
the tropical West Atlantic fauna are evident.
I n t r o d u c t io n
Dr. Peter E. Pickens of the University of Arizona, Tucson, sent us 44
species of opisthobranchs for classification, adding three prosobranchs
without shell (Titiscania) or with an internal one (Lamellaria), traditionally
collected and studied together with opisthobranchs. The very well preserved
material was accompanied by photographs of the living animals in natural
colors.
Except for the so-called nudibranchs, the malacofauna of the Gulf of
California is well known. Contrasting with the numerous references to the
prosobranchs (Parker, 1964: 151-156) and the Cephalaspidea (Keen,
1958: 496-499) those to the nudibranchs are poor (Steinbeck & Ricketts,
1941: 538-547). Since then, the range of several Californian species was
extended to the Gulf (Winkler, 1958; Lance, 1961; Farmer & Collier,
1963), and new species were described by Farmer (1963), Collier &
Farmer (1964), and Lance (1962b).
The inventory of the opisthobranchs is certainly still incomplete for
this sea extending over nine degrees of latitude. According to Parker
(1964: 3, 27) “the configuration of the shoreline of the Gulf is so variable
that a tremendous number of ecological niches are present for marine
and intertidal invertebrates. The great diversity of species in any one
environment is the outstanding feature of the shallow water assemblages
in the Gulf of California.”
For a fine set of kodachromes of opisthobranchs, chiefly from the area
of La Jolla, we are indebted to the generosity of Dr. James R. Lance.
To Professor Dr. Frederick M. Bayer we are very much indebted for
revising the language in our manuscript and for reading the proofs.
The material seen by the authors will be sent to the United States
National Museum. Dr. Peter E. Pickens has kept duplicates of most species
in the Department of Zoology, University of Arizona, Tucson.
141
E x p l a n a t io n of L e t t e r in g
L ist of G a st r o po d s C l a s s if ie d in t h e P r e se n t P aper
Order ASCOGLOSSA
Family Hermaeidae
7. Hermaea hillae, spec. nov. Figs. 9-11.
Family Elysiidae
8. Tridachiella diomedea (Bergh, 1894).
Order A N A SPID EA
Family Aplysiidae (Subfamily Aplysiinae)
9. A plysia (Pruvotaplysia) parvula Morch, 1863.
10. A plysia (Neaplysia) californica Cooper, 1863. Fig. 12.
11. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832. Figs. 13-15.
Family Aplysiidae (Subfamily Notarchinae)
12. Stylocheilus longicauda (Quoy & Gaimard, 1824). Figs. 16-17.
Order NOTASPIDEA
Family Pleurobranchidae
13. Berthellina engeli ilisima, subsp. nov. Fig. 18.
14. Pleurobranchus (Pleurobranchus) areolatus (Morch, 1863).
Fig. 19.
Order D ORIDOIDEA
Suborder E u d o r id o id e a
Tribe C r y p t o b r a n c h i a
Dorididae (Subfamily Conualeviinae)
15. Conualevia mizuna, spec. nov. Fig. 20.
Family Dorididae (Subfamily Chromodoridinae)
16. Cadlina evelinae Marcus, 1958.
17. Chrom odoris norrisi Farmer, 1963. Figs. 21-24.
18. Chromodoris banksi sonora subsp. nov. Figs. 25-29.
19. H ypselodoris californiensis (Bergh, 1879). Figs. 30-33.
20. Casella sedna, spec. nov. Figs. 34-37.
Family Dorididae (Subfamily Inudinae, subfam. n ov.).
21. Inuda luarna, gen. nov., spec. nov. Figs. 38-44.
Family Dorididae (Subfamily Doridinae)
22. D oris pickensi, spec. nov. Fig. 45.
Family Dorididae (Subfamily Discodoridinae)
23. D iscodoris mavis, spec. nov. Fig. 46.
24. Diaulula sandiegensis (Cooper, 1862).
25. Taringa aivica timia, subsp. nov. Figs. 47-51.
26. Tayuva ketos, gen. nov., spec. nov. Figs. 52-56.
143
Tribe P h a n e r o b r a n c h i a
Superfamily N o n s u c t o r ia
Family Gymnodorididae
27. N em brotha eliora, spec. nov. Figs. 57-58.
Family Polyceridae
28. Polycera atra MacFarland, 1905.
29. Polycera gnupa, spec. nov. Fig. 59.
Superfamily S u c t o r ia
Family Onchidorididae
30. Acanthodoris pina, spec. nov. Fig. 60.
Suborder P o r o s t o m a t a
Family Dendrodorididae
31. Dendrodoris krebsii (Morch, 1863). Fig. 61.
32. Doriopsilla albopunctata (Cooper, 1863).
33. Doriopsilla janaina Marcus, 1967.
34. Doriopsilla rowena, spec. nov. Fig. 62.
Order D EN D RO NO TO ID EA
Family Tritoniidae
35. Tritonia (Candiella) pickensi, spec. nov. Fig. 63.
Family Dendronotidae
36. Dendronotus nanus, spec. nov. Figs. 64-65.
Family Dotoidae
37. D oto amyra Marcus, 1961.
38. D oto lancei, spec. nov. Fig. 66.
Order AEOLIDOIDEA
Suborder P l e u r o p r o c t a
Family Flabellinidae
40. Coryphella iodinea (Cooper, 1862). Figs. 68-69.
41. Coryphella cynara, spec. nov. Figs. 70-75.
42. Flabellina telja, spec. nov. Figs. 76-82.
Suborder C l e i o p r o c t a
Family Facelinidae
43. Hermissenda crassicornis (Eschscholtz, 1831).
Family Aeolidiidae
44. Spurilla chromosoma Cockerell & Eliot, 1905.
Order SOLEOLIFERA
Superfamily O n c h id i a c e a
Family Onchidiidae
45. Onchidella binneyi Stearns, 1893. Fig. 83.
46. Onchidella hildae (Hoffmann, 1928). Figs. 84-86.
47. Hoffmannola hansi, spec. nov. Figs. 87-95.
S y s t e m a t ic S e c t io n
sc q aw rncL
Navanax inermis (Cooper); male copulatory organ.— F i g u r e s 9-11.
F ig u r e 8 .
Hermaea hillae, sp. nov. 9, diagram of alimentary organs; 10, radular tooth;
11, diagram of reproductive organs.
150
Tiburon Island and Puerto Penasco. Donald A. Thomson, February 25,
1966. “Intertidally among the rocks.”
Further distribution.— From Monterey Bay to southern California (original
locality: San Diego) and Lower California, Mexico (Lance, 1966: 71).
Rem arks. — The photograph (material 1) shows two specimens, 75-mm
long alive, of dark brown to black ground color. One of them is covered
with golden yellow dots, the other with longitudinal stripes of the same
color. The parapodia are edged with a yellow line followed by a row of
bright blue dots. In front of the tail there is a white transverse band, and
the inner side of the caudal flaps is bordered with white.
Dr. Pickens states that the color patterns are various. This is shown
also by the four preserved specimens of material (1), which are more or
less densely dotted or striped. One of them has stripes on the back and
dots on the ventral side. The body of the largest specimen is 40 mm long;
the left caudal flap is 15 mm long and the right one is shorter. In this
animal the head shield is longer than the mantle shield; in the three others
it is shorter.
We give a figure of the protruded penis (Fig. 8) of an animal 36 mm
long and nearly black. There are two flaps (/) of a collar-like dilatation
at the penial base, and the two prostates ( q ) are closely apposed to one
another (Bergh, 1894: 217).
Under the parapodia and on the gill, which is white with brown stipples,
many flat copepods were found. Those known from N. inermis on the
oceanic coast are Pseudomolgus navanacis Wilson, 1935.
The two preserved specimens (material 2) from Puerto Lobos are 60-70
and 37 mm long. They have yellow circular spots on a black ground; those
of the longer specimen are larger than those of the shorter one, but even
in the latter they are bigger than in the material collected in 1965 (1).
In the photograph there are also some blue dots on the border between the
yellow spots. The material collected in 1966 agrees with Bergh’s figures
(1894: PI. 11, Figs. 2-4) based upon Cooper’s original color sketches
(p. 232) better than does that obtained in 1965. One pharynx is everted
as frequently occurs in preserved Aglajidae. The length of the penis is
not correlated with that of the body.
As previously, we use the name Navanax, though a clear separation of
Navanax and Chelidonura is impossible.
F ig u r e s 25-29. Chromodoris banksi sonora, ssp. n ov. 25, living slug from
color photo; 26, gills; 27, jaw elem ent; 28, radular teeth; 29, diagram o f
reproductive organs.
174
ones form two spirals (Fig. 26). Asymmetrical defects occur, though only
nine gills (Farmer, 1963: 84) were not observed in the present material.
The anterior border of the foot is bilabiate. The sole is narrow. The hind
end is pointed and projects backwards.
The yellow labial plates (Fig. 27) bear curved hooklets with bifid tips.
The height of the labial elements reaches 50 the diameter 4 The
radula (Fig. 28) is 6X1.5 mm; it has 150 rows. The half-rows contain
38-41 lateral teeth. A rhachidian pseudo-tooth is present; its base measures
34 /A, the cusp 20-26 fi. The innermost lateral tooth, whose base is
50 n in height, bears a 32 ^ long cusp with 3-7 inner and 7-8 outer denti
cles. In the following lateral teeth the single cusp reaches 80 ^ and bears
7-10 outer denticles. In the outer third to fourth of the half-row the teeth
have no denticles.
The spermoviduct (Fig. 29, h) bifurcates within the female gland mass
(g). The male duct ( d ) begins straight, then forms thin, glandular coils
( q )y has a muscular sheath in the following section (ss), and bears a
broad, penial papilla ( p ) in its ectal course. Two retractor muscles (r)
insert on this papilla. The common atrium (ac) receives the nidamental
duct (ni) beside the ejaculatory duct ( e ). The vagina (v) begins farther
inwards. It leads to the thin-walled spermatheca (s) and the tubular, curved
spermatocyst (sc). These seminal receptacles lie one beside the other at
the inner end of the vagina, hence corresponding to the vaginal arrange
ment described by Odhner (1926: 51). Also the insemination duct (in)
goes out from the inner end of the vagina and enters into the gland mass
near to the ingress of the spermoviduct. A vestibular gland (vn) is
connected with the common atrium by a wide duct.
Material of 1966 .—In life the 15-mm-long slug “lacked the red trim on
the border of the mantle and the red dots on the body. The rhinophores
and gills were red-tipped, however” (Dr. P. Pickens). The preserved animal
was a brilliant purplish red with numerous black and some orange spots on
the notum. The tips of the rhinophores and gills were orange red. There
is a total of only 12 gills; probably the branchial tuft is in regeneration.
The radula agrees with that of the material of 1964-65.
Discussion .—The general character of the color pattern, the radular for
mula, and the shape of the rhachidian pseudo-tooth approximate the
present material to Chr. banksi, whose reproductive organs, however, were
not described. The gills of banksi banksi and banksi sonora differ, as reveal
ed in the preceding description. A second, possibly more weighty difference
refers to the first lateral tooth of b. banksi, figured without denticles
(Farmer, 1963: Fig. 1, j). Perhaps the figure is a back view of this tooth.
Less important differences are: the cusps of the inner lateral teeth of
b. banksi are shorter and bear fewer denticles than in b. sonora; the borders
175
of the branchial and rhinophoral pockets of b. banksi are not colored;
and its ground color is pure white.
Relatively short cusps of the inner lateral teeth as in Chr. b. banksi occur
in Chr. dalli Bergh (1879a: 109), a related species from Puget Sound.
The total of 15 branchial plumes, and the rhachidian pseudo-tooth divided
into two halves (PL 13, Figs. 11, 12) by a longitudinal groove, separate
dalli from banksi sonora; the rhachidian tooth differs also from banksi
banksi.
The radula and the gills of banksi sonora agree well with Chr. punctilu-
cens Bergh (1890a: 163) from the Straits of Florida and the Canary
Islands (Odhner, 1932: 33). Along the notal border, around and on the
rhinophores and gills, and on the hyponotum punctilucens has more black
elements than the present slugs. Its insemination duct originates far from
the inner end of the vagina (Odhner, 1932: Fig. 5), and a vestibular
gland is not developed (1934: 251).
19. Hypselodoris californiensis (Bergh, 1879)
Figures 30-33
References.—Bergh, 1879a: 112; 1894: 181; O’Donoghue, 1926: 211
(literature); 1927a: 90; MacFarland, 1966: 157; Lance, 1966: 72;
Farmer, 1967: 341.
Synonyms.— Chromodoris universitatis Cockerell, 1901b: 79, according
to Cockerell & Eliot, 1905: 37; Chromodoris agassizii Bergh, 1894: 182.
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, March, June,
July, and November 1964-1965. “It is the third most abundant doridid,
after no. 3 (Chromodoris banksi sonora) and no. 4 (Casella sedna), at
Puerto Penasco. It is a rocky intertidal form usually found singly.”—2.
Puerto Libertad, Sonora, Mexico. Joy Cooper, May 30, 1965. One
specimen.— 3. Guaymas, July 1965.
Further distribution.—Monterey to Angel de la Guarda Island (Farmer
& Collier, 1963: 62), Punta Penasco (MacFarland, 1966: 162), and the
Bay of Panama; rare at northern end of range (MacFarland, 1906: 130;
Steinberg, 1963b: 69). Original locality: Santa Catalina Island, California.
Description.— In life, the slugs (Fig. 30) are 32 mm long. They are dark
ultramarine blue with golden yellow spots on the dorsal surface and the
hyponotum, and have a lighter blue, not spotted, sole. In the specimen from
the second locality (Fig. 31) the notum is set off from the foot by an outer
whitish and an inner light blue line. A labeled photo received by courtesy
of Dr. James R. Lance from LaJolla shows one light blue line. In Dr.
Lance’s specimen the golden yellow elements are a few longish streaks;
in front and behind they are shorter, more spot-like. The rhinophores of
the Sonoran slugs are dark blue. The gills are dark blue, sometimes with
yellow dots, in the animals from the first locality; in the specimen from
176
F ig ures 30-33. Hypselodoris californiensis (Bergh). 30, living slugs from
color photo, locality 1; 31, living slug from color photo, locality 2; 32, jaw
elements; 33, radular teeth.
Puerto Libertad they are whitish and tipped with dark blue. In Bergh’s type-
specimen the roots of the branchial leaves were silver-white.
In preservation part of the blue pigment has passed into the surrounding
wet cotton and the label, as was observed in the original material, in
MacFarland’s (1966: 158), and in other blue Chromodoridinae (Mac
Farland, 1906: 129; Anderson & Lane, 1963). The yellow spots of the
smallest present specimens have faded out to white.
The largest of the six slugs from Puerto Penasco is 30 mm long, 10 mm
high, and 8 mm broad; the corresponding measurements of the smallest
animal from this locality are 17.5, 4, and 4 mm. The tentacles are rolled in
(locality 2) or appear as short cylinders (locality 1), which end with a
groove due to the contraction of the tip. The rhinophoral clubs have 17
leaves. There are 8-10 unipinnate gills. Of the semiglobular glands in the
mantle border, frequently mentioned in descriptions of the Chromodoridi
nae, six or more under the hind end of the notum are most conspicuous,
but there are also smaller ones on the level of the rhinophores. The anterior
border of the foot is transversely grooved.
177
The labial plates (Fig. 32) are yellow triangles; their elements are
unicuspidate hooklets. In the material from locality 1 they are about 50 ^
in length and occasionally have secondary points. Near the borders of the
labial plates they are shorter and broader. The latter type of labial hooklets
is the only one in the slug from the second locality. The radula (Fig. 33)
is blue and composed of bifurcated hooks, whose outer or posterior prong
is denticulated, especially in the middle of the half-row. The anterior prong
of the innermost tooth bears a single denticle on the inner side, present in
the material from Puerto Penasco and Puerto Libertad, but not mentioned
in the previous descriptions of califor niensis and the synonymized species.
The outer prong of the hook is shorter than the inner one, especially in the
middle of the half-row. The radula from locality 1 consists of 70 rows
with 93 teeth per half-row. The innermost tooth, 66 /x in height, has cusps
about 30 n long. In the middle of the half-row the inner cusp reaches
57 /jl\ the ten outer teeth are shorter. The small slug from the second
locality, 12 mm long in preservation, had 51 rows and 85 teeth in the
half-row.
Remarks.— The naked mid-line of the radula is limited by the converging
bases of the adjacent teeth. The illusion of a triangular rhachidian pseudo
tooth (O’Donoghue, 1927a: 91) is produced by focusing only the deepest
layer of the radula.
Chromodoris agassizii from the Bay of Panama (Bergh, 1894: 182) has
the same color pattern and radula as calif or niensis. The size of the single
specimen is the same as that of the present slug from the second locality,
and also the labial elements of both agree. Therewith no discriminatory
character remains.
Pruvot-Fol (1951: 152) approximates Chr. macfarlandi Cockerell,
1901, to Chr. universitatis Cockerell, 1901 ( —californiensis). Chromo
doris macfarlandi, however, has a radula of what we call the Chromodoris-
type (MacFarland, 1966: PL 34, Fig. 1-5), hence differs widely from
Hypselodoris californiensis.
In the two similar, though striped, Caribbean species of Hypselodoris,
H. edenticulata (White, 1952) (see Marcus, 1967b) and H. sycilla (Bergh,
1890a: 165), the half-row of the radula has 169-246 and about 290
teeth, respectively. In californiensis the maximum is 132.
20. Casella sedna, spec. nov.
Figures 34-37
Material.— 1. Puerto Penasco, Sonora, Mexico. Rocky intertidal zone.
P. Pickens, July, August, and November 21, 1964. 20 specimens.—2.
Guaymas, July, 1965. “The species is the most common nudibranch at
Puerto Penasco. It is present throughout the year, but may be locally
abundant during August.”
178
Description .—The color alive, from a photo of a 38-mm slug (Fig. 34) is
transparent white on the back and on the sides of the foot. The notum and
foot are bordered by an opaque white zone. That of the notum is thickened
and bears voluminous glands with broad openings. Outward to the white
zone of the notum follow an inner red and an outermost yellow line, while
that of the foot has only a yellow line. The clubs of the rhinophores and
the tips of the gills are red; the rhachis of the rhinophores is opaque white,
and probably also the quite narrow outer side of the gills. The opaque white
and yellow elements are still distinct in the preserved specimens.
The preserved slugs are 12-20 mm in length; the biggest is 11 mm broad
and 8 mm high. The notum is smooth, and so are the slightly protruding
rims of the rhinophoral and branchial pockets. The rhinophores of the
living animal lie 1/4-1/5 of the total length behind the anterior border;
their clubs bear 25-30 foliations. The distance between the gills and the
hind end of the notum is about the same as that between the rhinophores
and the fore end. The branchiae are unipinnate, the anterior ones longer
than the posterior ones. Their bases are arranged in two spiral lines to the
F ig u r e s 34-37. Casella sedna, sp. nov. 34, living anim al from color photo;
35, jaw elem ents; 36, radular teeth; 37, diagram o f reproductive organs.
179
sides of the anus. In the smallest present specimen there are 18, in the
largest 34 gills. Dr. Pickens observed that the animals continuously rotate
the gills back and forth, clockwise and counterclockwise. The border of the
mantle is undulated and about 2 mm broad. Also the edges of the foot are
undulated and projecting. The yellow border line ends where it reaches
the anterior border of the foot which is grooved transversely.
The labial cuticle is strengthened in a yellowish ring-shaped area
consisting of pegs about 65 ^ long (Fig. 35), 4 ^ in diameter, and cleft
into two pointed tips. The radula (Fig. 36) is 4 mm long, 1.6 mm broad;
its formula is 130 X 55.(1).55. The rhachidian pseudo-tooth is 30 ^ high
and has a cusp 16 ^ long; the corresponding measurements of the first
lateral tooth are 53 ^ and 32 p. The cusp of the second lateral tooth is
40 fx long; the cusps in the middle of the half-row measure 55 jjl. The cusp
of the innermost lateral tooth bears 1-2 inner and about 5 outer denticles.
The following teeth have only outer denticles, up to 7 in number, on their
cusps. The outer 25-35 teeth have smooth cusps; the outermost teeth are
small. The stomach and the small left caecum are covered by the digestive
gland. The strongly curved intestine courses first to the left side, then
forward, then to the right and backwards, and then curves to the anus in
the mid line.
The ramified ovotestis lies upon the liver. The quite thin hermaphrodite
duct (Fig. 37, h) opens into a spacious ampulla ( a ), whose outlet merges
into the female gland mass (g). Here the male and female ducts separate.
The spermatic duct emerges from the gland mass far ectally. Rather near
to its origin it widens, becomes glandular and much convoluted ( q ). When
these coils are spread out, they prove to be a prostatic section of the male
duct, not an annexed, massy prostate. The following part ( d ) of the
efferent duct is a thin, muscular tube. Entally it is coiled, ectally it
straightens. Then the muscular sheath is thickened, and the thin duct itself
winds through the sheath functioning as an acrembolic penis ( p ). In the
outermost section of the ejaculatory duct ( e ) the muscular sheath is thin
ner. The male opening ( ma ) is the foremost aperture of the common
genital vestibule. Immediately behind the male opening the thin vagina
(v) begins, describes a loop and opens entally into a spherical spermatheca
( s ) and a longish spermatocyst (sc). So the arrangement of the “vesiculae
seminales” (Odhner, 1926: 53) corresponds to the vaginal type recorded
for Casella and the other genera of the Miamirinae. The insemination
duct (in) goes out from the vagina at some distance from the seminal
vesicles and reaches the gland mass (g) near the origin of the spermatic
duct. The nidamental duct (ni), the outlet of the female gland mass, is
the hindmost canal that opens into the common atrium.
Discussion .—A slug shaped like a Chromodoris, but with a strongly undu
lated edge of the mantle and foot (Engel & Eeken, 1962: 25) can easily
be allotted to Casella. But a moderate undulation as in the present species
180
differs only by degree from what occurs in certain species of Chromodoris,
such as Chr. nona (Baba, 1955: PL 9, Fig. 23). The decreasing of the
denticles on the outer radular teeth belongs to the characters of the
subfamily Miamirinae, in contrast with the Chromodoridinae (Odhner,
1926: 53). If the pectinate outer teeth of a Chromodoris (Farmer, 1963:
Fig. 1, b) are compared with the smooth ones of the present Casella, the
difference is obvious. However, smooth outermost teeth occur in Chromo
doris too (ibid., Fig. 1, g), and denticulate ones in Casella (Vayssiere,
1912: 58; O’Donoghue, 1929: 825).
The tips of the labial rodlets, entire or cleft, and the presence or absence
of a rhachidian thickening or a pseudo-tooth are variable characters in
Casella.
So high a number of gills as in the present species is exceptional in
Chromodoris. But this character is not reliable; of Casella atromarginata
from the Red Sea there are indications of 15 (Vayssiere, 1912: 57),
25 (White, 1951: 248), or 22-32 gills (Gohar & Abul-Ela, 1959: 49),
and in specimens from the eastern Indian Ocean 12 (Bergh, 1905a: 162)
and about 30 (Risbec, 1956: Fig. 35) were recorded.
The color of the dorsal surface of the mantle varies in certain species of
Casella. In the original material of C. rufomarginata Bergh (1890b: 943)
from Amboina it is darker than the border-line, in a specimen from the
Red Sea lighter (Engel & Eeken, 1962: 25). Therefore we separate
C. sedna from the other species by a short discussion, not by an inclusion
in our previously published key of the genus (Marcus, 1960b: 904), where
C. rufomarginata was separated according to its original description. In
that species as well as in C. atromarginata, atromarginata pallida, and foxi
only the notum, not the foot, bears a colored border-line. In C. cincta and
C. sedna both are colored, blue in cincta, red and yellow and yellow in
sedna. C. cincta has 11, sedna 17 gills on either side. C. rubra is uniformly
yellowish red without border-lines.
The hitherto known range of the species of Casella extends from Lake
Timsah, 81 km north of Suez, to middle Japan, New South Wales, the
Solomon Islands, and Tahiti.
Inuda, gen. nov.
Diagnosis.—Dorididae with cleft hooks on the labial cuticle, multidentate
radula with bicuspid rhachidian tooth and denticulate lateral teeth.
Prostate well developed; male copulatory organ acrembolic, with spines;
vagina unarmed.
Type species.— lnuda luarna, spec. nov.
Apart from the shape of the rhachidian tooth, Inuda resembles Cadlina
but the well-developed prostate makes it impossible to allocate these
genera in the same subfamily.
181
We also pondered an affinity of lnuda with Carminodoris Bergh, 1889,
and therewith its systematic place in Odhner’s Baptodoridinae (1926: 54).
But the radula and the labial armature are less differentiated in all species
of Carminodoris (Bergh, 1889: 818; 1905a: 111; Risbec, 1928: 109;
1930: 283; 1937: 159; 1953: 47), so that it appears artificial to assign
lnuda to the same subfamily.
It requires a subfamily of its own, Inudinae, subfam. nov., to be placed
between the Chromodoridinae and the Cadlinellinae.
21. lnuda luarna, gen. nov., spec. nov.
Figures 38-44
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens. Found under
boulders in a large tide pool, together with Taringa aivica timia.
The two specimens at hand were preserved together with the mentioned
discodoridine which differs externally from I. luarna by its notal sculpture.
Description.—The preserved animals are whitish mottled with brown
(Fig. 38), due to numerous dots of dark brown pigment (Fig. 40, is)
lying in different layers of the notum. The rhinophores are brown, and
also the gills are darker than the notum. The measurements are: 27 (20)
mm long, 20 (18) mm broad, 11 (14) mm high. Measured over the back
the lengths are 37 (40) mm. The foot is 20 mm long, 8 mm broad, its
anterior border is bilabiate, slightly concave in the middle (Fig. 39). The
antero-lateral corners of the foot are distinct.
The rather stiff notum is smooth; the vertical bundles of spicules ( cu)
do not project over the surface, but produce some slight and irregularly
distributed elevations of the skin. The spicules are up to 0.6-0.8 mm
long and 30 ^ in diameter. Most of them lie in the deeper layers of the
body wall. The tentacles are broad and grooved on their outer side. The
rhinophoral clubs bear about 16 leaves. There are seven tripinnate gills.
The rims of the rhinophoral and branchial pockets are smooth.
The labial cuticle bears a broad yellow ring of stratified hooks (Fig.
41). These are cleft for half their length, up to 50 ^ long, and 10 ^
broad. The radula (Fig, 42) is 3.1 mm long, 1.8 mm broad. It comprises
145-150 rows, has a well-developed rhachidian tooth and 50 teeth per
half-row. The rhachidian tooth (r) is 50 ^ in height, 16 ^ in breadth,
and bears two 16 /x long cusps. Its base is tripartite. The innermost
lateral tooth has a secondary cusp on either side of the middle cusp
(50 fx). In the following teeth there is no inner secondary cusp, and the
outer one diminishes in size, constituting a denticle. Behind it smaller
denticles appear, increasing in number outward to about ten. In the different
rows the denticulation varies much. The length of the main cusp increases
outwards to 65 ^. The outermost teeth are imperfectly developed. The
stomach is covered by the intestinal gland.
182
F ig u r e s 38-44. Inuda luarna, gen. nov., sp. n ov. 38, sculpture o f notum ; 39,
fore end in ventral view ; 40, transverse section o f notum , clarified; 41, labial
rodlets; 42, radular teeth; 43, diagram o f reproductive organs; 44, tip o f penis.
The exit of the ampulla (Fig. 43, a) divides into the short inner
oviduct (o) which merges into the female gland mass (g), and the male
duct. The latter begins as an inner prostatic part ( q ) forming broad folds,
and continues as a tubular portion with high cells and large nuclei. The
muscle layers of the following ciliated efferent duct ( d ) separate into an
outer and an inner layer, the former is a sheath 0.7 mm long. The ectal
section of the duct bears about 20 longitudinal rows of spines up to
18 fji long, which end where the ejaculatory duct coalesces with the
atrium (ac). In Fig. 43 about half the penis ( p ) is everted, as is shown by
the spines on its outside and inside. Some of the spines are slightly
curved (Fig. 44).
The vagina (v) begins in the fundus of the common atrium (ac); it
is folded but not cuticularized. The spermatheca (s) and the spermatocyst
(sc) are both connected with the vagina by long canals. This arrangement
corresponds to that in Cadlina laevis (Odhner, 1939: Fig. 13); Odhner
calls it the semiserial type. The nidamental duct (ni) opens (aw) imme
diately behind the male aperture (ma).
22. Doris pickensi, spec. nov.
Figure 45
Material.— 1. Puerto Lobos, Sonora, Mexico. P. Pickens, April 16, 1965.
“Fairly common under boulders in the intertidal zone; also found in the
spring at Puerto Penasco.” We received nine specimens together with one
of Taringa aivica timia.—2. Puerto Penasco, Sonora, Mexico. Mary Anne
Hill, February 5 and March 5, 1966. February: “These are probably a dif
ferent color phase of Doris pickensi.” March: “A very small orange form
(7 mm) found among some gorgonians.”
Description .—In the photographs, the animals are 25 mm long, white
with a translucent border of the notum and more opaque white notal
papillae (Fig. 45, A). Dr. Pickens states “that some specimens have tinges
of brown or are speckled with brown.” The rhinophores, gills, and the
anterior border of the foot are light yellow. Evidently also Dr. Lance
collected this species in the Gulf, to judge from a kodachrome that he
kindly sent us.
The largest, much contracted, specimen is 23 mm long, 15 mm broad.
The breadth of the lateral borders of the notum is about 3 mm. The gills
are 5 mm from the hind end. The length of a smaller, straighter animal
is 12.5 mm, its width 8.5 mm, and the height 3.5 mm. The hyponotum is
2 mm broad; the foot measures 11X4.5 mm.
Of material from collection 2 we have three specimens collected in
February. The largest measures 19 X 13X7 mm. In the color photograph
the slugs are a gelatinous light orange with brick red rhinophores and gills.
The specimen collected in March was 7 mm long alive and preserved.
184
The notum bears small, mamillary warts whose tips are about 300 ju,
distant from one another in the central parts of the notum, 200-250 ^ on
the sides. Each papilla has a big spicule (Fig. 45, B), 400 ^ long and 35 ^
in diameter, and several smaller ones surrounding it. The spicules may
project on the surface. A mat consisting of thick spicules lies in the back,
extending into the hyponotum.
The tentacles are short, broad and grooved on their outer side. The
clubs of the rhinophores have 10 leaves. There are 13-14 unipinnate
gills. As far as the rims of the much contracted rhinophoral and branchial
pockets can be seen, they are smooth, not papillose. The anal papilla is
high and lies somewhat behind the center of the branchial circle. The
anterior border of the foot is bilabiate, sometimes notched, sometimes
entire.
The labial cuticle is smooth. The radula of a smaller animal has 35,
that of the largest 40 rows. The half-rows contain 50 and 64 teeth res
pectively. All teeth are hook-shaped, without denticles; their size increases
in the two inner thirds of the half-row, decreasing in the outer third. The
innermost teeth converge over the narrow rhachis. The salivary glands are
quite short and broad sacs which envelop a great part of the pedal com
missure. The free stomach is large; the roundish caecum lies beneath it.
F ig ure45. Doris pickensi, sp. nov. A , sketch of living slugs from color photos
1 and 33; B, transverse section of notal papillae; C, diagram of reproductive
organs.
185
The narrowed outlet of the sausage-shaped ampulla (Fig. 45, C, a)
enters the female gland mass (g) from which the male duct emerges. It
begins narrow but widens immediately and becomes prostatic ( q ). The
following portion ( ss ) runs winding within a muscular sheath, which is
entally continued into a retractor (r) and ectally opens into the atrium.
The efferent duct does not form a papilla. From the atrium the vagina (v)
courses apposed to the male sheath and leads first to the large spermatheca
(s) and then, with a circular loop, to the small spermatocyst (sc). Near
the spermatocyst the insemination duct (in) connects the seminal recepta
cles with the glandular oviduct (g). The nidamental oviduct (ni) opens
(aw) immediately behind the male aperture.
The species is named for Dr. Peter E. Pickens.
Discussion.— Though the acrembolic ejaculatory duct may project into
the male atrium (“preputium,” Bergh), it does not form a true penial
papilla (glans) as in Archidoris. The absence of a prostatic part of the
efferent duct is the principal character of Austrodoris; a muscular sheath
of the efferent duct occurs in Austrodoris and in Doris . As Hoffmann
(1938: 1070) said, thick and short salivary glands ( Austrodoris) and
long, band-like ones (Doris) cannot be used as generic characters, because
in A. nivium Odhner (1934: 269) both types were observed in one and
the same animal. In African material of D. verrucosa, Bergh (1907: 47)
found the salivary glands nodular and shorter than usually in this type
species of Doris. Papillae on the rims of the rhinophoral and branchial
pockets, which protect them like valves, are specific, not generic characters
(Eliot, 1903: 557).
Doris echinata O’Donoghue (1922a: 149) has brown spots on the
opaque white notum, but differs from Doris pickensi by six gills, four
anterior unipinnate and two posterior bipinnate ones, and 16-18 radular
rows of 13-15 teeth per half-row. It was found at San Juan Island (Wash
ington) and Vancouver Island (British Columbia), and called Doridigitata
maculata O’Donoghue (1926: 206) and Doris odonoghuei Steinberg
(1963a: 63). The reproductive organs of this species are not known, so
that its true generic place is not settled. In his family Doridigitatidae
O’Donoghue (1927a: 87-89) included a white slug from Laguna Beach,
for which he created the new genus Glossodoridiformia (species: alba). Its
notum is villous, there are five unipinnate gills, 17-18 radular rows, and
25-26 teeth per half-row.
Doris (Asteronotus) alabastrina Cooper (1862: 204) from San Diego
is alabaster white, opaque, and has 12 unipinnate gills. These characters
agree with the present species, while Conualevia alba Collier & Farmer
(1964: 383) which occurs abundantly at Newport Bay, hence one degree
north of San Diego, has eight tripinnate gills. But the description of
Cooper’s species is so poor that its name, indeed a “nomen dubium”
(Steinberg, 1961: 59), cannot be used for the present species.
186
23. Discodoris mavis, spec. nov.
Figure 46
Material.--Puerto Penasco, Sonora, Mexico. P. Pickens, March 5, 1966.
The slug was found crawling under a granite boulder. “When pulled olf
a rock it has a tendency to fold the edge of the mantle ventrally, so that
it covers the foot” (P. Pickens).
Description .—The living specimen was 24 mm long, 17 mm broad. Its
color was orange pink with scattered brownish spots (Fig. 46, A), denser
towards the sides. The color has completely disappeared in the preserved
specimen; only some dark rings around the openings of the cutaneous
glands are recognizable.
F ig ure 46. Discodoris mavis, sp. nov. A, pigment pattern of living slug from
color photo; B, spicule pattern on border of same; C, section of notum; D, end
of half-row of radula; E, diagram of reproductive organs.
187
There are no traces of spicules, but the color photo shows small
tubercles with white tips (Fig. 46, B). These were evidently supported by
vertical spicules. Cutaneous glands lie between the tubercles (Fig. 46, C).
The preserved specimen, that of the color photo, measures 19 X 13X8
mm. The underside is pigmentfree. The thin edge of the notum is curled
inwards. The foot is 7 mm in width and projects beyond the 5-mm-broad
hyponotum. The tentacles are digitiform; the anterior pedal border is
bilabiate, and the upper lip notched. The rhinophores have a white tip,
a pink shaft, and a brown club with 17 foliations. The rim of the
rhinophoral pocket is smooth. The 8-10 orange-pink gills are multi-
pinnate; the border of their pit is smooth.
The labial armature consists of two areas with stratified rodlets up to
100 n high. The radula measures 2.6 X 1.6 mm and comprises 25 rows.
Each half-row contains 15 marginal lamellae and 21-22 hooks (Fig. 46, D),
up to 230 /A in height. The innermost hook is slightly angled on the
rhachidian side under the cusp. The edges of the lamellae are irregularly
dentated, probably worn, not provided with distinct denticles.
The ampulla (Fig. 46, E, a ) is long and curves around the prostate,
which is composed of three parts. The acini of the inner part ( q ) have
small cells. Apposed to it lies the outer part ( qu ), whose acini are big-
celled. Then follows a smooth tube ( d ) which begins glandular, as third
prostatic section of the efferent duct, and continues muscular. The sheathed
muscular part (ss) of the efferent duct leads to a rather long penial
papilla ( p ) which projects from the male atrium (am ).
Behind the atrium lie the vagina (v) and the protruded nidamental
opening (aw). The vagina is muscular and seems to be coated with a thin
cuticle; its ectal part bears a large gland (vn) lined by a high epithelium
with basal nuclei. Entally the vagina narrows and opens into the longish
spermatheca (s). Near its entrance, but independent from it, the in
semination duct (in) leaves the spermatheca. The spermatocyst (sc) has
its own connection with this duct, which merges into the gland mass (g)
near the nidamental duct (ni).
Discussion.— D. mavis is related to D. heathi MacFarland, 1905, from
Vancouver Island to northern Lower California (Farmer & Collier, 1963:
62; Farmer, 1967: 341) and to D. pusae Marcus from the coast of S. Paulo,
Brazil, and from Miami, Florida. D. heathi is yellow ochre to raw umber
(MacFarland, 1966: 193), D. pusae orange or pinkish red. The darker
flecks occurring in both species are especially dense in the mid-region.
The skin of D. heathi (Marcus, 1961a: Fig. 62) is rather similar to that
of mavis, but the spicular tubercles are larger and more densely set in
heathi. The skin of D. pusae (Marcus, 1955: Fig. 154) has mainly
horizontal spicules, so that it is less similar to the skin of D. mavis.
The labial rodlets are about 42 ^ long in D. heathi, those of D. pusae
reach 60 In D. heathi the number of radular rows, 20-22, is inferior
188
to that of D. mavis; in D. pusae it is 21-26. The number of marginal
lamellae and lateral hooks per half-row does not furnish palpable dif
ferences; D. heathi has 9-17 lamellae and 20-31 hooks; D. pusae 8-18
lamellae and 15-27 hooks. The denticle-shaped shoulder of the innermost
hook and the irregular denticles of the lamellae noted for D. mavis occur
in D. pusae as well.
The prostates of all three species agree; the penial papilla of D. mavis
and D. heathi is longer than that of D. pusae. In large specimens of the
latter the duct of the vaginal gland is provided with cuticular spines; the
gland of D. heathi is angled, and so its shape is farther distant from that
of D. mavis than that of D. pusae.
24. Diaulula sandiegensis (Cooper, 1862)
References.—MacFarland, 1906: 122; 1966: 190; Marcus, 1961a: 18.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, February and
May, 1965. “Rare. Collected in the rocky intertidal.”
Further distribution.—From Lower California, Cape San Lucas (Lance,
1961: 66) to the Aleutian Islands and northern Japan. Original locality:
San Diego.
Descriptive notes.—The present specimens are small, 20 and 12 mm in
length. They have a series of brown rings on either side of the back.
Many paler rings are scattered chiefly on the sides of the white notum,
but also in the middle. The radula of the larger slug has 17 rows with
29 teeth per half-row.
Probably the “large, seal-brown nudibranch” from the western coast of
the Gulf (Steinbeck & Ricketts, 1941: 543) is this species. According
to Lance (loc. cit.) D. sandiegensis is uncommon intertidally, common
subtidally to 37 m.
25. Taringa aivica timia, subsp. nov.
Figures 47-51
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, January, March,
August 1964, January 15, and April 16, 1965. Generally found under
rocks in large tide pools together with Inuda luarna, Doris pickensi, and
Doriopsilla albopunctata.
Description.—The photographs of three of the five specimens (collection
No. 20, 20 mm alive; 61, 62) show a purplish brown ground color of the
notum with light marks around the back, the branchial pocket and the
bases of the papillae. These are brownish yellow, the rhinophores dark,
their tips white. In preservation the hyponotum and the sole have dark
brown flecks, especially numerous along the dorsal border of the foot.
In another photograph (no. 19, one specimen 35 mm long) the animal
is brownish pink with brown dots, light stellate marks around the bases of
189
the light orange papillae, light violet gills, dark brownish violet rhinophores
tipped with white, and with white papillae on the rims of their pockets.
Some of the notal papillae appear branched. In a third photograph (no.
14, six specimens) the notum is yellowish grey (Fig. 47). The rhinophores
are dark brown, the gills of the ground color, and the cutaneous papillae
yellowish. Around their bases they have dark brown blotches.
Measured over the back, the largest of the preserved specimens, one
of lot no. 14, is 70 mm long, its linear length is 35 mm, the height 26 mm,
the breadth 43 mm; that of the hyponotum is 10 mm. The much folded
foot measures 25 X 20 mm. The brown dots of the hyponotum are pre
served. A 17-mm slug (no. 1) has uniform caryophyllidia. Evidently it
had been quite white alive, because it was put into the vial with Doris
pickensi. In all other specimens the notum bears a few long, smooth
papillae between the uniform, densely set caryophyllidia. The papillae are
much contracted in the preserved specimens and only sometimes bicuspid.
F ig ures47-51. Taringa aivica timia, ssp. nov. 47, sketch of living slug from
color photo 14; 48, radular tooth from middle of row; 49, marginal tooth; 50,
diagram of reproductive organs, prostate simplified; 51, penial cuticle.
190
The tentacles are pointed, sometimes widened under the tip. The rhino
phoral clubs have 16 leaves. The rim of their sheaths bears about eight
lobular papillae. There are six multipinnate gills; the border of the
branchial pouch is undulate. The anterior border of the foot is bilabiate
and notched.
The labial cuticle is smooth. The radula of a 29-mm slug, belonging to
lot no. 20, comprises 29 rows with 48 teeth per half-row. The 44 inner
teeth are hook-shaped, the four outer ones pectinate. The inner teeth
increase in size to the 30th (Fig. 48), continue high to the 40th, and
decrease to the 44th. The cusp of the hook-shaped teeth bears a minutely
knobby ridge on the outer side, which corresponds to the distinct denticles
in aivica aivica. The radula of the largest specimen is 7 mm long, 6 mm
broad, has 41 rows, and 75-76 teeth per half-row. Of these the 70 inner
ones are hook-shaped, the 5-6 outer ones pectinate. The inner teeth have
only a ridge on the outer side of the cusps, but no knobs. The 17-mm slug
had no knobs at all; its radular formula is 30X45.0.45. In all examined
radulae the flat, pectinate teeth (Fig. 49) have a broad uppermost spine,
and the rest of the edge is feathery.
The prostate is a massy dilatation of the efferent duct; in the diagram
(Fig. 50, q) it is simplified. The distal part of the duct ( d ) has a folded
epithelium. The penial papilla is coated with a beaker-shaped thick cuticle
(Fig. 51). The wall of the male atrium ( am ) bears spineless papillae.
Between the male atrium and the glandular nidamental duct ( ni ) the
narrow vagina (v) begins tubular, not bulbar. The vaginal epithelium is
thrown into longitudinal folds. The entrance of the vagina into the
spermatheca (s) and the exit of the insemination duct (in) are neighboring,
but independent from one another. The entrances of the inner oviduct (o)
and the insemination duct into the female gland mass (g) almost meet.
Discussion.—The present animals represent a new subspecies of T. aivica
Marcus, 1967. Both have long papillae and small caryophyllidia on the
notum. The denticles of the lateral teeth are strong in aivica aivica, and
its marginal teeth have not the broad spine of aivica timia. This spine
resembles that of the pectinate teeth of Dictyodoris tesselata Bergh
(1880b: PI. C, Fig. 11), a doubtful doridid, whose prostate is not known.
The muscular bulb of the vagina with cuticularized folds in aivica aivica
and the absence of spines on the atrial papillae of a. timia might be due
to different states of reproductive development. The shape of the cuticular
coat is different in both subspecies. Alive, the color of the previously
described form is yellowish mottled with grey.
Tayuva, gen. nov.
Diagnosis.—Discodoridinae with pointed tentacles, labial plates with
rodlets, hook-shaped radular teeth, stout penial papilla, large vestibule
(atrium) stiffened by spicules and lodging the penial papilla and the
191
vaginal aperture. Nidamental opening independent from that of atrium.
Type species.— Tayuva ketos, spec. nov.
The anatomy of the present species evidences Bergh’s terms “preputium
or penial sac” as inappropriate for the terminal section of the male duct.
There are two genera of the Discodoridinae, Phialodoris and Audura,
with aberrant outer genital organs, but neither of them can receive the
present species.
Tayuva ketos is one more example to reveal that the classification
of many doridids is impossible without dissection of the reproductive
organs; the extraction of the radula is often insufficient.
26. Tayuva ketos, gen. nov., spec. nov.
Figures 52-56
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, November 2,
1963.—2. Puerto Lobos. P. Pickens, April 16, 1965. Under rocks in
large tide pools.
Description.—In a photograph of the specimen from locality 1 (Fig. 52)
the notum has brown blotches on a whitish ground, and between them
single brown papillae. These are more numerous towards the sides where
the blotches are smaller. The rhinophores and gills are brown. The blotches
of the slug from locality 2 are greyish brown, and several especially large
ones occur in the marginal zone. The rhinophores are brown with light
tips, the gills greyish.
The specimen from the first locality was 47 mm long alive; preserved,
it is 30 mm in length, 22 mm in width, and 9 mm in height; the second
slug measures 50, 40, and 8 mm. The sole of the smaller animal is
18 mm long, 9 mm broad. The hyponotum of both specimens is white with
small dark flecks. Those on the light sole are larger.
The papillae (Fig. 53) of the smaller slug are rather pointed, those of
the larger, evidently more contracted animal, blunt. The tips of the
papillae are ciliated. The spicules, which are up to 350 ^ in length and
30 /x in diameter, do not reach the tips of the papillae.
The white tentacles are 2.5 mm long, carrot-shaped, and have some dark
spots. The rhinophores have 24 leaves; the borders of their pits and that of
the branchial pocket bear papillae of the common type. There are six
pluripinnate gills. The anterior border of the foot is bilabiate and notched
in the middle.
The yellow labial cuticle bears up to 100 ^ long striped rodlets (Fig.
54) which are 10 /x in diameter at their thickened free end. The radula
of the smaller animal measures 2.7 X 2.4 mm, that of the larger one
4.0 X 4.0 mm. There are 24 (locality 1) and 29 rows (locality 2), and 38
and 41 teeth per half-row. The teeth (Fig. 55) are hooks whose cusps are
up to 220 jui long in the middle of the half-row and smaller to the sides.
192
The blood glands are dark. The oesophagus enters the completely free
stomach from the ventral side. The stomach is large; its dorsal, strongly
folded and glandular wall projects into the gastric lumen. The intestine
leaves the stomach on the left side, in front and dorsally, and curves to
the right and backwards. The pylorus conceals the small caecum.
Near the outlet of the winding ampulla (Fig. 56, a) the short inner
oviduct and the spermatic duct ( d ) separate. The former merges into the
female gland mass (g), the latter becomes prostatic. The longish ental
portion of the prostate ( q ) is soft and consists of radially disposed
glandular sacs. The ectal globular portion ( qu ) is more compact. Farther
outwards follows the efferent duct ( d ) containing sperm in its folded
lumen. The duct ends with a stout, crooked penial papilla (p), which
projects into the large vestibulum ( ac ). The vestibular wall is stiffened by a
layer of longitudinal spicules, 3 mm in length and 10 jjl in diameter.
F ig u r e s 52-56. Tayuva ketos, gen. nov., sp. nov. 52, living slug from color
photo; 53, spicules and m elanophores in notum papillae; 54, labial rodlet; 55,
radular tooth from m iddle o f half-row ; 56, diagram o f reproductive organs.
193
The vagina (v) whose epithelium is folded begins in the ental third of
the atrium. Quite near to its entrance into the spermatheca (s’) the insemi
nation duct (in) emerges from the latter, runs curved towards the female
gland mass (g), and receives the canal from the spermatocyst (sc) short
before it reaches the mass. The disposition of the sperm-storing vesicles
is of the serial type (Odhner, 1926: 51). The insemination duct opens
into the gland mass near to the entrance of the inner oviduct. The
aperture (aw) of the outer oviduct, the outlet of the gland mass, is inde
pendent from that (ma) of the penial and vaginal atrium.
27. Nembrotha eliora, spec. nov.
Figures 57-58
Material.— 1. Puerto Lobos, Sonora, Mexico. Paula Vreeland, April 16,
1965. Three specimens in a small boulder-strewn area in the intertidal
zone.—2. Same locality. Alex Kerstitch, February 25, 1966. One specimen
subtidally in about 10 m of water, on a rock.
Description .— (Material 1). A photograph shows that the slugs are black
with many green and blue longitudinal stripes (Fig. 57). To the sides of
the median blue line bordered with black, the green lines form two triangles
between the black rhinophores. The paramedian green stripes end in front
of the gills which are black with a green line along the efferent and afferent
vessels; the more lateral green stripes are confluent behind the gills. Each
of the two next pairs of green stripes is subdivided by a black streak which
begins in the anterior third. The most ventral stripes are undivided and
F ig u r e s 57-58. Nembrotha eliora, sp. nov. 57, living slug from color photo;
58, half-row o f radula.
194
uninterrupted from the anterior to the posterior end. In preservation the
blue color has disappeared from the black ground, but the green is pre
served. Ten light and 15 black stripes are distinguishable. The light
stripes unite on the back of the tail which is smooth, not cristate.
The two present specimens are 30 and 12 mm in length, 10 and 8 mm
in height, and 6 and 4 mm in width. The sole is folded. The anterior
border of the notum is somewhat thickened. The border of the rhinophoral
pits is light in one, dark in the other specimen. The clubs have about 22
leaves. The broad black tentacles have a light outer groove. There are five
and seven pluripinnate gills. Extra-branchial processes (as in Tambja
Burn, 1962) are not developed. The anus lies between the gills, the
gonopore at the end of the anterior fourth of the body. The anterior pedal
border is green, thick, neither grooved nor notched. The sole is white,
its border black.
The cuticle of the labial disc is folded and smooth, not composed of
rods. The buccal mass is 6 mm long in the larger specimen. The radula
(Fig. 58) is yellowish, except for the pale first lateral tooth. There are
15 rows of teeth and four outer lateral teeth per half-row. The rhachidian
tooth is a transversely striate plate, divided in the middle, 0.35 mm in
width and 0.29 mm high. Its anterior edge is thickened and smooth. The
inner lateral tooth is 0.8 mm or more in length, its pointed cusp 0.6 mm;
a strong spur projects in the middle. The outer teeth decrease in size
outwards; the first has a strongly crested inner edge.
In order to keep one specimen intact, the larger one with everted genital
atrium was dissected, because only the male organs are systematically
important. The spermoviduct which leaves the large and wide ampulla
divides within the female gland mass as in N. morosa Bergh (Macnae,
1958: Fig. 14). The prostatic part of the efferent duct begins with a thin
layer of muscles, its ciliated epithelium is high; its wide lumen contains
sperm. Farther outward the muscles become thicker and the lumen narrows;
the diameter of the duct continues the same. In the next section the duct
is thicker. Here its muscle layers are divided into that of the duct proper
and that of its sheath. The terminal part of the duct in the sheath bears
cuticular hooks and opens into the fundus of the atrium, whose epithelium
is greenish. In the present everted vestibulum the occurrence of a vestibular
gland, as indicated for morosa by Bergh (1907: 70), not by Macnae
(1958), could not be examined.
(Material 2). The living animal was about 80 mm long (Dr. P. Pickens).
Preserved it is 40 mm long, 20 mm high, and 12 mm broad. In the photo
graphs the ground color is green. The lateral crests which separate the
notum from the sides and unite behind the gills into a caudal crest are more
salient than in the specimens from lot no. 1. Their color is yellow, trimmed
with a thin black line. Also the yellow bulges on the notum which
correspond to the stripes of material 1 are seamed with black. The tentacles
195
are yellow; the clubs of the rhinophores violet, their sheath yellow with a
black basal rim. Two dark blue spots appear behind the rhinophores. The
gills are violet, their inner (afferent) and outer (efferent) vessels yellow.
The margin of the foot is broad and bright ochre.
Discussion.—This is the first Nembrotha from the Pacific coast of the
Americas. The two American Atlantic species are N. gratiosa Bergh
(1890a: 172) and N. divae Marcus (1958b: 47). The latter, found at
Cabo Frio (22° 57' S, 42° 01' W), in life is scarlet with white dots and
white gills, and therefore differs widely from the present species, as it does
also by its radula. The color of N. gratiosa, from off Key West in 52 m
depth, is similar to that of specimen 2 from Puerto Lobos, but the radulae
are different. The rhachidian plate of N. gratiosa is undivided, its anterior
edge is thin, and there are only three outer lateral teeth. The color of the
slugs of lot no. 1 differs from that of N. gratiosa. Among the four species
recorded from Japan (Baba, 1960: 72-73) N. sagamiana Baba (1955: 16,
44) has a similar divided rhachidian tooth; due to its tuberculate notum,
Burn ascribes sagamiana to Tambja (1962a: 98). N. eliora is similar to
N. morosa Bergh (1877: 457), recorded from South Africa to the Philip
pines, in Macnae’s concept of its synonymy (1958: 360). This species has
up to five light stripes and an undivided median tooth. Though the cleft tip
of the first lateral or intermediate tooth of morosa is absent in capensis,
Macnae united it with morosa; the simple cusp of capensis (Bergh, 1907:
PI. 12, Figs. 10-12) is much shorter than in eliora.
Bergh’s term (1883: 165) “prostata discreta nulla” (“gesonderte
Prostata fehlt” Thiele, 1931: 425) refers to the type-species ( nigerrima
Bergh) and others, such as morosa (Macnae, 1958: 361), divae Marcus,
1958, and eliora. In their intercalary prostates the widening of the prostatic
section of the efferent duct or an equal diameter of the efferent duct
throughout its length depends on its physiological state. An independent
prostatic gland annexed to the efferent duct as in N. lineolata (Eales, 1938:
Fig. 16) is a different type and can lead to a subdivision of the genus.
28. Polycera atra MacFarland, 1905
References .—MacFarland, 1906: 142; 1966: 115; Odhner, 1941: 17.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, May 2, 1965 and
March 5, 1966.
Further distribution.—From Tomales Bay, Marin County (Marcus, 1964:
131), to Los Coronados Islands, Lower California (Lance, 1961: 66).
Original locality: Monterey Bay.
Remarks .—The pale grey ground color bears narrow dark stripes in
MacFarland’s specimens (1966: PL 18, Figs. 2 & 3). In a photograph
(no. 58) which we owe to Dr. Lance, the dark stripes are broader. So
196
they agree with the original figure (MacFarland, 1906: PI. 29, Fig. 22),
which was “redrawn from an incomplete color sketch” (page opposite to
PL 22). Still broader are the dark stripes in another photograph (no. 40)
received from Dr. Lance, and even broader than these in Pickens’ speci
men of 1966. The darkest color pattern is shown by Pickens’ specimen of
1965, a dark brown animal not showing the light ground color.
The last-mentioned slug, as well as that of Pickens (1966), and that of
Lance’s photo No. 40 have red spots between the dark pigment, still
recognizable in the preserved material.
The tips of the velar processes and the rhinophores as well as the points
of the lateral tubercles are more or less intensely powdered with yellow.
The tubercles stand on the light stripes, except in Pickens’ specimen of
1965, which lacks these stripes. The maximum of 4 tubercles on either
pallial ridge (Marcus, 1964: 131) is exceeded by 5 (Pickens 1965 and
1966) and more (MacFarland, 1966: 116). The number of gills varies
from 7 (Lance, photos 58 and 40; Pickens, 1966) to 11 (MacFarland
1966: 116).
The determination of preserved specimens of polycerids is far from easy.
Certain colors, especially yellow tones, are generally not preserved. Con
tracted rhinophores and partially retracted ones are indistinguishable. The
dorso-lateral ridges or pallial margins and thickenings on them are some
times nearly evened in preservation. The color of the living animals is
valuable, but its intraspecific variation and change with age must be
considered.
The jaws and the radula are important and must be compared with the
published figures for every identification; for the jaw of P. atra one of
the original figures (MacFarland, 1906: PL 20, Fig. 67) should be used,
not the last one (id., 1966: PL 31, Fig. 29).
Until now, none of the species is known to occur on both the Atlantic
and Pacific coasts of the Americas, but as geographic criteria should not
be overrated in the classification of opisthobranchs, the following artificial
key refers to the American Atlantic and Pacific species of Polycera Cuvier,
1817, Greilada Bergh, 1894, and the farther distant Galacera Risso-
Dominguez, 1960.
1. Extrabranchial processes conspicuous lobes, or digitiform, club-
shaped or carrot-shaped appendages..................................................... 2
1. Extrabranchial processes in form of small tubercles, bosses, or
papillae, much shorter than the veil digitations....................................9
2. A single extrabranchial appendage on either side...............................3
2. More than one pair of extrabranchial appendages...............................4
3. Pallial ridge indistinct; rhinophores with 12 leaves.................................
.............................................................. marplatensis (Franceschi, 1928)
197
3. Low pallial ridge between veil and extrabranchial appendages
present; rhinophores with 25 leaves.................. chilluna Marcus, 1961
4. Jaws with upper wing-like expansion................................................... 5
4. Jaws without wing-like expansion........................ priva Marcus, 1959
5. Three gills................................................... zosterae O’Donoghue, 1924
5. Seven to nine gills.......................................................................................6
6. Extrabranchial processes shorter than gills, .hedgpethi Marcus, 1964
6. Extrabranchial processes longer than gills.......................................... 7
7. Extrabranchial processes carrot-shaped......... aurisula Marcus, 1957
7. Extrabranchial processes nearly cylindrical with pointed tips.........8
8. Rhinophores with 14-15 leaflets; foremost gill especially long;
3-4 extrabranchial processes on either side. . . .hummi Abbott, 1952
8. Rhinophores with 9 leaflets; foremost gill not especially long, 2
extrabranchial processes on either side.................... gnupa, spec. nov.
9. Second tooth of radula with basal spine.......................................... 10
9. Second tooth of radula without basal spine...............................................
..................................................................... alabe Collier & Farmer, 1964
10. First tooth half the size of the second............. herthae Marcus, 1963
10. First tooth smaller than second, but larger than half its size........... 11
11. Pallial ridges with closely set mamillary bulbs between rhino
phores and gills and behind gills; oral tentacles distinct........................
.................................................................................. odhneri Marcus, 1955
11. Pallial ridges with few, generally no more than five on either
side, compressed, pointed tubercles; oral tentacles very short.............
.............................................................................. atra MacFarland, 1905
29. Polycera gnupa, spec. nov.
Figure 59
Material.—Cholla Bay, Puerto Penasco, Sonora, Mexico. P. Pickens,
March 5, 1966. This species was found, together with P. atra “in a
branching bryozoan colony, growing on the sandstone reef in the middle
of Cholla Bay below the zero tide level.”
Description .—The two present specimens were 14 mm long in life, 7 and
8 mm preserved. The ground color is pale grey. This appears on the foot,
the pallial ridges, along the caudal crest, and on scattered tubercles on the
back and the sides. For the most part the dorsal and lateral skin is densely
stippled with brown pigment, which is concentrated to blackish brown
around the gills and on the head.
198
The two velar processes on either side are long, slender and pointed.
They are white with a bluish and an orange band around the middle
(Fig. 59, A). The rhinophores have a dark shaft, 9 lighter brown perfoli-
ations, and a white tip. The oral tentacles (Fig. 59, B, t) are quite short
and broad lobules. The two extrabranchial appendages are nearly cylin
drical with pointed tips. The tip is light and followed by a bluish and then
an orange ring. Both rings lie above the middle of the appendage. In the
larger specimen the outer, anterior, appendage is shorter, more pointed
and thinner than the stout inner posterior one; in the smaller specimen
it is longer. Both appendages are longer than the gills.
There are 7 unipinnate gills, the foremost of which is not much longer
than the others. The tip and the inner side of the gills are lighter than the
outer side. The knobbed median crest behind the gills ends near the tip
of the tail which bears an anterior orange and a pale blue posterior mark.
The antero-lateral corners of the foot are finger-shaped.
F ig ure 59. Polycera gnupa, sp. nov. A, right side view of preserved slug; B,
fore end of same in ventral view; C, jaws; D, half-row of radula, and one
innermost tooth in a different view.
199
The jaws (Fig. 59, C) are yellow and have broad, upper, wing-like
expansions. Their inner border is stronger than in P. hedgpethi (Marcus,
1964: Fig. 1). The radula (Fig. 59, D) com prises 9 rows of
3-4.1.1.0.1.1.3-4 teeth. The height of the teeth in micra is from the middle
outwards: 220, 300, 140, 120, 100, 50. Old teeth are reddish brown. The
innermost or first lateral tooth has a pointed cusp, and in its middle two
unequal spurs; the second lateral tooth has a basal spur. The innermost
marginal tooth has a rudimentary cusp, the outer marginal teeth have none.
The eversible part of the efferent duct is provided with very numerous
(about 1000) cuticular bristles.
r
Discussion .— P. gnupa is similar to two West Atlantic species, but cannot
be united with either of them. However, its taxonomic separation is
possibly not definitive, and more material of gnupa may diminish the
differences between it and the neighboring species. The color pattern of
adult specimens (Marcus, 1958b: 56) of the Brazilian-Floridian P. auri-
sula, not that of young slugs which are brighter (id., 1957: 432), agrees
with that of P. gnupa, but the carrot-shaped extrabranchial appendages
of aurisula separate it from gnupa.
In P. hummi Abbott, 1952, the rhinophores and the extrabranchial
appendages bear two brilliant blue rings, one above and one below the
orange ring. The rhinophores of 10-mm-long living slugs have 14-15
leaflets. Mrs. Sally D. Kaicher (St. Petersburg, Fla.), graciously sent us an
excellent color photograph of P. hummi, determined by Dr. R. Tucker
Abbott. This photograph shows that the differences between the anterior
and posterior gills, a general feature in Polycera, is accentuated in
P. hummi, so that in lateral view the branchial cup appears slanting
backwards, as if it were cut obliquely, even more than in Abbott’s figure
(1952: PI. 2, Fig. 1). In a single specimen of P. hummi from Beaufort,
N. C., two denticles were found at the base of the unworn cusp of the
second radular tooth (Marcus, 1961b: Fig. 5). This character must be
considered with prudence in comparing P. hummi with the only two
specimens from the Gulf of California.
The big P. hedgpethi Marcus, 1964, from Tomales Bay, California, is
grey with small black dots and yellow-orange marks on the velar and
extrabranchial appendages, on the pallial ridges, caudal crest, upper border
of the foot, and body tubercles. Blue elements were not recorded. This
feature and the black dots distinguish the color pattern of P. hedgpethi
from P. gnupa. As we strive for a key to preserved material, whose colors
may vary in several specimens, we separated P. hedgpethi by its short
extrabranchial appendages. Its tripinnate gills and more radular rows might
be correlated with its larger size, and perhaps with age. A short cusp of the
first tooth as in P. hedgpethi (Marcus, 1964: Fig. 3) can be seen also in
P. gnupa in the corresponding view (Fig. 59, D).
200
30. Acanthodoris pina, spec. nov.
Figure 60
Material.— 1. Puerto Penasco, Sonora, Mexico. Paula Vreeland, February
13, 1965. One specimen from the rocky intertidal zone.—2. Same locality.
Two more specimens found on February 5, 1966, by P. Pickens and Mary
Anne Hill.
Description .—As shown in a color photograph, a slug in movement is
rather flat (Fig. 60A); another animal, at rest, is higher. In life, the
length is about 18 mm; preserved, the animals are 8-11 mm in length,
7-8.5 mm in width, and 5-7 mm in height. The black dorsum is beset with
soft conical papillae, large red ones and numerous small white ones in
between. In preservation the bases of the large papillae are white and 1 mm
in diameter. The papillae are up to 2 mm in height. The notum, the
frontal veil, and the foot are bordered with red. The red stripe of the foot
is accompanied by an outer white line. In the photographs the rhinophores
are dark brown with white tips; the gills are brick red. The hyponotum is
sprinkled white and black; the sole is black.
The foot is rounded behind, where it stands out beyond the notum when
the slug is gliding. The anterior border of the foot is bilabiate due to a
F ig ure 60. Acanthodoris pina, sp. nov. A, living slug from color photo; B,
half-row of radula.— F ig ure 61. Dendrodoris krebsii (Morch). A, male duct,
specimen from Puerto Penasco (the section between the asterisks bears hooks);
B, male duct, specimen from Sao Sebastiao, Brazil.
201
transverse furrow; the upper lip is white. The veliform head has no median
notch. The clubs of the long rhinophores have 20 leaves; the sheaths bear
red papillae similar to those of the dorsum. In one case their number
differs on the two sides (3 and 5); in the other specimens there are 4 and 5
on both sides. There are 5-7 pluripinnate gills of equal size, one of them
in front. The anal papilla projects from behind into the center of the
branchial circle which is free from cutaneous papillae. One papilla com
pletes the circle of the gills. The peritoneum, correctly called pseudo
peritoneum by McFarland (1906; 1925-26), is colorless.
The labial armature consists of two lateral areas and a median plate of
thickened cuticle as in most species of the genus. The labial elements form
about 18 irregular rows. Their surface is rough, and they are widened
above and cleft into a variable number of lobules. The crop (ingluvies
buccalis) is less than half the volume of the buccal mass. The radula
(Fig. 60B) has 28 rows. There are 5-6 outer lateral teeth whose size
decreases outwards. The innermost lateral or intermediate tooth is 0.12 mm
high and 0.285 mm long. Its cusp is 0.12 mm in length, pointed and
curved. Its inner ridge bears 1-3 denticles. Sometimes there are up to
six denticles, and on some teeth none at all.
In the reproductive system the unarmed copulatory organ is a conspicu
ous feature, though penial hooks as well as their absence have been
related for A. pilosa, A. rhodoceras, and A. falklandica (Eliot, 1910: 16,
67; Marcus, 1961a: 27). The prostatic section of the male duct forms two
loops and is little shorter than the following muscular part, which runs
within a muscular sheath. The muscular atrium, Bergh’s “preputium,”
is distinctly set off from the efferent duct and lodges its everted, cylindrical
tip.
The genital aperture lies at the level of the anterior pedal border. The
external openings of the male and the two female canals are united.
The long vagina runs looping inwards and opens into a big, globular
spermatheca. From this point a long, thin duct leads to the narrow
spermatocyst filled with orientated sperm, and a short duct, the insemi
nation duct, to the female gland mass.
Essentially the genital organs agree with MacFarland’s figures (1925-26:
PI. 3, Figs. 1 and 6), except that the glandular part of the efferent duct
and the duct of the spermatocyst are longer in the present slug.
Discussion .—Leaving aside possible synonyms (Steinberg, 1963a: 64),
we have compared our specimens with the 12 species from the Pacific
coast of North America (O’Donoghue, 1927b: 6-7), two of which have
red elements in their color pattern. The back of A. pilosa purpurea Bergh
(1880a: 98) is purple brown, preserved reddish brown, not black, and
the shape of the first lateral tooth with its short and blunt cusp differs
from the present material. The dorsum of A. lutea MacFarland (1925-26:
12) is orange red with dots of lemon yellow between the papillae; there
202
are 34-39 radular rows, and the labial armature, especially the ventro-
median plate, is rudimentary.
Extending our comparison to the 7-10 further species of Acanthodoris,
we found it difficult to separate our slug from the geographically widely
distant A. metulifera Bergh (1905b: 98) from Tasmania. The different
characters of metulifera are: 40 radular rows; 3-4 denticles on the first
lateral tooth; head and foot yellowish; peritoneum with many violet-brown
flecks on the dorsal side. Possibly (Eliot, 1907: 350) Bergh’s species is
identical with the older A . globosa which has seven outer lateral teeth, and
whose labial elements are not divided.
31. Dendrodoris krebsii (Morch, 1863)
Figure 61
References.—Bergh, 1875: 87; Marcus, 1962: Fig. 19; 1963: 35; Collier
& Farmer, 1964: 389, Fig. 2 G-H, PI. 5; Marcus, 1967b: 95.
Synonyms.— Doridopsis Krebsii var. pallida Bergh, 1879b: 44; D. atropos
Bergh, 1879b: 49; Dendrodoris atropos Marcus, 1957: 443.
Material.— 1. Puerto Penasco, Sonora, Mexico. Rocky intertidal zone. In
1964 and part of 1965, April, June, July, and November.—2. Puerto
Lobos and Puerto Libertad. P. Pickens. “They are found infrequently.”—
3. Guaymas, Sonora, Mexico. Alex Kerstitch, March 12, 1966. Intertidal.
One specimen.
Further distribution.—Atlantic Ocean: Florida; Bahamas; Virgin Islands
(St. Thomas, St. Croix: original localities); Antilles; Curasao; coast of
Rio de Janeiro (original locality of atropos ) to Sao Paulo, Cananeia
(25° S).—East Pacific Ocean: Lower California (oceanic side); Gulf of
California from Puertecitos (30° 25' N) to the Bay of La Paz; mainland
of Mexico, Jalisco, Bay of Tenacatita (about 19° N, 105° W).
Remarks .—The color as shown by a photo of a 5 5-mm slug is velvety
black with a red-trimmed border and white tips of rhinophores and gills.
The latter are six in number, but as the two hindmost tufts are subdivided
from their bases, one might as well count eight branchiae. The largest of
the four preserved animals was 29 mm in length, 16 mm in width, and
10 mm in height.
Collier & Farmer stressed the thickened base of the penis in their East
Pacific material. In two specimens examined from Puerto Penasco we also
found this part thicker (Fig. 61A) than in two Atlantic specimens, one
from St. Kitts, Lesser Antilles, the other from Sao Sebastiao, coast of
Sao Paulo (Fig. 61B). These Atlantic slugs agree with one another.
For this comparison only completely retracted penes can be used. The
figures of Western Atlantic Dendrodoris, referring to D. krebsii (Bergh,
203
1875: PL 11, Fig. 15, 16) cannot be discussed, because the penes of the
three examined slugs (p. 90) were protruded in different degrees. In our
animals from the Gulf of California the ental penial hooks are up to 40 ^
high, while their maximum is 30 ^ in the Caribbean and Brazilian animals.
Neither this difference nor that of the diameter of the penial base is
sufficiently clear-cut to justify a subspecific separation. The length of the
section of the male duct between the prostate and the base of the penis is
variable. It measured 4 mm in one of the animals from Puerto Penasco
(Fig. 61 A), 2.5 mm in the other, 2.6 mm in the slug from St. Kitts, and
10 mm in that from Sao Sebastiao.
By re-examination of our Atlantic material and the study of that from
the Gulf of California we did not find more than individual differences,
e. g., the length of the cerebro-buccal connectives, and a shallow furrow
between the right and left cerebro-pleural ganglia.
Dr. Pickens observed that these slugs and a flatworm, we presume
Pseudoceros bajae Hyman, 1953 (Bull. Am. Mus. Nat. Hist., vol. 100,
art. 2, p. 365), have very similar colors. The polyclads generally hide
under stones during the daytime, and prey upon other animals in the dark,
while the dendrodoridids feed principally on sponges. So one can hardly
expect any biological significance of this resemblance. Presumably the
number of adult Polycladida, Opisthobranchia and other invertebrates with
planktonic stages depends greatly on the environmental factors, mainly
predation, acting upon the larvae.
32. Doriopsilla albopunctata (Cooper, 1863)
References.— O’Donoghue, 1927a: 93; Steinberg, 1961: 58; MacFarland,
1966: 196.
Synonyms.— Doriopsilla fulva MacFarland, 1905: 45; 1906: 130; 1966:
194 ( Dendrodoris fulva). Doriopsis reticulata Cockerell & Eliot, 1905: 41.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, January 15,
1965. A single specimen collected in the same tide pool as Taringa aivica
timia.
Further distribution .—California, from Salt Point, Sonoma County, about
38° 25' N, to Lower California, Point Eugenia (27° 51' N, 115° 05' W).
Gulf of California, Puertecitos (30° 25' N, 114° 40' W) (Farmer &
Collier, 1963: 62). Original locality: Santa Barbara.
Remarks.—In the photograph the animal is chestnut brown with a
purplish hue, the crenulate notal margin and the rhinophores are orange,
the gills and the tail light orange. Small opaque white dots are densely
set all over the notum, with exception of the margins. Those in the mid
line are quite small, the lateral ones a little larger, and those lying farther
outwards small again. The dots leave numerous round areas of the ground
204
color free, whose size is distinctly greater than that of a dot. In the
preserved slug these areas are slightly elevated; they correspond to sub
cutaneous glands. Alive, the specimen was 60 mm long; preserved, its
height, 8.5 mm, is small in comparison with the length (40 mm) and
breadth (21 mm). As in other Porostomata, the mouth is minute. The
contracted pharyngeal bulb together with the oral tube is only 4.5 mm
long.
33. D oriopsilla janaina Marcus, 1967
Reference.— Marcus, 1967b: 96.
M aterial.— Puerto Lobos, Sonora, Mexico. P. Pickens, April 16, 1965.
Two specimens in the intertidal zone.
Further distribution.— Panama, Pacific side of the Canal Zone (original
locality).
D escriptive notes.— The present preserved slugs are about 20 mm long,
14 and 10 mm broad, and 4-5 mm high. In the photograph the pink color
of the notum and the generally white, but sometimes dark red tips of the
tubercles are most conspicuous. The bases of the tubercles are surrounded
by black dots, especially large in one median and two lateral bands be
tween gills and rhinophores, and confluent in front. Two pairs of white
blotches, evidently consisting of secretion, lie on the sides. The lateral
tubercles are smaller than those in the middle. The rhinophores are red,
the gills orange, the borders of the rhinophoral pits and the branchial
pocket are white. The hyponotum is transparent, showing the bundles
of spicules.
34. Doriopsilla rowena, spec. nov.
Figure 62
M aterial.— Puerto Penasco, Sonora, Mexico. Mary Anne Hill, April 2,
1966. Seven specimens.
Description. — Alive, the animals are 8-10 mm in length. The biggest pre
served specimen is 10 mm long, 5 mm broad, and 3 mm high. The hypo
notum is about 2 mm in width, the foot 2.5 mm, pointed in front and
behind. The anterior pedal border is entire.
The spicules are dissolved except for traces in the hyponotum. Evi
dently the spicules are large and lie diagonally crossed in the connective
tissue of the back, parallel to the course of the cutaneous muscle fibers.
Those in the hyponotum are mostly radial, a few parallel to the border.
The color of the living animals varies; the collector noted whitish yel
low, pinkish orange, and yellow orange as fundamental colors. In all
specimens, small blackish brown melanophores, still present in the pre
served material, are uniformly scattered over the smooth back. In the
205
color photographs there are also white dots. Of these, especially large
spots occur in two lateral lines from the rhinophores to the gills, and
a few big spots externally to these lines. The white elements, evidently
secretion, are not preserved.
The ground color is more intense on the rhinophores and gills; these
organs are light yellow in the whitish yellow slugs.
The tentacles are coalesced in front, forming a small rim around the
mouth. The club of the rhinophores bears 8 thick, in part incomplete,
foliations; the brim of the inconspicuous sheath is smooth. There are 5-6
pluripinnate gills; the thick anal papilla and the renal pore in front of
it lie to the left of the branchial tuft.
The eyes (Fig. 62, B, y ) are uncommonly big: the pigment cup is 80-
100 ^ wide, the lens 60 ^ in diameter. The rhinophoral ganglia are
distinctly set off from the cerebro-pleural mass, whose two halves are
separated by a furrow. The buccal ganglia (cc) lie immediately behind
the nerve ring. The pharynx ( n x ) is, in the dissected slug, similar to that
of D. pharpa Marcus (1961b: Fig. 19, h )\ it begins wide with thin,
F ig u r e62. Doriopsilla rowena, sp. nov. A, living slug from color photo; B,
anterior part of alimentary tract; C, diagram of reproductive organs.
206
glandular walls, then becomes narrow and muscular, passes through the
nerve ring, and continues narrow in its post-cerebral course. The following
wider oesophagus (os) enters a large gizzard (iz), which is as long as the
next gastric section, the stomach (so), which is especially muscular in the
present species.
The pear-shaped ampulla (Fig. 62, C, a) receives the hermaphrodite
duct (h ) not quite at its inner end, but subapically. The bifurcation of
the spermoviduct lies in the female gland mass.
The prostatic part (q ) of the thin efferent duct bears many digitiform
glandular appendages. These are much longer than in D. janaina. The
following muscular part is sheathed by muscle layers (ss) and ends with
inconspicuous spines.
The vagina (v) leaves the atrium. The further female organs are rather
similar to those of D. janaina, though the proportions are somewhat dif
ferent. In the present species the insemination duct (in ) is much longer
than the canal which connects it with the spermatocyst ( s c ) ; in D. janaina
the insemination duct is shorter, and the stalk of the spermatocyst leaves
the spermatheca at the entrance of the vagina into the latter.
Discussion.— Besides the mentioned differences in the copulatory organs,
the size, the notal tubercles, the color, and the 18 leaflets of the rhi
nophores separate D. janaina from D. rowena.
Another species which must be mentioned in connection with D. rowena,
due to its range and the lobate prostate, is D. albopunctata. This large
species is quite different from D. rowena. Three species of Doriopsilla
from West Atlantic warm waters (areolata Bergh, leia Marcus, and
pharpa Marcus) have the prostatic part of the efferent duct thickened,
but not beset with digitiform appendages.
F igu re 63. Tritonia (Candiella) pickensi, sp. nov. A, living slug from color
photo, dorsal view; B, lateral view of right rhinophore; C, cones of masticatory
border; D, radular teeth; E, copulatory organs.
208
The genital apertures are located between the second and the third right
branchial tuft; the anus lies between the fourth and fifth gills, hence
behind the middle of the body in small animals but slightly in front of
the middle in larger ones. The renal pore is a little in front of the anus.
The masticatory border of the jaw (Fig. 63, C) bears about four rows
of smooth, pointed teeth. The radula (Fig. 63, D) has the formula 25-
28X10-11.1.1.1.10-11. The teeth are colorless or yellow. The tricuspid
rhachidian tooth is only a little broader than high, its middle cusp a little
longer than the lateral cusps. These fit into the concave inner side of the
first lateral teeth. The cusp of this intermediate tooth bears 6-8 smaller
denticles; one stronger point in front of them is not always distinct. All
following teeth are smooth. A malformed third row of two coalesced teeth
was noted in one of the radulae examined.
The ovotestis overlies the intestinal gland. The ampulla (Fig. 63, E, a)
is slender, fusiform. As systematically important organs of the reproduc
tive system, the copulatory organs were especially considered. The inner
muscular section ( d ) of the sperm duct is rather long, though shorter
than the following prostatic part ( q ). The penial part ( p ) of the male
duct ( e ), which lies in the atrium ( am) , is not quite as long as the pro
static section. In two preserved 5-mm slugs the prostates were 0.68 and
0.45 mm long, the penes 0.51 and 0.3 mm. In correlation with the
slender, relatively long penis the bursa canal (vagina, v) is longer than
the ampulla (s) of the bursa (spermatheca).
The species is named for Dr. Peter E. Pickens.
Discussion. — The editors of MacFarland’s posthumous work (1966)
intentionally left it in its original form (p. V III). Therefore Odhner’s
(1963) names of the Tritoniidae were not considered. In Odhner’s nomen
clature Duvaucelia is restricted to species with the anus far in front of the
middle and the genital apertures near the first gill. So MacFarland’s species
of Duvaucelia would be called Tochuina (tetraquetra) and Tritonia (the
3 other species).
Of the three subgenera of Tritonia, only Tritonidoxa Bergh, 1907, is
well characterized by its flagelliform penis (Odhner, 1926: 35). The dis
tinctly bilobed veil with numerous short processes of Tritonia and the
evenly rounded veil with at most a slight median notch and up to 10 digi
tate appendages of Candiella separate the type species well (Larsen,
1925: 12), principally because their sizes are very different (Tritonia
hom bergi up to 200 mm, Candiella plebeia up to 30 mm). However,
species of intermediate size, specimens not full grown, and preserved
material cannot always be allotted safely to one or other of these sub
genera, as already stated by Eliot (1905: 17; 1910: 146).
Tritonia pickensi is an indubitable Candiella. The only Candiella among
MacFarland’s species (1966) is Tritonia festiva (Stearns, 1873). Its in
nermost lateral tooth may bear denticles (MacFarland, 1966: 222).
209
However it differs from T. pickensi by reticulate lines on the sides and
dorsum; absence of an appendage on the rhinophore sheath; 10, ac
cording to MacFarland even 12, velar processes; and a completely dif
ferent penis (Marcus, 1961a: Fig. 112; MacFarland, 1966: PL 44, Fig. 2).
Another Californian tritoniid might belong to Candiella, the little known
(Marcus, 1961a: 32) Tritonia palm eri Cooper, 1862. If the comparison
is based upon Cockerell & Eliot’s specimen (1905: 33), the rugose back
with small warts and the roughly 10 velar appendages separate palmeri
from pickensi.
The species most nearly related to T. pickensi is T. (Candiella) bayeri
from Florida, also with an appendage of the rhinophore sheath. It has a
diffuse net of opaque white lines on the notum; the velar and rhinophoral
appendages are peppered with white; the mandibular cones have a longitud
inal sculpture; the rhachidian tooth is much broader than high; there is a
denticulate second lateral tooth besides the first, and the copulatory organs,
penis and vagina, are shorter than in the species from the Gulf of California.
D escription.— The expanded and the contracted slug in the photos are
semitransparent greyish with a slight reddish hue, few scattered opaque
white dots, and an opaque white line along the edge of the foot. The
tips of the outermost ramifications of the appendages are white; then
follows a blackish brown ring and an orange base (Fig. 64, A ).
The skin of the preserved slugs contains a little brown pigment. By local
contraction darker zones are brought about, e.g., on the edge of the foot,
where an inner dark line accompanies the outer white one. The larger of
the two animals is 13 mm long, 4 mm high without the cerata, and 3 mm
broad. The corresponding measurements of the smaller slug are 10, 3, and
2 mm. The fore end is semilunar, the body not sharply set off from the foot,
the tail end pointed; the anterior border of the foot is bilabiate and evenly
rounded to the sides. The rhinophores have about 15 leaves; their tips
are light.
The frontal veil bears three rows of papillae (Fig. 64, B ). That nearest
to the mouth comprises about 20 white nodules. Then follow about 10
papillae of different size. They are decorated with a dark ring and bear a
few small white knobs. The dorsal-most row of the frontal veil consists of
210
about three larger compound papillae of almost equal size on each side.
The white tips and orange bases have faded out; only the brown rings are
preserved. The rim of the rhinophoral sheath bears about five papillae,
some simple, some compound. The papilla on the outer posterior side of
the rim is highest. Two or three compound papillae occur along the back
of the sheath between rim and base (Fig. 64, A ). Immediately behind the
outer side of the base there is a fan-shaped appendage with a broad base
and about six compound papillae on its free border. The male opening
lies behind this appendage on the right side.
The body bears six pairs of flabellate appendages (cerata), the foremost
of which are the largest. The broad trunk emits about five branches
terminating with 4-10 brown bulbs beset with numerous light papillae. The
ramification of the cerata is not dichotomous. The female opening lies
under the first right ceras. Farther dor sally than the reproductive apertures
the anus is located in the pericardial interspace between the first and
second ceras. The second cerata lie a little behind the middle. Of the four
branches which go out from the thick trunk, the second from the middle is
a trifle longer than the others, and the same holds, though still less
pronounced, for the foremost ceras. The following cerata stand near to
F igure 64. Dendronotus nanus, sp. nov. A, lateral view of fore end, combined
from color photo and preserved specimen; B, head of larger preserved speci
men in ventral view.
211
one another and decrease in size backwards; the hindmost of the right side
is represented only by one bulbar papilla. This description refers to the
preserved 13-mm slug; in the smaller one the number of appendages,
especially on the veil and the rhinophores, is much smaller.
Thompson (1960) had only quite small (2 mm) and full-grown (90
mm) specimens of D. frondosus for the examination of the intestinal
diverticula in the cerata. In our 9- and 13-mm-long animals the liver
branches do not extend into the cerata.
The yellow mandibles of the larger specimen are 4.0 mm long, 1.3 mm
broad behind and 3 mm in front, where 1.7 mm belong to the superior
process, 1.3 mm to the masticatory border. The latter bears about 90
crested pointed denticles. The radula (Fig. 65, A) comprises 26 rows.
There are 9-14 lateral teeth per half-row. The rhachidian tooth has about
10 strong denticles on either side of the cusp. The lateral teeth increase in
size to the sixth and decrease outwards. The innermost lateral tooth is
shorter than the following ones. Its broad cusp bears up to 12 denticles.
In the old rows of the radula the second to sixth or even eighth lateral teeth
have up to six denticles; in the newer rows two or three, and in the newest
ones only a single denticle or none at all may appear. The salivary glands
are long and voluminous as in D. frondosus (Bergh, 1879a: PI. 3, Fig.
12). The posterior and the right and left anterior livers open by separate
ducts into the stomach, as drawn by Odhner (1936: 1064, Fig. 4).
The reproductive organs of the larger specimen (Fig. 65, B) are juvenile
but not primordial. The male duct ( d ) is very long and convoluted, with
beginning glandular differentiation of the ental half, hence similar to adult
D. iris (Odhner, 1936: Fig. 40). Also the cylindrical copulatory organ
agrees with iris. The rudimentary bursa copulatrix (Fig. 65, B, s) is the
same as in frondosus and iris; a distinct differentiation of the vagina into
vagina proper and vesicula seminalis is not developed, but in any case the
type is more like iris than like frondosus, robustus, and MacFarland’s
species (1966: PI. 50, Figs. 2-4), whose vesicle (MacFarland’s “sperma
theca” ) communicates with the vagina by a duct. The great length of the
ectal, non-glandular, efferent duct is peculiar in the present species, and
the difference between our Figure 65, B, and that of a young specimen of
D. iris (30 mm; Odhner, 1936: Fig. 41) is obvious.
R em arks. — If Odhner’s synopsis is followed, D. gracilis (Baba, 1949: 87,
167) and D. nanus must be assigned to the first group with “obsolete”
veil, hence not protruding laterally as in D. robustus (Bergh, 1894: 142).
The second character of group I, “velar processes about 3-4, largest to
wards the median line of the body, smaller laterally” does not apply to
D. nanus. The cerata of the hitherto described species of Dendronotus,
including also MacFarland’s (1966: 265-279), are arborescent, each
containing one or two main stems and looking very different from the
212
F igu re 65. Dendronotus nanus, sp. nov. A, innermost teeth of radula; B,
diagram of reproductive organs, the efferent duct drawn as a simple line.
F i g u r e 66. Doto lancei, sp. nov. A, living slug from color photo; B, ceras
with few tubercles; C, ceras with many tubercles; D, same seen from inner
side; E, radular teeth.
215
The species is named for Dr. James R. Lance of Scripps Institution of
Oceanography, La Jolla, California, who sent us a beautiful kodachrome
transparency (his No. 34) of a slug from the Gulf of California which
evidently belongs to the species described here.
Discussion.— D oto columbiana O’Donoghue (1921: 204) from Vancouver
Island and Dillon Beach (38° 15' N) agrees with D. lancei regarding the
subapical ring around the ceratal tubercles, an uncommon character in
D o to , and the shape of the gills (Marcus, 1961a: Fig. 128). In D. colum
biana, however, neither apical spots of the tubercles occur, nor black spots
on the inner side at the insertion of the cerata. Already the classification of
the specimens from Dillon Beach, which are smaller than those from
Vancouver and have more cerata, may be criticized. A further amplification
of the concept of D. columbiana to include slugs with still more cerata than
occur in the animals from the Vancouver region and from Dillon Beach,
with color marks not recorded from the original material and absent in all
specimens from Dillon Beach also, and with a different border of the
rhinophoral sheath, is in our opinion untenable. The ejaculatory duct
winds through the penis in D. columbiana (ratio: 3.5:1), while its course
is straight in that of D. lancei (ratio: 4.2:1).
D. kya Marcus (1961a: 39) from Pacific Grove (36° 38' N) has eight
pairs of cerata and black spots at their insertions, but neither rings nor
spots on the cerata. The latter insert axially in kya, obliquely in lancei
and columbiana.
Another species with a basal ceratal or notal spot and with a range
reaching the Caribbean is D. cinerea Trinchese, 1881 (Marcus, 1963: 38).
Its gills are indistinct. The cerata have light tips, and the ceratal tubercles,
which are set somewhat irregularly, have neither rings nor apical spots.
The variable shape of the border of the rhinophore sheath in D. lancei
cannot once and for all invalidate this character in all other species of D oto,
but it is not always reliable (Marcus, 1961a: 41).
tooth was compared with that of a European species, “except that the
lower corner is very much more produced.” As the general color differs
widely from that of cynara, and only the white tips of the cerata agree,
Cockerell’s description without figures is far from sufficient to approach
cynara to cooperi. As Steinberg said (1961: 62), only further collection
in the San Pedro area can establish the identity of C. cooperi. The Carib
bean C. dushia Marcus (1963: 41) is morphologically rather different
from cynara.
Steinberg united (1963a: 66) C. piunca Marcus (1961a: 47) with
C. trilineata O’Donoghue (1921: 197; 1924: 25). The single examined
specimen of the latter had a U-shaped median tooth, and the longer side
of the triangular lateral tooth (the hypotenuse, as O’Donoghue called it)
is denticulate. The jaw was not figured, only compared with that of
C. fusca O’Donoghue (1921: 195), in which the breadth is 76 per cent of
the length. The median tooth of piunca (loc. cit.: Fig. 165) is not U-
222
shaped, the lateral tooth bears denticles on one of the shorter sides of the
triangle, and the breadth of the jaws is 63-70 per cent of its length. If these
characters were found to be connected by intermediate types, and inhabi
tants of the type localities, the Vancouver region and Dillon Beach, agreed
in their reproductive organs, the proposed synonymy could be accepted.
Many species of Coryphella have similar buccal armatures. The color pat
tern varies intraspecifically in some cases, and in others that of different
species agrees.
In Bergh’s later lists (1891; 1892a), which comprise several problemat
ical species of Coryphella and Flabellina, cooperi and iodinea are not
included.
From the coasts of central and northern California two new species of
Coryphella were recently published (MacFarland, 1966: 313-322). Only
one of them, C. pricei, must be compared with C. cynara, because both
have high, slender cerata. The short conical copulatory organ of C. pricei
(loc. cit.: PL 66, Fig. 8) differs widely from the nearly globular one of
C. cynara.
42. Flabellina telja, spec. nov.
Figures 76-82
M aterial. — 1. Cholla Bay, Puerto Penasco, Sonora, Mexico. Dorothea
Caskey, June 30, 1965. Two specimens about 3 m from one another “were
found in about 3 feet of water. They swim with lateral bending movements
of the body in much the same manner as D endronotus nanus.”— 2. Puerto
Penasco, Sonora, Mexico. Mary Anne Hill, June 18, 1966. One specimen
on some algae in the rocky intertidal zone.
Description. — The color in the photograph of the specimen taken in 1965
is similar to that of Coryphella cynara. The body is bluish, the back and
the area between the rhinophores and tentacles bluish grey; the cerata
contain orange brown hepatic diverticula and have white tips. The knobs of
the rhinophores, the two distal thirds of the long tentacles, and the tips of
the basally broad, not much produced corners of the foot are white, and
a white crest covers the dark blue back of the tail which ends white.
The blue color has stained the alcohol in which the animals are pre
served. The preserved specimens are 11 and 8 mm in length; in the larger,
the body is 3 mm in breadth, the sole 2 mm. The further measurements
are: cerata up to 4 mm; foot corners ( z o ) 1 mm; tentacles (t) 3 mm;
rhinophores (io) 2.5 mm, with 30 leaves.
The slug of 1966 is white with a bluish tone, especially on the tentacles,
and with red hepatic diverticula in the cerata. Alive it was about 5 mm in
length, and the rhinophores are as long as the tentacles or a little longer.
The foot is set off from the sides, its anterior border is bilabiate and
notched. The gonopore lies under the right anterior group of the cerata; the
anus (ar) farther backwards between the socket bearing the hind-most cerata
223
of the right anterior, and that of the first row of the posterior liver.
The cerata stand in oblique rows (Fig. 76) which are more dorsal in
front, more ventral behind. The cerata are firmly inserted in groups on
common sockets (Fig. 77).
The following numbers refer to the large slugs taken in 1965. The
anterior livers (ri) have each 4 groups with 6, 5, 3, and 2 cerata; the
median cerata are, as usually, highest. The posterior liver (/) comprises
nine groups on either side, the two first each with 6, the two following each
with 5, the five last with 4, 3, 2, 1, and 1 cerata. Buds of cerata, which
were not counted, appear on the outer ends of the sockets as in many
Eolidacea. The cnidosacs are up to 0.3 mm long, hence less than 1/10 of
the highest cerata.
The colorless, delicate jaws (Fig. 79) are 1.0 mm in length, 0.75 mm in
breadth. The masticatory process is small; it bears about 10 rows of
denticles, the largest about 20 jjl in height, which stand along the border
(Fig. 80). There are 28 radular rows (Fig. 81). The rhachidian tooth is
100 fx high and 80 ^ broad. Its median cusp is flanked by 8-11 denticles.
The large uppermost of these meet over the median cusp; the following
ones are slightly narrower. The lateral teeth are 130 ^ in length, 42 ^ in
breadth. Their longest side is straight, the shortest provided with 6-8
denticles, the middle-sized side is concave. The pharynx is minute, 1.5 mm
in length in the 11-mm slug.
The mandibles and the radula of the small specimen collected in 1966
were compared and found to agree with the corresponding organs of the
larger slugs; the denticles flanking the cusp of the rhachidian tooth are
only 5-8 in number.
The gonad is dorsal to the main liver duct. The coiled, colorless ampulla
(Fig. 82, a) enters the female gland mass (g), from which the winding
prostatic section ( q) of the male duct emerges. The following portion, not
glandular, opens into the atrium ( am) . Its muscular folded walls can be
understood by the everted copulatory organ in the smaller specimen (Fig.
78). There the atrium forms a collar with a folded, warty peduncle; a
roundish penial papilla bearing the ejaculatory duct projects from the collar.
The male and female openings are united ( o a ) . A stalked spermatheca
(s) goes out from the nidamental duct (ni ). Far entally, immediately at the
entrance of the ampulla and the exit of the prostate, there is a second
seminal vesicle (sv) embedded in the mucus gland (g), similar to Odhner’s
finding (1939:61) in Coryphella pellucida (Alder & Hancock, 1846).
Discussion. — The insertion of the groups of cerata on peduncled supports
characterizes the genus Flabellina, This feature is obvious in Vayssiere’s
frequently copied figure (1888: PI. 2, Fig. 1) of the type species, F. affinis
(Gmelin, 1791). The presence of such supports led Baba (1955: 29, 52)
to transfer Coryphella ornata Risbec (1928: 266; 1953: 143; Baba, 1936:
44) to Flabellina. If this assignment is maintained, Risbec’s species must
224
F i g u r e s 76-82. Flabellina telja, sp . n o v . 76, r ig h t s id e v ie w w it h o u t c e r a ta ;
77, f o u r th r o w o f c e r a ta o f r ig h t liv e r ; 78, a n te r io r p a r t w ith e v e r te d c o p u la to r y
o r g a n ; 79, ja w ; 80, m a s t ic a to r y b o r d e r o f ja w ; 81, r a d u la r te e th ; 82, d ia g r a m
o f r e p r o d u c t iv e o r g a n s.
receive a new name, because F. ornata Angas (1864: 67) has priority.
While the peduncled supports of Risbec’s species are distinct in Baba’s
figures of 1955 and in Abe’s (1964: PI. 32, Fig. 114), they are not in the
earlier ones. Even F. affinis may exceptionally seem to have sessile cerata
(Odhner, 1939: 62). Possibly the peduncles are obscured by contraction
of the cutaneous muscles of the back.
225
Neither perfoliate rhinophores nor anterior corners of the foot produced
into tentaculiform appendages (O ’Donoghue, 1922b: 139) are peculiar
characters of Flabellina. Coryphella californica Bergh (1904: 6) has the
front corners of the foot longer than the tentacles or the rhinophores.
Perfoliation of the rhinophoral clubs occurs in C. trophina (Bergh, 1894:
134), C. fusca, C. trilineata (O’Donoghue, 1921: 196, 197), and C. piunca
Marcus (1961a: 47).
When Pruvot-Fol (1954a: 421, note 2) called the lateral teeth different
in Flabellina and Coryphella, she probably meant the approximately rec
tangular base of these teeth in F. affinis (Vayssiere, 1888: PL 4, Fig. 48),
from which the triangular cusp is set off. This is a specific character of
F. affinis, different only by degree from certain species of Coryphella, e.g.,
C. lineata (Loven, 1846) (Bergh, 1876a: PI. 15, Figs. 18, 20).
It is an open question whether a penial stylet is characteristic of
Flabellina (O’Donoghue, 1922b: 139). Coryphella ornata Risbec, 1928,
allotted to Flabellina by Baba (see above), has an unarmed penis (Risbec,
1953: Fig. 110, I). For F. affinis, and vaguely also for his F. iodinea
(1879a: 81), Bergh indicated a stylet (1876a: 651; 1886d: 50). Vayssi
ere, who knew Bergh’s papers (1888: 11), called the penis of F. affinis
unarmed (p. 79, 84; 1913: 294; 1928) without any reference to Bergh’s
observation, and Bergh repeated the occurrence of a stylet in his diagnoses
of the genus (1891: 39; 1892a: 1034). Bergh’s figure (1876b: PI. 16,
Fig. 4) is not incompatible with Vayssiere’s (1888: PI. 4, Fig. 49); pos
sibly the thick wall of the penial papilla shown by Vayssiere becomes cutic-
ularized in completely mature slugs of F. affinis. In Samla annuligera Bergh
(1900: 237) the penis is unarmed; the species seems to be a Flabellina.
The above-mentioned Flabellina from Puget Sound differs from the
present species by its iodinea-Ykt color, viz., violet purple body, orange
rhinophores, orange red cerata, and foot angles longer than the tentacles.
The lateral tooth of Flabellina telja is similar to that of Coryphella fusca
O’Donoghue (1921: 195). Along with other characters this species, in
preservation up to 19 mm in length, has 19 rows of radular teeth. The
median cusp of the rhachidian tooth bears 4-6, exceptionally 7, lateral
denticles which leave a great extension of the cusp free.
The teeth of Coryphella fisheri MacFarland (1966: PI. 65, Figs. 16-18)
are similar to those of F. telja. The color pattern, the annulate (not per
foliate) rhinophores which are longer than the tentacles, and the absence of
a groove in the anterior margin of the foot distinguish C. fisheri at once
from the present species.
43. H ermissenda crassicornis (Eschscholtz, 1831)
R eferences .— Marcus, 1961a: 52 (literature); MacFarland, 1966: 358.
M aterial. — 1. Puerto Penasco, Sonora, Mexico. P. Pickens, January 15,
1965.— 2. Puerto Lobos. P. Pickens, April 16, 1965. “In the rocky
226
intertidal zone, quite common at the second locality.”
Further distribution. — From Sitka, Alaska (original locality), to Lower
California, Point Eugenia, and in the Gulf at Angel de la Guarda Island
(Farmer & Collier, 1963: 62); northern part of the Gulf of California
(Lance, 1966: 79).
D escriptive notes. — The known color variation of this species is evidenced
by the present photographs which show a transparent bluish and a more
opaque orange yellow slug. The tips of the cerata are white, the intestinal
diverticula in the cerata are brown.
233
These cells contains granular and gelatinous secretion and open into cel
lular ductules which unite into wider ducts leading to the pores. The parcels
of these gland cells are embedded in diagonal fibers of the body muscula
ture and blood spaces. From the main duct a few branches go into the
perinotum, where they drain clusters of small gland cells.
The openings of the body are as in H. lesliei (Stearns); the male
aperture lies in the middle of the forehead hidden by the veil; the opening
of the mantle cavity (anus) behind the posterior end of the foot; the
female opening to the right of the anus; and the pneumostome on a
papilla between anus and notal border.
The body cavity of Hoffmannola is divided into a wider left compart
ment and a smaller right one by a septum (Fig. 89, se) formed by a fold
of the annular musculature protruding from the body wall. The septum
begins at the hind end of the body cavity and ends in front of the level of
the stomach (fe). On the ventral side it extends farther forward than
dorsally. The peritoneum is colorless, but the pericerebral conjunctive
tissue and that along some blood vessels is pigmented.
The salivary glands ( sa ) are branched; their fundi are fastened to the
nerve ring as in Onchidella (Plate, 1893: 112). The radula (Fig. 91)
comprises about 170 rows, each with about 600 teeth per half-row. The
tricuspidate rhachidian tooth, the shorter innermost lateral tooth, and the
long rectangular bases of the outer lateral teeth, their large, blunt mesocone,
and the small, pointed ectocone agree with H. lesliei (Hoffmann, 1928:
Text-fig. 12). Also the lobes of the intestinal gland, whose size decreases
from the right to the left and to the hind one, and the course (/) of the
intestine (ibid.: Text-Fig. 13) are the same.
The asymmetry of the reno-pulmonary complex ( k ) , much longer on
the left than on the right side, is fundamentally the same as in H. lesliei,
but the organs are much bigger in the present species.
The ovotestis ( o v ) consists of more than 10 follicles, whose efferent
ductules unite into the hermaphrodite duct. Its fine coils, the seminal
vesicle (Fretter, 1943: 648-649), and the caecum, Hoffmann’s vesicula
seminalis (1928: Text-fig. 14, ves. sem.), as well as the following parts,
the pouches of the mucus gland ( mu) , and the prostate in the middle,
the lobulate albumen glands on both sides, are the same as in Onchidella.
The brownish, globular bursa inserts far ectally, the coiled vaginal ap
pendage, Fretter’s flagellum (1943: 703), is quite short and inserted still
farther ectally.
The male duct (Fig. 92, d ) enters the body cavity beside the right
tentacle. It begins thick, 0.18 mm in diameter, and glandular; in the
preserved specimen this section is green. It loops on the bottom of the
visceral cavity backwards to the fore end of the septum {se), turns for
wards, becomes thin (80 /x), colorless, with single pigment cells on the
walls, and enters the penis, a straight muscular tube between the male
234
opening and the insertion of the penis retractor (r), which originates deep
between the two muscle layers of the septum. In some cases there is a
pigment spot at the insertion of the retractor. A loose outer layer of
muscles sheathes the central one around the coils of the male duct ( d ),
still 80 fi in diameter, which first descends and then ascends again (Fig.
95). The anterior efferent section (Fig. 94, as) of only 25 diameter
opens into a cuticular tube (ci) produced by a thin epithelium and sur
rounded by the outer muscle layer. A dilatation with conical glandular
villosities (Hoffmann, 1928: PI. 4, Fig. 16, dr. a.) is not developed. A
loose tissue, rich in nuclei ( nu) , lies between the outer muscle layer
and the inner one containing the male duct. The foremost section, the
male atrium (Fig. 93, a m ), is a muscularized invagination of the skin,
whose noncuticularized epithelium is thrown into longitudinal folds.
The species is named in remembrance of Hans Hoffmann (Jena).
R esu m o
Die Arbeit enthalt 47 Arten und Unterarten, die Dr. Peter E. Pickens
von der Universitat von Arizona (Tucson) im oberen Litoral der nordost-
lichen Kuste des Golfes von Kalifornien gesammelt hat. Von den 20 neuen
Formen stellen lnuda luarna, sp. n., und Tayuva ketos, sp. n., neue Gat-
tungen der Dorididen dar. Mehrere der Arten, die Dr. Frederick M. Bayer
vor Kurzem an der pazifischen Kuste von Panama gefunden hat, kommen
auch im Golf vor, unter ihnen die zu den Neritacea gehorige, im ubrigen
indowestpazifische Titiscania limacina. Gut zwei Drittel der festgestellten
Opisthobranchier sind Warmwasserformen. Beziehungen zur westatlan-
tischen Warmwasserfauna sind deutlich.
L is t of T ypes
The type specimens of the new species described in this paper have been
assigned catalogue numbers in the U.S. National Museum. As it was impossible
to insert these in the text, they are given below in the order in which the
species appear in the text.
7. Hermaea hillae, 678400 27. N em brotha eliora, 678410
13. Berthellina engeli ilisima, 678401 29. Polycera gnupa, 678411
15. Conualevia mizuna, 678402 30. Acanthodoris pina, 678412
18. Chromodoris banksi sonora, 34. Doriopsilla rowena, 678413
678403 35. Tritonia (Candiella) pickensi,
20. Casella sedna, 678404 678414
21. lnuda luarna , 678405 36. Dendronotus nanus, 678415
22. Doris pickensi , 678406 38. D oto lancei, 678416
23. Discodoris mavis, 678407 41. Coryphella cynara, 678417
25. Taringa aivica timia, 678408 42. Flabellina telja, 678418
26. Tayuva ketos , 678409 47. Hoffmannola hansi, 678419
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248
INDEX*
To Taxa Described or Mentioned
*Names in italic type are currently rejected for nomenclatural or taxonomic reasons; names
in roman type are in current use. Responsibility for this index rests with the editors,
not with the authors of the work.
249
atlanticus, Pleurobranchus (Susania), 44, 163 limbaughi, 169
atra, Polycera, 196, 198, 238 marginata, 170
atromarginata, Casella, 181 rumia, 51, 124, 169
atropos, Dendrodoris, 96, 203 scabriuscula, 51, 170
Doridopsis, 203 sparsa, 169
Doriopsis, 95 Cadlinella, 166
Audura, 192 calcarata, Bornella, 105, 124
aurantiaca, Dirona, 217 californica, Aplysia (Neaplysia), 155, 238
aurila, Discodoris, 80, 85 Coryphella, 226
aurisula, Polycera. 198, 237 californiensis, Hypselodoris, 59, 176, 238
Austraeolis, 114 Candiella, 104, 209
catina, 112, 124 plebeia, 209
fucia, 114 capensis, Aglaja, 20
ornata, 114 Nembrotha, 196
Austrodoris, 66, 74, 186 Pleurobranchaea, 47
nivium, 186 caramella, Doto, 106
Awuka, 166 carpenteri, Onchidella, 227, 229, 230
Onchidium, 227, 229, 230
Casella atromarginata, 181
bakeri, Aglaja, 21 cincta, 181
banksi banksi, Chromodoris, 173, 237 foxi, 181
banksi sonora, Chromodoris, 173, 237 rubra, 181
Baptodoris, 91 rufomarginata, 181
baratariae, Corambella, 94 sedna, 178, 237
bayeri, Feiimare, 62, 124 catina, Austraeolis, 112, 124
Tritonia (Candiella), 101, 124 cauze, Elysia, 28
berghi, Cadlina, 170 cauze scops, Elysia, 28, 123
Berghia, 117 cavernae, Discodoris, 80
bermudense, Umbraculum, 42, 123 cedrosensis, Aplysia (Aplysia), 158
Berthella, 162, 165 celtica, Onchidella, 121, 230
patagonica, 43 ceylonica, Aglaja, 19
plumula, 162 Chelidonura, 151
tupala, 43, 124 evelinae, 18
Berthellina, 162 hirundinina, 21, 124
amarillia, 163 chilluna, Polycera, 198
circularis, 163 Chromodoris, 56, 124
edwardsi, 162 agassizii, 56, 176, 178
engeli ilisima, 160, 237 amoena, 173
quadridens, 43, 123, 163 banksi banksi, 173, 237
Berthellinops, 162 banksi sonora, 173, 237
bicolor, Doris, 66 dalli, 176
binneyi, Onchidella, 227, 231, 237 macfarlandi, 178
bonairei, Petalifera, 39 neona, 51, 124
borealis, Onchidella, 2.29, 230 norrisi, 170, 237, 238
Boreodoris, 166 nyalya, 53, 58, 124
Bornella, 208 punctilucens, 176
calcarata, 105, 124 roseo-picta, 61
Bouvieria, 165 tura, 55
bovena, Doris, 66 universitatis, 176
branneri, Discodoris, 75, 78, 80 zebra, 60
braziliana, Spurilla, 118 chromosoma, Spurilla, 227, 237, 238
brevicauda, Phidiana, 109, 110 cincta, Casella, 181
Bulla antillarum, 22 cinerea, Doto, 216
gouldiana, 148, 237 circularis, Berthellina, 163
Bullaea guamense, 15 citrinus, Stylocheilus, 40, 124
clandestina, Thordisa, 91
Cadlina, 169 (key), 181 coelestis, Hypselodoris, 62
berghi, 170 coirala, Hermaea, 152, 237
evelinae, 123, 168, 170, 237, 238 columbiana, Doto, 216
flavomaculata, 169 Conualevia, 166, 237
juvenca, 169 alba, 186
laevis, 184 mizuna, 167
250
cooperi, Coryphella, 221, 222, 223 branneri, 75, 78, 80
Corambella baratariae, 94 cavernae, 80
Coryphella, 220, 226 edwardsii, 80
californica, 226 erubescens, 80
cooperi, 221, 222, 223 evelinae, 75, 78, 80, 124
cynara, 220, 222, 237 fragilis, 75
dushia, 222 heathi, 80, 82, 87, 188, 189, 288
falklandica, 221 hedgpethi, 75, 78, 80, 123
fisheri, 226 indecora, 80
fusca, 222, 226 maculosa, 80
iodinea, 218, 238 mavis, 187, 238
lineata, 226 mortenseni, 76, 80, 82
ornata, 224, 226 muta, 80
pellucida, 224 notha, 80, 87
piunca, 222, 226 notiperda, 75
pricei, 223 phoca, 78, 124
rufibranchialis, 219 purcina, 80, 81, 123
trilineata, 222, 226 pusae, 80, 82, 124, 189, 238
trophina, 226 rubens, 80
crassicornis, Hermissenda, 226, 238 spetteda, 75, 76
Cratena, 115 tristis, 80
kaoruae, 114, 124 voniheringi, 80, 87
crispa, Elysia, 34 diuda, Thordisa, 91
crispata, Tridachia, 33, 35, 37, 123, 124, 237 divae, Doto, 108
crossei, Pleurobranchus, 44 Nembrotha, 196
cristallina, Cyerce, 38 Dolabrifera dolabrifera, 38, 124
cucullata, Marionia, 104, 125
dolabrifera, Dolabrifera, 38, 124
Cyerce antillensis, 37, 123
cristallina, 38 Doridella obscura, 94
jheringi, 38 Dorididigitata maculata, 186
cynara, Coryphella, 220, 222, 237 Doridopsis atropos, 203
Krebsii var. pallida, 203
dalli, Chromodoris, 176 Doriopsilla, 99
dactylomela, Aplysia (Varria), 38, 124
albopunctata, 98, 99, 189, 204, 238
Dendrodoris, 99 areolata, 98
atropos, 203 fulva, 204
fulva, 204
janaina, 96, 124, 205, 207, 237
krebsii, 95, 123, 124, 203, 237, 238 leia, 207
subpellucida, 96
pharpa, 206, 207
Dendronotus, 208
rowena, 205, 237
frondosus, 212
Doriopsis atropos, 95
gracilis, 212
fulva, 99
iris, 210, 212, 213
krebsii var. pallida, 95
nanus, 210, 238
reticulata, 99, 204
robustus, 212
Doris, 186
depicta, Aglaja, 20
(Asteronotus) alabastrina, 186
depilans, Aplysia, 158 bicolor, 66
depressa, Phyllaplysia, 40
bovena, 66
Diaulula sandiegensis, 189, 238
echinata, 186
Dictyodoris tesselata, 191 odonoghuei, 186
diegoensis, Lamellaria, 147 pickensi, 184, 189, 237
digueti, Lamellaria, 166 verrucosa, 186, 237
Pleurobranchus, 165 Doto, 215, 216
diomedea, Aglaja, 20 amyra, 214, 238
Tridachiella, 34, 124, 154, 237 caramella, 106
Dirona albolineata, 217 cinerea, 216
aurantiaca, 217 columbiana, 216
picta, 216, 238 divae, 108
disa, Taringa telopia, 87, 124 lancei, 214, 216, 238
Discodoris, 69, 74, 76 (key), 124 pita, 106
alba, 80 uva, 108
aurila, 80, 85 varians, 214
dubia, Aglaja, 20 Noumea, 55
Thordisa, 91 flavomaculata, Cadlina, 169
duis, Elysia, 31, 124 flindersi, Anisodoris, 70
dura, Aplysia (Aplysia), 158
floridana, Onchidella, 119, 123
dushia, Coryphella, 222
Duvaucelia, 104, 209 foxi, Caselia, 181
insulae, 104 frangoisi, Risbecia, 62
lineata, 104 fragilis, Discodoris, 75
frondosus, Dendronotus, 212
fucia, Austraeolis, 114
echinata, Doris, 186 fulva, Dendrodoris, 204
edenticulata, Hypselodoris, 56, 123, 178 Doriopsilla, 204
edwardsii, Berthellina, 162 Doriopsis, 99
Discodoris, 80 fusca, Coryphella, 222, 226
elegans, Harminoea, 23, 24, 123 fuscovittatus, Stiliger, 154
Tritoniopsis, 103
eliora, Nembrotha, 194, 237 Galacera, 197
Elysia cauze, 124
gardineri, Susania, 44
cauze scops, 28, 123
gemmata, Aglaja, 20
crisp a, 34
glauca, Aeolidiella, 117
duis, 31, 124
Glaucus atlanticus, 119, 124
evelinae, 27, 123
globosa, Acanthodoris, 203
flava, 33
gracilis, 32 Glossodoridiformia, 166, 186
alba, 186
livida, 32
papillosa, 27, 123 Glossodoris aegialia, 61
papillosa, 29 gnupa, Polycera, 198, 237
picta, 32 Godiva rubrolineata, 112, 124, 125
pruvotfolae, 34 gouldiana, Bulla, 148, 237
sp., 32 gracilis, Dendronotus, 212
Elysia, 32
splendens, 32
tuca, 29, 123 Hypselodoris, 60
(Elysiopterus) verrilli, 34 gratiosa, Nembrotha, 196
greeleyi, Peltodoris, 66, 72, 124
engeli, Phyllaplysia, 40, 123
Greilada, 197
engeli ilisima, Berthellina, 160, 237
guadaloupensis, Haminoea antillarum, 22
enteromorphae, Phyllobranchopsis, 153 guamensis, Micromelo, 15
Eolidina, 116 Guyonia, 166
erubescens, Discodoris, 80
Erythrodoris, 166
Etidoris, 91 Haminoea, 21
ladislavii, 91 anlillarum, 22, 124
evelinae, Cadlina, 123, 168, 170, 237, 238 antillarum guadaloupensis, 22
Chelidonura, 18 elegans, 23, 24, 123
Discodoris, 75, 78, 80, 124 strongi, 149
Elysia, 27, 123 virescens, 149, 238
Onchidella, 236 virescens rosacea, 149
exigua, Phidiana, 110, 111 hamva, Pleurobranchaea hedgpethi, 48, 124
exsulans, Tritonia, 123 hansi, Hoffmannola, 232, 237
heathi, Discodoris, 80, 82, 87, 188, 189, 238
falklandica, Acanthodoris, 202 hedgpethi, Discodoris, 75, 78, 80, 123
Coryphella, 221 Polycera, 198, 199, 200
Felimare, 62, 166 hedgpethi hamva, Pleurobranchaea, 48, 124
bayeri, 62, 124 hedgpethi hedgpethi, Pleurobranchaea, 48, 49
festiva, Tritonia (Candiella), 104, 209 Hermaea coirala, 152, 237
Fiona pinnata, 109, 124 hillae, 151, 237
fisheri, Coryphella, 226 oliviae, 154
Flabellina, 219, 220, 224, 226 ornata, 154
affinis, 224, 225, 226 vancouverensis, 153
iodinea, 226 Hermaeina, 153
ornata, 225 oliviae, 154
telja, 223, 237 smithi, 153
Flabellinopsis iodinea, 218 Hermissenda crassicornis, 226, 238
flava, Elysia, 33 herthae, Polycera, 198
252
hilaris variety, Thordisa, 91 digue ti, 166
hildae, Onchidella, 230, 237, 238 orbiculata, 147
hillae, Hermaea, 151, 237 perspicua, 11, 124, 238
hirundinina, Chelidonura, 21, 124 perspicua mopsicolor, 11, 148
Hoffmannola, 234 perspicua perspicua, 145
rhombica, 147
hansi, 232, 237
stearnsii, 147
lesliei, 234, 237
lancei, Doto, 215, 216, 238
hombergi, Tritonia, 209
leia, Doriopsilla, 207
Homoiodoris, 66
hulli, Aeolidiella, 116 lesliei, Hoffmannola, 234, 237
hummelincki, Peltodoris, 74 limacina, Titiscania, 7, 124, 125, 145, 238
hummi, Polycera, 198, 200, 237 limbaughi, Cadlina, 169
Hydatina physis, 16 lineata, Coryphella, 226
quoy ana, 16 Duvaucelia, 104
lineolata, Nembrotha, 196
(Aplustrum) velum, 14
Lissodoris, 166
vesicaria, 16, 123, 124
livida, Elysia, 32
Hypselodoris, 56
Lobiger nevilli, 25
acriba, 60, 123 picta, 25
californiensis, 59, 176, 238 pilsbryi, 27
coelestis, 62
serradifalci, 27
edenticulata, 56, 123, 178
souverbiei, 25, 123, 124
gracilis, 60 viridis, 25
sycilla, 59, 178
longicauda, Stylocheilus, 41, 42, 159, 238
luarna, Inuda, 181, 182, 189, 237
ilisima, Berthellina engeli, 160, 237
ilo, Siraius, 66 lurana, Aeolidiella, 115, 125
inca, Phidiana, 110, 111 lutea, Acanthodoris, 202
indecora, Discodoris, 80 lynceus, Phidiana, 109, 110, 111, 123, 124
Tritonia, 103
indica, Aeolidiella, 117 macfarlandi, Chromodoris, 178
inermis, Navanax, 19, 149, 238 macleayi, Aeolidiella, 116
indolens, Onchidella, 121, 122, 230 maculata, Dorididigitata, 186
insulae, Duvaucelia, 104 maculosa, Discodoris, 80
insulae, Tritonia, 104 maderensis, Aglaja, 20
Inuda, 166, 181 Madrella, 217
luarna, 181, 182, 189, 237 marginata, Cadlina, 170
Inudinae, 182 Onchidella, 236
iodinea, Coryphella, 218, 238 Marionia cucullata, 104, 125
Flabellina, 226 occidentalis, 104
Flabellinopsis, 218 marmorata, Anisodoris, 69, 124
iris, Dendronotus, 210, 212, 213 marplatensis, Polycera, 197
Marsenina rhombica, 147
janaina, Doriopsilla, 96, 124, 205, 207, 237 mavis, Discodoris, 187, 238
japonica, Aeolidiella, 118 meckelii, Pleurobranchaea, 45, 47, 48
jheringi, Cyerce, 38 membranaceus, Pleurobranchus, 165
Jorunna, 166 metulifera, Acanthodoris, 203
juliana, Aplysia (Aplysia), 155, 238 Micromelo guamensis, 15
juvenca, Cadlina, 169 undata, 12, 123
Miesea, 215
kaoruae, Cratena, 114, 124 millegrana, Aporodoris, 88, 91
Kentrodoris, 166 minuta, Aglaja, 20
ketos, Tayuva, 192, 237 mizuna, Conualevia, 167
krebsii, Dendrodoris, 95, 123, 124, 203, 237, mopsicolor, Lamellaria perspicua, 11, 148
238 morosa, Pleurobranchaea, 48
krebsii var. pallida, Doriopsis, 95 Nembrotha, 195
Doridopsis, 203 mortenseni, Discodoris, 76, 80, 82
kyolis, Siraius, 64, 124 mulleri, Armina, 108
multicolor, Aeolidiella, 117, 118
ladislavii, Etidoris, 91 muta, Discodoris, 80
laevis, Cadlina, 184 Myrella, 101
lafontii, Phyllaplysia, 40
Lamellaria diegoensis, 147 nana, Aglaja, 21
253
nanus, Dendronotus, 210, 238 pallida, Casella atromarginata, 181
Navanax, 151 Doriopsis krebsii var., 95
aenigmaticus, 19 palmeri, Tritonia, 104, 210
inermis, 19, 149, 238 papilligera, Phyllidiopsis, 99, 123
neapolitana, Spurilla, 118, 124 papillosa, Elysia, 27, 123
Nembrotha capensis, 196 papillosa, Elysia, 29
divae, 196 Paratritonia, 103
eliora, 194, 237 parvula, Aplysia (Pruvotaplysia), 154, 238
gratiosa, 196 patagonica, Berthella, 43
lineolata, 196 Phidiana, 109, 110, 112
morosa, 195 pellucida, Coryphella, 224
nigerrima, 196 Peltodoris, 69, 74
Neodoris, 66, 166 greeleyi, 66, 72, 124
neona, Chromodoris, 51, 124 hummelincki, 74
nevilli, Lobiger, 25 noumeae, 72
nigerrima, Nembrotha, 196 perspicua mopsicolor, Lamellaria, 148, 238
nigra, Aplysia, 159 perspicua perspicua, Lamellaria, 11, 124, 145
nivium, Austrodoris, 186 Petalifera bonairei, 39
nobilis, Anisodoris, 71 ramosa, 39, 124
norrisi, Chromodoris, 170, 237, 238 pharpa, Doriopsilla, 206, 207
notha, Discodoris, 80, 87 Phialodoris, 192
notiperda, Discodoris, 75 Phidiana, 109, 112
Noumea flava, 55 brevicauda, 109, 110
noumeae, Peltodoris, 72 exigua, 110, 111
nyalya, Chromodoris, 53, 58, 124 inca, 110, 111
lynceus, 109, 110, 111, 123, 124
patagonica, 109, 110, 112
obscura, Doridella, 94 pugnax, 110
occidentalis, Aeolidiella, 117 selencae, 109
Marionia, 104 selenkai, 109, 110
Pleurobranchaea, 45, 123 phoca, Discodoris, 78, 124
ocelligera, Aglaja, 20 Phyllaplysia depressa, 40
odhneri, Polycera, 198 engeli, 40, 123
Risbecia, 62 lafontii, 40
odonoghuei, Doris, 186 papilligera, 99, 123
oliviae, Hermaea, 154 Phyllidiopsis (Phyllidiopsis)
Hermaeina, 154 papilligera, 99, 123
Onchidella, 121, 228, 231, 234 Phyllobranchopsis enteromorphae, 153
armadilla, 122 physis, Hydatina, 16
binneyi, 227, 231, 237 pickensi, Doris, 184, 189, 237
borealis, 229, 230 Tritonia (Candiella), 207, 237
carpenteri, 227, 229, 230 picta, Dirona, 216, 238
celtica, 121, 230 Elysia, 32
evelinae, 236 Lobiger, 25
boridana, 119, 123 pilosa, Acanthodoris, 202
hildae, 230, 237, 238 pilosa purpurea, Acanthodoris, 202
indolens, 121, 122, 230 pilsbryi, Lobiger, 27
marginata, 236 pina, Acanthodoris, 201, 237
steindachneri, 121, 122, 123 pinnata, Fiona, 109, 124
Onchidium, 121 pita, Doto, 106
carpenteri, 227, 229, 230 piunca, Coryphella, 222, 226
trans-Atlanticum, 122 Platydoris angustipes, 93, 124
orbiculata, Lamellaria, 147 argo, 94
orbignyana, Aglaja, 20 rubra, 93
plebeia, Candiella, 209
ornata, Austraeolis, 114
Pleurobranchaea agassizii, 49, 123
Coryphella, 224 226
capensis, 47
Flabellina, 225
Hermaea, 154 hedgpethi hamva, 48, 124
meckelii, 45, 47, 48
Otinodoris, 166 morosa, 48
Oxynoe aguayoi, 25 occidentalis, 45, 123
antillarum, 25, 123 Pleurobranchus (Pleurobranchus) areolatus,
254
44, 123, 163, 237 rufibranchialis, Coryphella, 219
(Susania) atlanticus, 44, 163 rufomarginata, Casella, 181
crossei, 44 rumia, Cadlina, 51, 124, 169
digueti, 165
membranaceus, 165 sagamiana, Tambja, 196
plumula, 162 saldanhensis, Aeolidiella, 117
tuberculatus, 165 Samla annuligera, 226
plicatulum, Umbraculum, 42, 123 sandiegensis, Diaulula, 189, 238
plumula, Pleurobranchus, 162 sargassicola, Spurilla, 118
Berthella, 162 scabriuscula, Cadlina, 51, 170
scops, Elysia cauze, 28, 123
Polycera, 197 (key)
Sebadoris, 166
alabe, 198
sedna, Casella, 178, 237
atra, 196, 198, 238
selencae, Phidiana, 109
aurisula, 198, 237
selenkai, Phidiana, 109, 110
chilluna, 198
semperi, Armina, 108
gnupa, 198, 237
serradifalci, Lobiger, 27
hedgpethi, 198, 199, 200
seurati, Aglaja, 20
herthae, 198
Siraius, 66, 166
hummi, 198, 200, 237
ilo, 66
marplatensis, 197
kyolis, 64, 124
odhneri, 198
smithi, Hermaeina, 153
priva, 198
zosterae, 198 sonora, Chromodoris banksi, 173, 237
souriei, Aporodoris, 91
prea, Anisodoris, 70, 124
souriei, Thordisa, 91
pricei, Coryphella, 223
priva, Polycera, 198 souverbiei, Lobiger, 25, 123, 124
sparsa, Cadlina, 169
pruvotfolae, Elysia, 34
spetteda, Discodoris, 75, 76
pugnax, Phidiana, 110
splendens, Elysia, 32
punctilucens, Aglaja, 20
Spongiodoris, 167
Chromodoris, 176
Spurilla, 117
punctuolata, Anisodoris, 69
braziliana, 118
purcina, Discodoris, 80, 81, 123
chromosoma, 227, 237, 238
purpurea, Aglaja, 20
neapolitana, 118, 124
pusa, Aglaja, 18, 124
sargassicola, 118
pusae, Discodoris, 80, 82, 124, 189, 238
stearnsii, Lamellaria, 147
steindachneri, Onchidella, 121, 122, 123
quadridens, Berthellina, 43, 123, 163
stelliferca, Archidoris, 69
quoyana, Hydatina, 16
Stiliger fuscovittatus, 154
vancouverensis, 153
ramosa, Petalifera, 39, 124 strongi, Haminoea, 149
reticulata, Doriopsis, 99, 204 Stylocheilus citrinus, 40, 124
reticulata, Risbecia, 62 longicauda, 41, 42, 159, 238
Tritonia, 102 subpellucida, Dendrodoris, 96
reticulopoda, Aplysia (Aplysia), 158 Susania gardineri, 44
rhodoceras, Acanthodoris, 202 sycilla, Hypselodoris, 59, 178
rhombica, Marsenina, 147
Lamellaria, 147
risbeci, Aporodoris, 91 takanosimensis, Aeolidiella, 117
Risbecia, 62, 166 Tambja sagamiana, 196
frangoisi, 62 Taringa, 91, 167, 237
odhneri, 62 aivica, 89, 124
reticulata, 62 aivica timia, 182, 184, 189, 204
versicolor, 62 armata, 88, 91
robustus, Dendronotus, 212 telopia disa, 87, 124
roseo-picta, Chromodoris, 61 Tayuva, 167, 191
Rosodoris, 166 ketos, 192, 237
rowena, Doriopsilla, 205, 237 telja, Flabellina, 223, 237
rubens, Discodoris, 80 telopia disa, Taringa, 87, 124
rubra, Aporodoris, 91 telopia telopia, Taringa, 87, 88
Platydoris, 93 tesselata, Dictyodoris, 191
rubra, Casella, 181 Thordisa, 91
rubrolineata, Godiva, 112, 124, 125 clandestina, 91
255
diuda, 91 tuca, Elysia, 29, 123
dubia, 91 tupala, Berthella, 43, 124
hilaris variety, 91 tura, Chromodoris, 55
souriei, 91 Tylodina, 21
Thorunna, 167
timia, Taringa aivica, 182, 184, 189 Umbraculum, 21
timorensis, Anisodoris, 70 bermudense, 42, 123
Titiscania limacina, 7, 124, 125, 145, 238 plicatulum, 42, 123
trans-Atlanticum, Onchidium, 122 undata, Micromelo, 12, 123
tricolorata, Aglaja, 19, 20 universitatis, Chromodoris, 176
Tridachia crispata, 33, 35, 37, 123, 124, 237 uva, Doto, 108
Tridachiella diomedea, 34, 124, 154, 237
trilineata, Coryphella, 222, 226 vaccaria, Aplysia, 158
tristis, Discodoris, 80 vancouverensis, Hermaea, 153
Tritonia, 101, 104, 209 vancouverensis, Stiliger, 153
(Candiella) bayeri, 101, 124 varians, Doto, 214
exsulans, 123 velum, Hydatina (Aplustrum), 14
(Candiella) festiva, 104, 209 verrilli, Elysia (Elysiopterus), 34
hombergi, 209 verrucosa, Doris, 186, 237
indecora, 103 versicolor, Risbecia, 62
insulae, 104 vesicaria, Hydatina, 16, 123, 124
palmeri, 104, 210 villafranca, Tritonia, 104
(Candiella) pickensi, 207, 237 virescens, Haminoea, 149, 238
reticulata, 102 virescens rosacea, Haminoea, 149
villafranca, 104 viridis, Lobiger, 25
(Tritonidoxa) wellsi, 99, 125 voniheringi, Discodoris, 80, 87
Tritonidoxa, 101, 209
Tritoniopsis, 103 Watsoniella, 236
elegans, 103 wellsi, Tritonia (Tritonidoxa), 99, 125
Tochuina, 209 worki, Anisodoris, 66, 124
trophina, Coryphella, 226
tuberculata, Archidoris, 68 zebra, Chromodoris, 60
tuberculatus, Pleurobranchus, 165 zosterae, Polycera, 198
256