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, NOS. 1-2 1983
MALACOLOGIA
MUS. COMP. ZOOL
LIBRARY
>
НАКУАКО
UNIVERSITY
ternational Journal of Malacology
¿A : 3
Revista Internacional de Malacologia
_ Internationale Malakologische Zeitschrift
MALACOLOGIA
Editors-in-Chief:
GEORGE M. DAVIS
ROBERT ROBERTSON
Editorial and Subscription Offices:
Department of Malacology
The Academy of Natural Sciences of Philadelphia
Nineteenth Street and the Parkway
Philadelphia, Pennsylvania 19103, U.S.A.
Associate Editors:
JOHN B. BURCH
University of Michigan, Ann Arbor
ANNE GISMANN
Maadi, A. R. Egypt
Editorial Assistants:
MARY DUNN
DAVID WATT
MALACOLOGIA is published by the INSTITUTE OF MALACOLOGY (2415 South Circle
Drive, Ann Arbor, Michigan 48103, U.S.A.), the Sponsor Members of which (also serving as
editors) are:
CHRISTOPHER J. BAYNE
Oregon State University, Corvallis
KENNETH J. BOSS
Museum of Comparative Zoölogy
Cambridge, Massachusetts
JOHN B. BURCH
MELBOURNE R. CARRIKER
University of Delaware, Lewes
GEORGE M. DAVIS,
Secretary and Treasurer
PETER JUNG, Participating Member
Naturhistorisches Museum, Basel, Switzerland
OLIVER E. PAGET, Participating Member
Naturhistorisches Museum, Wien, Austria
ROBERT ROBERTSON
CLYDE F. E. ROPER
Smithsonian Institution
Washington, D.C.
W. D. RUSSELL-HUNTER, President-Elect
Syracuse University, New York
NORMAN F. SOHL
United States Geological Survey
Washington, D.C.
SHI-KUEI WU, President
University of Colorado Museum, Boulder
J FRANCES ALLEN, Emerita
Environmental Protection Agency
Washington, D.C.
ELMER G. BERRY, Emeritus
Germantown, Maryland
Copyright © 1983 by the Institute of Malacology
1983
EDITORIAL BOARD
J. A. ALLEN
Marine Biological Station,
Millport, United Kingdom
E. E. BINDER
Muséum d'Histoire Naturelle
Genève, Switzerland
A. J. CAIN
University of Liverpool
United Kingdom
P. CALOW
University of Glasgow
United Kingdom
A. H. CLARKE, Jr.
Mattapoisett, Mass., U.S.A.
B. C. CLARKE
University of Nottingham
United Kingdom
С. J. DUNCAN
University of Liverpool
United Kingdom
Z. A. FILATOVA
Institute of Oceanology
Moscow, U.S.S.R.
E. FISCHER-PIETTE
Muséum National d'Histoire Naturelle
Paris, France
\. ERETTER
University of Reading
United Kingdom
E. GITTENBERGER
Rijksmuseum van Natuurlijke Historie
Leiden, Netherlands
A. N. GOLIKOV
Zoological Institute
Leningrad, U.S.S.R.
S. J. GOULD
Harvard University
Cambridge, Mass., U.S.A.
A. V. GROSSU
Universitatea Bucuresti
Romania
T. HABE
Tokai University
Shimizu, Japan
A. D. HARRISON
University of Waterloo
Ontario, Canada
K. HATAI
Tohoku University
Sendai, Japan
B. HUBENDICK
Naturhistoriska Museet
Göteborg, Sweden
S. HUNT
University of Lancaster
United Kingdom
A. M. KEEN
Stanford University
California, U.S.A.
R. N. KILBURN
Natal Museum
Pietermaritzburg, South Africa
M. A. KLAPPENBACH
Museo Nacional de Historia Natural
Montevideo, Uruguay
J. KNUDSEN
Zoologisk Institut & Museum
Kobenhavn, Denmark
A. J. KOHN
University of Washington
Seattle, U.S.A.
Y. KONDO
Bernice P. Bishop Museum
Honolulu, Hawaii, U.S.A.
J>EEVER
Amsterdam, Netherlands
A. LUCAS
Faculté des Sciences
Brest, France
N. MACAROVICI
Universitatea “Al. I. Cuza’’
lasi, Romania
C. MEIER-BROOK
Tropenmedizinisches Institut
Túbingen, Germany (Federal Republic)
H. K. MIENIS
Hebrew University of Jerusalem
Israel
J. E. MORTON
The University
Auckland, New Zealand
R. NATARAJAN
Marine Biological Station
Porto Novo, India
J. VKLAND
University of Oslo
Norway
T. OKUTANI
National Science Museum
Tokyo, Japan
W.L. PARAENSE
Instituto Oswaldo Cruz, Rio de Janeiro
Brazil
J. J. PARODIZ
Carnegie Museum
Pittsburgh, U.S.A.
W.F. PONDER
Australian Museum
Sydney
А. W. В. POWELL
Auckland Institute 4 Museum
New Zealand
R. D. PURCHON
Chelsea College of Science £ Technology
London, United Kingdom
O. RAVERA
Euratom
Ispra, Italy
N. W. RUNHAM
University College of North Wales
Bangor, United Kingdom
$. С. SEGERSTRÁLE
Institute of Marine Research
Helsinki, Finland
G. A. SOLEM
Field Museum of Natural History
Chicago, U.S.A.
F. STARMÜHLNER
Zoologisches Institut der Universität
Wien, Austria
У. |. STAROBOGATOV
Zoological Institute
Leningrad, U.S.S.R.
WSTREIRE
Université de Caen
France
J. STUARDO
Universidad de Chile
Valparaiso
T. E. THOMPSON
University of Bristol
United Kingdom
Е. TOFFOLETTO
Societa Malacologica Italiana
Milano
В. О. TURNER
Harvard University
Cambridge, Mass., U.S.A.
W.S.S. VAN BENTHEM JUTTING
Domburg, Netherlands
J. A. VAN EEDEN
Potchefstroom University
South Africa
J.-J. VAN MOL
Université Libre de Bruxelles
Belgium
N. H. VERDONK
Rijksuniversiteit
Utrecht, Netherlands
B. R. WILSON
National Museum of Victoria
Melbourne, Australia
C. M. YONGE
Edinburgh, United Kingdom
H. ZEISSLER
Leipzig, Germany (Democratic Republic)
A. ZILCH
Natur-museum und Forschungs- Institut
Senckenberg
Frankfurt-am-Main, Germany (Federal
Republic)
MALACOLOGIA, 1983, 24(1-2): 1-113
TAXONOMIC STUDIES ON GYRAULUS (GASTROPODA: PLANORBIDAE)'
Claus Meier-Brook
Tropenmedizinisches Institut der Universität, Wilhelmstrasse 31,
D-7400 Tübingen, Federal Republic of Germany
ABSTRACT
From 94 mostly Eurasiatic samples, alcohol material of Gyraulus snails was compared con-
chologically and 492 specimens were dissected for anatomical characteristics. Infraspecific
shell variation is shown to depend on, among other things, water movement. G. albus (Müller)
becomes larger in lenitic biotopes than animals with equal whorl numbers in lotic biotopes. In
animals killed by immersion in ethanol particularly the length of the penis sheath is shorter
than in snails relaxed with pentobarbital.
Characters highly valued for species discrimination are among others: the distribution of
pigment cells on the mantle to the right of the kidney; the presence or absence of distinct
transverse septa in the tubular portion of the kidney, rendering its margins “undulate” or
“straight,” respectively; the number of prostate diverticula and their shape and arrangement
in the gland; the width of the vas deferens; the length ratio penis sheath : preputium; the shape
of the penis tip and stylet, and the position of the penis pore. In certain groups peculiarities
occur in various other organs (radula, seminal vesicle, bursa copulatrix, bursa duct, etc.).
Europe harbours five indigenous species of the genus plus six endemic species in Macedo-
nian ancient lakes. North Asia is inhabited by Gyraulus acronicus (Férussac) only, while south
Asia harbours a ““Rassenkreis,” i.e. а. chinensis (Dunker) (commonly used junior synonyms:
G. convexiusculus (Hutton), G. spirillus (Gould)) with at least eight peripheral isolates. Two new
species are described, i.e. G. eugyne from Inner Mongolia and G. malayensis from Malaya,
also occurring in Java and Bali. Abandonment of planispiral growth and formation of several
angles or keels on the shell periphery in Gyraulus of the ancient lakes is certainly due to
convergent evolution. There is strong evidence from anatomy that a species endemic to Lake
Biwa in Japan has a history different from that in a group endemic to the Macedonian Lakes
Ohrid and Prespa. Two keys are presented: one for identification of the species living in Eu-
rope, the second a provisional one for the west, south, and east Asiatic species.
Analysis of relationships according to Hennig's ‘‘consequent phylogenetic systematics”’ re-
vealed the existence of seven subgenera, ¡.e. Torquis Dall (Nearctic and west Palaearctic),
Lamorbis Starobogatov (west Palaearctic), Armiger Hartmann (Holarctic), Caillaudia Bourguig-
nat (Ethiopic), Choanomphalodes Lindholm (Lake Biwa, Japan), Carinogyraulus Polinski (Lakes
Ohrid and Prespa, Macedonia), and a provisional ‘‘subgenus Gyraulus $. str. Charpentier”
(cosmopolitan except South America and the Ethiopic region). Gyraulus s. str., being a remnant
group merely characterized by symplesiomorphies and autapomorphies, cannot be established
as a monophyletic taxon. According to Hennig its paraphyletic character is not excluded. Hen-
nig's arguments appear to leave such remnant groups as a general problem of his systematics.
In the case of ‘‘Gyraulus $. str.’’ only chorological implications can help to support monophy-
letic status in Hennig's sense. Incompatibility is shown between Hennig's ‘‘deviation rule” and
biological reality in the case of the ‘‘Rassenkreis’’ of G. chinensis, which is considered to be
the ‘‘persisting stem species” of the marginal species that have evolved as peripheral isolates.
The centre of origin and the primary centre of differentiation is probably the west Palaearctic
to which all the genera closely related to Gyraulus are confined. A secondary centre of differ-
entiation is the Nearctic, a third one the Indo-Malayan region. A redescription of the genus
Gyraulus Charpentier, 1837, is given using present knowledge on variation of anatomical and
conchological characters.
INTRODUCTION of “species” in the planorbid genus Gyraulus
Charpentier, 1837. For the South African re-
As in other molluscan groups, diversity of gion, Brown & van Eeden (1969) have pub-
shell characters has led to excessive naming lished an outstanding revision of Gyraulus
' Habilitationsschrift, Fachbereich Biologie der Eberhard-Karls-Universitat zu Tübingen, 1977.
(1)
2 MEIER-BROOK
species; the species endemic to Lake Ohrid
(SE Europe) have been revised by Hubendick
& Radoman (1959); three species of central
Europe have been compared anatomically by
Meier-Brook (1964). A synoptic view of Eu-
rasiatic forms was, however, still lacking, al-
though repeatedly called for, particularly by
those interested in medical malacology (Burch
& Natarajan, 1965; Pace, 1973).
The health impact of Gyraulus is their röle
as intermediate hosts of the intestinal flukes
Echinostoma ilocanum Odhner, 1911 and E.
lindoense (Sandground & Bonne, 1940),
which are pathogenic to inter alia man.
The number of names created for Eura-
siatic species or subspecies of the genus, or
later assigned to it, is not far from 200, and
only recently three new species have been
described from Thailand (Brandt, 1974).
The goal of the present investigation was
to find out to what extent conchological vari-
ation is paralleled by anatomical variation.
Consequently the studies had to be confined
to instances where living or preserved ani-
mals were available. A revision of North
American forms must be postponed since the
huge American collections would require
years of study. Moreover, American species
have so far not been incriminated as hosts of
human helminths. In South America there is
only one species of this genus (Paraense,
personal communication) and the Australian
region will be covered by studies now being
undertaken by Dr. D. S. Brown. It would be
premature to expect a complete revision and
list of synonyms, but | hope that the present
study will serve as a basis for a revision where
results of further studies can find their appro-
priate places. Great gaps in knowledge re-
main from central Asia, viz. Siberia and the
Peoples Republic of China, where | could col-
lect some material but was prevented from
taking it out of the country.
Material to be studied and compared was
selected according to the following criteria:
planorbid snails with a hollow stylet on their
penis and having a shell with three to four
and a half (at most five) rapidly increasing
whorls. These features have been accepted
as characteristic of the genus Gyraulus by all
authors who have used this name (Baker,
1945; Hubendick, 1955, etc.).
MATERIALS AND METHODS
Snails were collected alive by various per-
sons at the localities indicated in the text un-
der ‘‘Material.’’ The animals were usually sent
alive and prepared for study at the laboratory
of the Institute of Tropical Medicine of Tü-
bingen University; in some cases animals
were relaxed, fixed and preserved by the col-
lectors, as indicated in the text. Where fresh-
ly collected material was not available, ¡.e.
from the U.S.S.R. and the Peoples Republic
of China, | had to rely on old museum collec-
tions.
Unless stated otherwise, living snails were
prepared as follows: They were relaxed in a
freshly prepared suspension of 5 mg or more
pentobarbital in 10 ml of tap water at room
temperature, as described by Meier-Brook
(1976a). After 6 to 12 hr they were killed and
fixed in 4% formalin at appr. 60°C and trans-
ferred to 70% ethanol until dissection. This
was done under a Wild Zoom Stereomicro-
scope М7 using pointed forceps, microscis-
sors after Vannas, ophthalmologic scissors,
and insect needles of the sizes 0 to 000.
Camera lucida drawings were done with a
Wild drawing tube. Whole mounts of organs
and organ sytems were made on microscope
slides in Eukitt. For histological studies or-
gans were removed from the body, dehy-
drated in a series of graded ethanol, soaked
with methyl benzoate (2 x 15 min), benzolum
(30 min), benzolum/histoplast-S (56-57°C,
Serva) (30 min) and embedded in histoplast-
S after 8 hr. Serial sections 7 um thick were
made with a Leitz-microtome and stained in
haemalum and azophloxine. When organs
were too small for exact orientation during
embedding, they were stained in an alcoholic
solution of safranin prior to dehydration.
Chromosome Counts.—For determination
of chromosome numbers, ovotestes and em-
bryos were used in some cases when living
snails were available. Ovotestes were ex-
cised in tap water where they remained for
15 to 30 min to make cells swell and thus
make better chromosome spreads. Embryos
were freed from their egg capsules and treat-
ed in the same way.
For fixation, tissues were transferred to a
mixture of acetic acid and 96% ethanol, 1:3
parts by volume (according to Patterson,
1971). After 5 min they were stained in a con-
centrated acetic orcein solution for up to 8 hr
and differentiated in 45% acetic acid. Prepa-
rations were examined in 45% acetic acid af-
ter squashing tissues in a screw clamp.
Counts were made under a Zeiss interfer-
ence microscope at 1200x.
SEM-Studies.—Snails were relaxed in a
suspension of pentobarbital for 1 hr, fixed at
GYRAULUS TAXONOMY (PLANORBIDAE) 3
room temperature in a 2% glutaraldehyde so-
lution, in which the body cavity was opened
immediately after immersion. For studies of
the penis tip, the penis sheath was cut and
the penis drawn out. The organs were fixed
for 2-4 days, rinsed in distilled water for 4 x
5 min, dehydrated in 30, 50, 70, 90 and 100%
acetone for 10 min each and dried in an ISI
critical point drying (CPD) apparatus over
carbon dioxide. Critical point drying was per-
formed in the Cytological Laboratory of the
Institute of Tropical Medicine, Tübingen, by
Dr. H. M. Seitz. SEM studies were performed
with a Cambridge Stereoscan Microscope
MK2A after coating with C + Pd/Au.
CHARACTERS USED IN
GYRAULUS TAXONOMY
The biological species concept (Scudder,
1974) is advocated in taxonomical revisions
at the infrageneric level. It is subject to a min-
imum of controversies aside from dealing with
fossils and problems stemming from unipa-
rental reproduction. As every taxonomist
knows, however, very few species would be
accepted at all if biologists relied on only this
species concept. In planorbid gastropods, the
fact that all members can self-fertilize poses
severe problems from crossing experiments.
Only in a few cases does a genetic marker,
e.g. recessive albinism, enable one to distin-
guish between the results of self- and cross-
fertilization (Richards, 1973b). In most groups,
such as Gyraulus, albinotic individuals have
not yet been found.
The biological species, however, is not de-
fined only as “the largest and most inclusive
reproductive community of sexual and cross-
fertilizing individuals that share in a common
gene pool” (Dobzhansky, 1950); “the es-
sence of the biological species concept is dis-
continuity due to reproductive isolation”
(Mayr, 1974a: 379). Evidence of reproductive
isolation between sympatric populations
serves to indicate that different species are
involved even if other types of data are un-
available.
Examples of reproductive isolation be-
tween species of Gyraulus hitherto charac-
terized as morphological species are regular-
ly encountered in Europe. In seven of the nine
localities where | collected G. laevis (Alder),
it lived together with G. albus (Müller), and
not only in the same water body, but in the
same square meter. In all of the eight central
European lakes where | found G. acronicus
(Férussac) this species coexisted with G. al-
bus. G. riparius (Westerlund), too, was found
associated with G. albus, and the only Euro-
pean species regularly living alone is the de-
cidedly stenoekous G. rossmaessleri (Auers-
wald) confined to temporary pools. Wherever
sympatric species of the genus were exam-
ined | never saw intermediate anatomical
traits indicating hybridization. Doubts arose
when only shell characters were considered.
But even if hybrids occurred and these were
fertile, this would not necessarily affect our
statement on reproductive isolation. ‘‘What
matters is not whether hybrids can be ob-
tained but whether the Mendelian popula-
tions do or do not exchange genes, and if
they do whether at a rate which destroys the
adaptive equilibrium of the population con-
cerned” (Dobzhansky, 1950: 415 et seq.).
Sympatry of pairs of European species of
course should not be taken as evidence for
sympatric speciation. On the contrary, in the
pairs Gyraulus albus-laevis and G. albus-ri-
parius relationships appear remote enough to
regard them as members of different subgen-
era. Although we are far from recognizing the
mechanism of reproductive isolation between
them, differences in form and size of the male
copulatory organs, as shown later, make
copulation unimaginable. This is also the case
in the pair G. albus-acronicus where differ-
ences in size (Tables 1 and 2) though not in
general morphology are pronounced. There
is no reason to assume that events leading
to separate species and subsequently to
separate subgenera did not occur in geo-
graphical isolation. Allopatric speciation with
a later reunion of geographic ranges may be
the rule in cases where closely related species
coexist, although the possibility of sympatric
speciation, for example by disruptive selec-
tion (Mayr, 1963: 156; Sperlich, 1973: 52-
53, 155), can no longer be denied (Scudder,
1974).
A different situation will be shown for a pair
of jointly occurring species that have virtually
no anatomical differences: Gyraulus chinen-
sis (Dunker)- tokyoensis (Mori). Here evi-
dence of reproductive isolation favours the
decision for separate species.
In the majority of samples from allopatric
populations comparison and evaluation of
morphological characters, supported by bio-
logical and ecological observations, remain
the only practical methods for species dis-
crimination. This approach is considered jus-
tifiable as long as the following, expressed by
Mayr (1974a: 381), is kept in mind: ““Repro-
4 MEIER-BROOK
FIG. 1.
Dimensions measured (A) and method of
counting whorls (B) in Gyraulus. Example: G. lae-
vis, Kuehren and Ansbach.
ductive isolation is effected by physiological
properties which have a genetic basis. Mor-
phological characters are the product of the
same gene complex,” and as long as priority
is given to reproductive isolation: ‘This infer-
ence method is by no means a return to a
С euphraticus, $ Iran
b n=73
D
EIGH2:
morphological species concept since repro-
ductive isolation always remains the primary
criterion and degree of morphological differ-
ence only a secondary indicator, which will
be set aside whenever it comes in conflict
with the biological evidence.”
Morphological characters are subject to
variation for various reasons: allometric and
seasonally different growth, preparation-de-
pendent changes, and, of course, genotypi-
cally- and ecophenotypically-determined ones.
Although the latter two reasons cannot be
separated in a morphological analysis, | later
include several paragraphs titled ‘’Non-Ge-
netic Variation,’ where undoubted cases of
externally induced aberrations are dealt with.
However, the inclusion of such paragraphs
does not imply that the preceding para-
graphs are confined to genetic variation!
Shell Dimensions
Absolute dimensions usually given in tax-
onomical studies of planorbids are the height
and the maximum diameter, height and width
of the aperture and the number of whorls. As
a measure of flatness the ratio maximum di-
ameter: height is generally indicated. One
problem arising in Gyraulus and some other
genera is caused by the tendency in some
groups to deflect the last whorl near the ap-
erture. A slight increase of the deflection an-
gle would result in a highly increased height
a
Dimension b plotted against a in a Gyraulus.
GYRAULUS TAXONOMY (PLANORBIDAE) 5
G. euphraticus .
S-Iran
1 2 3 mm
FIG. 3. Graph with a/b plotted against a and
regression line. Same sample as Fig. 2. п = 67 (+6
juv. neglected); y = 0.84 + 0.85х (y = = х =
а); г = 0.921.
figure and thus in а falsification of the “flat-
ness index.” This is avoided by indicating the
maximum diameter minus aperture and the
height of the last whorl in its middle as seen
in apertural view (Fig. 1A, a and b). These
dimensions were first used by Brown & van
Eeden (1969: figs. 16, 17), who, moreover,
plotted the ratio a/b against a, arguing that
the degree of flattening of the whorl during
(allometric) growth is thus best demonstrat-
ed. These authors have not commented on
the question of linearity of such a function.
Growth is allometric since individuals are
proportionally high in youth and become pro-
portionally lower during growth (Fig. 2). Plot-
ting absolute dimensions, viz. b against a,
yields a distribution of points probably follow-
ing a hyperbolic function (personal commu-
nication by Dr. W. U. an der Heiden, Inst.
Biomathematics, Tübingen). Although well-
fitting curves in such a graph would give a
clear idea of the course of growth, it is im-
practical to construct them or to find appro-
priate equations that help in comparing pop-
ulations. Calculating the ratios a/b and
plotting these against a indeed appears to be
better suited for these purposes, because at
Gyraulus spirillus
Taiwan
1 2 3 4 5 тт
а
FIG. 4. Same as Fig. 3 but with another species.
п= 45; y = 1.55 + 0.73x (y = = x =a); r= 0.924.
а > 1 mm, points apparently follow a linear
function. Hence regression lines can be drawn
and the corresponding equations, y = f + g-X,
can be tabulated and compared. The higher
the inclination, indicated by g, the lower the
relative increase of dimension b, or the angle
between upper and under sides in the grow-
ing shell. The higher f, indicating the position
of the line, the flatter the shell as a whole
(Fig. 109). Linearity of the regression line in a
coordinate system with a/b as the y-axis and
a as the x-axis was examined by the F-test
(Sachs, 1969: 421 et seq.). In the two ex-
amples (Figs. 3, 4), the critical F-values at the
5%-level (Fig. 3) or at the 10%-level (Fig. 4)
were not attained or exceeded; therefore the
test did not disprove the linearity of the
regression function. Regression lines are
compared in cases where critical F-values on
the 5%-level are not reached.
There has been some controversial use of
the terms upper and under side, due to dif-
ferent orientations—some authors handle
planorbid shells as sinistral, most as dextral.
It has long been known that all Planorbidae
are morphologically sinistral (compare Physi-
dae and Bulininae): genital openings and the
anus are on the left. Consequently all Gyrau-
lus snails are here considered sinistral.
The number of whorls is determined as
usual (Fig. 1B) (Mandahl-Barth, 1954, fig. 2;
Pan American Health Organization, 1968: fig.
7). Dimensions, proportions and whorl num-
bers are important for characterizing species,
particularly since they are the main concho-
logical features that, in earlier years, formed
the only basis for taxonomy.
As in other groups of fresh-water mol-
6 MEIER-BROOK
1 2 3 4 mm
a
FIG. 5. Variation of a/b plotted against a in two
similar biotopes (fish ponds in Holstein, N Ger-
many).
luscs, variation of shell characters within a
population is much less than between neigh-
bouring populations, populations being de-
fined as communities of individuals actually
interbreeding. Hubendick (1951) convincingly
demonstrated this for lymnaeid snails by his
“mean photographs.”
Intraspecific variation demonstrable with
this technique is much less in Planorbidae
than in high-spired snail groups, but propor-
tions in two populations of presumably simi-
lar biotopes may prove to be so different that
their ranges overlap only marginally (Fig. 5).
Interpopulational variation of proportions in
one species in lakes and in ponds or back-
waters is furthermore demonstrated in Fig. 7.
Restricted variation is seen particularly in
small water bodies where often one or a few
individuals have founded a population, which,
consequently, has a restricted gene pool (as
in cases where the founder-principle be-
comes effective). Genetic isolation resulting
in “a homogeneous genetic constitution” is
favoured by self-fertilization of which basom-
matophoran snails generally are capable (Hu-
bendick, 1951: 31, 32), but probably also by
a certain island character of small water bod-
ies (Lassen, 1975; Keddy, 1976; Aho, 1978).
The hostile land masses separating ponds
and pools may reduce gene flow between
limnic populations as compared with gene
flow in the vast and continuous populations
of many terrestrial animal species. Phenotyp-
ic similarity between individuals of such small
populations may, furthermore, be caused by
the influence of environmental factors that act
upon all of them largely to an equal degree,
either through selection (e.g. founder effect,
as understood by MacArthur & Wilson, 1967:
154, 188) or immediate influence (within the
range of reaction norm) upon the growth of
individuals.
Non-Genetic Variation of Dimensions
and Proportions
Non-genetic variation is due to environ-
mental factors. For any given population it is
difficult to assess the influence of selection,
as distinct from ecologically induced non-in-
herited influences, on shell phenotype. For
some factors, however, direct action on
growth is Known. For fresh-water molluscs
factors most interacting with shell develop-
ment are water movement and water chem-
istry, particularly the calcium content. Cal-
cium deficiency, if not inhibiting occurrence of
certain species (Meier-Brook, 1963), may
cause shells to become thin and fragile (Hu-
bendick, 1947: 503; Meier-Brook, 1978). Shell
thickness in the widely distributed species of
Gyraulus is, however, regarded not to be sig-
nificant, neither for taxonomic conclusions nor
merely for identification.
Water movement in standing waters gen-
erally varies with wind exposure. This means
that snails of larger lakes (lotic biotopes) are
usually more exposed to water movement
than are those in ponds and pools (lenitic bio-
topes). Reduced shell size in lakes, as shown
for Gyraulus albus (Figs. 6 and 7), is a plau-
sible Consequence, because the planispiral
shell offers a large surface to waves. Three
ways of size reduction in lotic biotopes are
imaginable: (1) Growth generally equals that
in lenitic biotopes but individuals die earlier.
In this case numbers of whorls must be re-
duced as well; (2) Snail growth is reduced,
because selection has favoured individuals
whose diameter grows more slowly; (3) Snail
growth is hampered by direct physiological
(=non-genetic) action of the environment on
each individual. In cases (2) and (3) it could
be that whorl numbers are not reduced as
compared with snails from still water; then
whorls must be narrower and less rapidly in-
creasing.
Fig. 6 depicts ratios of whorl number : di-
ameter in Gyraulus albus from some lakes
and ponds shows that in these samples sit-
uation (1) has not been realized. Whorl num-
bers are not smaller than in ponds. At least
when high whorl numbers (around 4) are
reached, pond dwellers have attained more
or less greater diameters than lake dwellers.
A decision on how far situations (2) or (3) are
GYRAULUS TAXONOMY (PLANORBIDAE) 7
A
6 G.albus > В
д - Westensee y
+ - Kührener + + а ss
5 Viehteich 4 5 ge A lab.offspr. A
+ - backwater RhineR. +} 0
+ +8
4 | y
) 5 A
5 2 >
© +
= ü 3| os
E - ой
: a7 :
E = $
Е Е
Е =
— +— == +
0 1 2 В 4
whorls
© D
ANCSIbUs G. albus
o - Selenter See x Bodensee: +
6 x - Kasseeteich Xu 6 | H
yx o - Untersee y
ххх + - Wollmat. Ried on
5 y 5 A
к” In.
A So”
о o o a
ae ei” Sd A
= о. 9 gt
© o Ae 8 oy y ы в
E ET Е A
Le) 3 AG x © 3 + я
Le) ge 19) do
x Bo e
© 2 a | |
E Br бо Я - o
= =
EMI za:
4 —+— + —+——_ —— + + + А
0 | 2 3 4 0 1 Ze 3 4 5
whorls whorls
FIG. 6A-D. Comparison of the ratio maximum diameter : whorl number in lotic (0) and lenitic (x, +) bio-
topes. According to the Hotelling-Test (Anderson, 1958) differences between the centroids are highly
significant (p < 0.001).
8 MEIER-BROOK
TO ID
1 2 3 4mm
1 2 3 4 mm
a
FIG. 7. Gyraulus albus. Comparison of shell proportions between lotic (——) and lenitic (----- ) biotopes;
A: a/b plotted against a; В: b plotted against a. 1—Haslacher See; 2—Selenter See; 3—Titisee; 4—
Zürichsee; 5—Lac St. Point; 6 —Westensee; 7—Ettenheim; 8—Kasseeteich; 9—Wollmatingen; 10—
Kuehren.
involved in size reduction is not possible 6B) suggest that reduction of shell size is to
without laboratory breeding experiments. Dif- some extent reversible and, thus, not inher-
ferences between snails collected in Zúrich- ited. How far genetic fixation of a reduced
see and their offspring reared in aquaria (Fig. shell size in lakes has taken place in other
GYRAULUS TAXONOMY (PLANORBIDAE) 9
populations cannot yet be stated. Neither can
the rôle of other ecological conditions, e.g.
food, be overlooked at present. Large lakes,
such as the Bodensee, offer an opportunity
for comparing samples from different habi-
tats of the same water body. In Fig. 6D a
sample from a wind-exposed shore in Unter-
see is compared with one from a bay very
rich in vegetation (Phragmition and Pota-
mion) and mud. The two biotopes, approxi-
mately 5 km from one another are in the same
water body. They are lotic and lenitic with
broad overlap. However, the difference visi-
ble in the other examples is also distinct here,
especially in snails with three and four whorls.
Probably a steady gene exchange between
the two localities takes place, at least in one
direction: from Wollmatinger Ried to Nieder-
zell/Untersee a strong current occurs, ¡.e. to-
ward the outflow of the Rhine River. The
strange aberration of six specimens that have
attained large diameters despite a low whorl
number must thus far remain unexplained.
The sample size was not sufficient to support
the real existence of a gap between two por-
tions or perhaps generations in the popula-
tion.
Fig. 6A-C shows, furthermore, that varia-
tion of the ratio is greater in lakes than in
ponds and other small water bodies. This can
have two causes: (1) a larger gene pool in
lakes as compared with that in small water
bodies (see above: founder principle!), and (2)
the fact that every lake has, besides lotic mi-
crohabitats, also lenitic ones where vegeta-
tion or stones protect animals from direct surf
or wave action. Therefore lakes have a great-
er variety of ecological conditions than ponds,
etc., and, consequently, a greater range of
ecophenotypes.
One of the other parameters underlying
variation is the flatness index (Fig. 7). A com-
parison of lakes (solid lines) with ponds and
similar lenitic biotopes (broken lines) does not
bring to light significant differences in the po-
sition of regression lines. But there is some
evidence that regression coefficients are
higher in lakes. What does this mean in terms
of shell morphology? In each two examples
of lake and pond populations the corre-
sponding curves in a system with a and b as
the coordinates (Fig. 7B) show that young
snails are slightly higher in lakes than in
ponds. The necessity for young and weak
snails to increase musculature for increased
resistance to moving water may be the cause:
bigger muscles in the cephalopedal mass re-
quire a larger volume of the last whorl (the
so-called body-whorl) since in danger the lat-
ter must be able to take up the whole head-
foot complex. For some time b grows well
with growing a, but later growth of b is nec-
essarily lowered (otherwise growth would re-
sult in snails with two completely conical um-
bilici). Reduced growth of b gradually takes
place when a certain number of whorls is
reached. Lake dwellers then usually have a
smaller diameter than pond dwellers, as
shown above (Fig. 6).
Extreme growth was attained in the labo-
ratory by reducing population density and of-
fering optimal nutrition (food according to
Standen, 1949). These giant forms are rarely
encountered in nature. Field-collected giant
individuals were in some cases trematode-in-
fected. Parasitic castration may be the cause
of giantism. Deformations such as open coil-
ing eventually occur. They may be caused by
external or internal (genetic) factors and are
left out of further consideration here.
Shell Surface
The presence or absence of a peripheral
angle and a periostracal fringe as well as the
colour and the presence or absence of spiral
striae have been used for species discrimi-
nation. Spiral striae, together with growth
lines, form a reticulate surface which is ap-
propriately termed “Gitterskulptur” in the
German literature. A keel, typical in Planorbis
carinatus Múller but also present in the ge-
nus Anisus, has rarely been developed in Gy-
raulus outside the old lakes (Lakes Prespa,
Ohrid, and Biwa).
Some species never produce an angle (ex-
ample: Gyrauius albus, G. laevis), others al-
ways have an angle (example: G. riparius) and
usually also a fringe at the same site (G. to-
kyoensis). Species are also found with shells
varying from entirely rounded to angulate at
the periphery. Thus the absence or presence
of an angle may be significant in one group
and insignificant in another. The same ap-
plies to reticulate sculpture, which is always
present e.g. in G. albus, always absent e.g.
in G. laevis, as a rule present in G. acronicus,
and rarely present in G. chinensis.
It must be stressed that embryonic shells
(=nuclear whorls of authors) have to be ex-
empted from consideration. On these shells
spiral striation occurs in all species examined
(Figs. 8 and 47b) including those that are
smooth in later stages. Moreover, this micro-
10 MEIER-BROOK
FIGS. 8-13. 8. Shell of freshly hatched individual showing spiral striae. G. spirillus, Taiwan. SEM. Scale =
0.1 mm. 9. Part of shell shown in Fig. 8. SEM. Scale = 0.01 mm. 10. Continuation of spiral striae from
embryonic to juvenile shell (arrow pointing to transition line between these) in G. albus, Zürichsee. SEM.
Scale = 0.1 mm. 11. Pigmentation of cephalopedal mass and mantle (diffuse). G. albus, Sjaelland. 12.
Mantle pigmentation with distinct pattern. G. chinensis, Hong Kong. 13. Undulate margins of kidney in G.
parvus, Speyer.
sculpture was present in all European species
examined of Anisus s. str., Planorbis s. str.,
as well as in Armiger and members of differ-
ent planorbid tribes (Hippeutis, Segmentina,
Planorbarius). The distribution of microsculp-
ture in other planorbid groups and in other
basommatophoran families has been re-
viewed by Walter (1962).
Striation on embryonic shells may consist
of spirally arranged nodules (Figs. 8, 9) visi-
ble in the periostracum, continuous periostra-
cal folds, as shown for Bulinus tropicus
GYRAULUS TAXONOMY (PLANORBIDAE) 11
(Krauss) by Hamilton-Attwell & van Eeden
(1971: fig. 5), round pits (“‘punctation”’ in oth-
er Bulinus spp., Walter, 1962: fig. 1) or reg-
ula Iy spaced transverse (=same direction as
growth lines) pits (e.g. in Biomphalaria gla-
brata (Say)).
Though | agree with Walter's (1962: 128)
statement “that punctae are lacking in most
planorbids,”’ it is likely that the various forms
of spiral striation, viz. nodules, folds, round
or transverse pits, are not so fundamentally
different that a different origin must be as-
sumed. Observations show that folds or ini-
tially continuous series of nodules gradually
become undulate, loosen to round pits and
into transverse pits, which finally become
weaker and weaker before totally disappear-
ing during juvenile growth. This can best be
observed in Hippeutis complanatus (L.)
(Meier-Brook, unpublished). Spiral striation of
embryonic shells is so universal in various
basommatophoran groups that it can be re-
garded as an ancestral structure. Conse-
quently, smooth or glossy adult shells con-
sistently present in planorbid groups may be
regarded as derived from spirally striated
ancestors. This view is supported by two
facts: (1) striation does not cease abruptly
after hatching in adult-smooth groups, but is
continued onto the first parts of the juvenile
shell; (2) in these as well as in adult-striated
groups postembryonic striae form an imme-
diate continuation of embryonic striae (e.g. in
Gyraulus albus, Fig. 10).
Gyraulus costulatus from Liberia was re-
ported as having “a total lack of punctation”
(Walter, 1962: 128). Examination of the same
species, also from Liberia, revealed that con-
tinuous striae are present on the nuclear
whorl, which later (but still before hatching)
break up into small nodules similar to those
in G. spirillus and other species.
Reticulate sculpture on the shell surface is
sometimes, though not necessarily, accom-
panied by hair-like protrusions formed by the
periostracum. They regularly occur in Gyrau-
lus albus (Fig. 36) and occasionally in G. ac-
ronicus (Fig. 62B).
Wherever peculiar periostracal structures
occur in Gyraulus, corresponding, but weak-
er, shell sculpture is found below these. A
peripheral fringe of periostracum never dec-
orates an equally rounded periphery; striated
periostracum does not cover a completely
smooth shell surface, as could be shown in
shells devoid of their periostracum. Below a
distinct fringe there is at least a distinct pe-
ripheral angle, if not even a keel (Fig. 87).
Thickness of shells was not measured. Un-
usual thickness probably plays a röle only in
ancient lakes where additional angles and
keels can be used to characterize species.
Shell color varies from light corneous to
reddish brown to light brown, but the color
range is too narrow to provide reliable char-
acters for species determination. The degree
of transparency is apparently also of equally
low value.
There were no apparent correlations be-
tween shell surface characters and environ-
mental conditions except where the reticu-
late sculpture was slightly worn in surf zones
of sandy lake shores.
Body Pigmentation
Pigmentation of the cephalopedal mass
shows slight variation, from colorless with
scattered spots (Fig. 89) to homogeneously
light grey. However, pigmentation of the
mantle roof in most cases can be categorized
(1) as diffuse and with poor contrast (exam-
ple Gyraulus albus, Fig. 11) or (2) with dis-
tinct patterns, and thus with rich contrast
(example G. chinensis, Fig. 12). These pat-
terns are most distinct on the right side be-
tween the kidney and the darker region where
the prostate gland, uterus and oesophagus
run along the inner side of the whorl. Al-
though pigment patterns are highly variable,
there were usually no doubts about which
category was involved. In G. acronicus the
pigment pattern is not as rich in contrast as
in G. chinensis; in some populations of G.
acronicus a clear decision was impossible and
identification had to be based on other char-
acters (shell, male copulatory organ). But as
a rule the nature of mantle pigmentation can
readily be categorized and serves as a help-
ful character.
Pallial Organs
The only pallial organ that proved to have
a shape characteristic for certain taxa is the
kidney. It has conspicuously ““undulate” mar-
gins (Fig. 13) in some species, whereas in
most species the margins are straight. In un-
relaxed individuals the margins of the shrunk-
en kidney seemed to be more or less undu-
late in all species (perhaps the reason that
the significance of this peculiarity has gone
unreported, e.g. Hubendick (1955: 456)
12 MEIER-BROOK
FIGS. 14-15. 14. Septate kidney in tangential section. G. laevis, Kuehren. Hematoxylin and eosin. (M
123). 15. Non-septate kidney in tangential section. G. euphraticus, S Iran. Hematoxylin and eosin. (M 140).
stressed ‘‘that details of the form and size of
the kidney are ... not reliable as taxonomic
characters”). In individuals fixed after relax-
ation as well as in living ones, however, the
difference is obvious. Histological investiga-
tion revealed that in Gyraulus laevis and G.
parvus (Say) real septa protrude into the ren-
al lumen alternating from both sides. Proba-
bly these septa have developed from folds of
the renal walls, but fusion (Fig. 14) has taken
place that is evidently irreversible. No such
septa can be detected in most other species
(Fig. 15), even when contraction yields a form
like “‘undulate’’ margins.
Alimentary Tract
The jaw and radula have long been the
main non-conchological characters used in
gastropod taxonomy. In rare cases, how-
ever, these parts have contributed to differ-
ential diagnosis at the infrageneric level. This
is the situation in Gyraulus. Except for a small
group of species endemic to ancient lakes in
Macedonia, radular teeth are uniform
throughout the genus. Should species differ-
ences really occur these must be so insignif-
icant that they are masked by individual vari-
ation. Although great hopes used to be placed
on the radula for species discrimination, many
authors (e.g. Annandale & Rao, 1925; Rosz-
kowski, 1929; Wagner, 1931) came to the
same conclusion: the value of the radula for
species discrimination has been exaggerat-
ed. Most regrettably variation of radular
characters is not quantifiable in our genus,
so that the above statements could only be
supported by a great number of additional
figures. SEM micrographs of radulae docu-
menting its low value have been deposited at
the Senckenberg-Museum, Frankfurt.
On this occasion it is necessary to caution
against misinterpretations of scanning elec-
tron micrographs. Owing to the great depth
of focus it is very difficult to tell the degree of
inclination of part of a radula. To demon-
strate the effect of varying directions of ob-
servation two micrographs of the same re-
gion of one individual radula is shown (Fig.
16). Hence, statements about “long” and
“short” cusps on radulae, judged from SEM
photos, should be regarded with caution.
Furthermore differences occur between
younger and older portions of a radula where
teeth are worn and, thus, may lack the sharp
edges of their cusps (Fig. 17). To save space
| refrain from giving descriptions of radular
teeth for all species except for the type-
species of the genus and the aberrant species
from ancient lakes. The number of teeth per
GYRAULUS TAXONOMY (PLANORBIDAE) 13
17 А ES р = 178
FIGS. 16-17. 16. Identical part of radula in perpendicular view (A) and with 37° inclination (B). G. parvus,
W Iceland. SEM. Scale = 0.01 mm. 17. Unworn (A) and worn (B) central and lateral teeth in different
regions of the same radula. G. a/bus, Sjaelland. SEM. Scale = 0.01 mm.
transverse row appears to be highly size-de- The oesophagus, crop, stomach, caecum,
pendent and is almost worthless for taxo- and digestive gland are devoid of taxonomi-
nomic purposes (Bertram & Meier-Brook, in cally significant characters within our genus.
preparation). Striking variation occurs in the size and form
14 MEIER-BROOK
FIG. 18.
Intestinal loop, viewed from the left (A),
ventral view (B) and viewed from the right (C). G.
albus, Sjaelland (M 286).
of the intestinal loop. Normally (Fig. 18) the
intestine, after leaving the pyloric region of
the stomach, runs anteriad on the right side
of the animal, bends to the left, passes over
the oesophagus, bends once more, runs
backward on the left side, forms an elongate
loop embedded in the large digestive gland,
and finally runs anteriorly after passing the
stomach on its left side again, to terminate
as the rectum. In portions of the lots exam-
ined of Gyraulus laevis and in all specimens
dissected of G. parvus the loop was absent,
and the intestine showed only a slight indi-
cation of flexure where the loop is usually in-
tercalated (Fig. 19A, arrow). Transitional
stages occur. Presence or absence of a loop
is not strongly correlated with more invaria-
ble species characters and, thus, is not con-
sidered to be a reliable character. In most
species presence and absence seems to be
randomly distributed, even within the same
population. In a few species percent omis-
sion is increased (e.g. G. parvus); in others
the loop is well developed in all individuals
examined (e.g. G. acronicus, G. rossmaess-
leri). Therefore, in certain cases presence or
absence of an intestinal loop may serve as a
supporting character, and in future studies
further attention should be devoted to it.
Richards (1973a), studying Biomphalaria gla-
brata with both absence and presence of a
ces crop
2 [2
FIG. 19. (A) Omission of intestinal loop, G. laevis,
Kuehren. The arrow indicates the site at which a
loop usually is intercalated. (B) Same; stomach re-
gion in ventral view. Key to lettering p. 27.
loop in the same stock, found that omission
is inherited. This may be the case in Gyraulus
also, but inheritance, of course, does not
necessarily mean taxonomic significance.
Reproductive Organs
The number of lobes in the ovotestis (her-
maphroditic gland), a character used in the
taxonomy of at least one group of planorbid
snails, viz. Biomphalaria (e.g. by Paraense,
1966; Pan American Health Organization,
1968), is difficult to determine, because the
lobes are insufficiently differentiated in the
most apical portion of the ovotestis. The val-
ue of lobe numbers is especially low if varia-
tion between species is low and there is
overlap; this is the case in Gyraulus (mostly
between 28 and 32).
GYRAULUS TAXONOMY (PLANORBIDAE) 15
The length of the spermoviduct (hermaph-
roditic duct) (Sod in Fig. 43) and of its parts
has not proven to be of value as a character,
particularly since the seminal vesicle can be
lengthened by stretching to more than twice
its dimension in the resting state. The shape
of the seminal vesicle coils may be charac-
teristic, being either rounded and bulbous or
more delicate and studded with spiny humps
(Fig. 20). Though in many cases a decision is
difficult, extreme formations can be found to
be constant.
The seminal vesicle (vs in Figs. 43, 57) can
be exceptionally voluminous and appear larg-
er than the prostate gland and uterus togeth-
er (e.g. Gyraulus riparius). The portion of the
spermoviduct between the seminal vesicle
and the carrefour (=distal spermoviduct),
which is usually very thin, in certain cases
can be wider than the vas deferens and may
then have diagnostic value (e.g. G. /aevis).
In the carrefour (car in Fig. 43) and the parts
of the female tract down to the vagina (vag
in Fig. 43), no characters useful for taxonom-
ic purposes have been found. The vagina in
many cases is much inflated, partly below the
zone of fusion with the duct of the bursa cop-
ulatrix. This inflation may be absent, but it
has, as yet, been impossible to recognize any
regularity. Intra- and interpopulational varia-
tion of the bursa copulatrix and its duct are
usually greater than interspecific variation.
Therefore it has lower value than in other
planorbid genera (e.g. Planorbis, Meier-Brook,
1976c), but an extremely slender bursa (e.g.
Gyraulus laevis, G. parvus) or an unusually
inflated one (e.g. G. eugyne n. sp.) is a good
species character.
In the male reproductive tract the length of
the free sperm duct (free spd in Fig. 48), rel-
ative to other parameters, is of restricted val-
ue, but the prostate gland has been judged
as bearing one of the most reliable charac-
ters for species discrimination in the genus
since counting the numbers of diverticula was
introduced into taxonomic work (Baker, 1945;
Meier-Brook, 1964; Brown & van Eeden,
1969; Pace, 1973). Numbers of diverticula
extend over a wide range in the genus, from
<10 to >30, and they can be determined
without trouble (Tables 1 and 2). They are
rendered invaluable, because they appear to
be age—or size—independent (Fig. 21). In
immature individuals diverticula may be diffi-
cult to recognize, but they are formed in de-
finitive number early in ontogeny. Variation
generally is low enough to permit the indica-
FIG. 20. Coiling of seminal vesicle: bulbous (A; G.
parvus, Speyer) (M 275); spinous (B; G. hieman-
tium, Hanechi-son) (M 260).
tion of mean diverticula numbers even in
smaller samples (Fig. 22). Diverticula are usu-
ally unbranched, except the most distal one,
two or three, which may incidentally be bi-
fid. The remaining diverticula were almost ex-
clusively found unbranched in most species
(e.g. Gyraulus acronicus, G. euphraticus
(Mousson)). Here and there, along the whole
prostate, bifid diverticula were regularly no-
ticed in G. parvus (Fig. 50). Another constant
feature in the prostate gland is the mode of
grouping. Diverticula are usually spaced
closely and regularly (Fig. 23), whereas in
certain groups they emerge from the pros-
tate duct with gaps and in changing direc-
tions (e.g. G. laevis, G. parvus, Figs. 48, 50).
The length of the vas deferens is occasion-
ally useful as a standard parameter for com-
parison with the proper male copulatory or-
MEIER-BROOK
16
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GYRAULUS TAXONOMY (PLANORBIDAE) 17
22
20
= 18 o o oo о oo
® о o о
2 1l6to о хо © о o
yo] o o oo o o о
=> 14 © 0 oo o o
a о © оо © o
= 1
Ze
9 1 40 50 60 70 mm
max. shell diam.
18
E s.d.
2 16
5 =
y
E
= 14
12
A em. ist
10 20 30 40 50
22 sample size
FIGS. 21-22. 21. Numbers of prostate diverticula in snails of various sizes. G. spirillus, Taiwan. y = 14.1
+ 0.33x; n= 51; r = 0.1245 (significance limit for р = 0.1 is 0.2329). 22. Progress of mean numbers of
prostate diverticula and standard deviations with increasing sample size. G. spirillus, Taiwan.
gan because it is a nonmuscular organ and,
thus, not subject to contraction during fixa-
tion (Meier-Brook, 1976b). In Gyraulus the
length could not be determined exactly in the
majority of cases. Delamination is easy in
Planorbis (Meier-Brook, 1976c; fig. 1), name-
ly between the upper end of the penis sheath
and the point of fusion of the sperm duct and
the prostate duct. In Gyraulus this point is
much less discernible.
The width of the vas deferens is uniformly
low through the vast majority of Gyraulus
species, but in some species from Lake Ohrid
and the Malayan Archipelago it is constantly
greater, and in one species (G. parvus) in the
proximal half it is Уз to V2 of that in the distal
half (both measured in the middle, Fig. 51). In
these few cases characters of the vas def-
erens can serve for the discrimination of taxa.
The form and total length of the male cop-
ulatory organ as well as the ratio of length of
penis sheath : length of preputium are of great
taxonomic value and have been highly valued
in planorbid taxonomy since they were used
for discriminating among G. acronicus, G.
rossmaessleri and G. albus (Meier-Brook,
1964; Hudec, 1967; Brown & van Eeden,
1969). This is in contradiction to Hubendick's
(1955: 457 et seq.) opinion that it is ‘‘worth-
less to make measurements of its (male cop-
ulatory organ) different parts and then to base
comparisons on details of proportion.” The
total length can be related to other dimen-
sions and expressed by indicating the posi-
tion of its upper end in relaxed individuals in
situ (Fig. 44; Meier-Brook, 1964: figs. 21, 22).
The ratio total length of the 4 copulatory
organ: greatest width of penis sheath, as
18
MEIER-BROOK
GYRAULUS TAXONOMY (PLANORBIDAE) 19
tentatively used in a previous paper (Meier-
Brook, 1964), is considered to be unneces-
sary, since drawings demonstrate differences
better than numerical data in this case.
The penis provides two significant charac-
ters: the position of the penis pore, i.e. the
aperture of the sperm canal and the shape
of the penis tip. The penis tip is usually
equipped with a bulbous thickening, resem-
bling the mammalian glans penis (Fig. 45,
etc.); in a few groups the thickening may be
less distinct, in others there is a conical nar-
rowing instead of a thickening (e.g. in Gyrau-
lus rossmaessleri, Fig. 55; Meier-Brook,
1964: figs. 24, 27). The penis pore almost
always is situated near or in the bulbous
thickening and generally has a well defined
site: (1) subterminal or in the distal half of the
thickening (G. albus), (2) near the proximal
end of the thickening or in its upper half (G.
acronicus, almost all groups from S and SE
Asia). Rarely it is situated above the middle
of the whole penis, as described for G. Iych-
nidicus by Hubendick & Radoman (1959: 236)
and observed in the present study only in G.
malayensis n. sp. from Bali, Java and Malay-
sia. In every case there is a more or less wide
open groove running down from the penis
pore to the proximal opening of the stylet (Fig.
24). During copulation the sperm traverses
the groove to the stylet. The stylet leads it
out of the everted copulatory organ through
the hole in the papilla (Fig. 29). The stylet is
so uniform in the whole genus as well as in
closely related genera that it is not attributed
interspecific significance, except in a few
cases where it is reduced in size and form:
the “hob-nail stylet” of G. rossmaessleri, G.
riparius, and G. crista (L.).
Hubendick & Radoman (1959: 231-234)
report on differences between two Species,
G. crenophilus Hubendick & Radoman and
G. fontinalis Hubendick & Radoman, in the
proximal end of the stylet. The basal part is
said to have chitinized internal bars in the one
and no such bars in the other species. When
these species are reexamined, attention
should be directed towards these alleged dif-
ferences. According to my experiences such
differences could occur depending on where
one happens to section. In the middle of the
wider basis (Fig. 25) a cross section may look
like a horseshoe, as stated for G. fontinalis
(Hubendick & Radoman, 1959: 234); a few
um towards the penis tissue the picture
should be quite different. A continuous series
of sections might have been impossible at that
time because of the resistant stylet material.
The basal opening may vary a little from
comparatively wide to narrower (Fig. 26). This
figure, by the way, clearly demonstrates that
the stylet is a hollow tube formed out of a
thin blade by rolling up in a clockwise sense.
The sperm leaves the stylet through a slit left
open near the stylet tip (Fig. 27), which in this
view appears extremely acute although it is
of lancet-shape when viewed after being
turned 90° (Fig. 28). In everted organs of in-
dividuals of various species only this tiny sty-
let tip penetrated the hole in the papilla
(Brown & van Eeden, 1969, fig. 6). This hole
(Fig. 29) is so small that most of the stylet
(including its base) probably cannot pass
through it, not to mention the penis tip itself.
However, a penis extending into the prepu-
tium has been reported by Wright (1963: 463)
as a characteristic of Gyraulus costulatus
from Angola. Unfortunately he did not men-
tion how frequently this condition was seen.
| have found no more than one such case,
which | could not explain, in adult Gyraulus.
In immature animals the stylet is posi-
tioned in the distal portion of the 3 copulatory
organ; this is the rule. In early stages of de-
velopment there is no clear distinction be-
tween the future penis sheath and the pre-
putium. The developing penis with the stylet,
which is formed in its definitive size (Huben-
dick, 1958: 429; Meier-Brook, 1964: 238),
extends distally to near the male genital pore.
E
FIGS. 23-28. 23. Prostate gland with closely spaced diverticula. G. spirillus, Taiwan. Most distal diver-
ticulum two-branched, others unbranched. CPD; SEM. Scale = 0.1 mm. 24. Penis pore (arrow) and groove
leading to stylet aperture. G. euphraticus, S Iran. CPD; SEM. Scale = 0.01 mm. 25. Stylet base is ‘“‘horse-
shoe-shaped” in cross section. G. chinensis, Chongpyong. SEM. Scale = 0.01 mm. 26. Stylet showing
apertures and edge of rolled up blade. G. euphraticus, S Iran. CPD; SEM. Scale = 0.1 mm. 27. Distal
aperture of stylet. G. euphraticus, S Iran, different individual. CPD; SEM. Scale = 0.01 mm. 28. Penis tip
showing penis pore (arrow) and stylet with basal aperture and lancet tip. G. euphraticus, S Iran, same
spec. as in Fig. 26. CPD; SEM. Scale = 0.1 mm.
20 MEIER-BROOK
FIG. 29. Papilla showing hole through which the
stylet tip is pushed for copulation. Copulatory or-
gan broken between papilla and diaphragm. G.
spirillus, Taiwan. CPD; SEM. Scale = 0.01 mm.
Later, when the copulatory organ begins to
differentiate the stylet is still seen in the pre-
putium surrounded by the tissue producing it
(Meier-Brook, 1964, fig. 35).
During longitudinal growth of the whole
organ the penis tip and the stylet then grad-
ually withdraw, before differentiation of pa-
pilla and diaphragm takes place. In trema-
tode-infected individuals, which were
externally adult, but occasionally retarded in
sexual development (microphally), | have ob-
served portions of the penis extended into
the preputium. Possibly an event like para-
site-induced retardation would also explain
Wright's (1963) statement. This is regarded
the more probable since Brown 4 van Eeden
(1969), investigating a large series of speci-
mens of G. costulatus (Krauss) from South
Africa, did not find the stylet position noticed
by Wright.
In immature snails the stylet may be unrec-
ognizable until a rather late developmental
stage. This was first observed in G. parvus
from Speyer (Germany). The first specimens
received appeared to be fully grown. Not-
withstanding, a stylet could not be detected,
and the snails were tentatively placed in the
genus Promenetus. In a few specimens the
distal area of the penis sheath was found chi-
tinized internally, a condition never reported
in the literature. After keeping the remaining
snails in aquaria for some weeks, all devel-
oped a normal stylet, while the chitinized re-
gion of the penis sheath was no longer ob-
served. The shells had not noticeably grown
in the meantime but body pigmentation had
turned dark. The phenomenon, subsequently
also found in G. parvus from Iceland and G.
laevis from Germany, has not been investi-
gated further. Observations suggest that the
development of the stylet in these two species
does not follow the pattern described for oth-
er species by Hubendick (1958) and con-
firmed in the majority of Gyraulus species in
the present and a previous study (Meier-
Brook, 1964). Possible differences in the site
and mode of stylet formation that should be
studied in detail may prove to support other
differences on the subgeneric level. They are
not so different that stylets in the differing
groups could be regarded to be non-homol-
ogous. Otherwise, G. parvus and G. laevis
could not be placed in the genus Gyraulus.
Any consistent lack of a distinct stylet in a
species would indicate that it does not be-
long in the genus Gyraulus. Several African
planorbid species were previously placed in
Gyraulus and its close relative, Anisus, on
conchological grounds, but in spite of the lack
of a stylet. They have later turned out to be
members of two separate genera, described
as new by Brown 8 Mandahl-Barth (1973).
In two of three new Gyraulus species de-
scribed by Brandt (1974: 241-243) the penis
is said to lack a stylet. Anatomical features
of these species are not figured; therefore it
is impossible to determine their generic po-
sition.
Other species previously regarded as Gy-
raulus, but devoid of a penial stylet, could be
placed in Planorbis, e.g. Gyraulus macedon-
icus Sturany (Hubendick 8 Radoman, 1959:
243), G. presbensis Sturany and G. intermix-
tus Mousson (Meier-Brook, 1976c).
The preputium usually contains two mus-
cular pilasters, formed as an invagination of
the preputial inner epithelium (cf. Meier-Brook,
1964: figs. 28-31). The interspace is filled
with parenchyma, sufficiently extensible to
permit eversion of the preputium for copula-
tion. Presence and site of pilasters are nor-
mally indicated in external view by a particu-
larly dense longitudinal pigmentation of the
preputium. This pigmentation is evidently a
result of folding of an outer epithelium which
is rich in pigment anyway. In species with
small ¿ copulatory organs pilasters may be
absent. Doubtful cases occur and are exclud-
GYRAULUS TAXONOMY (PLANORBIDAE) 21
G т
Е Е
FIG. 30. Shapes of уадта and bursa copulatrix in contracted (A-D) and extended (E-H) individuals of
G. spirillus, Taiwan.
ed from decisions. Clear absence may be
consistent and is then used for characteriz-
ing species (compare also definitions!).
Non-Genetic Variation in
Reproductive Organs
Effects of Fixation.—Since form and di-
mensions of soft parts underlie different
amounts of contraction depending on fixing
conditions it was necessary to evaluate the
röle of preparation methods commonly used.
Anatomical characters generally used in
taxonomical studies of planorbid snails were
compared in (1) individuals killed by immer-
sion in 70% ethanol and in (2) those killed in
warm (60-70°C) 4% formalin after relaxation
with pentobarbital (according to Meier-Brook,
1976a, 1976b) for 12 hr. Both cohorts were
selected at random from the same stock of
laboratory reared snails. Comparison was
made as to the shape of the bursa copulatrix
and vagina and to the proportions of parts of
the 3 copulatory organ.
The bursa copulatrix in the example used
(Gyraulus spirillus from Taiwan) has a more
irregular shape in contracted (unrelaxed) in-
dividuals than in extended (relaxed) ones (Fig.
30) and is smaller. These alterations are evi-
dently caused by external pressure exerted
by surrounding tissues of the shrinking ani-
mals. Active contraction is unlikely because
this organ lacks muscular tissue.
While the preputium (prp in Table 3) loses
an insignificant portion of its length, it is
mainly the penis sheath (psh in Table 3) that
is reduced in length by 25% on average. Con-
sequently the ratio psh:prp is also altered
significantly. These results differ from those
found in a Planorbis species (Meier-Brook,
1976b), where the penis sheath is smaller and
much less muscularized. Why the preputium
in Gyraulus contracted to a lesser degree than
expected after the experiences with Planor-
bis is not clear. Histologically these parts do
not differ fundamentally in the two genera.
There is good reason not to transfer exact
data of length reduction from Gyraulus spi-
rillus to other species of the genus.
Alterations not given in the table are seen
in the length and shape of the penis (Fig. 31).
The penis in most cases coils within the pe-
nis sheath, which, in its turn, shortens and
widens by contraction of the spirally ar-
ranged longitudinal musculature (Fig. 32).
Circulatory and Nervous Systems
The circulatory and nervous systems of all
groups have been examined for taxonomi-
cally valuable characters. | examined the
heart, sections of the aorta, and the central
nervous system. There were no unique char-
acter states enabling species discrimination.
Dense pigmentation of the heart is constant
in Gyraulus laevis, rendering the heart the
darkest part of the animal beside the stom-
ach and parts of the intestine. The heart has
little pigmentation or is unpigmented in all
other species studied.
Chromosomes
The karyotype has proved useful in some
planorbid groups, particularly in the problem-
22
TABLE 3. Dimensions (mm) and ratios of parts of the male copulatory organ in G. spirillus from Taiwan, fixed in relaxed (fully extended) and unrelaxed
(contracted) states.
Whorl number
Length of psh t-test Length of prp t-test Ratio psh:prp t-test
ES.Q.
Е 5.0.
Е Sd:
0.68 + 0.16
(range 0.4-1.0)
+ S.d.
1.21 + 0.20
x
(range 0.8-1.5)
4.11
Fixing state
1.85
(range 1.33-3.00)
+ 0.32
21
fully extended
<0.01
-0.05
<0.001
+ 0.36
1.42
(range 0.75-2.33)
0.66 + 0.15
(range 0.3-0.9)
+ 0.20
0.91
(range 0.6-1.2)
+ 0.28
4.11
20
contracted
MEIER-BROOK
atic taxonomy of Bulinus species, where the
usual planorbid chromosome number of 2n =
36 has undergone various multiplications
(Burch, 1964, 1967). Burch (1960) was the
first to show tetraploidy (2n = 72) па Gyrau-
lus species (G. parvus, in the original publi-
cation misidentified as G. circumstriatus
(Tryon) (Natarajan et al., 1965: 251)). In the
present study G. parvus from Mainz was
found to have п = 36 and 2n = 72. A cyto-
logical examination of G. laevis, considered
closely related to G. parvus on morphological
grounds, was not possible though most de-
sirable; animals did not survive transport to
the laboratory. G. euphraticus from southern
Iran and G. spirillus (Gould) from Taiwan had
n = 18 and 2n = 36. Burch et al. (1964: 221)
reviewed chromosome numbers of five Gy-
raulus species, stating that in G. tokyoensis
two of five specimens studied had 19 instead
of 18 bivalents in meiotic cells; one of them
even had 19 bivalents plus a univalent. Apart
from chromosome numbers, the karyotype
does not appear to differ between species
using my techniques.
Biochemical Approaches
Biochemical data are well suited to sup-
port or disprove hypotheses about relation-
ships between taxa including species (Davis
& Lindsay, 1967; Grossu, 1977). Electropho-
retic studies, for example, require living ma-
terial, because samples should, whenever
possible, be examined simultaneously. Since
living snails were available only from a re-
stricted number of species application of
chemotaxonomic methods had to be post-
poned until sufficient material is successfully
cultivated in the laboratory.
Parasitological Evidence
Parasitological data have been used to
support or weaken conclusions about rela-
tionships to an increasing extent in several
groups of animals (Fahrenholz's Rule, Eich-
ler, 1941). Echinostomatid rediae were regu-
larly observed during these studies, but a
species identification would have required
establishing life cycles in the laboratory. The
expected value of the findings, however,
would not have justified the expense.
Number of Eggs per Egg Mass
The number of eggs per egg mass was de-
termined for species reared in the laboratory.
Results so far seem to indicate that egg
GYRAULUS TAXONOMY (PLANORBIDAE) 23
vd
pen
Stylet |
FIG. 31. Penis sheath and penis in an individual
fixed without (A) and with (B) relaxation. G. spiril-
lus, Taiwan. Scale = 0.1 mm.
numbers per mass in general are correlated
with snail size and are, thus, of restricted val-
ue. Nevertheless, in certain cases, when
species of approximately equal sizes are
FIG. 32. Spirally arranged longitudinal muscles of
penis sheath in an unrelaxed, contracted individu-
al, fixed in ethanol. G. chinensis, Chongpyong.
CPD; SEM.
compared, such as Gyraulus albus, G. laevis,
and G. parvus, highly significant differences
in egg numbers per mass (Table 4) are found.
These support conclusions drawn anatomi-
cally. Aquarium conditions were kept con-
stant and as similar as possible for all species:
25°C, 12 hr/12 hr light-dark regime; 2 liters
of hard water (24° German hardness, =8.56
mval/l); fortnightly water change; twisted
fresh lettuce ad libitum daily; medium popu-
lation density (appr. 20 to 40 mature snails in
2 liters of water).
Differences reported here between G. al-
bus and G. parvus correspond well with those
TABLE 4. Numbers of eggs per egg mass in Gyraulus species.
Species, origin, counting date
albus, Zürichsee 23-IV-1974
laevis, Haarlem 21-1X-1977
parvus, Mainz 22-111-1974
dto. 17-X11-1974
dto. 30-IX-1975
costulatus, Liberia 8-1-1975
ehrenbergi, Cairo 7-X1-1974
euphraticus, S Iran 19-11-1975
chinensis, Kwekers 2-X1-1977
spirillus, Taiwan 12-\1-1976
(density: 105 ind./2 |)
dto., 7-X-1976 (7 ind./2 |)
n masses eggs/mass
counted range ER Sia!
105 1-10 piles es)
32 1-7 4.6: =='1.2
231 1-4 2.3.07
103 1-4 23 REO
140 1-5 2.4 + 0.9
253 2-6 ANO
119 2-8 5.0 1.2
167 1-11 E 2:0
152 2-12 5:8 E 1.9
139 2-14 5.8 202,2
88 3-14 TARN
t-test: albus/parvus, albus/laevis, parvus/laevis, and spirillus high/low density p < 0.001.
24 MEIER-BROOK
described in the literature. Bondesen (1950:
54) even observed up to 16 eggs per mass
in G. albus, and Krull (1931: 5), for G. parvus
from Michigan, wrote that ‘masses of two or
three are the most common but those of one
or four are also found.”
Population density greatly affects egg pro-
duction as shown by data obtained at high
and low densities of Gyraulus spirillus (Table
4). The differences between 5.8 and 7.4 are
very highly significant (p < 0.001). Mean val-
ues of 6.1, 6.4, and 6.5 were found at inter-
mediate densities.
Ecological Differences
Very little is known about ecological differ-
ences between Gyraulus species. In the case
of stenotopic snails, | attach taxonomic sig-
nificance to differences in physiological re-
quirements that now probably prevent gene
flow. Examples of such species are Gyraulus
rossmaessleri with its preponderance in
astatic water bodies, and G. /aevis which ap-
pears to be favoured by a high nitrate con-
tent and often reaches high abundance in
slightly brackish water. The importance of
such differences in chemical and physical re-
quirements as well as behavioural differences
‘in maintaining (possibly not in initiating) iso-
lation between populations” has been point-
ed out for most groups of organisms (Pros-
ser, 1974: 359). Of course, in the Planorbidae
the situation is certainly not as simple as in
Paramaecium aurelia, where Sonneborn &
Dippell (1943) distinguished no fewer than 16
physiological varieties, each of which has a
potentially common gene pool which is effec-
tively cut off from the gene pool of every oth-
er variety” and two of which are distin-
guished by the temperatures optimal for their
growth and reproduction (Sonneborn, 1974:
173).
Although we cannot yet define the specific
ecological preferences of most species, the
situation in the pair of sibling species, Gyrau-
lus parvus and G. laevis is an example of dif-
fering ecologies. My objections to the view of
conspecificity of these species as represent-
ed by Jaeckel (1962: 69) initially were based
on differences in ecological range, which is
much wider т G. parvus than т С. laevis. As
mentioned above, the latter species is both
rather stenotopic and rare, while the former
species is nearly ubiquitous (Clarke, 1973:
403). G. parvus regularly lives in pools of bo-
tanical and zoological gardens in Europe,
since 1973 constantly in a ground-water lake
near Speyer (W. Germany), and in May 1977
it was collected in a forest pool near Stutt-
gart (S. Germany) and hence outside man-
controlled water bodies in central Europe. It
readily propagates in aquaria. G. laevis, on
the other hand, is difficult to rear and is ob-
viously diminishing in its original distribution
area. These observations support the view
that environmental requirements differ be-
tween the two species.
Characterization of Anatomically
Examined Species
A detailed description of taxonomically rel-
evant organs including generic characters is
given for the type-species of the genus, Gy-
raulus albus. Descriptions of other species are
confined to characters of diagnostic impor-
tance. Quantifiable data are listed in tables
or demonstrated in graphs as much as pos-
sible.
Before conclusions are drawn as to species
delimitations, species are dealt with under
provisional names. These are mostly in con-
formance with common use in the respective
geographical areas. Type-localities are added
in parentheses.
For clarity, treatment of species is subdi-
vided according to its origin in the generally
accepted zoogeographical regions (‘‘realm’’)
(de Lattin, 1967, fig. 87): the Palaearctic, the
Oriental and, as isolated relics of Tertiary or-
igin and foci of advanced speciation, the an-
cient lakes. As the borderline between Pa-
laearctic and Orient, as found for most groups
of animals, does not fit the distribution pat-
terns in Gyraulus completely, chapters are
headed “Europe and North Asia” and “South
and East Asia.”
Synonyms, even those generally accepted
as correctly identified by authorities, are
omitted unless | could examine preserved
specimens or shells in original series, and
when their identification was beyond all doubt.
Under “material” collection localities are
written in short form. More detailed informa-
tion on the locations is listed alphabetically in
the Appendix. Spelling of names follows The
Times Atlas of the World (1967) if entered
there. Material preserved without relaxation
and hence in a contracted state is marked
“unrelaxed.'' All individuals without this note
were relaxed and fixed, as described above.
Symbols specifying material are included in
parentheses.
GYRAULUS TAXONOMY (PLANORBIDAE) 25
D
C
deflected last whorl
sph. el.
tadpole
pilaster (d>c) ©
sph. el.
club tapering
torus
FIG. 33. Definitions. A-C: outlines of shell periphery in cross section; D: deflected last whorl (example:
G. acronicus, Bodensee); E-F: preputial wall in cross section; G: shapes of bursa copulatrix (from top
left): tadpole spherical, tadpole elongate, club spherical, club elongate, tapering.
26 MEIER-BROOK
In the figures, scales measure 1 mm unless
stated otherwise. In the drawings, shaded
areas indicate sectioned tissues.
MUSEUM ABBREVIATIONS
ANSP—Academy of Natural Sciences of
Philadelphia
BMNH— British Museum (Natural History),
London
IRSNB—- Institut Royal des Sciences Natu-
relles de Belgique, Bruxelles
M or MG—no. in the author's collection of
microsc. preparations
NHMG—Naturhistoriska Museet Göteborg
NHRMS-—-Naturhistoriska Riksmuseet
Stockholm
MHNG—Musée d'Histoire Naturelle Geneve
NMC—National Museum of Canada, Ottawa
NMV—National Museum of Victoria, Mel-
bourne
RMNH—Rijksmuseum van Natuurlijke His-
torie, Leiden
SMF—Senckenberg-Museum Frankfurt/M.
ZMA—Zool. Museum Amsterdam
ZMZ—Zoologisches Museum Zurich
ZSI—Zoological Survey of India, Calcutta
DEFINITIONS AND SYNONYMOUS
TERMS
The following alphabetical list of terms used
here for morphological analysis, including
synonymous terms, is provided to enable
recognition of and the position of organs or
parts of organs, and to delimit them from
other organs connected or attached to them.
Definitions do not include functional aspects.
Terms in common and uncontroversial use in
planorbid taxonomy are exempted.
angle—any interruption of a regular rounding
(Fig. 33A).
club elongate (bursa copulatrix)—an elon-
gate bursa with gradual transition to bursa
duct, but not evenly tapering (Fig. 33G).
club spherical (bursa copulatrix)—a balloon-
like bursa with gradual transition to bursa
duct (Fig. 33G).
copulatory organ—penis + penis sheath +
preputium.
deflected whorl—a whorl leaving the plani-
spiral phase of growth toward the upper
side (=under side in the sense of many
previous authors, therefore: deflected =
bent down) (Fig. 33D).
diaphragm—the muscular ring proximally
terminating the preputium, resembling a
sphincter; =velum sensu Hubendick, 1955;
=Ringwulst sensu Buchner, 1891; =dia-
phragm or muscular ring sensu Baker,
1945.
distal—situated towards outside.
embryonic whorl—the whorl formed within the
egg shell; =nuclear whorl (Fig. 10, up to
arrow).
free sperm duct—the portion of the sperm
duct not covered by the prostate gland.
fringe (of periostracum)—a periostracal col-
lar on the periphery of a shell, attached to
an angle or keel (Fig. 33C).
height—the maximum dimension measured
perpendicular to the plane in which the
whorls grow (Fig. 1).
keel—a calcareous collar on the periphery of
a shell, resulting in a concave outline in
cross section (Fig. 33B).
maximum diameter—measured from peri-
stome (‘outer lip’’) of the aperture through
the innermost (nuclear) whorl (Fig. 1).
papilla—the conical to hemispherical tissue
distally terminating the penis sheath, car-
rying a hole which is penetrated by the sty-
let for copulation (Fig. 29); =sarcobelum
sensu Hubendick, 1955; Stilettscheide
sensu Buchner, 1891; =papilla sensu Bak-
er, 1945.
penis sheath—the proximal portion of the 4
copulatory organ harbouring the penis, de-
limited proximally by the end of the vas
deferens, distally by the widened lumen be-
tween papilla and diaphragm. This is the
hinge-point for eversion of the preputium;
=penis sac in Pan American Health Orga-
nization, 1968; =vergic sac sensu Baker,
1945; =phallotheca Il auctorum (P2 Hu-
dec, in Macha, 1963); =distale (!) Phallo-
theca sensu Hudec, 1967.
pilaster—a longitudinal ridge running along
the inner wall of the preputium and reach-
ing a width greater than at the site of con-
nection with preputial wall (Fig. 336);
=muscular pillar sensu Hubendick, 1955;
=Muskelpfeiler sensu Meier-Brook, 1964.
(The term pilaster is preferred to pillar, be-
cause there is always a connection with the
wall along its full length, whereas “pillar”
implies partial detachment from the wall.)
preputium—the distal portion of the 4 copu-
latory organ, delimited proximally by a weak
zone between papilla and diaphragm, dis-
tally by the body integument; =phallotheca
| auctorum; =proximale (!) Phallotheca or
Praeputium sensu Hudec, 1967.
GYRAULUS TAXONOMY (PLANORBIDAE) 27
proximal—situated towards the inner whorls.
tadpole elongate (bursa copulatrix)—an elon-
gate bursa with bursa duct distinctly set off
(Fig. 33G).
tadpole spherical (bursa copulatrix)—a bal-
loon-like bursa with bursa duct distinctly set
off (Fig. 33G).
tapering (towards the vagina: bursa copula-
trix)—a bursa evenly tapering, without a
clear distinction of bursa and bursa duct
(Fig. 33G).
under side—under side in a shell held with its
aperture on its left side (as in apertural
views of figures in the present paper);
=morphological under side or conchologi-
cal upper side or functional right side sen-
su Hubendick & Radoman, 1959; =right
side sensu Pan American Health Organi-
zation, 1968.
upper side—upper side in a shell held with
its aperture on its left side; =morphological
upper side or conchological under side or
functional left side sensu Hubendick & Ra-
doman, 1959; =left side sensu Pan Amer-
ican Health Organization, 1968.
KEY ТО LETTERING
ад! albumen gland
an anus
aur auricle
bc bursa copulatrix
bm buccal mass
cae caecum
car carrefour
cga cerebral ganglion
dgl digestive gland
dgld common duct of digestive gland por-
tions
dia diaphragm
free spd free sperm duct
h heart
int intestine
intl intestinal loop
ki kidney
mbo mantle border
mcol columellar muscle
mrp penis retractor muscle
ngl nidamental gland
od oviduct
oes oesophagus
ot ovotestis
pa papilla
pc pericardium
pen penis
pgm male genital pore
pnst pneumostome
pp penis pore
pre preputium
psb pseudobranch
psh penis sheath
pst prostate gland
pstd prostate duct
rec rectum
ret renal tube (tubular portion of kidney)
$9! salivary gland
ski saccular portion of kidney
sod spermoviduct
spd sperm duct
st stylet
sto stomach
uo external opening of ureter
ut uterus
vag vagina
vd vas deferens
ventr ventricle
vep pulmonary vein
ver renal vein
vs seminal vesicle
SPECIES OF EUROPE AND
NORTH ASIA
Gyraulus albus (Müller, 1774)
Planorbis albus Muller, 1774: 164, No. 350
(Fig. 34) (Fridrichsdal, Sjaelland, Denmark).
Planorbis hispidus Draparnaud, 1805.
Planorbis crosseanus Bourguignat, 1862: 42.
Gyraulus albus, Ehrmann, 1933: 170, pl. 8,
fig. 104 (but not including ‘‘G. a. acronicus
(=Planorbis deformis Hartmann)” listed
hereunder!).
Gyraulus albus, Hubendick, 1949: 43, figs.
83-85.
Gyraulus albus, Jaeckel, 1962: 68 (but not
including G. deformis (Hartm.) and G. te-
nellus (Hartmann)!).
P. hispidus Draparnaud was designated by
Dall (1870) as the type-species of Gyraulus.
Material —Jorlose, Sjaelland (Denmark), 18
specs.; Switzerland: Zürichsee, 3 specs.;
Greifensee, 3 specs.; Fischau (Austria), 4
specs. (unrelaxed); Germany: Kuehren, 2
specs.; Bodensee, 2 specs.; Dobersdorfer
See, 2 specs.; Ettenheim, 4 specs.; Wollma-
tingen, 2 specs.; Ursee, 9 specs.; Windgfäll-
weiher, 6 specs.
Shell (Fig. 35)— The shell is planispiral, of
medium size (4-7 mm maximum diameter),
relatively high (1.2-1.8 mm) and has 3/2 (at
most 4 or 4/4) whorls, which rapidly increase
28 MEIER-BROOK
350. PLANORBIS argus.
Рг.амокв!$ celta alba, utrinque umbilicata, aper-
> q > t
tura dilataca.
PLaNorsis minima duorum orbium.
вс во.
Berl, Mages. 4. В. р. 253. 1. $. £ 23.
Petiv, gazuphyl.
an. DEN HVIDE SKIVE.
diam ı - 2 lin, alt, ? lin,
id
Teffa albida, peilucida, rar > fut Виз, fupra planiufcula, fub-
tus convexa, utrinque æque umbilicata, Anfraéfns tereres , in
adulris ad fu mmum quatuor; lenticule oc ulari {triis Jongitudinali-
bus & rransverfalibus inf , he in qu busdam obfi er х o vix
detegendz, illæ faris confpicu: Anfracus extimus reliquis omni-
bus major & latior, ps ra ejus pars ultra infernam He exten-
fa eft, uti in P, Purpura. Are ertura rotundata, larga, obliqua
margin? inferiore antra Au m vicidum regi,
Limax grifeus, 13 lin. longus. Tentacula albida ejusdem
fere longitudinis Soli oculi nigri.
Vafculo vitreo fervari ER medio Julii jungebantur , fimul
agentes & parientes, Membra genitalia latore finiftro.
In plants aquatitis amnis Fridrichsdalenfis,
FIG. 34. Facsimile of O. F. Múller's description of
Planorbis albus (1774: 164).
in width. The last whorl expands towards the
aperture and ends in a broadening bend. The
whorls are a little wider than high, giving the
aperture an ovoid shape. Each whorl scarce-
ly embraces the preceding one; the whorls
are equally rounded on each side as well as
peripherally and rarely have a trace of an an-
gle. The shell is slightly concave on the under
side and deeply concave on the upper side.
The last whorl is not or very little deflected.
Growth lines are arcuate, crossed by spiral
striae, both forming a reticulate surface
sculpture (Fig. 36A, B). This sculpture is usu-
ally well developed; when weakly developed
it is visible at least on the upper side of the
shell. The surface is never glossy; the peri-
ostracum sometimes has short hair-like pro-
trusions; these are part of the periostracal
ridges formed on growth lines, the ridges
being stretched out to triangular laminae on
the nodules formed at the crossing points.
“Hairs” thus follow the spiral striae (Fig. 36C).
The color is light-corneous.
Animal (Fig. 37)— The animal is light to me-
dium grey. The pigment is distributed not quite
evenly on the cephalopedal mass that com-
prises body portions extended out of the shell
during crawling. The tentacles are lighter, ex-
cept for a central line that is darker; pigmen-
tation is cloudy on the remaining parts. The
anterior mantle roof (Figs. 11 and 37) is dif-
fusely pigmented, showing a cloudy pattern.
A
D
FIG. 35. С. albus, Sjaelland. Shell shape: A—lat-
eral view, B—upper side, C—apertural view, D—
under side (SMF 246 302).
The mantle roof is dark grey on the right side,
in a zone parallel to the columellar muscle.
Pigmentation is weaker and diffuse to almost
absent in between this densely pigmented
zone and the peripheral region. At the periph-
ery there is a narrow longitudinal unpigment-
ed stripe indicating the position of the pul-
monary vein (Fig. 37A, vep); next to it the
kidney roof is densely pigmented. The renal
vein (Fig. 37A, ver), again devoid of pigment,
delimits the kidney on its left side. The re-
maining portion of the left mantle side is
weakly pigmented, only a few scattered pig-
ment cells being present.
Pallial Organs (Fig. 38)—The pallial cavity
extends back from the thickened mantle bor-
der (mbo) to about 3 of the last whorl. The
dorsal part of the mantle roof is occupied by
the kidney. The kidney consists of the very
GYRAULUS TAXONOMY (PLANORBIDAE) 29
short saccular portion (ski) situated to the left
of the pericardium (in dorsal view) and the
long renal tube (ret). Anteriorly, the kidney ta-
pers off and ends in a reflected ureter, dis-
charging (uo) behind the pneumostome (pnst).
In extended individuals the kidney has straight
margins; there are no septa in the lumen of
the renal tube. None of the three pallial ridges
regularly found in other planorbid groups is
developed.
The mantle border is partially connected
with the roof of the cephalopedal mass by
connective tissue (Fig. 39). The pallial cavity
opens on the left side through the pneumo-
stome (pnst) formed by the mantle lobe, which
in the living snail is held against the mantle
border (mbo), together with the pseudo-
branch (psb). The pseudobranch is triangular
to rectangular and carries a longitudinal fold.
The rectum ends (an) near the base of the
pseudobranch on the right of this fold.
Alimentary Tract—The jaw is composed of
numerous chitinized bars, approximately 12
in the median part, and 15 to 20 in each of
the lateral portions (Fig. 40).
The radula of a sexually mature animal
consists of approximately 150 transverse
rows of teeth. The almost straight rows are
formed by 27 to 35 single teeth (Fig. 41); the
central tooth is bicuspid (Fig. 17). A small ad-
ditional denticle regularly occurs between the
cusps; similar denticles are found on both
sides of the tooth, but nearer to the base.
The base broadens posteriorly. The lateral
teeth are tricuspid. The mesocone is the
longest one; the endocone is a little shorter
and the ectocone the shortest. Again there
are additional denticles between each cusp
and on both sides of a tooth. The base of the
lateral teeth has parallel edges directed
obliquely towards the lateral posterior region
of the radula. About the 10" or 11" lateral
tooth shows the first signs of a conversion
to marginal teeth. These are pluricuspid, but
the three cusps of the lateral teeth, particu-
larly the mesocone, are still pronounced,
though narrower. The denticles increase in
number and size (Fig. 42). Cusps and denti-
cles together total a dozen or even more.
Variation of radula characters is slight.
The oesophagus emerges from the dorsal
part of the buccal mass and penetrates the
ring of the central nervous system as a nar-
row tube. A short distance behind the central
nervous system it widens abruptly and runs
backwards along the right side of the body,
FIG. 36. Shell surface of G. albus, Sjaelland, with
characteristic reticulate sculpture (А, В) or ‘‘hairs”’
(C). Scales = 0.1 mm.
30 MEIER-BROOK
aur EIN.) à
38 ре ER
ventr =“.
FIGS. 37-40. G. albus, Sjaelland. 37. Animal with shell removed, viewed from the left and right, to show
pigmentation and position of organs visible externally. 38. Ceiling of mantle cavity showing kidney region.
Ventral view (SMF 246 301). 39. Mantle border in frontal view. Cephalopedal mass and portion behind
mantle border cut off. Relaxed individual with widely dilated pallial opening. 40. Jaw (M 289).
usually attached to the exterior side of the cular part of the stomach (the gizzard) that is
prostate gland. Behind the end of the pallial always filled with sand grains.
cavity it turns to the middle and widens, The intestine leaves the stomach on its right
forming a crop. This leads to the strong mus- posterior side and bends anteriorly (Fig. 18).
280 -
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REF eo à
GYRAULUS TAXONOMY (PLANORBIDAE) 31
FIGS. 41-42. G. albus, Sjaelland. 41. Transverse rows of radular teeth. SEM. Scale = 0.01 mm. 42.
Marginal teeth nos. 13-15. SEM. Scale = 0.01 mm.
At the bending point the short duct of the
digestive gland merges into the intestine. A
caecum, in the anteriad direction, follows im-
mediately. The intestine then follows the
course described above as an intestinal loop
(Fig. 18). The rectum makes its way to the
pseudobranch on the extreme left side of the
floor of the pallial cavity.
The salivary glands insert on the dorsal part
of the buccal mass, close to the oesophagus.
They penetrate the nerve ring together with
the oesophagus, buccal retractor muscles and
cephalic arteries. The two unequal parts of
the digestive gland (Fig. 37B, dgl) almost fill
the whole space from the central part of the
stomach to the middle of the ovotestis. Only
the intestinal loop and the spermoviduct with
the seminal vesicle are embedded in it.
The intestinal loop may be more or less
well developed, i.e. longer or shorter. Com-
plete lack of the loop has not been observed
in Gyraulus albus.
Reproductive System (Fig. 43)—The ovo-
testis (ot) consists of a double row of 20 to
30 lobes. The anterior part of the ovotestis is
situated on the left side of the posterior end
of the digestive gland. The spermoviduct (sod)
is embedded ventrally in the middle of the lat-
ter. It leaves the ovotestis as an initially wide
and delicate tube, then narrowing and, after
a short distance, coiling up to form the sem-
inal vesicle (vs), where sperm is stored after
production. A narrow distal part, consider-
ably longer than the proximal one, follows and
enters the rather voluminous carrefour (car).
This is a complex hollow organ, entered also
by the tiny duct of the albumen gland (ag)).
The albumen gland is a compact organ, con-
vex dorsally, concave ventrally, composed of
numerous small acini.
Distal to the carrefour the female and the
male tracts of the reproductive system sep-
arate. The oviduct leaves the carrefour on the
left side as a wide and irregularly shaped tube.
FIG. 43. Reproductive system of G. albus, Sjael-
land. Max. shell diameter 5 mm, 4% whorls (М 286).
Key to lettering p. 27.
32 MEIER-BROOK
FIG. 44. Position of distal genitalia in a fully ex-
tended G. albus, Sjaelland. Roof of cephalopedal
mass and posterior portion of oesophagus re-
moved (SMF 246 301). Key to lettering p. 27.
Its transition to the part joined by the nida-
mental gland is mostly invisible from the out-
side. Under favourable circumstances the ni-
damental gland can be distinguished from the
oviduct by its opaque yellowish appearance.
The uterus can be told from the nidamental
gland by a more transparent and colorless
appearance. The uterus finally narrows
abruptly towards the vagina, which is unin-
flated or, at most, weakly inflated near the 2
genital pore. Here the bursa copulatrix
branches off with a duct of moderate width.
The bursa is club-shaped in Gyraulus albus,
usually elongate, rarely more spherical. It is
always narrower than the uterus.
The male tract emerges from the carrefour
on the right ventral side. The sperm duct is
always considerably narrower than any part
of the female tract. It lies close to the oviduct
and is joined by the duct of the prostrate
gland. Their fusion takes place gradually so
that it cannot be clearly located in many
cases. The prostate duct is longer than half
the sperm duct. A number of diverticula
merges into the duct. These are densely ar-
ranged in a single row and are not separable
histologically from the prostate duct. Diver-
ticula are unbranched in the vast majority of
examined individuals; the bifid condition rare-
ly occurs. A more scattered, irregular and less
dense arrangement of diverticula was ob-
served in several individuals of one popula-
tion (Sjaelland, Denmark). Numbers of pros-
tate diverticula usually vary from 13 to 20. In
one population, numbers as low as 7 were
counted (Table 1).
The vas deferens, extending between the
point of fusion of the prostate duct with the
sperm duct and the 3 copulatory organ, is
relatively wide.
The 3 copulatory organ is comparatively
short. In relaxed and well extended animals
of Gyraulus albus its proximal end at most
reaches up to the bursa copulatrix, but often
not even to the vagina (Fig. 44). The penis
sheath (psh) is 1.5 to 2.0 times the length of
the preputium (Table 2). It is club-shaped in
outline, being narrowest near its middle. A
knob-like thickening of the copulatory organ
in the region of the papilla and diaphragm in-
dicates the border between the two portions
in external view. A single retractor muscle in-
serts at the penis sheath near its distal end.
The preputium has relatively thin walls that
are folded inwards, but too little to form pi-
lasters (see definitions). The preputial lumen
is oval in cross section or slightly S-shaped.
The male copulatory organ opens outwards
a short distance behind the left tentacle (Fig.
37A, pgm).
The penis is enclosed in the penis sheath.
It consists of a long tube, uniform except at
its tip. Distally the penis is thickened on one
side (see Meier-Brook, 1964, figs. 25, 28). The
orifice of the sperm canal (pp) is in the distal
half of the thickening, and is often subtermi-
nal. Variation in the form of the penis tip and
position of the penis pore, is moderate (Fig.
45). The groove between the penis pore and
the opening of the stylet is wide and shallow,
not narrow as shown for Gyraulus euphrati-
cus (Fig. 28). The stylet varies a little in size
and shape, but is generally similar to that fig-
ured from other species (Figs. 25-28).
As to the histology of the 4 copulatory or-
gan, | refer to the excellent account by Buch-
ner (1891: 78-84, pl. V: 1) for Anisus vortex.
Remarks—Gyraulus albus is so well sep-
GYRAULUS TAXONOMY (PLANORBIDAE) 33
FIG. 45. Variation in shape of penis tip and position of penis pore in G. albus. A-F: Sjaelland (M 286);
G-H: Ettenheim (М 288); I: Kuehren (М 90). Scale = 0.1 mm.
arated from any other species of the genus
that problems arise neither in delimitation nor
in recognition. Shell characters alone are suf-
ficient to distinguish this species from others.
G. albus is distributed all over Europe; it is
most frequent in temperate climates. It ex-
tends far to the east. Khazannikov (1973: fig.
1) figured 4 copulatory organs clearly proving
correct identification of this species from the
River Terek Basin (northeast of the Cauca-
sus), but so far this is the most eastern rec-
ord of anatomically-identified specimens. All
records from Central or East Asia seem to
have been published by authors who only
knew G. albus from descriptions and insuffi-
cient illustrations, or had little experience with
this species. Thus, the species named Ani-
sus (Gyraulus) albus by Mori (1938: 294, pl.
16: 13) was doubtlessiy misidentified, as
judged from the description and figures. Ku-
roda's listing of G. albus (1963: 23) and of
G. spirillus as one of its synonyms was evi-
dently based on Mori's paper. Consequently
it is highly probable that this species does
not live in Japan.
Yen (1939: 69) claimed that Gyraulus al-
bus was frequently found in N and S China.
| examined the material in the Senckenberg-
Museum, Frankfurt. The shells in the three
lots named G. albus from Sagsagyr/Ordos,
Peak/Hongkong and Pok-Fulam/Hongkong
(Yen, 1939, pl. 6, fig. 6) are devoid of any
spiral striation and cannot be distinguished
from the material anatomically examined and
identified as G. chinensis in this study.
It must be conceded, however, that at that
time—and until recently (Jaeckel, 1962: 68) —
Planorbis deformis Hartmann was consid-
ered a variety of Gyraulus albus, and that in
E Asia there are indeed snails resembling P.
deformis at first glance. P. deformis has, in
the meantime, proven to be a synonym of G.
acronicus (Meier-Brook, 1964). Whether G.
albus is conspecific with North American
forms such as G. deflectus (Say, 1824) and
G. hirsutus (Gould, 1839) cannot yet be an-
34 MEIER-BROOK
QoS
pS Sy
FIG. 46. G. laevis, Kuehren. Shell. A—upper side,
B—apertural view, C—under side, D—lateral view
(SMF 246 308).
swered satisfactorily. Species identity of these
has been claimed by Dall (1905) and Baker
(1928), but other authors have questioned this
on conchological grounds (Miller, 1966;
A
FIG. 47.
Scale = 0.1 mm.
Clarke, 1973). Judging by Baker’s (1945: pls.
15, 16) anatomical figures, the two American
species have a ¢ copulatory organ at least
twice the length of that of G. albus.
Gyraulus laevis (Alder, 1838)
Planorbis laevis Alder, 1838: 337 (Whitley
Quarries near Newcastle).
Planorbis thermalis Westerlund, 1885: 83
(Bad Villach).
Gyraulus laevis, Ehrmann, 1933: 170, pl. 8,
fig. 105.
Gyraulus laevis, Jaeckel, 1962: 69.
Material—Kuehren (Germany, 16 specs.,
including 5 trematode-infected ones with
underdeveloped reproductive organs; Haar-
lem (Netherlands), 12 specs.; Bad Villach
(Austria), 10 specs. (Gyraulus thermalis).
Shell (Fig. 46) —The shell is a little smaller
than in Gyraulus albus (4 mm maximum di-
ameter), relatively high (1.3 mm); it has 32
whorls, which increase regularly in width, the
last one not expanding towards the aperture.
The whorls are scarcely wider than high and
separated by deep sutures; the aperture is
roundish. The periphery is rounded, never
keeted or angled. The shell is deeply concave
on the upper side; on the under side, the
whorls interior to the penultimate one are also
concave; the whorls are more rounded on the
under side than on the upper side. Growth
lines are less arcuate than in G. albus and
irregular. The surface is smooth (Fig. 47) and
С. laevis, Kuehren. Shell surface, SEM. A— whole shell. B—spiral striation on embryonic whorl.
GYRAULUS TAXONOMY (PLANORBIDAE) 35
FIG. 48. С. laevis, Kuehren. Reproductive system. Maximum shell diameter: A—3.4 mm (SMF 246 309),
B—3.6 тт (М 290), C—5.4 mm (М 66). Key to lettering р. 27.
slightly glossy; spiral striae are absent ex-
cept on the nuclear whorl, as in all species.
The color is brownish.
In the population at Bad Villach named Gy-
raulus thermalis shells are smaller, 2 to 3 mm
in diameter, 0.9 to 1.0 mm in height, with 3
to 3% whorls.
Animal—The animal is moderately grey and
scarcely pigmented; the mantle roof is vir-
tually unpigmented except on the kidney and
lateral borders of the accompanying veins
where scattered pigment cells are found. The
mantle border is unpigmented. The extreme
right portion of the mantle covering the re-
productive tract is more densely pigmented.
Conspicuously dark pigmentation is present
only in the heart, stomach and intestine be-
tween stomach and flexion (Fig. 19, arrow).
There are several distinct septa in the kid-
ney giving it an undulate shape (Fig. 14). The
jaw and radula are not fundamentally differ-
ent from those described for Gyraulus albus.
An intestinal loop was lacking (Fig. 19) in ten
individuals and well developed in six from the
population at Kuehren.
In the reproductive system (Fig. 48), the
distal portion of the spermoviduct is wider
(0.030-0.033 mm) than the two halves of the
vas deferens in their respective middle parts
(proximal half ~ 0.012 mm; distal half ~
0.018 mm) and also wider than the sperm
duct. The bursa copulatrix is inconspicuous,
elongate club-shaped or tapering. The pros-
tate gland is so delicate that it is almost im-
possible to detach it from the female tract
without damage. It shows two unusual fea-
tures: the scattered arrangement of divertic-
ula and their changing shape. The number of
36 MEIER-BROOK
diverticula seems to be slightly less than in
Gyraulus albus (Table 2). The vas deferens is
similarly narrow in both halves (see above).
The 3 copulatory organ is not longer than that
in G. albus (Table 2). Microphally was ob-
served in trematode-infected individuals (х =
0.60 + 0.2 mm). The length ratio penis
sheath : preputium is also similar to that in G.
albus. The preputium cannot be distin-
guished from that of G. albus.
The penis shows considerable variation in
shape. In larger individuals (e.g. Fig. 48C) it
is nearly congruent with those figured for Gy-
raulus albus in Fig. 45E and similar forms. In
smaller individuals 3.3-4.0 mm in diameter,
the distal thickening is regularly undevel-
oped; the tip is rather conical in shape, and
the penis pore is lateral rather than subter-
minal. This shape and penis pore position is
possibly a sign of immaturity although an or-
ange color in all individuals indicates that
copulation has taken place. The question de-
serves further attention. Smaller individuals
with an incompletely differentiated stylet and
a colorless bursa copulatrix have been ex-
empted from evaluation.
Remarks—Gyraulus laevis has been de-
fined as a distinct species conchologically for
a long time. Doubts can arise, where G. lae-
vis and G. albus occur together and the latter
has weak spiral striation. However, the sep-
tate kidney and the irregular prostate diver-
ticula provide good characters for safe dis-
tinction. G. laevis is closely related, though
not identical, with the North American G. par-
vus, which is included here for comparison
and because it lives on Iceland and has been
introduced to Europe. The only anatomical
description of G. laevis thus far published is
by Soos (1935: 26, fig. 3). From his figure it
can be seen that his Hungarian material re-
sembles that presented here in the irregular
prostate diverticula, their number (11), the
short 4 copulatory organ, and the club-shaped
bursa copulatrix. Soos, however, expressed
the opinion that С. /aevis differs anatomically
from G. albus only to a negligible degree.
Gyraulus thermalis has been given the rank
of a subspecies of G. laevis or a species en-
demic to the thermal spring of Villach (Klemm,
1960: 12; Jaeckel, 1962: 69; Jaeckel, 1967:
103). However, the small size of G. thermalis
appears to be the only character by which it
constantly differs from G. laevis. In this con-
nection it may be noted that there are more
instances in the genus where thermal springs
harbor snails significantly smaller than those
that occur in cooler waters. One is G. acroni-
cus from Paratunka (Kamchatka): the snails
were collected “near hot springs’; the big-
gest of 97 individuals was 4.7 mm in diame-
ter, whereas samples from other Siberian lo-
calities contained snails exceeding 6 or 6.5
mm in diameter. The other example is G. par-
vus from a warm spring in Iceland (about
30°C, as estimated by the collector). Snails
from two shipments were transferred to
aquaria at 25°C in attempts to raise larger
individuals for anatomical studies. They all
died before growing larger than 2.5-3.0 mm
diameter. As the sample also never con-
tained empty shells larger than these, | as-
sume that they do not exceed this size in their
natural habitat. The mechanism causing
dwarfism in warm springs is not yet under-
stood. At any rate it is unlikely that a process
of speciation has been initiated here. Only in
such a case would it be justified to raise a
dwarf form, such as thermalis, to subspecies
rank. Experimental analysis should clear up
how far in phenotypical dwarfism genetically
fixed characters are involved. It seems pref-
erable to deal with thermalis as merely an
ecological form.
Snails diagnosed as Gyraulus laevis have
been recorded from Europe and Asia as far
east as Kamchatka. According to Alzona
(1971) the species is said to live all over Italy,
and other records were published on its dis-
tribution in other Mediterranean countries,
sometimes under names such as G. agraulus
and G. numidicus. In one case an anatomical
figure was published of " Gyraulus agraulus.'
This figure clearly proves that the material
does not belong in Gyraulus (Giusti, 1968).
In attempts to obtain more material of Gy-
FIGS. 49-52.
o
С. parvus. 49. Ann Arbor (U.S.A.). Shell (SMF 246 310). A—upper side; B—apertural
view; C—under side. 50. Variation in prostate glands. A, B: Ann Arbor (M 30), C: W Iceland (M 198), D:
SE Iceland (MG 208). 51. ¿ copulatory organ (Speyer) and characteristic vas deferens being wide in the
distal half (0.050 mm in its middle) and narrow in the proximal На! (0.018 mm) (M 227). Scale = 0.1 mm.
52. Variation in shape of penis tips and position of penis pore. A, B: Ann Arbor (M 81, 83); C, D: W
Iceland (M 199); E, F: SE Iceland (M 274); G, H: Speyer (M 275). Scale = 0.1 mm.
37
GYRAULUS TAXONOMY (PLANORBIDAE)
MEIER-BROOK
tap.
Bursa copul.
sph.
el.
0.6
0.3
0.4
асе
2.3
2.1
2:03
1.8
Ratio psh:prp_ u
range
11-25
1.5 + 1.7 +
1.6 + 1.0
1.9-2.3
105225
115225
1.1
14 = 02
ЕО
0810/8512
1.4+1.8
Male copulatory organ
1.4 + 0.2
Total length (mm)
range
1.1-1.6
1.0-1.3
1.2-1.6
0.8-2.0
6
322
8.0
12.8
12027,
7-17
No. prost. divert.
7-10
range
12-14
n
Reproductive organs of Gyraulus parvus.
31
Locality
Ann Arbor
Abitibi-R'
? Excl. unrelaxed individuals from Canada.
Lesser Slave L.'
TABLE 5.
W Iceland
SE Iceland
Mainz
Speyer
Total
' Unrelaxed
raulus laevis, | have dissected snails identi-
fied as G. laevis. Moreover, | checked draw-
ings of material studied and kindly provided
by Dr. Giusti from various Mediterranean is-
lands. Specimens from Crete, Corfu, Rho-
dos, Elba, Montecristo, Corsica, Sardinia,
Mallorca and, moreover, from Madeira had
anatomical features of the genus Planorbis;
it is therefore doubted that G. /aevis lives in
the Mediterranean region. The occurrence of
this species in Asia must also be doubted un-
less anatomical evidence proves the con-
trary.
Gyraulus parvus (Say, 1817)
Planorbis parvus Say, 1817: pl. 1, fig. 5 (Del-
aware River, near Philadelphia).
Material—Ann Arbor (U.S.A.), 13 specs.;
Abitibi River (Canada), 3 specs. (unrelaxed);
Lesser Slave Lake (Canada), 2 specs. (unre-
laxed); W Iceland, 5 specs.; SE Iceland, 3
specs.; Germany (introduced): Speyer, 11
specs.; Mainz, 11 specs.
Shell—The shell is small, deeply umbilicate
on the upper side; on the under side the pen-
ultimate whorl is strikingly elevated (Fig. 49).
Both the inner whorls and the last whorl are
sunken or deflected respectively. This char-
acter state is more strongly developed in Gy-
raulus parvus than in G. laevis. The whorls
are rounded or nearly angled on the under
side; the sutures are deep. The surface is
smooth and glossy; no spiral striation is
present. The color is light-corneous to
brownish.
Animal—The animal is very weakly pig-
mented, like Gyraulus laevis. The kidney has
distinctly undulate margins. The intestinal loop
was lacking in all individuals seen. The sem-
inal vesicle is inconspicuous to slightly bul-
bous; the distal portion of the spermoviduct
is aS narrow as in G. albus. The vagina is not
or scarcely inflated. The bursa copulatrix is
as narrow (and relatively long) as in G. laevis,
mostly tapering or elongate club-shaped,
rarely spherical club-shaped. The number of
prostate diverticula is insignificantly less than
in G. laevis (Table 5). The shape of the diver-
ticula is still more irregular than in G. laevis
(Fig. 50A-C), often branched, but in two pop-
ulations (SE Iceland, Speyer), regularly and
closely spaced diverticula occasionally oc-
curred (Fig. 50D). The most consistent char-
acter state in G. parvus is the strikingly
changing width of the vas deferens (Fig. 51).
In its course it narrows in between as usual
GYRAULUS TAXONOMY (PLANORBIDAE) 39
in most species, but in its distal half it widens
so much that its diameter becomes twice that
of the proximal half (width ratio distal : prox-
imal half: x = 2.1 + 0.5, range 1.3-3; п =
22).
The 3 copulatory organ is similar to that of
Gyraulus laevis; its total length as well as
proportions are of the same order of magni-
tude. The preputium does not have pilasters,
but its lumen is S-shaped. The penis, in most
cases, has a slight terminal thickening (Fig.
52A-E), less distinct than that shown for G.
albus (Fig. 45), but incidentally such a thick-
ening may be indiscernible (Fig. 52F, H). The
penis pore lies within the terminal thickening
or corresponding region, but its position var-
ies to some degree.
Remarks— The question of species iden-
tity of Gyraulus laevis and G. parvus has
repeatedly been raised because of their con-
chological similarities. Anatomical investiga-
tions show that common shell characters are
evidently not a result of convergent evolu-
tion, as they share a number of derived char-
acters to the extent that they can be consid-
ered sibling species, one living in Europe, the
other in North America. The common char-
acters, most of which I regard as derived, are:
(1) whorls almost circular in cross section;
(2) lack of spiral striation; (3) deeply umbili-
cated upper side; (4) lack of mantle pigmen-
tation left and right of kidney; (5) black pig-
mentation of heart, stomach and intestine; (6)
lack of marked thickening of penis tip; (7) del-
icate, irregularly shaped and loosely ar-
ranged prostate diverticula; (8) narrow and
long bursa copulatrix, and (9) undulate kid-
ney margins.
Of these, characters 7 and 9 are unique in
the Planorbis-tribe; they have not been ob-
served in related genera, Anisus, Bathyom-
phalus, and Planorbis. Placement in their own
subgenus is justified. A subgenus name cre-
ated by Dall (1905: 86) with Planorbis parvus
Say as the type-species is available: Torquis.
A diagnosis will be given in a later chapter.
According to Dall there are some more
species in North America to be placed here.
| have not examined other North American
species. In Europe, Gyraulus laevis is the only
representative of Torquis. G. parvus is dis-
tributed all over North America; “its northern
limit corresponds fairly well with the tree line”
(Clarke, 1973: 403). As shown here, its range
includes Iceland as well. The occurrence in
Europe is without doubt due to a quite recent
introduction, probably together with aquari-
um plants. Characters separating G. parvus
from G. laevis are: (1) the elevated penulti-
mate whorl, and (2) the marked widening of
the distal half of the vas deferens as com-
pared with that of the proximal half. These
two traits of G. parvus are unique in the tri-
bus and thus they must be regarded as de-
rived. In addition, (3) the distal portion of the
spermoviduct is not wider than the widest
portion of the sperm duct, and (4) the euryoe-
kous character of G. parvus clearly distin-
guishes it from G. laevis.
My observation that all individuals exam-
ined lack an intestinal loop agrees with Bak-
er's (1945: 74) statement “no intestinal loop
in Torquis.” This may hold true for American
species of the subgenus, but not for Gyrau-
lus laevis.
Gyraulus rossmaessleri (Auerswald, 1851)
Planorbis rossmaessleri Auerswald, 1851:
179 (Leipzig).
Material—Beside the material dealt with
previously (Meier-Brook, 1964), further spec-
imens were studied from Drusenheim
(France), 3 specs. (unrelaxed); Sessenheim/
Soufflenheim (France), 1 spec.; and Pleistin-
ger Wörth (Germany), 7 specs. (unrelaxed).
Shell—The shell (Fig. 53) is small- to me-
dium-sized, usually not exceeding 4 mm in
diameter and 1.3 mm in height. It is similar to
Gyraulus laevis in shape, but less deeply um-
bilicate; the sutures are less deep, the whorls
are rounded, never angled or keeled. The ap-
erture has a characteristic thickened lip (that
does not necessarily terminate growth!).
Growth marks indicating growth interrup-
tions are regularly present (desiccation of
habitat!). The surface is not smooth, but
rather dull, with a very fine reticulate sculp-
ture (spiral striae very close to each other).
The color is red-brown.
Animal—The animal is similar to Gyraulus
albus insofar as diffuse pigmentation is con-
cerned. The heart is not dark, but so sparsely
pigmented that it contrasts with its grey sur-
roundings by its light appearance. The kidney
has undulate margins. The intestinal loop is
present in all individuals. The seminal vesicle
is voluminous, with spiny coiling. The distal
spermoviduct is long and very slender (Fig.
54). The bursa copulatrix is elongate club-
shaped or tapering, in one population (Pleis-
tinger Wórth) being almost of the elongate
tadpole type. The bursa duct is wide, not nar-
rower than the vagina. The sperm duct is
40 MEIER-BROOK
FIGS. 53-55.
mum shell diameter 4.3 mm, 3% whorls (M 291). 55. Variation in shape of penis tip and position of penis
pore. A: Soufflenheim (M 292); B-D: Pleistinger Wörth (M 293); E: Drusenheim (SMF 246 312). Scale =
0.1 mm.
wide. The prostate gland has a low to mod-
erate number of diverticula (Table 2), which
are closely spaced and regular in form. The
length of the 3 copulatory organ is about the
same as in G. albus or longer (Pleistinger
Wörth). The penis sheath which is extremely
short and narrow is scarcely set off from the
vas deferens. The preputium is considerably
wider and longer, the ratio psh:prp being far
below one. The penis is correspondingly tiny
and slender. The penis tip is tapering where
in G. albus and other species the thickening
rm
vd
G. rossmaessleri. 53. Shell, Drusenheim. 54. Reproductive system. Soufflenheim. Maxi-
occurs. The penis pore lies where tapering
begins. The stylet is hob-nail shaped (Fig. 55).
Remarks—Beside my previous studies on
Gyraulus rossmaessleri there is only one fur-
ther paper figuring the reproductive system
of this species, as found in a Czechoslova-
kian population by Hudec (in Macha, 1963:
87). In the shape of the bursa copulatrix, its
wide duct, and in the 4 copulatory organ there
is agreement with the present finding. As the
status of G. rossmaessleri as a separate
species had not been acknowledged for a
GYRAULUS TAXONOMY (PLANORBIDAE) 41
57
FIGS. 56-57.
234 whorls (SMF 246 313).
long time (compare Meier-Brook, 1964: 233,
238 and following), distribution data in the lit-
erature will have to be revised. This is a
strictly European species.
Gyraulus riparius (Westerlund, 1865)
Planorbis riparius Westerlund, 1865: 106
(Ronneby).
Material—Kolksee (Germany), 4 specs.
Shell —The shell (Fig. 56) is very small and
delicate; most do not exceed 2.3 mm in di-
ameter and 0.6 mm in height; it is discus-like,
resembling a young Hippeutis complanatus
in general shape. The three whorls increase
rapidly; they are flattened. The periphery is
angled; the aperture is spindle-shaped in its
exterior portion. The growth lines are ar-
cuate. The surface is silky in texture and has
very fine, but relatively remote spiral striae.
Animal— The animal is weakly pigmented;
the cephalopedal mass is light grey, and the
mantle including the kidney is almost unpig-
vd
G. riparius, Kolksee. 56. Shell. 57. Reproductive system. Maximum shell diameter 2.2 mm,
mented. The kidney margins are undulate. An
intestinal loop was present in the specimens
examined. The ovotestis is large as com-
pared with the remaining parts of the repro-
ductive system (Fig. 57). The proximal por-
tion of the spermoviduct is extremely short.
The seminal vesicle has bulbous coils and is
very voluminous, approaching or exceeding
the albumen gland in size. The bursa copu-
latrix is elongate club-shaped to cylindrical,
its duct being proximally narrow, but inflated
towards its orifice, giving it the same width
as the vagina itself. The prostate has 7 to 10
diverticula, which are closely spaced and
of regular shape. The ¿ copulatory organ is
comparatively long (0.7, 0.8, 0.8, 0.9 mm). The
preputium is wide; its lumen is S-shaped. The
penis (Fig. 58) is very similar to that de-
scribed for Gyraulus rossmaessleri, with al-
most no variation in the four individuals.
Remarks— This rare species was placed in
Hippeutis for a long time. Odhner (1929) was
the first to find a stylet and therefore included
42 MEIER-BROOK
7
N
| Pia NN
N Y AN—psh
N HAN
N N A ES pen
N \ MAN
NS 7 N y
BARS PP
NA ATEN
| XA EN
== NFS == IN
E VIE N st
FIG. 58. G. riparius, Kolksee, region of transition
between penis sheath/preputium and penis tip.
Same specimen as in Fig. 57. Horizontal shading:
columnar epithelium in distal area of penis sheath.
Scale = 0.1 mm. Key to lettering p. 27.
the species in Gyraulus. His figure (Odhner,
1929, reproduced in 1956: fig. 2d) shows the
general shape of the 3 copulatory organ found
in the German specimens, but (certainly due
to the restricted optical facilities then avail-
able) his drawing is too poor to permit fur-
ther comparison in detail.
Because of the 4 copulatory organ, which
undoubtedly was derived from the type seen
in Gyraulus by size reduction of the penis and
penis sheath, and retention of preputium size,
it appears justified to place Gyraulus riparius
in its own subgenus.
Starobogatov (1967: 296) named a new
subgenus of Choanomphalus (!) with Pla-
norbis riparius as the type-species: Lamor-
bis. He also wanted to place Gyraulus ross-
maessleri in Lamorbis but gave no reasons.
This will be discussed later.
Gyraulus crista (Linnaeus, 1758)
Nautilus crista Linnaeus, 1758: 709, no. 234.
Turbo nautileus Linnaeus, 1767: 1241, no.
654.
Planorbis imbricatus Múller, 1774: 165, no.
351.
Planorbis paladilhi Moitessier, 1867: 424, pl.
22, figs. 7-14.
Armiger crista, Ehrmann, 1933: 172, fig. 105.
Gyraulus (Armiger) crista, Soos, 1935: 28, fig.
4
Armiger crista, Baker, 1945: 75, pl. 18, figs.
6-11.
Material —Kuehren (Germany), 15 specs.;
Tübingen (Germany), 5 specs.; Wollmatingen
(Germany), 4 specs.
Shell (Fig. 59) —The shell is very small, 2.2
to 2.8 (rarely >3 mm) in diameter and 0.75
to 0.9 mm in height. It has 272 to 2% rapidly
increasing whorls. The under side of the
whorls are + flattened; the upper side is well
rounded. The periphery is angled, the angle
being nearer to the under side. The last whorl
does not embrace the penultimate whorl as
in all other planispiral species of the genus,
but is loosely attached to its upper side, the
peristome thus being continuous. The shell is
almost flat or very slightly convex on its un-
der side and deeply concave on its upper side.
The whorls are traversed by ridges, these
sometimes carrying periostracal fringes pro-
jecting peripherally (Fig. 59B). Reticulate
sculpture is absent.
Animal—The animal is weakly pigmented,
the cephalopedal mass being light grey and
the mantle showing a light and diffuse pig-
mentation. The kidney margins are distinctly
undulate. An intestinal loop was present in all
specimens studied. The ovotestis and sem-
inal vesicle vary in size. The female tract is
short and wide (Fig. 60). The bursa copulatrix
is long and slender and is club-shaped. The
prostate diverticula are reduced in length, so
that the gland merely consists of the prostate
duct (Fig. 60B); only in one of the individuals
studied were there vestiges of 7 diverticula
(Fig. 60A) that indicated that, before reduc-
tion, these must have been regularly and
densely arranged. The penis sheath is long
and slender, being always longer than the
preputium (compare Table 2). The diaphragm
is very muscular, giving the knoblike thick-
ening between the penis sheath and the pre-
putium a particularly pronounced look. The
penis has a conical tip and a hob-nail stylet
(Fig. 60C, D). The penis pore lies near the tip.
Remarks— The reproductive system of this
well-defined species was previously figured
GYRAULUS TAXONOMY (PLANORBIDAE) 43
727777
ААА E
7) / [22277
\
D
FIGS. 59-60. G. crista. 59. Shell. A—ditch near Wallnau/Fehmarn (SMF 248 548); B—form found in
lenitic biotopes, Wollmatingen (SMF 248 547); C—lake form, Dobersdorfer See (SMF 248 549). 60. A—
reproductive system, Kuehren (SMF 248 543); B—prostate gland of another specimen from Kuehren (SMF
248 543); C—penis tip with stylet, Tübingen (M 302); D—middle part of 4 copulatory organ, Kuehren,
shading as in Fig. 58 (M 313). Scales: 1 mm in A and B, 0.01 mm in C, 0.1 mm in D.
by Soos (1935: fig. 4) and Baker (1945, pl.
18). There is general agreement with the
present findings. Differences concerning the
armament of the penis tip and the form of the
prostate gland exist between Baker's (1945)
figures and my findings. Soos showed 9 and
Baker 13 prostate diverticula of normal length.
There is possibly greater variation in this
character than seen in the material | studied,
but considering the smallness of the animal,
| consider the possibility that the presence of
well-defined and long prostate diverticula in
all closely related taxa fired these authors'
imagination. This is more plausible consider-
ing that in contracted animals transverse folds
are commonly found in the female tract to
which the prostate gland adheres. Greater
concentrations of pigment cells at these sites
could indeed simulate the picture of prostate
diverticula.
Gyraulus crista has usually been regarded
as a member of a separate genus, Armiger.
This idea was based on the really aberrant
form of the shell and supported by the state-
ments of previous students who did not rec-
ognize a penial stylet. Baker (1945: 76) spoke
of a “pointed fleshy papilla” instead of a sty-
let, and Odhner (1956) observed what he
called a ‘small cuticular cap” or a ‘‘bulbous
tip.” The alleged lack of a stylet led both au-
thors to exclude the species from Gyraulus.
The only publication mentioning the presence
of a stylet in G. crista was the one by Huben-
dick (1955: 466), but Odhner (1956) believed
that Hubendick had mistaken a G. riparius
for G. crista, arguing that Armiger crista did
not occur in Hubendick's collecting place.
Whatever may be the truth, the smallness of
the two species in question may have ham-
pered thorough studies. In light section the
true nature of the stylet must indeed remain
obscure. Only an extraction of the penis from
44 MEIER-BROOK
FIG.61. G. acronicus, Bodensee near Überlingen
(loc. typ.), shell.
its sheath, which was successful for the first
time during the present studies, revealed that
the armament of the penis tip in G. crista does
not differ fundamentally from that in G. ripa-
rius and G. rossmaessleri. In one stylet of G.
crista (Fig. 60C) there is even a tinge of the
brown colour typical for the stylet of Gyrau-
lus. Other features, found with the aid of oil
immersion and phase contrast microscopy,
are the well-defined border between the pe-
nis tissue and the base of the stylet, two dou-
ble refractive areas in the base and a trace
of a longitudinal line indicating that the stylet
is formed by a rolled blade as in those other
taxa of the Planorbis tribe which are equipped
with a stylet. Therefore, there is no doubt that
FIG. 62.
the stylet of G. crista has the same origin as
the stylet in any other Gyraulus species.
The possession of such a stylet alone
would not necessarily place the species in the
genus Gyraulus, as | shall explain later. But,
besides the shell character state ‘‘rapidly in-
creasing whorls” there is at least one other
feature that leaves no alternative but placing
Nautilus crista in Gyraulus, ¡.e. the distinctly
undulate kidney margin. As will become
clearer in the cladistic analysis, the undulate
kidney margin is a character very probably
having evolved only once in the Planorbis
tribe. It does not occur in any others of its
genera. G. crista must have evolved from a
parent species carrying this character state.
This stem species must have been identical
or a descendant of the one from which the
species G. parvus, G. laevis, G. rossmaess-
leri, and G. riparius evolved. According to
Hennig's quite correct and convincing argu-
ments against Mayr's (1974b) criticism (Hen-
nig, 1974), a taxon can only be regarded as
monophyletic if all descendants of one parent
species are included in this taxon. According-
ly G. crista must not be grouped in a sepa-
rate genus.
De
ur s AÑ a
С. acronicus, shell surface. A—Bodensee, faint spiral striae and weak keel; B—-Vikarsjön,
“hairs’’ protruding from periostracal ridges on growth lines. Scale = 0.1 mm.
GYRAULUS TAXONOMY (PLANORBIDAE) 45
FIG. 63. С. acronicus, variation in shape of penis tip and position of penis pore. A—Tárna; B—Ragunda;
C—Tulom; D—Krasnojarsk; E—Valtjarn; F—Ragunda; (A-F unrelaxed; microscopic preparations under
respective reg. no. in museum collections, see Appendix); G—Siggen (M 46); H—Dobersdorfer See (M
41); |—Vikarsj6n (unrel.); K—Bodensee (М 48). Scale = 0.1 mm.
Gyraulus acronicus (Férussac, 1807)
Planorbis acronicus Férussac, 1807: 105
(Bodensee near Uberlingen).
Gyraulus deformis Hartmann, 1844: 95, pl.
27, figs. 1-5.
Planorbis gredleri Gredler, 1859: pl. 5, figs.
1a-d.
Planorbis borealis (Loven Ms) Westerlund,
1875: 77-79.
Gyraulus albus acronicus, Ehrmann, 1933:
170, pl. 7, figs. 107, 107a.
Gyraulus gredleri, Ehrmann, 1933: 171, pl. 8,
figs. 106, 106a.
Gyraulus albus deformis, Jaeckel, 1962: 68.
Gyraulus acronicus, Jaeckel, 1962: 69.
Material—In addition to the material pre-
viously studied (Meier-Brook, 1964: 235), un-
relaxed snails were available from the follow-
ing localities, partially under the name
Planorbis borealis Lovén: Sweden: Vikars-
jón, 2 specs.; Ragunda, 3 specs.; Valtjárn, 2
specs.; Vojmán, 3 specs.; Tärna, 3 specs.;
USSR: Karabella, 2 specs. (named Gyraulus
rossmaessleri, det. Westerlund); Tulom, 3
specs. (named borealis, det. Odhner); Dudin-
ska, 1 + 2 specs. (2 lots; named borealis, det.
Odhner); Krasnojarsk, 3 specs. (named bo-
realis, det. Odhner); Paratunka, 2 specs. (un-
named).
Shell—The shell (Fig. 61) is large, usually
reaching 7 mm in diameter or more and 2 mm
in height; it has 4-4% whorls, is not deeply
umbilicate, with sutures not deep and whorls
flattened. The periphery is angled or round-
ed, sometimes with a small periostracal fringe.
The aperture is ellipsoid, usually oblique. The
last whorl is regularly deflected, thus giving
the shell the appearance of being deformed.
The surface is dull, and is reticulately sculp-
46 MEIER-BROOK
G. acronicus
4mm
FIG. 64. G. acronicus, a/b plotted against a in
three central European and four N European or N
Asiatic populations. 1—Bodensee (n = 117); 2—
Siggen (n = 38); 3—Dobersdorfer See (n = 40);
4—Vojmán (n = 25); 5—Tulom (n = 13); 6—Ras-
no-volok (n = 14); 7—Krasnojarsk (n = 18). Cor-
relations significant: no. 5: p < 0.05, all other sam-
ples: p < 0.001.
tured. The sculpture sometimes is almost in-
discernible, sometimes stronger, but never
coarse as in Gyraulus albus. Spiral striae are
less remote than in G. albus. Fine periostra-
cal ‘‘hairs’’ can be present (Fig. 62). The col-
or is light-corneous, like G. albus.
Animal—The animal is moderately grey on
the cephalopedal mass. The mantle roof has
a distinct pattern, which is sometimes weakly
developed. The kidney margins are straight.
An intestinal loop was developed in all indi-
viduals checked. The seminal vesicle varies
in shape. The bursa copulatrix is elongate
club-shaped. The prostate gland has a high
number of diverticula (compare Table 2),
which are closely spaced, have a regular
shape and are unbranched. The 3 copulatory
organ is very long (Table 2) and usually bent
twice (Meier-Brook, 1964: figs. 6-10) The
penis tip can be faintly thickened. The penis
pore lies near the proximal end of the thick-
ening or in its proximal half (Fig. 63).
Remarks—Gyraulus acronicus was long
considered a peculiar lake form of G. albus
until its identity with G. gredleri was proved
by Odhner and Forcart (compare Meier-
Brook, 1964). G. borealis (Loven, Ms) (Wes-
terlund, 1875) was for a long time recognized
as a larger boreal-subarctic form of G. gred-
еп, and now of G. acronicus. It cannot be
overlooked that G. borealis is persistently dif-
ferent in shell characters from the temper-
ate European forms of G. acronicus. In cen-
tral Europe the whorls of G. acronicus are
flattened, the last whorl is more deflected, the
periphery is distinctly angled or weakly keeled.
In N Europe and boreal-subarctic Asia, the
whorls are more rounded, the last whorl is
less deflected, the periphery slightly angled
though often decorated by a delicate perios-
tracal fringe.
A latitudinal cline is demonstrated in Fig.
64, where flatness and relative height in-
crease are expressed by regression lines for
3 central European (1-3) and 4 boreal-sub-
arctic samples.
Gyraulus acronicus is the largest European
species of the genus. It is characterized by
the long, slender and bent 3 copulatory organ
and by its mantle pigmentation, showing a
distinct pattern although this is less strong
than in G. chinensis (Fig. 12). It is the only
European indigenous species with such a
pattern. Hubendick's (1955: 479) misidentifi-
cation of G. albus as G. acronicus was point-
ed out previously (Meier-Brook, 1964: 236).
The only study of the anatomy of this species
from outside central Europe is the one pub-
lished by Khazannikov (1973, fig. 4) from Riv-
er Erek Basin, Caucasus. In the shape of the
figured ¿ copulatory organs there is close
agreement with my previous figures of G. ac-
ronicus (Meier-Brook, 1964, figs. 6-10). Kha-
zannikov's printed drawings, however, are too
poor to allow comparison of internal details.
Final Notes on European Species
The main results of anatomical studies on
the European Gyraulus species, except the
endemic species from Macedonia, which will
be dealt with in a separate chapter on an-
cient lakes species, are summarized in Table
2. Provisional grouping into subgenera will be
substantiated after discussion of relation-
ships between all taxa studied (see below). A
key is included as an aid to identification of
species indigenous to or introduced to Eu-
rope outside the Macedonian lake basin. It is
based on diagnostic characters. These are
GYRAULUS TAXONOMY (PLANORBIDAE) 47
not necessarily indicators of relationship. The phylogenetic relationships are suitable for
dendrograms (Figs. 107, 111, 112) showing identification as well.
ЗВ.
4A.
4B.
5A.
5B.
6A.
6B.
TA.
ZB:
8A.
8B.
Key to the Gyraulus Species Living in Europe Excluding Macedonia
РЕЛЕ A te de tdo PR PR AS ео 2
х LENPNELYTOUNdedL..- ns о Пена 5
. Shell diameter not exceeding 3 mm; mantle pigmentation diffuse or lacking; kidney
А en e ee Se ee N Ac once sie eee ee 3
. Shell diameter exceeding 4 mm, mantle pigmentation with distinct pattern, kidney
O A A A à CR RE e Sl PRET TER 4
. Angle approximately in the middle between upper and under side; shell therefore
discus-shaped; surface with silky lustre; penis sheath shorter than preputium, not
widened proximally; prostate gland with 7 to 10 regular diverticula which are densely
AENA A A O re ee tora ela: G. (L.) riparius
Angle near under side, therefore shell + flat on under side, whorls well rounded on
upper side; last whorl not embracing the penultimate, but loosely attached to its upper
side; whorls traversed by ridges (costae) at greater intervals; sometimes ridges pe-
ripherally projecting to spiny lamellae; penis sheath longer than preputium; prostate
diyertieulallacking'orvestigial 7 ... ns NN RES G. (A.) crista
Shell rough and not glossy, usually with distinct reticulate sculpture, spiral striae
relatively distant; prostate gland with 20 to 40 diverticula; pigmentation of mantle
ОЕ О О o о и а С. (G.) acronicus
Shell smooth and glossy, at most faintly reticulate sculpture, then spiral striae very
fine and dense; prostate gland with 8 to 20 (exceptionally up to 24; in Europe so far
only between 12 and 18) diverticula; pigmentation pattern on mantle distinct and rich
in contrast; introduced from Asia to rice fields in N Italy and Camargue; also found
mevvestirrisia.(Netherlands)' ©... e ©... esse ee AO G. (G.) chinensis
Shell surface smooth, glossy, aperture rounded, kidney margins undulate, 3 copula-
tory organ 1 to 2 mm long, penis sheath distinctly longer than preputium, widening
proximally, prostate diverticula mostly irregular and not closely spaced ............. 6
Shell surface with fine or coarse spiral striation or, if appearing glossy, aperture with
white lip; kidney margins straight or undulate; ¿ copulatory organ of varying length;
penis sheath distinctly shorter and narrower than preputium, not widening proximally,
prostate diverticula regular and closely Spaced: "Rd oe 7
Penultimate whorl distinctly elevated, distal portion of spermoviduct slender, not wider
than widest portion of sperm duct; distal half of vas deferens much wider (2:1 on an
average) than proximal half (introduced from N America) ................. G. parvus
Penultimate whorl not or not distinctly elevated, distal portion of spermoviduct wider
than widest portion of sperm duct and vas deferens, distal half of vas deferens not
CONSPICUOUS IVAWIGEMeC m CPR a Ie oo G. laevis
Aperture circular, with white callous lip, shell surface with very fine spiral striation,
almost glossy, color red-brown, penis sheath distinctly shorter and narrower than
preputium, not widening proximally, penis narrowing distally ........ G. rossmaessleri
Aperture ovoid, without callous lip, shell surface with fine or rough spiral striation,
color whitish to light-corneous, penis sheath distinctly longer than preputium, widen-
ndiproxmaly; penisithickening Чат. 8
Shell with fine spiral striation, prostate gland with 20 to 40 diverticula, 3 copulatory
organ longer than 2 mm, usually bent twice (S-form), penis pore near proximal end
of thickened penis tip or in its proximal half ......... non-angled form of G. acronicus
Shell with coarse spiral striation, prostate gland with less than 22 diverticula, 3 cop-
ulatory organ shorter than 2 mm, bent at most once, penis pore subterminal in thick-
enedipenisttiprorimntsrdistallhalf' er... seco re ee nenn u G. albus
48 MEIER-BROOK
FIG. 65.
521 181.
G. euphraticus, Samava, lectotype ZMZ
SPECIES OF SOUTH AND
EAST ASIA
Before conclusions are drawn as to species
delimitations material is treated under prelim-
inary names that are widely used in the liter-
ature.
Gyraulus euphraticus (Mousson, 1874)
Planorbis (Gyraulus) devians Porro var. eu-
phratica Mousson, 1874: 40 (Samava/Eu-
phrates).
The original series on which Mousson's
description is undoubtedly based (Mousson's
handwriting kindly confirmed by Dr. Zilch)
consists of two lots with identical labels. One
contains hundreds of shells, obviously un-
sorted (ZMZ 521 180) and mixed with mud
particles, in the other one (ZMZ 521 181)
Mousson has sorted out 33 specimens
(probably the ‘‘beautiful’’ ones, as usual at
that time). A lectotype has been selected from
the latter one (Fig. 65). A second specimen
from lot no. 521 180 is also figured here (Fig.
66) to demonstrate the extreme conchologi-
cal variation in the original series. There are
all transitional shell forms between the fig-
ured ones, and the full range of variation is
even wider. No gap is visible within the range.
Material —S Iran, 30 specs.
Dr. Massoud, who kindly put at my dispos-
al living snails and preserved specimens from
the collecting site in S Iran was unable to give
more precise information on the locality. The
stock had been kept in his laboratory for par-
asitological research since previous epidemi-
ological surveys in Khuzistan. At least this is
not far distant from the type-locality of Gy-
raulus euphraticus. The shell form in this
stock resembles Mousson's original series
and can therefore be identified with his
species (Fig. 85, lines 7 + 8). The following
description is based on both series, as far as
shell characters are concerned.
Shell —The shell is of medium size, the
maximum diameter scarcely reaching 7 mm;
it is rather flat (<1.5 mm high). As the last
whorl is only exceptionally deflected, it is
possible to give a reliable figure of the ratio
maximum diameter : height: in 100 randomly
selected shells between 2.5 and 5.2 mm di-
ameter and mean values of 3.76 and 1.10
mm (ZMZ 521 180) it is 3.42. Three and a
half to 4/2 whorls are rapidly (Fig. 66) or
slowly (Fig. 65) increasing, in the latter case
nearly approaching Anisus in appearance.
The whorls are slightly or considerably wider
than high. The periphery is angled. The shell
is not deeply umbilicate. The surface is
smooth, glossy; no reticulate sculpture is
present. The color is light-corneous.
Animal— The animal is light grey, the man-
tle pigmentation showing a distinct pattern,
similar to that figured in Gyraulus chinensis
(Fig. 12). The kidney margins are straight. An
intestinal loop is absent or reduced in size.
The seminal vesicle is bulbous (Fig. 67) or
slightly spinous. The bursa copulatrix has a
spherical club shape and is sometimes taper-
GYRAULUS TAXONOMY (PLANORBIDAE) 49
67
FIGS. 66-67.
G. euphraticus. 66. Samava, paralectotype (ex ZMZ 521 180). 67. S Iran. Reproductive
system. Maximum shell diameter 3.7 mm, 32 whorls (М 121).
ing though inflated at its upper end. The va-
gina is inflated in most specimens. The pros-
tate gland bears 9 to 18 diverticula (cf. Table
6). The 4 copulatory organ is similar to that
of G. acronicus in general, but shorter, bent
no more than once with the penis sheath rel-
atively longer (cf. Tables 2 and 6). The penis
tip and the position of the penis pore are in-
distinguishable from those in G. acronicus.
Preputial pilasters are present.
Remarks—Mousson’s description is so in-
sufficient (‘'Fere regularis, pallide corneo-hy-
alina’’), like most attempts at verbal charac-
terizations of shells, and moreover, the
description suggests similarity to Gyraulus
devians Porro (which is probably a synonym
of G. acronicus) that misidentifications are
understandable. The first and as yet only an-
atomical account of a snail called G. euphra-
ticus was published in 1919 by Annandale 8
Prashad. These authors, complaining about
frequent confusions, “because conchologists
have rarely seen specimens from the original
localities” (Annandale & Prashad, 1919: 52)
probably succumbed to the same error. The
shell figured from Quetta, Baluchistan,
(30°15’N; 67°00’E), and said to “agree closely
with shells of Mousson's G. devians var. eu-
phratica recently collected by Captain C. L.
Boulenger in Mesopotamia,’’ measures 8.9
mm in max. diameter and 2.55 mm in
height(!). It does not fall in the range of vari-
ation shown in Mousson's original series.
Moreover, the distance between Quetta and
Samava is 2160 km, and there are large des-
erts between the localities, so that great
doubts arise as to their identification. The fig-
ures of the reproductive system prove that
the animals dissected were indeed Gyraulus
but further details are lacking due to the in-
sufficient optical facilities of that time. The
fact, by the way, that Annandale & Prashad's
fig. 5D—of a penis sheath of G. euphraticus
(according to the caption)—is not that of a
Gyraulus, is evidently due to an accidental
exchange of letters D and F. The shell figured
by Annandale & Prashad (1919: fig. 7A) is so
similar in size, shape and proportions to shells
in the British Museum of Natural History (no
registration number) under the name of Plan-
MEIER-BROOK
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GYRAULUS TAXONOMY (PLANORBIDAE) 51
FIG. 68. G. hebraicus, Bahr el Houle, Syria; lec-
totype (MHN Genève).
orbis nanus that species identity is probable.
The lot is labelled: ‘‘Planorbis nanus Sower-
by, Loc. Ganges, India. H. Cuming colin. 3
specs, Асс. no. 1829—Apparently not the
specimens figured in Reeve, Conch. Icon. XX,
Planorbis, PI. 9, Sp. 75.” The largest of the
3 shells measures 8 mm in diam. and 2.5 mm
in height. The anatomy of P. nanus, however,
could not be examined for comparison.
The question of a relationship between G.
euphraticus and G. chinensis will be dis-
cussed later.
Gyraulus hebraicus (Bourguignat, 1852)
Planorbis hebraicus Bourguignat, 1852: 23,
Nr. 3 (Bahr-el-Houlé, Syria).
A lectotype is designated here from Bour-
guignat's original lot in Musée d'Histoire Na-
turelle Genéve (no registration number) la-
belled ‘‘Planorbis hebraicus Bourg., type,
FIG. 69. G. piscinarum, Baalbeck, Lebanon; lec-
totype (MHN Genève).
Bahr-el-Houlé (Зупе)” (Bourguignat's hand-
writing kindly identified by Dr. Zilch). The lot
contains 2 shells of 5.9 mm diameter and 1.55
mm height, and 4.85 mm diameter and 1.50
mm height; the smaller one has been select-
ed (Fig. 68).
Material —Since alcohol material from Syr-
ia or Lebanon was not obtainable, material
tentatively identified as G. hebraicus from the
geographically least distant localities in Tur-
key was examined: Kumluca, 2 specs., Li-
myra, 8 specs. (all unrelaxed).
Shell —The shell (Fig. 68) is small to medi-
um-sized, rather flat, usually 3 to 5 mm in
diameter and 1.2 to 1.6 mm in height with 3%
to 4 whorls which increase rather rapidly,
flattened and wider than high. The shell is
equally concave on both sides. The periphery
is slightly angled. The surface is smooth,
glossy; reticulate sculpture is absent. The
color is light-corneous.
Animal—The animal is light grey, the man-
tle pigmentation showing a distinct pattern
52 MEIER-BROOK
that is marbled but not as contrasting as in
Fig. 12. The kidney character state was not
determined in the contracted animals. An in-
testinal loop was present in all individuals. The
bursa copulatrix has a club shape. The pros-
tate gland bears 11 to 15 closely spaced and
regular diverticula. The ¿ copulatory organ,
as judged from the contracted material, is not
much shorter than in Gyraulus euphraticus
(Table 6), the ratio penis sheath : preputium
similar to that in G. euphraticus. The position
of the penis pore was not discerned due to
poor fixation.
Remarks— There are few records of Gy-
raulus hebraicus in the literature. Beside the
type lot there is one other lot in MHNG iden-
tified by Bourguignat from ‘fossés pres du
camp Deer (almost illegible) pins, pres de
Beyrouth. Mahmoud,” containing 7 shells.
The shells are similar to those of G. euphra-
ticus, though slightly higher. If the material
anatomically examined were from the type-
locality | should not hesitate in considering
euphraticus a junior synonym of G. hebrai-
cus. But before a widely used name such as
G. euphraticus is withdrawn, one should wait
for an opportunity to study topotypes of G.
hebraicus.
Gyraulus piscinarum (Bourguignat, 1852)
Planorbis piscinarum Bourguignat, 1852: 22,
no. 2 (Baalbeck, Lebanon).
A lectotype is designated here (Fig. 69) from
the lot labelled ‘‘type’’ by Bourguignat
(MHNG, no registration number), from Baal-
beck. The lot includes 63 paralectotypes.
Material—Diyarbakir (Turkey) 5 specs.
(unrelaxed).
Shell—The shell is of medium size, 5 to 6
mm in diameter, and relatively high due to a
largely deflected last whorl, with a height often
up to 2.5 mm. Three to four whorls rapidly
increase. The whorls are not flattened (Fig.
85, regression line 2). The periphery is round-
ed or at most slightly angled (Fig. 70). The
under side is convex and hardly umbilicate.
The upper side is deeply umbilicate. The last
whorl is slightly to strongly deflected. The ap-
erture is oblique. The surface is moderately
smooth and silky. In the sample anatomically
examined there is very faint reticulate sculp-
ture. The color is dark-corneous to light
brown. Gyraulus piscinarum is pronouncedly
thick-shelled.
Animal—The animal is dark grey, with dif-
fuse and dark mantle pigmentation. The kid-
ney margins are straight. An intestinal loop
was present in the individuals seen. The ovo-
testis is large (Fig. 71). The seminal vesicle is
of the usual size and spinous. The bursa cop-
ulatrix is of the spherical tadpole type, with a
long and narrow duct. The prostate gland
bears 12-16 closely spaced, fleshy diverticu-
la. The vas deferens is equally narrow over
its full length. The 4 copulatory organ is rel-
atively long (Table 6), with a varying ratio pe-
nis sheath : preputium. The penis tip has nor-
mal thickening, the penis pore being situated
in its middle part or the proximal half (Fig.
72).
Remarks—Gyraulus piscinarum is a well-
defined species, even on purely conchologi-
cal grounds. Its convex under side makes it
one of the most easily recognizable species
in the genus. It is the only Asiatic species
seen in this study having diffuse mantle pig-
mentation. Its bursa copulatrix of the tadpole
type seems to be a species-specific charac-
ter state. Nevertheless, G. piscinarum has
often been confused in the literature and in
collections. Bourguignat himself erroneously
used this name for material of two lots. One
contained one specimen from ‘‘fossés pres
de Beyrouth, Mahmoud” (probably from the
same locality as G. hebraicus, see above),
which is G. hebraicus. The other lot from “le
Danube a Ibraila’’ contained one shell of G.
rossmäessleri. Records of G. piscinarum in
the literature are quite common, but in my
opinion Bourguignat's types were insuffi-
ciently known to many authors, and G. pis-
cinarum probably has a wider distribution in
the literature than in nature.
Gyraulus ehrenbergi (Beck, 1837)
Planorbis ehrenbergi Beck, 1837: 119.
Planorbis mareoticus Innes, 1884: 339.
Material— Cairo (Egypt) 13 specs.; Alex-
andria (Egypt) 4 specs. (unrelaxed).
Shell (Fig. 73)—The shell is small, usually
not exceeding 4 mm in diameter, but relative-
ly high: 1.3 mm (Fig. 85, line 1); 372 whorls
increase rapidly. The periphery is slightly an-
gled, with a small, but distinct fringe of peri-
ostracum. The last whorl is scarcely deflect-
ed. The surface is smooth and silky. Growth
lines are very fine. No reticulate sculpture is
present. The color is pale-whitish.
Animal—The animal is almost hyaline, al-
most lacking pigment. The cephalopedal
. mass is almost white. The mantle pigmenta-
GYRAULUS TAXONOMY (PLANORBIDAE) 53
FIGS. 70-72. G. piscinarum, Diyarbakir. 70. Shell (SMF 246 318). 71. Reproductive system; maximum
shell diameter 5.0 mm, 3% whorls (SMF 246 317). 72. Penis tips (M 294). Scale = 0.1 mm.
tion has a distinct, though weakly developed,
pattern. The kidney, heart, stomach and pre-
putium which in most other Gyraulus species
are darkly pigmented, bearing only scattered
black spots. The kidney margins are straight.
An intestinal loop is present. The bursa cop-
ulatrix is of the elongate club type, with a duct
about half as wide as the bursa itself (Fig.
74). The prostate gland bears 14 to 19 diver-
ticula (Table 6), which are closely spaced. The
vas deferens is equally narrow over its full
length. The 4 copulatory organ is relatively
long. The ratio penis sheath: preputium is
high (Table 6). The penis tip is normally thick-
ened, the penis pore lying in its proximal half.
Remarks—The snails studied of the Egyp-
tian populations belong to a well-defined
species, without close affinity to the W Asiat-
ic species dealt with on the preceding pages.
Apart from their relative height they do not
have many character states in common with
Gyraulus piscinarum, as might be implied by
Jaeckel's (1962: 104) considering G. pisci-
narum as a subspecies of G. ehrenbergi. G.
ehrenbergi differs from G. piscinarum not only
in shell characters, but also in pigmentation,
width of the bursa duct and numbers of pros-
tate diverticula.
A close relationship of Gyraulus ehrenbergi
with other African species, anatomically
known as G. costulatus (Krauss) and G. con-
nollyi Brown 8 van Eeden, is improbable. For
comparison, G. costulatus is briefly charac-
terized below.
54 MEIER-BROOK
FIGS. 73-75.
Е
5 E Е
7 (
f
cg g PrP
w
G. ehrenbergi, Cairo, shell (SMF 246 319). 74. G. ehrenbergi, Cairo, shape of 3 bursae
copulatrices and 2 4 copulatory organs. 75. C. costulatus, Liberia: reproductive system; maximum shell
diameter 4.9 mm, height 1.3 mm, 3% whorls.
Gyraulus costulatus (Krauss, 1848)
Planorbis costulatus Krauss, 1848: 83, pl. 5,
fig. 3 (Umgeni Valley, Natal).
Material —Liberia, five specs.
Shell —The shell is small- to medium-sized,
scarcely reaching 5 mm in diameter and of
moderate height (1.3 mm). Three and two
thirds whorls increase rapidly. The periphery
is angled, with a small fringe of periostracum.
The surface is glossy, with regular costae in
the direction of the growth lines.
Animal—The animal is uniformly light grey.
The mantle has many well-defined small pig-
mented areas. The kidney has some tiny sep-
ta which, however, do not render its margins
undulate. An intestinal loop is present. The
entire reproductive system (Fig. 75) is deli-
cate and slender: the ovotestis is very small.
The spermoviduct is long and slender. The
seminal vesicle is inconspicuous. The albu-
men gland is composed of few long acini. The
bursa copulatrix is tiny and is of an elongate
club shape or tapering. The sperm duct is ex-
tremely narrow. The prostate gland bears 5
to 8 diverticula, which are irregular in shape
and arrangement, the most distal one not
touching the bursa copulatrix as in all Pa-
laearctic species, but distant from it. The 3
copulatory organ is of the normal form, but
very short, scarcely exceeding 1 mm in length.
The penis sheath is approximately the same
length as the preputium or is slightly longer.
The proximal end of the penis sheath does
not reach the vagina in fully extended ani-
mals.
Remarks— The anatomy of this widely dis-
tributed tropical and southern African species
‘
GYRAULUS ТАХОМОМУ (PLANORBIDAE) 55
has been well known since various authors
have dealt with it (Fraga de Azevedo et al.,
1961; Wright, 1963; Brown, 1965). A partic-
ularly comprehensive study has at the same
time shown that there is a South African
species resembling Gyraulus costulatus in
many details, but clearly separated (Brown &
van Eeden, 1969).
A comparison of the two African species,
Gyraulus costulatus and G. connollyi with
Palaearctic species shows several striking
deviations from character states usually seen
in the genus, namely the extremely small
ovotestis, the tiny seminal vesicle, the very
narrow sperm duct, the number of prostate
diverticula that is by far lower (in Ethiopia no
more than three, according to Brown, 1965),
and the long distance between the most dis-
tal diverticulum and the bursa copulatrix.
These character states, evidently commonly
derived in G. costulatus and G. connollyi,
strongly suggest the existence in Africa of a
separate subgenus. Bourguignat (1883: 99)
has established a genus Caillaudia with the
type-species G. angulata Bourguignat. G. an-
gulata is commonly accepted as a junior syn-
onym of P. costulatus. Thus Caillaudia is
available as a subgenus name for at least the
two African species mentioned above.
Gyraulus convexiusculus (Hutton, 1849)
Planorbis convexiusculus Hutton, 1849: 657
(Candahar, Afghanistan).
Hutton's notes on habitats begin with ““Oc-
curs plentifully at Candahar in tanks.” Local-
ities further listed by him are Quettah, Kojuck
Pass, river Helmud at Girishk (all in Afghani-
stan) and, furthermore, the Gangetic prov-
inces, at Tope Chancey, Pinjore below Simla.
| consider Candahar the type-locality.
Material —Sayedabad, 3 specs. (unre-
laxed); Gawargin (both Afghanistan), 2 specs.
(unrelaxed), Mauritius, 3 specs. (named G.
mauritianus (Morelet) by G. Mandahl-Barth)
(unrelaxed); Bangalore, 6 specs. (unrelaxed);
Bharatpur (both India), 4 specs. (unrelaxed);
Thailand, 2 localities: 4 + 4 individuals (un-
relaxed); further alcohol material was seen
from Ceylon, but authorities of the Nat. Hist.
Mus. Vienna did not permit dissection.
Shell (Fig. 76)—The shell is small- to me-
dium-sized, 4 to 5 mm in diameter and rela-
tively high (1.2 to 1.8 mm). Three and a half
to 334 whorls increase rapidly. The umbilicus
on the upper side is deep, on the under side
shallow. The sutures are not very deep. The
last whorl is slightly deflected. The periphery
is rounded or angled, sometimes with a fringe
of periostracum. The surface is smooth and
glossy, without reticulate sculpture. The col-
or is light to dark corneous.
Animal—The animal is light to very dark
grey. The mantle has a distinct pigmentation
pattern, in intensity varying from very dark
(Sayedabad, Mauritius) to light (Gawargin,
Bangalore, Thailand). The kidney has straight
margins. An intestinal loop is absent or pres-
ent. The bursa copulatrix is small, spherical
or elongate, its duct in most individuals being
wide and not distinctly set off; the duct is as
wide as the bursa itself (Fig. 77). The pros-
tate gland bears 10 to 15 (X = 13.1 + 1.7) (in
Bangalore 20-24) closely spaced and regu-
larly shaped diverticula, which are occasion-
ally bifid. The vas deferens is as narrow as in
most species. The 3 copulatory organ is of
the usual shape and 1-2 mm long. The penis
sheath is as long or slightly longer than the
preputium. Preputial pilasters are present.
The penis tip has a thickening, with the penis
pore in varying positions, in two individuals
from Thailand virtually terminal (Fig. 78D), in
the other ones subterminal, in the distal half
of the thickening or near its middle.
Remarks—| have not seen the full range
of characters in the few (unrelaxed) individu-
als | had available and better knowledge of
variation will require study of additional ma-
terial. Original material of Hutton's collection
was to be found neither in ZSI, Calcutta, nor
in BMNH, London. Annandale & Prashad
(1919: 53) figured G. convexiusculus from
Quetta, one of the localities indicated in Hut-
ton’s description. Their shell (Annandale &
Prashad, 1919, fig. 7B) measures about 10
mm in maximum diameter, whereas Hutton
(probably measuring one of the largest indi-
viduals, as usual at that time) speaks of 6.25
mm (“4 of an inch”). The differences are
striking, and the question may be raised
again, whether Annandale & Prashad exam-
ined Planorbis nanus Sowerby, as has been
suggested with their “*G. euphraticus.” Star-
mühlner (1974: 168-171) figured organs of a
snail from Ceylon, which is evidently conspe-
cific with those anatomically examined in the
present study. The shape and proportions of
the 4 copulatory organ as well as the narrow
width of the vas deferens agree with those
observed by me.
A more flattened and strongly angled form,
usually considered Gyraulus convexiusculus
“var. compressus Hutton,” has not been ex-
56 MEIER-BROOK
Е
ce)
76
pen
78
PP
St
FIGS. 76-78. G. convexiusculus. 76. Variation in shell shape. A—Thailand (SMF 193 798); B—Thailand
(SMF 197 350/15); C—Bangalore; D—Sayedabad; E—Gawargin. 77. Variation in shape of bursa copu-
latrix. A—Sayedabad (M 297); B, C—Gawargin (M 298); D—Bangalore; E—Thailand (SMF 193 798). 78.
Variation in penis tip and position of penis pore. A—Gawargin (M 298); B—Sayedabad (M 297); C—
Bangalore (M 299); D—Thailand (SMF 193 798).
amined anatomically. Their affinity can, thus,
not yet be judged.
Character states of the reproductive sys-
tem studied from Bangalore (India) are in good
accordance with Baker’s (1945, pl. 19, fig. 2)
drawing of a specimen from Calcutta. Con-
gruence even includes the number of pros-
tate diverticula (23), which, as seen in the
present study, is higher in Bangalore (20-24)
than in Afghanistan (12-15) and Thailand (10-
14). The data are, however, too restricted to
generalize.
Gyraulus chinensis (Dunker, 1848)
Planorbis chinensis Dunker, 1848: 41 (Hong
Kong).
Material —Hong Kong, 13 specs. (unre-
laxed).
Shell (Fig. 79)—The shell is small- to me-
dium-sized, 3.5-4 mm in maximum diameter
and 1.2-1.3 mm in height. Three and three
quarter whorls increase rapidly in width. The
last whorl is not or slightly deflected. The pe-
riphery is rounded to moderately angled. A
small periostracal fringe is present on some
of the 28 specimens available from this pop-
ulation. The surface is not glossy, with growth
lines irregular and pronounced. Fine spiral
striae are present in most individuals. The
colour is light to brownish-corneous.
Animal—The animal is light grey, the ceph-
alopedal mass with small scattered black
patches. There is a distinct pattern in the
GYRAULUS TAXONOMY (PLANORBIDAE) 57
FIGS. 79-81.
mantle pigmentation (Fig. 12). The kidney
margins are straight. An intestinal loop is
present. The seminal vesicle is bulbous to
slightly spinous. The bursa copulatrix (Fig. 80)
is elongate or spherical club-shaped and
sometimes large. The bursa duct is as wide
as the vagina. The prostate gland bears 11
to 15 closely spaced diverticula. The vas def-
erens is equally narrow over its full length.
The 3 copulatory organ has the usual shape
(Fig. 80) and is between 1 and 2 mm long.
Preputial pilasters are present. The penis
sheath is slightly longer than the preputium.
The penis has a distal thickening, in which
G. chinensis, Hong Kong. 79. Shell (SMF 246 321). 80. Variation in shape of bursa copu-
latrix and 4 copulatory organ (SMF 246 322 and М 300). 81. Variation in shape of penis tip and position
of penis pore (M 300). Scale = 0.1 mm.
the penis pore is situated laterally at various
sites (Fig. 81).
Remarks—Dunker's original material could
be located neither in the Zoological Museum
of Humboldt-University, Berlin (personal
communication, Dr. R. Kilias) nor in BMNH,
London. According to Dunker's description.
snails in the type-series reached 4.75 mm in
diameter and 1.15 mm in height, which is only
slightly larger than in the present sample. The
animals examined correspond to the original
description except for the weak spiral stria-
tion that is not mentioned by Dunker, prob-
ably due to his poor optics.
58 MEIER-BROOK
FIG. 82. G. spirillus, variation in shell shape. A—
Onna-son (ANSP A453); B—Chongpyong (SMF
246 323); C—Kunsan (SMF 246 324); D— Taiwan
(SMF 246 325).
Gyraulus spirillus (Gould, 1859)
Planorbis spirillus Gould, 1859: 40 (Ousima;
-Amami-O-Shima, Ryukyu Islands, appr.
28°08’М, 129°19’E).
Material—Motobu-cho (Okinawa; as the
locality nearest to Ryukyu-Islands), 5 specs.
(unrelaxed); Taiwan, 60 specs. (20 of which
unrelaxed); Korea: Chongpyong, 15 specs.;
Kaejong, 5 specs.; Kunsan, 8 specs.
Shell (Fig. 82)— The shell is small to medi-
um in size, rarely exceeding 5 mm in diame-
ter and variable in height: rather flat, 1.2 to
1.4 mm in height (Fig. 85, lines 8 and 10), or
less flat, 1.3 to 1.5 mm in height (lines 3 and
4). Three and a half to 4/2 whorls, slowly
(Taiwan) or more rapidly (other localities) in-
creasing in size. The last whorl is more or
less deflected. The periphery is angled, usu-
ally with a distinct fringe of periostracum. The
FIG. 83. G. spirillus, Taiwan; shell surface on up-
per side and periphery. SEM. Scale = 0.01 mm.
surface is smooth (Fig. 83) (Taiwan. Only one
of several hundred shells from a laboratory
stock had clear spiral striation!) or with retic-
ulate sculpture. Spiral striae are faint to pro-
nounced; in the latter case the surface has
periostracal lamellae on growth lines, pro-
truding in spiral rows (Fig. 82C) like the
“hairs” of Gyraulus albus (Fig. 36C). The col-
our is light to brownish-corneous.
Animal— The animal is light grey, the ceph-
alopedal mass being uniformly light grey or
with many small black dots. The mantle pig-
mentation has a distinct pattern. The seminal
vesicle is bulbous or spinous. The bursa cop-
ulatrix is extremely variable (Fig. 84), from
slender and tapering to spherical club-shaped.
The bursa duct likewise varies from narrow
to wide. The vagina is usually inflated. The
prostate gland bears 11 to 21 (Taiwan) or 8
to 19 (Korea) closely spaced diverticula (Ta-
ble 6). The diverticula are two-branched. The
vas deferens is narrow. The 3 copulatory or-
gan is of the usual shape, around 2 mm in
length. The length of the penis sheath is 1.5
to 2 times that of the preputium. Pilasters are
present in the latter. The penis tip is thick-
ened, the position of the penis pore situated
at various sites laterally in thickening.
Remarks— The lectotype of Gould's Pla-
norbis spirillus, designated and figured by
Johnson (1964, pl. 44, fig. 7), is very similar
to the flat population from Taiwan or Kae-
jong; it carries some spiral striae ('“lirae” in
Gould’s description).
GYRAULUS TAXONOMY (PLANORBIDAE) 59
УИ
vag
FIG. 84. G. spirillus, variation in shape of bursa copulatrix. A—Motobu-cho; B—Onna-son; C, D—Kun-
san; E—Kaejong; F—Chongpyong. For Taiwanese specimens see Fig. 30.
Pace (1973: 79) figured the reproductive
system of a species under this name from
Taiwan, noting that the Taiwanese speci-
mens have fewer than 20 prostate diverticula
(cf. Table 6); his fig. 16 is completely in line
with my observations. Pace, at the same time,
poses the question again that has frequently
been asked: are Gyraulus spirillus, G. chinen-
sis, G. convexiusculus, and G. euphraticus
forms of the same species? This question
must be considered anew in the light of an-
atomical characters. As Table 6 shows, there
are no essential differences between these
four species in (1) mean numbers of prostate
diverticula, (2) length of the copulatory organ,
(3) ratio penis sheath : preputium. Moreover
the shapes of the bursa copulatrix and a usu-
ally wide bursa duct as well as the penis tip
do not show differences so consistent that
species discrimination could be based on
them. Nevertheless essential differences in
shell characters exist, as Rensch (1934: 210-
211) already pointed out. He expressed the
height of the last whorl in its middle as a per-
centage of the maximum diameter, stating
(translated): “In 24 typical euphraticus this
percentage was 16.9-23.0%, mean 20.3%,
in 57 typical convexiusculus, on the other
hand, it was 21.9-35.5%, mean 27.4%.” My
own measurements of G. euphraticus from
southern Iran yielded results similar to
Rensch's (Figs. 85, 88). It is interesting to
note that the other extremely flat shells of the
group in question occur at the other end of
the Asiatic continent: in Taiwan and Korea
(Fig. 85, lines 9 and 10). Both the Iranian and
the Taiwanese snails have smooth shells
without spiral striation, and it would be diffi-
cult to tell one from another, if the Taiwanese
shells did not regularly carry a tiny, but dis-
tinct, fringe of periostracum, which is always
absent in G. euphraticus. In the whole south-
ern Asiatic region between Iran and the Far
East flat and smooth forms with anatomical
features of this group seem to be unknown.
In view of the slight differences in the group
ale
27
zn 1 == = 1 = 1 =
2 3} 4mm
a
FIG. 85. a/b plotted against a in samples from S
and E Asia. The right upper end of the regression
lines indicates the maximum value of a in the sam-
ple.
60 MEIER-BROOK
FIG. 86. G. tokyoensis, Sashiki-son, shell shape,
apertural view.
it would seem reasonable to treat Gyraulus
euphraticus, G. convexiusculus, G. chinensis,
and G. spirillus as geographical races or sub-
species of one species. One exception, how-
ever, must probably be made. According to
observations made by Dr. Mandahl-Barth and
co-workers, Charlottenlund, G. euphraticus
is sympatric with G. convexiusculus in Iran,
where they are said to occur in the same
water bodies (J. E. Jelnes, personal commu-
nication, VIII-1977). This, of course, would be
a strong argument for reproductive isolation
between these groups and, thus, for sepa-
rate species status of G. euphraticus. Hence,
the group dealt with under the names of G.
convexiusculus, G. chinensis and G. spirillus
and perhaps some others form a separate
network of races in south and east Asia. |
want to apply the term Rassenkreis for this
chain, as coined by Rensch (1929: 13, trans-
lated): ‘А Rassenkreis is a complex of geo-
graphical races having immediately devel-
oped from each other, geographically
vicariating and showing unrestricted fertility
between neighbouring races.” We do not yet
know whether G. convexiusculus, G. chinen-
sis and G. spirillus are reproductively iso-
lated. If they are reproductively isolated they
would have to be considered sibling species.
It would, in this case, not be justified to call
the group of these three a superspecies, as
defined by Mayr who replaced Rensch's ‘‘Ar-
tenkreis’’ by this term: "А superspecies con-
sists of a monophyletic group of entirely or
essentially allopatric species that are mor-
phologically too different to be included in a
single species’ (Mayr, 1963: 499). As shown
above, morphological differences between the
three groups are negligible.
A similar observation was made by Huben-
dick (1951) who studied a great number of
races of the lymnaeid snail Radix auricularia
(L.), which have a distribution like Gyraulus
chinensis, etc. The races, previously regard-
ed as species, are defined on shell charac-
ters, but show virtually no consistent differ-
ences in anatomy. He found geographic
; AS F 4
d 7 .
j FA 4 и |
<. tokyoensis, Tomigusuku-son, shell
periphery with keel and weak reticulate sculpture;
periostracum removed. SEM. Scale = 0.01 mm.
y
7
ЕС. 87.
overlap of races with transitional forms oc-
curring here and there. In western Burma and
Assam, for example, the Indian race A. a.
rufescens appears to grade into the Chinese
race R. a. swinhoei (Hubendick, 1951: 154).
In his discussion, Hubendick saw most rea-
son for steering a middle course between the
two alternatives: “geographical races con-
stituting one species’ or ‘‘separate though
closely related species, all of which ... form
one superspecies.” Finally he apparently de-
cided to speak of a superspecies, whereas |
am tending to consider the Gyraulus widely
distributed over south and east Asia mem-
bers of one Rassenkreis. The reason is that
the Gyraulus taxa called races are hardly dis-
cernible from each other, even on the basis
of shell characters. The oldest available name
for this taxon is G. chinensis (Dunker).
In the material collected by G. M. Davis on
Okinawa there are snails that | should iden-
tify as Gyraulus hiemantium. They are almost
congruent with Mori's figure of this species
(Mori, 1938, pl. 15, fig. 9). Anatomically they
do not show any substantial deviations.
GYRAULUS TAXONOMY (PLANORBIDAE) 61
FIG. 88A-C. G. malayensis п. sp., shell. Holotype
(SMF 246 333).
Should topotypes of Planorbis hiemantium
Westerlund, 1883 from Hiroshima show the
same characters, G. hiemantium may also be
included in the chinensis-Rassenkreis as a
very flat form of the race G. ch. spirillus.
Gyraulus tokyoensis (Mori, 1938)
Anisus (Gyraulus) tokyoensis Mori, 1938: 290
(Tokyo).
Material —Tomigusuku-son (Okinawa), 13
specs. (unrelaxed); Sashiki-son (Okinawa), 7
specs. (unrelaxed).
88
FIG. 88D-F. G. malayensis n. sp., shell. Para-
type, Bukateja, Java (RMNH 9056/alcohol).
Shell (Fig. 86) —The shell is large, regularly
exceeding 7 mm in diameter, relatively flat (2
mm in height), with 474 to 42 rapidly increas-
ing whorls. The last whorl is not or only
slightly deflected. The periphery is distinctly
angled and keeled (Fig. 87), carrying a wide
fringe of periostracum (removed in Fig. 87 to
show the keel). The surface shows weak re-
ticulate sculpture. Anatomical characters
(Table 6) are like those of G. chinensis.
Remarks—The lack of anatomical differ-
ences between Gyraulus tokyoensis and the
G. chinensis-group suggests very close rela-
tionship. There are two reasons for maintain-
ing G. tokyoensis as a separate species: (1)
Size and a strongly keeled periphery distin-
guish G. tokyoensis from the G. spirillus race.
Extreme forms within the chinensis-Rassen-
kreis differ from each other not to the same
extent. (2) The two species live in the same
region, e.g. on Okinawa. Here Davis & Ya-
maguchi (1969: 147) collected Gyraulus at 29
localities. Of these, 23 had G. ch. spirillus and
7 had G. tokyoensis, while at one site the
two forms were found together. This could
also mean that they are ecophenotypes of
one species, but according to these authors
62 MEIER-BROOK
FIG. 89.
G. malayensis n. sp., paratype, Kuala
Lumpur, body pigmentation (shell removed).
“there was no gradation of G. spirillus into
G. tokyoensis.'' Consequently G. tokyoensis
apparently complies with the biospecies cri-
terion of reproductive isolation from its clos-
est relative.
Gyraulus malayensis Meier-Brook, n. sp.
Diagnosis—A species of the genus Gyrau-
lus which differs from all known species by
its extremely wide vas deferens, the penis
sheath being not much wider than the vas
deferens, and the penis pore being situated
near the middle or in the proximal half of the
penis.
Shell (Fig. 88A-F)—The shell is medium-
sized to large, 3.5 to 6.5 mm in diameter, of
varying height, the smaller snails being rela-
tively high, 1 to 1.25 mm (Bukateja, Tasik
Madu), the larger relatively flat, 1.2 to 1.55
mm (Kuala Lumpur). Three and three quar-
ters to 5 whorls increase moderately fast. The
periphery is rounded or slightly angled, an-
gled forms having a wide fringe of periostra-
cum (Kuala Lumpur). The last whorl is hardly
deflected. The aperture is round to ovoid. The
upper side is almost flat with shallow su-
tures. The under side is deeply concave with
deep sutures. The surface is smooth to
glossy, occasionally (Kuala Lumpur) carrying
faint spiral striae. Growth lines are + irregu-
lar. The colour is pale corneous.
Animal—The animal is light grey, the ceph-
alopedal mass has distinct black spots (Fig.
89). The mantle pigmentation shows a dis-
tinct pattern, either with large black areas (Fig.
89) or smaller patches like those in Gyraulus
chinensis (Fig. 12). The pseudobranch is large.
The kidney margins are straight. An intestinal
loop is present. Ovotestis, seminal vesicle,
and albumen gland are of the usual size and
form (Fig. 90). The female tract is long and
narrow. The bursa copulatrix is elongate club-
shaped. The bursa duct is long and relatively
wide (about the width of the vagina). The va-
gina is slightly inflated. The free sperm duct
is conspicuously long. The prostate gland
bears 17 to 25 (Kuala Lumpur) or 21 to al-
most 30 (Tasik Madu, not exactly determined
in contracted specimens) or 29 and 33 (Bu-
kateja) diverticula, which are densely and
regularly arranged. The vas deferens is
equally wide throughout its length (Fig. 90,
91A-B); its width is 0.12 to 0.20 (Kuala Lum-
pur), 0.10 to 0.14 (Tasik Madu) or 0.08 to
0.11 mm (Bukateja). The penis sheath is not
considerably wider than the vas deferens, its
widest portion being 0.14 to 0.24 (Kuala
Lumpur), 0.16 to 0.20 (Tasik Madu) or 0.14
to 0.16 mm (Bukateja) wide. The preputium
is of the usual form. Preputial pilasters are
present. The penis sheath is longer than the
preputium: the ratio penis sheath : preputium
in Kuala Lumpur is x=1.5 + 0.4 (n = 8);
Tasik Madu (unrelaxed!) x = 1.8, (n = 3); Bu-
kateja (unrelaxed!) x = 1.5 (n = 4). The penis
gradually tapers from about the middle of its
length. The penis pore lies near the middle of
the penis length or in its proximal half (Fig.
91C). The stylet has the normal form.
Type-locality —Kuala Lumpur (Malaysia), 5
km from the city, large abandoned tin-mining
pool. On grass and Eichhornia. J. K. Lie legit,
V1-1973.
Holotype (Fig. 88A)—6.1 mm in diameter,
1.5 mm high; a:b = 4.6:1.25 mm, 47 whorls.
Deposited in SMF 246 333/1.
Paratypes—SMF 246 334/4 and 246 326
(microscopica! preparation); RMNH, Leiden
(alcohol), no. 9055/4 from Kuala Lumpur.
SMF 246 335/3 from Tasik Madu, N Bali:
Lovina Beach, legit K. Y. T. Tjhen, VI-1976,
RMNH Leiden (alcohol) no. 9056/2 from Bu-
kateja, south-central Java, Tjhen legit, VI-
1976 (Fig. 88B)
Additional paratypes are in the author's
collection. Material examined anatomically:
Kuala Lumpur (Malaysia) 11 specs., Tasik
Madu (Bali) 4 specs. (unrelaxed); Bukateja
(Java) 4 specs. (unrelaxed).
Conchologically, this new species appears
to fall in the range of variation of what has
usually been called Gyraulus convexiusculus.
The expert for Indonesian mollusks, Dr. van
der Feen van Benthem-Jutting of Domburg
GYRAULUS TAXONOMY (PLANORBIDAE) 63
90
FIGS. 90-91.
G. malayensis n. sp. 90. Paratype, Kuala Lumpur, reproductive system (SMF 246 326).
91. Paratypes. A, B—-3 copulatory organs with vas deferens from Tasik Madu, Bali and Bukateja, Java.
C—penis of a specimen from Tasik.
(Netherlands), was kind enough to check the
shells of the Javanese and Balinesian sam-
ples and to give me her opinion (in litt. 22-XI-
1976), before | began to study their anatomy.
She used to attribute the name G. convex-
iusculus to snails of this form. In her mono-
graph of Javanese mollusks (van Benthem-
Jutting, 1956: 463) she mentions only two
Gyraulus species: G. convexiusculus and G.
terraesacrae Rensch. G. terraesacrae is a tiny
species not exceeding 2.2 mm in diameter and
is probably a separate species. Whether the
Indonesian material in the various collections
named G. convexiusculus really belongs to
this species or rather to G. malayensis п. sp.
cannot be judged without extensive anatom-
ical studies. It is hoped that future studies will
also reveal reliable conchological differences
between G. chinensis and G. malayensis n.
sp.
Most characters of the animal of the new
species shared by Gyraulus chinensis are
symplesiomorphous within the subgenus Gy-
raulus and thus of little value, but the distinct
pattern of mantle pigmentation and perhaps
also the relatively wide bursa duct are syn-
apomorphous within the subgenus. These
synapomorphies indicate close relationship
between G. chinensis and G. malayensis.
Moreover, there is a high probability that in
Indo-China there has never been any other
taxon from which G. malayensis could have
split off than the parent species G. chinensis.
And the presence of G. malayensis in Malay-
sia and in Indonesia strongly suggests that
isolation took place somewhere in or near the
Malayan peninsula.
Reproductive isolation between Gyraulus
chinensis and G. malayensis is not only un-
derstandable because of the fundamental dif-
ference in the copulatory organs but also be-
cause their geographical ranges overlap in at
least one region: a sample from Malacca, ap-
proximately 120 km SSE of Kuala Lumpur,
64 MEIER-BROOK
FIG. 92. G. eugyne n. sp., holotype (NHRM
Stockholm 2198).
recently received from Mr. Sigurdsson, con-
tained the species common in S Asia: G. chi-
nensis. From this finding it is probable that
the two species are sympatric in Malaysia.
One of the most exciting questions now 1$
whether they are sympatric also in Sumatra,
Java, Bali, and Borneo, which according to
Rensch (1936: 267) and other zoogeogra-
phers (de Lattin, 1967, fig. 58; Illies, 1971:
53-54) were separated from Java, Sumatra
and the continent at the end of the Pleisto-
cene.
Gyraulus eugyne Meier-Brook, n. sp.
Diagnosis—A species of the genus Gyrau-
lus which differs from all known species by
its extraordinarily inflated bursa copulatrix.
Shell (Figs. 92, 93)— The shell is medium-
sized to relatively large and has 3% to 4%
whorls with the periphery rounded. The last
whorl is deflected. The aperture is slightly
oblique and round to ovoid. The shell is deep-
ly umbilicate on the upper side and slightly
concave on the under side. Fine growth lines
are present. The surface is almost smooth,
with at most a few faint spiral striae. The su-
tures are deep and the whorls are rounded
laterally.
Animal—The cephalopedal mass is almost
dark grey. The mantle pigmentation consists
of a weak pattern, similar to that in Gyraulus
acronicus. In the reproductive system of three
paratypes the prostate gland had 17, 15 and
14 diverticula; the 4 copulatory organ (1.8,
3.4 and 2.2 mm long) is of the ordinary type,
the length ratio penis sheath : preputium (1.4,
1.6, 1.9) being in the range of G. acronicus.
The penis pore has not been located exactly
but seems to be at the base of the bulbous
thickening. The bursa copulatrix in all three
individuals showed enormous inflation, com-
parable to the “spherical tadpole type” in
some Planorbis-species (cf. Meier-Brook,
1976c), which has to this extent never been
observed in any other Gyraulus species. This
amount of inflation cannot solely be caused
by recent copulation. It greatly exceeds the
amount observed in freshly mated individuals
of other species. Moreover, the characteris-
tic orange color of the bursa contents, as is
usually seen in freshly mated individuals, was
absent here.
Type-locality—Inner Mongolia (‘‘Lager XI”
Honnen-tjaggan-tschollogol; Sven Hedin-Ex-
pedition, 1927).
Holotype—NHRM Stockholm no. 2198
(Fig. 92); 5.9 mm maximum diameter, 4%
whorls; 2.3 mm height; a:b = 4.25:1.55 mm.
Paratype no. 1—6.2 mm diameter; 2.1 mm
height; a:b = 4.5:1.35; Paratype no. 2—-5.7
mm diameter; 2.2 mm height; a:b = 4.15:1.5;
Paratype no. 3—-5.7 mm diameter; 2.2 mm
height; a:b = 3.9:1.5 (Fig. 93); Paratype no.
4—5.6 mm diameter, 2.0 mm height; a:b =
4.1:1.5; Paratype no. 5—6.6 mm diameter;
2.1 mm height, a:b = 4.3:1.45.
Relationship—Conchologically, there is
superficial resemblance between Gyraulus
eugyne and G. piscinarum, both having a
mean ratio a:b of 4.2:1.5 in adult specimens.
Anatomically the size and shape of the bursa
copulatrix (Figs. 71 and 94) also show some
affinity. The mean number of prostate diver-
ticula lies within the range of that in G. pis-
cinarum and G. chinensis with subgroups,
while shell characters are quite different from
those in the latter species.
So far as we know, Gyraulus piscinarum
has a restricted distribution in the Near East.
Its occurrence, as an eastern race, in Mon-
golia, approximately 7000 km distant from the
Near East, appears improbable. The shells
could be compared with the east Siberian
forms of G. acronicus, but the number of
prostate diverticula does not fall in its range.
However, close relationship with G. acroni-
cus cannot be excluded. Of the many Gyrau-
lus species described from China there is
none that is conchologically similar except,
perhaps, G. zilchianus Yen. As long as the
GYRAULUS TAXONOMY (PLANORBIDAE)
65
FIGS. 93-94 G. eugyne n. sp. 93. Paratype no. 3; (NHRMS 2198). 94. Reproductive system, including
shapes of two further bursae copulatrices. Paratypes (NHRMS 2198).
chance of studying Chinese material anatom-
ically is virtually zero (my attempt to collect
in Soochow, the type-locality of G. zilchi-
anus, was obstructed) | dare not identify the
only anatomically known species from China
with a conchologically defined species from
a locality more than 1300 km distant. Thus, |
prefer to run the risk of creating another syn-
onym in order to avoid misidentification of this
species.
Concluding Remarks on South
Asiatic Species
Our knowledge of anatomical characters in
Gyraulus species must still be called sporadic
with regard to the dimensions and geomor-
phological heterogeneity of the southern part
of Asia. We may state that there is one Ras-
senkreis extending from Middle to Far East:
G. chinensis. There is also no doubt that the
Near East harbours at least two species
clearly separated from G. chinensis, viz. G.
piscinarum and G. ehrenbergi. There is prob-
ably also a third one, G. euphraticus. In the
Far East there is one taxon considered here
as a separate species though with close af-
finity to G. chinensis: G. tokyoensis. Possibly
Japan is inhabited by more than these two
species, apart from the endemic species of
Lake Biwa. It is, of course, premature to claim
that in the entire south Asiatic region any Gy-
raulus must be a member of the Rassenkreis
of G. chinensis. Conchological variation, e.Q.
in the Chinese Gyraulus fauna (Yen, 1939, pl.
6, figs. 1-8), suggests that there are a few
more separate species, as the discovery of
G. eugyne n. sp. from Inner Mongolia implies.
No further conclusions, however, are reason-
66 MEIER-BROOK
able without anatomical studies of more ma-
terial from this region.
Although | am far from having full knowl-
edge of the species inhabiting the Orient, |
provide a provisional key. Users should be
aware of its incompleteness and of possible
weaknesses, e.g. a sure distinction between
Gyraulus chinensis and G. euphraticus. AS
pointed out above there are forms of G. chi-
nensis, such as the Taiwanese population
dealt with here under the name G. spirillus,
that are almost indistinguishable from G. eu-
phraticus. Unfortunately there are not even
anatomical characters constant enough to
support a decision, so that only geographical
distribution offers help.
A Provisional Key to the Gyraulus Species Inhabiting
Continental Southwest, South and East Asia
1A. Mantle pigmentation diffuse, without a distinct pattern and poor in contrast, underside
of shell flat or slightly concave or convex, upper side deeply concave, aperture +
oblique; 3 to 4 whorls rapidly increasing, diameter 5 to 6 mm, periphery rounded;
Saco AU se ee
BEER REN RES G. piscinarum
1B. Mantle pigmentation with a distinct pattern rich in contrast, though sometimes weakly
developed (+ pale), surface + shiny ...
2A. Vas deferens wide, not much narrower than penis sheath, not well set off against it;
penis tapering towards its tip; penis pore in middle or proximal half of penis ......
RE ee ace a G. malayensis
2B. Vas deferens slender, considerably narrower than penis sheath, always well set off
against it; penis with + distinct distal thickening with the penis pore in or close to it
ЗА. Shell scarcely >4 mm in diameter, relatively high: 1.3 mm; 3Y2 whorls very rapidly
increasing, periphery slightly angled, with a small fringe of periostracum; animal poor
in pigment, but pattern, though poor in contrast, discernible on mantle
... а. ehrenbergi
3B. Shell larger, >4 mm in diameter, mantle pigmentation rich in contrast .............. 4
4A. Bursa copulatrix much inflated (balloon-like), wider than uterus, bursa duct narrow;
shell 5 to 6 mm in diameter, scarcely concave on under side, deeply concave on
upper side; about 4 whorls rapidly increasing, periphery rounded .......... G. eugyne
4B. Bursa copulatrix + elongate, not wider than uterus, bursa duct + wide; shell periph-
ervioften angled =. = u a... 000 eked So a rl o e a A 5
5A. Shell large, >6 to 7 mm in diameter, 44 to 4/2 whorls rapidly increasing, periphery
keeled, with a conspicuous fringe of periostracum ................... G. tokyoensis
5B. Shell small, usually <5 to 6 mm in diameter, periphery rounded or angled ........... 6
6A. Shell flat, height at most 23% of diameter; surface glossy, spiral striation absent
O Na G. euphraticus (? = G. hebraicus?)
6B. Shell usually biconcave, height usually >25% of diameter, periphery rounded or an-
gled, fringe of periostracum often present, surface usually glossy; fine spiral striation
sometimes: presento vs bibi a Ds aa ss
SPECIES OF ANCIENT LAKES
Molluscs are one of those groups of or-
ganisms that first drew biologists’ attention
to highly distinctive endemic taxa in lakes of
Tertiary origin. Examples of lakes with well
known endemic molluscan faunas are Lake
Tanganyika, Lake Titicaca, Lake Biwa, Lake
Baikal, and Lake Ohrid. Regarding the Pla-
norbis-tribe (sensu Hubendick, 1955), en-
en A GENRE EN FEN G. chinensis
demic taxa have been described from the
three last mentioned lakes, all situated in the
Palaearctic region. The genus Choanom-
phalus was established as the first represen-
tative of ancient lake planorbids (Gerstfeldt,
1859). A striking character of this genus is its
pseudo-dextrality caused by hyperstrophy.
Subsequently, the first planorbid snail en-
demic to Lake Ohrid was described as Pla-
norbis (Gyrorbis) macedonicus by Sturany
GYRAULUS TAXONOMY (PLANORBIDAE) 67
(1894) who pointed out its great similarity to
Choanomphalus. Finally, anew species was
discovered in Lake Biwa and relegated to
Choanomphalus (Preston, 1916). The possi-
bility that a peculiar planorbid genus was
common to three ancient lakes so far distant
from one another opened up exciting aspects
as to the origin and evolution of such a group.
Studies on the Baikalian snails revealed that
Choanomphalus is anatomically distinct from
Gyraulus, because of the acquisition of a dis-
tal accessory gland complex at the male cop-
ulatory organ, which gives the genus a unique
status in the family (Hubendick, 1954, 1955).
There are other features, however, such as
the penial stylet, that leave no doubt that it
is the group around Gyraulus that must have
given rise to the seven Baikalian Choanom-
phalus species. Surprisingly, Lake Baikal has
no endemic species of the genus Gyraulus
itself. The Gyraulus species inhabiting the lake
has been identified by previous authors as G.
gredleri(=G. acronicus) (Kozhov, 1963: 159).
Unfortunately, no material from Lake Baikal
was available for my anatomical studies.
Species of Gyraulus from Lake Ohrid have
been examined thoroughly as regards mor-
phology (Hubendick & Radoman, 1959).
There are four species of Gyraulus in the
Ohrid basin similar to each other and strik-
ingly distinct from European species outside
the Ohrid basin. A fifth species resembles the
usual Gyraulus species, but has been de-
scribed as G. albidus (Radoman, 1953). The
first of the four unusual Ohrid species rele-
gated to a new subgenus, Carinogyraulus,
was G. trapezoides (Polinski, 1929). A new
species described from neighbouring Lake
Prespa (G. stankovici) was considered to be
Carinogyraulus (Hadzis¢ée, 1953). No doubt
the remaining three species of the Ohrid ba-
sin, G. lychnidicus Hesse, G. crenophilus, and
G. fontinalis (but not including G. albidus) can
be united in this subgenus. Probably a fifth
species, G. paradoxus Sturany, 1894, be-
longs here. Hubendick & Radoman's (1959)
information was so ample that further inves-
tigations on Ohrid species did not appear ur-
gent, and my studies are thus restricted to
the dissection of one individual of G. Iychni-
dicus (see below).
The species described as Choanomphalus
japonicus and Gyraulus biwaensis by Pres-
ton (1916) have not been previously exam-
ined anatomically. Information on these forms
is given below.
N a == a a /
a
5 /
\ ; /
F nber red | YA /
a О
N У /\
N 4 / \
О / \
FIG. 106. Probable phylogenetic relationship in
Macedonian Gyraulus species. Arrows are from
plesiomorphous (open squares) to apomorphous
characters (black squares). Shaded squares—di-
rection of change uncertain. Open circles—hypo-
thetical stem species. Black circles—contempo-
rary species.
within the Planorbis-tribe. These are the gen-
erally similar structure of the 4 copulatory or-
gan, including the hollow penial stylet formed
by a rolled blade. Moreover, a lack of pallial
folds in Choanomphalus (Hubendick, 1954:
504) relegates this species to the Planorbis-
tribe. The genera of this tribe are united by
the synapomorphy “loss of pallial folds.”
Characters giving Choanomphalus a unique
state in the family are an additional gland
complex opening immediately beside the 3
genital pore, a greatly concentrated central
nervous system, and the lack of a reflected
ureter. None of these characters run counter
to assuming its origin in the Anisus-Gyraulus
group. Also Hubendick (1954: 508; 1955:
531-532) concluded that “the morphology of
Choanomphalus seems to indicate that the
genus is most closely related to Anisus and
Gyraulus but has undergone a further evolu-
tion.” This view is shared by Kozhov (1963:
133) and Starobogatov (1958).
2 We will call the characters or character conditions from which transformation started in a monophyletic group plesio-
morphous, and the derived conditions apomorphous. . . . We will call the presence of plesiomorphous characters symple-
siomorphy, the presence of apomorphous characters synapomorphy, always with the assumption that the compared
characters belong to one and the same transformation series” (Hennig, 1966: 89).
74 MEIER-BROOK
Planorbis Anisus Bath Gyraulus
ne - —^ Sha
G re ' м 9 : = :
2 3 ; 5 5, 5 5 A E 58 5 Sy Eu
5 A и D = >5 = o 5 9 Go as = 93
es a 2 ur o £3 Е Е 5 5 25 Su 33 22
= “i 5 4 у e] oO a 4 5 © 3 Se Ou Ons oa
e o ® o o ig o e ® air >
| | | | Ш 0 h nl) - ae |
| > / 1 thick as =
| thick мое fiatte ne nn !
Le AE o ii
Е un > À
e A \ radular ee num /
| =: о Vers reel я
y /
\ —>B т ©. /
| | / я
\ | в 2 /
т ys
| | NA pe
\ | O /
kidney mar 4% 7
| | ındulate er 5%
B-- ===
| a
| RD
IEA 74
Ji od sees SO’ 7
\ laterally 7
\ DO-——>B we
=
\ x | nn Pile
O elonsted y р
MH
\ >07
\ nial Stylet
0 Set М
\ mn
o-
FIG. 107. The probable kinship relations in the Planorbis-tribe. For symbols see Fig. 106.
The question whether the Choanomphalus
clade diverged before or after the parent
species common to all recent Gyraulus
species came into being, cannot yet be an-
swered. If it diverged after, then it must show
synapomorphies that are shared with a
subgroup of Gyraulus. In this case Choa-
nomphalus would have to be included in Gy-
raulus as a subgenus according to Hennig's
(1966: 73) definition of monophyly: “А mono-
phyletic group is a group of species descend-
ed from a single (‘‘stem’’) species, and which
includes all species descended from this stem
species.” Which group of Gyraulus species
could have given rise to formation of a new
clade in the Baikal basin? Torquis and La-
morbis occur in the Palaearctic but do not
reach as far as Central Asia (Fig. 116). These
two subgenera are sufficiently recognizable
from shell characters that their past distribu-
tion in the Baikal region can be excluded as
well. The only subgenera having reached
Central Asia are Armiger and Gyraulus s. str.
Armiger has too aberrant traits to be consid-
ered derived from a stem species shared with
Choanomphalus. Thus only Gyraulus s. str.
is left. This subgenus, however, is character-
ized exclusively by synapomorphies, thus
forming a ‘‘remnant group” whose monophy-
letic status sensu Hennig cannot be proven.
Should, one day, synapomorphous character
states be found in Gyraulus species occur-
ring, in past and present, in the Baikal re-
gion and should these synapomorphies be lo-
cated in Choanomphalus, too, there would be
no argument against including Choanom-
phalus in Gyraulus as a subgenus (Fig. 107,
straight broken line). The genus Gyraulus
would otherwise not keep its monophyletic
status in Hennig's sense, as was also argued
in the case of Armiger. If Choanomphalus
branched off before the parent species of all
recent Gyraulus species existed it must fur-
thermore be regarded as a separate genus
(Fig. 107, curved broken line), as all authors
have done (Lindholm, 1927; Baker, 1945;
Hubendick, 1954, 1955; Zilch, 1960).
Let us briefly consider the possibility that
Choanomphalus branched off from a trunk
that led to other genera of the tribe. Planor-
bis, Anisus, and Bathyomphalus presently
extend to the Baikal region. The past exis-
tence of a species ancestral to—and only
to—Planorbis and Choanomphalus giving
GYRAULUS TAXONOMY (PLANORBIDAE) 75
them the status of sister genera would mean:
a structure as complex as the penial stylet in
all genera of the tribe except Planorbis, would
have developed twice independently. This is
so unlikely that | can exclude it. Anisus and
Bathyomphalus possess a derived character
state not shared by Choanomphalus: an ex-
treme elongation of the total body. In a branch
leading to Choanomphalus this elongation
would have had to revert to the shortened
state. Such a reversion is not impossible, as
Gyraulus biwaensis demonstrates. But it is
improbable that all organs re-shorten to the
same extent. The unusually long 3 copulatory
organ in G. biwaensis is an example of this
partial re-shortening. In Choanomphalus there
is no indication of a partial re-shortening (Hu-
bendick, 1954). Consequently | should think
it equally improbable to assume a common
ancestor for Choanomphalus and Anisus or
Bathyomphalus that is shared only by them
(as a condition for finding sister groups).
The question of why there is no endemic
subgroup of the genus in Lake Titicaca and
Lake Tanganyika is, in the former case, eas-
ily answered: The Andes region has evidently
been devoid of members of the genus Gyrau-
lus at all times. From Lake Tanganyika a few
records of ‘‘Anisus’’ or ‘‘Gyraulus’’ species
are found in the literature. At least one such
species was assigned to a different genus by
Brown & Mandahl-Barth (1973), namely АЕ
rogyrorbis natalensis (Krauss).” Lake Tan-
ganyika was probably outside the geograph-
ical range of real Anisus or Gyraulus during
most of its history. It is of interest, though,
that a strongly carinate form, described as
Gyraulus bicarinatus Mandahl-Barth (1954:
91) and somewhat ressembling ancient lake
forms from other parts of the world, has been
discovered in Lake Albert. However, G. bi-
carinatus belongs to Afrogyrorbis, too (Brown
8 Mandahl-Barth, 1973).
Finally the question of an affinity between
ancient lake Gyraulus and fossil groups de-
serves attention. Shells from the Upper Mio-
cene of Steinheim/Albuch, Württemberg,
Germany, cover a wide range of forms from
flat and almost planispiral to turreted and ex-
tremely pseudo-dextral. These fossil taxa
have been placed in a single species, Gyrau-
lus trochiformis (Stahl, 1824) (synonym: Poe-
cilospira multiformis (Zieten, 1830)). Compar-
ing shell characters of the Steinheim species
with those in the Planorbis tribe, particularly
with thick-shelled and angled forms of an-
cient lakes, I do not object to grouping them
in this tribe. However, the similarity of P. mul-
tiformis to Gyraulus is not greater than simi-
larity to Paleocene Carinulorbis Yen, 1949.
Even if we consider that P. multiformis may
really have had anatomical characters in
common with Anisus and Gyraulus, concho-
logical differences between the Steinheim-
group and any ancient lake Gyraulus would
be greater than between distantly related
standard Gyraulus species.
The probability substantiated above of an
independent evolution of multicarinate and
non-planispiral Gyraulus species in different
parts of the world is one more reason not to
overemphasize the possibility of a common
evolution of Ohridan species and the Stein-
heim series, as was done by Polinski (1932)
and accepted by Stankovié (1960: 244).
CLADISTIC ANALYSIS
Character Weighting
My study of morphological characteristics
has revealed a number of qualitative or quan-
titative character states enabling me to dis-
tinguish species from each other. The use of
such character states for characterizing and,
thus, for identifying species, as discussed
above, does, however, not necessarily mean
that they are suited as tools in the analysis
of genealogical relationship. One such char-
acter, to mention an example, is size. Within
a certain range, enlargement or reduction of
size appears to have happened independent-
ly in various groups where one observes
giants or dwarfs. In a discussion on weight-
ing of the ‘‘hob-nail-stylet’’ | shall show that
isolated occurrences of dwarf-forms in the
genus are encountered in Europe, Australia,
Java, and Samoa. Of course, this does not
preclude that splitting up of a tiny parent
species may result in two sibling species
maintaining their small size. Whether evolu-
tion took this course must be decided on two
bases: (1) the occurrence of other derived
characters common to these and only to
these, and (2) a contact or overlapping of their
geographical ranges at least in the past.
There are a number of processes besides
convergent evolution that invalidate the re-
3 Afrogyrorbis Starobogatov, 1967, described with the same type-species, Planorbis natalensis Krauss, has priority over
Ceratophallus Brown 8 Mandahl-Barth, 1973.
76 MEIER-BROOK
sulting characters or lower the value of such
characters for phylogenetic conclusions, al-
though not for the discrimination of species.
These are, among others, mosaic evolution,
and gradual and sporadic reappearance of
traits once lost during phylogeny.
Recognition of mosaic evolution (Mayr,
1975: 208) requires a great variety of char-
acters undergoing transformation during evo-
lution after the existence of the stem species
of a taxon. In Gyraulus few characters of this
kind have been observed, so that it cannot
be judged whether or not mosaic evolution
happened.
The problem arising through the second
event mentioned above as limiting the value
of a character may be elucidated by examin-
ing reticulate sculpture. Spiral striae, which in
adult snails form this structure together with
transverse striae (growth lines), are encoun-
tered in embryonic shells of all Planorbidae
studied so far, as stated above. Since they
occur in many other basommatophoran taxa,
too, one may suspect that this sculpture once
was common in adult shells of a taxon at the
stem of the Basommatophora or one of their
higher subordinate taxa. If this were the case
(fossil records in support of this have not
systematically been examined) then its com-
plete absence in most planorbid shells includ-
ing those of the closest relatives of Gyraulus
must be secondary and its presence in sev-
eral Gyraulus species either is due to reap-
pearance of the ancestral structure or to a
new acquisition. The observation that spiral
striae on adult shells are an immediate con-
tinuation of embryonic striae supports the
assumption of a reappearance. This would
also facilitate an explanation of the obviously
independent occurrence of spiral striae here
and there (sg. Lamorbis, Choanomphalodes,
Gyraulus s. str., Carinogyraulus), even in
usually smooth-shelled groups, such as G.
chinensis spirillus.
A new acquisition, however, can not be en-
tirely excluded. The new striae could have
started from the embryonic striae because the
shell-forming mantle edge was functionally
prepared for the formation of striae just at
these sites. If my concept, established on
several reliable characters (Fig. 107), is cor-
rect, new formations of striae, however,
would have had to arise several times inde-
pendently, since Caillaudia, Torquis, and Ar-
miger always lack spiral striae.
The above speculation shows how difficult
a decision is on what presence or absence of
reticulate sculpture tells us about phyloge-
netic relationships. As long as there is no pa-
laeontological evidence for spiral striation
being ancestral, the problem must remain
open. The value of the structure for cladistic
analysis is, thus, zero. The suitability of a dis-
tinct reticulate sculpture for characterization
and recognition of species, such as Gyraulus
albus and G. acronicus, is not affected by this
difficulty.
Characters particularly useful in the evalu-
ation of relationship are all those that, be-
sides a high stability, show (1) clear alterna-
tives (example: undulate/straight kidney
margins), (2) can be categorized as ancestral
or derived with as little doubt as possible (ex-
ample: pluricuspid/unicuspid radula teeth) and
(3) tie in well with the geographical distribu-
tion of their bearers (example: the hob-nail-
stylet in Armiger and Lamorbis).
Analytical Procedure
| have attempted to group species in su-
praspecific taxa according to a system ex-
pressing relationships and resulting in mono-
phyletic groups. By relationship one means
“overall similarity’' as discussed by most au-
thors (e.g. Hennig, 1966: 74). This can even
be expressed in terms of mathematics and,
consequently, meets the demand of modern
biology for quantification, e.g. as in numerical
taxonomy (Sokal 8 Sneath, 1963). Proceed-
ing in this way one may establish a reason-
able classification, but this does not neces-
sarily yield groups united by relationship in a
genealogical sense. Moreover, groups estab-
lished in this way are '“monophyletic” only in
a sense that is not unequivocally defined. It
was Hennig’s (1950 and later) great contri-
bution to recognize a logical and consequent
way of phylogenetic research permitting clear-
cut and reproducible decisions. Of his many
thoughts at least one has been generally ac-
cepted: the superiority of synapomorphous
over symplesiomorphous characters in judg-
ing relationship, i.e. for the reconstruction of
the actual course of the evolutionary pro-
cess. Hennig's arguments in favour of this
procedure are well known and need not be
repeated here; they have not seriously been
contradicted. Hennig's conception of mono-
phyly (Hennig, 1950: 307 et seq.; 1966: 207;
1969: 17; 1974: 283), however, has not found
general agreement. Inasmuch as the first
thought in his definition is concerned, there
is little controversy: “Only groups of species
GYRAULUS TAXONOMY (PLANORBIDAE) dol
that can ultimately be traced back to a com-
mon stem species can be called monophylet-
С” (Hennig, 1966: 207). Similar definitions
have been in use a long time, with one minor,
though important, difference. The generally
accepted use of the term monophyly was
sketched by Simpson (1961: 124): “Mono-
phyly is the derivation of a taxon through one
or more lineages ... from one immediately
ancestral taxon of the same or lower rank.”
Hennig defended his conception against this
definition by pointing out that “the device of
bisexual reproduction makes the species the
real unit. New species can arise only through
the breakdown of individually existing
species” (Hennig, 1966: 207).
There has been much argument about the
second part of Hennig's definition: “... it
must be added that not only must a mono-
phyletic group contain species derived from
a common stem species, but it must also in-
clude all species derived from this stem
species” (Hennig, 1966: 207). Acceptance of
this postulate means that one has to break
with familiar views about "natural group-
ings’ as they appear to every unprejudiced
observer. Reptilia, for example, cease to be
a monophyletic taxon, whereas extremely
dissimilar organisms, such as crocodiles and
birds, are united in one taxon as Mayr (1975:
72-75) pointed out. Hence, the overwhelm-
ing majority of taxonomists disagreed with
Hennig, i.e. all those who concentrated on the
degree of divergence and not the stem group.
Their sticking to a definition of monophyly
excluding the demand for all species derived
from the stem species being included was
criticized by Hennig (1974: 283) with the sim-
ple, but irrefutible comment (translated):
“Each pair of whatever species has a com-
mon ancestor. Following this criterion one
may establish arbitrary grouping. It is only the
postulate that animal species must have an
ancestor common only to them in order to be
members of a monophyletic group that makes
definition clear and workable.”
Actually, Hennig's conception has the ad-
vantage of being logical and allowing objec-
tive and reproducible conclusions and state-
ments on phylogenetic relationships. It avoids
decisions about the degree of overall similar-
ity or dissimilarity which, in my opinion, must
always be subjective because there will never
be general agreement about which yardstick
has to be used. Even mathematical methods
will not be able to help in this respect, as
drastic differences result depending on which
method of computation is employed using
identical data (according to Mayr, 1975: 187).
The decision between (1) the logically con-
sequent concept held by the “cladists” as
Mayr calls them and based on a biological
theory and (2) the ‘‘evolutionists’’’ concep-
tion of monophyly, taking into account the
“dual nature of evolutionary change,” but up
to now based on no clear theory, is perhaps
a matter of confession. For the analysis of
relationship between taxa dealt with here |
accept Hennig's principle and his definition of
monophyly, although | acknowledge that there
are some weaknesses in his works as, for
example, the ‘‘deviation rule.” Concerning the
deviation rule Hennig himself (in Schlee, 1971:
28) more recently explained how his thoughts
should be understood. He did not refuse
Schlee's (1971: 27) remark, that this rule
contributes to mastering a merely semantic
problem.
Plesiomorphy and Apomorphy
“Recognition that species or species
groups with common apomorphous charac-
ters form a monophyletic group rests on the
assumption that these characters were taken
over from a stem species that only they share
in common, and which already possessed
these characters prior to the first cleavage”
(Hennig, 1966: 90). What criteria can be used
to distinguish between plesiomorphous and
apomorphous characters? (1) Apomorphous
characters by definition appear later in geo-
logical time, ¡.e. in younger fossils, than ple-
siomorphous characters. (2) Plesiomorphous
characters also occur in closely and distantly
related taxa. Apomorphous characters are
unique or rare and restricted to relatively small
groups. (3) In groups with low vagility a
character is probably apomorphous if it pre-
dominates in a restricted geographic area
whereas taxa with the corresponding sym-
plesiomorphous character have a wider dis-
tribution.
In the literature, a number of other criteria
are offered, for example, in a paper dealing
with relationships in advanced snakes (Marx
8 Rabb, 1972). In their work, the authors
enumerate a total of 10 criteria for derivative-
ness. Several of them more or less repeat
each other so that their number can be re-
duced. At least one of them, in my opinion,
is open to circular reasoning, viz. ‘‘Correla-
tion of derived states. A character state is
derived if its occurrence is positively correlat-
MEIER-BROOK
78
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GYRAULUS TAXONOMY (PLANORBIDAE) 79
ed with derived states of other characters.
Such correlation of derived states of different
characters probably results from common
genetic history of taxa” (Marx & Rabb, 1972:
5). One should at least add “... correlated
with undoubted derived states of other char-
acters.”
Of the three criteria mentioned above, the
one based on fossil records does not help
much in our case. The only characters pre-
served in the fossil state are those of the shell,
and as shown in these studies, almost all re-
liable characters in this group are hidden in
the animal.
Concerning the second criterion, i.e. the
distribution of characters in Gyraulus and in
other genera of the same tribe, | refer to Ta-
ble 7. A character common to all of them is
the lack of pallial folds. This synapomorphy
does not appear in the table.
Most of the unique character states con-
fined to small groups within the Planorbis-tribe
occur solely in Gyraulus but these are not
even in the majority of species. The following
character states are regarded as apomor-
phous: undulate kidney margins, a patchy
mantle pigmentation, a loose and irregular
arrangement of prostate diverticula, an ex-
tremely wide vas deferens, the penis pore
lying remote from the penis tip, an inflated
bursa copulatrix. As to the number of pros-
tate diverticula (Table 7, column 3), this is a
quantitative character that may be subject to
repeated reduction. Reduction or disappear-
ance of an organ is therefore considered by
many authors to greatly lower its value for
phylogenetic analyses. But diverticula num-
bers in the Planorbis-tribe are not so unsta-
ble that extreme reduction would lack any
significance for relationship. For example, the
two African Gyraulus species, G. costulatus
and G. connollyi, which are very similar in
other characters, have extremely reduced di-
verticula numbers. It is very improbable that
reduction took place independently after
branching of their common stem species.
Most probably their stem species already had
reproductive organs greatly reduced in size
and complexity and, among others, ex-
pressed in very low diverticula numbers.
The problem arising with an interpretation
of shell surface structure (Table 7, column 11)
has been discussed above. The completely
smooth surface in postlarval shells of all gen-
era of the tribe except in some subgroups of
Gyraulus and Choanomphalus is striking. All
other Recent planorbid genera have a smooth
shell as well. Only few exceptions occur (e.Q.
Bulinus reticulatus Mandahl-Barth, 1954).
Since spiral striation incidentally appears
or reappears in various species there is no
clear decision in what direction transforma-
tion took place. Consequently the question
has been left open (Fig. 112, shaded squares
between G. albus and G. piscinarum).
In another case (Table 7, column 6) a rare
character is seemingly shared by Planorbis
and subgroups of Gyraulus. There is strong
evidence that the cylindrical penis sheath in
G. riparius, G. rossmaessleri, and moreover
less regularly in G. crista and G. malayensis
had their origin in the club-shaped penis
sheath typical of Gyraulus, Anisus, and
Bathyomphalus: in G. crista and G. malay-
ensis, transitions between shapes are fre-
quently found. On the other hand, the cylin-
drical shape of the penis sheath in Planorbis
(Meier-Brook, 1976c, figs. 1, 6a), corre-
sponds to the shape found throughout all
other tribes of the family (Hubendick, 1955).
The form of the penis tip shows the same
phenomenon (Table 7, column 7). A distal
thickening of the penis in groups of the Pla-
norbis-tribe as described above is unique in
the family. Its lack in Planorbis is consistent
with the state common to all other planorbid
tribes. Within the genus Gyraulus intermedi-
ary stages are observed (Figs. 52, 63) be-
tween presence (Figs. 45, 72, 78, 81) and ab-
sence of the thickening (Figs. 55, 58, 60, 91).
This suggests that its absence in certain Gy-
raulus species is secondary.
A penial stylet (Table 7, column 9) is ab-
sent in Planorbis. It is absent in the vast ma-
jority of Planorbidae though not in all of them.
Its presence in various planorbid groups sug-
gests that the question be examined whether
stylets more probably have a common origin
or have developed independently. A tenden-
cy to sclerotize the penis tip is observed in
one or the other group, according to Huben-
dick (1955) in Polypylyis (Segmentina-tribe),
Planorbula and Promenetus (Helisoma-tribe)
as well as in Physastra (Physastra-tribe).
Comparison of structure shows that all these
stylets are solid and evenly transient from the
penis tissue, both in form and in consistency.
A picture of the penis tip of a Physastra
species demonstrates this (Fig. 108). Stylets
have also been reported from quite different
groups, as in sacoglossan Opisthobranchia,
where, according to Gascoigne (1974), there
is a variety of stylets serving for transfer of
sperm, either by conducting it to the bursa
80 MEIER-BROOK
FIG. 108. Stylet in Physastra sp., L. Barracuta.
copulatrix or simply by hypodermic injection.
None of these stylets has the exact charac-
teristics of structure found in the four genera
of the Planorbis-tribe listed in the table, i.e.
the sharp delimitation against the penis tis-
sue where it is, to a certain extent, movable
like a door on a hinge. The broad base and
the stylet itself consist of a rolled blade, leav-
ing a tube inside and one proximal and one
distal opening (Figs. 25 to 27). It is highly im-
probable that a stylet of such a complex con-
struction has been formed more than once.
Of all synapomorphies encountered in the
tribe, the stylet is the one with the highest
probability of being derived.
Probably two other genera with a sclero-
tized penis tip belong in the Planorbis-tribe:
Afrogyrus Brown & Mandahl-Barth, 1973, and
Afrogyrorbis Starobogatov, 1967. Pallial
ridges are not mentioned; their lack would
clearly place them here. The sclerotized tip of
the penis in these two genera (Brown & Man-
dahl-Barth, 1973: fig. 4) is again so different
that | conclude that it must have formed in-
dependently. An origin from a stem species
that had the Gyraulus stylet is unthinkable.
Let us assume that Afrogyrus and Afrogy-
rorbis belong in the Planorbis-tribe. Then they
must have branched off somewhere between
the stem species of the tribe (Fig. 107, basal
circle) and the stem species of all Planorbis
Species.
The hob-nail stylet undoubtedly arose from
the usual stylet by size reduction, associated
with a narrowed penis sheath and penis, and
a loss of the distal thickening of the latter.
These reductions in size could be understood
as a consequence of extreme size reduction
of the whole animal as seen in the tiny
species, Gyraulus riparius and G. crista. Ac-
cordingly, one might conclude that the origin
of hob-nail stylets in these species took place
independently, by convergent evolution, were
there not a Gyraulus species with normal di-
mensions, i.e. G. rossmaessleri, which also
has a hob-nail stylet. The fact that a species
has these features although it could have
maintained the normal penis, makes it likely
that the following happened. Size reduction
culminating in the evolution of G. crista and
G. riparius took place independently (Fig.
111). So the hob-nail stylet came into being
before such size reduction. Or, at least, hob-
nail stylet formation was not an immediate
consequence of an overall reduction in ani-
mal size.
Initially | hesitated to place Gyraulus ross-
maessleri in the same subgenus as G. ripa-
rius. The reason was that conchologically G.
rossmaessleri is so similar to G. laevis that
paleontologists would find it difficult to tell
them apart. On the other hand, there was the
aberrant 3 copulatory organ also shared by
G. riparius. The undulate kidney also seemed
to support the conchological decision. Later,
when the undulate kidney was found to be
present in G. riparius and G. crista, too, it
became clear what had probably happened.
The formation of prominent kidney septa took
place when the parent species common to
the subgenera Torquis, Lamorbis, and Armi-
ger came into being. This parent species had
the normal size of a Gyraulus and the normal
rounded whorls. These shell characters are,
therefore, symplesiomorphous to G. ross-
maessleri and the subgenus Torquis, and are,
thus, not suited to prove a close relationship
between these. Moreover, abandonment of
the closely and regularly arranged prostate
diverticula, a process leading to the synapo-
morphy of the Torquis species, did not take
place in G. rossmaessleri. Size reduction of
the whole animal, including the shell plus flat-
tening towards a peripheral angulation on the
GYRAULUS TAXONOMY (PLANORBIDAE) 81
other hand, is a process occurring repeatedly
in species in different parts of the world, so
that its independent occurrence in G. crista
and G. riparius is not too unlikely. The differ-
ent types of angulation (=formation of an
edge) in the two species (Figs. 56 and 59)
support independent formation of edges.
It would be of great interest to see how
other species of the genus that have drasti-
cally reduced their size have managed the
problems of housing the penis in the в copu-
latory organ. There are tiny snails—evidently
members of the genus Gyraulus—in different
parts of the world, e.g. Planorbis singularis
Mousson on Samoa (1.6 mm in diameter),
Gyraulus terraesacrae Rensch in central Java
(2.2 mm in diameter) and a species in Austra-
lia (2.5 mm in diameter). The only character
common to these is their small size, so that
independent reduction of size is highly prob-
able. Do they possess a narrowed penis
sheath and a conical penis tip with a hob-nail
stylet?
Regarding the Australian species, here
called G. sp. 1, | could examine specimens
from two samples recently provided by Dr. B.
J. Smith, of Melbourne. My prediction that
size reduction does not necessarily cause
formation of a conical penis tip and a hob-
nail stylet, proved true. The distal part of the
penis has a distinct though slight thickening
with the penis pore near its middle and a
“miniature edition’ of a stylet in the normal
form. The space problem in the actually nar-
row penis sheath has evidently been solved
by stretching in length: the distal thickening
is proportionally longer than usual in the ge-
nus.
The bursa copulatrix is of an elongate form
in most Planorbidae. Inflated bursae are much
rarer than in the Lymnaeidae, for example.
There is a reasonable explanation for differ-
ences in abundance of the inflated bursa. The
body whorl in lymnaeid snails leaves suffi-
cient space for increased volume of organs,
whereas in planorbid snails it is simply too
narrow. When whorls are not so much flat-
tened one can indeed find a more rounded
bursa, e.g. in G. piscinarum (Fig. 71) and the
more or less globose snails of ancient lakes
(Fig. 96, 100, 103). On the other hand, as
seen in G. ehrenbergi (Fig. 74) increase of
whorl height does not automatically lead to
inflation of the bursa. And an extreme infla-
tion, as in G. eugyne (Fig. 94), certainly goes
far beyond what is plausible as a simple con-
sequence of whorl height increase. When an
external or functional causative agent for an
extreme morphological change is not ob-
vious, the value of an otherwise apparently
insignificant character is raised.
Two of the character states considered
apomorphous on the strength of rarity within
the Planorbis-tribe require some comment.
These are the undulate kidney margins and
the patchy mantle pigmentation. These char-
acters force us to reconsider the problem of
irreversibility. The question has already been
discussed in connection with reticulate sculp-
ture.
As shown above the undulate kidney mar-
gins are caused by distinct septa protruding
into the lumen of the tubular kidney. Accord-
ing to Baker (1945: 12) in the Planorbidae the
“central tube ... is regularly marked by in-
ternal septa which project into the lumen for
a greater or less distance.” This would mean
that the presence of septa is the rule in this
family, and thus their absence rendering mar-
gins straight is the derived state. Baker's re-
mark could imply that the formation of septa
in subgroups of Gyraulus is merely a reversal
of a loss, which, from time to time, occurred
during evolution. Baker's figures of kidneys
from a great variety of planorbid genera
(Baker, 1945, pls. 44-47) do not show any
sign of transverse septa except in a Gyraulus
(G. circumstriatus (Tryon) (which is a Torquis
species, where it has been placed by other
authors on a conchological basis)) and two
species of Helisoma. In the latter case | am
not even sure whether it is not an artifact
caused by fixation. | myself have never ob-
served distinct septa in any living planorbid
snail except in the case detected during the
present studies. Should real septa be discov-
ered in other planorbid groups these will
probably prove to be non-homologous to
those described here. The undulate kidney
margins in some Gyraulus species are justi-
fiably called apomorphous.
A probable reversal in evolutionary change
is seen in the patchy pattern of mantle pig-
mentation. It is the rule in Planorbidae as well
as in other basommatophoran families and
also in various prosobranch groups. The
patchy pigment is probably ancestral. Since
a pigment pattern is not a complex structure
only a small step is necessary from a distinct
pattern to diffuse pigmentation and back
again. Nevertheless this step has not oc-
curred as frequently as one might suspect.
Otherwise, the two alternate states should
be distributed at random. In Lymnaeidae we
82 MEIER-BROOK
see two patterns. These are (1) a distinct
pattern or (2) a more uniform pigmentation
with small cutouts. These alternatives are
usually clearly distributed and consistent
within genera (genera have been generally
retained by authors in spite of Hubendick's
(1951) opinion), e.g. Radix and Stagnicola,
respectively. The diffuse pigmentation of
Planorbis, Anisus, Bathyomphalus and sev-
eral Gyraulus species is at least so constant
that it may be considered ancestral within the
Planorbis-tribe. Accordingly the reappeared
pattern is derived. This conclusion is sup-
ported by the geographic distribution of
species with the patchy pigment pattern as it
relates to the present systematic analysis of
species. The patchy pigment pattern is miss-
ing in Armiger, Lamorbis, Torquis, Choa-
nomphalodes, and Carinogyraulus. In the
African subgenus Caillaudia the patchy pig-
ment consists of a great number of unusually
small spots, so that its independent deri-
vation from diffuse pigmentation is not im-
probable. In the remaining species, tentative-
ly united in a subgenus Gyraulus s. str. a
patchy pigment pattern occurs only in N Eu-
rope, Asia and Indonesia. Of the two Austra-
lian species | have seen none* having mantle
pigmentation at all, so that a decision is im-
possible. Whether the patchy pigment pat-
tern occurs in the North American species is
an important question that has not yet been
examined. So far we may assume that reap-
pearance of the patchy mantle pigmentation
in Gyraulus has probably happened only once,
in western Eurasia. From this point the first
species with the patchy pattern spread and
split up into a northern (G. acronicus) and a
southern branch (the stem species of all
species living in southwest, south, southeast
and east Asia including Indonesia: G. ehren-
bergi, G. euphraticus etc.) (Fig. 112). These
two branches may have dispersed mainly in
an easterly direction. While G. acronicus evi-
dently did not evolve into isolated species,
the southern stem species, on its way to the
east, branched off several times. The latter
process may have been favoured by the ir-
regular shape of the continent south of the
great mountain chain (Caucasus, Elburz, Hin-
dukush, Himalaya). The many large peninsu-
las facilitated formation of peripheral iso-
lates, of which Mayr (1963: 513) says: “
most peripheral isolates do not evolve into
new species, but when a new species
evolves, it is almost invariably from a periph-
eral isolate.'' This is likely to be the case in
S Asiatic Gyraulus. Secondary contact of pe-
ripheral isolates with the parental popula-
tions proved that reproductive isolation had
not been completely achieved and thus we
see the races of G. chinensis. In a few cases
isolation was complete enough, and sibling
species of G. chinensis (G. euphraticus, G.
tokyoensis), or morphologically more dissim-
ilar species evolved (G. malayensis and the
diverse species of Sumatra and New Guinea)
(Rensch, 1934; van Benthem-Jutting, 1963).
In summary, the more important apomor-
phous characters in the Planorbis-tribe may
be categorized as follows: (1) Characters of
high frequency in the tribe (but absence or
rarity in other tribes of Planorbidae and relat-
ed families), e.g. the hollow penial stylet; the
thickened penis tip; the club shaped penis
sheath. (2) Characters of low frequency in the
tribe as well as in the family and order, or of
low frequency in the tribe and scattered oc-
currence in the higher taxa indicating reversal
of evolutionary change, e.g. the undulate kid-
ney margins, loosely and irregularly arranged
prostate diverticula, the penis pore being re-
mote from the penis tip, the extremely wid-
ened vas deferens, the patchy mantle pig-
mentation. (3) Quantitative characters that lie
beyond the usual range of variation, e.g. an
extremely low or high number of prostate di-
verticula, an extremely inflated bursa copu-
latrix, a constant and distinct reticulate sculp-
ture in the adult shell.
Characters listed under (1) serve to define
relationships between the subgroups of the
Planorbis-tribe. Characters listed under (2)
mainly contribute to defining subgroups of the
genus Gyraulus, whereas the characters of
category (3) predominantly (though not ex-
clusively) are autapomorphies? of species.
The nearly complete restriction of the cat-
egory (3) characters to autapomorphies in this
study is fortuitous. Each apomorphous fea-
ture, when appearing for the first time, is aut-
apomorphous in one species (Hennig, 1966:
90). When it is stable enough, as required,
this feature becomes the synapomorphy of
* After completion of the manuscript | found the patchy pattern in G. essingtonensis (Smith) from the Kimberley region,
West Australia (J. Walker legit 1979).
° An autapomorphy is a derived character state confined to one individual taxon. A synapomorphy is a derived character
connecting two or more taxa.
GYRAULUS TAXONOMY (PLANORBIDAE) 83
all succeeding species. The extremely low-
ered number of prostate diverticula in the two
(or more?) species of the genus Caillaudia is
one example. Even genera can be estab-
lished from quantitative characters alone, as
will be substantiated in the next chapter. In
any case the point is: what is the probability
of having the character because the common
stem species had it? The probability should
be high.
Cladograms
In his “scheme of argumentation of phylo-
genetic systematics’ Hennig (1966, fig. 22;
1969, fig. 2) establishes a phylogeny by
working backwards from the synapomor-
phous character states seen in recent taxa
to the character states in stem taxa. The pro-
cedure consists of three steps. (1) Collect in-
formation on all characters that occur in more
than one state in a group. (2) Decide what
state is plesiomorphous and what is apomor-
phous. (3) Arrange species (or higher taxa)
so that every two or more species lie togeth-
er if they share apomorphous character states
(Hennig, 1966, fig. 22). A result of proceeding
this way is a picture of the sequence of
changes of character states (‘transformation
of characters” in Hennig, 1966). In the clado-
grams (Figs. 106, 107, 111) squares indicate
pairs of character states in the positions re-
sulting from Hennigian arguments. White
squares indicate the plesiomorphous state,
black squares the apomorphous character
state. All Recent species branching off one
trunk carry the total number of character
states (squares) indicated on the lines lead-
ing to them in the cladograms. In some cases
where the direction of character transforma-
tion is not clear, squares are shaded. If two
characters are uncorrelated, transformations
of states are depicted by two pairs of
squares, e.g. Fig. 106, transformation of plu-
ricuspid to unicuspid radula teeth and of uni-
serial to multiserial prostate diverticula. When
two apomorphous character states are prob-
ably linked, there is only one pair of squares,
e.g. Fig. 107, transformation of thickened to
conical penis tip and of the standard stylet to
hob-nail stylet during evolution of the stem
species of Armiger and Lamorbis.
It is evident that transformations of char-
acter states could only have happened in a
sequence as shown in Figs. 106, 107, 111.
One exception is made, ¡.e. when multiple new
character states appear in the same stem
species it is impossible to tell in what se-
quence. But finding the sequence of trans-
formation events between two branching
points is irrelevant for relationship research.
The sequence of transformations deter-
mined in a branch does not necessarily indi-
cate absolute points of time that can be com-
pared with those in a neighbouring branch.
This can only be found in palaeontological
studies. Hence the ordinate in Figs. 106, 107,
111, and 112 does not provide absolute mea-
sures; it indicates only the relative sequence
within each single branch.
Hennig's method of phylogenetic reason-
ing does not involve measures of divergence
or overall similarity. The positioning of taxa
in a cladogram has no meaning as regards
closeness of relationship. Each two branches
originating from the same stem species can
be exchanged, and the direction of arrows in
the cladogram is accordingly random. In or-
der not to express degrees of similarity, Re-
cent taxa are shown at equal distances.
Subgroups in the Planorbis Tribe
and in the Genus Gyraulus
To determine if Gyraulus is monophyletic
(sensu Hennig) one must study its closest rel-
atives. The Planorbis-tribe is unquestionably
a monophyletic taxon. The synapomorphies
of its species are the lack of the three pallial
ridges present in the other Planorbidae, short
marginal radula teeth, and the presence of a
separate prostate duct.
The hollow penial stylet is a distinct syn-
apomorphy characterizing the majority of tribe
members (Fig. 107). Other synapomorphies,
not shown in the cladogram, due to lack of
space, are the club-shaped penis sheath and
the thickened penis tip. There is a limited
number of species living in Europe and west
Asia that share a character state belonging
in the third category of apomorphies listed
above i.e. extreme elongation of the body.
This is not unique in the family. There are
species similar in external appearance in
South America, but remotely related to. the
Planorbis-tribe, namely Drepanotrema
species. Besides some Drepanotrema species
and those of Anisus and Bathyomphalus
there are, as far as | know, no other planor-
bid groups with an extremely elongated body.
Although convergent changes to an elon-
gated body in species from South America
and Eurasia is proved, there is no evidence
of a selective value for this character state
84 MEIER-BROOK
o|w
FIG. 109. ‘‘Flatness indices” in Gyraulus (Gy) and
related genera, Anisus (An = 2x vortex, 1 X spi-
rorbis, 2 x leucostomus) and Bathyomphalus con-
tortus (Ba).
that could explain the convergence. If elon-
gation and slenderness had a selective value,
convolution of such an animal would, in my
opinion, invalidate any advantage. On the ba-
sis of similarity of relative body length | con-
clude that this transformation happened only
once in the Planorbis-tribe. In the next clade
one daughter species retained its roundish
body cross section while the other became
flattened laterally, yielding an unusually thick
snail with a relatively low diameter, namely
Bathyomphalus (Fig. 109, Ba). The daughter
species with the plesiomorphous body form
then divided once more resulting in the stem
species of the two (three?) Recent species of
the subgenus Anisus s. str. and that of the
species Anisus vortex and A. vorticulus (sg.
Disculifer C. Boettger). In the latter subgenus
whorls are flattened as well but perpendicular
to the direction seen in Bathyomphalus. Re-
ferring to the “flatness-index,” all of the four
(five?) Anisus species lie beyond the range of
variation in Gyraulus (Fig. 109, An).
The width of the last whorl as a percent-
age of the total diameter is a good parameter
to illustrate differences. Fig. 110 shows a
broad gap between Bathyomphalus plus An-
al
number of species
15 20 25 30 35 40
width of last whorl as %o of total diameter
FIG. 110. Relations between whorl width and snail
diameter in the stylet-bearing genera of the Pla-
norbis-tribe.
isus and Gyraulus. Class frequencies are dis-
tributed not only bimodally but also without
contact or overlapping. The genus Planorbis
is not inserted in the figure to avoid confu-
sion. Planorbis lies well within the range of
Gyraulus species with 27% (Planorbis plan-
orbis), 28% (P. intermixtus) and 33% (P. ca-
rinatus).
| see no objection to conceiving the whole
group of stylet-bearers as one genus, Ani-
sus, as has been done by some authors. The
group is clearly a monophyletic taxon. On the
other hand, the gap (Fig. 110) separating Gy-
raulus and Anisus/Bathyomphalus justifies
acceptance, at least of Gyraulus, as a sepa-
rate genus (Mayr, 1969: 233). If Gyraulus has
monophyletic status | prefer to leave it at the
rank of a genus as most authors have done.
A practical reason is the following. Indication
of “Gyraulus sp.” in publications provides
much more information than ‘‘Anisus sp.’ or
even ‘‘Planorbis sp.”
Finding an apomorphy that is shared by all
Gyraulus species but not by Anisus and
Bathyomphalus would help to clearly state
the monophyletic character of Gyraulus. Such
an apomorphy would prove that Anisus and
Bathyomphalus must have branched off the
common trunk before the stem species of
Gyraulus existed. One character state that
should be synapomorphous to all Gyraulus
species has not entered the cladogram (Fig.
107), because | am not sure enough about
its reliability, but it should at least be consid-
ered. Two of the three Planorbis species have
an angle or keel that is quite close to the up-
per side of the shell. This side is more or less
plane, the whorls being flattened. The Anisus
species have this flattened upper side as well;
GYRAULUS TAXONOMY (PLANORBIDAE) 85
moreover they have an angle or keel at ex-
actly the same place as Planorbis (except A.
vorticulus that is so extremely flattened that
the angle appears to be in the middle). Thus
the asymmetric shell seems to be plesiomor-
phous. Whenever a Gyraulus forms an angle
or keel, this is situated symmetrically in the
middle of the shell. Even in multicarinate
species of old lakes, where additional angles
or keels may appear near the under and up-
per side, their position is always different from
that in Planorbis and Anisus, because the
upper side is not plane, but marginally con-
vex and centrally deeply concave (Figs. 99,
102). If one were able to accept this central
angulation or carination or at least a tenden-
cy to form it as an apomorphy, there would
be no question that Gyraulus is a monophy-
letic group in Hennig's sense, even when An-
isus and Bathyomphalus are regarded as
separate genera.
Summarizing the above discussions, ac-
cording to the concept of so-called evolution-
ists (a higher taxon is an aggregate of related
species separated from others by a discon-
tinuity; Mayr, 1969: 88) there would be no
doubt that Gyraulus, Anisus and Bathyom-
phalus are well separated genera. The strict
monophyly concept of Hennig postulates
synapomorphies uniting taxa to monophylet-
ic higher taxa: ‘The possession of plesio-
morphous characters (symplesiomorphy)
does not justify the conclusion that the bear-
ers of these characters form a monophyletic
group” (Hennig, 1966: 90). The proof could
perhaps be the central angulation or carina-
tion or a tendency towards this formation, if
this can be accepted as a synapomorphy of
Gyraulus species. If this character is not ac-
cepted as a synapomorphy (because there
are Gyraulus groups constantly lacking an
angled shell periphery), this would of course
not be evidence against the monophyletic
status of Gyraulus. A lack of synapomor-
phies merely means that proof of monophyly
cannot be furnished.
Proceeding on the assumption that Gyrau-
lus is in fact a monophyletic group, the fur-
ther progress of evolution in most lines ap-
pears to be conclusive on the basis of
synapomorphies discussed above (Fig. 107).
Several of the probable evolutionary events
have been presented in the remarks follow-
ing descriptions of individual species. One
problematic group, however, is left, i.e. the
group of Gyraulus species placed in the pro-
visional subgenus Gyraulus s. str. Again, the
sg. Torquis sg. Lamorbis sg. Armiger
Г Г
parvus laevis rossm riparius crista
) o o } e
N ii \ tiny, Sngul-
a<—oO 0—u
4 vas deferen d penis sheath.
Sr ni
O O
\ ape
A a
O— +в
— 8
hs e |
Sn
O
FIG. 111. The probable kinship relations in the
Gyraulus species bearing undulate kidney margins.
monophyly of this group cannot be proved.
Its pathway is marked by nothing but plesio-
morphous character states. Following this
pathway it is evident that in one lineage there
is the group acquiring an undulate kidney
(Armiger, Lamorbis, Torquis; Figs. 107, 111).
After another branching a drastic reduction
in volume and complexity of almost all repro-
ductive organs occurred, i.e. in Caillaudia. The
peculiar ancient lake species with distinct
synapomorphies (Carinogyraulus) or an aut-
apomorphy (Choanomphalodes) originated in
two other such lineages. Consequently, the
group called Gyraulus s. str. is a remnant
group. The subgenus formation, shown in Fig.
107, namely Carinogyraulus and then
Choanomphalodes and Gyraulus, could thus
also have taken place in a different se-
quence. The sequence shown in Fig. 107 is,
however, most likely from a geographical
standpoint.
The ancestral stem species of Recent Gy-
raulus spread over wide parts of Laurasia af-
ter the stylet-bearing species diverged into
two sister lineages, one producing elongated
animals, the other keeping its normal body
proportions (or perhaps producing the apo-
morphy of a central angle or keel). In the west
part of Laurasia an organism evolved undu-
86 MEIER-BROOK
lating kidney margins, i.e. the stem species
of Recent Armiger, Lamorbis, and Torquis. In
the marginal southern zone of Laurasia a
species branched off forming the stem
species of Caillaudia with small and simple
reproductive organs. In or near the present
Ohrid basin a lineage appeared with thick-
shelled and multicarinate snails reducing their
radula cusps, i.e. the stem species of Recent
Carinogyraulus. In Japan, finally, the stem
species of Choanomphalodes branched off.
What remained was Gyraulus s. str. retaining
the plesiomorphous character states corre-
sponding to the apomorphies mentioned
above.
When Hennig’s principle is applied the
problem arises again and again that some
species remain that carry only autapomor-
phies and are connected only by symplesio-
morphies. ‘That a common stem form is
shared by a group of species (a condition for
a ‘‘топорпуейс group” ...) can be proved
only by means of synapomorphous charac-
ters, not with symplesiomorphous charac-
ters” (Hennig, 1966: 90). Gyraulus s. str.
therefore is considered here only a provision-
al subgenus. There is one reason why Gyrau-
lus s. str. has probably to be considered even
a "paraphyletic group. The paraphyletic
groups ‘‘have no ancestor in common only to
them, and thus also no point of origin in time
common only to them in the true historical
course of phylogeny” (Hennig, 1966: 146-
147). Gyraulus s. str. could be monophyletic
(sensu Hennig) in two cases, namely (1) if
Carinogyraulus and Choanomphalodes had
a synapomorphy and, together, formed the
sister group of the remaining Gyraulus
species, or (2) if both Carinogyraulus and
Choanomphalodes had split off from one and
the same ancestor species. Concerning case
(1), the derived character states in Carino-
gyraulus and Choanomphalodes turned out
to have formed by convergence very proba-
bly. They are not sister groups. Concerning
case (2), regarding the long distance from
Europe to Japan it is probable that the
species migrating eastward split into several
lineages. The stem species of Choanom-
phalodes would then be not the same as the
stem species of Carinogyraulus. These are
the problems as regards the monophyletic
status of a subgenus Gyraulus s. str. in a
cladistic analysis according to Hennig. In
contrast, the ‘‘evolutionists’’ and the numer-
ical taxonomists accept paraphyletic groups
(sensu Hennig) as monophyletic groups in
their sense. Basing their taxa on morpholog-
ical similarity and separating their taxa on
morphological divergence they would not face
troubles in conferring monophyletic status to
a subgenus Gyraulus s. str. as characterized
in my study.
Within my provisional subgenus Gyraulus
s. str. a monophyletic subgroup can be dis-
tinguished, namely the group of species with
the synapomorphy ‘‘patchy mantle pigmen-
tation’ (Fig. 112, the seven species from the
right). The two species carrying the plesio-
morphous character state ‘diffuse mantle
pigmentation,” however, form the remnant
group in this case, i.e. G. albus and the Near
East species G. piscinarum. (A species with
diffuse mantle pigmentation was, by the way,
collected by Dr. H. Schütt in Turkey recently.
It will have to be described as a new species
showing that the remnant group consists of
more than G. albus and G. piscinarum.)
Should this group prove to be monophyletic
it would have to bear the subgenus name
Gyraulus s. str., whereas the group with the
synapomorphy “patchy mantle pigmenta-
tion’’ would have to be in a new subgenus.
For the time being, | prefer to wait for oppor-
tunities to examine more material from
southwest, south and east Asia, hoping es-
pecially that samples from the Near East and
North America will yield further insights. Only
then will it be time to draw further conclu-
sions about the remnant group. Finally the
question must be asked: why not leave Ar-
miger in genus rank? Mayr's (1974b) call that
divergent evolution be considered in the re-
construction of dendrograms would favor a
genus Armiger. Hennig replying to Mayr em-
phasized what he had explained in detail in
previous works (among others Hennig, 1950:
308, and 1969: 17-20): “In reality there is a
categorial difference between the monophy-
letic groups ..., possessing a stem species
common only to them, and the non-mono-
phyletic (paraphyletic and polyphyletic)
groupings whose components also possess
a stem species common to them but not only
to them” (Hennig, 1974: 284, translated).
Consequently raising Armiger to generic rank,
although it has synapomorphous characters
in common with the Gyraulus subgenera Tor-
quis and Lamorbis, would mean that the ge-
nus Gyraulus would cease to be a monophy-
letic taxon in Hennig's sense. It would mean
that Gyraulus in this case becomes a para-
phyletic taxon. The distinction between para-
phyletic and polyphyletic groupings, on the
GYRAULUS TAXONOMY (PLANORBIDAE) 87
albus piscin. acron. ehrenb. euphr. malay. eugyne tokyo. chin
/
/ \
o) giant shell И
Ye с
/ B-<0
ON O
bursa copulatrix
inflated
mo
penis pore distant =
from penis tip /
Me —
widened vas Mey
flattened shell /
en
shel] height
increased A
a<0
high number of O
prost.divert.
Шо
м
7
patchy mantle pigmentat.
>B
A SS
O
2<7>12
ми
ve
albus Iscin. acron. ehrenb euphr. chın. tokyo. eugyne malay.
y
O O O 6 8 O ” ©
giant shell
2 >E N
2=->g D
O O
flatten-
ed shell
вп
O bursa copula-
N trix inflated
O=B
pigment decreased \
\ O
9 ———=g
shell height
increased YA :
B-<0 penis pore distant
from penis tip
-- —
\\ widened vas deferens
0>R
NES,
high number of
prost. divert.
B<-D
NY
B O
patchy mantle
pigmentation
DD"
a
4
FIG. 112. Two of the several possible kinship relations in the provisional subgenus Gyraulus s. str.
88 MEIER-BROOK
other hand, is only a methodological one and
does not mean fundamental differences in
genealogical relations, as has been demon-
strated by Hennig (1969: 19; 1974: 284). As
to the former genus name Armiger, there is
no objection against its use as a subgenus.
As illustrated in the dendrogram (Fig. 107)
there is good reason to accept Armiger as
the sister subgenus of Lamorbis.
Hennig (1966: 154) advanced the demand
“that objects to which the same label is given
must be comparable in some way” against
the arbitrary treatment of ranking generally
performed in the literature. Seeing the enor-
mous difficulties in applying this principle to
groups as different as annelids and mammals
one will welcome his confinement to the de-
mand that at least “sister groups must have
the same rank” (Hennig, 1966: 159). This is
reasonable and feasible. In practice, how-
ever, when interpreting Fig. 107, problems
arise. The cladogram shows Lamorbis and
Armiger as subgenera; they are sister groups.
Together they form a group whose sister
group is named Torquis. Thus Torquis does
not rank at the same level as Armiger. It is
desirable not to suppress the independence
of the monophyletic group Torquis being ex-
pressed. Consequently Torquis could be
raised to generic rank. lts sister group would
be a genus “Armiger s.l. The two genera
“Armiger s.!.' and Torquis would perhaps
present a group intermediate between a ge-
nus and a tribe. The sister group of this ge-
nus group would be the one comprising the
groups named Caillaudia etc. through Cari-
nogyraulus. In this group there would be a
genus Caillaudia, whose sister genus would
comprise the groups named Choanompha-
lodes, ‘’Сугашиз $. str.,'” and Carinogyrau-
lus. The next lower taxon not necessarily
carrying a category name would be the group
consisting of Choanomphalodes and ''Gy-
raulus $. str.'* whose sister group is Carino-
gyraulus. И one sees a need for retaining the
subgenera Choanomphalodes and ''Gyrau-
lus $. str.,'” Carinogyraulus could no longer
stay at subgeneric rank because it does not
hold the same level as e.g. Choanomphalo-
des. It would, strictly speaking, have a lower
rank than generic and a higher one than
subgeneric. This would, however, not contra-
dict speaking of an unnamed category with a
single subgenus, which has to carry the name
Carinogyraulus. As ranking in one group is
not dependent on how often branching oc-
curred in its sister group, one must do what
has unexpressediy been done by system-
atists since long ago: (1) acknowledge a se-
ries of hierarchical intermediate categories (as
many as branchings maximally occur), which
may be named or unnamed; (2) in branches
that have not split up as frequently as their
sister branches insert an assumed additional
“stem species” at the same level where in
the sister group a branching point occurs.
When doing so the difficulty seen above is
easily solved: Armiger and Lamorbis are
subgenera, together forming a taxon higher
by one level, say “taxon of order Y.” Torquis
is a subgenus and that the only one of the
sister ‘‘taxon of order Y.' The three subgen-
era form a “taxon of order X,’’ which, togeth-
er with its sister “taxon of order X,’ forms
the genus Gyraulus. Proceeding in this way,
one avoids contradictions arising from strict
consequences of applying Hennig's quite jus-
tified rule for equal ranking of sister groups.
This is the procedure usual in handling mono-
typic taxa.
Absolute Age of Subgenera
The question as to how old the subgenera
are arises because Hennig (1950: 255-261)
had postulated that the absolute age of ori-
gin of a taxon should determine its rank. In
the meantime manifold criticism apparently led
him to abandon this concept or at least to
mitigate it: . phylogenetic systematics
must be content with a much coarser time
scale for its correlations'' (Hennig, 1966: 183).
Determination of fossil snails should, of
course, only be relied on when shells are
characteristic enough to preclude errors. As
confusion of many snail groups (Planorbis,
Afrogyrus, Promenetus, and others) in Re-
cent material demonstrates, a safe identifi-
cation of Gyraulus in fossil samples seems
to be impossible. This may explain fossil rec-
ords of “Gyraulus $. str.'' from as early as
the “Jurassic, ? Upper Cretaceous, Paleo-
cene” (Zilch, 1960: 110). At least the two
Mesozoic references are presumably erro-
neous. Subgenera that can more reliably be
determined have been indicated from the
Pleistocene (Caillaudia), Pliocene (Carinogy-
raulus), and Miocene (Armiger). Zilch (1960:
111) moreover records Torquis from the Mio-
cene, but Torquis is often misidentified (con-
fused with small Planorbis), even in Recent
samples, as shown above.
Given current information on sea-floor-
spreading (Scrutton, 1976; Thenius, 1977:
GYRAULUS TAXONOMY (PLANORBIDAE) 89
99), the disintegration of Laurasia into Pa-
laearctic and Nearctic became complete in the
Eocene. Accordingly, the ancestors of the
European and American Torquis have lived
in the Eocene at the latest. A migration, e.g.
during the Pleistocene, via the Bering-bridge
is highly improbable because there is no evi-
dence that Torquis ever lived in Central and
NE Asia.
As in the preceding case snails from other
tribes are sometimes misidentified as Pla-
norbis, because Planorbis has the plesiomor-
phous shell form that is shared by many oth-
er Planorbidae. According to Zilch (1960: 108)
the oldest record for Planorbis is from the
Upper Oligocene. Other close relatives of Gy-
raulus were recorded from Miocene (Bathy-
omphalus), Upper Miocene (subgenus Ani-
sus s. str.) and Upper Pliocene (subgenus
Disculifer). In view of the scattered nature of
palaeontological data there is, at least for the
time being, no chance of tracing the succes-
sive origin of taxa in geological time.
Speciation in Gyraulus
The probable course of speciation in the
Ohrid basin group and in the group having
the “undulate kidney margin’ was previously
discussed. The genus Lamorbis appears to
consist of only two species, Gyraulus ross-
maessleri and G. riparius. North America has
many Gyraulus species not examined by me,
but Baker (1945, pl. 15-19) figured the repro-
ductive organs of most of them, and there is
no evidence that any one of them has the
features of the ¿ copulatory organ peculiar to
Lamorbis.
The situation is apparently different in the
subgenus Torquis. There is at least one more
species in North America. Of the two sibling
species, Gyraulus parvus and G. laevis, the
North American one (G. parvus) has a wide
distribution and a wide ecological range while
the vicariant species in Europe (G. laevis) has
a limited distribution and a narrow ecological
range. A hypothetical explanation for such a
difference could be the following. The com-
mon stem species of G. parvus and G. laevis
was widely distributed in the now North
American portion of Laurasia, where it was
adapted to a wide range of habitats. A small
peripheral section of the population split off
and became reproductively isolated (either
before or after complete separation of North
America and Europe). This isolate spread over
parts of Europe and invaded a limited num-
ber of habitat types, and that either (1) be-
cause of its restricted gene pool as com-
pared with the large remaining population of
their stem species or (2) because it found the
majority of suitable ecological niches occu-
pied already, perhaps by other Gyraulus
species. Both factors may have contributed.
The phenomenon of pairs of vicariant sub-
species or species strikingly different in dis-
tribution and environmental requirements is
not rare. Hennig (1966: 59) quotes the situ-
ation in insects where cases are commonly
encountered “in which, in addition to a com-
mon widely distributed euryoekous species,
there is a second species that is very little
different, less widely distributed and stenoe-
kous.” This is exactly the situation in the pair
Gyraulus parvus and G. laevis.
Unlike the species of the subgenera Tor-
quis, Lamorbis and Armiger, most species of
Gyraulus s. str. lack distinct apomorphies. In
this group at least two species have main-
tained the diffuse mantle pigmentation, G. al-
bus (Europe, W Asia) and G. piscinarum (SW
Asia). G. albus has distinct reticulate sculp-
ture, G. piscinarum does not. It is not yet
possible to decide which of these characters
is more derived. All species of the Asiatic
continent have patchy mantle pigmentation. |
regard this to be their synapomorphy. Unfor-
tunately all other apomorphies hitherto ob-
served in the group are autapomorphies of
species. The problematic situation for the
phylogeneticist who deals with a group hav-
ing only symplesiomorphies and species-aut-
apomorphies can be well demonstrated with
this group (Fig. 112). The sequence of evo-
lutionary changes cannot be determined from
morphology, and theoretically none of the n!
(where n is the number of species with noth-
ing but symplesiomorphies and autapomor-
phies) possibilities (5040 different sequences
in our case, where n = 7) would have a higher
probability than the other ones. My assump-
tion is that a species most probably living in
Europe or W Asia once developed patchy
mantle pigmentation. This assumption in-
creases the probability that the first splitting
led to a northern branch, which did not split
any more (at least not into reproductive iso-
lates), i.e. G. acronicus, and a southern
branch. We may thus assume that G. acroni-
cus had its origin near the base of the tree of
patchily pigmented species, as depicted in Fig.
112. The remaining six species could have
arisen, theoretically, via 720 alternatives of
90 MEIER-BROOK
sequences, two of which are represented as
Fig. 112A and В.
If my assumption is correct that the stem
species of the southern branch spread to S
Asia from W to E, two main alternative modes
of further speciation are conceivable. (1) Dur-
ing dispersal in an easterly direction the
southern branch split up several times leav-
ing behind a series of species with limited dis-
tribution: Gyraulus ehrenbergi, G. euphrati-
cus, G. malayensis, etc. (as shown in Fig.
112A, or something similar). This sequence
would be consistent with the observation that
“speciation apparently always goes parallel
with a progression in space” (Hennig, 1966:
134). (2) Initially the southern branch spread
into a vast area, as seen presently, and in
the course of time peripheral isolates arose
as has been shown for many animal groups,
e.g. the tree snake Dendrophis pictus, the
sandfly Phlebotomus papatasii (Hennig, 1966:
figs. 16-17), and several groups of birds
(Mayr, 1963: 496). It is unimportant whether
the peripheral isolates are “only” subspecies
or “already” reproductively isolated commu-
nities. If the stem species of all Asiatic Gy-
raulus at first spread from W to E Asia, we
cannot even say whether an eastern isolate
arose before a western, and the sequence
may be completely random (Fig. 112B). At
any rate it is obvious that the widespread
species (the Rassenkreis or polytypic species
G. chinensis) has maintained the most plesio-
morphous characters and, thus, has di-
verged from its stem species to a lesser de-
gree than the other S Asiatic species.
Reflexion about the first alternative, i.e.
splitting of several species during easterly
dispersal, brings us again to a discussion of
the deviation rule. Hennig established it by
Stating: A species ceases to exist as a
species when it breaks up into two species
by partial ending of tokogenetic relations be-
tween its individuals” (Hennig, 1950: 102,
translated). This rule has meantimes been
heavily attacked by other taxonomists
(among others Mayr, 1974b). In 1966, Hen-
nig largely withdrew from this formulation, but
he was still speaking of the ‘‘deviation rule,
which is derived from the similarity distribu-
tion within species groups and which says
when a species splits, one of the daughter
species tends to deviate more strongly than
the other from the common stem species (or
from the common original condition)” (Hen-
nig, 1966: 207). This is self-evident, and no-
body has ever believed that two daughter
species would deviate to an exactly equal de-
gree from their stem species! But Hennig
sticks to a change of both daughter species
in any case. There is one fact that makes it
impossible to disprove Hennig's deviation rule.
Our most objective species concept, the
biospecies concept, can only be applied to
contemporary organisms, not to organisms
from different geological periods, as Hennig's
disciple, Schlee, points out: ‘‘Concerning the
species definition of the remaining population
there is an uncertainty whether it is really
identical—in a very strict sense—with the
original population of the stem species.
Species identity or unidentity, however, can-
not be proved, because a biological species
definition as a potential reproductive com-
munity is valid only for a point of time (tem-
poral cross-section) and populations of suc-
cessive times are, naturally, not open to
crossing experiments” (Schlee, 1971: 28-29,
translated).
The question whether a stem species can
continue to exist, although new species have
split off, is answered by Hennig as follows:
“Stem species, from which two or more re-
cent species have arisen (by whatever type
of speciation), do not occur in the hierarchic
system of recent species. They can be pro-
vided only by paleontology’’ (Hennig, 1966:
64). | do not concur because | consider Gy-
raulus chinensis as the continuously existing
stem species of G. ehrenbergi, G. euphrati-
cus, G. malayensis, etc. (Fig. 112). According
to Hennig’s deviation rule the stem species A
of the S Asiatic Gyraulus species would have
split at first into, let us say, G. ehrenbergi and
a different new stem species A1. In a second
cleavage the two species G. euphraticus and
A2 would have appeared. Thus, in the five or
more splitting processes the former stem
species A would have changed five or more
times. If unrestricted fertility between Gyrau-
lus from Afghanistan and Korea should prove
true, the existence of a biospecies G. chinen-
sis would be clear. This would disprove the
justification of Hennig's deviation rule, unless
one assumes that the product of the latest
splitting, A5 (or A6 or so) is the Recent
species G. chinensis. G. chinensis then would
have had to spread back from E Asia to W
Asia covering the vast area now inhabited,
which appears extremely improbable.
Would Hennig have voided the “rule” in our
case? Probably not. Rather, he would warn
us not to mix biological and genealogical sys-
tems (Hennig in Schlee, 1971: 28). In my
GYRAULUS TAXONOMY (PLANORBIDAE) 91
opinion, this is the point where dialectics be-
gin.
Schlee apparently has a more realistic view
of the situation at the species level such as
in the Gyraulus group, saying ‘Чп monophy-
letic groups containing a taxon that is char-
acterized by nothing but plesiomorphies, this
taxon can be the stem species” (Schlee,
1971: 34, translated). This obviously applies
to our G. chinensis and the group of its prob-
able descendants. And this perhaps applies
even to the relation between Gyraulus albus
or G. piscinarum and the whole genus. (That
G. albus has been designated type-species
of the genus and is probably the species with
the greatest similarity to the stem species of
the genus, is, of course, a mere coincidence!)
Briefly returning to our alternatives of evo-
lutionary processes in the S Asiatic region,
one should see that there is fundamentally
the same incompatibility of biological reality
and the consequence of Hennig's deviation
rule. A series of peripheral isolates would
have to be graphically represented in the
same way, for example like that in Fig. 112B.
It would likewise raise the question: why
should the stem species of the marginal new
species have ceased to exist?
DISTRIBUTION AND CHOROLOGICAL
ASPECTS
A combination of morphological and distri-
butional features has long been used in phy-
logenetic arguments. The geographical impli-
cations in taxonomic research climax in the
books of Rensch, particularly in his opus
founding the ‘‘Rassenkreislehre’’ (1929). Our
knowledge has recently been summarized in
Hennig's (1966: 133) sentence: ‘‘Every
species originally occupies a certain area, and
the breaking up of a species into several re-
productive communities usually, if not al-
ways, is closely related to the dispersal of the
species in space.” | will first discuss the dis-
tribution of Eurasiatic species of Gyraulus, as
far as it is known, and of the species of the
closely related genera.
The distribution maps (Fig. 113A-C) clear-
ly indicate that the genera related to Gyrau-
lus are restricted to the Palaearctic. The
species of the genera Planorbis and Anisus
primarily inhabit Europe; they only marginally
extend into the central areas of Asia. The only
species reaching as far as the Pacific coast
is Bathyomphalus contortus (Linnaeus). They
all have in common a southern limit in the
Mediterranean region, Turkey or the Black
Sea, the Caucasus or Transcaucasia, the re-
gions of the Caspian and Aral Seas and a
line roughly crossing Kazakhstan and Mon-
golia. Fossil records remote from the Recent
distribution areas have become known in
Planorbis planorbis (Linnaeus), viz. from the
Sudan and Ethiopia (Brown, 1965: 67). Pla-
norbis species not mapped here are P. pres-
bensis Sturany and P. macedonicus Sturany,
both endemic to two Macedonian ancient
lakes, and the group of P. atticus Bourguig-
nat and P. intermixtus Mousson which live in
the circummediterranean countries, east-
ward to Iran. They are probably not separate
from each other (Meier-Brook, 1976c) but well
isolated from P. planorbis, with which they
share great parts of their geographical
ranges.
As more or less all Planorbidae were once
included in the genus ‘‘Planorbis’’ there was
also a time when a great heterogeneous
group of Planorbidae was named “‘Anisus.”’
It might, therefore, appear as if | had not in-
cluded the entire group of Anisus in the maps.
Bollinger (1914) described an endemic
species, Planorbis sarasinorum, from Lake
Lindoe, Celebes, for example, which later
usually was called '*Anisus'' sarasinorum. Its
shell does not show any similarity with real
Anisus species. The few whorls increase very
rapidly, even more than usual for Gyraulus.
Its aperture is trumpet-like. A sharp angle with
a keel is situated in the middle of the periph-
ery (material studied in ZMA). Moreover, Hu-
bendick (1955) found a big gland in the pre-
putium, which has not been found in any
member of the Planorbis-tribe. Unfortunately
| cannot discern from his semi-schematic fig-
ures the nature of the stylet. As has been
shown above, formation of stylets has oc-
curred independently in several groups of the
family. It is highly improbable that * Planor-
bis’ or ‘‘Anisus’’ sarasinorum is closely re-
lated to any Anisus species. Another ques-
tion which cannot be answered is whether
““Anisus” sarasinorum could have originated
in the genus Gyraulus.
Another species recently referred to the
genus Anisus is A.” pauxillus van Benthem-
Jutting, 1963. This tiny snail from New
Guinea, which its author classified in Anisus
“only tentatively, because no soft parts are
available, indeed has less rapidly increasing
whorls than does Indonesian Gyraulus but still
MEIER-BROOK
92
nd Bathyomphalus.
FIG. 113A. World distribution of Planorbis, Anisus a
ulus.
ctic Gyra
nd Palaear
FIG. 113B. World distribution of Anisus a
GYRAULUS TAXONOMY (PLANORBIDAE) 93
D E
FIG. 113C. World distribution of Palaearctic Gyraulus.
decidedly more than in any of the European
Anisus species. | expect that anatomical ex-
amination will reveal its close relationship with
Gyraulus species. The nature of mantle pig-
mentation could supply the clue: if it shows
a distinct pattern (besides other features of
Gyraulus) it belongs in Gyraulus, because
hitherto all Gyraulus snails seen from S and
E Asia and the islands S of Asia have patchy
mantle pigmentation. All four species of Ani-
sus, on the other hand, lack this pattern.
Some other snails listed as ‘‘Anisus species”
by Baker (1945: 60) may either turn out to
have only superficial resemblance with or to
be merely synonyms of the European Anisus
species. Presumably, the only other good
species is Anisus strauchianus (Clessin).
Being confined to S Russia, it fits within the
distribution area of the genus. Fossil records,
also of extinct species, do not lie beyond the
Recent geographical limits of the genus.
We can thus state that all but one genus
belonging to the Planorbis-tribe are restricted
to the Palaearctic. The only genus that has
spread to nearly every continent is Gyraulus.
Should further studies reveal that the African
genera Afrogyrus and Afrogyrorbis are also
members of this tribe these would be further
exceptions. North America, the Oriental, the
Malay Archipelago, Australia, New Zealand
and the Pacific Islands harbour Gyraulus only.
Of the Gyraulus species living in Europe
only one has a Holarctic distribution: G. cris-
ta. The species with the second widest dis-
tribution is G. acronicus. lt extends farthest
to the N and displays a typical boreo-alpine
disjunction, as is known for many organisms.
Representatives of limnic animals with a sim-
ilar distribution (accord. to Thienemann, 1950)
are the water bug Arctocorisa carinata Sahlb.,
the chironomid midges Paratrichocladius al-
picola Zett., Pseudodiamesa nivosa
(Goethgh.), the copepod Heterocope borealis
(S. Fischer), and several turbellarians. The
boreal portion of the range of G. acronicus
extends farther southward than that of the
examples mentioned. It comprises the cir-
cum-Baltic countries as far as the Baltic ter-
minal moraines reach. This area is shared by
the cladoceran Holopedium gibberum Zada.,
and two plant species typically associated
with it: Lobelia dortmanna L. and /soetes la-
custris L. (Thienemann, 1950: 161). The only
difference is that these three organisms ex-
clusively inhabit lakes poor in lime, whereas
G. acronicus requires a higher calcium con-
centration. Thienemann joins Ekman in ap-
plying the term glacial ‘‘pseudo-relics’’ to
these species, previously often labelled ‘‘gla-
cial relics.’’ Since “glacial relics' should be
restricted to species whose present distribu-
tion coincides with that during glaciation, Ek-
94 MEIER-BROOK
man introduced his term, defining it as fol-
lows (according to, and slightly modified, by
Thienemann, 1950: 194, translated): ‘‘Pseu-
do-relics are faunal elements that are not rel-
ics in a certain area but have immigrated to
it in the past when natural conditions, pres-
ently no longer existing, allowed access to
the area.” As to fresh-water molluscs this is
true not only for G. acronicus, but also for
some pill clams: Pisidium lilljeborgii Clessin,
P. conventus Clessin, and probably also P.
hibernicum Westerlund.
The two species of the subgenus Lamor-
bis, Gyraulus rossmaessleri and G. riparius,
are not only stenoekous (or rather stenotop-
ic, since the determining ecological factors are
unknown) but also have a very limited distri-
bution. G. laevis may be called central Euro-
pean. G. crista is apparently the only species
of the Planorbis-tribe with a Holarctic distri-
bution (Fig. 113C). There is, however, a
strange difference in distributional character
in the continents. G. crista is almost ubiqui-
tous in the Palaearctic. In central Europe it is
encountered in almost every water body (if
not extremely soft); in constancy and abun-
dance it is outdone at most by the lymnaeid
snails Radix peregra and R. ovata, and per-
haps also by Anisus vortex. In the Nearctic it
is extremely scattered and rare. Clarke (1973:
406-409) surveying the fresh-water mollusc
fauna of the Canadian Interior Basin found it
in only two localities (of a total of 577 sta-
tions). He states that the species is “rare and
local in distribution’ in his investigation area
“and probably in North America as a whole.”
He quotes La Rocque who lists 18 eastern
North American localities from which this
species has been recorded, but 15 of these
are from Pleistocene sediments and only 3
represent living specimens. From this he
concludes that ‘‘the relative abundance of
Armiger crista in North America appears to
have declined substantially since the Pleis-
tocene.' In eastern North America G. crista
is particularly sporadic. It does not extend into
the milder and warm climates, whereas the
same species in the Old World inhabits cir-
cum-Mediterranean regions as far S as NW
Africa. Its northern distribution, low abun-
dance and low constancy in North America,
together with its continuous distribution over
N Asia, the Chukotskiy Peninsula, Alaska,
and Canada strongly suggest that G. crista
has reached the Nearctic continent via NE
Asia. G. crista is certainly one of the fresh-
water molluscs with the highest vagility. Its
extremely small size, its costae and spines
and its quasi-omnipresence in water vegeta-
tion render it particularly suited for passive
dispersal, e.g. by birds, to whose feathers it
probably adheres easily. Since no older fossil
records than from the Pleistocene seem to
be known in North America (according to
Clarke, 1973: 406) it may well be possible
that the species was introduced to the North
American continent after the connection
(Bering-bridge) in the last glacial period in the
Pleistocene had been formed that permitted
considerable exchange of animals and plants
(Thenius, 1977: 22). If this hypothesis is true,
two (alternative or, more probably, joint) fac-
tors may determine the limited distribution and
ecological range of G. crista in North Ameri-
ca. (1) Only those individuals of the Palaearc-
tic populations that were sufficiently adapted
to a cold climate crossed the periglacial en-
vironment (tundra, steppe) of the Bering-
bridge. Hence this would mean a restricted
gene pool of the American population, but not
by chance (as responsible for the founder
principle) but by selection. (2) Similarly, as
hypothetically assumed for the European
member of Torquis (G. laevis), the newcom-
er, G. crista, found most ecological niches
occupied and could not settle as well as it
might have done earlier.
The actual distribution of the SW, S and E
Asiatic species is far too little known to pres-
ent maps for them. | merely wish to demon-
strate the relative uniformity of one taxonom-
ically significant organ in the majority of
samples (Fig. 114). | have omitted only those
cases where several samples from neigh-
bouring localities were checked, e.g. from
Korea and Okinawa. All animals in these
samples had the patchy mantle pigmentation
indicating their evidently common origin. The
majority of them have been included in Gy-
raulus chinensis in this study. Some marginal
populations are considered separate species
although the form of the penis sheath, penis
tip, vas deferens and the position of the pe-
nis pore are virtually indistinguishable. These
species, regarded as having originated as pe-
ripheral isolates (Fig. 115), are G. euphrati-
cus (Fig. 116A; 115, 2), a species probably
endemic to the Seychelles (nomenclature not
as yet clear) (Fig. 114, D; Fig. 115, 3), G. to-
kyoensis (Fig. 115, 5; not drawn in Fig. 114,
due to lack of space), G. brongersmai (Fig.
114, N; Fig. 115, 8); G. tondanensis (Fig. 116,
Q; the penis was not recognizable, due to
poor preservation, but the vas deferens and
GYRAULUS TAXONOMY (PLANORBIDAE) 95
the penis sheath suggest conformity with the
species mentioned so far; Fig. 115, 7). G. eu-
gyne (Fig. 115, 4) is probably also a periph-
eral isolate of the G. chinensis-Rassenkreis;
but one should take into account that G. ac-
ronicus in this region probably almost reach-
es the range of chinensis (cf. Agöcsi & Pinter,
1971); it could, thus, also be that G. eugyne
is a peripheral isolate of G. acronicus; this
should be examined in future studies.
The species examined from Mauritius un-
der the name G. mauritianus shown to be
identical with G. chinensis is obviously a re-
cent introduction to the island through hu-
man activities. According to Starmühlner
(personal communication) the fauna and flora
of Mauritius is rich in anthropochorous ele-
ments from India. Brown (personal commu-
nication) claimed that Mauritius has in fact an
endemic Gyraulus species, which has to car-
ry the name G. mauritianus (Morelet) and
which is said to be identical with the species
of the Seychelles. | have not followed this
question further; but in any case the Gyrau-
lus inhabiting Mauritius and the Seychelles are
distinctly different from any African Gyraulus,
while anatomically it is very similar to the S
Asiatic group. This suggests a close relation-
ship that agrees with other biogeographic
observations. Mani (1974: 645) lists a num-
ber of insect species endemic to the Sey-
chelles but belonging in genera indigenous to
India. Faunistic affinities to the Madagascan
region are much smaller. According to Scrut-
ton (1976) the Seychelles (unlike Mauritius)
are most probably a fragment of the conti-
nental lithosphere that was left when India
broke off and started to drift northward, about
60 to 70 millions of years ago. Up to that
time, however, India was part of Gondwa-
naland and there would be no faunal relation-
ship to taxa living in Laurasia. The present
Gyraulus species of India certainly immigrat-
ed to the subcontinent only after India's fu-
sion with the Asiatic continent. As long as
Pangaea was intact, on the other hand, up
to the Jurassic, Gyraulus certainly did not yet
exist. The question concerning where the
Seychelles received the stem species of their
endemic Gyraulus species thus far remains
unanswered.
The Gyraulus fauna of the Malay Archipel-
ago deserves special attention. Since the
pioneer work by Wallace this region has
aroused biogeographers' interest. A sharp
borderline, the Wallace line between faunal
regions, runs between Bali and Lombok, be-
tween Borneo and the Celebes. West of it
there are nearly exclusively Oriental elements
with a very low proportion of Australian-Pap-
uan elements; from Bali to Lombok there is
an increase in Australian-Papuan elements
and a decrease in Oriental elements. A less
sharp line, Lydekker's line, between Austra-
lia/New Guinea and the Lesser Sunda Is-
lands characterizes the eastern limit of Ori-
ental elements, e.g. of the flying lizard Draco
lineatus (Hennig, 1966: 136). Both lines co-
incide with the seacoast lines during periods
of glaciation in the Pleistocene associated
with a eustatic dropping of sea level. In be-
tween there are gradients in either direction.
This region, named “indoaustralisches Zwi-
schengebiet” by Rensch (1936: 251), is gen-
erally assigned the status of a different faunal
region, the Wallacea (de Lattin, 1967: 273;
Illies, 1971: 53). A line designating about equal
portions of the respective faunal elements,
Weber's line, is without interest here. It must
be stated, however, that Wallacea has a
number of genera and species of its own, in-
cluding non-marine mollusks (Bollinger, 1914;
Rensch, 1936).
While in Wallacea elements of Indo-Malay-
an and of Australian origin used to meet and
overlap in many groups of organisms, Aus-
tralia cannot be regarded as a source of
Planorbis-tribe members. Consequently,
spreading of Gyraulus in the Malayan Archi-
pelago should have taken place mainly in
easterly and southerly directions. The Aus-
tralian Gyraulus fauna could have originated
from only two continents: Asia and Africa.
From South America, S of Venezuela, there
are known neither Recent nor fossil Gyraulus
species. One mode of extension of S Asian
Gyraulus obviously was via Malacca to the
Sreater Sunda Islands. At present only four
samples have been examined, one of which
being G. chinensis from Malacca (Fig. 114,
H), the remaining three belong to species with
the aberrant characters in the 4 copulatory
organ, Я. malayensis (Fig. 114, В, $, T). It is
important to learn whether G. chinensis itself
or a close relative with the corresponding
plesiomorphous characters entered the
Greater Sunda Islands and Bali at all. If not,
the other Gyraulus species endemic to Su-
matra (G. proclivis von Martens, G. suma-
tranus von Martens, G. feuerborni Rensch)
and to Java (G. terraesacrae Rensch) prob-
ably share the striking features of G. malay-
ensis. Spread of G. malayensis is likely to
have ended at Wallace's line (distance be-
96 MEIER-BROOK
e 3
EN
So
GYRAULUS TAXONOMY (PLANORBIDAE) 97
. ?
A
FIG. 115. A tentative map of the approximate distribution of the southern branch of Gyraulus species
with a patchy mantle pigmentation. White encircled area: G. chinensis. Shaded areas: peripheral isolates.
1 = G. ehrenbergi; 2 = G. euphraticus; 3 = a species endemic to the Seychelles; 4 = G. eugyne п. sp.;
5 = G. tokyoensis; 6 = G. malayensis n. sp. plus endemic spp. of Sumatra and Java; 7 = G. tondanensis;
8 = G. brongersmai plus endemic spp. of New Guinea.
tween Bali and Lombok scarcely 30 km, but
depth of sea 300 m). Australia has evidently
not received its Gyraulus fauna this way. The
copulatory organ in the two Australian
species studied is of the normal, ¡.e. plesio-
morphous, form. The second route by which
Gyraulus could have arrived at the Australian
continent is the one via New Guinea. Firstly,
G. chinensis itself has been encountered here
(Fig. 114, P) (van Benthem-Jutting, 1963:
494), and secondly | note the similarity in es-
sential anatomical features between (1) G.
chinensis, (2) a species endemic to New
Guinea, and (3) one of the Australian species
examined. The New Guinean species in
question is G. brongersmai van Benthem-
Jutting (Fig. 115, N); the Australian species |
wish to provisionally call G. sp. 2 (loc.: Bom-
bala, New South Wales). These two share a
E
rather unusual shell form, recalling at first
glance a Segmentina or Hippeutis in outline
and even in surface character. | expect that
study of more material will reveal that G.
brongersmai and the Australian G. sp. 2 are
sister species. Material of the other species
described as endemic to New Guinea was not
available for dissection.
How Gyraulus chinensis reached New
Guinea from the Asian continent is a ques-
tion not easily answered. G. chinensis is
present in the Philippines (Fig. 114, O). G.
tondanensis from North Celebes (Fig. 114, Q)
has the general shape of the 4 copulatory or-
gan of G. chinensis as well. Moreover, Bollin-
ger reported Planorbis compressus Hutton
from the Celebes, which is probably a syn-
onym of G. chinensis. Pleistocene land
bridges between New Guinea and the Philip-
FIG. 114. Shapes of penis sheath and vas deferens and penis plus positions of penis pore in Gyraulus
samples from $ and E Asia. A—G. euphraticus; В, С, E through М, O, P—G. chinensis; D—G. sp.,
probably endemic to the Seychelles; N—G. brongersmai van Benthem-Jutting, 1963 (paratype); Q—G.
tondanensis (Quoy 8 Gaimard, 1834); R, S, T—G. malayensis n. sp. Equal magnification in all figures.
Localities: A—S Iran; B—Sayedabad; C— Mauritius; D—Seychelles; E—Bharatpur; F—Bangalore; G—
Thailand; H—Malacca; I—Hong Kong; K—Kaejong; L—Motobu-cho; M— Taiwan; N—L. Tage, West
ап; O—Mindanao; P—Merauke, West Irian; Q—Tondano, Sulawesi; R—Kuala Lumpur; S—Bukateja,
Java; T—Tasik Madu, Bali.
98 MEIER-BROOK
NW
: N
PET HA eh ZA.
À PAZ Torquis ©
, ae AA
On!
ААА
# А
= ‘Caillaudia
FIG. 116. World distribution of Gyraulus subgenera.
pines cannot have existed considering the
great depth of the sea between the islands.
Moreover, the period since the Pleistocene
seems to be rather short in view of the ex-
tensive speciation that has evidently taken
place in New Guinea. Hence it follows that
passive dispersal, probably by birds, remains
the only explanation for the existence of Gy-
raulus in New Guinea.
Particularly close biogeographic relations
between New Guinea and Australia on the
one hand, and a very low affinity between the
Australian fauna and that of the Greater Sun-
da Islands on the other have been well estab-
lished in various groups of plants and ani-
mals (Keast, 1959; McMichael & Iredale,
1959: 241). My preliminary data presented so
far as a by-product seem to be in line with
these statements.
The second possible origin of Australian
Gyraulus to be discussed is that from Africa.
A prerequisite of this assumption would be
(1) that transfer from Africa at least to the
transantarctic continent happened before the
Upper Jurassic/Lower Cretaceous, and (2)
that snails were able to reach Gondwanaland
from Laurasia despite their separation by the
Tethys Sea. The probability of both is ex-
tremely low.
Surprisingly, however, | found a great re-
duction of portions of the reproductive sys-
tem (e.g. two prostate diverticula!) recalling
the apomorphous characters of the African
species (Subgenus Caillaudia) in one of the
Australian species examined. It is the species,
provisionally named G. sp. 1, from Benam-
bra, E Victoria, mentioned above. Although
shell characters are quite dissimilar, future
students should pay particularly thorough at-
tention to further similarities between this
species and the African ones before the con-
clusion is drawn that similarities are merely a
result of convergent evolution.
Finally, where were the centre of origin and
the centre of differentiation? The following
hypothesis is given. As the genera branching
off near the base of the tribe (Fig. 107) are
exclusively Palaearctic in distribution far be-
yond the present limits (except Planorbis
planorbis in Africa), there can be no doubt
that the stem species of all recent Gyraulus
species lived in the Palaearctic region (Fig.
116). The stem species of the subgenera Ar-
miger, Lamorbis, and Torquis split off, split
once more, and the stem species of Torquis
spread westward, joined by the plesiomor-
phous remnant group (with ‘‘straight kidney
margins''), far beyond the limits of the pres-
ent Palaearctic. In the Nearctic part of the
then Laurasian continent speciation began,
both in ‘‘Gyraulus $. str.‘ and in Torquis. The
European Torquis species, G. laevis, spread
GYRAULUS TAXONOMY (PLANORBIDAE) 99
in the counter-direction settling in the Pa-
laearctic. Lamorbis remained limited to Eu-
rope; Armiger primarily inhabited the Pa-
laearctic only and, if the above reflexions are
correct, reached the Nearctic only during the
Pleistocene.
A southern portion of the remnant plesio-
morphous species, having reached Africa,
was isolated by the formation of the Sahara
desert. The remaining species having spread
over the whole Palaearctic continent gave rise
to a peculiar group in the Balkan region, and,
independently, to that of another taxon in Ja-
pan. When, by the end of the Pleistocene,
glaciations began, the stem species of Cari-
nogyraulus and Choanomphalodes withdrew
in the refugia of the Recent Lakes Ohrid/
Prespa and Lake Biwa respectively.
The group that remained then split off a
species developing the patchy mantle pig-
mentation. After another cleavage the two
branches spread eastward separately, N and
S of the great mountain chains. The southern
branch cleft several times resulting in a great-
er number of peripherally isolated species.
A relatively basic (=early) diversification
leading to the Recent subgenera took place
in the European/West Asiatic region (3 sub-
genera, plus a subgenus in 2 Macedonian an-
cient lakes plus a subgenus probably immi-
grated from the Nearctic plus a marginal one
in the south: Caillaudia (see Fig. 116)).
Another, though minor, centre of differentia-
tion can be located in the North American re-
gion (two subgenera plus one, probably more
recent, immigrant). The large area of S and
E Asia plus the Malayan Archipelago and
Australia harbour species of only one sub-
genus, although extensive speciation (=dif-
ferentiation) occurred.
We may, therefore, state that the centre of
origin of the genus Gyraulus was probably
the West Palaearctic. A primary centre of dif-
ferentiation was in the same area, a second-
ary one in the Nearctic, and another one in
the Indo-Malayan region.
Two of the main tasks of future research
in taxonomy of the group will, consequently,
be an elaboration of phylogenetic relations
between the Palaearctic and the -Nearctic
taxa, and a study of speciation and dispersal
in the Malayan Archipelago. Moreover, it is
hoped that further knowledge of certain char-
acters (especially the mantle pigmentation) in
North American species will help to solve the
question of a further division of the species
group provisionally united in a ‘‘subgenus
Gyraulus s. str.”
DIAGNOSES OF SUBGENERA
1. Sg. Gyraulus s. str.
Type-species: Planorbis hispidus. Drapar-
naud 1805, designated by Dall (1870) (=Pla-
norbis albus Müller, 1774).
The nominate subgenus, comprising the
vast majority of Gyraulus species, differs from
the other subgenera by the lack of charac-
ters typical for these other subgenera. The
shell is always planispiral although the last
whorl may descend towards the aperture. The
aperture is more or less oval. The periphery
is equally rounded, angled or keeled and may
carry a periostracal fringe. The shell surface
may be smooth, but in most species shows
spiral striation, resulting in reticulate sculp-
ture, usually visible at least on the upper side.
The kidney has straight margins. The pros-
tate diverticula, between 8 and 40 in number,
are regularly and closely spread in a single
row. The monophyletic status of this subge-
nus is uncertain because the character states
common to all of its members are plesiomor-
phous ones. Of the Eurasiatic species ex-
amined the following are assigned to the
nominate subgenus: G. albus, G. acronicus,
G. chinensis with diverse races, G. euphrati-
cus, G. ehrenbergi, G. piscinarum, G. tokyo-
ensis, G. eugyne п. sp., G. malayensis п. sp.
2. Sg. Torquis Dall, 1905
Genus Planorbis subgenus Gyraulus sec-
tion Torquis Dall, 1905: 83, 86.
Type-species: Planorbis parvus Say, by
Original designation.
Torquis differs from all other subgenera by
the following characters: A planispiral shell
with a nearly round aperture and the periph-
ery neither angled, keeled nor fringed. The
shell surface is always smooth, due to lack
of spiral striation. The kidney is distinctly
septate, thus appearing to have undulate
margins. The prostate diverticula are ar-
ranged irregularly and usually scattered in a
single row.
Eurasiatic species assigned here are: G.
laevis, G. parvus.
3. Sg. Lamorbis Starobogatov, 1967
Genus Choanomphalus subgenus Lamor-
bis Starobogatov, 1967: 296.
Type-species: Planorbis riparius Wester-
lund, by original designation.
Lamorbis differs from all other subgenera
by a cylindrical penis sheath which is, more-
100 MEIER-BROOK
over, distinctly shorter than the preputium, by
a tapering penis tip with a hob-nail shaped
stylet, a position of the penis pore relatively
remote from the distal end of the penis, and
a distinctly septate kidney as in Torquis.
Species assigned here: G. riparius, G. ross-
maessleri.
4. Sg. Armiger Hartmann, 1844
Genus Armiger Hartmann, 1844: 172, 219.
Type-species: Nautilus crista Linnaeus, by
original designation.
Armiger differs from all other subgenera by
its last whorl not embracing the penultimate
whorl, but rather being + loosely attached to
the upper side of the latter, the peristome thus
being continuous. Armiger has undulate kid-
ney margins, a tapering penis tip with a hob-
nail shaped tiny stylet. There is probably only
one species: G. crista.
5. Sg. Carinogyraulus Polinski, 1929
Genus Gyraulus subgenus Carinogyraulus
Polinski, 1929: 161.
Type-species: Gyraulus trapezoides Polin-
ski, by original designation.
Carinogyraulus differs from the other sub-
genera by a non-planispiral pseudodextral
shell, a spire raised or flat, the left side car-
rying a funnel-like deep umbilicus delimited by
an angle or keel. The whorls of the thick-
shelled species of the subgenus are some-
times reinforced by up to three keels. Central
and lateral teeth of the radula are unicuspid,
Or cusp numbers at least partially reduced.
Prostate diverticula are as in sg. Gyraulus s.
str. or closely spaced in more than one row,
totalling more than 40. The vas deferens is
wider than in most species of the other sub-
genera.
Species assigned here are G. stankovici
from Lake Prespa and G. crenophilus, G.
fontinalis, G. lychnidicus, G. trapezoides from
Lake Ohrid.
6. Sg. Choanomphalodes Lindholm, 1927
Genus Choanomphalus subgenus Choa-
nomphalodes Lindholm, 1927: 182.
Type-species: Choanomphalus japonicus
Preston (=G. biwaensis), by original desig-
nation.
Choanomphalodes differs from the other
subgenera by a non-planispiral, pseudodex-
tral shell, whose spire is flat to weakly raised.
The periphery is tricarinate to rounded. The
radula, vas deferens, and prostate are as in
the sg. Gyraulus s. str., but the male copu-
latory organ differs from that in all other
groups by its enormous length.
If G. amplificatus (Mori) should prove to be
identical with G. biwaensis, the subgenus is
monotypic.
7. Sg. Caillaudia Bourguignat, 1883
Genus Caillaudia Bourguignat, 1883: 99.
Type-species: Caillaudia angulata Bourg.
(=Planorbis costulatus Krauss) by original
designation.
The subgenus Caillaudia differs from all
other subgenera by reproductive organs
largely reduced in size and complexity: a tiny
ovotestis, inconspicuous seminal vesicle,
prostate diverticula reduced in length and
number (3 to 12), irregular in shape and ar-
rangement, the most distal one lying distinct-
ly remote from the bursa copulatrix. Species
assigned here: G. costulatus, G. connollyi,
both in Africa south of the Sahara (Australian
species ?).
REDESCRIPTION OF GYRAULUS
The genus name was introduced in a pub-
lication by Charpentier (1837). He lists a sub-
genus “Gyraulus Ag. (Msc.)” of the genus
Planorbis on p. 21 of his Catalogue des Mol-
lusques terrestres et fluviatiles de la Suisse.
A description or diagnosis is not added. The
first species listed thereunder is Planorbis
hispidus Drap.; аз a synonym “Pl. albus
Stud.” is mentioned. PI hispidus Drapar-
naud, 1805, as the type-species was formally
chosen by Dall in 1870.
The planorbid genus Gyraulus is concho-
logically and anatomically briefly defined by
the following characters.
Shell—The shell is planispiral or, excep-
tionally (only in some ancient lakes), pseu-
dodextral with elevated spire. The shell is
small, 2 to 10 mm in maximum diameter. In
planispiral species the mean height is % to Y,
of the maximum diameter. The shell has 3 to
5 rapidly increasing whorls. The shell is deep-
ly concave on the upper side, less concave
or nearly flat on the under side. The aperture
is roundish, ovoid or spindle-shaped at its
outer margin. The surface is smooth to dull,
with or without spiral striation. The color is
light to dark corneous.
Animal—The animal is of a light to mod-
GYRAULUS TAXONOMY (PLANORBIDAE) 101
erate grey. The mantle pigmentation is dif-
fuse to conspicuously patchy (‘‘distinct pat-
tern’). The kidney is long and narrow, with
straight or undulate margins. The ureter is
reflected. The pseudobranch is triangular to
rectangular, with one longitudinal dorsal fold.
The anus lies immediately to the right of this
fold. The jaw is composed of a number of
chitinized platelets. The radula formula is 12-
1-12 to 27-1-27; the central teeth are biscus-
pid, with 3 additional denticles (2 lateral, 1
intermediate); the lateral teeth are tricuspid
with usually 4 (2 lateral, 2 interstitial) addi-
tional denticles; the marginal teeth have up
to 12 cusps formed by the 3 cusps of the
lateral teeth and additional denticles which are
increased in size and number. Cusp numbers
are reduced (to one in central and lateral
teeth) in forms endemic to Lake Ohrid basin,
Macedonia. The alimentary tract has a more
or less developed intestinal loop which is
sometimes omitted. The ovotestis is com-
posed of up to 40 lobes, which are arranged
in 2 (rarely 3) rows. The proximal end of the
spermoviduct is shorter than that distal to the
seminal vesicle. The seminal vesicle has bul-
bous or spinous coils. The albumen gland is
elongate triangular, convex dorsally, concave
ventrally (the concavity harbouring the stom-
ach). The female tract has no externally visi-
ble distinctions into oviduct, nidamental gland
and uterus. The vagina is narrower, but often
inflated close to the $ genital pore. The bursa
copulatrix is narrow and cylindrical to inflat-
ed, the bursa duct being usually long and of
varying width. The sperm duct is narrower
than the oviduct. The prostate gland is sep-
arated from the sperm duct, with 3 to 40 di-
verticula emerging from the prostate duct. The
diverticula are densely or loosely arranged in
one row (exceptions with up to 70 or more
diverticula, arranged in several rows, occur in
species of the Lake Ohrid basin). The vas
deferens usually narrows not far from the
zone where the prostate duct branches off
the sperm duct, then slightly widening, but
still narrow towards the 4 copulatory organ.
In at least one species the vas deferens wid-
ens to approximately twice the diameter of
the proximal half. In some species the vas
deferens does not narrow, but is equally wide
down to the 4 copulatory organ; the 4 copu-
latory organ is in shape well divided into a
penis sheath and a preputium by a knob-like
thickening. The penis sheath usually has a
club-shaped proximal end, which is well set
off against the vas deferens (exceptions in
forms of the Malay Archipelago, where it is
scarcely wider than the unusually wide vas
deferens). The penis sheath is more cylindri-
cal and lacks a distinct club-like proximal
thickening, but is nevertheless well distin-
guished from the vas deferens, in two Euro-
pean species (G. riparius, G. rossmaess-
еп). The distal end of the penis sheath is
nearly closed by a muscular papilla of hemi-
spherical shape. The preputium begins with
a muscular ring called the diaphragm. The lu-
men of the preputium distal to the diaphragm
is wide; towards the middle its lumen is usu-
ally narrowed by longitudinal, introverted folds
which usually form pilasters; folds can be so
weak that the preputial lumen appears nearly
round. The penis sheath is usually of once to
twice the length of the preputium (only in the
two European species mentioned above dis-
tinctly shorter than the preputium). The penis
is as long as the penis sheath, usually with a
more or less conspicuous distal thickening
(instead of the thickening there may be a
conical tapering). The penis tip is equipped
with a chitinized hollow stylet, which is clear-
ly delimited against the penis tissue. The pe-
nis pore usually lies in varying portions of the
thickened penis tip, exceptionally near the
middle of the penis (forms of the Malay Ar-
chipelago, probably—according to Huben-
dick & Radoman, 1959—also in a species
from Lake Ohrid).
SUMMARY
1. An approach is made towards basing
the classification of the planorbid snail genus
Gyraulus on as many characters as possible.
The study is focussed on taxa inhabiting Eu-
rope and Asia with observations pertaining
to other continents.
2. Shell proportions, as expressed by a
“flatness-index” (introduced by Brown & van
Eeden) and the ratio maximum shell diame-
ter: whorl number are demonstrated in dia-
grams to show interspecific and infraspecific
(both genotypical and ecophenotypical) vari-
ation. In one species, G. albus, shells of equal
whorl numbers in lenitic biotopes are larger
in diameter than in lotic biotopes (Fig. 6).
3. Influence of the usual fixing techniques
on anatomical data is elaborated in one ex-
ample. The length of the penis sheath is
greatly affected by immersion of living Gyrau-
lus snails in 70% ethanol, as compared with
values for snails fixed after relaxation with
102 MEIER-BROOK
pentobarbital. The preputium does not ap-
pear to be significantly shortened.
4. Characters highly valued for species dis-
crimination are among others: the distribu-
tion of pigment cells on the mantle to the right
of the kidney; the presence or absence of
distinct transverse septa in the tubular por-
tion of the kidney rendering its margins "un-
dulate” or “straight,” respectively; the num-
ber of prostate diverticula, their shape and
arrangement in the gland; the width of the
vas deferens; the length ratio penis sheath :
preputium; the shape of the penis tip and
stylet, and the position of the penis pore. In
certain groups peculiarities may occur in var-
ious other organs (radula, seminal vesicle,
bursa copulatrix, bursa duct, etc.).
5. Europe outside Macedonia harbours five
indigenous species. For their identification a
key is presented which also permits recog-
nition of two species recently introduced to
Europe.
6. In Asia the number of species is much
greater. N Asia is inhabited by G. acronicus
(Fér.) only. A huge area of S Asia, from Iran
to Japan and New Guinea, harbours forms
so little different both conchologically and an-
atomically that they are probably not repro-
ductively isolated and must, thus, be regard-
ed as races of one polytypic species or
“Rassenkreis”: G. chinensis (Dunker) (syn-
onyms: G. convexiusculus (Hutton), G. spiril-
lus (Gould) and others). Species anatomically
indistinguishable, but conchologically more or
less distinctly different, are G. tokyoensis Mori
and G. euphraticus (Mousson). They are con-
sidered separate species because they are
said to be sympatric with G. chinensis, thus
indicating reproductive isolation. There are
some more species whose anatomical simi-
larity suggests close relationship with G. chi-
nensis; they are conchologically divergent
species mostly endemic to island groups
(Seychelles, Celebes, New Guinea). One
species, on the other hand, conchologically
so far indistinguishable, has quite aberrant
anatomical features giving it a unique status.
It is known from Malaya, Java and Bali and
is described as a new species: G. malayen-
sis. Another species named here is G. eu-
gyne n. sp. from Inner Mongolia.
7. Abandonment of planispiral growth and
formation of several angles or keels on the
shell periphery in Gyraulus of ancient lakes is
certainly due to convergent evolution. There
is strong evidence that a species endemic to
Lake Biwa in Japan has a history different
from that in a group endemic to the Mace-
donian Lakes Ohrid and Prespa.
8. For an analysis of phylogenetic relation-
ships, Hennig's (1950 and later) recommen-
dations are followed т a “search for the sis-
ter group” and a hierarchic system expressing
the “recency of common ancestry” of taxa.
Tools used in the analysis are common de-
rived characters (‘‘synapomorphies’’). More-
over, Hennig's concept of monophyly is fol-
lowed in this study.
9. Criteria for judging the apomorphous
character states are discussed. Highly valued
as being derived are characters of unique-
ness or low abundance in the genus or in the
family or higher taxa. Qualitative characters
meeting these requirements are relatively rare
in the genus, so that quantitative ones must
be used in addition. In some cases, these are
even used as synapomorphies of subgenera
or genera, provided they display extreme val-
ues and appear to be sufficiently stable, e.g.
the extreme elongation of the body in the
genera Anisus and Bathyomphalus.
10. The systematic position of Gyraulus in
the Planorbis-tribe is examined (Fig. 107). The
species group bearing a hollow penial stylet
(Anisus, Bathyomphalus, Gyraulus) is clearly
a monophyletic group within the Planorbis-
tribe. Since Gyraulus is likely to be a mono-
phyletic group, too, maintenance of the three
genera mentioned above is recommended
because of clear gaps in their range of vari-
ation. Anisus and Bathyomphalus together
form the sister group of Gyraulus.
11. Phylogenetic analysis in the genus Gy-
raulus reveals the existence of seven sub-
genera: Torquis Dall (Nearctic and W Pa-
laearctic), Lamorbis Starobogatov (W
Palaearctic), Armiger Hartmann (Holarctic),
Caillaudia Bourguignat (Ethiopic), Choanom-
phalodes Lindholm (Lake Biwa, Japan), Cari-
nogyraulus Polinski (Lakes Ohrid and Pres-
pa, Macedonia), and a provisional ‘‘subgenus
Gyraulus $. str.” (cosmopolitan except South
America and the Ethiopic region) (Figs. 107,
116). Gyraulus s. str. being a remnant group
merely characterized by plesiomorphous
character states is likely to be a paraphyletic
group in Hennig's sense. Diagnoses of sub-
genera are given.
12. The impossibility of deriving sequences
of evolutionary steps in groups sharing only
in plesiomorphous characters is demonstrat-
ed in the remnant group, the provisional
GYRAULUS TAXONOMY (PLANORBIDAE) 103
“subgenus Gyraulus $. str." Sequences may,
with a certain probability, be supposed from
dispersal pathways only.
13. Incompatibility is shown between Hen-
nig's ‘‘deviation rule” and biological reality in
the case of peripheral isolates. The example
advanced is the polytypic species (‘‘Rassen-
kreis”) Gyraulus chinensis, with marginal
species such as G. euphraticus, G. tokyoen-
sis, G. malayensis and others. G. chinensis
is considered to be their “persisting stem
species” although, according to this “rule,”
it ought to have changed its species status
once at each time a peripheral isolate split
off. A “persistent stem species of Recent
species” is inconsistent with the deviation
rule.
14. A limited geographical distribution and
a narrow ecological range of the only Torquis
species indigenous to Europe suggests that
the subgenus primarily inhabited the western
part of the former Laurasian continent and
that Gyraulus laevis, the sister species of the
Nearctic G. parvus, was split off from their
common stem species as a small marginal
portion of a large reproductive community.
This is similar to phenomena encountered in
insect groups where in pairs of vicariant
species or subspecies one is often much less
widely distributed and displays a much nar-
rower ecological range than the other.
15. The centre of origin of the genus Gy-
raulus is probably the W Palaearctic, to which
all its closely related genera are confined. With
three subgenera of wider distribution plus one
subgenus endemic to two ancient lakes in
Macedonia plus one marginal subgenus in the
Ethiopic plus one subgenus probably origi-
nating in the Nearctic, the W Palaearctic is at
the same time regarded as the primary centre
of differentiation. In the Nearctic there is a
secondary one: two subgenera plus one sub-
genus probably acquired not before the
Pleistocene. A third centre of differentiation,
though only on the species level in a single
subgenus, is the Indo-Malayan region (Fig.
116). Expansion probably took place radially
from the W Palaearctic to the Nearctic, Afri-
ca, N Asia, and S Asia. Australia received its
poor Gyraulus fauna evidently from South-
east Asia via New Guinea. An African off-
shoot in the Australian Gyraulus fauna,
though unlikely, cannot yet be completely ex-
cluded.
16. A redescription of the genus Gyraulus
Charpentier, 1837, is given using present
knowledge on the variation of anatomical and
conchological characters.
ACKNOWLEDGEMENTS
During these studies | had the help and
support of several private persons and insti-
tutions. Technical assistance or advice was
received from Mrs. A. Schmelzeisen (histol-
ogy, darkroom), Mrs. B. Löffel and Ms. S.
Renkhold (shell measuring, computing), Ms.
|. Hermann (graphs, darkroom), Mrs. В. Klett
(scanning electron micrographs), Doz. Dr. H.
M. Seitz (critical point drying). | am further
indebted to Dr. W. U. an der Heiden and Dr.
G. Deichsel who gave advice on mathemati-
cal handling of growth phenomena and other
data, Dr. B. Hubendick and Doz. Dr. K. E.
Lauterbach for critically reading parts of the
manuscript, Мг. D. Hilgeman, Dipl.-Biol., Mr.
D. Kelly, and Dr. G. M. Davis for help in im-
proving the English, and Mrs. Willbold for
typing the manuscript. Texts were translated
from foreign languages by Mrs. N. Шдеп
(Russian), Ms. В. Шдеп, Dipl.-Biol. (Serbo-
Croat), Mr. Iguchi, Túbingen (Japanese), and
Prof. Hudec, Praha (Czechoslovak). Dr. A.
Zilch, Frankfurt/M., was indefatigable in
sending copies of original descriptions.
Donations or loans of material were re-
ceived from the following collectors and mu-
seum curators: Мг. À. Andersson, Fil. lic.,
Naturhist. Riksmus. Stockholm; Mr. R. Bank,
Haarlem, Netherlands; Dr. E. Binder, Musée
d'Histoire Naturelle Genéve; Dr. H. D. Boe-
ters, Múnchen; Mr. A. J. Brandt, Hong Kong;
British Museum (N.H.) authorities; Dr. J. B.
Burch, Mus. of Zool., Ann Arbor, Mich.; Mr.
N. Burgemeister, Túbingen; Dr. A. H. Clarke,
formerly of National Mus. of Canada, Otta-
wa; Dr. H. E. Coomans, ZMA, Amsterdam;
Dr. С. М. Davis, Acad. Nat. Sci. Philadelphia;
Dr. E. Gittenberger, RMNH, Leiden; Dr. J. van
Goethem, IRSNB, Bruxelles; Dr. L. Hásslein,
Lauf/Pegnitz; Prof. Dr. W. Höfler, Inst. Trop.
Med. Túbingen; Dr. B. Hubendick, Naturhist.
Mus. Góteborg; Dr. J. Jungbluth, Zool. Inst.
Heidelberg; Dr. H. Jungen, Zool. Mus. Zü-
rich; Prof. Dr. R. Kinzelbach, Zool. Inst.
Mainz; Prof. Dr. H. J. Knúttgen, Inst. Trop.
Med. Tübingen; Prof. Dr. K. J. Lie, Hooper
Found., Kuala Lumpur; Mr. W. Maassen,
Duivendrecht; Dr. G. Mandahl-Barth, Dan.
Bilh. Lab., Charlottenlund; Dr. J. Massoud,
School of Pbl. Hith., Teheran; Dr. P. Mildner,
104 MEIER-BROOK
Klagenfurt; Dr. T. Miura, Otsu Hydrobiol. Sta.,
Kyoto Univ.; Mr. P. Mordan, Brit. Mus. Nat.
Hist., London; Dr. W. Rähle, Zool. Inst. Tü-
bingen; Dr. N. V. S. Rao, Zool. Survey of In-
dia, Calcutta; Dr. H. Schütt, Düsseldorf-Ben-
rath; Mr. J. B. Sigurdsson, Newcastle upon
Tyne; Dr. B. J. Smith, NMV, Melbourne; Dr.
К. Y. T. Tjhen, Ammerbuch; Мг. J. Walter,
Zool. Inst. Zürich; Dr. A. Zilch, Senckenberg-
Museum, Frankfurt/M. To all | express my
sincere gratitude.
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APPENDIX. Alphabetical List of Collecting Sites (short forms listed in the text)
Nearest
Locality place
(reg. no. of Other data about local.; collector, entered in Lat. Long.
mus. coll.) sampling date Times Atlas ME умы
Abitibi R. (NMC Canada: Ontario: A. River at Hwy. Clute 49.13N 81.00W
25150) 579, appr. 27 km N of Cochrane;
А. H. Clarke VII-1961
Alexandria Egypt: Ezbet Belal 25 km S of Alexandria 31.13N 29.55Е
Alex.; Demian 111-1966
Ann Arbor USA: Michigan. Woods-pool at Ann Arbor 42.18N 83.43W
Zeeb Rd., Te V-1974
Ansbach Germany: Bavaria: Scheerweiher, 3 Ansbach 49.18N 10.36E
km W of Ansbach; H. Stocker
|-1966
Baalbeck Lebanon (collector and date not in- Ba'albek 34.00N 36.12E
dicated)
Bad Villach see Villach
Bahr-el-Houle Syria: running and standing waters ?
near Bahr-el-Houle (Lake Sama-
chonitides); Bourguignat (no date)
Bangalore India: Bangalore; Muraleedharan, Bangalore 12.58N 7:7.35E
X11-1973 Mandahl-Barth dedit
Barracuta (L.) Australia: East Victoria; Neboiss 2
1-1975
Locality
(reg. no. of
mus. coll.)
Benambra
Bharatpur
Biwa (L.)
Bodensee
Bombala
(NMV 2305)
Bukateja
Cairo
Chongpyong
Diyarbakir
Dobersdorfer
See
Drusenheim
Dudinska
Ettenheim
Fehmarn
Fischau
Gawargin
Greifensee
Haarlem
Hanechi-son
(ANSP A288)
GYRAULUS TAXONOMY (PLANORBIDAE)
APPENDIX. Continued
Other data about local.; collector,
sampling date
Australia: Corryong Rd., 8km N of
B.; in water beside road; Dart-
mouth Survey X-1973
India: Bird sanctuary near Bh.; D.
Haas 1-1974
Japan: Lake Biwa; at Shiotsu and
Oura: Т. Miura VI-1975; at Eizan:
Н. О. Boeters 11-1975
Germany: Lake of Constance at
Sipplingen/Uberlingen; Meier-Br.
IX-1961
Australia: New S Wales: Saucy
Ck.; 10 km S of B., on Cann Val-
ley Hwy.; A.B.R.S. Snail Survey
IV-1975
S-central Java; Tjhen VI-1976
Egypt: Drain in Giza near pyramids;
Demian X-1974
Rep. of Korea: Kyong-gi-do: fish
pond; Meier-Brook VI/XI1-1970
Turkey: Vilayet Diyarbakir; Paydak
VI-1972; H. Schütt dedit
N Germany: Holstein: Probsteier-
hagen; Meier-Brook IX-1960
France: Bas Rhin; Meier-Brook
111-1961
U.S.S.R.: Siberia: Dudinka/Yenisey,
Swed. Sib. Exped. VII-1876
SW Germany: Baden; fishpond;
Meier-Brook VI-1961
see Wallnau
Austria: Therme Bad Fischau;
Meier-Br. IX-1968; Jungbluth
X-1974
Afghanistan: Prov. of Helmand
(31.26N 64.20Е), alt. ‘750 т”
(according to Times Atlas, coordi-
nates are between 200 and 500
m in alt.!); Kawata IX-1968; Man-
dahl-Barth dedit
Switzerland: E of Zürich; Meier-
Brook VII-1961
Netherlands: “Lake” in Kennemer-
duinen; В. Bank VIII-1977
Japan: Okinawa: channel and ditch
along rt. 124 betw. Maeshine and
Gabusoka; Davis III-1968
Nearest
place
entered in
Times Atlas
Corryong
Bharatpur
Kinomoto
Otsu
Überlingen
Bombala
Purbolinggo
Giza Pyram.
Kapyong
Diyarbakir
Schönberg
Drusenheim
Dudinka
Ettenheim
Bad Fischau
Zaras
Greifensee
Haarlem
Nakaoshi
Lat.
о
36.11$
27.14N
35.33N
35.00N
47.46N
36.555
7.225
29.59N
37.53N
37.55N
54.24N
48.46N
69.27N
48.16N
47.50N
31.18N
47.18N
52.23N
26.43N
109
Long.
147.58E
77.28E
136.12E
135.50Е
9.10Е
149.16Е
109: 15Е
31.07Е
127.30E
40.14E
10.23E
И ЭВЕ
86.13E
T2S2E
ls
64.13E
8.42E
4.38E
12757
Locality
(reg. no. of
mus. coll.)
Haslacher See
Hong Kong
Iceland, SE
Iceland, S
Iceland, W
Inner Mongolia
Iran, S
Kaejong
Karabella
Kasseeteich
Kolksee
Krasnojarsk
(NHRMS 404)
Kuala Lumpur
Kuehren
Kumluca
Kunsan
Kwekerslaan
Lake...
Läger XI
(NHRMS 2198)
Lesser Slave
Lake (NMC
29922)
MEIER-BROOK
APPENDIX. Continued
Other data about local.; collector,
sampling date
Germany: Bavaria: near Burggen;
Meier-Brook VIII-1959
Hong Kong: Island peak in moun-
tain stream; A. J. Brandt IV-1974
A. Skaftaffellssysla; pond on Stapa-
sandur, nr. Thveit; Einarsson VIII-
1976; dedit Sigurdsson
Arnessysla: Opnur, Stora-saurbae,
Olfus; Sigurdsson VII-1976
Snaefellsnessysla: Stadarsveit,
Kúka & Hofgardatjörn, Yovi-Gar-
dar; Sigurdsson VII-1976
See Läger XI
locality? 111-1975 Massoud dedit
Rep. of Korea: Cholla-puk-do; ditch
W of rd. Iri-Kunsan; Meier-Brook
IX-1970
U.S.S.R.: Kola Peninsula; on Ribat-
ski; Sandebergs Exped. VII-1877
Germany: Holstein: near Probsteier-
hagen; Meier-Brook 111-1960
N Germany: Holstein: forest lake
near rd. Preetz-Plön; Meier-Br.
VII-74
U.S.S.R.: Siberia; Swed. Sib.
Exped. 1876
Malaysia: 5 km from K.L., large un-
used tin-mining pool. On grass
and Eichhornia. J. К. Lie VI-1973
N Germany: Holstein: Kührener
Viehteich; Meier-Brook VIII-1974
Turkey: well NE Kumluca; Kinzel-
bach VIII-1975
Rep. of Korea: Cholla-puk-do: res-
ervoir betw. K. and Air base;
Meier-Brook VIII-1970
Netherlands: greenhouses in Kw.,
Haarlem; В. Bank VII-1977
see proper name
Р. В. China: Inner Mongolia: Hon-
nentjaggan-tschollogol, a northern
tributary of Hoang-ho; Sven Hed-
in Exped.; Hummel VIII-1927
Canada: Alberta: Slave Lake;
Athearn; VII-1965 A. H. Clarke id.
et dedit
Nearest
place
entered in
Times Atlas
Schongau
Lech
Victoria
Bjarnarnes
Kaldadarnes
Stadastadur
?
Kunsan
Kiel
Preetz
Karsnojarsk
Kuala Lumpur
Preetz
Kumluca
Kunsan
Haarlem
(?) Ch'ing-tai
(?) Slave
Lake
Lat.
o
47.49N
22.16N
64.19N
63.56N
64.49N
35.57N
54.20N
54.14N
56.05N
3.08N
54.14N
36.23N
35.57N
52.23N
41.46N
55.17N
Long.
10.54E
114.13E
15.13W
21.10W
23.00W
126.42E
10.08E
10.17E
92.46E
101.42E
1OA7E
S0'17E
126.42E
4.38E
113.29Е
114.43W
GYRAULUS TAXONOMY (PLANORBIDAE)
APPENDIX. Continued
a
Locality
(reg. no. of
mus. coll.)
Liberia
Limyra
Mainz
Malacca
Mauna-gawa
Mauritius
Merauke
(RMNH 5322)
Mindanao
(NHRMS 4166)
Motobu-cho
(ANSP A266)
Niederzell/Unter-
Ses
Ohrid (L.)
Onna-son
(ANSP A 453)
Paratunka
(NHRMS 3727)
Pleistinger Wörth
Prespa (L.)
Ragunda
(NHMG 11249)
Rasno-volok
(NHRMS 347)
Rhine River
Backwater
Other data about local.; collector,
sampling date
Centra Liberia; Hófler et Knúttgen
V-1974
Turkey: mouth of a creek 5 km E of
Limyra near Finike; Kinzelbach
VII-1975
W Germany: pond in Bot. Garden
Mainz Univ.; Kinzelbach 1973-
1974
Malaysia: 20 km inland from M.,
stream running through paddy-
field; J. В. Sigurdsson II-1977
Japan: Okinawa; branch of Mauna-
gawa, E of Teguchi-village; Davis
111-1968
Beau Bassin, canal; C. N. Courtois
V1-1973
West Irian (New Guinea): pool N of
M. in swampy area (cf. v.
Benth.-J., 1963: 494); Holthuis
111-1955
Philippines: Davao Penal Colony,
40 km N of Davao City, creek;
В. Hubendick VII-1952
Japan: Okinawa: E of Teguchi-vil-
lage Davis 111-1968
Germany: Lake of Constance: Is-
land of Reichenau (also called Un-
terzell)
Yugoslavia: Macedonia: Lake Ohrid
near Sveti Zaum; Rähle VI-1973
Japan: Okinawa: paddy field W of
Atsuta, along Rt. 1; Davis IV-
1968
U.S.S.R.: Kamchatka: hot springs
near P.; Hulten 11-1921
S Germany: Bavaria: island in
Donau; Hásslein VIII-1965
Yugoslavia: Macedonia: Lake Pres-
pa, Rähle VI-1973; Maassen
V-1976
Sweden: Jmt.: SW Stormoberget,
Ammeräns N strand; Walden et
Wanntorp VII-1961
U.S.S.R.: Kola-peninsula; turbid
marsh; Sandebergs Exped. VII-
1877
Germany: Baden: Plittersdorf;
Meier-Brook 111-1961
Nearest
place
entered in
Times Atlas
?
Finike
Mainz
Malacca
Toguchi
Rose Hill
Merauke
Davao
Tohuchi
Radolfzell
Ohrid
Atsuta
Paratunka
Vilshofen
Resen
Ragunda
i
Rastatt
Lat.
o
36.18N
50.00N
2.14N
26.40N
20.14S
8.305
7.05N
26.40N
47.44N
41.06N
26.30N
52.58N
48.38N
41.05N
63.06N
48.51N
Long.
30.08E
8.16E
102.14E
127.53E
NE
140.22E
125.38E
127.53E
8 5ЭЕ
20.49Е
127.50Е
158.14E
13.12Е
2101
8.13E
112
Locality
(reg. no. of
mus. coll.)
St. Point (Lac)
Samava
Sashiki-son
(ANSP A374)
Sayedabad
Selenter See
Seychelles
(IRSNB 1.G.
25.536)
Siggen
Sjaelland
Soufflenheim
Speyer
Stuttgart
Tärna
Tage (L.) (RMNH
alc. 4985)
Taiwan
Tasik Madu
Thailand (SMF
193798)
MEIER-BROOK
APPENDIX. Continued
Other data about local.; collector,
sampling date
France: Dep. Jura: Lac de St. Point
F. Geissert 1959
Iraq: AS Samawah/Euphrates;
Schlaefli 1873
Japan: Okinawa; ditch in Fusozaki
along Kaneku-ko Bay; Davis III-
1968
“Afghanistan” (30.49N 67.51Е);
(according to Times Atlas, 1967,
these coordinates are in West Pa-
kistan, Quetta Prov.), alt. 2500
m; Kawata VIII-1968; Mandahl-
Barth dedit
Germany: Holstein; Meier-Br.
111-1960
lle de Mahé, Riv. Seche (Route de
Misere), Sta. 40; G. Marlier
X-1976
S Germany: Allgäu: Krs. Wangen;
Neuweiher near Siggen; Meier-
Brook IV-1974
Denmark: lake 1 km N of Jorlose,
E of rd. Kalundborg-Slagelse;
Burgemeister VII-1976
France: Bas Rhin; inundated mead-
ow NE rd. S Sessenheim; Meier-
Brook V-1963
SW Germany: Autobahnsee NE
Speyer; Fischenich IIl-1974; Kin-
zelbach ded.
Germany: Württemberg: forest pool
near Stuttgart-Büsnau; Meier-
Brook V-1977
Sweden: Ly Lpm., SE Granäs; 500
m SE of Granäs; H. Walden,
Svensson, and Wanntorp VIII-
1962
West Irian (New Guinea): Lake
Tage, near missionary post Tage,
Wissel Lake region (cf. v. Benth.-
J., 1963: 497); L. D. Brongersma
X11-1954
Rice fields near Hualien-Airport; Lo
VI-1975
N Bali: Lovina-beach; Tjhen
VI-1976
Bangkok: Zoolog. Garden; R.
Brandt VI-1965
Nearest
place
entered in Lat.
Times Atlas o Y
St. Point 46.49N
As Samawah 31.18N
Baten 26.12N
?Hindubagh 30.51N
Lütjenburg 54.17N
Mahé I. 4.415
Isny 47.42N
Jerslev 55.37N
Soufflenheim 48.50N
Speyer 49.18N
Stuttgart 48.47N
2
between 3.46S
Uwapa and 3.425
Mbambawa
Hualien 23.58N
Singaradja 8.06S
Bangkok 13.44N
Long.
GAME
45.18E
127.46E
67.50E
10.36E
55.30E
10.02E
11.15Е
T.51E
8.26E
9.12E
135.31E
137.31E
121.53E
115:07E
100.30E
GYRAULUS TAXONOMY (PLANORBIDAE) 113
APPENDIX. Continued
Nearest
Locality place
(reg. no. of Other data about local.; collector, entered in Lat. Long.
mus. coll.) sampling date Times Atlas a CURE
Thailand (SMF Klong Killard, 4.7 km from Grabi to Krabi 8.04N 98.52E
197350) Khao Thong; R. Brandt X-1969
Titisee Germany: Baden: Schwarzwald; Titisee 47.55N 8.10E
Meier-Brook X-1961
Tomigusukuson Japan: Okinawa; ditch near lraha, Oroku/Naha 26.10N 127.40E
(ANSP A416) 1.6 km S Naha air base; Davis
111-1968
Tondano (ZMA) Sulawesi (Celebes): River near Ton- Tondano ION 125.56Е
dano; Siboga Expedition
Tübingen S Germany: Quarry pool near Wen- Tübingen 48.32N 9.04E
delsheim, 6 km W of T.; Meier-
Brook VI-1973
Tulom River U.S.S.R: Kola peninsula: Tulom В. Kola 68.53N 33.01E
(NHRMS 350) 10 to 20 verst from Kola; Sande-
bergs Exped. VII-1877
Untersee see Niederzell
Ursee SW Germany: bog in Schwarzwald Lenzkirch 47.52N 8.13E
between Feldberg and Lenzkirch;
Meier-Brook IV/VIII-1963
Valtjärn (NHMG Sweden: Jmt, 1300 m N of Mattmar Mattmar 63.19N 13.45E
66-11.853) K:a, creek just W of Valtjärn;
Walden 1X-1966
Vikarsjön (NHMG Sweden: Hrj, Vikarsjón S. str. v. ?
64-11490) Kvarnáns utfl.; Walden, Svensson
and Wanntorp VIII-1962
Villach Austria: Kärnten: Therme Bad Vil- Villach 46.37N 13:51
lach; Mildner VIII-1974
Vojmán (NHMG Sweden: As Lpm, northern part of ?
67-11.982) Gotajaure; Walden and Hultin
VIII-1968
Wallnau/Fehmarn Germany: Holstein: Island of Feh- Orth 54.27N 11.04E
marn; Meier-Brook 111-1960
Westensee Germany: Schleswig; J. Rentner Flemhude 54.20N 9.58E
V-1961
Windgfállweiher SW Germany: tarn in Schwarzwald Lenzkirch 47.52N 8.13E
between Feldberg a. Lenzkirch;
Meier-Brook VI/IX-1964
Wollmatingen S Germany: Wollmatinger Ried, Konstanz 47.40N 9.10E
part of Bodensee near Reich-
enau; Meier-Brook VI-1961
Zurichsee Switzerland; J. Walter IV-1974 Zurich 47.23N 8.33E
MALACOLOGIA, 1983, 24(1-2): 114-276
A REVIEW OF THE NUDIBRANCHS OF THE CALIFORNIA COAST
Gary R. McDonald'
Moss Landing Marine Laboratories of the Central California State Universities and Colleges,
P.O. Box 223, Moss Landing, CA 95039, U.S.A.
ABSTRACT
All of the named species of nudibranchs reported from the California coast are described,
including five unnamed species and one previously reported but unnamed species, a total of
101 species comprising 50 genera and 32 families. The species descriptions use the same
terminology and present information in the same order to facilitate comparison. An extensive
synonymy is given for each species, along with a detailed bibliography. A dichotomous key to
aid in identification and drawings of the radula of each species are presented. Location of type-
specimens, geographic ranges on the west coast of North America, type-locality, habitat, and
food data are also presented. Recent publications and study have resulted in a number of
generic reallocations and in changed names due to the law of priority.
INTRODUCTION
This study describes all of the valid species
of nudibranchs presently known along the
California coast. Five species are included
which have not previously been reported from
California. It is quite possible that these five
species are described from other geographic
areas; presently they are identified only to
genus until further investigation determines if
they are assignable to an already described
species. Listing them thus would seem best,
as has been done by other workers (e.g.
Marcus 8 Marcus, 1967a: 32), rather than to
name them as new species at this time.
At present no publication describes all of
the California nudibranch species. Two pub-
lications, MacFarland (1966), and Marcus
(1961) each treated many, but not nearly all
of the species for California. Unfortunately,
neither publication presents a key to aid in
identification of species.
MacFarland (1966) studied specimens col-
lected over many decades and from numer-
ous localities along the coast. The majority
of species were seen alive and color paint-
ings were made. The volume by MacFarland
(1966) is based on his extensive notes which
were organized by his wife and workers at
the California Academy of Sciences after his
death in 1951. In the 15 years between his
death and the publication of his work, six of
his new species were collected and named
by other workers. Roller (1970a) found six
other species to be junior synonyms of older
species. Since MacFarland's work was unfin-
ished at the time of his death, the coverage
of species is highly variable; some were de-
scribed in great detail, while others were only
briefly described.
The basis for the study by Marcus (1961)
was a collection of preserved specimens col-
lected over a relatively short span of time,
mostly from Dillon Beach, Marin Co., and San
Diego, California, and sent to Marcus in Bra-
zil. Observation of living specimens is most
desirable when preparing descriptions of new
species of nudibranchs. Unfortunately, Mar-
cus received only preserved material and
never saw any of the species alive. The study
is thus of limited usefulness and is deficient
in species coverage since only 37 of the total
101 California species are described.
METHODS AND MATERIALS
My personal collection forms the major ba-
sis for the present study. The collection con-
tains several thousand specimens collected
over a period of ten years. The majority of
specimens in the collection is from the central
California coast (Pescadero Point, San Ma-
' Present address: Long Marine Laboratory, Center for Coastal Marine Studies, University of California, Santa Cruz, CA
95064, U.S.A.
(114)
CALIFORNIA NUDIBRANCHS 115
teo Co., to Pismo Beach, San Luis Obispo
Co.), but localities all along the California
coast are represented. In addition, the collec-
tions of the Department of Invertebrate Zo-
ology, California Academy of Sciences (CAS-
IZ) and the collections of the Moss Landing
Marine Laboratories were frequently used.
Most of the live specimens examined were
from the central California coast (viz. Mon-
terey and San Luis Obispo counties); how-
ever, some specimens from northern and
southern California were also examined alive.
Only material which was preserved is listed
under material examined.
For two species (viz. Aeolidiella takanosi-
mensis and Сайта limbaughi) neither live nor
preserved specimens were available for study.
In these cases published descriptions were
relied upon for information, and color photo-
graphs were available for study.
Live specimens were relaxed in 2-
phenoxyethanol and preserved in ethanol and
butylated hydroxytoluene.
The radula from at least one specimen of
each available species was examined. The
buccal mass was removed and macerated
successively in 10% potassium hydroxide and
Clorox (5.25% sodium hypochlorite), cleaned
with forceps and teasing needles, rinsed in
distilled water and mounted in Turtox CMC-
9AF on a microscope slide. Radulae were
usually mounted whole to facilitate determi-
nation of the radular formula. For certain
species it was necessary to separate out in-
dividual teeth to facilitate observation of tooth
shape. The radula of each species was drawn
with the aid of a drawing tube on a Wild M-
20 microscope. The radulae are arranged in
the figures roughly in taxonomic groups so
that similarities and differences may readily
be seen. Drawings consist either of an entire
half row of the radula, or in cases where teeth
are numerous and very similar (e.g. Eudori-
dacea) of only a few representative teeth.
The body of this work is organized so that
a brief generic diagnosis is given first, fol-
lowed by descriptions of the species, based
on observations of both live and preserved
specimens and on the literature. In some
cases it was not possible to discern certain
characters such as the location of the anus
(certain aeolids). The descriptions present as
complete information about external mor-
phology and coloration as is practicable. In-
formation concerning internal morphology and
anatomy is presented only where it may be
useful in identification of species. Odhner
(1934: 229), in discussing the nudibranchs of
the ‘Terra Nova’ Expedition, said:
“... think it necessary to give good fig-
ures of the whole animal, and not of de-
tails only in the ancient manner of Bergh.
His method of description is followed by
later authors with the result that one is
often very little informed as to the essen-
tial external appearance of a species de-
scribed, and this is the more to be re-
gretted as external shape and features
provide not only an immediate means of
recognition, but often also important
systematic characters."
The descriptions are arranged in alphabetical
order so that they may be quickly found. Be-
cause the higher taxa are still in a state of
flux, it was felt that this arrangement would
be more useful than a taxonomic arrange-
ment which may become obsolete in a few
years. The species descriptions present in-
formation in the same sequence for each
species and use the same term for a given
structure throughout. Many of the terms used
are illustrated in Figs. 1 to 28.
The range limits are listed as exactly as
possible in the hope that future collections
will show whether the present known ranges
are increasing or are still imperfectly Known.
It must be kept in mind that published ranges
for animals frequently better reflect the ranges
of collectors than those of the animals.
The total lengths (indicated by T.L.) given
are a rough average for California speci-
mens, based on personal observations and
the literature. However, individuals may vary
widely from the average, and individuals from
other areas may regularly be larger or smaller
than these averages. A case in point is Dis-
codoris sandiegensis which regularly reaches
lengths over 100 mm in Elkhorn Slough, Cal-
ifornia, while 20 miles north and south of this
area specimens average 25 mm in length.
The habitat and food notes are based upon
the literature and personal observation. While
the food habits of some species have been
studied in some detail (e.g. Aeolidia papillo-
sa), the food of some other species is totally
unknown. As a result, the food data given
herein are highly variable, but | feel that even
stating that a species feeds upon an uniden-
tified ascidian is better than no data. The
same sort of variability applies to the habitat
information. Some species are very limited
(e.g. Cumanotus beaumonti is found almost
exclusively on Tubularia crocea in bays) while
116 MCDONALD
Ex. br. app. Rh.
Bh: Br. pl.
Br. pl. Vin 2
о. cor Er
ER aay za < Ам
= Vas “7 — Lab. ten.
| Ex. rh. app.
Br. pl 2 A SS
: Rh.
Et >
> Br. pl. Dor. pro.
: > À N ge — Fr. veil
à nn 8
u 5 AS Per
Pal. rid.
ym rl
D Cer. Eh.
Br. pl.— | u |
S ANN) AU) Ze
7 Site er Le Pr vell
ABR RCN RAS
a PANAM Leann KR
Cer. WHYS WN AN A
Rh. 8
« PC
Le x IF Cer.
| 2: Fr. veil | Rh.
10 # | эт ze hood
Cer. The г
| Rh. 4
A il Cer.
ía : Rh.
$ _ Fr veil | /
a © A Ve X Ag у
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[7 19 20 22 24
Ех. br. app.
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LA
6
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Г Fr. veil
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25 $ /
26 28
FIGS. 1-28. External morphology. 1, Trapania. 2, Acanthodoris. 3, Polycera. 4, Anisodoris. 5, Triopha.
6, Chromodoris. 7, Corambe. 8, Hancockia. 9, Dendronotus. 10, Dirona. 11, Melibe. 12, Tritonia. 13, Doto.
14, Eubranchus. 15, Phidiana. 16, digitiform labial tentacles. 17, auriculate labial tentacles. 18, labial ten-
tacles absent. 19, crescent-shaped oral veil. 20, clavate ceras. 21, spindle-shaped ceras. 22, ceras with
sail-like expansion (Fiona pinnata). 23, ceras showing plume-like gill (Doto). 24, smooth rhinophore. 25,
verrucose rhinophore. 26, annulate rhinophore. 27, perfoliate rhinophore. 28, rhinophore of Rostanga
pulchra. Abbreviations: Br. app.: branchial appendage; Br. pl.: branchial plumes; Cer.: cerata; Dor. pro.:
dorsal process; Ex. br. app.: extra branchial appendage; Ex. rh. app.: extra-rhinophoral appendage; Fo.
cor.: foot corner; Fr. veil: frontal veil; Lab. ten.: labial tentacle; Lat. pro.: lateral process; Or. hood: oral
hood; Or. ten.: oral tentacle; Pal. rid.: pallial ridge; Rh.: rhinophore; V. app.: velar appendage.
CALIFORNIA NUDIBRANCHS ий
others are found т many areas (e.g. Phidi-
ana crassicornis).
Each synonymy lists all the literature deal-
ing with the species, especially publications
of taxonomic interest. The synonymies are
derived from those previously published, with
the addition of new publications and new
synonyms. While it is virtually impossible to
cite every paper which mentions a given
species, the synonymies are as complete as
time and facilities would allow, and are the
most complete yet published for any of the
species discussed. Popular books (viz. John-
son & Snook, 1927; Ricketts 8 Calvin, 1968;
Miner, 1950) which mention various species,
have not been cited in the synonymies, ex-
cept in cases where they give important
taxonomic information. They are, however,
cited in the bibliography. Publications dealing
with the biology of molluscs or opistho-
branchs in general (viz. Hoffman, 1932-1940;
Fischer et al., 1968; Thiele, 1931; Hyman,
1967; Purchon, 1968; Wilbur & Yonge, 1964,
1966; Morton, 1967) are likewise not listed in
the synonymies as they would cause an in-
ordinate increase in the length of the synon-
ymies, but are listed in the bibliography. It is
presumed that anyone seeking information
about a given species will consult these gen-
eral references as well as those listed in a
synonymy.
The location of type-specimens is listed for
those species for which it could be deter-
mined.
The key presented should enable the
nonspecialist, with a minimum of difficulty, to
identify any named species of nudibranch
found along the California coast. The key is
based on the one in Light's Manual (Mc-
Donald, 1975a), but with minor corrections
and the addition of eight more species. Using
the key requires living specimens, as it is
based almost entirely on external morpholo-
gy and coloration. Preserved specimens are
unsuitable as they quickly lose their color and
become shrivelled and contorted. Preserva-
tion also renders observation of important
characters difficult, or even impossible (e.g.
number of branchial plumes or number of
rhinophore lamellae). Some species are high-
ly variable in color and may not always per-
fectly fit a key couplet. Further, there are al-
ways new and undescribed species being
found, as well as occasional specimens with
damaged cerata or rhinophores, or other ab-
normalities.
Four species of nudibranchs which have
been described from California are not de-
scribed herein. Janolus coeruleopictus Cock-
erell 8 Eliot, 1905, corresponds very closely
to Antiopella barbarensis (Cooper, 1863b), the
only important difference being that in A. bar-
barensis the masticatory margin of the man-
dibles is denticulate, while in J. coeruleopic-
tus it is non-denticulate. This seems sufficient
reason to distinguish the two species. Stein-
berg (1963a: 66) has suggested that J. coe-
ruleopictus be considered a nomen dubium,
based on the meager description.
Tritonia palmeri Cooper, 1863a comes
close to Tritonia diomedea Bergh, 1894, but
since the description of T. palmeri is so in-
complete, and since T. palmeri was found in
the intertidal and T. diomedea is not reported
from the intertidal in southern California, it
would be very difficult to consider the two
conspecific. Cockerell (1903: 117) and
Thompson (1971: 334) consider T. palmeri a
nomen dubium.
Doris alabastrina Cooper (1863a: 204) is
described as alabaster white with 12 simple
branchial plumes. The only named species
from California which approaches D. alabas-
trina is Cadlina modesta MacFarland, 1966.
However, the description of D. alabastrina is
so poor that it is best to consider it a nomen
dubium as suggested by Steinberg (1961: 59).
Chromodoris glauca Bergh, 1879e is dis-
cussed with Hypselodoris californiensis (q.v.).
RESULTS AND DISCUSSION
In attempting to review any group of ani-
mals, systematic, nomenclatorial, and other
difficulties arise. With the nudibranchs one is
confronted at the outset with many prob-
lems: several different systems of classifica-
tion, vast numbers of synonyms, improperly
described species, many poorly described
Species, preoccupied names, a relatively large
and widely scattered literature (‘There is also
the great hydra of literature to be wrestled
with...” Winckworth, 1932: 217), and a vast
and often confusing terminology in which
several different terms may refer to the same
entity. Furthermore, nudibranchs have few
hard parts which can be used as taxonomic
characters. This review is intended to help
clarify some of these problems.
There are two points which require discus-
sion. First, the use of the radula as a taxo-
nomic character, and second, the difficulties
presented by the current classification.
118 MCDONALD
The radula is one of the very few seeming-
ly conservative characters in nudibranchs. As
one of the very few hard structures which will
remain almost unchanged by preservation, it
has been much used in classifying nudi-
branchs. Radulae have a number of param-
eters which may be used: the number of
transverse rows of teeth, the number of teeth
per row, the number of denticles on the teeth,
and the shape of the teeth. At present it would
appear that the number of denticles and the
shape of the radular teeth are useful taxo-
nomic characters at the species level in the
Aeolidoidea and Arminoidea and are slightly
less useful in the Dendronotoidea and Ana-
doridacea. In the Eudoridacea the teeth are
usually all hamate and little different among
species and therefore of little use in separat-
ing species. Bertsch (1976b) discusses and
documents variation in nudibranch radulae. In
comparing the radulae of a number of speci-
mens of Triopha maculata, | found that both
the number of teeth per transverse row and
the number of rows increased with size of the
animal. Ferreira (1977) found that the radular
teeth increase in size with increased animal
size but did not find a significant correlation
between number of rows of teeth and animal
size. Ontogenetic variation such as found in
the radulae of T. maculata would account for
the discrepancies between previously pub-
lished radular formulae and those reported
here. Another cause of discrepancy is that
the rachidian teeth may be vestigial and not
obvious unless the radula is properly stained
when mounted. More detailed studies are
necessary to determine how much the radula
varies within a species and within an individ-
ual over time. Only then can the real value of
radulae as taxonomic characters be deter-
mined.
While the classification used herein is not
perfect, | feel that it probably best represents
the present understanding of the relation-
ships of the nudibranchs. O'Donoghue (1926:
22) in listing the species occurring along the
Pacific coast of North America stated:
“Classifications are by their nature ten-
tative, for they express to a large extent
the opinions of their authors regarding
relationships and are subject to revision
in the light of additional knowledge."
Baba (1937a: 195), in describing the Opis-
thobranchia of Japan, summarized quite well
the state of systematics in the Opisthobran-
chia when he said:
“The classification and nomenclature of
the Opisthobranchia are serious prob-
lems still not satisfactorily solved. Sub-
sequent investigations therefore will ren-
der it necessary to alter some of the
accounts of the species, genera, or even
families contained in this paper, but care
has been taken to make them as accu-
rate as possible. The systematic ar-
rangement here adopted is mainly that
of Dr. J. Thiele (1931), with some nec-
essary emendations in accordance with
the current workers’ opinions and on the
basis of my study.”
More than 40 years later, this statement is
still largely true and applies equally well to
the present paper except that the systematic
arrangement is mainly that of Fischer et al.
(1968) rather than of Thiele (1931).
Certain of the taxa in the present classifi-
cation seem ambiguous or inappropriate, for
example, the tribes of the Aeolidoidea (viz.
Pleuroprocta: anus lateral, outside and be-
neath the liver branches; Acleioprocta: anus
between the right liver and the left liver
branches, in the interhepatic space; and
Cleioprocta: anus more or less within or be-
hind the second group of cerata). While
Odhner (1939: 50) felt that these tribes are
clearly distinguished, it now appears that such
is not the case. Edmunds (1964: 7) described
the new genus Selva which has the anus in
the cleioproct position but has an armed pe-
nis, cuspidate radular teeth, smooth rhino-
phores, rounded foot, and 2 short rows of
cerata to the anterior liver, which are all char-
acteristic of the acleioproct family Cuthoni-
dae. Miller (1971: 312), in discussing the aeo-
lids of the Flabellinidae and Eubranchidae
from New Zealand, stated that although
Odhner's tribes show evolutionary stages,
they cut across the aeolid families rather than
separating them. He further stated that at
least 2 of the 3 anal positions occur in sev-
eral families and that there is considerable
variation of each of the anal positions. Thus
it would seem that the position of the anus
may not be a good taxonomic character for
division of the Aeolidoidea into tribes. This is
especially true since it is a character which is
very difficult to observe and is hence not re-
corded for some species. Schmekel (1970:
134) in an extensive study of the genital or-
gans of nudibranchs concluded that the
Acleioprocta form a distinct group, while the
Cleioprocta and Pleuroprocta form another
CALIFORNIA NUDIBRANCHS iS
distinct group. For this latter group Schmekel
formed the infraorder Heteroprocta. While
Schmekel's system seems more realistic at
present than Odhner's system, the resulting
two groups are still not as clearly separated
as one might wish. It would thus appear that
until the characters used for division of higher
taxa are reevaluated and perhaps changed,
certain higher taxa are apt to remain in a state
of flux. While the existing classifications are
not perfect, it is not my intention to propose
a better system, but only to place the Cali-
fornia species into the existing system as best
as possible. Ideally, in order to improve and
somewhat stabilize the classification it is
necessary to study all species of nudibranchs
worldwide, decide which characters best dis-
tinguish the various taxa, and then to ar-
range them in a taxonomic hierarchy.
CLASSIFICATION OF THE
NUDIBRANCHS KNOWN TO OCCUR
IN CALIFORNIA
*. . . there is nothing to be done in the
wide boundless field of natural history
without a system.’’—Gilbert White
This classification is taken largely from
Fischer et al. (1968), with addition of some
new taxa and corrections due to nomencla-
torial changes.
ORDER—NUDIBRANCHIA
SUBORDER—Doridoidea Odhner, 1934
(=Holohepatica Bergh,
1881)
Superfamily—Anadoridacea Odhner,
1939
(=Phanerobranchia
Fischer, 1883)
Tribe—Suctoria Bergh, 1892
Family—Corambidae Bergh, 1892
Corambe Bergh, 1869
Doridella Verrill, 1870
Family—Goniodorididae H. & A.
Adams, 1854
Subfamily—Okeniinae Iredale &
O'Donoghue, 1923
Okenia Menke, 1830, ex Leuc-
kart in Bronn, Ms.
Subfamily—Anculinae Pruvot-Fol,
1954
Ancula Loven, 1846
Trapania Pruvot-Fol, 1931
Subfamily—Hopkinsiinae Odhner,
1968
Hopkinsia MacFarland, 1905
Family—Onchidorididae Alder &
Hancock, 1845
Acanthodoris Gray, 1850
Onchidoris Blainville, 1816
Tribe—Nonsuctoria Bergh, 1892
Family—Triophidae Odhner, 1941
Subfamily—Triophinae Odhner,
1968
Crimora Alder & Hancock, 1862
Triopha Bergh, 1880
Family—Aegiretidae Fischer, 1883
Aegires Loven, 1844
Family—Polyceratidae Alder & Han-
cock, 1845
Laila MacFarland, 1905
Polycera Cuvier, 1817
Superfamily—Eudoridacea Odhner, 1934
(=Cryptobranchia Fischer,
1883)
Family—Cadlinidae Bergh, 1891
Subfamily—Cadlininae
Cadlina Bergh, 1878
Family—Chromodorididae Bergh,
1891
Chromodoris Alder & Hancock,
1855
Hypselodoris Stimpson, 1855
Family—Actinocyclidae Pruvot-Fol,
1934
Hallaxa Eliot, 1909
Family—Conualeviidae Collier &
Farmer, 1964
Conualevia Collier & Farmer,
1964
Family—Aldisidae Odhner, 1939
Aldisa Bergh, 1878
Family—Rostangidae Pruvot-Fol,
831
Rostanga Bergh, 1879
Family —Dorididae Rafinesque, 1815
Doris Linnaeus, 1758
Family —Archidorididae Bergh, 1892
Archidoris Bergh, 1878
Atagema Gray, 1850
Family —Discodorididae Bergh, 1891
Subfamily —Discodoridinae Bergh,
1891
Anisodoris Bergh, 1898
Discodoris Bergh, 1877
Thordisa Bergh, 1877
Family —Asteronotidae Thiele, 1931
Sclerodoris Eliot, 1904
Family —Platydorididae Bergh, 1891
Subfamily —Platydoridinae Bergh,
1891
Platydoris Bergh, 1877
120 MCDONALD
Superfamily —Porodoridacea Odhner,
1968
(=Porostomata,
Bergh, 1892)
Family —Dendrodorididae O'Dono-
ghue, 1924
Dendrodoris Ehrenberg, 1831
SUBORDER—Dendronotoidea Odhner,
1936
Family —Tritoniidae Lamarck, 1809
Tochuina Odhner, 1963
Tritonia Cuvier, 1798
Family —Hancockiidae MacFarland,
1923
Hancockia Gosse, 1877
Family —Dendronotidae Sars, 1878
Dendronotus Alder & Hancock,
1845
Family—Tethyidae Fischer, 1883
Melibe Rang, 1829
Family —Dotidae Gray, 1853
Doto Oken, 1815
SUBORDER—Arminoidea Odhner, 1934
Superfamily—Euarminacea Odhner,
1968
Family —Arminidae Pruvot-Fol, 1927
Armina Rafinesque, 1814
Superfamily—Metarminacea Odhner,
1968
Tribe—Pachygnatha
Family—Dironidae MacFarland,
1912
Dirona Eliot in Cockerell & Eliot,
1905 ex MacFarland, Ms.
Family—Zephyrinidae
Antiopella Hoyle, 1902
SUBORDER—Aeolidoidea Odhner, 1934
Superfamily —Eueolidacea Odhner, 1968
Tribe—Pleuroprocta Odhner, 1939
Family—Coryphellidae Bergh, 1892
Coryphella Gray, 1850
Tribe—Acleioprocta Odhner, 1939
Family—Eubranchidae Odhner,
1934
Subfamily—Cumanotinae Odhner,
1968
Cumanotus Odhner, 1907
Subfamily—Eubranchinae
Odhner, 1934
Eubranchus Forbes, 1838
Family—Cuthonidae Odhner, 1934
Subfamily—Precuthoninae
Odhner, 1968
Precuthona Odhner, 1929
Subfamily—Cuthoninae Odhner,
1934
Cuthona Alder & Hancock, 1855
Tenellia Costa, 1866
Family—Fionidae Alder & Hancock,
1855
Fiona Alder & Hancock, 1855
Tribe—Cleioprocta Odhner, 1939
Family—Phidianidae Odhner, 1968
Subfamily—Phidianinae Miller,
1974
Phidiana Gray, 1850
Subfamily—Babakininae Roller,
1973
Babakina Roller, 1973
Family—Aeolidiidae Orbigny, 1834
Aeolidia Cuvier, 1797
Aeolidiella Bergh, 1867
Cerberilla Bergh, 1873
Family —Spurillidae Odhner, 1939
Spurilla Bergh, 1864
KEY TO THE SPECIES OF NUDIBRANCHS KNOWN TO OCCUR IN CALIFORNIA
“If one does not know the names, one's knowledge of things is useless.’’—Isidorus
1. Gills (branchial plumes) located posteriorly on dorsum (Figs. 1-6) or in 3 groups
directed'Posterionly нео a NN 2
1% Gills (branchial plumes) not located posteriorly on dorsum; respiratory surface
OIDEIWISEHN nee ae Re aie Re SERRES 52
2.(1) | With extra-branchial.appendages (Figs: 1,3)». 22.2.2... man. той 3
2% without extra-branchial appendages. „аа a E AN 9
3.(2) With extra-rhinophoral appendages (Fig. 1); branchial plumes 3 ............... 4
3’. Without extra-rhinophoral appendages; branchial plumes usually more than3 ...6
4.(3) Each rhinophore shaft with 1 extra-rhinophoral appendage (Fig. 1); ground color
whitish; longitudinal brownish lines running most of the length of body; distal
tips of rhinophores, oral tentacles, tips of branchial plumes, extra-branchial and
extra-rhinophoral appendages orange-yellow
Ча Trapania velox
CALIFORNIA NUDIBRANCHS 121
Each rhinophore shaft with 2 extra-rhinophoral appendages ................. 5
With 1 extra-branchial appendage on each side; ground color translucent tan to
whitish; head, tail, and sides of body with irregular patches of dark reddish-
BROWN A, A A NS A NE ba eh Ancula lentiginosa
With 4 or more extra-branchial appendages on each side; ground color trans-
lucent yellowish-white; 3 longitudinal yellow lines running most of the length of
body; rhinophores and extra-branchial appendages with subapical orange-yel-
[IMD A En A ee MO li Ancula pacifica
Body covered with numerous tubercles; yellow spots on body and on tips of low
tubercles; body color dark brownish; branchial plumes 3-5 .... Polycera zosterae
BodyÿmotcoverediWithinumerousitubercles 12. ad a a ee 7.
Overall body color translucent grayish-white; velar appendages (Fig. 3) and ex-
tra-branchial appendages with black on proximal third and a yellow ring imme-
diately distal of black pigment; branchial plumes 5-6, blackish with yellow lines
D E ee Dm te Polycera tricolor
Фета оч соо АК ео nr en 8
With oblong orange spots occurring between longitudinal black lines running the
length of the body; branchialplumes: 7-11 72... Ce 855e5 > Polycera atra
With closely set small black dots on body; rhinophores, corners of foot, velar
and extra-branchial appendages with yellowish-orange; caudal crest and upper
edge of foot with streaks of yellow-orange; branchial plumes 7-9............
Gy Meee Ne teh fit ee Е. Polycera hedgpethi
Rhinophores smooth (Fig. 24); ground color opaque white .................. 10
Rhinophores not:smeoth; ground color variable... 2.2 225.4 aumas ee ns oe es 11
Dorsum with large cylindrical tubercles; usually with many irregularly scattered
dark brown to black spots; branchial plumes 3; rhinophore sheaths with 5-6
hignrroundeditubercles ren... Aare aa aes een Sarna = Aegires albopunctatus
Dorsum nearly smooth, tubercles minute; lacking any brown or black; branchial
plumes about 8; rhinophores long and tapered (Fig. 24); lateral edge of dorsum
Withropaque white GlaMGS: „ее оно tans tes вооон Conualevia alba
With dorsal processes (Fig. 5) other than tubercles or papillae on dorsum or
НЕО ОЕ ee re ER fee ee To 12
Dorsum without dorsal processes, but with tubercles (Fig. 4) or papillae (Fig. 5)
Or entire smooth (BIO). ar nano OR Nee ee Acanthodoris hudsoni
Dorsum with black or brown pigment, not including rhinophores or branchial
ME RE N A ANR CE RE RE CR EE She mee eee 38
Dorsümslackingiblack.or browm pigment”. 70 eme Re ee 46
Dorsum with a lateral series of small brown to black dots with yellow centers;
ground color salmon to yellowish-pink; branchial plumes 12 ...... Cadlina sparsa
Dorsum-lackingiblack-dots with yellow Centers) 225 2... wu ms O a 39
Rhinophores tipped with maroon to brownish-black; dorsum with small tuber-
cles, largest of which are tipped with brown; ground color yellowish ........
Rhinophores not tipped with maroon to brownish-black .................... 40
Branchial plumes 8-10, tripinnate, sprinkled with minute brownish flecks, whitish
distally; blackish pigment in form of many very small dots, with a single major
concentration just anterior of branchial plumes; ground color yellowish to yellow-
ish=onowne tubencles:simalle 6. 72.5 Sarton ee en ee ee Discodoris heathi
Branchial plumes less than 8; blackish pigment not in a single major concentra-
tiomitormingrasblotchiondorsums 2... A ee sees 41
Dorsum with small tubercles which give dorsum a velvety appearance ........ 42
Dorsimiwithlarge Conspicuous tubercles] 2 m ya A en des 43
With numerous, minute flecks of opaque white on margin of dorsum, forming a
diffuse band; dorsum with irregularly oval, light brown spots and numerous mi-
nute light brown flecks; branchial plumes 6 .................... Doris (s.l.) sp.
Lacking diffuse white band around margin of dorsum; dorsum with brown to
almost black rings, or occasionally blotches, of various sizes; branchial plumes
OTI eee TIERE en ERA Discodoris sandiegensis
Dorsum with numerous large, irregular tubercles which bear smaller tubercles;
ground color light yellowish-tan; dorsum with irregularly round, dark tan to brown
blotches between the large tubercles; ventral surface of foot and mantle margin
with flecks ofibrown: branchialiplumess/ (se. Br Sclerodoris tanya
Dorsum with numerous large rather regular tubercles which do not bear smaller
tUDEICIOS catas ot. Ae IAS AS eerie nnd 44
Dorsum with 2 brown to blackish blotches formed of smaller spots on midline;
ground color whitish to yellow or orange; dorsum with many papillae; branchial
PINES OA os A SOI IN EN: Thordisa bimaculata
58.(55’)
McDONALD
Dorsum lacking 2 brown to blackish spots on midline ...................... 45
Blackish pigment in blotches on both dorsum and tubercles; branchial plumes
7, yellowish; ground color light yellow to orange-yellow ... Archidoris montereyensis
Blackish blotches on dorsum only, not on tubercles; branchial plumes 6, whitish;
ground color light yellow to yellow-orange .................. Anisodoris nobilis
Glavusvof rhinophores:brown to black ©. ое. a ee AA 47
Glavus of rhinephores;, white to yellow >... cios a ee 48
Branchial plumes brown to black; ground color whitish; dorsal and ventral sur-
faces of foot and sides of body with minute black spots ....... Cadlina limbaughi
Branchial plumes white to yellow; ground color whitish to yellow; lateral edges
of dorsum with 6-10 yellow dots on each side, anterior-most dot usually pos-
teTOPOLNMODRO RSR ia acre E O none CI Cadlina flavomaculata
Dorsum with yellow band around edge and with yellow tipped tubercles; ground
color whitish to pale yellow; branchial plumes 6, tipped with yellow .........
NE N A NN A SRT RTS Cadlina marginata
Dorsum lacking yellow band around edge ...:..:..-........ er 49
Dorsum with many long, slender papillae giving animal a fuzzy appearance;
branchial plumes 9-12, unipinnate; ground color white to yellowish-white; dor-
sum with many opaque white flecks .................... Onchidoris hystricina
Dorsum lacking long, slender papillae: ;.:.....:.... с. vo ne PRA 50
Dorsum with many bulbous tubercles which appear nearly spherical; ground
color yellowish-white to yellow-orange; rhinophores orange-yellow, with yellow
tips;tubercles.tipped Withrorange: = =. 222 Mecca de ees ee Onchidoris sp.
Tubercles, if present, not bulbous, not appearing spherical .................. 51
Ground color pure white with white tubercles of various sizes on dorsum; bran-
chial plumes 7, white, rather fluffy in appearance ............ Archidoris odhneri
Ground color pale yellow; branchial plumes 10-12, yellowish-white; dorsum with
several small, yellow dots along lateral edges, anterior-most dots extending an-
LEMOlZzOTNMIMOPMONES, Er 6 oe AN ee Cadlina modesta
With cerata (Figs. 8-11, 13-15), or with branched branchial plumes arranged
laterally-om.dorsum (Fig: 12) оао Pe A SE ER 53
Withouticeratan TN PER Ad E a TL le MT ВИ 98
Cerata or branchial plumes branched (Figs. 8, 9, 12) ...................... 54
Cerata not branched (Figs 10,11 1315) $2,021.55 . ооо т 62
Cerata palmately branched, each with 4-26 digitiform projections (Fig. 8); cerata
in 4-7 pairs; head with a broad palmate velar lobe on either side, each with 6-
10 or more unequal digitiform processes; ground color reddish-brown in mature
individuals, younger individuals translucent greenish-brown .... Hancockia californica
Cerata notipalmately branched ©... ;»....2 sr Re 55
Processes on frontal veil simple (Fig. 12) or lacking: .. 2.0... AAA 56
Processes on frontal veil branched (Fig: 9) зо 58
Dorsum distinctly tuberculate, each tubercle tipped with white; ground color
deep orange-yellow to yellow-brown; body margins with an irregular series of
low white branchial plumes; foot edged with narrow white band ............
RN IO ROS A М И Tochuina tetraquetra
Dorsum not distinctly-tuberculate to a EE 57
Ground color white to yellowish-white; frontal veil (Fig. 12) with 7-12 digitiform
processes; body margins with 11-15 branchial plumes on each side; fine retic-
ulate opaque white lines on dorsum; foot edged with a narrow band of opaque
CI MA ER E O A TE, MO КЕ Tritonia festiva
Ground color rose pink; frontal veil with 10-30 white processes; body margins
with about 20-30 branchial plumes; margins of foot, margins of dorsum, and
edge of rhinophore sheath bear a narrow white line .......... Tritonia diomedea
Rhinophore shaft with a lateral: process (Fig. 9)" #2. 2.2... e 59
Rhinophore shaft lacking a lateral process; frontal veil with 4-6 stout, branched
processes; margin of rhinophore sheath with 5-7 short, blunt processes which
are shorter than clavus; dorsum with 4 distinct longitudinal light brown lines
59.(58)
CALIFORNIA NUDIBRANCHS 125
running from posterior of rhinophore shaft to tip of tail; ground color extremely
variable, may be yellow, brown, orange, greenish, or white; 3-6 pairs of cerata
м Dendronotus subramosus
Posterior face of rhinophore shaft with vertical row of 3-6 small, slightly branched
processes; lateral process arising near base of rhinophore shaft; usually 4 pairs
of branched velar processes; dorsal edge of foot usually with an opaque white
line; ground color may be white, gray, brownish, orange-red, or maroon; 4-7
pairs of cerata which may be tipped with orange, yellow, or purple .........
A NE AI OO: Dendronotus iris
Posterior face of rhinophore shaft lacking vertical row of small, slightly branched
processes; margin of rhinophore sheath usually with 5 long processes ........ 60
With a narrow, median, opaque white line extending posteriorly from between
ledasticeratasto mp Oftalmol Anne hate Lee PE EE ER 61
Without a white line dorso-medially on tail; ground color variable, translucent
grayish-white, greenish, or even brownish; cerata in 3-9 pairs; frontal veil usu-
ally with 3 pairs of branched processes ................ Dendronotus frondosus
Frontal veil with 2 pairs of branched processes; general ground color translucent
grayish-white; 4-8 pairs of cerata ............:.......... Dendronotus albus
Frontal veil with 3 pairs of branched processes; ground color translucent gray-
ish-white or lilac; 4-5 pairs of cerata ................ Dendronotus diversicolor
With large oral hood (Fig. 11), having 2 rows of cirri on margin; about 5-6 pairs
of petal-like cerata; ground color yellowish-brown to greenish-brown ........
Eackingilange:oralihoodiwithreinnine A a E eee 63
Rhinophores perfoliate (Fig. 27) or annulate (Fig. 26), not smooth or verrucose ... 64
Rhinophores smooth (Fig. 24) or verrucose (Fig. 25), not perfoliate or annulate ... 75
Cerata extending well in front of rhinophores and same color as rhinophores ...65
Cerata not extending well in front of rhinophores ......................... 67
Cerata spindle-shaped (Fig. 21); lacking frontal veil; oral tentacles present; cera-
ta with a subapical band of yellow-orange, tips blue or white; an orange-red
crest between rhinophores; ground color translucent grayish-white .........
0-0 F050 cai р о Ne PO ee Antiopella barbarensis
Cerata lanceolate (Fig. 10); with a distinct frontal veil; oral tentacles absent ....66
Cerata usually with irregular bumps; ground color light brown to greenish-gray
with fine yellowish-white, olive green, and pink dots; a pale red spot on outer
SIMGIOMCACHUCCHAS eset tee it тЫ Dirona picta
Cerata smooth, lateral margin of each with an opaque white line; anterior edge
of frontal veil and median crest of tail with opaque white line; ground color
transiucent-grayish-Whlte E. a Satine ee Dirona albolineata
Rhinophores arising from a single median stalk, Y-shaped; cerata pinkish-red
on proximal half, followed distally by a wide band of opaque white, then a narrow
band of yellow-orange below translucent tip; ground color whitish to pinkish . .
E A on AES N HA EL Ad Babakina festiva
Rhinophores not ona single common опа. 68
With an opalescent blue line along edge of foot; double opalescent blue median
lines running the length of body and enclosing a bright orange-yellow elongate
spot between and posterior of rhinophores and another in cardiac region; cerata
with a subapical band of orange-yellow, cores extremely variable in color
E LADEN RAS IIA Phidiana crassicornis
Backingiopalescemtspluevliness 22 о WEINE INA 69
al AAA AE PMR AMOR IT EIN EA 70
Rhinophoresperfoliates =... 8.9.71. ооо мои о. 72
Oral tentacles brilliant vermilion to scarlet-red or yellow to deep cadmium orange
STANS ET N EEE EEE MALER EN 71
Oral tentacles encrusted with opaque white dots distally; rhinophores pale yel-
lowish-green distally; cerata with subterminal band of rich brown, cores deep
olive-green toro WNISA=GEena NI I a ae Coryphella pricei
126
71.(70)
FT
80.(76’)
84.
MCDONALD
With an opaque white line dorso-medially and similar lines dorso-laterally; tips
of cerata whitish, cores usually reddish to orange, occasionally brownish
С RA RT RE EEE ee Owe TREE Coryphella trilineata
Lacking opaque white lines on dorsum; ground color light pinkish; with an irreg-
ular band of scarlet-orange to vermilion on either side of head and a band of
similar color below: tip ofi.each'ceras 3... еее: Phidiana stearnsi
Ground color deep purple; rhinophores perfoliate, lamellae deep maroon; cerata
violet proximally, grading to flaming scarlet or orange distally .... Coryphella iodinea
Ground: color not deep ‘purples... 4208 4 404.5 ha me поднос ба о 73
With a narrow orange-red line between and on the proximal half of the oral
tentacles; cerata with pink tinge throughout, cores dark brown to black .....
О RINES AA aie sen Hc ade ar Phidiana hiltoni
Lacking narrow red line between oral tentacles :.................:.. 9.0050: 74
Cerata orange-red with white tips; ground color grayish-white; dorsum along
midline free of cerata; head usually with a pale orange spot ....... Spurilla oliviae
Cerata cores greenish-brown, cerata with pink tinge, tipped with white; ground
color dull orange; midline of dorsum with irregular patches of opaque white; oral
CAES O A ee re Near Spurilla chromosoma
Oral ‘tentacles present: and digitiform: : . 2.045, cocida ee AT
Oral tentacles rudimentary or absent or in the form of a frontal veil ........... 76
Rhinophore sheath calyciform (Fig. 13); cerata large and bulbous with a plume-
like gill;ontinner surface:(Fig. 13) 2.22: 3 e ee 78
Rhinophore sheath absent; cerata lacking plume-like gill ................... 80
Each ceras with thin sail-like expansion on posterior edge (Fig. 22); ground color
HrahiSiicCene gray. TO DIOWAISI SE. 2. ars reste se Stas Fiona pinnata
Cerata lacking posterior sail-like expansion .............................. 81
With: brown:to black On: body” 2... 7 22 2... а edie AN 79
Lacking brown to black on body or cerata; cerata cores orange to pinkish,
cerata rather bulbous; ground color pale yellowish-white; tubercles on cerata
CIOMG ALG geri acs tots vate ne Re O A Doto amyra
Cerata with black rings at base of tubercles; upper border of rhinophore sheaths
smooth; ground color whitish with brown on head, dorsum, and sides .......
CSE ae Se eens ee NE а Е Doto columbiana
Cerata lacking black rings at base of tubercles; upper border of rhinophore
sheaths somewhat scalloped; ground color white to yellowish; cerata long with
YeNOWISNAO DIOWNICOreS cara rc Se eh nn cas a ee ay ae Doto kya
Cerata usually in groups of 1-2, longest cerata shorter than body length; ground
color white to yellowish-white; a few black specks on dorsum ... Tenellia adspersa
Cerata very long, longest about equal to body length, some groups with more
than 2 cerata; ground color grayish-green to light brownish; cerata cores brown-
ish-yellow to bluish-green, with network of similar colored lines on dorsum be-
o o AM ete Bata Sete ee ey ne Me A Cumanotus beaumonti
Dorsum with a distinct (sometimes discontinuous) opaque white line medially . . .82
Dorsum without a distinct opaque white line medially ...................... 83
Usually with a greenish-blue patch anterior and to the left of cardiac region;
cerata with few, very minute, opaque white dots; cerata cores reddish-brown;
ground color translucent grayish-white with pink tinge ....... Coryphella cooperi
Lacking a greenish-blue patch in cardiac region; cerata with numerous opaque
white dots; cerata cores pinkish-brown distally and greenish-brown proximally;
ground color translucent grayish-white without pink tinge ......... Coryphella sp.
Rhinophores with various amounts of orange to red ....................... 84
Fininophores lacking orange:to led)... ое Cy Ae ee 87
Dorsum with distinct opaque white blotches medially; foot corners distinct; ver-
milion on oral tentacles, rhinophores, head, and dorsum; cerata tipped with white
and with a narrow white band below tip, cores brownish .................
BEF Mae NT A ee Ra O EE Aeolidiella takanosimensis
Dorsum lacking distinct opaque white blotches medially; foot corners indistinct .. .85
85.(84’)
85’.
86.(85)
86’.
87.(83’)
87’.
88.(87)
88’.
89.(88’)
89’.
90.(87’)
90’.
91.(90’
—>
91’.
92.(91’)
92’.
93.(92’)
93’.
94.(93)
94’.
95.(93’)
95’.
96.(95’)
96.
97.(96)
CALIFORNIA NUDIBRANCHS 127
Oral tentacles with white on at least distal half; tail with median opaque white
ÜBEL NEAR A Ra I A a 86
Oral tentacles orange-red with white tips; tail lacking median opaque white line;
cerata recumbent with broad, opaque white band extending from base to tip of
each ceras, covering distal third, cores brownish .......... Cuthona columbiana
Rhinophores white distally; light orange covering front of head, extending one
quarter of the way up rhinophores and on bases of oral tentacles; cerata with
small, opaque white spots, cores brownish to dark green .... Cuthona flavovulta
Rhinophores orange-red for nearly entire length; large orange-red spot between
oral tentacles and anterior of rhinophores; cerata with orange-red tips and black-
NCAA o IE N ee A Cuthona lagunae
Cerata irregularly inflated, warty in appearance (Fig. 14) .................... 88
Gerata mot irregulanly inflated; mot warty’ NT MN a 90
Anterior foot corners tentaculiform; rhinophores frequently with 2-3 brown rings;
oral tentacles frequently with a subterminal, brown ring; numerous distinct brown
dots on dorsum, extending up onto cerata; each ceras with a subapical yellow
N A A E Eubranchus misakiensis
Anterior foot corners not produced into tentaculiform processes ............. 89
With a fairly broad, wavy band of deep olive green dorso-medially on the dor-
sum; cerata cores deep olivaceous green; rhinophores frosted with small, opaque
white dots distally and with a subterminal band of brownish to olivaceous green;
body color pale, translucent yellowish-green, with numerous light brown spots
A E EEE. Eubranchus olivaceus
Lacking a band of deep olive green dorso-medially on dorsum; rhinophores and
oral tentacles with subapical band of brownish to light gray or greenish; sides
ombody with: smalli vENOWNdOIS. = ance ee. Geet epee | cee Eubranchus rustyus
With 2 median light orange oval spots, one spot anterior and one posterior of
base of rhinophores; opaque white dots on distal third of rhinophores and oral
tentacles; an irregular, longitudinal series of white blotches medially on dorsum;
cerata with 3 dark brownish-black bands, cores ochre ...... Phidiana morroensis
Lacking median light orange spots anterior and posterior of rhinophores ...... 91
With band of purple midway on oral tentacles and rhinophores; cerata cores
yellowish proximally, olive green to brown medially, and yellow distally ......
Ste eden cede Pa TO Oa TE ces сек Cuthona abronia
Lacking purple band midway on rhinophores and oral tentacles .............. 92
Cerata frosted with opaque white on outer two thirds; dorsum encrusted with
Opaque white; cerata cores pale green to raw umber; distal third of rhinophores
enerustediwithwhlter 3. 2 aa 5 Aalto ee ae Cuthona albocrusta
Cerata not frosted with opaque white on distal two thirds .................. 93
With yellow band near tips and bases of cerata, or with yellow on frontal margin
DEIWEOMMONLACIGS! Hr. ео een 94
Lacking yellow bands on cerata or between oral tentacles .................. 95
With yellow on frontal margin between oral tentacles; cerata with orange flecks,
cores green with some brown; rhinophores and oral tentacles with white dots
Sa ESA ty ge a O E E EN: ИИ Cuthona virens
Lacking yellow on frontal margin; cerata with yellow band near tip and another
near base, cores yellow-brown to dark brown, surface of cerata with white dots
IPMN 5/58) A ee A Are ere Cuthona fulgens
With band of opaque white subapically on each ceras; cerata cores greenish-
brown to brown; dorsum and cerata with opaque white dots ........ Cuthona sp.
Lacking band of opaque white subapically on each ceras ................... 96
Cerata slightly flattened, not round in cross section ....................... 97
Cerata not flattened, but round in cross section, cores dark chocolate brown to
readish-Drowmsstorpinkr 12. a. me Ne certo te A eI DER EBEN EIERN 100
With a triangular to crescentic patch of encrusting white to cream in the area
between the rhinophores and oral tentacles and in cardiac region; body color
dirty white, аи! gray, mauve, rose, or pink .................. Aeolidia papillosa
128 MCDONALD
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FIGS. 29-65. Radulae. 29, Acanthodoris brunnea; (a) sixth (outermost) marginal; (b) fifth marginal; (c)
fourth marginal; (d) third marginal; (e) second marginal; (f) first marginal; (g) lateral. 30, Acanthodoris
hudsoni: (a) fifth (outermost) marginal; (b) fourth marginal; (c) third marginal; (d) second marginal; (e) first
marginal; (f) lateral. 31, Acanthodoris lutea: (a) fourth (outermost) marginal; (b) third marginal; (c) second
marginal; (d) first marginal; (e) lateral. 32, Acanthodoris nanaimoensis: (a) fourth (outermost) marginal; (b)
third marginal; (c) second marginal; (d) first marginal; (e) lateral. 33, Acanthodoris pilosa: (a) fourth (out-
ermost) marginal; (b) third marginal; (c) second marginal; (d) first marginal; (e) lateral. 34, Acanthodoris
rhodoceras: (a) fifth (outermost) marginal; (b) fourth marginal; (c) third marginal; (d) second marginal; (e)
CALIFORNIA NUDIBRANCHS 129
97’. Lacking a patch of white to cream between rhinophores and oral tentacles or in
cardiac region; ground color translucent grayish-white, encrusted with brown to
KECGISI=DNOW Mate ee оо yan San ee Cerberilla mosslandica
98.(52’) Dorsum with undulating longitudinal ridges of white on a pinkish-brown back-
ground; gills located laterally below dorsum edge on either side; edge of dorsum
with an antero-median notch through which the rhinophores project; clavus of
rhinophores with many longitudinal grooves ................ Armina californica
98’. Dorsum without undulating longitudinal ridges; gills located postero-ventrally (Fig.
7); ground color pale gray, with small yellowish-brown blotches and reticulate
HIROSE RE ое А реа 99
99.(98') Dorsum with posterior notch through which gills may be seen; rhinophore shaft
Witiplate-IIke expansion ee a le o Corambe pacifica
99’. Dorsum without posterior notch; rhinophores smooth and tapered ..........
ео Doridella steinbergae
100.(96’) Distal half of rhinophores and dorsal surface of oral tentacles encrusted with
opaque white; ground color translucent white ........... Cuthona cocoachroma
100’. Lacking opaque white on rhinophores and oral tentacles; ground color translu-
centicreamito Drowmish-=pinke* NC seo iaa da Precuthona divae
—
first marginal; (f) lateral. 35, Onchidoris bilamellata: (a) marginal; (b) lateral; (c) rachidian. 36, Onchidoris
hystricina: (a) marginal; (b) lateral; (c) rachidian. 37, Onchidoris sp.: (a) fifth (outermost) marginal; (b) fourth
marginal; (c) third marginal; (d) second marginal; (e) first marginal; (f) lateral. 38, Doridella steinbergae: (a)
fifth (outermost) marginal; (b) fourth marginal; (c) third marginal; (d) second marginal; (e) first marginal; (f)
lateral. 39, Corambe pacifica: (a) sixth (outermost) marginal; (b) fifth marginal; (c) fourth marginal; (d) third
marginal; (e) second marginal; (f) first marginal; (g) lateral. 40, Okenia angelensis: (a) marginal; (b) lateral.
41, Okenia plana: (a) marginal; (b) lateral. 42, Trapania velox: lateral. 43, Ancula pacifica: (a) marginal; (b)
lateral. 44, Ancula lentiginosa: (a) marginal; (b) lateral. 45, Laila cockerelli: (a) first lateral; (b) second
lateral; (c) first marginal; (d) third marginal. 46, Triopha catalinae: (a) fourteenth marginal; (b) thirteenth
marginal; (c) twelfth marginal; (d) eleventh marginal; (e) tenth marginal; (f) ninth marginal; (g) first marginal;
(h) fourteenth (outermost) lateral; (i) thirteenth lateral; (j) first lateral; (К) second rachidian; (I) first rachidian.
47, Triopha occidentalis: (a) twelfth (outermost) marginal; (b) eleventh marginal; (c) tenth marginal; (d)
ninth marginal; (e) eighth marginal; (f) seventh marginal; (g) sixth marginal; (h) fifth marginal; (i) fourth
(outermost) lateral; (|) first lateral; (К) second rachidian; (I) first rachidian. 48, Triopha maculata: (a) eighth
(outermost) marginal; (b) seventh marginal; (c) sixth marginal; (d) second marginal; (e) first marginal; (f)
fifth (outermost) lateral; (9) second lateral; (h) first lateral; (i) second rachidian; (|) first rachidian. 49, Cri-
mora coneja: (a) fourteenth marginal; (b) eleventh marginal; (c) second marginal; (d) first marginal; (e)
second lateral. 50, Hopkinsia rosacea: (a) lateral; (b) marginal. 51, Aegires albopunctatus: (a) twentieth
lateral; (b) first lateral. 52, Polycera atra: (a) first lateral; (b) second lateral; (c) first marginal; (d) second
marginal. 53, Polycera hedgpethi: (a) first lateral; (b) second lateral; (c) first marginal; (d) second marginal;
(e) third marginal; (f) fourth (outermost) marginal. 54, Polycera tricolor: (a) first lateral; (b) second lateral;
(c) first marginal; (d) second marginal; (e) third marginal; (f) fourth (outermost) marginal. 55, Polycera
zosterae: (a) first lateral; (b) second lateral; (c) first marginal; (d) second marginal; (e) third marginal; (f)
fourth marginal; (g) fifth (outermost) marginal. 56, Aldisa sanguinea: (a) fourth lateral; (b) distal portion of
lateral. 57, Rostanga pulchra: (a) sixth-seventh lateral; (b) fortieth lateral; (c) fourteenth lateral; (d) third
lateral; (e) second lateral; (f) first lateral. 58, Cadlina flavomaculata: (a) rachidian; (b) first lateral; (c) twenty-
first lateral; (d) twenty-fifth (outermost) lateral. 59, Cadlina limbaughi: (a) rachidian; (b) first lateral; (c)
thirtieth lateral; (d) forty-fourth (outermost) lateral; after Lance (1962a). 60, Cadlina modesta: (a) rachidian;
(b) first lateral; (c) second lateral; (d) eighteenth lateral; (e) twenty-seventh lateral; (f) twenty-eighth (out-
ermost) lateral. 61, Cadlina sparsa: (a) rachidian; (b) first lateral; (c) second lateral; (d) twelfth lateral; (e)
nineteenth lateral. 62, Cadlina marginata: (a) rachidian; (b) first lateral; (c) thirty-seventh lateral; (d) fifty-
eighth lateral; (e) fifty-ninth lateral; (f) sixtieth (outermost) lateral. 63, Hallaxa chani: (a) lateral; (b) first
marginal; (c) sixth marginal; (d) fourteenth marginal; (e) fifteenth (outermost) marginal. 64, Hypselodoris
californiensis: (a) first lateral; (b) twenty-ninth lateral; (c) one-hundred and tenth lateral; (d) one-hundred
and eighteenth (outermost) lateral. 65, Chromodoris macfarlandi: (a) rachidian; (b) first lateral, dorsal view;
(c) second lateral, dorsal view; (d) first lateral, lateral view; (e) second lateral, lateral view; (f) fifty-third
(outermost) lateral. Scale lines 50 um long.
130 MCDONALD
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FIGS. 66-98. Radulae. 66, Chromodoris porterae: (a) first lateral; (b) seventh lateral; (c) twenty-third
lateral; (d) twenty-sixth lateral; (e) twenty-ninth lateral. 67, Atagema alba: (a) first lateral; (b) second lateral;
(c) third lateral; (d) ninth lateral; (e) nineteenth lateral; (f) twentieth lateral; (g) twenty-first (outermost)
lateral. 68, Conualevia alba: (a) first lateral; (b) thirty-sixth lateral; (c) sixty-sixth lateral; (d) sixty-seventh
(outermost) lateral. 69, Thordisa bimaculata: (a) sixth (outermost) marginal; (b) second marginal; (c) first
marginal; (d) twenty-seventh lateral; (e) fifth lateral; (f) first lateral. 70, Archidoris montereyensis: (a) first
lateral; (b) second lateral; (c) third lateral; (d) fourth lateral; (e) fifth lateral; (f) sixth lateral; (g) twenty-sixth
CALIFORNIA NUDIBRANCHS 131
SYSTEMATIC ACCOUNT
“Nomina se pereunt, perit et cognitio re-
rum.’’—Linnaeus
Acanthodoris Gray, 1850
Body rather soft and doridiform, broadly
rounded anteriorly, somewhat tapered pos-
teriorly; dorsum densely covered with numer-
ous large, conical papillae. Margin of dorsum
covers foot everywhere except posteriorly
where postero-dorsal tip of foot is visible
hooks below with projecting thickenings of
the cuticle. Radular formula n.1.0.1.n, lateral
large and hooked distally, with or without
denticles. Marginals small and frequently
pointed distally.
Penis unarmed, vagina usually very long.
Buccal crop with pharyngeal teeth.
Type-species: Acanthodoris pilosa (Abild-
gaard in Muller, 1789).
Acanthodoris brunnea MacFarland, 1905
Acanthodoris brunnea MacFarland, 1905: 52.
when animal is actively crawling. Labial ten-
tacles large and auriculate, extending nearly
to edge of dorsum. Rhinophores rather long
and perfoliate, retractile into low sheaths bor-
dered by low papillae. Branchial plumes usu-
ally 5-9, usually bi- or tripinnate, arranged in
a circle around postero-dorsal anus, they are ghue, 1926: 218. O'Donoghue, 1927a: 6.
non-retractile below dorsum but are contrac- Smith & Gordon, 1948: 180. LaRocque,
tile. 1953: 256. MacFarland, 1966: 118-120, pl.
Armature of labial disc consists of minute 20, figs. 5-6. Hurst, 1967: 255 ff, text fig.
MacFarland, 1906: 146-147, pl. 20, figs.
81-88a; pl. 21, fig. 104; pl. 29, figs. 20-
21. Berry, 1907: 35. O'Donoghue, 1921:
171-172, pl. 4, figs. 41-42. O'Donoghue,
1924: 24. MacFarland, 1925: 53-55.
MacFarland, 1926: pl. 2, fig. 7. O'Dono-
—
lateral; (h) thirty-first lateral; (i) thirty-second lateral; (j) thirty-third lateral; (k) thirty-fourth lateral; (I) thirty-
fifth lateral; (m) thirty-sixth (outermost) lateral. 71, Archidoris odhneri: (a) first lateral; (b) twenty-sixth
lateral; (с) fortieth lateral; (а) forty-first lateral; (e) forty-second lateral; (f) forty-third lateral; (9) forty-fourth
(outermost) lateral. 72, Sclerodoris tanya: (a) thirty-fourth (outermost) lateral; (b) thirty-third lateral; (c)
thirty-second lateral; (а) ninth lateral; (e) first lateral. 73, Doris (s.l.) sp.: (a) twenty-fifth (outermost) lateral;
(b) twenty-fourth lateral; (c) twenty-third lateral; (d) fourteenth lateral; (e) first lateral. 74, Discodoris san-
diegensis: (a) thirty-first (outermost) lateral; (b) thirtieth lateral; (c) twenty-ninth lateral; (d) twenty-eighth
lateral; (e) eleventh lateral; (f) second lateral; (g) first lateral. 75, Platydoris macfarlandi: (a) seventieth
(outermost) lateral; (b) sixty-ninth lateral; (c) sixty-eighth lateral; (d) seventh lateral; (e) first lateral. 76,
Anisodoris nobilis: (a) forty-third (outermost) lateral; (b) forty-second lateral; (c) forty-first lateral; (d) fif-
teenth lateral; (e) first lateral. 77, Discodoris heathi: (a) nineteenth (outermost) lateral; (b) eighteenth lateral;
(c) seventeenth lateral; (d) eleventh lateral; (e) third lateral; (f) second lateral; (g) first lateral. 78, Dendron-
otus albus: (a) rachidian; (b) first lateral; (c) second lateral; (d) third lateral; (e) fourth lateral; (f) fifth lateral;
(9) sixth lateral; (п) seventh lateral; (i) eighth (outermost) lateral. 79, Dendronotus diversicolor: (a) rachidian;
(b) first lateral; (c) second lateral; (d) third lateral; (e) fourth lateral; (f) fifth lateral; (g) sixth lateral; (h)
seventh lateral; (i) eighth (outermost) lateral. 80, Dendronotus frondosus: (a) rachidian; (b) first lateral; (c)
second lateral; (d) third lateral; (e) fourth lateral; (f) fifth lateral; (g) sixth lateral; (h) seventh (outermost)
lateral. 81, Dendronotus subramosus: (a) rachidian; (b) first lateral; (c) second lateral; (d) third lateral; (e)
fourth lateral; (f) fifth (outermost) lateral. 82, Dendronotus iris: (a) rachidian; (b) first lateral; (c) second
lateral; (d) third lateral; (e) fourth lateral; (f) fifth lateral; (g) sixth lateral; (h) seventh lateral; (i) tenth lateral;
(j) fifteenth lateral; (к) sixteenth lateral; (I) seventeenth lateral; (m) eighteenth (outermost) lateral. 83, Doto
amyra, rachidian. 84, Doto columbiana, rachidian; after O'Donoghue (1921). 85, Doto kya, rachidian. 86,
Hancockia californica: (a) rachidian; (b) lateral. 87, Tochuina tetraquetra: (a) one-hundred sixty-second
(outermost) marginal; (b) one-hundred twenty-fourth marginal; (c) seventeenth marginal; (d) lateral; (e)
rachidian. 88, Tritonia diomedea: (a) eighty-fourth (outermost) marginal; (b) eighty-third marginal; (c) sev-
enty-third marginal; (d) first marginal; (e) lateral; (f) rachidian. 89, Tritonia festiva: (a) twenty-fourth (out-
ermost) marginal; (b) sixth marginal; (c) lateral; (d) rachidian. 90, Armina californica: (a) sixty-third (outer-
most) marginal; (b) sixty-second marginal; (c) fifty-fourth marginal, distal portion; (d) fifth-second marginal,
distal portion; (e) fiftieth marginal, distal portion; (f) thirty-eighth marginal; (g) tenth marginal, distal portion;
(h) ninth marginal, distal portion; (i) eighth marginal, distal portion; (j) lateral; (k) rachidian. 91, Antiopella
barbarensis: (a) fourteenth (outermost) lateral; (b) thirteenth lateral; (c) twelfth lateral; (d) fourth lateral; (e)
third lateral; (f) second lateral; (g) first lateral; (h) rachidian. 92, Dirona albolineata: (a) marginal; (b) lateral;
(c) rachidian; (d) rachidian, lateral view. 93, Dirona picta: (a) marginal; (b) lateral; (c) rachidian; (d) rachidian,
lateral view. 94, Aeolidia papillosa, rachidian. 95, Aeolidiella takanosimensis, rachidian; after Ferreira &
Bertsch (1973). 96, Cerberilla mosslandica, rachidian. 97, Spurilla chromosoma, rachidian. 98, Spurilla
oliviae, rachidian. Scale lines 50 um long.
132 MCDONALD
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FIGS. 99-123. Radulae. 99, Cumanotus beaumonti: (a) rachidian; (b) lateral. 100, Coryphella cooperi:
(a) rachidian; (b) lateral. 101, Coryphella iodinea: (a) rachidian; (b) lateral. 102, Coryphella pricei: (a) rachidian;
(b) lateral. 103, Coryphella trilineata: (a) rachidian; (b) lateral. 104, Coryphella sp.: (a) rachidian; (b)
lateral. 105, Eubranchus misakiensis: (a) rachidian; (b) lateral. 106, Eubranchus olivaceus: (a) rachidian;
(b) lateral. 107, Eubranchus rustyus: (a) rachidian; (b) lateral. 108, Phidiana crassicornis: (a) rachidian; (b)
rachidian, median cusp, lateral view of distal portion. 109, Phidiana hiltoni, rachidian. 110, Phidiana mor-
roensis, rachidian. 111, Phidiana stearnsi, rachidian. 112, Fiona pinnata, rachidian. 113, Tenellia adspersa,
rachidian. 114, Babakina festiva, rachidian. 115, Precuthona divae, rachidian. 116, Cuthona abronia,
rachidian. 117, Cuthona albocrusta, rachidian. 118, Cuthona columbiana, rachidian. 119, Cuthona flavovul-
ta, rachidian. 120, Cuthona fulgens, rachidian. 121, Cuthona lagunae, rachidian. 122, Cuthona virens,
rachidian. 123, Cuthona sp., rachidian. Scale lines 50 um long.
brown, to almost black in occasional speci-
mens; dorsum sprinkled with irregular blotch-
es of black and numerous minute flecks of
lemon yellow. T.L.: 12 mm.
Radular formula of specimen examined
13(5-6.1.0.1.5-6), MacFarland (1966: 119)
reports 24-28(6-7.1.0.1.6-7). Laterals (Fig.
33e) bear 8-12 denticles in specimen exam-
ined, MacFarland (1966: 119) reports 14-19
denticles on laterals.
Type-locality: Monterey Bay, California.
1, pl. 26, fig. 1; fig. 24-5. Haderlie, 1968:
333, 339. Sphon 4 Lance, 1968: 75. Roller
& Long, 1969: 429. Haderlie, 1969: tab. 2.
Bernard, 1970: 85. Robilliard, 1971a: 164,
165. Abbott, 1974: 361, text fig. 4306.
McDonald & Nybakken, 1978: 111. Had-
erlie 8 Donat, 1978: 52, 60.
Type-specimens: U. S. Nat. Mus., no.
181293.
Foot oval, bluntly rounded and bilabiate
anteriorly, rounded posteriorly. Rhinophores
with 20-28 lamellae, deep blue-black to
brownish, tipped with pale yellow. Branchial
plumes 7, bipinnate, light brown, with 2 lon-
gitudinal dark brown lines on inner surface,
tipped with yellow. General ground color
Range and habitat: Nanaimo, British Co-
lumbia, Canada (Robilliard, 1971a), to Santa
Monica Bay, Los Angeles Co., California
(personal observation). Intertidal to 110 m,
rare in rocky intertidal in California. Occa-
sionally found in rocky subtidal or dredged
from muddy bottom in deeper water. Kozloff
CALIFORNIA NUDIBRANCHS 133
(1973: 93) states that it feeds upon bryozo-
ans.
Remarks: The specimens reported by
Haderlie (1968, 1969) and by Haderlie & Do-
nat (1978), based on the rarity of A. brunnea
and on the habitat to which their specimens
were found, may prove to be Onchidoris bi-
lamellata.
Acanthodoris hudsoni MacFarland, 1905
Acanthodoris hudsoni MacFarland, 1905: 51.
MacFarland, 1906: 144-146, pl. 20, figs.
73-80. O’Donoghue, 1921: 170-171.
O’Donoghue, 1922b: 164. MacFarland,
1925: 51-53. MacFarland, 1926: pl. 2, fig.
1. O'Donoghue, 1926: 218. O'Donoghue,
1927a: 6. Smith & Gordon, 1948: 180.
LaRocque, 1953: 256. Steinberg, 1963b:
70. Hurst, 1967: 255 ff, text fig. 2, pl. 26,
fig. 2. Lee 8 Brophy, 1969: 220. Robilliard,
1969a: 290. Bernard, 1970: 85. McDonald,
1970: 375. Roller, 1970b: 482. Robilliard,
1971a: 164. Bertsch et al., 1972: 302.
Sphon, 1972a: 154. Gosliner & Williams,
1973b: 352-354. Abbott, 1974: 362. Lam-
bert, 1976: 294, 296. Thompson, 1976a:
79, text figs. 38a-b. Nybakken, 1978: 135.
Cadlina luteomarginata MacFarland. Kozloff,
1973: pl. X.
Type-specimens: U. S. Nat. Mus., no.
181289.
Foot oval, bluntly rounded and bilabiate
anteriorly, rounded posteriorly. Rhinophores
bear 22-24 lamellae, clavus lemon yellow.
Branchial plumes 5, bipinnate, grayish white,
tipped with lemon yellow. General ground
color translucent yellowish white. Dorsal pa-
pillae tipped with lemon yellow; mantle mar-
gin with band of same color. T.L.: 15 mm.
Radular formula of specimens examined
18(5-6.1.0.1.5-6), MacFarland (1925: 51) re-
ports 27(5-6.1.0.1.5-6). Laterals (Fig. 30f)
large and hooked, bearing 5-8 denticles on
CUSP.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976), to Gavi-
ota, Santa Barbara Co., California (Lee &
Brophy, 1969). Intertidal to 215 m, rare in
rocky intertidal in California.
Acanthodoris lutea MacFarland, 1925
Acanthodoris lutea MacFarland, 1925: 60-65.
MacFarland, 1926: pl. 2, figs. 2, 6, 8; pl. 3,
figs. 3, 6. O'Donoghue, 1927a: 7. Lance,
1961: 67. Steinberg, 1963b: 70. Mac-
Farland, 1966: 120-121, pl. 32, fig. 15.
Marcus & Marcus, 1967a: 202. Farmer,
1967: 342. Sphon 8 Lance, 1968: 76.
Turner et al., 1969: 132, append. 1, 2. Roll-
er £ Long, 1969: 425. Roller, 1970b: 482.
Gosliner 8 Williams, 1970: 176. Bertsch et
al., 1972: 303. Abbott, 1974: 362. Michel,
1976: 46, fig. 4. Nybakken, 1978: 144.
Type-specimens: not listed, may be at
CASIZ.
Foot oval, bluntly rounded and bilabiate
anteriorly, rounded posteriorly. Rhinophores
bear about 26 lamellae, orange-red to red.
Branchial plumes 9, bipinnate, grayish white.
General ground color orange to red-orange,
papillae of same color. Dorsum sprinkled with
numerous minute lemon yellow flecks. T.L.:
22 mm.
Radular formula of specimens examined
33-36(4-5.1.0.1.4-5), MacFarland (1966:
120) reports 34-39(5-6.1.0.1.5-6). Laterals
(Fig. 31e) bear 5-7 denticles on cusp,
MacFarland (1966: 120) reports 2-4 denti-
cles on laterals.
Type-locality: Cayucos, San Luis Obispo
Co., California.
Range and habitat: Dillon Beach, Marin
Co., California (Steinberg, 1963b), to 12.9 km
S of Cabo Colonet, Baja California, Mexico
(Farmer, 1967). Intertidal to 500 m.
Remarks: This species frequently gives off
a quite pungent odor of sandalwood or ce-
dar.
Acanthodoris nanaimoensis
O'Donoghue, 1921
Acanthodoris nanaimoensis O'Donoghue,
1921: 172-174, pl. 10, figs. 43-44. O'Don-
oghue, 1922a: 126. O'Donoghue, 1922b:
164. O'Donoghue & O'Donoghue, 1922:
139. O'Donoghue, 1926: 218. MacFarland,
1926: 100. O'Donoghue, 1927a: 7. La
Rocque, 1953: 256. Steinberg, 1963a: 64.
Steinberg, 1963b: 70. MacFarland, 1966:
123. Hurst, 1967: 255 ff, text fig. 3, pl. 26,
fig. 3; fig. 24-8. Lance, 1968: 11. Lee 8
Brophy, 1969: 220. Roller 8 Long, 1969:
425. Roller, 1970a: 371. Bernard, 1970: 85.
Gosliner & Williams, 1970: 176. Holleman,
1972a: 60. Bertsch et al., 1972: 303.
Sphon, 1972a: 154-155. Abbott, 1974:
362. Lambert, 1976: 294, 296. Thompson,
1976a: 79, text fig. 38c. Thompson, 1976b:
42. McDonald 8 Nybakken, 1978: 111.
Acanthodoris columbina MacFarland, 1926:
94-100, pl. 2, figs. 5, 9-10; pl. 3, figs. 1-
134 MCDONALD
2, 5. O'Donoghue, 1927a: 3, 4, 7. Stein-
berg, 1963a: 64. MacFarland, 1966: 121-
123, pl. 32, fig. 16. Roller, 1970a: 371.
Type-specimens: Mus. Dominion Biol.
Stat., Nanaimo, B. C., Canada.
Foot oval, bluntly rounded and unilabiate
anteriorly and rounded posteriorly. Rhino-
phores bear 22-26 lamellae, shaft brownish,
clavus deep wine red to maroon, entire rhi-
nophore sprinkled with lemon yellow flecks.
Branchial plumes 7-9, bipinnate, light gray-
brown, tipped with wine red to maroon. Gen-
eral ground color translucent grayish-white to
dirty brownish-mauve in larger specimens;
dorsal papillae grayish-white to brownish,
tipped with lemon yellow. Mantle margin with
band of lemon yellow all around. T.L.: 30 mm.
Radular formula of specimens examined
36-37(4.1.0.1.4), O'Donoghue (1921b: 173)
reports 35(6-7.1.0.1.6-7), while MacFarland
(1966: 122) reports 40-43(5.1.0.1.5) for A.
columbina. Laterals (Fig. 32e) bear 2 denti-
cles on cusp in specimens examined, Mac-
Farland (1966:122) reports 6-8 denticles on
laterals, while O'Donoghue (1921: 173) re-
ports no denticles.
Type-locality: Jesse Island, Vancouver Is-
land, British Columbia, Canada.
Range and habitat: Wales Island, British
Columbia, Canada (Lambert, 1976), to Puri-
sima Point, Santa Barbara Co., California (Lee
& Brophy, 1969). Intertidal to 10 m, quite rare
south of Pigeon Point, San Mateo Co., Cali-
fornia, more common northward. Feeds upon
an unidentified encrusting ascidian.
Acanthodoris pilosa
(Abildgaard in Müller, 1789)
Doris pilosa Abildgaard in Müller, 1789: 3, pl.
85, figs. 5-8. Gmelin in Linnaeus, 1791:
3106. Cuvier, 1804: 451, 470. Blainville
1819: 449. Lamarck, 1819: 312. Rapp,
1827: 517. Bosc, 1830: 111. Cuvier, 1834,
115. Bouchard-Chantereaux, 1835: 139-
140. Lamarck, 1836: 464. Johnston,
1838a: 54. Johnston, 1838b: pl. 2, figs. 9-
10. Thompson, 1840: 86. Hassall, 1842:
133. Thompson, 1844: 250. Alder & Han-
cock, 1851c: fam. 1, pl. 1, figs. 1, 3-5, 12;
pl. 2, figs. 2-6; pl. 15. Hancock & Emble-
fon, 1852: 208 ff, pl. 11, figs. 4, 6; pl. 13,
figs. 6-12; pl. 14, fig. 4; pl. 16, figs. 1, 3-
7; pl. 17, fig. 8. Gosse, 1853: 62. Byerly,
1854: 44. Alder & Hancock, 1855: 4, 27,
31, 32, 43. Gosse, 1856: 105. Thompson,
1856: 274. Collingwood, 1859: 463-464.
E. Wright, 1859: 88. Collingwood, 1860:
202. Collingwood, 1861: 114. Hancock 8
Norman, 1864: 49, 51, 53. Meyer 8 Mö-
bius, 1865: 63-67, pl. 5. Mcintosh, 1865:
390. Robertson, 1868: 205. Jeffreys 1869:
93. Dall, 1870: 249. Gould, 1870: 232.
Sauvage, 1873: 29. Mcintosh, 1874: 432.
McIntosh, 1875: 89. Friele 8 Hansen, 1876:
71, pl. Il, fig. Il. Tiberi, 1880: 207. Leslie &
Herdman, 1881: 312. Hertzenstein, 1885:
709. Locard, 1886: 27. Higgins, 1886: 25.
Haddon, 1886: 530. Herdman & Clubb,
1892: 148. Hecht, 1893: XIV, XVI. Cooke,
1899: 64. Colgan, 1908: 110. Colgan,
1909: 174. Pelseneer, 1911: 55, pl. 15, fig.
24; pl. 16, figs. 1-4. Chumley, 1918: 55,
169. White, 1938: 15. Volodchenko, 1955:
250, pl. 48, fig. 6.
Doris stellata Gmelin in Linnaeus, 1791: 3107.
Cuvier, 1804: 449 ff, 470. Blainville, 1819:
449. Lamarck, 1819: 311. Rapp, 1827: 517.
Bosc, 1830: 113. Cuvier, 1834: 114. Bou-
chard-Chantereaux, 1835: 139. Philippi,
1836: 105. Lebert, 1846: 444. Alder, 1850:
110.
Doris nigricans Fleming, 1820: 618. Fleming,
1828: 283. Johnston, 1838a: 55. Hassall,
1842: 133. Alder, 1850: 110. Herdman,
1886: 269.
Doris flemingii Forbes, 1838: 3, pl. 1, figs. 2-
3.
Doris sublaevis Thompson, 1840: 87, pl. 2,
fig: dE
Doris similis Alder & Hancock, 1842: 32-33.
Alder, 1850: 111.
Doris subquadrata Alder & Hancock, 1845c:
313-314. Alder & Hancock, 1851c: fam. 1,
pl. 16. Forbes & Hanley, 1851: 571. Alder
& Hancock, 1855: 31, 43. Collingwood,
1860: 200, 202. Collingwood, 1861: 113.
Collingwood & Byerley, 1862: 189. Mc-
Intosh, 1865: 390. Jeffreys, 1869: 93.
Doris fusca Müller. Lovén, 1846: 136.
Doris tomentosa Cuvier. Lovén, 1846: 136.
Doris laevis. Gray, 1850: pl. 214, fig. 6.
Doris rocinela Leach in Gray, 1852: 19-20.
Acanthodoris pilosa (Abildgaard in Müller).
Adams & Adams, 1854: 56, pl. 63, fig. 4.
Chenu, 1859: 404, fig. 3045. Mörch, 1868:
203. Sars, 1878: 308-309, 364, 383, 390,
402, pl. XIV, fig. 4. Bergh, 1879a: 359.
Bergh, 1880a: 240-246, pl. 10, figs. 12-
15; pl. 11, figs. 1-2; pl. 12; pl. 13, figs. 2-
5. Bergh, 1880b: 91-97, pl. 10, figs. 12-
15; pl. 11, figs. 1-2; pl. 12; pl. 13, figs. 2-
5. Bergh, 1881a: pl. L, figs. 1-5. Verrill,
1882b: 549. Herdman, 1886: 269, 277.
Fischer, 1887: 519, text fig. 282. Herdman
CALIFORNIA NUDIBRANCHS 135
& Clubb, 1889: 226. Garstang, 1889: 178.
Carus, 1889-1893: 224. Garstang, 1890:
446-447. Norman, 1890: 72. Bergh, 1890b:
989-990. Bergh, 1892: 1151 (159). Herd-
man & Clubb, 1892: 133, 146, pl. 6, figs.
4-5. Garstang, 1894: 226. Tregelles, 1896:
220. Herdman et al., 1896: 446. Gamble,
1896: 132, 133. Cooke, 1899: 66. Beau-
mont, 1900: 849. Nichols, 1900: 596.
Knight, 1901: 207. Conchol. Soc., 1901:
26. Johansen, 1902: 387. Farran, 1904: 3.
Mar. Biol. Assoc., 1904: 284. Cockerell &
Eliot, 1905: 32. Norman & Scott, 1906: 217.
Nordgaard, 1907: 33. Odhner, 1907: 72.
Eliot, 1907: 327 ff. Elmhirst, 1908: 228.
Walton, 1908: 238. Balch, 1909: 36. Far-
ran, 1909: 17. Eliot, 1910: 4, 8, 13, 28, 155.
Colgan; 1911: 25. Walton, 1913: 110.
Sumner et al., 1913: 705. Farran, 1915:
10, 67. Johnson, 1915: 173. Evans 8 Ev-
ans, 1917: 110. Bardarson, 1919: 73. Bar-
darson, 1920: 108. Odhner, 1922: 24. Elm-
hirst, 1922: 42. Iredale & O'Donoghue,
1923: 222-223. O'Donoghue, 1924: 30-31.
Larsen, 1925: 28-30, text figs. 20-21, pl.
figs. 4a-c. Odhner, 1926b: 25. O'Dono-
ghue, 1926: 219. Jutting, 1927: LXXXVII!.
Loyning, 1927: 246, 247, 250-252, 262.
O'Donoghue, 1927a: 6. Derjugin, 1928:
320. Lemche, 1928: 15-16. O'Donoghue,
1929: 776. Fisher, 1931: 198. Mar. Biol.
Assoc., 1931: 274. Monod & Dollfus, 1932:
163, 191. Winckworth, 1932: 234. Leigh-
Sharpe, 1933: 114. Renouf, 1934: 400.
Fisher, 1935: 249. Baba, 1935b: 115, 117-
119, pl. 7, figs. 5-7. Baba, 1937a: 199.
Baba, 1937b: 294. Moore, 1937: 190. Mil-
lott, 1937: 406 ff. White, 1938: 15, 17.
Lemche, 1938: 21-22 ff. Odhner, 1939: 39.
Volodchenko, 1941: 60, 61. McMillan,
1944: 161. Jutting, 1947: 64. Purchon,
1947: 292 ff. Cornet & Marche-Marchad,
1951: 32. Jaeckel, 1952: 28 ff. Stock, 1952:
58. Forrest, 1953: 232, text fig. 5b. Wil-
liams, 1954: 106. Graham, 1955: 152, Clark
& Milne, 1955: 180. Mar. Biol. Assoc.,
1957: 310. Baba, 1957: 8, 13, text fig. 5.
Thompson, 1959: 240. Marcus, 1959: 61.
Swennen, 1959: 57, 58. Buznikov, 1960:
374. Thompson, 1960b: 126-127, text fig.
1. Buznikov & Manukhin, 1961: 226, text
fig. 2. Thompson, 1961: 236. Marcus,
1961: 26. Miller, 1961: 107, 114. Swen-
nen, 1961: 200. Roginskaya, 1962a: 88, 92,
figs. 1.4-1.5. Roginskaya, 1962b: 203, 212,
213, fig. 1.4. Miller, 1962: 552-553, text
fig. 6. Zenkevitch, 1963: 112. Bruce et al.,
1963: 202-203. Steinberg, 1963b: 70.
Thompson, 1964: 280 ff. Marcus & Mar-
cus, 1967a: 202. Miller, 1967: 9. Thomp-
son, 1967: 9. Morse, 1967: 770. Morse,
1968: 305-319, text figs. 1-9. Edmunds,
1968b: 131. Barrett, 1969: 69. Loveland et
al., 1969: 418. Franz, 1970: 172 ff. Hughes,
1970b: 81, 82. Kress, 1971: 326, 336-338,
text figs. 8-10. Meyer, 1971: 137-139.
Holman, 1972: 179-183. Platts, 1973: 383,
385. Rasmussen, 1973: 267-268, 447.
Harris, 1973: 217, 221, 246-247. Abbott,
1974: 361, pl. 16, fig. 4305. Roginskaya,
1974b: 998. Rozsa, 1974: 8. Clark, 1975:
36-37. Franz, 1975a: 81. Thompson &
Brown, 1976: 90, fig. 46. Thompson,
1976a: 36. Thompson, 1976b: 70, 92, 80,
fig. 7. Crampton, 1977: 49. Garlo, 1977:
24, 25, 27. Bleakney & Saunders, 1978:
82. McDonald & Nybakken, 1978: 111.
Clark & Goetzfried, 1978: 290. Ortea, 1979:
47.
Acanthodoris subquadrata (Alder & Han-
cock). Adams & Adams, 1854: 56. Abra-
ham, 1877: 228. Bergh, 1879a: 360. Bergh,
1880a: 240. Bergh, 1880b: 91. Bergh,
1892: 1151 (159). Cooke, 1899: 66. Con-
chol. Soc., 1901: 26. Eliot, 1910: 155. Ire-
dale 8 O'Donoghue, 1923: 222. Mac-
Farland, 1925: 49. Winckworth, 1932: 234.
White, 1938: 17. Thompson, 1964: 290.
?‘‘Lamellidoris Elfortiana Blv.?’’ Mórch, 1868:
204.
Doris quadrangulata Alder & Hancock. Jef-
freys, 1869: 93-94.
Doris bifida Verrill, 1870: 406-407. Verrill,
1872: 210. Verrill, 1873: 307, 333, 664-
665, pl 25 10. 176:
Doris pilosa var. stellata Gmelin т Linnaeus.
Sauvage, 1873: 30.
Acanthodoris citrina Verrill, 1879: 313-314.
Verrill, 1882b: 549. Bergh, 1892: 1151
(159).
Acanthodoris ornata Verrill, 1879: 314. Ver-
rill, 1882b: 549. Bergh, 1892: 1151 (159).
Acanthodoris stellata (Gmelin in Linnaeus).
Verrill, 1879: 313. Verrill, 1882b: 549.
Acanthodoris pilosa var. albescens Bergh,
1880a: 246-247, pl. 10, figs. 14-15; pl. 11,
fig. 2; pl. 12, figs. 13-16. Bergh, 1880b:
97, pl. 10, figs. 14-15; pl. 11, fig. 2; pl. 12,
figs. 13-16. Bergh, 1892: 1151 (159).
O’Donoghue, 1921: 168. O'Donoghue,
1922b: 164. O'Donoghue, 1926: 219.
O'Donoghue, 1927a: 6.
Acanthodoris pilosa var. purpurea Bergh,
1880a: 247-252, pl. 12, figs. 1-9. Bergh,
1880b: 98-103, pl. 12, figs. 1-9. Bergh,
1892: 1151 (159). O'Donoghue, 1924: 31.
136 MCDONALD
O'Donoghue, 1926: 219. O'Donoghue,
1927a: 6.
Acanthodoris quadrangulata (Alder & Han-
cock). Herdman, 1886: 269. Herdman,
1896: 49. Herdman et al., 1896: 446. Con-
chol. Soc., 1901: 26.
Doris (Acanthodoris) pilosa. Herdman, 1890a:
pl. 6, fig. 2.
Acanthodoris pilosa var. nov. zealandiae
Bergh, 1905a: 94-97, pl. 6, figs. 23-26, pl.
Fong. t:
Acanthodoris pilosa var. pallida Bergh, 1905a:
97-98, pl. 6, figs. 27-29; pl. 7, fig. 2. Eliot,
1907: 329.
Acanthodoris pilosa ornata Verrill. Johnson,
1915: 173. Johnson, 1934: 157. La-
Rocque, 1953: 257.
Acanthodoris pilosa pilosa. Johnson, 1934:
157. LaRocque, 1953: 256.
Acanthodoris pilosa var.
Odhner, 1939: 39.
Acanthodoris pilosa albescens Bergh. La-
Rocque, 1953: 256.
Acanthodoris pilosa purpurea Bergh. La-
Rocque, 1953: 257. Marcus & Marcus,
1967a: 202.
fusca Loven.
Type-specimens: not listed.
Foot oval, bluntly rounded and bilabiate
anteriorly and rounded posteriorly. Rhino-
phores bear 18-25 lamellae, translucent
grayish-white with numerous small brown
flecks which are most concentrated on cla-
vus, with a few small yellow flecks, tips whit-
ish. Branchial plumes 5-9, tripinnate, gray-
ish-white, densely covered with small,
brownish flecks, and a lesser number of yel-
low and white flecks. General ground color
translucent grayish-white; dorsum densely
covered with numerous brown flecks and a
lesser number of yellow flecks. Dorsal pa-
pillae tipped with black. Color quite variable
in specimens from Europe and Atlantic coast
of North America. T.L.: 25 mm.
Radular formula of specimens examined
28(3-4.1.0.1.3-4), Baba (1935b: 118) re-
ports 24(4.1.0.1.4). Laterals (Fig. 33e) bear
4-7 denticles on cusp, Baba (1935b: 118) re-
ports 4 denticles on laterals.
Type-locality: ‘Mari Norvegico.”
Range and habitat: Kiska Island, Alaska
(Bergh, 1880a) to Morro Bay, San Luis Obis-
po Co., California (personal observation); cir-
cumboreal; Atlantic coast of North America
(Franz, 1975a); Japan (Baba, 1935b); Eu-
rope (Pruvot-Fol, 1954); Okhotsk Sea (Vo-
lodchenko, 1955). Intertidal to 270 m, rela-
tively uncommon in California. Usually found
in rocky areas and on floating docks in bays.
Feeds upon the bryozoans Alcyonidium ge-
latinosum, Alcyonidium hirsutum, Alcyoni-
dium polyoum, Alcyonidium spp., Callopora
dumerili, Cryptosula pallasiana, Electra pilo-
sa, Flustrellidra hispida, Membranipora, Po-
rella concinna, Schizoporella unicornis, Smit-
tina reticulata (Barrett, 1969; Bleakney &
Saunders, 1978; Bruce et al., 1963; Jaeckel,
1952; Meyer, 1971; Miller, 1961; Morse,
1968; Swennen, 1961; Thompson, 1964;
Thompson & Brown, 1976).
Remarks: This is a highly variable species
on the Atlantic coast of North America and
in Europe. Further investigation may well
show several California species of Acantho-
doris to be junior synonyms of A. pilosa (viz.
A. hudsoni, A. nanaimoensis, and A. rhodo-
ceras).
Dr. T. E. Thompson (personal communi-
cation) states that Gregory H. Brown has ex-
amined the types of Doris subquadrata, and
found that they present no significant differ-
ences from A. pilosa.
Acanthodoris rhodoceras Cockerell
in Cockerell & Eliot, 1905
Acanthodoris rhodoceras Cockerell in Cock-
erell & Eliot, 1905: 38. MacFarland, 1925:
55-60. MacFarland, 1926: pl. 2, figs. 3-4;
pl. 3, fig. 4. Marcus, 1961: 26-27, 58, pl.
5, figs. 84-88. Lance, 1961: 67. Paine,
1963a: 4. Farmer & Collier, 1963: 62.
Steinberg, 1963b: 70. Paine, 1964: 385.
Paine, 1965: 607. Marcus 8 Marcus,
1967a: 202. Sphon 8 Lance, 1968: 76.
Roller & Long, 1969: 425. Roller, 1970b:
482. Gosliner 8 Williams, 1970: 176. Ab-
bott, 1974: 362. Haderlie & Donat, 1978:
60.
Acanthodoris rhodocera Cockerell 8 Eliot.
O'Donoghue, 1926: 219 (lapsus).
Acanthodoris sp. MacGinitie 8 MacGinitie,
1949: 363.
Type-specimens: not listed.
Foot oval, bluntly rounded and bilabiate
anteriorly and rounded posteriorly. Rhino-
phores bear about 13-17 lamellae, translu-
cent grayish-white, clavus yellowish to brown,
tipped with white, occasionally with a subter-
minal black band. Branchial plumes 5, bi- and
tripinnate, translucent grayish-white, tipped
with yellow to brown. Posterior-most bran-
chial plume on either side bears an accessory
branch nearly as large as an independent
plume. General ground color translucent
grayish-white; dorsum may be finely sprin-
CALIFORNIA NUDIBRANCHS 137
kled with tiny flecks of yellow and dark brown
to black. Dorsal papillae tipped with dark
brown to black. Mantle margin with yellow
band and with a black band just inside yellow
band. One or both of these marginal bands
may be discontinuous or indistinct in some
specimens. T.L.: 15 mm.
Radular formula of specimens examined
28-31(5.1.0.1.5), MacFarland (1925: 57) re-
ports 32-36(5-6.1.0.1.5-6). Laterals (Fig. 34f)
bear 3-6 denticles on cusp.
Type-locality: San Pedro, Los Angeles Co.,
California.
Range and habitat: Dillon Beach, Marin
Co., California (Marcus, 1961), to Punta
Mesquite, Baja California, Mexico (Farmer &
Collier, 1963). Intertidal to 25 m. Usually found
on floating docks in bays and in rocky inter-
tidal areas, frequently in association with an
unidentified ascidian upon which it may feed.
Aegires Loven, 1844
Body rather limaciform and somewhat rig-
id, dorsum with numerous tubercles. Margin
of dorsum indistinct and tuberculate; frontal
margin narrow and tuberculate. Labial tenta-
cles small and lobiform. Rhinophores smooth
and retractile into oblique, tuberculate
sheaths. Branchial plumes few, retractile, tri-
pinnate, arranged around the anus, each pro-
tected by a peculiar individual lobe.
Armature of labial disc consists of a band
of minute rods close to buccal aperture be-
low; large mandible present above. Radular
formula 0.n.0.n.0, laterals numerous and uni-
formly hooked.
Penis armed with hooks.
Type-species: Aegires punctilucens (Or-
bigny, 1837).
Aegires albopunctatus MacFarland, 1905
Aegires albopunctatus MacFarland, 1905: 45.
Cockerell & Eliot, 1905: 32, 44-45. Mac-
Farland, 1906: 133, pl. 19, figs. 41-44.
Guernsey, 1912: 75, fig. 39F. O’Dono-
ghue, 1926: 213. O'Donoghue, 1927a: 7-
9, pl. 1, figs. 7-9. O'Donoghue, 1927b: 95-
96. Costello, 1938: tabs. 1-5. Marcus,
1961: 20-21, pl. 4, figs. 65-66. Lance,
1961: 66. Paine, 1963a: 4. Farmer 8 Col-
lier, 1963: 62. Steinberg, 1963b: 70. Paine,
1964: 385. MacFarland, 1966: 101-103, pl.
18, figs. 5-8; pl. 31, figs. 1-5. Lance, 1966:
73-76. Farmer, 1967: 341. Sphon 4 Lance,
1968: 76. Haderlie, 1968: 339. Lance,
1969: 34, 37. Long, 1969c: 232. Roller 8
Long, 1969: 425. Haderlie, 1969: tabs. 1,
2. Turner et al., 1969: 132. Gosliner & Wil-
liams, 1970: 176. Bernard, 1970: 85. North,
1971757. Keen, 1971.-829, pl 21: #9; 1.
Bertsch et al., 1972: 304. Harris, 1973: 281,
289. Abbott, 1974: 358. Haderlie et al.,
1974: tab. 4. Michel, 1976: 4b, fig. 1. Lam-
bert, 1976: 294, 296. Nybakken, 1978: 135,
144. Haderlie 8 Donat, 1978: 52, 60.
Bertsch, 1980: 223-224, figs. 1-6.
Cregires albopunctatus MacFarland. Berry,
1907: 35 (lapsus).
Aegires albopunctata. O'Donoghue, 1927b:
77 (lapsus).
Aegirus (Aegires) albopunctatus Mac-
Farland. Smith & Gordon, 1948: 180.
Type-specimens: U. S. Nat. Mus., no.
181282.
Dorsum densely covered everywhere with
large, cylindrical tubercles which may be
slightly expanded apically, and are arranged
in irregular longitudinal rows; they may be
somewhat reduced in occasional specimens,
especially in small individuals from the rocky
intertidal. Foot narrow and elongate, distinct-
ly set off from sides of body. Labial tentacles
small and lobe-like. Rhinophores completely
smooth, cylindrical, and truncated apically;
white to lemon yellow with small, opaque
white flecks. Borders of rhinophore sheaths
bear 5-6 rounded tubercles which are higher
on lateral surface than on medial surface.
Branchial plumes 3, non-retractile, white to
cream. A large irregularly tuberculate lobe
occurs lateral of branchial plumes, and a sim-
ilar lobe occurs anterior of median branchial
plume. General ground color white to pale
yellowish-white or cream, dorsum and sides
of body generally sprinkled with small, irreg-
ularly scattered dark brown to black flecks,
though these may be lacking in some individ-
uals. Dorsum also bears small, opaque white
flecks scattered between the tubercles. T.L.:
13 mm.
Radular formula of specimens examined
17-18(22-24.0.22-24), MacFarland (1966:
103) reports 16-22(17.0.17). Laterals (Fig. 51)
hamate.
Type-locality: Monterey Bay, California.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976), to Islas
Coronados, Baja California, Mexico (Lance,
1961). Intertidal to 30 m. Most common on
docks and pilings in bays, occasional in rocky
intertidal and subtidal. Bertsch (1980: 223)
reports that it feeds upon the sponge Leucilla
nuttingi. It is also frequently found upon the
138 MCDONALD
sponge Leucosolenia eleanor which it prob-
ably also eats.
Remarks: Smith & Gordon (1948: 180) list
one of the localities at which A. albopuncta-
tus has been found as: “extreme low water,
on stones, at Santa Cruz (Cooper).” This ref-
erence appears to be taken from Cooper
(1870: 56) who cited Doris albopunctata (an
old name for Doriopsilla albopunctata) from:
“Santa Cruz, rare on stone, extreme l.w..”'
Aeolidia Cuvier, 1797
Body aeolidiform, broadest anteriorly and
tapered posteriorly. Wide foot rounded and
bilabiate anteriorly, lateral margins extend
beyond body margins; anterior foot corners
slightly elongate. Cerata flattened and lan-
ceolate, pointed apically and broadest near
base; they occur in numerous transverse
rows, with little space between rows. Cni-
dosacs present. Rhinophores smooth and
non-retractile.
Masticatory border of mandibles non-den-
ticulate. Radula uniseriate, with pectinate
teeth.
Penis unarmed. Anus cleioproct.
See ICZN (1966: opinion 779), Aeolidia
placed on Official List.
Type-species: Aeolidia papillosa (Lin-
naeus, 1761).
Aeolidia papillosa (Linnaeus, 1761)
Doris spinis mollibus hirsuta Baster, 1760: 81,
pl. 10, fig. 1.
Limax papillosus Linnaeus, 1761: 508, no.
2093. Linnaeus, 1767: 1082. Mohr, 1786:
115. Cuvier, 1817: 16. Bruguiere & La-
marck, 1830: 114. Lemche, 1938: 1.
Lemche, 1964b: 116, 117.
Doris bodoensis Gunnerus, 1770: 170, un-
numbered pl., figs. 11-16. Cuvier, 1817: 16.
Bruguiere 8 Lamarck, 1830: 114. (Non)
Möller, 1842: 5.
Doris papillosa Múller, 1776: 229, no. 2775.
Fabricius, 1780: 345. Stewart, 1801-1802:
336. Montagu, 1815: 16, pl. 4, fig. 3. Cu-
vier, 1817: 16, 25. Bruguiere 8 Lamarck,
1830: 114, 115. Lemche, 1964b: 116. (Non)
Kelaart, 1858: 101. (Non) Pease, 1860: 30.
Tritonia papillosa (Linnaeus). Pennant, 1777:
81. Bosc, 1802: 91. Jameson, 1809: 556.
Fleming, 1820: 619. Bosc, 1830: 105.
Doris vermigera Turton, 1807: 132.
Eolis симеп Lamarck, 1819: 302. Start, 1828:
69. Bouchard-Chantereaux, 1835: 33. La-
marck, 1936: 450. Thompson, 1844a: 250.
Eolida papillosa Fleming, 1823a: 295. Flem-
ing, 1828: 285.
Eolidia bodoensis (Gunnerus). Cuvier, 1834:
122.
Eolidia papillosa (Linnaeus). Cuvier, 1834:
122. Johnston, 1835: 376-378, text fig. 35.
Sars, 1835: 68. Johnston, 1838b: 118-120.
Thompson, 1840: 89. Kroyer, 1847: 116.
Gray, 1857: 225. Herdman & Clubb, 1892:
146. Herdman et al., 1896: 446. Daro,
1969: 136.
Eolis Cuverii Lamarck. Bouchard-Chanter-
eaux, 1835: 129-132 (lapsus).
Eolidia cuvierii. Blainville, 1825: 486. Bru-
guiere & Lamarck, 1830: 115. Johnston,
1838b: 120-121, pl. 3, figs. 9-11.
Eolida zetlandica Forbes & Goodsir, 1839:
647.
Eolis rosea Alder & Hancock, 1842: 34. Al-
der, 1850: 114.
Eolis obtusalis Alder & Hancock, 1842: 34.
Alder, 1850: 114. Collingwood, 1859: 466.
Aeolis murrayana MacGillivray, 1843: 193.
Aeolis lesliana MacGillivray, 1843: 194.
Aeolis papillosa (Linnaeus). MacGillivray
1843: 192. Loven, 1846: 139. Sars, 1850:
193. Adams & Adams, 1854: 73, pl. 65, fig.
8. Norman, 1860: 7243. Meyer & Möbius,
1865: 29-32, pl. 2. Hogg, 1868: pl. 10.
Mörch, 1868: 205. Gould, 1870: 238-240.
Verrill, 1873: 486, 495, 499, 666. Verrill,
1874b: 132. Sars, 1878: 318-319 ff, pl. XV,
fig. 8. Collin, 1884: 50. Locard, 1886: 43-
44. Fischer, 1887: 532, 540, text figs. 290,
298. Garstang, 1890: 432-433. Norman,
1890: 81. Hecht, 1893: XIII. Garstang
1894: 226. Tregelles, 1896: 222. Mansel-
Pleydell, 1898: 27. Nichols, 1900: 587. Al-
len & Todd, 1900: 165 ff. Whiteaves, 1901:
204-205. Grosvenor, 1903: 468 ff. Todd,
1903: 544, 556. Farran, 1904: 6. Odhner,
1907: 77. Eliot, 1907: 327. Colgan, 1908:
108. Colgan, 1911: 23. Retzius, 1914: 13-
15, pl. 4, figs. 1-15. Colgan, 1914: 177-
178. Chumley, 1918: 74, 169. Bardarson,
1919: 72. Bardarson, 1920: 110. Loyning,
1922: 2. Odhner, 1926a: 25. Derjugin,
1928: 321. Lonnberg, 1931: 19-20. Ellis,
1933: 189. Linke, 1937: 232-236, text figs.
1-6. Volodchenko, 1941: 60. Kepner, 1943:
305. Homans 8 Needler, 1944: 37, 38.
Schafer, 1950: 9-14, text figs. 1-5. Fisch-
er, 1950: 234 ff. Schonenberg, 1969: 288.
(Non) Hagg, 1905: 104.
Eolis papillosa (Linnaeus). Thompson, 1844:
250. Hancock 8 Embleton, 1845: 2 ff, pl.
1, figs. 1, 5-11; pl. 2, figs. 1-2, 4-5, 7, 9,
CALIFORNIA NUDIBRANCHS 139
12, 14; pl. 3, figs. 5-7; pl. 4, figs. 4-9; pl.
5, figs. 7-14, 16. Alder & Hancock, 1847:
fam. 3, pl. 8. Alder, 1850: 105, 114. Forbes
8 Hanley, 1851-1852: 590. Alder 8 Han-
cock, 1852: fam. 3, pl. 9. Hancock 8 Em-
bleton, 1852: 236, 237. Gosse, 1853: 12,
16. Dalyell, 1853: 314-317, pl. 45, figs. 23-
27. Gosse, 1854: 105, 125. Byerley, 1854:
45. Alder 8 Hancock, 1855: 3, 23, 26, 31,
32, 48. Thompson, 1856: 277. Colling-
wood, 1859: 466. E. Wright, 1859: 87. T.
Wright, 1859: 39. Collingwood, 1860: 199,
202. Collingwood, 1861: 114. Kinahan,
1861: 31. Wright, 1863: 53. Mcintosh,
1865: 392. Hogg, 1868: 103. Robertson,
1868: 206. Jeffreys, 1869: 37-38, pl. 2, fig.
3. Gould, 1870: pl. 18, figs. 258, 261. Sau-
vage, 1873: 25, 34-35. Mcintosh, 1874:
428. Mcintosh, 1875: 85. Friele & Hansen,
1876: 74. Leslie & Herdman, 1881: 310.
Verrill, 1882a: 340. Herdman, 1886: 273,
277. Higgins, 1886: 26. Herdman 8 Clubb,
1889: 232, pl. 12, figs. 4-6. Garstang,
1889: 190-191. Hecht, 1896: 543 ff. Gam-
ble, 1896: 132. Boutan, 1898: XXXVII-XLII.
Cooke, 1899: 60. Liverpool Mar. Biol.
Comm., 1899: 55. Knight, 1901: 206. Con-
chol. Soc., 1901: 24. Grosvenor, 1903: 462.
Cuenot, 1907: 87, 94. Fleure 8 Walton,
1907: 219. Elmhirst, 1908: 227. Pelseneer,
1911: 55-57, pl. 16, figs. 5-14. Colgan,
1913: 165-166. Farran, 1913: 6. Farran,
1915: 10, 67. Engel, 1925: 33. Pelseneer,
1928: 179. White, 1938: 16. Arvy, 1950:
159. Humes & Stock, 1973: 167, 168.
Eolidia cuvieri (Lamarck). Leach, 1852: 23, pl.
Mg э. Gray, 1857: 225.
Eolis farinacea Stimpson ex Gould Ms,
1853: 25. Gould, 1870: pl. 18, figs. 257,
259, 260, 262, 263.
Eolis plumata Dalyell, 1853: 300, pl. 44, figs.
1-2.
Aeolis Cuvieri (Lamarck). Adams & Adams,
1854: 73. Chenu, 1859: 411, fig. 3070. Lo-
card, 1886: 44.
Eolis cuvierii (Lamarck). Thompson, 1856:
РТА
Aeolis farinacea. Stimpson, 1862: 4.
Eolis papillata. Baudelot, 1863: 212-213 ff,
pl. 5, figs. 5-6.
Aeolidia serotina Bergh, 1873a: 618-620, pl.
9, figs. 14-17; pl. 10, figs. 4-12. Bergh,
1879b: 74. Bergh, 1879c: 130. Bergh,
1892: 1019 (27). Bergh, 1898: 541-544,
pl. 31, figs. 26-31. Eliot, 1907: 327, 351-
352. Odhner, 1921: 225. Odhner, 1926a:
25, 29. Powell, 1951: 54.
Aeolidia papillosa (Linnaeus). Bergh, 1860:
309-331, pl. 8. Bergh, 1868: 174, 184,
200-203, pl. 1, figs. 1-20. Dall, 1870: 249.
Bergh, 1874: 396. Bergh, 1877c: 822.
Bergh, 1879b: 74-77, pl. 1, figs. 1-6.
Bergh, 1879c: 130-131, pl. 1, figs. 1-6.
Trinchese, 1881: 33. Dall, 1884: 341.
Bergh, 1890a: 28. Bergh, 1892: 1019 (27).
Hecht, 1893: XVI. Lundbeck, 1893: 175.
Bergh, 1894: 127. Bergh, 1898: 540-541.
Cooke, 1899: 65. Beaumont, 1900: 833.
Vayssiere, 1901: 301. Johansen, 1902:
387. Mar. Biol. Assoc., 1904: 278. Cock-
erell & Eliot, 1905: 32. Eliot, 1907: 327, 352.
Walton, 1908: 228-229. Balch, 1909: 37.
Раггап, 1909: 3=4. Eliot; 1910: 5, 7, 11),
36, 50-59, 175, text fig. 7. Sumner et al.,
1913: 705. Walton, 1913: 110. Johnson,
1915: 162. Evans & Evans, 1917: 109.
O'Donoghue, 1921: 199-201, pl. 3 (9), fig.
31. Odhner, 1921: 225. O'Donoghue &
O'Donoghue, 1922: 135-136, pl. 3, fig. 1.
O'Donoghue, 1922b: 165. O'Donoghue,
1922d: 141. Loyning, 1922: 70-73, 94, text
figs. 51-54, pl. 4, figs. 14-15. Elmhirst,
1922: 40. Iredale 8 O'Donoghue, 1923:
201. Odhner, 1926a: 29. Odhner, 1926b:
28. O'Donoghue, 1926: 232. Jutting, 1927:
LXXXVIII. Loyning, 1927: 262. Carter,
1927: 5 ff. Cuenot, 1927: 261-262. Carter,
1928: 97 ff. O'Donoghue, 1929: 742. Rus-
sell, 1929: 210 ff, text figs. 2-3, pls. 6, 8—
10. Lemche, 1929: 18-19. Carter, 1929:
561. Mar. Biol. Assoc., 1931: 268. Rous-
seau, 1931: 954-956. Winckworth, 1932:
238. Nobre, 1932: 28, pl. 9, fig. 2. Monod
8 Dollfus, 1932: 138 ff. Renouf, 1934: 400.
Rousseau, 1934: 677. Chambers, 1934:
602 ff. Johnson, 1934: 154. MacGinitie,
1935: 740. Baba, 1935b: 116, 121-123, pl.
8, figs. 3-5. Leigh-Sharpe, 1935: 48. Nobre,
1936: 20. Baba, 1937a: 199. Baba, 1937b:
335. Millott, 1937: 406 ff. Moore, 1937: 11,
194. Hewatt, 1937: 200. White, 1938: 16,
17. Lemche, 1938: 28 ff. Graham, 1938:
267 ff, text fig. 11. Odhner, 1939: 84-85,
text figs. 49-50. Cockerell, 1940: 503.
Russell, 1942: 80-81. McMillan, 1942a:
324. McMillan, 1942b: 327. McMillan,
1944: 162. Jutting, 1947: 65. Pruvot-Fol,
1951а: 54. Cornet 8 Marche-Marchad,
1951: 37-38. Stehouwer, 1952: 161-170.
Jaeckel, 1952: 24 ff. Stock, 1952: 58, 59.
Braams 8 Geelen, 1953: 242 ff. LaRocque,
1953: 248. Williams, 1954: 107. Pruvot-Fol,
1954: 426-427, text fig. 167. Graham,
1955: 153. Franzén, 1955: 428. Mar. Biol.
140 MCDONALD
Assoc., 1957: 318. Baba, 1957: 9. Aboul-
Ela, 1959: 442. Swennen, 1959: 58.
Thompson, 1960b: 125. Haaften 8 Ver-
wey, 1960: 493 ff. Pruvot-Fol, 1960: 160,
194. Swennen, 1961: 223-225. Miller,
1961: 98 ff. Thompson, 1961: 237. Mar-
cus, 1961: 54 ff, pl. 10, figs. 193-195.
Robson, 1961: 685-690. Lance, 1961: 68.
Buznikov & Manukhin, 1961: 226. Kohn,
1961: 299. McLean, 1962: 111. Rogin-
skaya, 1962a: 88, 104, 106, figs. 5.6-5.7.
Roginskaya, 1962b: 209-211, 213, 214.
Sakharov, 1962: 311. Miller, 1962: 562-
563. Roginskaya, 1963: 1179. Bruce et al.,
1963: 209. Paine, 1963a: 4. Paine, 1963b:
71. Steinberg, 1963b: 72. de Vries, 1963:
137 ff. Thompson, 1964: 279 ff. Gonor,
1965: 231. Baba 8 Hamatani, 1965: 107.
Ghiselin, 1965: 345 ff. Sakharov, 1966:
957. Edmunds, 1966: 28 ff. Ross, 1967:
313. Hurst, 1967: 255 ff, text figs. 10a-d,
pl. 29, fig. 13, pl. 33, fig. 29; fig. 24-10.
Wolter 1967: 277 ff, text figs. 6, 19, 34—
36. Franz, 1968: 11. Mauzey et al., 1968:
617. Beeman, 1968b: 268. Bebbington 8
Thompson, 1968: 10. Sphon 8 Lance,
1968: 76. Streble, 1968: 357 ff. Thompson
& Hinton, 1968: pl. 2, figs. A-C. Burn,
1968: 91. Rosin, 1969: 74, 76. Tardy, 1969:
34. Barrett, 1969: 68-69. Daro, 1969: 147.
Roller £ Long, 1969: 425. Loveland et al.,
1969: 419. Franz, 1970: 172 ff. Hughes,
1970b3 584; 82: text 1655 78 29,
Schmekel, 1970: 158. Rosin, 1970: 47.
Tardy, 1970: 352. Bernard, 1970: 85. Gos-
liner & Williams, 1970: 176. Daro, 1970:
168-169. Waters, 1971a: 32. Waters,
1971b: 33. Harris, 1971a: 79-80, 81. Har-
ris, 1971b: 699. Harris, 1971c: 22. Meyer,
1971: 148-149. Keen, 1971: 839, fig. 2385.
Kress, 1971: 326, 342-345, text fig. 14.
Salvini-Plawen, 1972: 394, 396. Korobtsov
8 Sakharov, 1972: 198. Bertsch et al.,
1972: 304. Holleman, 1972a: 60. Humes &
Stock, 1973: 167, 168. Thompson 4 Beb-
bington, 1973: 148, 149. Navoni, 1973:
1334, 1352. Waters, 1973: 174-192.
Morse, 1973: 1334. Harris, 1973: 217 ff.
Mariscal, 1974: 163. Roginskaya, 1974b:
998, text fig. 1. Bertsch, 1974: 3-6, figs.
1-4. Nybakken, 1974: 371. Hinegardner,
1974: 452. Edmunds et al., 1974: 939-947.
Abbott, 1974: 380, pl. 16, fig. 4470. Ro-
zsa, 1974: 7, 8. Rozsa, 1975: 3. Belcik,
1975: 2270, 21 Ta ‘Franz. 19/99: -81, 102.
Zack, 1975a: 271. Clark, 1975: 37. Kalker
8 Schmekel, 1976: 42 ff. Roginskaya,
1976: 26. Edmunds et al., 1976: 65 ff. Har-
ris, 1976: 301. Lambert, 1976: 296. Mich-
el, 1976: 48, fig. 10. Thompson, 1976a: 36,
39, 50, text figs. 21b-h. Thompson, 1976b:
64, 71, 77, 80, 87, fig. 1. Thompson &
Brown, 1976: 160, fig. 85. Barletta 8 Me-
lone, 1976: 204, 205, 230-231, 233. Bar-
letta & Melone, 1977: 320-324, text figs.
1-2. Hoc 8 Fricke, 1977: 362, text fig. 9b.
Dunn, 1977: 70. Garlo, 1977: 24, 26, 27.
Moreteau, 1977: 369 ff. Tardy, 1977: 255-
258. Nybakken, 1978: 133, 135. Bonar,
1978: 187. McDonald & Nybakken, 1978:
115. Haderlie & Donat, 1978: 52, 60. Day
8 Harris, 1978: 105 ff. Howe 8 Harris,
1978: 551 ff. Brewer, 1978: 8. Williams,
1978: 58. Harris 8 Howe, 1979: 138 ff.
Russo, 1979: 46, 47. Boyle 8 Cohen, 1980:
2130. Cockburn & Reid, 1980: 275 ff. Gos-
liner, 1980: 72. Porter 8 Rivera, 1980: 217
ff, figs. 1, 2, 6-8, 11-12. Williams, 1980:
99 ff, text fig. 1b.
Aeolidia papillosa var. pacifica Bergh, 1879b:
75-77, pl. 1, figs. 1-6. Bergh, 1879c: 131-
133, pl. 1, figs. 1-6.
Aeolidia herculea Bergh, 1894: 128-129, pl.
1, figs. 8-12. Bergh, 1898: 540. Cockerell
8 Eliot, 1905: 32. O'Donoghue, 1922d: 141.
Marcus, 1961: 54.
Aeolidiella papillosa. Balch 1909: 36.
Eolis papillosa var. albina Dautzenberg 4 Du-
rouchoux, 1913: 8.
Aeolidida papillosa. O'Donoghue, 1921: 199.
Jaeckel, 1952: 45 (lapsus).
Aeolidia hercules Bergh. Smith 8 Gordon,
1948: 181 (lapsus).
Aeolidia papillosa var. serotina Bergh. Mar-
cus, 1959: 3 ff, 81-84 ff, figs. 191-196.
Marcus, 1961: 56.
Aeolidia pappilosa (Linné). Almaca, 1960:
209-211 (lapsus).
Aeolidia papillosa herculea Bergh. Mac-
Farland, 1966: 370-373, pl. 72, figs. 1-8.
Aaeolidia papillosa. Daro, 1969: 139 (lap-
sus).
Aerolidia papillosa. Platts, 1973: 384 (lap-
sus).
Aeolodia papillosa. Kalker 8 Schmekel, 1976:
41 (lapsus).
Type-specimens: not listed.
Body tapered to a short, broad, pointed tail
which extends posteriorly just beyond the
cerata. Oral tentacles slightly shorter than
rhinophores, same color as body. Rhino-
phores usually a slightly darker shade of body
color. Cerata occur in numerous (19-24 or
more) transverse rows, usually about one
third of the cerata rows occur anterior of the
CALIFORNIA NUDIBRANCHS 141
rhinophores. Cardiac region lacks cerata, re-
mainder of dorsum so densely covered with
cerata as to make individual rows almost in-
distinguishable. Cerata recumbent when ani-
mal is actively crawling. Color of body and
cerata highly variable, most commonly dirty
white, but individuals which are dull gray,
mauve, rose, or pink have been collected.
Area between oral tentacles and rhinophores
with triangular to crescentic patch of encrust-
ing white to cream, this patch may be pro-
longed onto dorsal surface of oral tentacles.
Area of similar pigment occurs posterior of
rhinophores and extends to cardiac region.
Core of cerata usually brownish to mauve or
umber, but quite variable. T.L.: 40 mm.
Anus located dorso-laterally on right, be-
tween the ninth and tenth rows of cerata.
Renal pore ventral of first post-anal row of
cerata. Genital aperture on right side, be-
tween fifth to tenth rows of cerata. Penis
conical and unarmed.
Radular formula of specimen examined
23(0.0.1.0.0), with 36-38 denticles on each
tooth (Fig. 94). MacFarland (1966: 371) re-
ports 18(0.1.0), with 38-43 denticles on each
tooth, while Marcus (1961: 54) reports
21(0.1.0), with up to 58 denticles on each
tooth. Masticatory border of mandible
smooth.
Type-locality: ‘Mari Norvegico.”'
Range and habitat: Widely distributed in the
northern hemisphere (Lance, 1961), Atlantic
(Franz, 1975a) and Pacific (Lance, 1961)
coasts of North America; Japan (Baba,
1935b), Europe (Pruvot-Fol, 1954). Intertidal
to 870 m. Most commonly found on docks
and pilings in bays; occasionally found in
rocky intertidal. Feeds on various species of
sea anemones: ‘Actinia, Actinia equina, Ac-
tinothoe sphyrodeta, Aiptasia couchi, ‘Ап-
thea,' Anemonia sulcata, Anthopleura arte-
misia, Anthopleura balli, Anthopleura
elegantissima, Anthopleura xanthogrammi-
ca, Cereus pedunculatus, Corynactis califor-
nica, Corynactis viridis, Diadumene cincta,
Diadumene luciae, Epiactis prolifera, Me-
таит dianthus, ‘Metridium marginatus,’
Metridium marginatum, Metridium senile, Sa-
gartia elegans, Sagartia troglodytes, Sagar-
tiogeton undata, Stomphia coccinea, Tealia
coriacea, Tealia crassicornis, Tealia felina,
Tealiopsis stella (Bertsch et al., 1972; Bruce
et al., 1963; Clark, 1975; Day & Harris, 1978;
Dunn, 1977; Eliot, 1910; Harris, 1971b; Har-
ris, 1973; Howe 8 Harris, 1978; Harris 8
Howe, 1979; Kalker 8 Schmekel, 1976;
McMillan, 1942b; Miller, 1961; Robson, 1961;
Russell, 1942; Swennen, 1961; Thompson,
1964; Thompson 8 Brown, 1976; Waters,
1973). In addition, Miller (1961) and Salvini-
Plawen (1972) list the hydroid Tubularia in-
divisa as a food item. Jaeckel (1952) also lists
worms, tunicates, and young Mytilus as prey.
Alder 8 Hancock (1855) found several small
specimens of the common mussel in the
stomach. Mcintosh (in Grosvenor, 1903)
found that the cod (Gadus callarius) would
eat Aeolidia papillosa. Homans 8 Needler
(1944: 37) found the stomachs of a fairly large
sample of young haddock (Melanogrammus
aeglifinus) filled exclusively with A. papillosa.
Aeolidiella Bergh, 1867
Body aeolidiform, tapered posteriorly. Foot
somewhat rounded anteriorly, anterior foot
corners somewhat produced into triangular
lobes. Cerata numerous, cylindrical, slightly
clavate, and decumbent, arranged in slightly
oblique rows which are very close together;
the cerata bear cnidosacs. Rhinophores non-
retractile, smooth or with a few shallow,
oblique lamellae.
Masticatory border of mandibles smooth or
striate, nondenticulate. Radula uniseriate,
rachidian teeth rather bilobed, with a some-
what prominent median denticle, with lateral
denticles largest on medial portion of either
lobe.
Penis unarmed. Anus cleioproct.
Type-species: Aeolidiella soemmerringii
(Leuckart, 1828).
Aeolidiella takanosimensis Baba, 1930
Eolis sp. Ijima, 1925: 615, fig. 740B.
Aeolidiella takanosimensis Baba, 1930: 122,
124, text figs. 4a-b, pl. 4, figs. 5a-c. Ko-
mori, 1932: 395-397, text figs. 3-4. Baba,
1949: 20, 111-112, 183-184, text figs.
154-155, pl. XLIX, fig. 167. Baba 8 Ha-
matani, 1952: 10. Abe, 1964: 72-73, pl.
36, fig. 129. Marcus & Marcus, 1967a: 117.
Schmekel, 1968a: 122, 123, 145. Usuki,
1969: 8. Schmekel, 1970: 135, 155, 156,
fig. 16b. Schmekel, 1971: 122, text fig. 2b.
Sphon, 1971a: 368-369. Salvini-Plawen,
1972: 394. Mulliner, 1972a: 38. Mulliner,
1972b: 4, figs. Abbott, 1974: 380. Keen 4
Coan, 1975: 45. Ferreira 8 Bertsch, 1975:
329, fig. 19. McDonald & Nybakken, 1978:
115. Bertsch, 1979a: 59. Baba, 1979a: 12-
18, figs. 1-6. Baba, 1979b: 7.
Eolidina (Eolidina) takanosimensis (Baba).
Baba, 1937b: 336. Baba, 1938: 3.
142 MCDONALD
Aeolidiella takanosimiensis Baba. Gosliner,
1980: 63-64, 72, text figs. 16A-B (lapsus).
(Non) Aeolidiella (?takanosimensis К. Baba).
Risbec, 1956: 31-32, pl. 21, figs. 110-115.
Marcus, 1961: 56 (=Spurilla risbeci Mar-
cus, 1961).
Type-specimens: missing (Baba, personal
communication).
Foot elongate, rather rounded and bila-
biate anteriorly, tapered posteriorly to mod-
erate, pointed tail. Oral tentacles long, cylin-
drical, and slightly tapered to blunt tips,
vermilion distally with an opaque white tip.
Rhinophores smooth, cylindrical, and ta-
pered to bluntly pointed tips; colored as oral
tentacles. Cerata arranged in about 28 or
more oblique rows dorso-laterally on either
side of dorsum leaving dorsum free antero-
medially, posterior groups of cerata less sep-
arated medially. General ground color trans-
lucent grayish-white with a vermilion tinge. A
broad band of vermilion occurs dorso-later-
ally on either side, between base of oral ten-
tacles and base of rhinophores. Dorsum ver-
milion in cardiac region; with about 3 or more
rather symmetrical, white blotches dorso-me-
dially. Each ceras bears a subapical band of
opaque white below light vermilion tip; core
light brown. T.L.: 25 mm.
Anus dorso-lateral on right, between ninth
and tenth rows of cerata. Genital aperture on
right side, ventral of third row of cerata.
Baba (1930: 122) reports the radular for-
mula 16(0.1.0), Baba (1979: 14) reports 12-
20(0.1.0), and Gosliner (1980: 63) reports 15-
17(0.1.0), while Ferreira 8 Bertsch (1975: 329)
report 32(0.1.0). Baba (1930: 122) reports 24—
34 denticles on either side of median denticle
of the rachidian tooth, Baba (1979a: 14) re-
ports 15-20 denticles, and Gosliner (1980:
63) reports 16-19 denticles, while Ferreira 8
Bertsch (1975: 329) report approximately 67
denticles per tooth. Masticatory border of
mandibles smooth.
Type-locality: Takanosima, Tateyama Bay,
Japan.
Range and habitat: Palos Verdes Penin-
sula, Los Angeles Co., California (Sphon,
1971a), to San Diego, San Diego Co., Cali-
fornia (Sphon, 1971a); Bahía San Marte, Baja
California, Mexico (Ferreira 8 Bertsch, 1975);
Japan (Baba, 1930); Gulf of Naples, Italy
(Schmekel, 1971); Hawaii (Gosliner, 1980);
Fanning Atoll, Line Islands (Gosliner, 1980).
Usually found on floating docks in bays.
Schmekel (1968a: 145) and Salvini-Plawen
(1972: 394) report that it feeds upon the sea
anemone Sagartia Sp.
Aldisa Bergh, 1878
Body doridiform, dorsum covered with nu-
merous small papillae. Labial tentacles small
and auriform or tuberculiform. Rhinophores
perfoliate and retractile into sheaths. Bran-
chial plumes retractile, uni- to tripinnate, ar-
ranged in a circle around anus.
Labial disc unarmed. Radular formula
0.n.0.n.0, laterals quite elongate and bear nu-
merous fine denticles distally, appearing
whisk-like.
Penis armed with small hooks.
Type-species: Aldisa zetlandica (Alder &
Hancock, 1854).
Aldisa sanguinea (Cooper, 1863)
Doris (Asteronotus) sanguinea Cooper,
1863a: 204. Carpenter, 1864: 608. Stein-
berg, 1961: 59.
Doris sanguinea Cooper. Cooper, 1867: 14.
Abraham, 1877: 203. Orcutt, 1885: 545.
Kelsey, 1907: 39.
Asteronotus (?) sanguinea (Cooper). Bergh,
1890b: 917.
Asteronotus sanguineus (Cooper). Bergh,
1891: 137. Pruvot-Fol, 1954: 269.
Asteronotus (?) sanguineus (Cooper). Bergh,
1892: 1103 (111).
Aldisa sanguinea (Cooper). MacFarland,
1905: 42. MacFarland, 1906: 123-125, pl.
18, figs. 25-26; pl. 21, figs. 112, 114; pl.
24, fig. 7. Kelsey, 1907: 33. O'Donoghue,
1926: 206, 209. O'Donoghue, 1927a: 10.
O'Donoghue, 1927b: 84-85, pl. 1, figs. 16-
19. Boone, 1929: 38. Costello, 1938: tabs.
1-3, 5. Smith & Gordon, 1948: 181. Pru-
vot-Fol, 1954: 269. Steinberg, 1961: 59.
Marcus, 1961: 16, 57, pl. 3, figs. 50-53.
Lance, 1961: 66. Cook, 1962: 196. Stein-
berg, 1963b: 70. MacFarland, 1966: 169-
171,,-pl: 25,. fig: 8; pl. -29; figs take plese.
figs. 17-22. Sphon & Lance, 1968: 76.
Long, 1969c: 232. Fournier, 1969: 74.
Roller, 1969a: 280-281, text fig. 1. Roller
& Long, 1969: 425. Robilliard & Baba,
1972: 409-413, fig. 1A. Abbott, 1974: 354,
text fig. 4236. Ferreira & Bertsch, 1975:
327-328, figs. 11-14. Belcik, 1975: 276.
Keen & Coan, 1975: 43. Bloom, 1976: 293,
295. Nybakken, 1978: 135. McDonald &
Nybakken, 1978: 110, 112. (Non) Farmer
& Collier, 1963: 62. (=Thordisa bimaculata
Lance, 1966.)
CALIFORNIA NUDIBRANCHS 143
Type-specimens: not listed; Geological
survey coll. (Cooper, 1863a).
Foot oval, bluntly rounded and bilabiate
anteriorly, slightly more acutely rounded pos-
teriorly. Labial tentacles short and auriform,
with a distinct external groove. Rhinophores
reddish, with 12-15 lamellae. Branchial
plumes 8-10, unipinnate, light red to crim-
son. General ground color light red to dark
crimson, occasionally yellow-orange to yel-
low-green; dorsum sprinkled with very mi-
nute dark brown to black dots; with 1-2 larg-
er black spots on median line of dorsum, one
just posterior of rhinophores and the other
just anterior of branchial plumes, one or both
spots may be absent, especially in small in-
dividuals. Laterally on either side of dorsum,
about two thirds of the distance from anterior
end occur (especially in large individuals)
patches of yellow to yellow-green; a similar
patch frequently occurs just posterior of
branchial plumes. T.L.: 17 mm.
Radular formula of specimen examined
55(89-107.0.89-107), MacFarland (1966:
170) reports 70(75-100.0.75-100). Laterals
(Fig. 56) long and rod-like, with numerous fine
denticles distally.
Type-locality: San Diego Bay, California.
Range and habitat: Bodega Bay, Sonoma
Co., California (Marcus, 1961), to San Diego,
San Diego Co., California (Cooper, 1863a);
Punta Prieta, W. Isla Espíritu Santo, Mexico
(Ferreira 8 Bertsch, 1975). Intertidal to 10 m.
Usually found in the rocky intertidal under
rocks. Feeds upon the sponge Hymendesmia
brepha (McDonald & Nybakken, 1978: 110),
also found on Ophlitaspongia pennata (Four-
nier, 1969).
Comments: This species should be closely
compared to Aldisa banyulensis Pruvot-Fol,
1951. Aldisa sanguinea cooperi differs from
A. sanguinea sanguinea by being lemon yel-
low to cadmium orange, and usually less
densely dotted with black points than A. s.
sanguinea. The black spots on the midline of
the dorsum are usually less prominent and
more numerous in A. s. cooperi than in A. s.
sanguinea.
A subspecies has recently been named and
the synonymy for that subspecies is given so
that it may be separated from Aldisa sangui-
nea sanguinea.
Aldisa sanguinea cooperi
Robilliard 8 Baba, 1972
Doris sanguinea Cooper. (Asteronotus.)
Cooper, 1863b: 58.
Doris sanguinea Cooper. Carpenter, 1864:
609.
Aldisa sanguinea (Cooper). Baba, 1940: 103-
104, figs. 1-2. Baba, 1949: 62-63, 150,
text fig. 75, pl. 24, fig. 86. Baba, Hamatani
8 Hisai, 1956: 211, fig. 6. Baba, 1957: 9.
Aldisa sanguinea cooperi Robilliard 8 Baba,
1972: 409-413, figs. 1B, 2-4. Lambert,
1976: 294, 296.
Ancula Lovén, 1846
Body limaciform and smooth, highest just
anterior of branchial plumes. Foot abruptly
rounded anteriorly, rather narrow and elon-
gate, tapered to narrow tail. Labial tentacles
short and lobe-shaped. Rhinophores non-re-
tractile, perfoliate; base of each rhinophore
bears 2 digitiform extra-rhinophoral append-
ages. Branchial plumes 3, non-retractile, bi-
and tripinnate, arranged in semi-circle around
anus. On either side, lateral of branchial
plumes are one to several slightly clavate,
extrabranchial appendages.
Armature of labial disc consists of rows of
separate, imbricated hooks. Radular formula
1.1.0.1.1, lateral large and broad with a den-
ticulate inner margin, marginals much smaller
and subtriangular.
Penis armed with very small hooks. Buccal
crop sessile.
Type-species: Ancula gibbosa (Risso,
1818).
Ancula lentiginosa Farmer in
Farmer 8 Sloan, 1964
Ancula lentiginosa Farmer in Farmer & Sloan,
1964: 148-150, pl. 18, figs. 1-2; text figs.
1-2. Lance, 1966: 78. Sphon & Lance,
1968: 76. Roller & Long, 1969: 425. Keen,
1971: 829. Marcus, 1972b: 299. Abbott,
1974: 364. Nybakken, 1978: 135. Jaeckle,
1981: 240.
Type-specimens: Calif. Acad. Sci., 1Z no.
13.
Labial tentacles short, slender, and blunt.
Rhinophores with 4-7 lamellae, translucent
tan to off-white, with various amounts of red-
dish-brown dots. The 2 extra-rhinophoral ap-
pendages about half as long as rhinophores
and tan to off-white with various amounts of
reddish-brown dots. Branchial plumes 3, tri-
pinnate, tan to off-white with various amounts
of reddish-brown dots. On either side is a sin-
gle extra-branchial appendage which is col-
144 MCDONALD
ored as the extra-rhinophoral appendages.
General ground color tan to off-white, with
various amounts of reddish-brown dots on
dorsum, head, tail, and sides of body; these
dots are usually clustered in groups forming
a pattern. T.L.: 5 mm.
Radular formula of specimen examined
23(1.1.0.1.1); Farmer & Sloan (1964: 148) re-
port 36(1.1.0.1.1). The laterals of the speci-
men examined (Fig. 44b) bear 12-14 denti-
cles, Farmer & Sloan (1964) report 13
denticles. The marginals (Fig. 44a) are rough-
ly triangular and bear a stout hook apically.
Type-locality: Salt water system (pipes) of
Scripps Institution of Oceanography, La Jol-
la, San Diego Co., California.
Range and habitat: Frontier Arts Nature
Reserve, Marin Co., California (Jaeckle,
1981), to La Jolla, San Diego Co., California
(Farmer & Sloan, 1964); also reported from
Bahía de los Angeles, Baja California, Mexi-
co (Lance, 1966). Intertidal and subtidal, usu-
ally in bays on floating docks and pilings.
Quite uncommon, but when found usually oc-
curs in numbers.
Ancula pacifica MacFarland, 1905
Ancula pacifica MacFarland, 1905: 53.
MacFarland, 1906: 148-149, pl. 20, figs.
89-92; pl. 21, figs. 93-96; pl. 30, fig. 23.
Guernsey, 1912: 75, fig. 39G. Hilton, 1919:
34. Odhner, 1926a: 46. O'Donoghue, 1926:
221. Smith & Gordon, 1948: 180. Pruvot-
Fol, 1951a: 28. Marcus, 1961: 28-29, 58,
pl. 6, figs. 97-102. Lance, 1961: 67. Stein-
berg, 1963b: 71. Farmer 8 Sloan, 1964:
148 ff. MacFarland, 1966: 123-124, pl. 21,
fig. 1; pl. 29, figs. 2-3. Sphon 8 Lance,
1968: 82. Long, 1969c: 232. Roller & Long,
1969: 425. Gosliner 8 Williams, 1970: 176.
Behrens, 1971a: 297-298. Robilliard,
1971a: 162-163, 164. Abbott, 1974: 364,
fig. 4336. Nybakken, 1978: 135. McDonald
& Nybakken, 1978: 110, 111.
Type-specimens: U. S. Nat. Mus., no.
181280.
Labial tentacles short, slender, blunt and
slightly flattened. Rhinophores perfoliate with
about 8-10 lamellae, lamellae yellowish, tip
of clavus orange. The 2 extra-rhinophoral ap-
pendages nearly as long as rhinophores;
white with subapical ring of yellow to orange.
Branchial plumes 3, bi- and tripinnate, yel-
lowish-white with orange tips. On either side
are 4-5 or more extra-branchial appendages
which are yellowish-white on proximal two-
thirds and yellow on distal third, with orange
tips. General ground color yellowish-white. A
narrow, orange line extends postero-medially
from between the rhinophores to branchial
plumes and continues from posterior of bran-
chial plumes to tip of tail. A similar orange
line on each side extends posteriorly along
the dorso-lateral margin, from the rhino-
phores to the extra-branchial appendages;
this same line continues between extra-bran-
chial appendages and for a short distance
posterior of them. T.L.: about 10 mm.
Radular formula of specimen examined
32(1.1.0.1.1), MacFarland (1966: 124) and
Marcus (1961: 29) both report 35(1.1.0.1.1).
The laterals (Fig. 43b) of the specimen ex-
amined bear 11-14 denticles; MacFarland
(1966: 124) reports 11-17 denticles on lat-
erals.
Type-locality: Monterey Bay, California.
Range and habitat: San Juan Island, Puget
Sound, Washington (Robilliard, 1971a), to
Point Loma, San Diego Co., California (Lance,
1961). Intertidal and subtidal, more common-
ly found on floating docks and pilings in bays
than on the rocky open coast in California.
Feeds upon the entoproct Barentsia ramosa
(McDonald & Nybakken, 1978: 110).
Remarks: The description of Ancula pacif-
ica in MacFarland (1966: 123-124) agrees al-
most perfectly with that of Ancula cristata in
Alder & Hancock (1847: fam. 1, pl. 25). The
external coloration is identical except for the
yellow-orange line on the dorso-lateral mar-
gin of A. pacifica, which is not mentioned by
Alder & Hancock (1847) for A. cristata. This
line is sometimes so discontinuous in A. pa-
cifica that it is almost absent (Robilliard,
1971a: 162-163), and Herdman & Clubb
(1892: 134) state that the variation in color-
ation of A. cristata “is very great, and the
larger specimens are almost invariably white,
light grey or almost colourless, while the
smaller ones are more or less conspicuously
ornamented with bright yellow.” The number
of branchial plumes (3) is the same in both
species, as is the number of rhinophore la-
mellae (8-10). A. pacifica has 4-5 extra-
branchial appendages and A. cristata usually
has 5 but may have 4-7 (Alder & Hancock,
1855; Meyer & Móbius, 1865). The radular
formula of A. pacifica is 35(1.1.0.1.1)
(MacFarland, 1966) while that for A. cristata
is 25-27(2.0.2) (Alder 8 Hancock, 1855).
MacFarland (1966) reports 11-17 denticles
on the laterals of A. pacifica while Alder 8
Hancock (1855: pl. 46 suppl.) show 12 den-
ticles on the laterals of A. cristata, and Meyer
8 Móbius (1865: pl. 4) show 13-15 denticles.
CALIFORNIA NUDIBRANCHS 145
There seem to be no significant differences
between these two species. Meyer (1971:
140) considers both A. cristata and A. sul-
phurea Stimpson, 1853 as junior synonyms
of A. gibbosa (Risso, 1818). However, spec-
imens from Britain, the Mediterranean, and
New England must be compared to those
from California before the synonymy is cer-
tain.
Anisodoris Bergh, 1898
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum covered with
numerous large, low, rounded tubercles. La-
bial tentacles digitiform. Rhinophores perfo-
liate and retractile into low sheaths which are
bordered by low tubercles. Branchial plumes
tri- or quadripinnate, retractile, arranged in a
circle around anus.
Labial disc smooth. Radular formula
0.n.0.n.0, laterals numerous and hamate.
Penis unarmed. Prostate gland large, set
off from the efferent duct.
Type-species: Anisodoris punctuolata (Or-
bigny, 1837).
Remarks: Thompson (1975) synonymized
Anisodoris with Discodoris. | here retain An-
isodoris for reasons discussed under Disco-
doris (q.v.).
Anisodoris nobilis (MacFarland, 1905)
Montereina nobilis MacFarland, 1905: 38-39.
Abbott, 1974: 351, pl. 17, no. 4215.
Anisodoris nobilis (MacFarland). MacFarland,
1906: 116-118, pl. 18, figs. 6-11; pl. 22,
figs. 1-2. Eliot, 1907: 338. Berry, 1907: 34.
Bovard & Osterud, 1918: 133-134.
O'Donoghue, 1921: 156-158, pl. 1 (7), figs.
9-10. O'Donoghue, 1922a: 126. O'Dono-
ghue & O'Donoghue, 1922: 137. O'Dono-
ghue, 1922b: 163. O'Donoghue, 1924: 1,
22-23, 28-29. O'Donoghue, 1926: 207.
O'Donoghue, 1927b: 81, pl. 1, figs. 4-5.
Fraser, 1932: 67. Hewatt, 1937: 178 ff.
Hewatt, 1938: 287. Costello, 1938: 325,
329-332, tabs. 2, 5, pl. 1, fig. 16. Smith &
Gordon, 1948: 181. Rigg 4 Miller, 1949:
343. Fischer, 1950: 199. LaRocque, 1953:
259. Bousfield, 1958: 109. Marcus, 1959:
45, 47. Eyerdam, 1960: 45. Marcus, 1961:
17-18, pl. 3, figs. 56-58. Lance, 1961: 66.
McLean, 1962: 111. Paine, 1963a: 4.
Farmer 8 Collier, 1963: 62. Steinberg,
1963b: 70. Pequegnat, 1964: 279. Wil-
lows, 1965: 707 ff. MacFarland, 1966: 188-
190, pl. 28, figs: 1, 3; pl. 29, figs. 16-17;
pl. 37, figs. 22-27. Marcus & Marcus,
1967a: 71. Gorman et al., 1967: 329.
Sphon & Lance, 1968: 76. Turner et al.,
1969: 133. Roller & Long, 1969: 425. Gor-
man & Mirolli, 1969: 615 ff. Marmor & Gor-
man, 1969: 293. Robilliard, 1969a: 290.
Mirolli, 1970: 141a. Gosliner & Williams,
1970: 177. Marmor & Salmoiraghi, 1970:
834. Marmor & Gorman, 1970: 65. Ber-
nard, 1970: 84. McBeth, 1970: 28. Gor-
man & Marmor, 1970a: 897, 898. Gorman
& Marmor, 1970b: 919. Gorman & Mar-
mor, 1971: 323. Gorman & Mirolli, 1971:
137a. Marmor, 1971: 575. North, 1971: 57.
Bertsch et al., 1972: 304. Holleman, 1972a:
60. Baker, 1972: 45. McBeth, 1972a: 55
ff. Gorman & Marmor, 1972: 319. Par-
tridge, 1973: 349. Gosliner & Williams,
1973b: 352. Mirolli 8 Gorman, 1973: 423.
Gorman & Marmor, 1974a: 36. Gorman &
Marmor, 1974b: 50. Marmor, 1975: 169 ff.
Belcik, 1975: 276. Schuler, 1975: 33.
Bloom, 1975: 311-314. Bloom, 1976: 289
ff. Guy & Connor, 1976: 146. Partridge 8
Stevens, 1976: 315, 316. Thompson &
Smith, 1976: 153. Williamson 4 Crill,
1976a: 217, 219. Bloom & Bloom, 1977:
296-299. Eckert et al., 1977: 1748. Eyer-
dam, 1977: 110. Hargens, 1977: 363. Ny-
bakken, 1978: 134 ff. Haderlie & Donat,
1978: 60. Eckert & Tillotson, 1978: 178a.
McDonald 8 Nybakken, 1978: 110, 112.
Partridge & Connor, 1978: C155. Fuhrman
et al., 1979: 290 ff. Partridge et al., 1979:
70 ff. Connor, 1979: 41 ff. Harris & Howe,
1979: 145. Aldrich et al., 1979a: 508 ff. Al-
drich et al., 1979b: 532 ff. Bertsch, 1980:
224. Fuhrman et al., 1980: 193.
Anisodoris (Montereina) nobilis. Eliot, 1907:
338.
Archidoris nobilis. Michel, 1976: 46, fig. 3.
Type-specimens: U. S. Nat. Mus., no.
181284.
Foot oval, bluntly rounded and bilabiate
anteriorly, slightly more acutely rounded pos-
teriorly; foot slightly lighter in color than dor-
sum. Rhinophores bear 18-24 lamellae, yel-
low on shaft and yellow-orange on clavus.
Branchial plumes 6, tri- and quadripinnate,
grayish-white, tipped with opaque white.
General ground color varies from light yellow
to deep orange-yellow, dorsum sprinkled with
numerous irregular, dark brown to black
blotches which do not extend up onto any of
the tubercles. T.L.: 35 mm, but specimens up
to 200 mm have been collected.
Radular formula of specimens examined
146 MCDONALD
20-22(40-44.0.40-44), MacFarland (1966:
189) reports 26(55-60.0.55-60), while Mar-
cus (1961: 18) reports 23-27(55-62.0.55-62).
Laterals (Fig. 76) hamate.
Type-locality: Monterey Bay, California.
Range and habitat: Washington Bay, Kuiu
Island, Alaska (Eyerdam, 1977), to Ensena-
da, Baja California, Mexico (Farmer 8 Collier,
1963). Intertidal to 250 m. Common in rocky
intertidal and subtidal zones; occasional small
specimens may be found on docks and pil-
ings in bays. Feeds upon the sponges: Hali-
chondria panicea, Haliclona permollis, Lisso-
dendoryx firma, Mycale adhaerens, Mycale
macginnitiei, Myxilla agennes, Paresperella
psila, Prianos sp., and Zygerphe hyaloderma
(Bloom, 1976: McBeth, 1971; McDonald &
Nybakken, 1978).
Remarks: Additional references which
mention Anisodoris and presumably refer to
A. nobilis are: Both et al., 1976; Connor &
Stevens, 1971a, 1971b, 1971c; Gutknecht,
1970; Williamson 8 Crill, 1976b.
Antiopella Hoyle, 1902
Bogy rather flattened, broadest anteriorly.
Cerata spindle-shaped, rather inflated. Rhi-
nophores non-retractile, perfoliate, with an
inter-rhinophoral crest (or caruncle).
Masticatory border of mandibles denticu-
late. Radular formula 0.n.1.n.0, laterals slightly
hooked distally and may bear denticles.
Penis generally thick and conical, un-
armed. Anus located postero-medially on
dorsum.
Type-species: Antiopella cristata (Delle
Chiaje, 1841).
Antiopella barbarensis (Cooper, 1863)
Aeolis barbarensis Cooper, 1863b: 59-60.
Carpenter, 1864: 609. Cooper, 1867: 14.
Yates, 1890: 41. O'Donoghue, 1922d: 140.
Steinberg, 1963a: 65.
Janolus (Aeolis) barbarensis (Cooper).
O'Donoghue, 1922d: 140-141. O'Dono-
ghue, 1924: 19.
Janolus barbarensis (Cooper). O'Donoghue,
1922d: 140. O'Donoghue, 1926: 228. Mar-
cus, 1958: 40. Lance, 1961: 68. Steinberg,
1963a: 65-66. Abbott, 1974: 373.
Janolus fuscus O'Donoghue, 1924: 16-19, pl.
2, figs. 18-20. O'Donoghue, 1926: 228.
Steinberg, 1963a: 65.
Antiopella aureocincta MacFarland, Mss.
Johnson & Snook, 1927: 500-501. Stein-
berg, 1963a: 66.
Antiopella aureocincta Johnson & Snook.
Costello, 1938: 321, tabs. 1, 5. Marcus,
1958: 40.
Antiopella (Janolus) fuscus O'Donoghue.
Pruvot-Fol, 1954: 375.
Antiopella fusca (O'Donoghue). Marcus,
1958: 40. Steinberg, 1963a: 65. Lambert,
1976: 294-295, 296. Robilliard 8 Barr,
1978: 153.
Antiopella barbarensis (Cooper). Steinberg,
1963a: 65-66. Paine, 1963a: 4. Farmer,
1967: 342. Sphon 8 Lance, 1968: 76.
Lance, 1969: 36. Roller 8 Long, 1969: 425.
Turner et al., 1969: 134. Burn & Miller,
1969: 30. Roller, 1970a: 372. Gosliner 8
Williams, 1970: 177. Keen, 1971: 835, pl.
21, fig. 4. Bertsch et al., 1972: 304. Sphon,
1972a: 155. Nybakken, 1974: 371. Birke-
land, 1974: 218. Ferreira & Bertsch, 1975:
328, figs. 15-16. Lambert, 1976: 294. Ny-
bakken, 1978: 135. McDonald 8 Nybak-
ken, 1978: 110, 114. Russo, 1979: 44, 48.
Antiopella aureocincta MacFarland, 1966:
303-308, pl. 57, figs. 1-5: pl. 63, figs. 13-
30; pl. 64, figs. 11-17. Sphon 8 Lance,
1968: 76. Roller, 1970a: 372. Bernard,
1970: 86. Lambert, 1976: 294.
Hermissenda crassicornis. Buchsbaum &
Milne, 1967: pl. 62.
Type-specimens: not listed; state coll.
species 978 (Cooper, 1863b).
Body somewhat aeolidiform, rounded and
broadest anteriorly, tapered posteriorly to
pointed tail. Foot bilabiate and widest ante-
riorly, and tapered posteriorly to tail. Labial
tentacles cylindrical, short, and blunt. Margin
of dorsum covers foot everywhere, resulting
in a distinct groove between margin of dor-
sum and foot. Rhinophores bear about 10-
14 lamellae, clavus usually bears a subter-
minal ring of orange-gold, with tip either white
or blue. Cerata spindle-shaped, arranged in
longitudinal rows along margin of dorsum,
leaving medial area of dorsum free of cerata;
they extend anterior of rhinophores. General
ground color translucent grayish-white to
pinkish-gray. In specimens from northern
California and Washington, cerata are tipped
with opaque white, below which is a subter-
minal band of orange. In specimens from Cal-
ifornia south of Santa Barbara, cerata tipped
with bright, metallic blue, below which is a
subterminal band of metallic gold. In central
California, in the area of Monterey Bay, both
variations in color of cerata are found, as well
as occasional specimens whose cerata are
CALIFORNIA NUDIBRANCHS 147
intermediate in color, with very light blue tips.
Cerata cores are raw umber. Caruncle is yel-
low-orange. T.L.: 20 mm.
Anus located medially, well back on dor-
sum. Genital aperture located midway on right
side. Penis cylindrical and tapered distally,
unarmed.
Radular formula of specimen examined
16(14-21.1.14-21), MacFarland (1966: 305)
reports 20-22(18-22.1.18-22). Rachidian
tooth (Fig. 91h) bears a single, median den-
ticle, with 6-8 small denticles on either side.
Laterals (Fig. 91a-g) are slightly hooked dis-
tally, inner laterals with 12-17 minute denti-
cles. Masticatory border of the large, thick
mandibles bears 8-14 denticles.
Type-locality: Santa Barbara, Santa Bar-
bara Co., California.
Range and habitat: Klu Bay, Revillagigedo
Island, Alaska (Robilliard & Barr, 1978), to
Bahía San Quintín, Baja California, Mexico
(Keen, 1971); between Isla San Diego and
Isla San José, Gulf of California, Mexico (Fer-
reira 8 Bertsch, 1975); Bahía de los Angeles,
Baja California, Mexico (Keen, 1971). Inter-
tidal to 30 m. Found in rocky intertidal as well
as on floating docks and pilings in bays.
Feeds upon the hydroid Corymorpha palma
and the bryozoan Bugula californica (Mac-
Ginitie & MacGinitie, 1949: McDonald 8 Ny-
bakken, 1978).
Remarks: O'Donoghue (1924: 16-19)
named Janolus fusca from Galiano Island,
British Columbia, Canada. In comparing
O'Donoghue's description of this species with
MacFarland’s (1966: 303-308) description of
Antiopella aureocincta (a junior synonym of
A. barbarensis) | note a number of similari-
ties. The number of denticles of the masti-
catory border of the mandible is 10-12 for J.
fusca (O'Donoghue, 1924: 18) and 8-14 for
A. aureocincta (MacFarland, 1966: 305).
O'Donoghue (1924: 17) reports the radular
formula 21(4-22.1.4-22) for the latter species.
O'Donoghue (1924: 18) reports 10-14 mi-
nute serrations on the inner side of the first
3-4 laterals and 2-3 denticles on the outer
side of same, while MacFarland (1966: 305)
reports 12-17 small pointed denticles on the
inner hook of the first 3 inner laterals and
occasionally a few similar denticles on the
outer face of the first lateral. O'Donoghue
(1924: 16) states that a line of intense,
opaque white spots runs from the last cerata
to the tip of the tail of J. fusca; while Mac-
Farland (1966: 306) reports the dorsum of
tail with a narrow, median, longitudinal band
of white in A. aureocincta. O'Donoghue
(1924: 16) states that ‘‘пеаг the end on the
cerata where the core stops is a very strik-
ing, Opaque, bright orange band. The pointed
tip is transparent for a short distance and then
is covered with opaque, intensely white
spots.” MacFarland (1966: 306) states “tips
of the papillae pure white encircled below by
a band of yellow orange.” O'Donoghue's
(1924: pl. 2, figs. 18-20) drawings of the
mandibles and radula correspond very well to
those by MacFarland (1966: pl. 63, figs. 13—
25). In comparing the mandibles and radula
of a specimen from Friday Harbor, Washing-
ton with those of a specimen from Hazard
Canyon, San Luis Obispo Co., California, |
can see no significant differences between the
two. Therefore, Janolus fusca is a junior sub-
jective synonym of Antiopella aureocincta,
and since Sphon 4 Lance (1968: 76) consid-
er the latter a junior synonym of Antiopella
barbarensis, the correct name for the species
which has been known by the above three
names is Antiopella barbarensis by the law
of priority.
O'Donoghue (1922d: 141) felt that Janolus
coeruleopictus Cockerell 8 Eliot, 1905, should
be considered a synonym of A. barbarensis,
and stated that Eliot agreed with him. The
description of J. coeruleopictus is quite brief
but is fairly close to A. barbarensis, the only
major difference being that the former has
tessellated, nondenticulate mandibles (Cock-
erell & Eliot, 1905) while the latter has dentic-
ulate mandibles. Unless this difference in the
mandibles can be explained, | feel that Stein-
berg (1963a: 66) was correct in stating that
because of the poor description of J. coeru-
leopictus it should be declared a nomen du-
bium.
Archidoris Bergh, 1878
Body somewhat soft, doridiform, rather
evenly rounded anteriorly and posteriorly;
dorsum covered with numerous low, rounded
tubercles of various sizes. Labial tentacles
short and thick, with an external longitudinal
groove. Rhinophores perfoliate and retractile
into low sheaths with tuberculate margins.
Branchial plumes retractile, tri- to quadripin-
nate, arranged in a circle around anus.
Labial disc unarmed. Radular formula
0.п.0.п.0, laterals hamate.
Penis unarmed, pleurembolic; glans penis
distinct. Vas deferens glomerate.
148 MCDONALD
Type-species: Archidoris tuberculata (Cu-
vier, 1804) (non Müller, 1778).
Archidoris montereyensis (Cooper, 1863)
Doris montereyensis Cooper, 1863a: 204.
Cooper, 1863a: 58. Carpenter, 1864: 608,
609. Cooper, 1867: 14. Cooper, 1870: 56.
Dall, 1871: 137.
Doris tuberculata Cuvier. Abraham, 1877:
203-204. O'Donoghue, 1926: 206. O'Don-
oghue, 1927b: 80.
Archidoris montereyensis (Cooper). Bergh,
1878b: 624, pl. 68, fig. 24. Bergh, 1879b:
107-108. Bergh, 1879c: 163-164. Bergh,
1880a: pl. 8 (16), figs. 10-11. Bergh,
1880b: pl. 8(16). Bergh, 1880c: 34. Bergh,
1891: 128. Bergh, 1892: 1092 (100). Bergh,
1898: 501. MacFarland, 1905: 37. Cock-
erell & Eliot, 1905: 34. MacFarland, 1906:
114-116, pl. 18, figs. 1-5; pl. 23, fig. 4.
Berry, 1907: 34. Cockerell, 1908: 106.
O'Donoghue, 1921: 154-156, pl. 1 (7), figs.
7-8. O'Donoghue, 1922a: 126. O'Dono-
ghue 8 O'Donoghue, 1922: 137, pl. 3, fig.
5. O'Donoghue, 1924: 22, 28. O'Dono-
ghue, 1926: 206-207. Odhner, 1926a: 64,
67. O'Donoghue, 1927b: 80. Fraser, 1932:
67. Costello, 1938: 324 ff, tabs. 1-5; pl. 1,
fig. 5; pl. 2, figs. 28, 41. Smith & Gordon,
1948: 181. LaRocque, 1953: 259. Mc-
Gowan 4 Pratt, 1954: 261-276. Goodwin
& Fox, 1955: 1086. Marcus, 1957: 471.
Marcus, 1961: 16-17, pl. 3, figs. 54-55.
Lance, 1961: 66. Cook, 1962: 194-196.
Paine, 1963a: 4. Steinberg, 1963b: 70.
deVries, 1963: 124. Willows, 1965: 707 ff.
MacFarland, 1966: 181-182, pl. 27, fig. 8;
pl. 37, figs. 1-10. Thompson, 1967: 9.
Hurst, 1967: 255 ff, text figs. 4a—b; pl. 27,
fig. 5; pl. 34, fig. 33; fig. 24-18. Sphon 8
Lance, 1968: 76. Burn, 1968: 91, 92. Lee
8 Brophy, 1969: 220. Roller & Long, 1969:
426. Nicaise, 1969: 2601, pl. 1, fig. 1. Rob-
illiard, 1969a: 290. Gosliner & Williams,
1970: 177. Bernard, 1970: 85. Crane,
1971: 57. Harris, 1971a: 84. North, 1971:
57. Holleman, 1972a: 60. Bertsch et al.,
1972: 305. Sphon, 1972a: 155. McBeth,
1972a: 56. Harris, 1973: 219 ff. Burn, 1973:
40, fig. 7c. Partridge, 1973: 349. Chia 4
Skeel, 1973: 153. Crane, 1973: 17. Had-
erlie et al., 1974: tab. 4. Abbott, 1974: 351,
pl. 17, no. 4211. Robilliard, 1974b: 989.
Hinegardner, 1974: 452. Belcik, 1975: 276.
Bloom, 1975: 311-314. Lambert, 1976:
296. Bloom, 1976: 289 ff. Elvin, 1976: 194.
Partridge & Stevens, 1976: 315, 316.
Thompson & Smith, 1976: 153. Williamson
8 Crill, 1976a: 217, 219. Bloom & Bloom,
1977: 296-299. Connor, 1977a: 43a. Con-
nor, 1977b: 487 ff. Connor & Ahmed, 1978:
186a. Nybakken, 1978: 134 ff. Haderlie &
Donat, 1978: 60. McDonald & Nybakken,
1978: 112, 116. Idler et al., 1978: 163 ff.
Partridge & Connor, 1978: C155. Clark 8
Goetzfried, 1978: 290. Dehnel 8 Kong,
1979: 1843. Aldrich et al., 1979a: 508 ff.
Aldrich et al., 1979b: 532 ff. Fuhrman et
al., 1979: 291 ff. Partridge et al., 1979: 70
ff. Connor, 1979: 41 ff. Ahmed 8 Connor,
1979a: 61 ff. Ahmed & Connor, 1979b:
265a. Connor & Ahmed, 1979: 265a.
Ahmed & Connor, 1980a: 2038. Ahmed &
Connor, 1980b: 403 ff.
Doris (Archidoris) montereyensis (Cooper).
Bergh, 1880a: 232. Bergh, 1880b: 83. Or-
cutt, 1885: 545.
Archidoris nyctea Bergh, 1900: 222-224, pl.
21, figs. 70-72. MacFarland, 1966: 182.
Burn, 1968: 91.
Archodoris montereyensis (Cooper). O'Don-
oghue, 1922b: 163 (/apsus).
Anisodoris nobilis (MacFarland). Guberlet,
1962: 255.
Type-specimens: not listed; Geological
survey coll. (Cooper, 1863a).
Foot elliptical, bluntly rounded and bila-
biate anteriorly, bluntly rounded posteriorly.
Labial tentacles relatively small and auricu-
late. Rhinophores bear 20-30 lameliae, yel-
low to orange. Branchial plumes 7, tri- to
quadripinnate, yellow to orange. General
ground color variable, light yellow to dark or-
ange, dorsum bears many irregular blotches
of brown to black which extend up onto the
tubercles. T.L.: 25 mm, but specimens up to
150 mm have been collected.
Radular formula of specimen examined
20(34-37.0.34-37), MacFarland (1966: 181)
reports 33(42-49.0.42-49), while Marcus
(1961: 17) reports 33(42-65.0.42-65), and
Bloom (1976: 292) gives the range 27-36(42-
70.0.42-70). Laterals (Fig. 70) hamate.
Type-locality: Monterey Bay, California.
Range and habitat: Jackson Point, Port
Valdez, Alaska (Robilliard, 1974b), to La Jol-
la, San Diego Co., California (Cockerell 8
Eliot, 1905). Intertidal to 256 m. Fairly com-
mon in rocky intertidal, also found on docks
and pilings in bays; also occurs well up into
Elkhorn Slough, Monterey Co., California,
where it feeds upon a yellow sponge which
CALIFORNIA NUDIBRANCHS 149
grows on muddy bottom. This sponge has
been tentatively identified as Halichondria
bowerbanki (McDonald & Nybakken, 1978).
Cook (1962: 196) reports that it feeds upon
Halichondria panicea.
Remarks: This species should be closely
compared to Archidoris tuberculata (Cuvier,
1804), which occurs in England and very
closely resembles A. montereyensis.
Additional references which mention Archi-
doris and are presumably referring to A.
montereyensis are Both et al. (1976) and
Stevens (1969).
Archidoris odhneri (MacFarland, 1966)
Austrodoris odhneri MacFarland, 1966: 173-
179%pl 26, fig: 1; pl. 29, fig. 14; pl. 36;
figs. 1-19. Hurst, 1967: 255 ff, text fig. 5,
pl. 26, fig. 4; pl. 33, figs. 31-32; fig. 24-
24. Sphon 8 Lance, 1968: 77. Burn, 1968:
90-92. Roller, 1970a: 371. Robilliard,
1971a: 164. Partridge, 1973: 349-350, fig.
1c-d.
Archidoris odhneri(MacFarland). Burn, 1968:
90-92. Roller £ Long, 1969: 425. Roller,
1970a: 371. Robilliard, 1971a: 164, 165.
Bertsch et al., 1972: 305. Burn, 1973: 40,
fig. 7f. Robilliard, 1974b: 989-990. Abbott,
1974: 351. Belcik, 1975: 276. Lambert,
1976: 296. Bloom, 1976: 289 ff. McDonald
8 Nybakken, 1978: 112. Robilliard 8 Barr,
1978: 153. Fuhrman et al., 1979: 291 ff.
Bertsch, 1980: 224. Andersen & Sum,
1980: 797-800.
Archidoris (=Austrodoris) odhneri (Mac-
Farland). Robilliard, 1971a: 164.
Type-specimens: type material at Calif.
Acad. Sci.
Dorsal tubercles larger medially and small-
er near edge of dorsum. Foot elliptical, blunt-
ly rounded and bilabiate anteriorly, and blunt-
ly rounded posteriorly. Labial tentacles slightly
triangular to digitiform. Rhinophores bear 20-
24 lamellae and are white to very light dusty
yellow. Branchial plumes 7, tri- to quadripin-
nate, very finely divided, appearing quite
feathery and delicate, white to very light yel-
low. General ground color pure white, occa-
sional individuals may be very light dusty yel-
low. Dorsum dusted with numerous minute,
opaque flecks of the ground color. T.L.: 100
mm, but specimens over 200 mm have been
collected.
Radular formula of specimen examined
20(44.0.44), MacFarland (1966: 176) reports
34(55.0.55). Laterals (Fig. 71) hamate.
Type-locality: Monterey Bay, California.
Range and habitat: Port Dick, Kenai Pen-
insula, Alaska (Robilliard & Barr, 1978), to
Point Loma, San Diego Co., California (Ro-
billiard, 1974b). Intertidal to 25 m. Most com-
monly found subtidally in kelp (Macrocystis
pyrifera) bed areas, quite rare intertidally in
California. Feeds upon the sponges: Craniel-
la sp., Halichondria panicea, Mycale adhae-
rens, Myxilla incrustans, Stylissa stipitata,
Syringella amphispicula, Tedania sp. and a
hexactinellid (possibly Rhabdocalyptus sp.)
(Bloom, 1976; Robilliard, 1971a).
Armina Rafinesque, 1814
Body ovate, rounded anteriorly and ta-
pered posteriorly to a pointed tail. Dorsum
bears numerous longitudinal ridges which oc-
cur along the entire length of dorsum; sculp-
ture of dorsum is different in a few species,
consisting of a few widely separated ridges,
or of numerous tubercles. Foot broadly
rounded anteriorly and tapered posteriorly to
pointed tail. Anterior margin of dorsum gen-
erally has a rounded median notch through
which the rhinophores project. Rhinophores
closely appressed, nearly united at base,
retractile beneath mantle margin, and bear
vertical perfoliations. Branchiae consist of
longitudinally directed lamellae, arranged an-
tero-laterally just below edge of dorsum; there
may be smaller lamellae between the larger
lamellae.
Masticatory border of mandibles denticu-
late. Radular formula usually n.1.1.1.n; ra-
chidian tooth bears a strong denticle, with
smaller denticles on either side; lateral den-
ticulate or not; marginals more elongate and
denticulate or not.
Penis unarmed. Anus on right side, below
mantle margin, usually posterior of bran-
chiae.
Type-species: Armina tigrina Rafinesque,
1814.
Armina californica (Cooper, 1863)
Pleurophyllidia californica Cooper, 1863a:
203-204. Bergh, 1866: 33, pl. 1. Cooper,
1867: 14. Bergh, 1869: 229. Bergh, 1876a:
2. Bergh, 1890a: 3-8, pl. 1, figs. 1-6; pl.
2, figs. 1-2. Bergh, 1892: 1063 (71). Bergh,
1894: 154-157, pl. 3, figs. 14-15; pl. 4,
figs. 7-12. MacFarland, 1897: 227, 229-
150 MCDONALD
244, pl. 18, figs. 1-10; pl. 19, figs. 11-20;
pl. 20, figs. 21-23. Bergh, 1904: 19-20, pl.
4, figs. 23-26. Kelsey, 1907: 47. O'Dono-
ghue, 1921: 178-180, pl. 5 (11), figs. 49-
50. O'Donoghue, 1922а: 124. O'Dono-
ghue, 1926: 222. Fraser, 1932: 67. Norris
8 Rao, 1935: 787. MacFarland, 1966: 205.
Hinegardner, 1974: 452.
Pleurophyllidea californica Cooper. Carpen-
ter, 1864: 608, 647 (lapsus).
Pleurophyllidia vancouverensis Bergh, 1876a:
3, 5-9, pl. 1, figs. 8-17. MacFarland, 1966:
205.
Armina columbiana O'Donoghue, 1924: 11-
14, pl. 2, figs. 13-17. O'Donoghue, 1926:
222. LaRocque, 1953:260. Marcus, 1961:
43-44, 58, pl. 8, figs. 151-154. Lance,
1962c: 54. Steinberg, 1963a: 64-65.
MacFarland, 1966: 206.
Armina vancouverensis (Bergh). O'Dono-
ghue, 1926: 222. LaRocque, 1953: 260.
Marcus, 1961: 44. Lance, 1962c: 54.
Steinberg, 1963a: 64-65. MacFarland,
1966: 206. Marcus 8 Marcus, 1967b: 215.
Armina californica (Cooper). O'Donoghue,
1926: 222. O'Donoghue, 1927a: 11. La-
Rocque, 1953: 260. Marcus, 1961: 41-43,
44, 56, pl. 8, figs. 147-150. Lance, 1961:
67. Lance, 1962c: 54. Steinberg, 1963a:
64-65. Steinberg, 1963b: 71. Lance, 1966:
69. MacFarland, 1966: 198-206, pl. 38,
figs. 1-6; pl. 43, figs. 37-44; pl. 44, figs.
6-7. Marcus & Marcus, 1967b: 215. Hurst,
1967: 255 ff, text figs. 11a-c, pl. 29, fig.
14; pl. 34, figs. 34-35; fig. 24-20. Farmer,
1967: 342. Sphon 8 Lance, 1968: 77.
Mauzey et al., 1968: 606. Bertsch, 1968:
440-441. Lance, 1969: 34. Roller & Long,
1969: 429. Turner et al., 1969: 133. Ber-
nard, 1970: 85. Keen, 1971: 834. Holle-
man, 1972a: 60. Thompson & Bebbington,
1973: 148, pl. 10, figs. a-d. Thompson,
1973: 167, 172, 179, 186, text fig. 9. Gos-
liner & Williams, 1973b: 354. Bertsch, 1973:
110. Abbott, 1974: 372, fig. 4400. Birke-
land, 1974: 211 ff. Kastendiek, 1975: 784.
Thompson, 1976a: 39, pl. 2, fig. d. Thomp-
son, 1976b: 16, 47. Kastendiek, 1976: 519
ff, text fig. 7. Poorman & Poorman, 1978:
373. McDonald & Nybakken, 1978: 114.
Dorsett, 1978: 307-324.
Armina (Pleurophyllidia) californica (Bergh).
Smith & Gordon, 1948: 181.
Armina digueti Pruvot-Fol, 1955: 464-465,
text figs. 8-10. Marcus, 1961: 44. Stein-
berg, 1963a: 65. Marcus 8 Marcus, 1967b:
216.
Armina (=Pleurophyllidia) californica (Coo-
per). Willows, 1965: 707 ff.
Type-specimens: not listed, Geological
Survey coll. (Cooper, 1963a).
Dorsum bears about 18-25 longitudinal
ridges. Head consists of a transverse head
shield which is broadly rounded anteriorly,
with undulating margins. Rhinophores bear
30-40 lamellae and are grayish-pink with
white on tip and on margins of lamellae.
Branchiae 15-30 on either side. General
ground color light pinkish-brown, longitudinal
dorsal ridges light pinkish-brown to cream,
and area between ridges gray to grayish-
brown. Margin of dorsum edged with white
to cream. Dorsal surface of head shield gray
to grayish-brown. T.L.: 30 mm.
Radular formula of specimen examined
25(39-44.1.1.1.39-44), MacFarland (1966:
202) reports 30(45.1.45) to 52(81.1.81), and
Marcus (1961: 42) reports 41(60.1.1.1.60).
Rachidians (Fig. 90k) bear 3-4 (rarely up to
8) denticles on either side of median denticle.
Type-locality: San Diego Bay, California.
Range and habitat: Hecate Strait, British
Columbia, Canada (O'Donoghue, 1921), to
Panama (Bergh, 1894). Found on sandy-mud
bottom in 1 to 230 m, usually in association
with the sea pansy Renilla kollikeri (vide
Bertsch 1968: 440) or the pennatulacean Pti-
losarcus gurneyi, both of which it feeds upon
(Bertsch, 1968; Birkeland, 1974; Kastendiek,
1976; MacFarland, 1966; Turner et al., 1969;
McDonald and Nybakken, 1978).
Additional references: Anderson 8 Chase
(1975) mention Armina, presumably referable
to A. californica.
Atagema Gray, 1850
Body rather firm, doridiform, rather evenly
rounded anteriorly and posteriorly. Dorsum
spiculose, bears a medial ridge. Labial ten-
tacles somewhat digitiform or triangular.
Rhinophores perfoliate and retractile into
sheaths. Branchial plumes few, tri- or quad-
ripinnate, located just anterior of anus.
Radular formula 0.n.0.n.0, laterals hook-
shaped.
Penis unarmed.
Type-species: Atagema carinata (Quoy &
Gaimard, 1832).
Atagema alba (O'Donoghue, 1927)
Glossodoridiformia alba O'Donoghue, 1927b:
87-89, pl. 1, figs. 29-32. Lance, 1961: 66.
CALIFORNIA NUDIBRANCHS Jo
Atagema quadrimaculata Collier, 1963: 73-
75, text figs. 1-5. Smith, 1964: 172. Sphon
& Lance, 1968: 82. Roller & Long, 1969:
429. Roller, 1970a: 371. Abbott, 1974: 351.
Petelodoris spongicola MacFarland, 1966:
183-187, pl. 27, figs. 1-5; pl. 30, fig. 16;
pl. 37, figs. 11-21. Lee & Brophy, 1969:
220. Roller, 1970a: 371.
Type-specimens: not listed.
Dorsum densely covered with numerous
minute, hispid papillae. An irregular ridge ex-
tends posteriorly on dorsum from between
rhinophores to just anterior of branchial
plumes; a few smaller irregular ridges extend
laterally from a median prominence on the
ridge. Edge of dorsum somewhat crenulate
or undulating. Foot elliptical, somewhat elon-
gated; bluntly rounded and bilabiate anterior-
ly, and bluntly rounded posteriorly. Labial
tentacles triangular, thin, and tapered to blunt
tips. Rhinophores bear 15-18 lamellae, cla-
vus very pale green. Rhinophore sheaths with
bluntly rounded margins and covered with
small, hispid papillae. Branchial plumes 3-5,
bi- to tripinnate, projecting posteriorly from
beneath 3 thick, bluntly triangular lobes on
anterior and antero-lateral margins of bran-
chial sheath; branchial plumes pale raw um-
ber. General ground color raw umber, with
small, dark brown to black spots between the
papillae on dorsum, and with very small flecks
of pink along mid-dorsal ridge. T.L.: 22 mm.
Radular formula of specimen examined
15(17-20.0.17-20), MacFarland (1966: 185)
reports 15(20-23.0.20-23), while Collier
(1963: 74) reports 18(18-19.0.18-19), and
O'Donoghue (1927b: 88) reports 17-18(25-
26.0.25-26). Laterals (Fig. 67) strongly
hooked.
Type-locality: Laguna Beach, Orange Co.,
California.
Range and habitat: Point Pinos, Monterey
Co., California (MacFarland, 1966), to San
Diego, San Diego Co., California (Collier,
1963). Intertidal to 210 m. Quite rare
throughout most of its range. It quite proba-
bly feeds upon encrusting sponges upon
which it has been found. Most commonly
found in extreme low intertidal zone, in areas
of heavy surf.
Remarks: O'Donoghue (1927b: 87-89) de-
scribed Glossodoridiformia alba from Laguna
Beach, California, but the species has not
been recognized since. However, upon closely
comparing O'Donoghue's original description
with that of Atagema quadrimaculata Collier
(1963: 73-75) and that of its junior synonym
Petelodoris spongicola MacFarland (1966:
183-187), it seems quite certain that they are
synonymous. The general ground color (white
to raw umber) is essentially the same in all
three species. The dorsum bears small pa-
pillae in all three cases. The number of rhi-
nophore lamellae is comparable, about 15 in
G. alba, 16-18 in P. spongicola, and 16 in A.
quadrimaculata. The foot is anteriorly bila-
biate and deeply notched in all three cases.
The number of branchial plumes is compa-
rable, five in G. alba, three with two basal
branches in P. spongicola, and five in A.
quadrimaculata. The branchial sheath of G.
alba was described by O'Donoghue (1927b:
88) as: ‘'... a large pocket with a raised rim,
which is papillated right up to the margin like
the rhinophore sheath, but this part of the
animal had been damaged in life and part of
the rim and sides of the sheath torn off and
partially healed, giving the animal a superfi-
cial likeness to a phanerobranchiate form.”
Certainly the branchial lobes and posteriorly
projecting branchial plumes of A. quadrima-
culata and P. spongicola could give the ap-
pearance described by O'Donoghue. The
radular formulae are comparable in all three
cases: 17-18(25-26.0.25-26) in G. alba,
15(20-23.0.20-23) in P. spongicola, and
18(18-19.0.18-19) in A. quadrimaculata. The
shape and relative sizes of the teeth are also
comparable in all three cases.
From the above facts it seems obvious that
G. alba is conspecific with A. quadrimacula-
ta, the correct name being Atagema alba
(O’Donoghue, 1927) by the law of priority.
Glossodoridiformia O'Donoghue, 1927, is
therefore a junior synonym of Atagema Gray,
1850.
Babakina Roller, 1973
Body aeolidiform, broadest anteriorly and
tapered posteriorly to pointed tail. Foot cor-
ners very produced. Cerata fusiform, ar-
ranged in numerous oblique rows on either
side of dorsum. Rhinophores non-retractile,
joined together basally for half their length.
Masticatory border of mandibles denticu-
late. Radula uniseriate, rachidians bear
prominent median cusp, with smaller denti-
cles on either side.
Penis unarmed. Anus pleuroproct.
Type-species: Babakina festiva (Roller,
1972).
152 MCDONALD
Babakina festiva (Roller, 1972)
Babaina festiva Roller, 1972: 416-418, text
figs. 1-9. Miller, 1974: 40.
Babakina festiva (Roller). Roller, 1973: 117-
118.
Type-specimens: Calif. Acad. Sci., no. 486.
Foot rather narrow, linear, somewhat
emarginate anteriorly and tapered posteriorly
to tail. Foot corners produced into rather long,
tentaculiform processes with a shallow
groove. Oral tentacles long and cylindrical,
slightly tapered to blunt points; dark pinkish-
red on proximal half and very light yellow on
distal half. Rhinophores perfoliate with 30-35
lamellae. Rhinophore shaft light pink, clavus
yellow to brownish with a pink vertical line on
proximal half of anterior face and with a num-
ber of small, encrusting, yellow dots distally,
which extend in a postero-medial line on dis-
tal half. With a small, yellow to brownish knob
projecting above clavus. Cerata cylindrical,
slightly clavate, somewhat decumbent, and
arranged in about 22 oblique rows from just
anterior of rhinophores, nearly to tip of tail.
Antero-medial area of dorsum free of cerata.
General ground color translucent grayish-
white with a pinkish tinge, which is more pro-
nounced in some specimens. Head region
may be quite pinkish-red in some specimens,
and very pale in others. A medial line of yel-
low-white passes from base of rhinophores
to between oral tentacles. Cardiac region
bears numerous spots of opaque white. Dor-
sal tip of tail yellowish. Cerata pinkish-red on
proximal half, with a subapical band of cad-
mium yellow, and just below this, a wide band
of rather diffuse, opaque white dots; the tips
are translucent grayish-white; cores dark
pinkish-red. T.L.: 15 mm.
Anus on right, just posterior of cardiac re-
gion. Genital aperture on right side, ventral of
fourth row of cerata. Penis unarmed.
Radular formula of specimen examined
14(0.0.1.0.0), Roller (1972: 418) reports
16(0.0.1.0.0). Rachidian tooth (Fig. 114) bears
6-11 small denticles on either side of large,
median cusp. Masticatory border of mandi-
bles bears several rows of long denticles.
Type-locality: White's Point, Palos Verdes
Peninsula, Los Angeles Co., California.
Range and habitat: Malibu Reef, Los An-
geles Co., California (Roller, 1972), to La Jol-
la, San Diego Co., California (Roller, 1972);
Matenchén, Nayarit, Mexico (personal obser-
vation); Japan (Roller, 1972). Intertidal to 5
m in rocky areas. Quite rare in California.
Cadlina Bergh, 1878
Body doridiform, somewhat elongate,
bluntly rounded anteriorly and less bluntly
rounded posteriorly. Dorsum frequently mi-
nutely granular, occasionally tuberculate. La-
bial tentacles more or less triangular. Rhi-
nophores perfoliate and retractile into
sheaths. Branchial plumes uni- to tripinnate,
retractile, arranged in a circle around anus.
Labial disc armed with minute hooks. Rad-
ular formula 0.n.1.n.0, rachidians each bear
a few denticles, laterals usually denticulate.
Penis usually armed with hooks.
Type-species: Cadlina laevis (Linnaeus,
1767).
See ICZN (1967, Opinion 812), Cadlina
validated.
Cadlina flavomaculata MacFarland, 1905
Cadlina flavomaculata MacFarland, 1905: 43.
Cockerell & Eliot, 1905: 35. MacFarland,
1906: 126-128, pl. 19, figs. 32-37; pl. 21,
fig. 110; pl. 25, fig. 9. Kelsey, 1907: 35.
Cockerell, 1908: 106. O'Donoghue 4
O'Donoghue, 1922: pl. 3, fig. 4. O'Dono-
ghue, 1922b: 154-155, 165, pl. 6, figs. 16-
18. O'Donoghue, 1924: 23, 29. O'Dono-
ghue, 1926: 210. O'Donoghue, 1927a: 10.
O'Donoghue, 1927b: 85-86, pl. 1, figs. 20—
24. Fraser, 1932: 67. Baba, 1935a: 343.
Baba, 1937c: 75. Costello, 1938: 324-326,
tabs. 1-3, 5. Smith & Gordon, 1948: 180.
LaRocque, 1953: 259. Marcus, 1955: 122.
Marcus, 1961: 14, pl. 3, figs. 41-42. Lance,
1961: 65. Lance, 1962a: 157. Paine,
1963a: 4, 7. Steinberg, 1963b: 69. Farmer,
1964: 24. MacFarland, 1966: 140, 143,
144-147 ff, pl. 23, fig. 1; pl. 29, fig. 12; pl.
33, figs. 12-21. Marcus 8 Marcus, 1967a:
169. Sphon & Lance, 1968: 77. Lee & Bro-
phy, 1969: 220. Roller 8 Long, 1969: 426.
Gosliner & Williams, 1970: 177. Bernard,
1970: 85. Schmekel, 1970: 194. North,
1971: 57. Keen, 1971: 823. Bertsch et al.,
1972: 305. Abbott, 1974: 356, fig. 4262.
Belcik, 1975: 276. Thompson, 1976b: 92.
Nybakken, 1978: 135. Haderlie & Donat,
1978: 60. McDonald & Nybakken, 1978:
110, 112. Fuhrman et al., 1979: 291.
Bertsch, 1980: 224. Chamberlain & Beh-
rens, 1980: 284.
Cardlina flavomaculata MacFarland. Ingram,
1935: 48 (lapsus).
Type-specimens: U. S. Nat. Mus., no.
181279.
CALIFORNIA NUDIBRANCHS 153
Dorsum covered with numerous very small,
low tubercles. Foot narrow and linear, broad-
ly rounded anteriorly and bluntly pointed pos-
teriorly. Labial tentacles short and bluntly au-
riculate, with a groove on external margin.
Rhinophores bear 8-12 lamellae, which are
brown to almost black, causing entire clavus
to appear brown. Branchial plumes 10-12,
uni- to bipinnate, light cream to yellow. Gen-
eral ground color light cream to light yellow.
A series of 6-10 or more small, lemon yellow
dots occurs longitudinally on either side of
dorsum, from just posterior of rhinophores to
just posterior of branchial plumes. T.L.: 15
mm.
Radular formula of specimen examined
51(28.1.28), MacFarland (1966: 145) reports
77(23.1.23), while Marcus (1961: 14) reports
68-77(22-27.1.22-27). Rachidian teeth (Fig.
58a) may bear 2-3 denticles on inner margin
and 4-7 denticles on outer margin. Outer lat-
erals (Fig. 58b-d) may bear 11-15 denticles
on margin.
Type-locality: Monterey Bay, California.
Range and habitat: Vancouver Island, Brit-
ish Columbia, Canada (O’Donoghue, 1922b),
to Punta San Eugenio, Baja California, Mex-
ico (Lance, 1961); northern end of Gulf of
California, Mexico. Intertidal to 220 m. Not
uncommon in rocky intertidal zone in central
California. Feeds upon the sponge Aplysilla
glacialis (McDonald 4 Nybakken, 1978: 110).
Cadlina limbaughi Lance, 1962
Cadlina sp. Lance, 1961: 66.
Cadlina limbaughi Lance, 1962a: 155-157,
text figs. 1-3. Shields, 1966: 189. Marcus
& Marcus, 1967a: 169. Sphon & Lance,
1968: 77. North, 1971: 57. Abbott, 1974:
357. Michel, 1976: 48, fig. 11.
Type-specimens: Calif. Acad. Sci., Paleo.
type coll. no. 12’396.
Dorsum nearly smooth, bearing very small
tubercles. Foot rather linear, broadly rounded
and slightly bilabiate anteriorly, and tapered
posteriorly to bluntly rounded tail. Labial ten-
tacles thick, triangular and auriform, with a
groove on external margin. Rhinophores bear
11-18 lamellae, clavus dark reddish-brown.
Branchial plumes 6-8, bi- and tripinnate,
whitish at base and dark reddish-brown on
distal two thirds, except for the most poste-
rior plume which is white. General ground
color whitish, with numerous opaque white
dots on dorsum. Sides of body and dorsal
and ventral surfaces of foot bear а few mi-
nute black dots. T.L.: 15 mm.
Lance (1962a: 156) gives the radular for-
mula 102(0.44.1.44.0). Rachidian teeth (Fig.
59a) bear 4-6 denticles. Innermost laterals
(Fig. 59b) bear 3 denticles on inner margin
and 6 denticles on outer margin. Outer later-
als (Fig. 59c) somewhat hooked with about
14 denticles below cusp; the outermost lat-
erals (Fig. 59d) lack a cusp and bear about 9
denticles.
Type-locality: La Jolla, San Diego Co., Cal-
ifornia.
Range and habitat: Coal Oil Point, Santa
Barbara Co., California (Sphon 8 Lance,
1966), to Islas Coronados, Baja California,
Mexico (Lance, 1961). Subtidal, 9-43 m, in
rocky areas. This species is relatively uncom-
mon.
Cadlina marginata MacFarland, 1905
Cadlina marginata MacFarland, 1905: 43.
Cockerell & Eliot, 1905: 35. MacFarland,
1906: 125-126, pl. 18, figs. 27-31; pl. 25,
figs. 10-12. Kelsey, 1907: 35. Berry, 1907:
35. Cockerell, 1908: 106. O'Donoghue,
1921: 161-162, pl. 1 (7), figs. 13-14.
O'Donoghue 8 O'Donoghue 1922: 138.
O'Donoghue, 1922b: 164. O'Donoghue,
1924: 24, 29. O'Donoghue, 1927a: 10.
O'Donoghue, 1927b: 86-87, pl. 1, figs. 25—
28. Fraser, 1932: 67. Baba, 1935a: 343.
Hewatt, 1937: 200. Baba, 1937c: 75. Cos-
tello, 1938: 324 ff, tabs. 1-3, 5, pl. 1, figs.
4, 6, 10, 22; pl. 2, fig. 32. Smith 8 Gordon,
1948: 180. Rigg 8 Miller, 1949: 343. La-
Rocque, 1953: 259. Marcus, 1955: 122.
Marcus, 1961: 15. Lance, 1961: 66. Lance,
1962a: 157. McLean, 1962: 110. Stein-
berg, 1963b: 69. Paine, 1963a: 4. Willows,
1965: 707 ff. MacFarland, 1966: 140. Mar-
cus & Marcus, 1967a: 170. Robilliard,
1969a: 290. North, 1971: 57. Abbott, 1974:
357, fig. 4263. (Non) Doris marginata Mon-
tagu, 1804: 79, pl. 7, fig. 7 (=Cadlina laevis
Linnaeus, 1767).
Cadlina luteomarginata MacFarland, 1966:
140-144, pl. 23, figs. 2-4; pl. 29, figs. 13-
13a; pl. 33, figs. 1-11. Sphon & Lance,
1968: 77. Roller 8 Long, 1969: 426.
Bertsch, 1969: 231. Gosliner & Williams,
1970: 177. Bernard, 1970: 85. Schmekel,
1970: 194. Bertsch et al., 1972: 305. Ro-
billiard, 1974b: 990. Belcik, 1975: 276.
Bloom, 1976: 289 ff. Thompson, 1976b: 92.
Nybakken, 1978: 135, 144. Haderlie 8 Do-
154 MCDONALD
nat, 1978: 60. McDonald 8 Nybakken,
1978: 112. Dehnel 8 Kong, 1979: 1835 ff.
Type-specimens: U. S. Nat. Mus., no.
181287.
Dorsum covered with numerous low,
rounded tubercles. Foot nearly linear, broadly
rounded and bilabiate anteriorly, and bluntly
pointed posteriorly. Labial tentacles short,
flattened, and triangular, with a groove on
external margin. Rhinophores bear 15-18 la-
mellae which are cream to light yellow. Bran-
chial plumes six, bipinnate, cream to light yel-
low. General ground color cream to very pale
yellow. Dorsal tubercles tipped with lemon
yellow, and a band of similar color occurs on
both the dorsal and ventral surfaces of man-
tle margin and also on edge of foot. T.L.: 25
mm.
Radular formula of specimen examined
89(57-63.1.57-63), MacFarland (1966: 142)
reports 90-114(47-58.1.47-58). Rachidian
teeth (Fig. 62a) bear 4-7 denticles. Inner lat-
erals (Fig. 62b) hooked and bear 4-14 denti-
cles.
Type-locality: Monterey Bay, California.
Range and habitat: Auke Bay, Alaska
(Robilliard, 1974b), to Punta San Eugenio,
Baja California, Mexico (Lance, 1961). Inter-
tidal to 45 m in rocky areas. Feeds upon the
sponges: Halichondria panicea, Higginsia Sp.,
and Myxilla incrustans (Bloom, 1976: 294).
Remarks: MacFarland (1966: 140) emend-
ed the specific epithet to /uteomarginata,
based on Doris marginata Montagu, 1804,
being a junior synonym of Cadlina laevis (Lin-
naeus, 1767). He felt that this made the epi-
thet marginata unavailable in the genus Caa-
lina. However his emendation is unjustified
because the specific epithet marginata is not
currently used in the genus Cadlina. The
present species therefore should be referred
to as Cadlina marginata MacFarland, 1905,
not as Cadlina luteomarginata MacFarland,
1905.
Cadlina modesta MacFarland, 1966
Cadlina modesta MacFarland, 1966: 140,
143, 147-152, pl. 30, figs. 14-15; pl. 33,
figs. 22-31. Sphon & Lance, 1968: 77.
Bertsch, 1969: 231-232. Roller & Long,
1969: 426. Gosliner & Williams, 1970: 170.
Schmekel, 1970: 194. Robilliard, 1971a:
163, 164. Bertsch et al., 1972: 305. Ab-
bott, 1974: 357. Thompson, 1976b: 43, 92.
Nybakken, 1978: 135, 144. Haderlie & Do-
nat, 1978: 60. McDonald & Nybakken,
1978: 110. 112:
Type-specimens: type material at Calif.
Acad. Sci.
Dorsum covered with numerous relatively
inconspicuous, low tubercles of various sizes.
Foot narrow and nearly linear, broadly round-
ed and bilabiate anteriorly, and less rounded
posteriorly. Labial tentacles triangular and
lobe-like, with a groove on external margin.
Rhinophores bear 10-12 lamellae which are
cream to light yellow. Branchial plumes 10-
12, unipinnate, cream to light yellow. General
ground color cream to light yellow, occasion-
ally tinted with orange and pink. A series of
16-20 or more small, lemon yellow dots oc-
curs longitudinally on either side of dorsum
extending anterior of rhinophores and pos-
terior of branchial plumes. T.L.: 15-20 mm.
Radular formula of specimen examined
43(28.1.28), MacFarland (1966: 148) reports
70-94(21-24.1.21-24). Rachidian teeth (Fig.
60a) bear 5-7 denticles, the median of which
is largest. First laterals (Fig. 60b) have 5-7
denticles, innermost of which is usually larg-
est. The most external laterals (Fig. 60f) bear
11-15 denticles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Nanaimo, Vancouver
Island, British Columbia, Canada (Robilliard,
1971a), to La Jolla, San Diego Co., California
(MacFarland, 1966). Intertidal to 10 m, in
rocky areas. Feeds upon the sponge Aplysil-
la glacialis (McDonald & Nybakken, 1978:
110).
Cadlina sparsa (Odhner, 1921)
Juanella sparsa Odhner, 1921: 225-226, text
fig. 3, pl. 8, figs. 13-14. Odhner, 1926a:
56-57.
Cadlina sparsa (Odhner). Odhner, 1926a: 56-
57. Marcus, 1955: 122. Marcus, 1958: 21.
Marcus, 1959: 3, 7, 27-29, 87, 93, text figs.
39-44. Lance, 1961: 66. Marcus, 1961: 15,
pl. 3, figs. 43-45. Lance, 1962a: 157. Mar-
cus 8 Marcus, 1967a: 169. Sphon 8 Lance,
1968: 77. Roller 8 Long, 1969: 426. Keen,
1971: 823. Abbott, 1974: 357. Nybakken,
1978: 135.
Type-specimens: not listed.
Dorsum covered with numerous small, low,
rounded tubercles. Foot narrow and nearly
linear, broadly rounded and bilabiate ante-
riorly, and bluntly pointed posteriorly. Labial
CALIFORNIA NUDIBRANCHS 155
tentacles tapered, somewhat triangular-con-
ical, with a groove on external margin. Rhi-
nophores bear about 8-12 lamellae, which are
light yellow to yellowish-tan. Branchial plumes
12, unipinnate, light yellow to light yellowish-
tan. General ground color cream to very light
yellow. An irregular series of 7-13 or more
small, brown to black dots with yellow cen-
ters occurs longitudinally on either side of the
dorsum, from just posterior of rhinophores to
just posterior of branchial plumes. T.L.: 15-
20 mm.
Radular formula of specimen examined
37(24.1.24), Marcus (1961: 15) reports 56-
69(28.1.1.1.28), and Odhner (1921: 225) re-
ports 54(20.1.1.1.20). Rachidian teeth (Fig.
61a) bear 3-4 relatively small denticles. First
laterals (Fig. 61b) have 6 denticles, innermost
of which is usually largest. The most external
laterals (Fig. 61e) bear up to 19 denticles.
Type-locality: Juan Fernandez Island, Chile.
Range and habitat: Monterey, Monterey
Co., California (personal observation), to San
Pedro, Los Angeles Co., California (Marcus,
1961); Juan Fernández and Chiloé Islands,
Chile (Marcus, 1961). Intertidal to 40 m, in
rocky areas. Relatively rare in California.
Cerberilla Bergh, 1873
Body aeolidiform, rather compressed dor-
so-ventrally. Foot wide, anterior foot corners
tentaculiform. Cerata usually somewhat flat-
tened and decumbent. Oral tentacles usually
quite long, cylindrical and tapered distally.
Rhinophores non-retractile and smooth.
Masticatory border of mandibles smooth.
Radula uniseriate, rachidian teeth very dis-
tinctive, each bears numerous unequal larger
denticles, between which (or on which) are
smaller denticles.
Penis unarmed. Anus pleuroproct.
Type-species: Cerberilla longicirrha Bergh,
1873.
Cerberilla mosslandica McDonald &
Nybakken, 1975
Cerberilla mosslandica McDonald 8 Nybak-
ken, 1975: 378-382, text figs. 1-2. Mc-
Donald, 1975b: 55. McDonald 8 Nybak-
ken, 1978: 115.
Type-specimens: Calif. Acad. Sci., no.
CASIZ 596.
Foot broad, rounded anteriorly and slightly
tapered posteriorly to short, abruptly pointed
tail; foot corners produced into tentaculiform
processes. Oral tentacles quite long, cylindri-
cal, and tapered to pointed tips; translucent
grayish-white and encrusted with brownish to
reddish-brown proximally, and encrusted with
numerous opaque white dots on distal half.
Rhinophores non-retractile, smooth, and ta-
pered to blunt tips; encrusted with brownish
to reddish-brown proximally and with opaque
white on tips. Cerata are slightly flattened and
decumbent, arranged in 8-10 crescent-
shaped, transverse rows on dorsum. Ante-
rior 3-4 rows have a median space which
leaves cardiac region free of cerata. General
ground color translucent grayish-white. Dor-
sum, dorsal surface of foot margin, and head
are encrusted with brownish to reddish-
brown. Cerata encrusted with brownish to
reddish-brown and tips encrusted with white.
TIE? zmm:
Anus on right, approximately ventral of
fourth row of cerata. Genital aperture on right
side, just ventral of second row of cerata. Pe-
nis unarmed.
Radular formula of specimen examined
10(0.0.1.0.0). Rachidian tooth (Fig. 96) bears
11-16 large denticles, between which occur
17-27 small denticles. Masticatory border of
mandibles smooth.
Type-locality: Monterey Bay, California.
Range and habitat: Monterey Bay, Califor-
nia (McDonald & Nybakken, 1975). Found on
a mud-sand bottom in 16-63 m. May feed
upon a burrowing sea anemone (McDonald
8 Nybakken, 1975b: 380).
Chromodoris Alder & Hancock, 1855
Body doridiform, evenly rounded anteriorly
and posteriorly, sometimes rather elongate;
usually with an ample projecting mantle mar-
gin. Dorsum soft and smooth, without tuber-
cles or papillae. Labial tentacles small and
rather conical. Rhinophores perfoliate, re-
tractile into sheaths with smooth borders.
Branchial plumes retractile, usually unipin-
nate, arranged in a circle around anus. Most
species are brightly colored.
Labial armature consists of minute rods
which are usually bifid at tip. Radula broad,
without a strongly developed rachidian tooth
(vestiges may be detectable). Laterals more
or less hamate, first laterals are each dentic-
ulate on both inner and outer margins, re-
maining laterals denticulate on outer margins
only. Outermost laterals denticulate at apex
only. Teeth do not have bifid cusps.
Penis unarmed.
156 MCDONALD
Type-species: Chromodoris magnifica
(Quoy 4 Gaimard, 1832).
The genera Chromodoris and Hypselodor-
is are sometimes difficult to separate, as
characters can overlap. | use here the dis-
tinctions put forward by Odhner (1957) and
further elaborated by Thompson (1972b).
Bertsch (1977) erected the genus Mexi-
chromis and included in it Chromodoris por-
terae. While Chromodoris is a large genus, in
need of study and possible division into sev-
eral subgenera or new genera, it would seem
unwise to erect new genera based on study
of species from a limited geographic area (i.e.
the Pacific coast of North America). The
Chromodoridinae are largely tropical and the
15 species found on the Pacific coast of North
America are but a fraction of the number of
species found worldwide. Bertsch (1977: 113)
distinguished Mexichromis with acuspid,
pectinate radular teeth from Chromodoris with
unicuspid, serrate radular teeth. However, a
cursory perusal of the literature shows that
there is gradation from one type of radular
tooth to the other. Chromodoris tryoni Gar-
rett, 1873 (Bergh, 1877a: pl. 54, fig. 2) is very
distinctly unicuspid, while Chromodoris pau-
pera Bergh, 1877 (Bergh, 1976b: pl. 53, fig.
21) and Chromodoris carnea Bergh, 1889
(Bergh, 1889: pl. 83, fig. 26) are slightly less
unicuspid. Chromodoris vicina Eliot, 1904
(Edmunds, 1971: text fig. 16E) is even less
unicuspid and Chromodoris petechialis
(Gould, 1852) (Kay & Young, 1969: text fig.
440) is acuspid. A range of variation such as
this would seem to make it difficult or impos-
sible to separate these species into two dis-
tinct genera based on this character.
Chromodoris macfarlandi Cockerell, 1901
Chromodoris mcfarlandi Cockerell, 1901c:
79-80. Cockerell, 1902: 19-21. Costello,
1947: 232. Costello, 1950: 430-431.
MacFarland, 1966: 158. Bertsch, 1976a:
157-158. Bertsch, 1977: 114, text fig. 3D.
Bertsch, 1978a: 313-317, 315, figs. 7, 13-
15, 37-40. McDonald 8 Nybakken, 1978:
112.
Chromodoris macfarlandi (Cockerell). Cock-
erell £ Eliot, 1905: 36. Kelsey, 1907: 37.
Cockerell, 1908: 106. Hewatt, 1946: 198.
Costello, 1947: 232. Costello, 1950: 430-
431. Marcus & Marcus, 1967a: 178. Lance,
1969: 37. Roller & Long, 1969: 425, 429.
Roller, 1970a: 371. Roller, 1970b: 482.
Long, 1970: 19. Hertz, 1970: 6. McBeth,
1970: 28. Schmekel, 1970: 194. North,
1971: 57. Keen, 1971: 822. Bertsch et al.,
1973: 287. Schuler, 1975: 33. Michel, 1976:
47, fig. 8. Hargens, 1977: 363, 365.
Bertsch, 1978a: 314. Bertsch, 1979b: 44,
fig. 3D.
Chromodoris (sp.?). Berry, 1907: 35.
Glossodoris macfarlandi (Cockerell). Cocker-
ell, 1908: 106. O'Donoghue, 1926: 212.
O'Donoghue, 1927a: 10. O'Donoghue,
1927b: 89-90, 116, pl. 2, figs. 33, 37. Pru-
vot-Fol, 1951b: 120. Pruvot-Fol, 1951c:
152. Lance, 1961: 66. McLean, 1962: 111.
Paine, 1963a: 4 ff. Farmer & Collier, 1963:
62. Steinberg, 1963b: 69. Farmer, 1964:
24. MacFarland, 1966: 153-157, pl. 22,
figs. 1-5; pl. 34, figs. 1-11. Sphon & Lance,
1968: 79. Roller, 1970a: 371. Abbott, 1974:
354, fig. 4239. Bloom, 1976: 292, 294.
Glossodoris mcfarlandi (Cockerell). Costello,
1947: 232.
Type-specimens: not listed.
Lateral edges of dorsum thin and undulat-
ing, projecting well beyond foot. Foot rather
elongate, rather truncate and bilabiate ante-
riorly, tapered posteriorly to blunt pointed tail.
Labial tentacles short and lobe-like, with an
obscure groove on external margin. Rhino-
phores bear 16-20 lamellae, and are pale vi-
olet on shaft and deep red-violet on clavus.
Branchial plumes 10-14, unipinnate, violet
proximally and deep violet-garnet distally.
General ground color deep, vivid violet
(somewhat paler in occasional specimens). A
narrow median stripe of yellow-gold extends
posteriorly from just anterior of rhinophores
to just anterior of branchial plumes; a similar
stripe occurs antero-laterally on either side of
dorsum, from just posterior of rhinophores to
just posterior of branchial plumes, where both
stripes unite. A similar yellow-gold band oc-
curs along margin of dorsum, and a white
band occurs just inside the yellow-gold band
around dorsum. Ventral edge of dorsum bears
a white band. T.L.: 25-35 mm.
Radular formula of specimen examined
57(50-55.1.50-55), MacFarland (1966: 155)
reports 62(47-50.1.47-50), and Bertsch
(1978a: 316) reports 36-76(16-50.1.16-50).
Rachidian teeth (Fig. 65a) smaller than later-
als, each bears a single, median denticle, and
is roughly triangular in shape. First laterals
(Fig. 65b, d) bear 3-6 denticles; the number
of denticles increases on outer laterals to
about 15 or more.
CALIFORNIA NUDIBRANCHS 157
Type-locality: San Pedro, Los Angeles Co.,
California.
Range and habitat: Monterey, Monterey
Co., California (MacFarland, 1966), to Bahía
Magdalena, Baja California, Mexico (Bertsch,
1978a). Intertidal to 27-34 m, in rocky areas.
Quite rare in northern part of range, more
common in southern California. Bloom (1976:
294) reports that it is found in association
with and may feed upon the sponges Gellius
sp. and Haliclona sp.
Remarks: See ICZN (1954, Opinion 221)
for emendation to Chromodoris macfarlandi.
Chromodoris porterae Cockerell, 1901
Chromodoris porterae Cockerell, 1901c: 79.
Cockerell, 1902: 19-21. MacFarland, 1905:
44-45. Cockerell 8 Eliot, 1905: 36. Mac-
Farland, 1906: 129-130, pl. 26, figs. 13-
14. Kelsey, 1907: 37. Baily, 1907: 92.
Guernsey, 1912: 74-75, fig. 39B. Crozier,
1922: 304. O’Donoghue, 1926: 212.
O’Donoghue, 1927b: 91-92. Costello,
1947: 232. ICZN, 1954: 118. MacFarland,
1966: 163-165, pl. 24, figs. 4-5; pl. 34,
figs. 24-31. Roller, 1970a: 371. Bertsch 8
Ferreira, 1974: 344. Schuler, 1975: 33.
Bertsch, 1976a: 158. Michel, 1976: 48, fig.
13. Bertsch, 1977: 113. Bertsch, 1978b:
80. McDonald 8 Nybakken, 1978: 112.
Glossodoris porterae (Cockerell). O'Dono-
ghue, 1926: 212. O'Donoghue, 1927a: 11.
O'Donoghue, 1927b: 91-92. Pruvot-Fol,
1951c: 134. Lance, 1961: 66. Steinberg,
1963b: 70. Sphon & Lance, 1968: 79.
North, 1971: 57. Abbott, 1974: 355, pl. 17,
no. 4252.
Glossodoris (Chromodoris) porterae (Cock-
erell). Smith & Gordon, 1948: 180.
Hypselodoris porterae (Cockerell). Lance,
1969: 37. Roller 8 Long, 1969: 425, 429.
Roller, 1970a: 371. Harris, 1973: 240.
Bertsch et al., 1973: 287.
Mexichromis porterae (Cockerell). Bertsch,
1977: 113, 114, text fig. 3J. Bertsch,
1978b: 80, 82-84, figs. 55-56. Bertsch,
1979b: 44, fig. 3J.
Type-specimens: not listed.
Foot broadly rounded and deeply notched
anteriorly, tapered posteriorly to long, point-
ed tail. Rhinophores bear 12-14 lamellae, and
are ultramarine blue on shaft and darker on
clavus. Branchial plumes 9-12, unipinnate,
ultramarine blue. General ground color ultra-
marine blue. A median stripe of pale blue ex-
tends from between the rhinophores to just
anterior of branchial plumes and continues for
a short distance just posterior of branchial
plumes; a similar stripe occurs dorso-medi-
ally on posterior portion of foot which ex-
tends beyond dorsum. A stripe of yellow-or-
ange extends longitudinally on either side of
dorsum from just posterior of rhinophores to
posterior of branchial plumes, where the
stripes meet; a line of similar color forms an
arc just anterior of rhinophores. Dorsum
edged with a narrow, white line. A number of
small, hemispherical, submarginal glandular
prominences occur posteriorly. T.L.: 15 mm.
Radular formula of specimen examined
50(38.0.38), Bertsch (1978b: 83) reports 38-
68(23-36.0.23-36). Laterals (Fig. 66) bear 5-
8 denticles on one edge, the terminal one or
two frequently longer than the remaining
denticles which are all about equal in size.
Type-locality: La Jolla, San Diego Co., Cal-
ifornia.
Range and habitat: Monterey Bay, Califor-
nia (MacFarland, 1966), to Isla Cedros, Baja
California, Mexico (Lance, 1961). Intertidal to
18 m, in rocky areas. Occasionally found on
the sponge Dysidea amblia upon which it may
feed (McDonald & Nybakken, 1978).
Conualevia Collier & Farmer, 1964
Body soft and doridiform, equally rounded
anteriorly and posteriorly. Dorsum smooth or
minutely papillate. Labial tentacles short and
stout. Rhinophores smooth, retractile below
dorsum but no sheath margin is obvious.
Branchial plumes uni- to tripinnate, arranged
in a circle around anus.
Labial cuticle thin or absent. Radular for-
mula (0.n.0.n.0); laterals hamate.
Penis unarmed.
Type-species: Conualevia marcusi Collier
8 Farmer, 1964.
Conualevia alba Collier & Farmer, 1964
Conualevia alba Collier & Farmer, 1964: 383-
385, pl. 3, text figs. 2a-d. Lance, 1966: 75.
Marcus 8 Marcus, 1967a: 186. Sphon 4
Lance, 1968: 77. Keen, 1971: 821, pl. 19,
fig. 8. Ferreira, 1972: 53-54. Abbott, 1974:
358.
Type-specimens: Calif. Acad. Sci., no. 29.
Dorsum varies from smooth to minutely
papillate. Foot rounded and bilabiate ante-
riorly and rather rounded posteriorly. Rhi-
nophores appear white, but are faintly ringed
158 MCDONALD
with minute black dots. Branchial plumes 8,
tripinnate, white. General ground color white.
An irregular row of opaque white glands oc-
curs around lateral edges of dorsum on either
side. T.L.: 12 mm.
Radular formula of specimen examined
47(68.0.68), Collier & Farmer (1964: 385) re-
port 34(56.0.56). Laterals (Fig. 68) simple
hooks.
Type-locality: Newport Bay, Orange Co.,
California.
Range and habitat: Cypress Point, Pacific
Grove, Monterey Co., California (Ferreira,
1972), to Bahia Tortugas, Baja California,
Mexico (Lance, 1966); Galapagos Islands,
Ecuador (Sphon, 1972b). Intertidal to 17 m,
in rocky areas, usually under rocks.
Corambe Bergh, 1869
Body elliptical in outline, quite flattened and
disc-like. Margin of dorsum wide and thin, ex-
tending well beyond foot; a median notch oc-
curs in posterior margin. Dorsum smooth.
Foot emarginate anteriorly and rounded pos-
teriorly. Labial tentacles short, rather blunt
lobes. Rhinophores bear an inner pair of wing-
like expansions and are retractile into
sheaths. Branchial plumes unipinnate, ar-
ranged on either side of the postero-median
anus, between dorsum and foot.
Labial cuticle bears two lateral thickenings.
Radular formula n.1.0.1.n; laterals large and
bear denticles below cusp. Marginals small
and lack denticles below cusp.
Penis unarmed.
Type-species:
Bergh, 1871.
Corambe sargassicola
Corambe pacifica MacFarland &
O'Donoghue, 1929
Corambe pacifica MacFarland & O'Dono-
ghue, 1929: 1-27, pls. 1-3. Smith & Gor-
don, 1948: 180. Marcus, 1955: 164 ff.
Marcus, 1959: 62. Steinberg, 1960: 49.
Lance, 1961: 67. Lance, 1962b: 36, 37.
Paine, 1963a: 4. Steinberg, 1963b: 71.
MacFarland, 1966: 130-132 ff, pl. 22, figs.
6-8; pl.¿29, fig. 20, pl. 32, figs. 13-14.
Marcus & Marcus, 1967b: 206 ff. Sphon 4
Lance, 1968: 77. Roller & Long, 1969: 426.
Gosliner & Williams, 1970: 177. Bernard,
1970: 84. Anderson, 1972: 19. Abbott,
1974: 365, text fig. 4342. Seed, 1976: 2.
McDonald & Nybakken, 1978: 111.
Carambe pacifica MacFarland & O'Dono-
ghue. Haderlie & Donat, 1978: 60 (lapsus).
Type-specimens: Calif. Acad. Sci., no. 634.
Foot equally rounded anteriorly and pos-
teriorly, with an antero-lateral notch. Labial
tentacles short and blunt. Rhinophores bear
a plate-like expansion on anterior and lateral
sides, extending from distal portion anterior-
ly, downward to posterior face of shaft; with-
in area enclosed by this expansion is another
pair of smaller plate-like expansions, one on
either side of shaft; a third, unpaired ridge
occurs postero-medially on shaft within area
enclosed by smaller plate-like expansions.
Rhinophores translucent grayish, stalk may
have a few spots of yellow, garnet, or black.
Branchial plumes 6-14, translucent grayish,
and may be seen through postero-median
notch in dorsum. General ground color trans-
lucent grayish. Dorsum bears numerous ir-
regular lines of pale yellow which become
small spots on central area of dorsum. Be-
tween yellow lines are indian red spots edged
with black. Scattered small black to dark
green flecks also occur, especially centrally
on dorsum. A thin, white line occurs around
margin of foot. In all, the color pattern of the
dorsum very closely resembles the colonies
of the bryozoan Membranipora spp. upon
which the animal feeds. T.L.: 5 mm.
Radular formula of specimen examined
26(6.1.0.1.6), MacFarland (1966: 132) re-
ports 38-40(4-5.1.0.1.4-5). Laterals (Fig.
39g) are large and bear 3-7 denticles below
large distal cusp. Marginals (Fig. 39a-f) de-
crease in size outward, each having a single,
simple hook.
Type-locality: Pacific Grove, Monterey Co.,
California.
Range and habitat: Nanaimo, Vancouver
Island, British Columbia, Canada (Mac-
Farland 8 O'Donoghue, 1929), to Punta San
Eugenio, Baja California, Mexico (Lance,
1961). Intertidal to 5 m, almost always found
on the kelp Macrocystis pyrifera which is en-
crusted with the bryozoan Membranipora
spp. Feeds upon Membranipora villosa
(MacFarland, 1966; MacFarland & O'Dono-
ghue, 1929); also found on Membranipora
serrilamella (Lance, 1961).
Coryphella Gray, 1850
Body aeolidiform and gradually tapered
posteriorly to tail. Foot corners somewhat
produced. Cerata cylindro-conical, pointed
apically, arranged transversely in simple or
irregularly divided rows along edge of dor-
sum, and bear cnidosacs. Edge of dorsum
CALIFORNIA NUDIBRANCHS 159
usually well defined. Oral tentacles usually
relatively long. Rhinophores non-retractile,
may be smooth, verrucose, annulate, perfo-
liate, or papillate.
Masticatory border of mandibles denticu-
late. Radula triseriate, with denticles on ra-
chidian and usually on laterals.
Penis unarmed. Anus pleuroproct.
Type-species: Coryphella rufibranchialis
(Johnston, 1832).
Coryphella cooperi Cockerell, 1901
Coryphella cooperi Cockerell, 1901a: 85-86.
O’Donoghue, 1926: 229. Marcus, 1961: 48.
Steinberg, 1961: 62. Lance, 1961: 68.
Marcus & Marcus, 1967a, 221 ff. Farmer,
1967: 342. Abbott, 1974: 374. Cooper,
1978: 8. Cooper, 1980: 284.
Type-specimens: not listed.
Foot widest and bilabiate anteriorly and ta-
pered posteriorly to pointed tail. Foot corners
prolonged into tentaculiform processes. Oral
tentacles about as long as rhinophores, cylin-
drical and tapered distally to blunt points;
translucent grayish-white, very pale orange
distally, with a few minute, opaque white dots.
Rhinophores verrucose, cylindrical, and
slightly tapered to tips; colored as oral ten-
tacles. Cerata arranged in about 5-7 groups
on dorso-lateral body margin, leaving dorsum
free medially; one group occurs anterior of
cardiac region, the others are posterior. Gen-
eral ground color translucent grayish-white
with a pink tinge, head in area of rhinophores
pale pinkish-orange. A greenish-blue patch
occurs between first and second cerata rows,
just anterior and to left of cardiac region. A
distinct, sometimes discontinuous, dorso-
medial white line occurs on dorsum. Cerata
greenish at base with white tips and a few
minute, opaque white flecks on surface, cores
reddish-brown. T.L.: 20 mm.
Anus on right, ventral of first post-cardiac
group of cerata. Genital aperture on right side,
ventral of first group of cerata.
Radular formula of specimen examined
11(0.1.1.1.0). Rachidian teeth (Fig. 100a) bear
7-10 denticles on either side of median cusp.
Laterals (Fig. 100b) roughly triangular with 7-
10 denticles below cusp.
Type-locality: San Pedro, Los Angeles Co.,
California.
Range and habitat: Elkhorn Slough, Mon-
terey Co., California (personal observation),
to Bahía San Quintín, Baja California, Mexico
(Farmer, 1967). Intertidal in bays and estu-
aries on floating docks and mudflats. Usually
on the hydroid Tubularia crocea. Cooper
(1978) states that it eats the polyps and go-
nophores of Tubularia crocea. Very rare in
California.
Coryphella iodinea (Cooper, 1863)
Aeolis (Phidiana?) iodinea Cooper, 1863a:
205. O'Donoghue, 1922d: 139.
Phidiana iodinea (Cooper). Cooper, 1863b:
60. Carpenter, 1864: 609. Cooper, 1867:
14. Cooper, 1870: 57. Bergh, 1873a: 615.
Yates, 1890: 41. O'Donoghue, 1922d: 139.
Aeolis (?Phidiana) iodinea Cooper. Carpen-
ter, 1864: 608.
Flabellina iodinea (Cooper). Bergh, 1873a:
615. Bergh, 1879b: 79-81, pl. 1, figs. 15-
17; pl. 2, fig. 16. Bergh, 1879c: 135-137,
pl. 1, figs. 15-17; pl. 2, fig. 16. Hilton, 1919:
34. O'Donoghue, 1922d: 139. O’Dono-
ghue, 1927a: 11. O'Donoghue, 1927b: 103.
Coe, 1932: 63. Ingram, 1935: 48, 49. Coe
& Allen, 1937: 126. Smith & Gordon, 1948:
181. LaRocque, 1953: 249. Goodwin &
Fox, 1955: 1086. Lance, 1961: 68. Pe-
quegnat, 1963: 427. Farmer & Collier,
1963: 63. Paine, 1963a: 4. Steinberg,
1963b: 72. Paine, 1964: 385. Farmer, 1967:
342. Marcus & Marcus, 1967a: 226. Har-
ris, 1971a: 84. North, 1971: 57. Nybakken,
1978: 133.
Coryphella iodina (Cooper). Cockerell, 1901b:
121 (lapsus).
Aeolis iodinea Cooper. Kelsey, 1907: 33.
Flabellina (Aeolis) iodinea Cooper. O'Dono-
ghue, 1922d: 138.
Coryphella iodinea (Cooper). O'Donoghue,
1922d: 139. Marcus & Marcus, 1967a:
218-220, 238, text figs. 68-69. Farmer,
1970: 78. Schmekel, 1970: 141. Abbott,
1974: 374. Bertsch & Ferreira, 1974: 347.
Keen & Coan, 1975: 45. Thompson, 1976a:
28. Thompson, 1976b: 58. Cowen & Laur,
1978: 292-294. Poorman & Poorman,
1978: 373.
Flabellinopsis iodinea (Cooper). MacFarland,
1966: 308-313, pl. 58, figs. 1-2; pl. 65,
figs. 1-8; pl. 66, figs. 1-6. Sphon & Lance,
1968: 79. Roller & Long, 1969: 427. Lance,
1969: 33. Turner et al., 1969: 136. Du-
Shane & Brennan, 1969: 361. Gosliner &
Williams, 1970: 178. Grigg & Kiwala, 1970:
151. Bernard, 1970: 86. McBeth, 1970: 28.
McBeth, 1971: 158, 159. Keen, 1971: 836-
837, 1p15 22. figs 1. Baker: 1972744 Ti.
McBeth, 1972a: 55 ff. Harris, 1973: 246,
160 MCDONALD
281. Hinegardner, 1974: 452. Kitting, 1974:
32. Michel, 1976: 47, fig. 9. McDonald 8
Nybakken, 1978: 114. Russo, 1979: 44 ff.
Flabellinopsis iodinia (Cooper). Lance, 1968:
7. McBeth, 1972a: 56 (lapsus).
Flabellina (Flabellinopsis) iodinea (Cooper).
McBeth, 1972a: 65.
Flabellina (=Flabellinopsis) iodinea (Cooper).
Harris, 1973: 239.
Flabillinopsis iodinea (Cooper). Kastendiek,
1976: 531 (lapsus).
Coryphella sabulicola Cowen & Laur, 1978:
292-294.
Type-specimens: not listed; Geological
Survey coll. (Cooper, 1863a).
Dorsum separated from foot by well de-
fined, longitudinal groove; body high and
much compressed laterally. Foot linear, quite
narrow, slightly rounded anteriorly and ta-
pered posteriorly to rather long, bluntly point-
ed tail. Oral tentacles elongate, slender, and
tapered, up to one third of body length, col-
ored as body, pale distally. Rhinophores per-
foliate with 40-80 lamellae, shaft colored as
body, clavus maroon to orange-red, with
Opaque white, antero-median ridge. Cerata
pedunculate, borne upon 5-8 or more pairs
of low, crescent-like longitudinal elevations of
the dorsum edge. General ground color
translucent purple. Cerata purple on proxi-
mal one quarter to one third, shading gradu-
ally to brilliant orange distally; cores burnt
sienna. T.L.: 30 mm.
Anus on right, postero-ventral of second
group of cerata. Genital aperture on right side,
just postero-ventral of first group of cerata.
Radular formula of specimen examined
16(0.1.1.1.0), MacFarland (1966: 310) re-
ports 17(0.1.1.1.0), and Marcus & Marcus
(1967a: 219) report 22(0.1.1.1.0). Rachidian
teeth (Fig. 101a) bear 12-16 denticles on
either side of median cusp. Laterals (Fig.
101b) have 11-22 denticles below cusp.
Masticatory border of mandibles bears nu-
merous, tubercle-like denticles.
Type-locality: San Diego Bay, California.
Range and habitat: Vancouver Island, Brit-
ish Columbia, Canada (Bernard, 1970), to
Cabo San Quintin, Baja California, Mexico
(Farmer, 1967). Intertidal to 39 m, found in
rocky areas and on pier pilings along relative-
ly open coast. McBeth (1971: 158) reports
that it feeds upon the hydroid Eudendrium
ramosum and MacGinitie & MacGinitie (1949:
76) state that it feeds to some extent on the
ascidian Diplosoma pizoni. C. iodinea is ca-
pable of swimming.
Remarks: Marcus & Marcus (1967a: 220)
state that the peduncles upon which the ce-
rata are borne, are not visible on the single
preserved specimen available to them, nor
could they see the peduncles on two color
photographs. On this basis they assigned this
species to the genus Coryphella. In the more
than 20 specimens which | examined the pe-
duncles were quite obvious; however, | feel
that the character pedunculate cerata is not
sufficient to distinguish a separate genus.
Since pedunculate cerata are the only char-
acter which separates Flabellinopsis Mac-
Farland, 1966, from Coryphella Gray, 1850,
the former is a junior synonym of the latter.
C. iodinea was also once placed in Flabellina
Voigt, 1834. However, the type-species of
Flabellina, F. affinis (Gmelin, 1791), has a
penial stylet (Bergh, 1875a: 647-651), is
acleioproct, and has a somewhat different
radula than does Coryphella which lacks a
penial stylet and is pleuroproct. Therefore,
while Coryphella and Flabellina are some-
what similar in general shape and may have
pedunculate cerata, they are not closely re-
lated.
Cowen 8 Laur (1978: 292-294) described
Coryphella sabulicola which they distin-
guished from C. iodinea almost solely on the
basis of starch-gel electrophoresis data. They
used the heads of starved animals of both
species, and found that 11 of the C. iodinea
had a cathode attracted protein which was
absent from all 12 C. sabulicola. They do not
mention what happened in the case of the
other specimen of C. iodinea. A morphomet-
ric comparison was made of 8 specimens of
both species in which total numbers of cerata
on the left side, total length, oral tentacle
length, number of cerata groups, and lengths
of longest and shortest cerata were record-
ed. Only the total number of cerata relative
to length showed any significant difference
between the two species. Number of cerata
is not normally (if ever) used to differentiate
species, as it is highly variable. The number
of denticles on the rachidian and on the lat-
eral teeth, as well as number of rhinophore
lamellae are not significantly different in the
two species. The somewhat lighter body col-
or of C. sabulicola and the other slight differ-
ences, including habitat, certainly do not seem
sufficient to separate two species. In addi-
tion, information concerning the reproductive
system, presence or absence of a penial sty-
let, location of anus, or nature of masticatory
border of the mandibles, all of which are
CALIFORNIA NUDIBRANCHS 161
characters normally used to differentiate
species of aeolids, are not given. They also
place C. sabulicola in the acleioproct family
Flabellinidae rather than in the pleuroproct
family Coryphellidae which contains all the
other species of Coryphella.
Certainly, the intraspecific variation in elec-
trophoretic data is not yet very well under-
stood. It therefore seems quite unadvisable
to use such data as virtually the only basis
for separating two species. Certainly the slight
morphological differences between the two
species are well within the limits of variation
found in many aeolids, and could easily be
the result of different habitats or food. There-
fore, C. sabulicola is a junior synonym of C.
iodinea.
Coryphella pricei MacFarland, 1966
Coryphella pricei MacFarland, 1966: 313-318
ff, pl. 58, fig. 6; pl. 65, figs. 9-13; pl. 66,
figs. 8-9. Marcus & Marcus, 1967a: 223.
Gosliner & Williams, 1970: 177. Schmekel,
1970: 141. Miller, 1971: 315. Abbott, 1974:
374. Lambert, 1976: 295, 296.
Type-specimens: type material at Calif.
Acad. Sci.
Foot widest anteriorly and tapered poste-
riorly to pointed tail; margins of foot rather
wide and distinct from sides of body. Foot
corners rather stout, tapered, and pointed
distally, ventral surface of each grooved. Oral
tentacles slender and tapered to blunt tips,
longer than rhinophores, translucent grayish
with encrusting, opaque white dots. Rhino-
phores annulate with 8-11 complete annuli
which alternate with more or less incomplete
annuli which occur on posterior face of cla-
vus; rhinophores taper to blunt tip and are
translucent grayish with pale yellow-green on
distal half. Cerata slightly flattened laterally
and directed obliquely posteriorly, arranged
in 9-12 or more transverse rows dorso-lat-
erally on dorsum. Rows posterior of cardiac
region well separated from each other in
crescent-shaped rows, each of which is on a
common, elevated base. The 5-8 rows which
are anterior of cardiac region are less dis-
tinctly separated. General ground color
translucent grayish-white. An irregularly bro-
ken band of opaque white occurs dorsally on
edge of foot. Occasional specimens have a
pale orange spot dorsally on head. Cerata
tipped with opaque white and bear a subter-
minal band of rich brown which grades into
yellow below; cores deep olive-green to
brownish-green. T.L.: 22 mm.
Anus located dorso-laterally on right, just
above lowermost cerata bases of first post-
cardiac row of cerata. Genital aperture on
right side, ventral of pre-cardiac group of cer-
ata. Penis short and bluntly conical.
Radular formula of specimens examined
21-25(0.1.1.1.0), MacFarland (1966: 315) re-
ports 19(0.1.1.1.0). Rachidian teeth (Fig.
102a) bear 6-9 denticles on either side of
median cusp. Laterals (Fig. 102b) without
denticles and in form of broadly triangular
plates. Masticatory border of mandibles bears
a row of approximately 26 blunt denticles.
Type-locality: Monterey Bay, California.
Range and habitat: Pearse Island, British
Columbia, Canada (Lambert, 1976), to Point
Fermin, San Pedro, Los Angeles Co., Califor-
nia (Sphon, 1972b). Intertidal to 19 m. A rel-
atively rare species in California, taken oc-
casionally in rocky intertidal areas. One
specimen was found crawling on hydroids
upon which it probably feeds.
Coryphella trilineata O'Donoghue, 1921
Coryphella trilineata O'Donoghue, 1921: 197-
198, pl. 3 (9), figs. 29-30. O'Donoghue,
1924: 25. O'Donoghue, 1926: 229. La-
Rocque, 1953: 250. Marcus, 1961: 48.
Steinberg, 1963a: 66. Steinberg, 1963b:
72. Lance, 1966: 69. Marcus & Marcus,
1967a: 222, 226. Hurst, 1967: 263. Sphon
8 Lance, 1968: 77. Haderlie, 1969: tab. 1.
Roller & Long, 1969: 426. Long, 1969c:
232. Gosliner & Williams, 1970: 177. Ber-
nard, 1970: 86. Keen, 1971: 836. Bridges
8 Blake, 1972: 293-297. Bertsch et al.,
1972: 305. Sphon, 1972a: 155. Bertsch 8
Ferreira, 1974: 347. Nybakken, 1974: 371.
Abbott, 1974: 374. Lambert, 1976: 297.
McDonald 8 Nybakken, 1978: 114. Coo-
per, 1978: 8. Cooper, 1980: 284.
Coryphella piunca Marcus, 1961: 47-49, 57,
pl. 9, figs. 161-167. Lance, 1961: 68.
Farmer & Collier, 1963: 63. Steinberg,
1963a: 66. Lance, 1966: 69. Marcus 4
Marcus, 1967a: 222, 226. Schmekel, 1970:
141. Miller, 1971: 315.
Coryphella fisheri MacFarland, 1966: 318-
322, pl. 58, figs. 3-5; pl. 65, figs. 14-18;
pl. 66, figs. 10-20. Marcus & Marcus,
1967a: 226. Roller, 1970a: 372. Edmunds,
1970: 47. Miller, 1971: 315. Schuler, 1975:
33.
Coryphella fischeri MacFarland. Marcus,
1970: 213 (lapsus).
162 MCDONALD
Type-specimens: Museum of Dominion
Biol. Stat., Nanaimo, British Columbia, Can-
ada.
Foot narrow, widest anteriorly; margins of
foot narrow, distinct from sides of body. Foot
corners short and rather triangular. Oral ten-
tacles of moderate length and tapered to blunt
tips, translucent grayish, yellow to deep cad-
mium orange on distal half of most speci-
mens. Rhinophores annulate with 7-10 com-
plete annuli which alternate with more or less
incomplete annuli which occur on posterior
face of clavus; rhinophores tapered to blunt
tips and translucent grayish with yellow to
deep cadmium orange on distal half. Cerata
arranged in 6-8 transverse rows dorso-lat-
erally on either side of dorsum, leaving mid-
dorsal area free; first row occurs anterior of
cardiac region, while remaining rows occur
posterior of cardiac region. General ground
color translucent grayish-white. A narrow,
opaque white line extends longitudinally from
tip of each oral tentacle, along its mid-dorsal
surface to its base, where the two unite and
pass posteriorly, mid-dorsally to tip of tail;
similar lines extend longitudinally on either
side of body, just ventral of cerata, from first
row of cerata to just posterior of last row of
cerata, where they unite with the median
white line. Cerata cores usually red-orange,
deepening in intensity distally to cadmium
yellow tip. T.L.: 22 mm.
Anus located laterally on right, just ventral
of first post-cardiac row of cerata. Genital
aperture on right side, just ventral of most
anterior row of cerata. Penis bluntly conical.
Radular formula of specimen examined
20(0.1.1.1.0), MacFarland (1966: 320) re-
ports 18(0.1.1.1.0), and Marcus (1961: 47)
reports 16-25(0.1.1.1.0). Rachidian teeth (Fig.
103a) bear 5-8 denticles on either side of
median cusp. Laterals (Fig. 103b) broadly tri-
angular with 5-12 denticles below cusp.
Masticatory border of mandibles bears nu-
merous blunt, conical denticles.
Type-locality: Nanoose Bay, Vancouver Is-
land, British Columbia, Canada.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976), to Los
Coronados Islands, Baja California, Mexico
(Lance, 1961). Intertidal to 50 m. Frequently
found in rocky intertidal, often on the hydroid
Eudendrium sp. Also sporadically abundant
on floating docks in bays, usually on the hy-
droid Tubularia crocea, the polyps and gono-
phores of which it eats according to Cooper
(1978).
Coryphella sp.
Foot narrow, rather truncated anteriorly and
tapered posteriorly to rather long tail. Foot
corners prolonged into tentaculiform pro-
cesses. Oral tentacles rather long, cylindrical,
and tapered to pointed tips, translucent gray-
ish-white, encrusted with opaque white flecks
distally. Rhinophores verrucose, translucent
grayish-white with a slight brownish-orange
hue and encrusted with minute white flecks
distally. Cerata arranged in 6-8 transverse
rows on dorsum, borne upon low, longitudi-
nal ridges. Anterior rows of cerata separated
medially, leaving cardiac area free. General
ground color translucent grayish-white. An
opaque white line extends from base of either
oral tentacle to between rhinophores, where
they unite and continue dorso-medially to tip
of tail as an irregular, often discontinuous line
which is rather broad in cardiac region. Dor-
sum also bears a number of irregular, opaque
white dots. Cerata bear numerous opaque
white dots, tips translucent grayish-white,
cores pinkish-brown distally and greenish-
brown proximally. T.L.: 25 mm.
Anus on right side, ventral and just poste-
rior of cardiac region. Genital aperture on right
side, midway between first group of cerata
and cardiac region.
Radular formula of specimens examined
15-17(0.1.1.1.0). Rachidian teeth (Fig. 104a)
bear 7-9 denticles on either side of median
cusp. Laterals (Fig. 104b) have 11-13 denti-
cles below cusp.
Range and habitat: This species has been
found only during October and November of
1972, in Elkhorn Slough, Monterey Co., Cal-
ifornia, on the hydroid Tubularia crocea on
floating docks (personal observation).
Remarks: These specimens may be found
to be a slight variation of Coryphella cooperi.
However, they lack the greenish-blue patch
between the first and second cerata rows
which is typical of С. соореп; they also have
11-13 denticles on the laterals while C.
cooperi has 7-10.
Crimora Alder & Hancock, 1862
Body limaciform and elongate. Dorsum
bears dorsal processes. Frontal veil bears
numerous, usually bifid or compound pro-
cesses. Labial tentacles short and blunt.
Rhinophores perfoliate and retractile into
sheaths. Branchial plumes 3, non-retractile,
bi- or tripinnate, arranged in an arch around
anus.
CALIFORNIA NUDIBRANCHS 163
Mandibles absent, but labial cuticle pres-
ent. Radula bears a number of types of teeth.
Rachidian tooth spurious or absent. Inner lat-
erals triangular, laterals hooked distally, inner
marginals roughly rectangular and bear a
cusp, outer marginals very elongate and bear
small denticles.
Penis armed with small hooks.
Type-species: Crimora papillata Alder &
Hancock, 1862.
Crimora coneja Marcus, 1961
Crimora coneja Marcus, 1961: 25-26, 57, pl.
5, figs. 77-83. Lance, 1961: 67. Paine,
1963a: 4. Lance, 1969: 36. Abbott, 1974:
361.
Type-specimens: Dept. Zool., Fac. Philos.,
Sci., Let., Univ. Sáo Paulo, Brazil (Marcus,
personal communication).
Dorsum and dorsal surface of tail bear nu-
merous rather long processes of various sizes
which may be simple, bifid, or slightly
branched distally. Frontal margin slightly ex-
panded into a veil-like process which bears
along its margin about 10 processes which
are slightly branched distally, and about 10
shorter processes which are very slightly or
not at all branched distally; both sets of pro-
cesses are orange distally. Foot rather nar-
row, bilabiate anteriorly and tapered poste-
riorly to long, pointed tail. Rhinophores bear
about 9-12 lamellae, clavus inclined slightly
posteriorly and orange, shaft translucent
grayish-white. Branchial plumes tripinnate,
translucent grayish-white. General ground
color translucent grayish-white. Some dorsal
processes tipped with dark brown to black,
and may bear a subterminal band of orange,
others tipped with orange. T.L.: 10 mm.
Radular formula of specimen examined
45(15.7-8.1.(1).1.7-8.15), Marcus (1961: 25)
reports 53(9.6.2.(1).2.6.9). Spurious rachidi-
an plate does not constitute a true tooth and
is quite difficult to see. First laterals, which
are small, rounded triangles, were not visible
on the single radula available for study. Sec-
ond laterals (Fig. 49e) relatively large, hooked
with a large, blunt denticle near base. Inner
marginals (Fig. 49c, d) roughly rectangular
with a blunt cusp. Outer marginals (Fig. 49a,
b) very elongate, rod-like, slightly curved, and
minutely denticulate.
Type-locality: Point Loma, San Diego Co.,
California.
Range and habitat: Point Loma, San Diego
Co., California (Marcus, 1961). Also, a spec-
imen identified from a photograph, collected
near Humboldt Bay, Humboldt Co., California
(personal observation). Intertidal in rocky
areas, very rare.
Cumanotus Odhner, 1907
Body rather aeolidiform, rather high and
short. Foot rather wide, and somewhat trun-
cate anteriorly and tapered posteriorly to
rather long tail. Anterior foot corners slightly
produced. Cerata quite long, cylindro-conical,
rather wide at base and tapered distally; first
rows occur anterior of rhinophores. A short,
truncated frontal veil bears the nearly rudi-
mentary, short oral tentacles at its antero-
distal corners. Rhinophores non-retractile,
arising close together on a slight prominence,
long, cylindrical, smooth, and tapered to
pointed tips.
Masticatory border of mandibles denticu-
late. Radula triseriate, rachidian teeth each
bear a rather long median denticle with small-
er denticles on either side, laterals denticu-
late on inner margin.
Penis unarmed. Bursa copulatrix bears on
the upper and lower margins a circular pad
armed with about 12 small cones which ter-
minate in small hooks. Anus pleuroproct.
Type-species: Cumanotus beaumonti (Eliot,
1906).
See ICZN (1966, Opinion 783), Cumanotus
placed on Official List.
Cumanotus beaumonti (Eliot, 1906)
Coryphella beaumonti Eliot, 1906c: 361-363,
pl. 11, figs. 15-16. Lemche, 1964c: 125,
126.
Cumanotus laticeps Odhner, 1907: 26-29.
Odhner, 1910: 82-84. Eliot, 1910: 6. White,
1938: 17. Lemche, 1964c: 125.
Cumanotus beaumonti (Eliot). Eliot, 1908:
313-314. Eliot, 1910: 125-127, 169, pl. 8,
figs. 1-5. Odhner, 1910: 82-84. Iredale 8
O'Donoghue, 1923: 209. Cuenot, 1927:
264. Mar. Biol. Assoc., 1931: 270. Winck-
worth, 1932: 237. White, 1938: 17.
Lemche, 1938: 38. Pruvot-Fol, 1954: 424—
425. Mar. Biol. Assoc., 1957: 315. Thomp-
son, 1964: 294. Hurst, 1967: 255 ff, text
fig. 14, pl. 30, fig. 17; pl. 35, fig. 39; fig.
24-6. Bebbington & Thompson, 1968: 10.
Roller, 1970b: 482. Gosliner & Williams,
1970: 177. Robilliard, 1971a: 164. Thomp-
son & Brown, 1976: 172, fig. 92. Thomp-
son, 1976a: 28, text fig. 38f. Thompson,
1976b: 55, 60, 62, 80, fig. 4. McDonald 8
164 MCDONALD
Nybakken, 1978: 114, 116. Cooper, 1978:
8. Cooper, 1980: 284. Tardy 8 Gantes,
1980: 204 ff.
Cumanotus species. Sphon & Lance, 1968:
78.
Type-specimens: not listed.
Foot rather wide and somewhat truncate
anteriorly, tapered posteriorly to rather long,
pointed tail. Foot corners produced into ex-
tremely short processes. Rhinophores trans-
lucent grayish-white, with slight orange tint.
Cerata quite long, cylindrical, and tapered to
fine points; arranged in about 7-8 poorly de-
fined rows dorso-laterally on dorsum. Gen-
eral ground color translucent grayish-white.
A few reticulate bands of yellowish-brown to
bluish-green occur on dorsum. Cerata trans-
lucent grayish-white, with a subapical band
of opaque white; cores yellowish-brown to
bluish-green. T.L.: 8 mm.
Anus dorso-lateral on right, anterior of sev-
enth cerata row.
Radular formula of specimen examined
12(0.1.1.1.0). Rachidian teeth (Fig. 99a) bear
8-12 denticles on either side of median cusp.
Laterals (Fig. 99b) somewhat triangular with
a hooked cusp distally, and about 18 denti-
cles. Masticatory border of mandibles bears
2-3 rows of denticles.
Type-locality: Barn Pool, Plymouth, En-
gland.
Range and habitat: San Juan Islands, Pu-
get Sound, Washington (Hurst, 1967), to San
Diego, San Diego Co., California (Roller,
1970b); England (Eliot, 1906c). This species
was probably introduced along with Tubular-
га crocea. Intertidal to 5 m, usually on bay
boat landings. Almost always found upon the
hydroid Tubularia crocea upon which it feeds,
and whose polyps it rather resembles (Mc-
Donald & Nybakken, 1978; Cooper, 1978).
Remarks: The specimens of Cumanotus
which are found in California do not perfectly
match the description of Cumanotus beau-
топи, but until specimens from England can
be compared with specimens from the Pacif-
ic coast of North America, it seems best to
refer the specimens from North America to
C. beaumonti, rather than name a new
species.
Cuthona Alder 4 Hancock, 1855
Body aeolidiform, somewhat compressed,
and tapered posteriorly. Foot narrow, linear,
rounded anteriorly and tapered posteriorly;
foot corners rounded, usually slightly ex-
panded. Cerata cylindrical, tapered to point-
ed tips, and arranged in transverse rows dor-
so-laterally on dorsum. Oral tentacles
cylindrical and tapered to blunt tips. Rhino-
phores non-retractile, usually smooth, and
slightly tapered to blunt tips.
Masticatory border of mandibles bears
denticles or numerous minute bristles. Rad-
ula uniseriate, rachidian teeth each bear a
strong median cusp which may or may not
be raised above the lateral denticles; smaller
denticles may be present between the lateral
denticles.
Penis may be armed with a stylet or un-
armed. Anus acleioproct.
Miller (1977) combined Catriona Winck-
worth, 1941, and Trinchesia lhering, 1879,
and a number of smaller genera under the
name Cuthona, stating that the characters
used to separate the two genera (mandible
dentition, cusp of rachidian tooth, penial sty-
let, etc.) overlap in the various species such
that there is no good set of characters to
separate Trinchesia from Catriona.
Type-species: Cuthona nana (Alder & Han-
cock, 1842).
See ICZN (1966a, Opinion 773), Cuthona
placed on Official List.
Cuthona abronia (MacFarland, 1966)
Cratena abronia MacFarland, 1966: 347-351,
pl. 59, figs. 3-4; pl. 68, figs. 18-22; pl. 70,
figs. 1-5. Long, 1969b: 281. Roller, 1970a:
372.
Trinchesia abronia (MacFarland). Roller,
1969b: 421, text fig. 1. Roller & Long, 1969:
428. Roller, 1970a: 372. Gosliner & Wil-
liams, 1970: 179. Robilliard, 1971a: 164,
165. Bertsch et al., 1972: 308. Sphon,
1972a: 156. Nybakken, 1974: 371. Abbott,
1974: 376, text fig. 4449.
Trinchesia (=Cratena) abronia (MacFarland).
Robilliard, 1971a: 164.
Cuthona abronia (MacFarland). Williams &
Gosliner, 1979: 208, 215.
Type-specimens: type material at Calif.
Acad. Sci.
Sides of body distinct from margins of foot.
Oral tentacles bear a band of purple midway
along their length, with a pale yellow ring
proximal of this and also encrusting distal one
third of tentacles, frequently with diffuse pur-
ple proximally. Rhinophores smooth, colored
as oral tentacles. Cerata erect, cylindrical, and
tapered to blunt tips, arranged in about 9
transverse rows dorso-laterally on body mar-
CALIFORNIA NUDIBRANCHS 165
gin. General ground color translucent gray-
ish-white, dorsum encrusted with a few
opaque white to pale yellow dots, a blotch of
the same color occurs just posterior of rhi-
nophores and a line of similar color occurs
dorsomedially on tail. Cerata bear about 3,
sometimes incomplete, narrow bands of en-
crusting white to pale yellow which form
boundaries between colored segments of
cores. Cores pale yellow in distal third, me-
dial third olive green to brown, and proximal
third maroon to carmine. T.L.: 8 mm.
Anus on right, just posterior of cardiac re-
gion. Genital aperture on right side, just ven-
tral of second row of cerata. Penis armed with
chitinous stylet.
Radular formula of specimen examined
25(0.0.1.0.0), MacFarland (1966: 348) re-
ports 33(0.0.1.0.0). Rachidian teeth (Fig. 116)
bear 4-9 large denticles on either side of me-
dian cusp, with lesser denticles between the
larger denticles. Masticatory border of man-
dibles bears a single series of 18-28 denti-
cles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Mukkaw Bay, Wash-
ington (Robilliard, 1971a), to Pismo Beach,
San Luis Obispo Co., California (Long,
1969b). Usually found in rocky intertidal zone
on hydroids.
Cuthona albocrusta (MacFarland, 1966)
Cratena albocrusta MacFarland, 1966: 340-
344, pl. 61, figs. 1-4; pl. 67, figs. 13-22;
pl. 69, figs. 4-5a. Hurst, 1967: 255 ff, text
figs. 26a-b, 29; fig. 25-1. Haderlie, 1968:
333 ff. Sphon & Lance, 1968: 78. Roller,
1970a: 372. Robilliard, 1971a: 164.
Trinchesia albocrusta (MacFarland). Roller,
1969b: 421, text fig. 1. Roller & Long, 1969:
428. Haderlie, 1969: tab. 1. Roller, 1970a:
372. Gosliner & Williams, 1970: 179. Ro-
billiard, 1971a: 165. Bertsch et al., 1972:
308. Sphon, 1972a: 156. Nybakken, 1974:
371. Abbott, 1974: 376, text fig. 4443.
Haderlie & Donat, 1978: 52, 60. Cooper
1978: 8.
Trinchesia (=Cratena) albocrusta (Mac-
Farland). Robilliard, 1971a: 164.
Catrena albocrusta. Thiriot-Quiévreux, 1977:
186 (lapsus).
Cuthona albocrusta MacFarland. Williams &
Gosliner, 1979: 208, 215. Cooper, 1980:
284.
Type-specimens: type material at Calif.
Acad. Sci.
Sides of body distinctly separated from foot
by a longitudinal groove. Oral tentacles
translucent grayish-white with a few scat-
tered flecks of opaque white. Rhinophores
smooth, bearing various amounts of en-
crusting white distally. Cerata cylindrical,
somewhat inflated, pointed at tips, and de-
cumbent; arranged in 6-7 transverse rows
dorso-laterally on body margin. General
ground color translucent grayish-white. Dor-
sum encrusted with opaque white extending
from head to posterior row of cerata, and
from there postero-medially, as a broken line,
to tail. Irregular spots of opaque white are
scattered on sides of body. Cerata encrusted
with opaque white on distal two thirds, leav-
ing tips free, cores vary from pale or deep
green to pale or deep raw umber. T.L.: 5 mm.
Anus on right, just anterior of innermost
ceras of first post-cardiac group. Genital ap-
erture on right side, between rhinophores and
first row of cerata. Penis armed with chitin-
ous stylet.
Radular formula of specimen examined
49(0.0.1.0.0), MacFarland (1966: 342) re-
ports 56-70(0.0.1.0.0). Rachidian teeth (Fig.
117) bear 4-5 denticles on either side of me-
dian cusp. Masticatory border of mandibles
bears a single row of about 18 triangular
denticles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Friday Harbor, Wash-
ington (Hurst, 1967), to Point Fermin, Palos
Verdes Peninsula, Los Angeles Co., Califor-
nia (Sphon, 1972b). Intertidal to 30 m, usually
in rocky intertidal areas, especially on hy-
droids at base of the algae Laminaria spp.;
also occasional on floating docks in bays.
Cooper (1980) reports that it eats Tubularia
crocea.
Cuthona cocoachroma
Williams 8 Gosliner, 1979
Cuthona cocoachroma Williams & Gosliner,
1979: 203-215, text figs. 1A, 3, 4, 5, 6A.
Type-specimens: Calif. Acad. Sci., no.
CASIZ 00715, 00716.
Oral tentacles encrusted with opaque white
dorsally. Rhinophores smooth, with opaque
white on the distal half. Cerata digitiform,
elongate, and somewhat acute apically, ar-
ranged in 9-11 rows on each side of the body
with 1-7 cerata per row. General ground col-
or translucent white. Cerata tipped with
opaque white, cores vary from deep reddish-
brown to dark chocolate brown. T.L. 12 mm.
166 McDONALD
Anus on right, antero-dorsal of fourth or
fifth row of cerata, posterior of inter-hepatic
space. Genital aperture on right side be-
tween and ventral of the first and second
rows of cerata. Penis unarmed.
Radular formula 20-39(0.0.1.0.0), rachidi-
an teeth bear 4-5 lateral denticles on either
side of median cusp (Williams & Gosliner,
1979). Masticatory border of mandibles bears
a single row of approximately 16 triangular
or rounded denticles.
Type-locality: Coleman Beach, Sonoma
Coast State Park, Sonoma Co., California.
Range and habitat: Known only from Dux-
bury Reef, Marin Co., California, and Cole-
man Beach, Sonoma Co., California. Found
in lower rocky intertidal zone.
Cuthona columbiana (O'Donoghue, 1922)
Amphorina (?) sp. Eliot, 1913: 4, 43. Baba,
1937b: 328.
Amphorina columbiana O'Donoghue, 1922b:
160-161, 165, pl. VI, figs. 23-24. Ed-
munds, 1968a: 207. Williams & Gosliner,
1979: 212.
Galvina columbiana O'Donoghue, 1922b: 161.
Cuthona (Cuthona) sp. Eliot. Baba, 1937b:
328.
Cratena columbiana (O'Donoghue). La-
Rocque, 1953: 249. Bernard, 1970: 85.
Catriona columbiana (O'Donoghue). Marcus,
1958: 50. Marcus, 1961: 52. Edmunds,
1968a: 207. Williams & Gosliner, 1979: 208,
212213244.
Cuthona alpha Baba 8 Hamatani, 1963b:
339-343, pl. 11, figs. 1-12. Lance, 1966:
79. Sphon & Lance, 1968: 78. Edmunds,
1968a: 207. Roller, 1969b: 422. Miller,
1977: 197, 198, 207-211, text figs. 3-4,
pl. 1, fig. В. Williams 8 Gosliner, 1979: 212.
Cratena spadix MacFarland, 1966: 351-354,
pl. 60, fig. 4; pl. 68, figs. 12-17; pl. 69,
figs. 6-7a. Sphon 8 Lance, 1968: 78. Ed-
munds, 1968a: 207. Roller, 1969b: 422-
423. Roller, 1970a: 372. Williams & Gosli-
ner, 1979: 212.
Catriona alpha (Baba & Hamatani). Ed-
munds, 1968a: 207. Roller, 1969b: 422-
423. Roller & Long, 1969: 426. Roller,
1970a: 372. Gosliner & Williams, 1970: 177.
Robilliard, 1971a: 163-165. Sphon, 1972a:
155. Nybakken, 1974: 371. Abbott, 1974:
376, fig. 4439. Lambert, 1976: 295, 296.
McDonald & Nybakken, 1978: 114. Coo-
per, 1978: 8. Williams & Gosliner, 1979:
212.
Cuthona columbiana (O'Donoghue). Cooper,
1980: 284.
Type-specimens: not listed.
Dorsum separated from foot by a shallow,
longitudinal groove. Oral tentacles about
same length as rhinophores, tapered to blunt
points and white distally and proximally with
a wide, red-orange band on medial three
quarters. Rhinophores smooth, distal two
thirds red-orange, with white tips, proximal
third white. Cerata arranged in 9-14 trans-
verse rows, recumbent and overlapping. Four
rows of cerata occur anterior of cardiac re-
gion and remainder occur posterior of car-
diac region. General ground color translucent
grayish-white. Opaque white flecks occur on
head and extend in a line along dorsal sur-
face of oral tentacles. A broad band of en-
crusting white occurs from base to tip of each
ceras, becoming wider distally until it covers
distal third of each ceras; core brown (burnt
sienna to deep burnt umber). Occasional
specimens collected from floating docks in
bays and estuaries may be somewhat paler,
especially with respect to red-orange of rhi-
nophores and oral tentacles and opaque
white of cerata. T.L.: 8 mm.
Anus dorso-lateral, on right, just posterior
of cardiac elevation. Genital aperture on right
side, ventral of first and second rows of ce-
rata. Penis conical and somewhat curved,
armed with a very small, non-protruding chi-
tinous stylet.
Radular formula of specimen examined
66(0.0.1.0.0), MacFarland (1966: 352) re-
ports 127 (0.0.1.0.0), and Baba & Hamatani
(1963b: 340) report about 80(0.0.1.0.0), while
O'Donoghue (1922b: 161) reports 65-
69(0.0.1.0.0). Median denticle of rachidian
teeth (Fig. 118) raised well above level of lat-
eral denticles which number about 2-6 on
either side, there may be very small denticles
between the larger lateral denticles. Masti-
catory border of mandibles bears numerous,
minute, rod-like structures, the surface of
which bears minute spines.
Type-locality: Gabriola Pass, Gabriola Is-
land, British Columbia, Canada.
Range and habitat: Pearse Island, British
Columbia, Canada (Lambert, 1976), to Mis-
sion Bay, San Diego, San Diego Co., Califor-
nia (Lance, 1966); Japan (Baba 8 Hamatani,
1963b); New Zealand (Miller, 1977). Intertidal
to 30 m, most frequently found on floating
docks in bays, frequently on the hydroid Tu-
bularia crocea, upon which it feeds (Cooper,
CALIFORNIA NUDIBRANCHS 167
1980). Feeds upon Tubularia marina and
Tubularia sp. (Robilliard, 1971a), also found
on Tubularia larynx, Obelia sp., and Synco-
ryne eximia (Miller, 1977; Robilliard, 1971a).
This species is rather rare in California.
Remarks: The radula of Cuthona alpha
Baba & Hamatani (1963b: pl. 11, fig. 3) is
virtually identical to that of Amphorina colum-
biana O'Donoghue (1922b: pl. 6, fig. 24). Al-
though O'Donoghue's description of the col-
oration of A. columbiana is very incomplete,
it approaches the description of C. alpha.
O’Donoghue (1922b: 161) states that the pe-
nis of A. columbiana is armed with a small,
somewhat flattened spine. The penis of C.
alpha was originally described as unarmed;
however, upon re-examination of paratype
material by Dr. Baba (Roller, 1969b: 422) a
short non-protruding stylet was found. Since
there are no significant differences between
the two, A. columbiana is conspecific with C.
alpha, the latter being a junior subjective syn-
onym of the former, the proper name is
therefore Cuthona columbiana (O'Donoghue,
1922) by the law of priority. Williams & Gos-
liner (1979: 212-213) also agree that A. co-
lumbiana and С. alpha are conspecific.
Cuthona flavovulta (MacFarland, 1966)
Cratena flavovulta MacFarland, 1966: 336-
ЗОБ: 2 pi 67, 805: 7, 12269,
figs. 2-2a. Roller, 1970а: 372.
Trinchesia flavovulta (MacFarland). Roller,
1969b: 421, text fig. 1. Roller & Long, 1969:
428. Roller, 1970a: 372. Gosliner & Wil-
liams, 1970: 179. Nybakken, 1974: 371.
Abbott, 1974: 376, fig. 4446.
Cuthona flavovulta (MacFarland). Williams &
Gosliner, 1979: 208, 215.
Type-specimens: type material at Calif.
Acad. Sci.
Sides of body distinctly separated from foot
by longitudinal groove. Oral tentacles light or-
ange to red on proximal third, and distal two
thirds encrusted with white. Rhinophores
smooth, light orange to red on proximal
fourth, and distal three fourths encrusted with
white. Cerata cylindrical, slightly inflated, ta-
pered to blunt tips, and rather erect; ar-
ranged in 9 or more transverse rows dorso-
laterally on body margin. General ground col-
or translucent grayish-white to cream. A light
orange blotch covers front of head, extend-
ing onto rhinophores and oral tentacles. An
opaque white line occurs dorso-medially on
tail. Cerata encrusted with opaque white
flecks distally, and a line of orange occurs
longitudinally on lateral edge of each ceras;
cores usually brownish, rarely green. T.L.: 8
mm.
Anus located on right, just anterior of in-
nermost ceras of first post-cardiac group.
Genital aperture on right side, ventral of sec-
ond row of cerata. Penis armed with chitin-
ous stylet.
Radular formula of specimen examined
66(0.0.1.0.0), MacFarland (1966: 336) re-
ports 70(0.0.1.0.0). Rachidian teeth (Fig. 119)
bear 5-7 denticles on either side of median
cusp. Masticatory border of mandibles bears
a single row of rounded denticles.
Type-locality: Monterey Bay, California.
Range and habitat: Palomarin, Marin Co.,
California (Gosliner 4 Williams, 1970), to Shell
Beach, San Luis Obispo Co., California (Rol-
ler £ Long, 1969). Intertidal in rocky areas,
usually on hydroids at base of the algae
Laminaria spp.
Cuthona fulgens (MacFarland, 1966)
Cratena fulgens MacFarland, 1966: 337-340,
pl. 60, fig. 3; pl. 67, figs. 8-11; pl. 69, figs.
3-3a. Roller, 1970a: 372.
Trinchesia fulgens (MacFarland). Roller,
1969b: 421, text fig. 1. Long, 1969a: 10.
Roller, 1970a: 372. Roller, 1970b: 483.
Gosliner 8 Williams, 1970: 179. Nybakken,
1974: 371. Abbott, 1974: 376, fig. 4447.
Cuthona fulgens (MacFarland). Williams 4
Gosliner, 1979: 208, 215.
Type-specimens: type material at Calif.
Acad. Sci.
Sides of body distinct from margins of foot.
Oral tentacles translucent grayish-white and
encrusted with opaque white on distal por-
tion. Rhinophores smooth, colored as oral
tentacles. Cerata cylindrical, tapered to blunt
tips, and slightly decumbent; arranged in 6-
8 transverse rows dorso-laterally on either
side of dorsum. General ground color trans-
lucent grayish-white to pale cream. An irreg-
ular line of opaque white may occur dorso-
medially on tail. Cerata bear encrusting white
dots on surface; cores yellow-brown, raw
umber, or dark brown medially with a yellow
band both proximal and distal of medial brown
area, tips white. T.L.: 6 mm.
Anus on right, just anterior of innermost
ceras of first post-cardiac group. Genital ap-
erture on right side, ventral of first and sec-
168 MCDONALD
ond groups of cerata. Penis armed with chi-
tinous stylet.
Radular formula of specimens examined
16-25(0.0.1.0.0), MacFarland (1966: 338) re-
ports 59(0.0.1.0.0). Rachidian teeth (Fig. 120)
bear 5-6 denticles on either side of median
cusp. Masticatory border of mandibles bears
single row of about 21 small, angular denti-
cles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Duxbury Reef, Marin
Co., California (Long, 1969a), to Shell Beach,
San Luis Obispo Co., California (Long,
1969a). Found in rocky intertidal, usually on
hydroids at base of the algae Laminaria spp.
Cuthona lagunae (O'Donoghue, 1926)
Hervia sp. ? Guernsey, 1912: 78, fig. 391.
Steinberg, 1961: 62. Steinberg, 1963a: 66.
Hervia lagunae O'Donoghue, 1926: 232.
Steinberg, 1961: 62. Steinberg, 1963a: 66.
Catriona ronga Marcus, 1961: 52, pl. 10, figs.
185-187. Steinberg, 1963a: 66.
Catriona lagunae (O'Donoghue). Steinberg,
1963a: 66. Steinberg, 1963b: 72. Farmer,
1967: 342. Sphon & Lance, 1968: 77.
Cratena rutila MacFarland, 1966: 332-336,
pl. 60, fig. 1; pl. 67, figs. 1-6a; pl. 69; figs.
1-1a; pl. 71, fig. 21. Sphon & Lance, 1968:
77. Roller, 1969b: 421, text fig. 1. Roller,
1970a: 372.
Trinchesia lagunae (O'Donoghue). Roller,
1969b: 421. Roller & Long, 1969: 428.
Roller, 1970a: 372. Gosliner & Williams,
1970: 179. Bertsch et al., 1972: 308. Ny-
bakken, 1974: 371. Abbott, 1974: 376, fig.
4445.
Cuthona lagunae (O'Donoghue). Williams 8
Gosliner, 1979: 208, 215.
Type-specimens: not listed.
Sides of body distinctly separated from foot
by a longitudinal groove. Oral tentacles
translucent grayish-white with encrusting
opaque white on distal two thirds. Rhino-
phores smooth, orange-red with extreme tips
slightly lighter. Cerata cylindrical, slightly cla-
vate, and somewhat decumbent; arranged in
8-9 transverse rows dorso-laterally on dor-
sum. General ground color translucent gray-
ish-white to cream. Top of head bears a large,
rhomboidal, orange-red blotch. An irregular,
opaque white line occurs dorso-medially on
tail. Cerata bear a few opaque white dots,
tips orange-red; cores vary from ochre to al-
most black. T.L.: 8 mm.
Anus on right, just anterior of innermost
ceras of first post-cardiac group. Genital ap-
erture on right side, just ventral of third row
of cerata. Penis armed with chininous stylet.
Radular formula of specimen examined
71(0.0.1.0.0), MacFarland (1966: 333) re-
ports 81(0.0.1.0.0). Rachidian teeth (Fig. 121)
bear 5-6 denticles on either side of median
cusp. Masticatory border of mandibles bears
a single series of rather coarse denticles.
Type-locality: Laguna Beach, Orange Co.,
California.
Range and habitat: Palomarin, Marin Co.,
California (Gosliner & Williams, 1970), to Ro-
sarito Beach, Baja California, Mexico (Farmer,
1967). Intertidal to 8 m, in rocky areas, fre-
quently in association with hydroids at base
of the algae Laminaria spp.
Cuthona virens (MacFarland, 1966)
Cratena virens MacFarland, 1966: 344-347,
pl. 61, fig. 5; pl. 68, figs. 8-11; pl. 70, figs.
6-8. Roller, 1970a: 372.
Trinchesia virens (MacFarland). Roller, 1969b:
421. Long, 1969a: 9, fig. 1. Roller, 1970a:
372. Roller, 1970b: 483. Abbott, 1974: 376,
fig. 4448. Gosliner 8 Williams, 1973b: 353,
354.
Cuthona virens (MacFarland). Williams 8
Gosliner, 1979: 208, 215.
Type-specimens: type material at Calif.
Acad. Sci.
Sides of body distinctly separated from foot
by a shallow, longitudinal groove. Oral ten-
tacles translucent grayish-white with minute
white dots on distal half. Rhinophores
smooth, translucent grayish-white, with pale
yellow on proximal half and distal half paler
and encrusted with minute, white dots. Cera-
ta cylindrical, somewhat spindle-shaped, ta-
pered to pointed tips and somewhat decum-
bent; arranged in about 7 transverse rows
dorso-laterally on dorsum. General ground
color translucent grayish-white, with a tinge
of yellow around and posterior of bases of
rhinophores. Anterior margin between oral
tentacles bordered with light yellow. Cerata
bear occasional flecks of orange or yellow;
cores grass green with orange-yellow tips.
TES mm.
Anus on right, just anterior of innermost
ceras of first post-cardiac group. Genital ap-
erture on right side, just ventral of second
row of cerata. Penis armed with chitinous
stylet.
Radular formula of specimen examined
CALIFORNIA NUDIBRANCHS 169
33(0.0.1.0.0), MacFarland (1966: 346) re-
ports 46(0.0.1.0.0). In examining Mac-
Farland's microscope slide of the radula of
the holotype, | found only 33 teeth; however,
some may have been lost in mounting since
MacFarland states that the radula had 46
teeth. Rachidian teeth (Fig. 122) bear 4-6
denticles on either side of median cusp. Mas-
ticatory border of mandibles bears a few tri-
angular denticles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Duxbury Reef, Marin
Co., California (Gosliner & Williams, 1970), to
Shell Beach, San Luis Obispo Co., California
(Long, 1969a). Found in rocky intertidal areas.
Remarks: This is the rarest member of the
genus in California. Only two or three speci-
mens have been recorded thus far.
Cuthona sp.
Oral tentacles translucent grayish-white
with small, opaque white flecks. Rhinophores
smooth, translucent grayish-white with nu-
merous small, opaque white flecks which are
more concentrated distally. Cerata cylindri-
cal, slightly inflated, and tapered to blunt tips;
arranged in 6-10 transverse rows dorso-lat-
erally on dorsum. General ground color
translucent grayish-white to cream. Dorsum
bears numerous minute, opaque white flecks.
Cerata bear numerous opaque white flecks
and a band of similar color well below tips,
which may be frosted with opaque white;
cores olivaceous-green to brownish. T.L.: 7
mm.
Anus on right, just posterior of cardiac re-
gion. Genital aperture on right side, just ven-
tral of second group of cerata.
Radular formula of specimen examined
28(0.0.1.0.0). Rachidian teeth (Fig. 123) bear
5-10 denticles on either side of median cusp.
Range and habitat: San Francisco Bay,
California (personal observation). Found on
floating docks, in association with the sea
anemone Haliplanella luciae upon which it
may feed.
Remarks: Additional study is needed to
determine if this species is a named species
which has been introduced into San Francis-
co Bay, or whether it is a species new to
science.
Dendrodoris Ehrenberg, 1831
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum may be
smooth or tuberculate, and is usually rather
translucent. Labial tentacles very rudimen-
tary, mouth pore-like. Rhinophores perfoliate
and retractile into sheaths. Branchial plumes
tri- to quadripinnate, retractile, arranged in a
circle around anus.
Radula absent.
Penis armed with spines.
Eliot (1906b: 663) states that in Dendro-
doris the buccal ganglia beneath the esoph-
agus are located at a constriction of the
esophagus, some distance posterior of the
main body of the central nervous system, and
are united to the nerve-collar by rather long
connectives, while in Doriopsilla Bergh, 1880,
the buccal ganglia beneath the esophagus lie
immediately posterior of the main body of the
central nervous system. This seems to be the
only recognizable difference between Den-
drodoris and Doriopsilla. The location of the
buccal ganglia seems a rather poor and in-
sufficient character for use in separating two
genera. Steinberg (1961: 58) states that “Until
it can be shown that the condition of the cen-
tral nervous system may be successfully used
in separating genera in this very difficult fam-
ily, I do not consider it wise to maintain Do-
riopsilla as generically distinct from Dendro-
doris.’’ Further, Thompson (1975: 500)
synonymizes Doriopsilla with Dendrodoris,
stating: ‘The distinction is based upon sev-
eral features of the morphology which ap-
pear to me to be inadequate.”
Type-species: Dendrodoris lugubris Ehren-
berg, 1831.
Dendrodoris albopunctata (Cooper, 1863)
Doris albopunctata Cooper, 1863b: 58. Car-
penter, 1864: 609. Cooper, 1867: 14. Coo-
per, 1870: 56. Abraham, 1877: 209. Or-
cutt, 1885: 545. Yates; 1890: 41.
O'Donoghue, 1922d: 143. Steinberg, 1961:
58.
Doriopsis reticulata Cockerell in Cockerell 8
Eliot, 1905: 41-42, pl. 7, fig. 5. Eliot, 1906a:
366. O'Donoghue, 1922d: 143.
Doriopsis fulva MacFarland, 1905: 45.
MacFarland, 1906: 130-131, pl. 19, figs.
38-40; pl. 22, fig. 3. Berry, 1907: 35.
Guernsey, 1912: 77, fig. 38B. O'Dono-
ghue, 1922d: 142-144. O'Donoghue, 1926:
212. O'Donoghue, 1927b: 92-93. Shelford
et al., 1935: 283. Norris 8 Rao, 1935: 787.
Costello, 1938: 330, pl. 1, fig. 3. Steinberg,
1961: 62. Fuhrman et al., 1979: 290.
170 MCDONALD
Doriopsilla reticulata (Cockerell & Eliot). Eliot,
1906b: 665.
Doridopsis fulva MacFarland. Eliot, 1907:
330, 349. Steinberg, 1961: 58.
Doris sp. Guernsey, 1912: 78, fig. 38C.
O'Donoghue, 1927b: 78. Steinberg, 1961:
62.
Doriopsis albopunctata (Cooper). O'Dono-
ghue, 1922d: 143.
Dendrodoris fulva (MacFarland). O'Dono-
ghue, 1922d: 142. O'Donoghue, 1926: 212.
O'Donoghue, 1927b: 92-93. Costello,
1938: 324, 327, 329, tabs. 1-3, 5. Worley
& Worley, 1943: 367, pl. 3, fig. 13. Stein-
berg, 1961: 57. Marcus, 1961: 30, 58, pl.
6, figs. 107-108. McLean, 1962: 111.
MacFarland, 1966: 194-196, pl. 28, fig. 2;
pl. 29, figs. 18-19. Roller, 1970a: 371.
Hertz, 1970 6. Michel, 1970: 7. McBeth,
1970: 28. McBeth, 1971: 158, 159. Mc-
Beth, 1972a: 55 ff. Harris, 1973: 264.
Poorman 8 Poorman, 1978: 373. Fuhrman
et al., 1979: 290.
Doriopsilla albopunctata (Cooper). O'Dono-
ghue, 1922d: 142-144. O'Donoghue, 1926:
205. O'Donoghue, 1927b: 93-95. Ingram,
1936: 48. Steinberg, 1961: 57. Marcus 8
Marcus, 1967a: 98-99, 189, 204-205, 238.
Roller, 1970a:372. McBeth, 1970: 28.
Gosliner £ Williams, 1970: 178. Keen, 1971:
830. Bertsch et al., 1972: 306. McBeth,
1972a: 55 ff. Abbott, 1974: 366, fig. 4351.
Haderlie et al., 1974: tab. 4. Bloom, 1976:
295. Nybakken, 1978: 134 ff. McDonald 8
Nybakken, 1978: 113. Haderlie & Donat,
1978: 60. Poorman 8 Poorman, 1978: 373.
Fuhrman et al., 1979: 290 ff. Russo, 1979:
44, 48. Bertsch, 1980: 224. McCosker,
1980: 30.
Doriopsilla fulva (MacFarland). Andrews,
1945: 26, 34. Pequegnat, 1963: 427. Mar-
cus & Marcus, 1967a: 204. Hargens, 1977:
363.
Dendrodoris (Doriopsis) fulva (MacFarland).
Smith & Gordon, 1948: 181.
Dendrodoris albopunctata (Cooper). Lance,
1961: 67. Farmer & Collier, 1963: 62.
Steinberg, 1963b: 71. Paine, 1963a: 4.
Paine, 1964: 385. Ghiselin, 1965: 345.
Lance, 1966: 69. MacFarland, 1966: 196-
197, pl. 28, fig. 4. Sphon & Lance, 1968:
78. Turner et al., 1969: 133. Roller, 1970a:
371. North, 1971: 57. Fuhrman et al., 1979:
290.
Dendrodoris albopunctatus (Cooper). Farmer,
1964: 24. Paine, 1965: 607 (lapsus).
Dendrodoris (Doriopsilla) albopunctata (Coo-
per). Ghiselin, 1964: 45-46.
Dendrodoris (Doriopsila) albopunctata (Coo-
per). Ghiselin, 1965: 335 (lapsus).
Doriopsilla ? albopunctata (Cooper). Roller &
Long, 1969: 427.
Type-specimens: not listed; state coll.
species 1000 (Cooper, 1863b).
Body rather elongate. Foot elongate and
elliptical, anterior margin bilabiate. Rhino-
phores bear 11-20 lamellae, and are pale
yellow to yellow-orange on shaft, clavus rath-
er brownish-yellow. Rhinophore sheaths have
low, smooth, thin margins. Branchial plumes
5, tripinnate, whitish to very pale yellow.
General ground color varies from pale yellow
to yellow-orange or chestnut brown, low tu-
bercles on dorsum each bear a small, apical,
white dot. Some specimens are brownish
medially on dorsum. T.L.: 25 mm.
Type-locality: Santa Barbara, Santa Bar-
bara Co., California.
Range and habitat: Van Damme, Mendo-
cino Co., California (Gosliner & Williams,
1970), to Punta San Eugenio, Baja Califor-
nia, Mexico (Lance, 1961). Intertidal to 46 m.
One of the most common dorids found in the
rocky intertidal zone in central California.
McBeth (1971: 158) reports that it feeds upon
the sponges: Acarnus erithacus, Cliona ce-
lata, Ficulina suberea, and Suberites sp.
Remarks: Roller (1970a: 371) and Stein-
berg (1961: 58) synonymized Dendrodoris
fulva with Dendrodoris albopunctata. This
may well be warranted as it is very difficult to
determine from Cooper's brief description
what D. albopunctata really is. There are,
however, other species of yellow porostomes
which occur along the California coast which
are very much like D. albopunctata as de-
scribed by Cooper, but yet are not like D.
fulva as described by MacFarland, and are
different from the animals which are currently
accepted as D. albopunctata (vide Dendro-
doris sp. a). In addition, there are occasion-
ally found specimens of a eudoridacean which
externally resembles D. albopunctata. It may
be distinguished from D. albopunctata by the
8-10 bipinnate, rather vertical branchial
plumes, the digitiform labial tentacles, and the
rather narrow foot; it is firmer than D. albo-
punctata, and is usually less than 20 mm in
length.
Dendrodoris nigromaculata (Cockerell in
Cockerell & Eliot, 1905)
Doridopsis vidua (?), Bergh. Cockerell & Eliot,
1905: 40-41. O'Donoghue, 1922d: 143.
CALIFORNIA NUDIBRANCHS NA
Doris nigromaculata Cockerell in Cockerell 8
Eliot, 1905: 40-41.
Doridopsis nigromaculata Cockerell 8 Eliot
(vidua Bergh, var. (?)). Cockerell, 1908: 106.
O'Donoghue, 1922d: 143.
Doriopsis nigromaculata Cockerell & Eliot.
O'Donoghue, 1922d: 143.
Dendrodoris vidua (Bergh). O'Donoghue,
1926: 212.
Doriopsis vidua Bergh. Kelsey, 1907: 39.
O'Donoghue, 1926: 212-213.
Doridopsis nigromaculata Cockerell. O'Don-
oghue, 1926: 213.
Dendrodoris nigromaculata (Cockerell).
Steinberg, 1961: 59.
Doriopsilla nigromaculata (Cockerell 8 Eliot).
Abbott, 1974: 366. Bertsch, 1977: 112.
Type-specimens: not listed.
Dorsum smooth. Foot bluntly rounded an-
teriorly and tapered posteriorly to blunt tail.
Rhinophores bear about 5-6 inclined lamel-
lae and are grayish-white. Branchial plumes
5-6, bi- and tripinnate, whitish. General
ground color translucent grayish-white, dor-
sum bears numerous minute, brownish flecks
which are more concentrated medially. About
4-5 opaque white blotches occur in a line
dorso-laterally on either side of dorsum, be-
tween rhinophores and branchial plumes.
12 s0 mm:
Type-locality: La Jolla, San Diego Co., Cal-
ifornia.
Range and habitat: La Jolla, San Diego Co.,
California (Cockerell 8 Eliot, 1905). This
species occurs in rocky intertidal, and is quite
rare.
Remarks: Doriopsilla rowena Marcus &
Marcus, 1967a is quite probably a synonym
of D. nigromaculata. However, since no
specimens of D. rowena or D. nigromaculata
from the known geographic range of the for-
mer were available for study, | hesitate to
synonymize the two.
Dendrodoris sp. a
Body quite broad and ovate, foot elliptical.
Rhinophores bear 13-18 inclined lamellae and
are orange to brownish-orange. Branchial
plumes 5, tripinnate, orangish. General ground
color varies from orange to brownish-orange,
dorsum somewhat darker medially. Dorsum
bears numerous small, opaque white dots
which occur over all the dorsum and are not
confined to the apex of each tubercle as in
Dendrodoris albopunctata. T.L.: 40 mm.
Range and habitat: Elkhorn Slough, Mon-
terey Co., California (personal observation),
to southern California. Vast majority of spec-
imens are subtidal, usually in rocky areas, but
also found in sloughs.
Remarks: The nidosome of this species is
very different from that of Dendrodoris albo-
punctata. That of D. albopunctata is typical
of dorids; it is a yellow spiral which is at-
tached to the substrate by the narrow edge
of the ribbon. The nidosome of Dendrodoris
sp. a is also a spiral, but is usually more yel-
low-orange than that of D. albopunctata, and
it is attached to the substrate by the broad
surface of the ribbon.
Dendrodoris sp. b
Dendrodoris sp. Lee 8 Brophy, 1969: 220.
Body quite elongate with nearly parallel
sides, and equally rounded anteriorly and
posteriorly; dorsum nearly smooth. Foot quite
elongate and rounded anteriorly and poste-
riorly. Rhinophores bear 8-10 lamellae and
are white to cream. Rhinophore sheaths have
low, smooth margins. Branchial plumes 5, tri-
pinnate, white to cream. General ground col-
or white to very pale cream. Dorsum bears a
number of irregular, chocolate-brown blotch-
es of various sizes, usually with the larger
blotches concentrated in three groups, one
just anterior of branchial plumes, one mid-
dorsally, and one just posterior of rhino-
phores; occasionally another concentration
may occur anterior of rhinophores near an-
terior margin of dorsum. Sides of body, be-
tween mantle margin and foot, bear an irreg-
ular, longitudinal row of about 4-8 small,
chocolate-brown dots. T.L.: 20 mm.
Range and habitat: Pescadero Point, Car-
mel Bay, Monterey Co., California (personal
observation), to Point Loma, San Diego Co.,
California (personal observation). Intertidal to
46 m, usually in rocky areas.
Dendronotus Alder & Hancock, 1845
Body limaciform. Cerata arborescent, ar-
ranged in a longitudinal row along either edge
of dorsum, usually in 3-8 opposite pairs; cni-
dosacs absent. Oral tentacles as such ab-
sent. Anterior margin of body bears a veil
which usually has 2-5 pairs of velar process-
es which are normally branched. Rhino-
phores perfoliate, clavus contained within a
campanulate sheath which bears a number
of more or less branched processes.
Labial disc armed with small rodlets or fil-
aments. Masticatory border of mandibles
normally denticulate. Rachidian teeth usually
172 MCDONALD
denticulate but may be smooth; laterals nar-
row and usually denticulate.
Penis unarmed. Anus located between first
and second cerata on right side.
Type-species: Dendronotus frondosus
(Ascanius, 1774).
Dendronotus albus MacFarland, 1966
Dendronotus albus MacFarland, 1966: 256,
272, 274, 275-279, pl. 40, fig. 1; pl. 46,
figs. 1-4; pl. 47, figs. 8-11; pl. 48, figs. 7—
8; pl. 49, fig. 5; pl. 50, fig. 4; pl. 51, figs.
6-7. Sphon 8 Lance, 1968; 78. Long,
1969c: 232. Roller & Long, 1969: 426.
Robilliard, 1970: 466-470, pl. 64, fig. 34,
text figs. 2, 4-6, 22-24. Schmekel, 1970:
180. Bertsch et al., 1972: 305. Robilliard,
1972: 421 ff. Nybakken, 1974: 371. Had-
erlie et al., 1974: tab. 4. Abbott, 1974: 368.
Lambert, 1976: 296, 297. Thompson,
1976b: 92. Nybakken, 1978: 135. Мс-
Donald & Nybakken, 1978: 113. Robilliard
8 Barr, 1978: 153.
Type-specimens: type material at Calif.
Acad. Sci.
Body rather high, and tapered to pointed
tail. Foot narrow, rounded anteriorly and ta-
pered posteriorly to pointed tail. Frontal veil
bears 4 long, branched, tapered processes
on a low, horseshoe-shaped ridge, 2 pro-
cesses on either side of median line; medial
pair longer; below this series may occur a
variable number of small, less branched ap-
pendages. Rhinophores bear 12-14 lamellae.
A short, simple process is directed obliquely
upward midway on cylindrical rhinophore
shaft. Margin of rhinophore sheath bears 5
slender, tapered, branched processes, the
longest of these processes is posterior. Cla-
vus and distal half of the processes of rhi-
nophore sheath usually orange-red to brown-
ish, rarely opaque white; rhinophore shaft
translucent grayish-white. The 4-8 opposite
pairs of cerata have rather long branches,
anterior pairs more branched than posterior
pairs. First pair of cerata occurs immediately
anterior of cardiac region, second pair occurs
immediately posterior of cardiac region. He-
patic diverticulae in 3-5 pairs of cerata. Gen-
eral ground color tanslucent grayish-white. A
narrow, median, opaque white line extends
posteriorly from between fourth cerata to tip
of tail. Velar processes and cerata branches
opaque white, usually tipped with orange-red
to brownish. T.L.: 25 mm.
Anus on right, about midway between first
and second cerata. Genital aperture on right
side, ventral and anterior of first ceras.
Radular formula of specimen examined
37(7-8.1.7-8), MacFarland (1966: 278) re-
ports 36-38(7-9.1.7-9), and Robilliard (1970:
468) reports 32-38(6-8.1.6-8). Rachidian
teeth (Fig. 78a) bear 16-20 small, blunt den-
ticulations on either side. First laterals (Fig.
78b) have 5-11 denticles on external margin.
Outermost 2-3 laterals (Fig. 78h, i) bear only
a single cusp and no denticles. Masticatory
border of mandibles bears 70-80 transverse,
plate-like ridges.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Port Dick, Kenai Pen-
insula, Alaska (Robilliard 8 Barr, 1978), to Is-
las Coronados, Baja California, Mexico (Ro-
billiard, 1970). Intertidal to 30 m, usually found
in rocky intertidal and subtidal. Feeds upon
the hydroid Thuiaria argentea Robilliard,
1970: 470), and also found on Abietinaria
amphora and Plumularia sp. (Robilliard 8
Barr, 1978; McDonald 8 Nybakken, 1978).
This species is capable of swimming.
Remarks: Dendronotus albus is frequently
quite difficult to distinguish from Dendrono-
tus diversicolor and may possibly be conspe-
cific with it, but present data are insufficient
to determine this.
Dendronotus diversicolor Robilliard, 1970
Dendronotus diversicolor Robilliard, 1970:
470-475, pl. 64, figs. 35-36, text figs. 4-
6, 25-28. Gosliner & Williams, 1970: 177.
Robilliard, 1972: 421 ff. Robilliard, 1974a:
335-336. Abbott, 1974: 368. Lambert,
1976: 296, 298. Thompson, 1976b: 92.
Michel, 1976: 47, fig. 7. McDonald & Ny-
bakken, 1978: 113.
Type-specimens: Calif. Acad. Sci., no. 416.
Body quite compressed laterally. Foot nar-
row, rounded anteriorly and tapered poste-
riorly to sharply pointed tail. Frontal veil bears
6 slender, slightly branched processes, me-
dial pair is longest. Rhinophores bear 16-21
lamellae. A small, simple lateral process oc-
curs between one third and one half way up
rhinophore shaft, it is usually about one quar-
ter the length of rhinophore shaft, but may
be shorter or even absent. Margin of rhino-
phore sheath bears 5 tall, slender, simply
branched processes, longest of which is pos-
terior. Distal one third of posterior rhinophore
sheath processes and posterior surface of
rhinophore sheath may be orange or opaque
CALIFORNIA NUDIBRANCHS 173
white; rhinophore shaft translucent grayish-
white to lilac. The 4-5 opposite pairs of cera-
ta are tall, slender, and sparsely branched.
First pair of cerata occurs immediately ante-
rior of cardiac region, second pair occurs im-
mediately posterior of cardiac region. First 3
pairs about equal in height, fourth pair about
one half to two thirds as high, and fifth pair,
if present, consists of short papillae. Hepatic
diverticulae in anterior 2 pairs of cerata. Gen-
eral ground color translucent grayish-white or
lilac. A narrow, median, opaque white line ex-
tends posteriorly from between last pair of
cerata to tip of tail. Velar processes and ce-
rata branches tipped with orange or opaque
white. T.L.: 40 mm.
Anus on right, midway between first and
second cerata. Genital aperture on right side,
ventral and anterior of first ceras.
Radular formula of specimen examined
38(6-7.1.6-7), Robilliard (1970: 472) reports
33-38(6-9.1.6-9). Rachidian teeth (Fig. 79a)
bear 13-25 small, blunt denticulations on
either side. First four laterals (Fig. 79b) have
4-11 denticles on external margin. Fifth lat-
eral (Fig. 79f) rarely bears any denticles be-
low the single distal cusp. Outermost 1-3 lat-
erals (Fig. 79g-i) have only a single cusp and
no denticles. Masticatory border of mandi-
bles bears 40-61 transverse ridges.
Type-locality: San Juan Island, Puget
Sound, Washington.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976), to Point
Loma, San Diego Co., California (Robilliard,
1974a). Intertidal to 19 m, in rocky areas.
Found on various hydroids: Abietinaria spp.,
Hydrallmania distans, and Sertularella tricus-
pidata (Robilliard, 1970: 474). This species is
capable of swimming.
Remarks: The white phase of this species
is very difficult to differentiate from Dendro-
notus albus.
Dendronotus frondosus (Ascanius, 1774)
Amphitrite frondosa Ascanius, 1774: 155, pl.
Sy ile ler
Doris arborescens Miller, 1776: 229. Fabri-
cius, 1780: 346. Mohr, 1786: 116. Cuvier,
1804: 449. Thienemann, 1824: 156. Bosc,
1830: 112. Odhner, 1936: 1109.
Doris cervina Gmelin in Linnaeus, 1791: 3105,
MO E
Doris frondosa. Cuvier, 1804: 449. Bosc,
1830: 112.
Tritonia arborescens Cuvier, 1805: 435, pl.
61, figs. 8-10. Cuvier, 1817: 28, figs. 8-
10. Lamarck, 1819: 304. Fleming, 1823b:
254. Grant, 1826: 165, 185-186. Fleming,
1828: 284. Stark, 1828: 68. Cuvier, 1834:
119. Lamarck, 1836: 454. Johnston,
1838a: 46. Johnston, 1838b: 115. Gould,
1841: 5. Thompson, 1844: 276. Alder &
Hancock, 1846b: 65. Kroyer, 1847: 116.
Alder 8 Hancock, 1855: 4.
Tritonia cervina. Bosc, 1830: 106.
Tritonia reynoldsii Couthouy, 1838: 74, pl. 2,
figs. 1-4.
Tritonia lactea Thompson, 1840: 88, pl. 2, fig.
3. Thompson, 1856: 275, 276, 485.
Tritonia pulchella Alder & Hancock, 1842: 33.
Alder, 1850: 114.
Tritonia felina Alder & Hancock, 1842: 33. Al-
der, 1850: 114.
Tritonia ascanii Möller, 1842: 78.
Tritonia reynoldsi Couthouy. DeKay, 1843: 8,
pl. 5, fig. 94.
Dendronotus arborescens (Müller). Alder &
Hancock, 1845a: 644. Alder & Hancock,
1845b: fam. 3, pl. 3. Reid, 1846: 377 ff.
Alder & Hancock, 1846a: fam. 3, pl. 2. Lo-
ven, 1846: 138-139. Alder, 1850: 105, 113.
Stimpson, 1853: 26. Adams & Adams,
1854: 65, pl. 64, fig. 7. Byerley, 1854: 45.
Alder & Hancock, 1855: 21, 31, 32, 47.
Gosse, 1856: 103. Thompson, 1856: 485.
Carpenter, 1857: 313. Gray, 1857: 219.
Chenu, 1859: 407, fig. 3059. Collingwood,
1859: 464-465. Hyndman, 1859: 286. E.
Wright, 1859: 88. Collingwood, 1860: 202.
Norman, 1860: 7243. Collingwood, 1861:
114. Collingwood & Byerley, 1962: 189.
Mcintosh, 1865: 391. Meyer & Möbius,
1865: 43-47, pl. 5, pl. 3. Robertson, 1868:
206. Mörch, 1868: 204. Hogg, 1868: pl. 10,
fig. 38. Jeffreys, 1869: 62-63, pl. 2, fig. 2.
Gould, 1870: 234-236, pl. 22, figs. 311-
313. Dall, 1870: 250. Sauvage, 1873: 25,
33-34. Verrill, 1873: 495, 499, 665. Verrill,
1874a: 43. Mcintosh, 1874: 430. Mc-
Intosh, 1875: 86. Friele 8 Hansen, 1876:
73. Smith & Harger, 1876: 8, 13. Mörch,
1877: 436. Garner, 1878: 93. Jones, 1878:
326. Sars, 1878: 314-315 ff, pl. XV, fig. 3.
Bergh, 1879b: pl. 2, figs. 13-15; pl. 3, fig.
1; pl. 4, figs. 1-4. Bergh, 1879c: pl. 2, figs.
13-15; pl. 3, fig. 1; pl. 4, figs. 1-4. Verrill,
1880: 385, 386. Leslie 8 Herdman, 1881,
310. Verrill, 1882a: 339. Bush, 1883: 245.
Hertzenstein, 1885: 710. Krause, 1885:
295. Becher, 1886: 14. Locard, 1886: 40-
41. Higgins, 1886: 26. Haddon, 1886: 530.
Herdman, 1886: 271, 277. Holm, 1887: 156
174 MCDONALD
ff. Fischer, 1887: 535, text fig. 292. Bergh,
1887: 25-35, pl. 2, figs. 12-28. Herdman
& Clubb, 1889: 228-231, pl. 12, figs. 1-3,
7. Garstang, 1889: 185. Herdman, 1890a:
46, 54, pl. 7, figs. 16-21. Herdman, 1890b:
202. Bergh, 1890a: 56. Garstang, 1890:
425. Bergh, 1892: 1051 (59). Herdman 4
Clubb, 1892: 136 ff, pl. 6, fig. 14. Lund-
beck, 1893: 175. Herdman, 1894: 14.
Bergh, 1894: 137-139. Clubb, 1895: 220-
234, pls. 14-15. Herdman et al., 1896: 446.
Sumner, 1896: 49. Vanhoffen, 1897: 188,
193. Posselt, 1898: 248. Cooke, 1899: 61-
62. Liverpool Mar. Biol. Comm., 1899: 55.
Beaumont, 1900: 847. Whiteaves, 1901:
206. Vayssiere, 1901: 296. Johansen,
1902: 387. Todd, 1903: 544, 556. Cock-
erell & Eliot, 1905: 32. Walton, 1908: 237-
238. Colgan, 1903: 106, 112. Farran, 1909:
14. Eliot, 1910: 13. Pelseneer, 1911: 63-
64, pl. 18, figs. 2-13. Sumner et al., 1913:
704. Chumley, 1918: 103, 149, 169. Bar-
darson, 1919: 72. O'Donoghue, 1921: 184,
pl. 4 (10), fig. 45; pl. 5 (11), figs. 51-53.
O'Donoghue, 1922a: 124. Scott, 1922: 49.
White, 1938: 14. Volodchenko, 1955: 248,
pl. 48, fig. 1. MacFarland, 1966: 254 ff.
Storch 4 Welsch, 1969: 528 ff. Daro, 1969:
141, 147. Zaitseva, 1978: 498 ff.
Amphitridea facrici. Kroyer, 1847: 114.
Dendronotus reynoldsii (Couthouy). Alder 8
Hancock, 1855: 28. Mörch, 1857: 78.
Stimpson, 1862: 4. Mórch, 1875: 125.
Mörch, 1877: 436.
Tritonia (Dendronotus) arborescens. Carpen-
ter, 1857: 218.
Dendronotus pulchella (Alder & Hancock).
Gray, 1857: 219.
Dendronotus felina (Alder & Hancock). Gray,
1857: 219.
Campaspe pusilla Bergh, 1863: 471-478, pl.
12, figs. 28-35. Mörch, 1857: 125. Mörch,
1877: 437. Bergh, 1892: 1050 (58). Vanof-
fen, 1897: 193. Posselt, 1898: 251. Odhner,
1907: 64. Loyning, 1927: 249. Odhner,
1936: 1108.
Dendronotus luteolus Lafont, 1871-1872:
207. ple 17, 1192 1. Fischer, 1872: 14. Lo=
card, 1886: 41. Bergh, 1892: 1051 (59).
Odhner, 1936: 1108.
Dendronotus purpureus Bergh, 1879b: 89-
94, pl. 1, figs. 18-20; pl. 3, figs. 7-12.
Bergh, 1879c: 145-150, pl. 1, figs. 18-20;
pl. 3, figs. 7-12. Bergh, 1892: 1051 (59).
Bergh, 1903: 15-18, pl. 2, figs. 10-12.
Odhner, 1936: 1108. LaRocque, 1953: 253.
MacFarland, 1966: 257.
Dendronotus arborescens var. aurantiaca
Friele, 1879: 284.
Dendronotus elegans Verrill, 1880: 385-386.
Verrill, 1882b: 551.
Dendronotus pulchellus (Alder & Hancock).
Locard, 1886: 41.
Dendronotus lacteus (Thompson). Becher,
1886: 14. Bergh, 1892: 1051 (59). Eliot,
1910: 112, 151. Odhner, 1936: 1108.
White, 1938: 18.
Campaspe major Bergh, 1887: 21-24, pl. 1,
figs. 23-26; pl. 2, figs. 1-11. Bergh, 1892:
1051 (58). Odhner, 1936: 1108. Loyning,
1927: 249.
Dendronotus frondosus (Ascanius). Norman,
1890: 78. Tregelles, 1896. 221. Cooke,
1899: 66. Herdman et al., 1900: 46. Nich-
ols, 1900: 592. Conchol. Soc., 1901: 25.
Knight, 1901: 207. Farran, 1904: 5. Mar.
Biol. Assoc., 1904: 282. Odhner, 1907: 66.
Balch, 1909: 36. Eliot, 1910: 161. Craws-
hay, 1912: 372. Vayssiere, 1913: 7-10, pl.
figs. 7-18. Johnson, 1915: 169. Evans 8
Evans, 1917: 109. Bardarson, 1920: 109.
Elmhirst, 1922: 41. Iredale & O'Donoghue,
1923: 216. Larsen, 1925: 38-42, text figs.
30-33, pl. figs. 9a-d. Odhner, 1926b: 17-
19, text fig. 13. O'Donoghue, 1926: 223,
Cuenot, 1927: 266. Jutting, 1927:
LXXXVIII. Loyning, 1927: 246-249, 262.
Derjugin, 1928: 319-320. Lemche, 1929:
8-9. Lonnberg, 1931: 20. Mar. Biol. As-
soc., 1931: 272. Winckworth, 1932: 235-
236. Chambers, 1934: 636. Odhner, 1936:
1063 ff, text figs. 3-4, 39. Moore, 1937:
191-192. White, 1938: 14, 18. Lemche,
1938: 13-14 ff. Graham, 1938: 300. Odh-
ner, 1939: 45-46. Lemche, 1941a: 23-24.
Lemche, 1941b: 15-18. Jutting, 1947: 64.
Fischer, 1950: 199. Jaeckel, 1952: 24 ff.
La Rocque, 1953: 252. Williams, 1954: 106.
MacGinitie, 1955: 66, 97, 101, 102, 175.
Franzén, 1955: 428, text fig. 96. Graham,
1955: 153. Baba, 1957: 9. Mar. Biol. As-
soc., 1957: 313. Swennen, 1959: 58. Mac-
Ginitie, 1959: 144-145, pl. 3, fig. 1.
Thompson, 1960a: 24-26, text figs. 1-2.
Thompson, 1960b: 126-127. Buznikov,
1960: 373. Buznikov 8 Manukhim, 1960:
1414-1416. Miller, 1961: 100, 105.
Thompson, 1961: 236. Swennen, 1961:
207-209. Marcus, 1961: 34-36 ff, pl. 7,
figs. 121-124. Lance, 1961: 67. Buznikov
8 Manukhim, 1961: 226, 232. Miller, 1962:
562. Sakharov, 1962: 310, 311. Roginska-
ya, 1962a: 88, 93-95, 106, figs. 2.1-2.3.
Roginskaya, 1962b: 205-206, 211-212 ff,
CALIFORNIA NUDIBRANCHS 175
fig. 1.5. Paine, 1963a: 4. Steinberg, 1963b:
71. Arronet, 1963: 11. Zenkevitch, 1963:
112. Bruce et al., 1963: 205. Thompson,
1964: 281 ff. Buznikov, 1964: 1243. Abe,
1964: 57, 87, pl. 29, fig. 100. Ghiselin,
1965: 351 ff. MacFarland, 1966: 254 ff.
Brattegard, 1966: 20. Sakharov, 1966: 957.
Hurst, 1967: 255 ff, text fig. 15, pl. 30, fig.
18; fig. 25-5. Marcus & Marcus, 1967a:
212. Thompson, 1967: 12. Carefoot, 1967:
628 ff. Buchsbaum & Milne, 1967: pl. 63.
Sphon & Lance, 1968: 91. Bebbington &
Thompson, 1968: 9. Marcus & Marcus,
1969: 27. Turner et al., 1969: 135, append.
1, 2. Loveland et al., 1969: 418. Haderlie,
1969: tab. 1. Long, 1969c: 232. Roller &
Long, 1969: 426. Morse, 1969: 38. Roller,
1970a: 372. Robilliard, 1970: 441-446, text
figs. 7-9, pl. 63, fig. 29. Gosliner & Wil-
ams51970; 177. Franz, 1970: 172 ff.
Hughes, 1970b: 81, 82. Bernard, 1970: 85.
Buznikov et al., 1970: 550, 557. North,
1971: 57. Meyer, 1971: 142. L. Williams,
1971: 166-171, text figs. 1-5. Keen, 1971:
832. Bertsch et al., 1972: 305. Holleman,
1972a: 60. Sphon, 1972a: 155. Gas-
coigne, 1972: 259. Robilliard, 1972: 421 ff.
Salvini-Plawen, 1972: 393. Voogt, 1973:
479 ff. Rasmussen, 1973: 27, 268, 460.
Platts, 1973: 384. Harris, 1973: 217, 292-
293. Thompson 8 Bebbington, 1973: 148,
149, pl. 9, figs. c-d. Robilliard, 1974a: 335.
Rozsa, 1974: 8. Nybakken, 1974: 371. Ab-
bott, 1974: 367, pl. 16, fig. 4357. Rogin-
skaya, 1974b: 998, text fig. 1. Harris et al.,
1975: 264. Robilliard, 1975a: 59. Robil-
liard, 1975b: 44-47. Sneli 8 Steinnes,
1975: 13. Clark, 1975: 40-41. Franz,
1975a: 81. Rozsa, 1975: 3. Belcik, 1975:
276. Michel, 1976: 49, fig. 14. Lambert,
1976: 296. Thompson 8 Brown, 1976: 66,
fig. 29. Thompson, 1976a: 39. Thompson,
1976b: 8, 34, 72, 92. Thiriot-Quiévreux,
1977: 178. Edmunds, 1977: 306, 307. Gar-
lo, 1977: 23 ff. McDonald 8 Nybakken,
1978: 113. Chamberlain 8 Behrens, 1980:
283. (Non) Odhner, 1936: 1105-1109
(=Dendronotus dalli Bergh, 1879, Dendro-
notus rufus O'Donoghue, 1921).
Dendronotus frondosus elegans Verrill. John-
son, 1915: 170. Johnson, 1934: 156.
Dendronotus frondosa. Dahlgren, 1925: 436
(lapsus).
Dendronotus frondosus frondosus. Johnson,
1934: 156.
Dendronotus purpurascens Bergh. Odhner,
1936: 1109 (/apsus).
Dendronotus frondosus var. purpureus
Bergh. MacFarland, 1966: 255.
Dendronotus venustus MacFarland, 1966:
271-275, pl. 40, fig. 2; pl. 46, figs. 9-12;
pl. 47, figs. 1-2; pl. 49, fig. 6; pl. 50, fig. 3;
pl. 52, figs. 3-6. Roller, 1970a: 372.
Schmekel, 1970: 180.
(Non) Dendronotus frondosus var. dalli. Zen-
kevitch, 1963: 134 (=Dendronotus аа!
Bergh, 1879).
Type-specimens: not listed.
Foot narrow, rounded anteriorly and ta-
pered posteriorly to short, pointed tail. Fron-
tal veil bears 4-8 branched, tapered process-
es; of these, two on either side of median line
are larger and relatively constant; between
these may occur smaller, less branched pro-
cesses. Rhinophores bear 6-14 lamellae. A
short process with short branches distally
occurs near base of rhinophore shaft on ex-
ternal surface. Rhinophore sheath bears 4-5
branched processes, postero-medial process
longest and most branched. Rhinophores
translucent grayish-green, with irregular
patches of yellow-green to brown-green, cla-
vus slightly darker shade of grayish-green.
The 3-9 opposite pairs of cerata bear rather
short branches, anterior pairs slightly more
branched than posterior pairs. First pair of
cerata occurs immediately anterior of cardiac
region, second pair occurs immediately pos-
terior of cardiac region. General ground color
translucent grayish-white, tending toward
greenish or brownish in most specimens. Ir-
regular patches of yellow-green to brown-
green occur on body, with smaller spots of
olive to pale green and pale yellow to bright
orange. Low conical papillae on body are
tipped with yellow. On occasional specimens,
a series of opaque white blotches occurs me-
dially on dorsum between successive pairs of
cerata. Distal ends of cerata branches flecked
with numerous small, yellow dots, cores light
green to brown. T.L.: 25 mm, may attain 100
mm.
Anus on right, dorso-laterally, between first
and second cerata. Genital aperture on right
side, ventral and anterior of first ceras.
Radular formula of specimen examined
32(8-9.1.8-9), MacFarland (1966: 272) re-
ports 28-34(6-7.1.6-7), and Robilliard (1970:
443) reports 33-48(7-11.1.7-11). Rachidian
teeth (Fig. 80a) bear 10-13 small denticles
on either side. Laterals (Fig. 80b-h) have 3-
6 denticles on external margin, innermost
denticles usually longest. Masticatory border
176 MCDONALD
of mandibles bears 27-40 transverse, ridge-
like denticles.
Type locality: Norway?
Range and habitat: Nearly cosmopolitan in
northern hemisphere (Lance, 1961); east
(Franz, 1975a) and west (Robilliard, 1970)
coasts of North America, British Isles (Mar.
Biol. Assoc., 1957), Europe (Cuenot, 1927),
China (Thompson & Brown, 1976), Japan
(Baba, 1957). Intertidal to 400 m. Occasion-
ally found in rocky intertidal; more commonly
found on floating docks and pilings in bays.
Feeds upon the hydroids: Bougainvillia glo-
rietta, Coryne sp., Dynamena pumila, Hy-
dractinia echinata, Hydrallmania falcata, Obe-
lia commissuralis, Obelia spp., Sertularia
argentea, Sertularia cupressina, Sertularia
dichotoma, Sertularia pumila, Syncoryne ex-
imia, Tubularia crocea, Tubularia indivisa, and
Tubularia larynx (Carefoot, 1967; Clark, 1975;
Collingwood, 1859; Grant, 1826; Harris,
1973; Harris et al., 1975; Herdman, 1886;
Kozloff, 1973; Meyer, 1971; Miller, 1961;
Robilliard, 1970; Swennen, 1961; Thomp-
son, 1964; Thompson 8 Brown, 1976). Also
found on the hydroids: Abietinaria abietina,
Aglaophenia sp., Laomedea geniculata, and
Obelia flabellata (Garlo, 1977; Herdman,
1890a; Marcus, 1961; Rasmussen, 1944,
1973). Morse (1969: 38) reports that it has
been observed eating the ascidian Botryllus
schlosseri. This species is capable of swim-
ming. Volodchenko (1955: 248) states that it
is preyed upon by fish.
Dendronotus iris Cooper, 1863
Dendronotus iris Cooper, 1863b: 59. Car-
penter, 1864: 609. Cooper, 1867: 14.
Yates, 1890: 41. Bergh, 1892: 1052(59).
O'Donoghue, 1926: 224. Odhner, 1936:
1105, 1107, 1109, pl, fig. 9; text figs:
40-41. Lance, 1961: 67. Marcus, 1961: 36.
Steinberg, 1963b: 71. MacFarland, 1966:
254-265 ff, pl. 47, figs. 12-18; pl. 48, figs.
1-6; pl. 49, fig. 4; pl. 50, fig. 1; pl. 51, figs.
1-5. Powell, 1966: 115. Marcus 8 Marcus,
1967a: 210, 212, 213. Hurst, 1967: 255 ff,
pl. 30, fig. 19; pl. 38, figs. 49-50; fig. 25-
2. Sphon & Lance, 1968: 78. Lee & Bro-
phy, 1969: 220. Roller & Long, 1969: 429.
Wobber, 1970: 383-387, pl. 55, figs. 1-2;
pl. 56, figs. 3-4; pl. 57, fig. 5. Roller, 1970b:
482. Robilliard, 1970: 446-450, pl. 63, fig.
30, text figs. 4-6, 10-12. Bernard, 1970:
85. Schmekel, 1970: 180, 181. Crane,
1971: 57. Robilliard, 1972: 421 ff. Gosliner
& Williams, 1973b: 352, 353. Harris, 1973:
221 ff. Thompson, 1973: 167 ff, text fig. 8.
Abbott, 1974: 367. Lambert, 1976: 296.
Thompson, 1976a: 28, pl. 3, figs. c, d.
Thompson, 1976b: 13, 58. McDonald &
Nybakken, 1978: 113.
Dendronotus giganteus O'Donoghue, 1921:
187-190, pl. 4 (10), fig. 47; pl. 5 (11) figs.
57-59. O'Donoghue, 1922b: 165. Agers-
borg, 1922a: 257. Agersborg, 1922b: 439.
O'Donoghue, 1924: 24. Agersborg, 1925:
167. O'Donoghue, 1926: 223. Fraser, 1932:
67. Odhner, 1936: 1105 ff. Hewatt, 1937:
178 ff. Smith & Gordon, 1948: 181. La-
Rocque, 1953: 253. Marcus, 1961: 36.
Kohn, 1961: 293 ff. MacFarland, 1966: 255,
258. Farmer, 1970: 79. Harris, 1973: 260.
Dendronotus iris (=giganteus). Thompson,
1976b: 92.
Type-specimens: not listed; state coll.
species 959 (Cooper, 1863b).
Foot relatively wide, rounded anteriorly and
tapered posteriorly to short, bluntly rounded
tail. Frontal veil bears 4-8 processes, which
are relatively sparsely branched. Rhino-
phores bear 15-31 lamellae. A stout,
branched process occurs near base of rhino-
phore shaft. Two to six (usually 4) small,
branched papillae are arranged vertically on
posterior face of rhinophore shaft. Rhino-
phore sheath bears 2-5 long, branched pro-
cesses, the longest of which is posterior. Cla-
vus translucent grayish white to orange or
maroon, usually tipped with opaque white.
The 4-7 opposite pairs of cerata bear nu-
merous long branches, anterior pairs larger
and somewhat more branched than posterior
pairs. First pair occurs immediately anterior
of cardiac region. Second pair occurs imme-
diately posterior of cardiac region. There are
two color phases commonly found in Califor-
nia. In the first, general ground color trans-
lucent grayish-white and cerata tipped with
opaque white with subterminal band of
brownish-orange, below which a band of
lighter orange. In the second, general ground
color deep maroon or reddish-purple, cerata
tipped with orange. In both phases, an
opaque white line extends around dorsal
margin of foot, and velar processes tipped
with metallic orange, maroon, yellow or pur-
ple, or white. T.L.: 60 mm, but specimens up
to 200 mm have been collected.
Anus on right, midway between first and
second cerata. Genital aperture on right side,
ventral and just anterior of first ceras.
CALIFORNIA NUDIBRANCHS 177
Radular formula of specimen examined
35(12-20.1.12-20), MacFarland (1966: 262)
reports 34-36(11-20.1.11-20), and Robil-
liard (1970: 448) reports 41-61(11-21.1.11-
21). Rachidian teeth (Fig. 82a) bear 9-18
pointed denticles on either side. First laterals
(Fig. 82b) have 1-8 denticles, while remaining
laterals (Fig. 83c—m) lack denticles, and bear
only the single, terminal cusp. Masticatory
border of mandibles bears 85-100 large den-
ticles.
Type-locality: Santa Barbara, Santa Bar-
bara Co., California.
Range and habitat: Unalaska, Alaska
(Robilliard, 1970), to Islas Coronados, Baja
California, Mexico (Lance, 1961). Intertidal to
215 m. Most commonly found subtidally (ca.
10 m) in association with the tube-dwelling
sea anemone Pachycerianthus fimbriatus
upon which it feeds (Powell, 1966; Wobber,
1970). This species is capable of swimming.
Dendronotus subramosus MacFarland, 1966
Dendronotus subramosus MacFarland, 1966:
255, 265-270 ff, pl. 40, fig. 3; pl. 46, figs.
5-8; pl. 47, figs. 3-7; pl. 49, figs. 1-3; pl.
50, fig. 2; pl. 52, figs. 1-2. Sphon & Lance,
1968: 78. Roller & Long, 1969: 426. Had-
erlie, 1969: tab. 1. Gosliner & Williams,
1970: 177. Robilliard, 1970: 462-466, pl.
64, fig. 33, text figs. 3, 4, 6, 19-21.
Schmekel, 1970: 180. Robilliard, 1972: 421
ff. Nybakken, 1974: 371. Abbott, 1974:
367-368. Thompson 1976b: 92. Mc-
Donald & Nybakken, 1978: 113.
Type-specimens: type material at Calif.
Acad. Sci.
Foot narrow, rounded anteriorly and blunt-
ly pointed posteriorly. Frontal veil bears 4-6
short, slightly branched processes, medial pair
longest. Rhinophores bear 9-14 lamellae. No
processes on rhinophore shaft, as opposed
to other species of this genus in California.
Margin of rhinophore sheath bears 5 short,
blunt processes, longest of which is posterior
and may bear small tubercle-like branches,
but remaining processes are unbranched.
Clavus and shaft translucent grayish-white to
brownish-white. The 3-6 opposite pairs of
cerata are stout, anterior pairs more branched
than posterior pairs. First pair occurs imme-
diately anterior of cardiac region, second pair
occurs immediately posterior of cardiac re-
gion. General ground color translucent gray-
ish-white to brownish-white to white or even
dark brown. Body sprinkled with numerous
lemon yellow or orange dots, especially on
cerata and velar processes; with a smaller
number of brown, red-brown, gold, green, and
white dots. Four distinct, light to dark brown
stripes occur longitudinally on dorsum. Out-
ermost stripe on either side runs from lateral
base of rhinophore shaft to lateral base of
each ceras, and continues on to tail; inner-
most stripe on either side runs from medial
base of rhinophore shaft to medial base of
each ceras, and continues on to tail where
the 4 stripes unite. These stripes may be
lacking in occasional specimens. T.L.: 25 mm.
Anus on right between first and second
cerata. Genital aperture on right side, ventral
and anterior of first ceras.
Radular formula of specimen examined
31(5-6.1.5-6), MacFarland (1966: 266) re-
ports 56-72(5-7.1.5-7), and Robilliard (1970:
464) reports 54-62(2-5.1.2-5). Rachidian
teeth (Fig. 81a) bear 7-12 long, blunt, heavy
denticles on either side. First laterals (Fig. 81b)
have 1-3 irregular denticles, remaining later-
als (Fig. 81c-f) bear 0-8 irregular denticles.
Masticatory border of mandibles bears about
120 transverse, smooth, crescentic ridges.
Type-locality: Monterey Bay, California.
Range and habitat: San Juan Islands, Pu-
get Sound, Washington (Robilliard, 1970), to
Islas Coronados, Baja California, Mexico
(Robilliard, 1970). Intertidal to 107 m. Most
commonly found in rocky intertidal, occasion-
ally on floating docks and pilings in bays.
Feeds upon the hydroid Aglaophenia stru-
thionides (Robilliard, 1970: 466).
Dirona Eliot in Cockerell & Eliot, 1905,
ex MacFarland, MS.
Body limaciform, rather broad, bluntly
rounded anteriorly and tapered posteriorly to
short, pointed tail. Cerata lanceolate, pointed
apically; arranged in closely set, irregular
dorso-lateral rows on dorsum, leaving mid-
dorsal area free. Cerata extend anterior of
rhinophores, but do not meet antero-medial-
ly, and they extend posteriorly where they
meet medially. Cnidosacs absent. Labial ten-
tacles as such absent, in their place a broad,
undulating frontal veil. Rhinophores non-re-
tractile, perfoliate.
Masticatory border of massive mandibles
smooth. Radular formula 1.1.1.1.1, rachidian
teeth bear a single, non-denticulate, median
denticle. Laterals somewhat hooked and bear
a few denticles. Marginals large and hooked,
178 MCDONALD
with no denticles. Labial cuticle covered with
thick-set, hair-like processes.
Penis unarmed. Anus on a papilla far back
on right side, just ventral of cerata.
Type-species: Dirona picta Eliot in Cock-
erell & Eliot, 1905, ex MacFarland, Ms.
Dirona albolineata Eliot in Cockerell & Eliot,
1905, ex Mac Farland, Ms.
Dirona albolineata MacFarland in Cockerell &
Eliot, 1905: 46. MacFarland, 1912: 518-
533, pl. 30, fig. 2; pl. 31, figs. 11-19; pl.
32, fig. 21. O'Donoghue, 1921: 181-183,
pl. 2 (8), figs. 23-24. O'Donoghue, 1922a:
124. O'Donoghue & O'Donoghue, 1922:
132-134, pl. 3, fig. 2. O'Donoghue, 1922b:
164. O'Donoghue, 1924: 24, 31. O'Dono-
ghue, 1926: 224. O'Donoghue, 1927b:
102-103, pl. 3, figs. 64-67. Smith & Gor-
don, 1948: 181. LaRocque, 1953: 260.
Marcus, 1961: 46-47, pl. 9, figs. 159-160.
Lance, 1961: 68. Steinberg, 1963b: 71.
Paine, 1963a: 4. Willows, 1965: 707 ff.
Hurst, 1966: 9 ff. MacFarland, 1966: 298-
302, pl. 30, fig. 13; pl. 56, figs. 1-4; pl. 63,
figs. 9-12; pl. 64, figs. 4-10. Lance, 1966:
79. Hurst, 1967: 255-ff; text figs. 16, 31,
pl. 30, fig. 20; pl. 36, fig. 41; fig. 24-2. Mar-
cus & Marcus, 1967a: 217. Sphon 8 Lance,
1968: 82. Mauzey et al., 1968: 609. Roller
& Long, 1969: 426. Robilliard, 1969a: 290.
Robilliard, 1969b: 23. Turner et al., 1969:
135, append. 1, 2. Gosliner & Williams,
1970: 178. Bernard, 1970: 86. Schmekel,
1970: 188. Robilliard, 1971c: 429 ff. Sphon,
1972a: 155. Blair & Seapy, 1972: 119.
Goddard, 1973: 9. Abbott, 1974: 373. Bel-
ск, 1975: 276% Schuler, 1975: 33. Lam-
bert, 1976: 296, 298. Thompson, 1976a:
39, pl. 3, fig. 6, text fig. b. Thompson,
1976b: 17, 47, 71. Nybakken, 1978: 135.
McDonald & Nybakken, 1978: 114. Mc-
Cosker, 1980: 31. (Non) Baba, 1935b: 116,
120-121, pl. 7, figs. 11-16; pl. 8, figs. 1-
2. (Non) Baba, 1937b: 321. (Non) Volod-
chenko, 1941: 60. (Non) Volodchenko,
1955: 251-252, pl. 48, fig. 10. (=Dirona
akkeshiensis, Baba, 1957).
Dirona sp. MacGinitie & MacGinitie, 1949:
364.
Type-specimens: Mus. Invert. Zool., Stan-
ford Univ.
Rhinophores bear 16-20 lamellae, conical
clavus occupies about half the rhinophore;
rhinophore shaft clear white, lamellae deli-
cate ochre; inner face of rhinophore shaft
bears an opaque white line, starting just be-
low clavus, passing down shaft, between
rhinophores and up opposite shaft. Cerata
smooth, and somewhat flattened antero-pos-
teriorly. Medial cerata are longest, reaching
about one half body length. General ground
color translucent whitish. Cerata also trans-
lucent whitish, with an opaque white line on
lateral margin of each. Frontal veil bears a
band of opaque white along anterior margin,
and crest of tail bears a similar line medially.
Large individuals may have a few small,
opaque white flecks on body and cerata. T.L.:
25 mm, but specimens up to 180 mm have
been collected.
Anus on right, near posterior end, in out-
ermost row of cerata. Genital aperture on
right side, below cerata, about one third of
the distance from anterior end. Penis un-
armed, but bears a spiral row of soft, conical
prominences.
Radular formula of specimens examined
23-25(1.1.1.1.1), MacFarland (1966: 299) re-
ports 29-32(1.1.1.1.1), and Marcus (1961: 47)
reports 22(2.1.2). Rachidian teeth (Fig. 92c,
d) small, and each bears a single, long, thin,
median denticle. Laterals (Fig. 92b) some-
what hooked, with 2-5 denticles. Marginals
(Fig. 92a) large and greatly hooked with no
denticles. Mandibles quite large, masticatory
border smooth.
Type-locality: Monterey Bay, California.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976), to La
Jolla, San Diego Co., California (Lance, 1966).
Intertidal to 37 m, usually in rocky areas.
Rather uncommon in central California. Ro-
billiard (1971c: 419) reports that it is an un-
selective predator, scraping up bryozoans,
hydroids, small crustaceans, sponges, bar-
nacles, and tunicates, and that it feeds upon
the snails Margarites pupillus, Margarites
helicinus, and Lacuna carinata.
Dirona picta Eliot in Cockerell & Eliot,
1905, ex MacFarland, Ms.
Dirona picta MacFarland in Cockerell & Eliot,
1905: 46-48, pl. 7, figs. 6-11. Mac-
Farland, 1912: 517, 518, 520-533, pl. 30,
fig. 1; pl. 31, figs. 1-10; pl. 32, 1195220:
22-24. Hilton, 1919: 34. O'Donoghue,
1926: 224-225. O'Donoghue, 1927b: 101,
102, pl. 3, figs. 60-63. Baba, 1937b: 320.
Hewatt, 1937: 188, 200. Smith & Gordon,
1948: 181. Marcus, 1961: 44-46, 47, pl. 9,
figs. 155-158. Lance, 1961, 68. Farmer 8
CALIFORNIA NUDIBRANCHS 179
Collier, 1963: 62. Steinberg, 1963b: 71.
Paine, 1963a: 4. Paine, 1964: 385. Paine,
1965: 604, 607. Hurst, 1966: 9 ff. Lance,
1966: 69. MacFarland, 1966: 295-299, pl.
56, figs. 5-7; pl. 63, figs. 1-8; pl. 65, figs.
1-3. Farmer, 1967: 342. Marcus & Mar-
cus, 1967a: 216-218, 238, fig. 67. Sphon
& Lance, 1968: 78. Haderlie, 1968: 339.
Roller & Long, 1969: 426. Gosliner & Wil-
liams, 1970: 178. Schmekel, 1970: 188.
Salvini-Plawen, 1972: 393. Holleman,
1972a: 60. Goddard, 1973: 9. Abbott,
1974: 373, fig. 4406. Belcik, 1975: 277.
Thompson, 1976a: 35. Nybakken, 1978:
135. McDonald & Nybakken, 1978: 114.
Dirona picta Eliot in Cockerell & Eliot, 1905,
ex MacFarland, Ms. Keen, 1971: 835, pl.
22, fig. 4.
Type-specimens: Mus. Invert. Zool., Stan-
ford Univ.
Rhinophores bear 8-10 lamellae, clavus
occupies one third to one half of total rhi-
nophore; rhinophore shaft light yellowish-
brown to burnt umber, lamellae a lighter
shade of same color. Cerata inflated and
rather tuberculate. Medial cerata are longest,
reaching about one half of body length. Gen-
eral ground color light yellowish-brown to
burnt umber, occasional specimens may be
greenish. Small pink and cream dots or dark
brown, dull green, and yellow dots may occur
on body. Cerata same color as body, with a
pale red spot on outer side of each ceras,
about one third of the distance from base.
ТЕ: 25 mm:
Anus on right, near posterior end, т out-
ermost row of cerata. Genital aperture on
right side below cerata, about one third of the
distance from anterior end. Penis unarmed,
but bears a spiral row of soft, conical promi-
nences.
Radular formula of specimen examined
24(1.1.1.1.1), MacFarland (1966: 298) re-
ports 32-35(1.1.1.1.1), and Marcus (1961: 45)
reports 24(2.1.2). Rachidian teeth (Fig. 93c,
d) small, each bears a single, thin, median
denticle which is shorter than that in Dirona
albolineata (Fig. 92c, d). Laterals (Fig. 93b)
somewhat hooked, with 3-7 denticles. Mar-
ginals (Fig. 93a) largest and greatly hooked,
with no denticles. Mandibles quite large,
masticatory border slightly roughened.
Type-locality: Dead Man's Island, San Pe-
dro, Los Angeles Co., California.
Range and habitat: Dillon Beach, Marin
Co., California (Marcus, 1961), to Puerto
Rompiente, Baja California, Mexico (Farmer
& Collier, 1963); Puerto Peñasco, Sonora,
Mexico (Farmer, 1967); Puertecitos, Baja
California, Mexico (Farmer 8 Collier, 1963):
also reported from Japan (Baba, 1937b). In-
tertidal to 10 m, usually in rocky intertidal
areas, occasional on floating docks in bays.
May feed upon bryozoans such as Thauma-
toporella sp. Marcus (1961: 45) found sev-
eral cheilostomatous bryozoans, e.g. Celle-
porella hyalina, in the digestive tract. Bertsch
et al. (1972: 306) found the avicularia of a
“Bugula-type bryozoan” in the stomach of
one specimen. Salvini-Plawen (1972: 393) re-
ports that it feeds upon the hydroid Aglao-
phenia sp.
Remarks: Volodchenko (1955: 252) re-
ports the radular formula (4.1.4). Since
MacFarland (1912: 516) states that the rad-
ular formula in the genus Dirona is (2.1.2),
this casts doubt on the identification of Vo-
lodchenko's specimens with D. picta. Fur-
ther, Volodchenko's specimens have a dis-
tinct circle of processes at the base of each
rhinophore, which is absent in specimens of
D. picta from California. In addition, Volod-
chenko's figure (1955: pl. 48, fig. 11) appears
to be copied from MacFarland (1912: pl. 30,
fig. 1), with the addition of the processes at
the bases of the rhinophores. Therefore, it is
difficult to know how closely Volodchenko's
specimens resemble D. picta. This makes the
reports of D. picta by Volodchenko (1941:
60; 1955: 252) rather dubious.
Discodoris Bergh, 1877
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum densely cov-
ered with numerous very small tubercles. La-
bial tentacles cylindro-conical. Foot broadly
rounded and bilabiate anteriorly and some-
what more acutely rounded posteriorly. Rhi-
nophores retractile, perfoliate. Branchial
plumes retractile, tripinnate, arranged in a cir-
cle around anus; branchial aperture slightly
crenulate, stellate, or bilabiate.
Radular formula 0.n.0.n.0, inner laterals
hamate, those nearer margin less hooked.
Penis unarmed; prostate large.
Type-species: Discodoris boholiensis
Bergh, 1877.
Remarks: Thompson (1975: 481, 487)
synonymized Diaulula Bergh, 1880, and An-
isodoris Bergh, 1898, with the older genus
Discodoris Bergh, 1877, stating that Disco-
doris has hemispherical papillae and that all
180 MCDONALD
three genera “contain species of similar size,
shape, texture, mode of life, radula and re-
productive organs and must be merged, be-
cause these similarities far outweigh the dif-
ferences between them (chiefly the presence
of rod-like differentiations in the labial cuticle
of some forms, absent in others). However,
Bergh (1877b: 518) described the dorsum of
Discodoris as finely granulate. (... “die
Rückenseite ist fein granulirt;'') rather than
bearing hemispherical papillae as stated by
Thompson (1975: 487). This would seem suf-
ficient to exclude Anisodoris nobilis, which
has hemispherical papillae, from inclusion in
the genus Discodoris. The type-species of
Anisodoris [Anisodoris punctuolata (Orbigny,
1847)] also has hemispherical or short cone-
shaped papillae on the dorsum (“... der
Rücken uberall bis an den Rand mit Kleinen,
halbkugelformigen oder kurz kegelformigen
Papeln bedeckt... .'' (Bergh, 1898: 509). Un-
til specimens of the types of the genera Dis-
codoris and Anisodoris can be compared, it
seems best to maintain them as distinct gen-
era.
In the case of Diaulula Bergh, 1880, it is
much more difficult to differentiate from Dis-
codoris. Both genera have basically the same
texture of the dorsum, Dicodoris is described
as finely granulate (Bergh, 1877b: 518) and
Diaulula as villous (Bergh, 1880a: 189). The
radular teeth of Diaulula sandiegensis (Coo-
per, 1863) have virtually the same form as
Discodoris boholiensis (Bergh, 1877b: pl. 61,
figs. 8-12), the type of the genus. Bergh
(1880a: 189) states: ‘‘In their general form
the Diaululae somewhat resemble the Dis-
codorides... and that “The radula nearly
agrees with that of the Discodorides ....”
Furthermore, Diaulula presently contains only
about 4 species. On these bases, | agree with
Thompson (1975) that Diaulula is a junior
synonym of Discodoris.
The Eudoridacea contains a number of
poorly defined and often small genera which
further study may show should best be com-
bined into a few broader, well defined genera.
Discodoris heathi MacFarland, 1905
Discodoris heathi MacFarland, 1905: 39-40.
MacFarland, 1906: 118-119, pl. 18, figs.
12-17; pl. 23, fig. 6. O'Donoghue, 1922b:
151-152, 165, pl. 5, figs. 8-11. O'Dono-
ghue, 1924: 23. O'Donoghue, 1926: 207.
O'Donoghue, 1927b: 82, pl. 1, figs. 6-9.
Hewatt, 1937: 200. Costello, 1938: 324,
tabs. 2, 3, 5,.pl. 1, figs. 9,23; pl. 2719929:
Smith & Gordon, 1948: 181. LaRocque,
1953: 259. Marcus, 1955: 150. Marcus,
1961: 17, 19-20, pl. 3, figs. 62-64. Lance,
1961: 66. Farmer & Collier, 1963: 62.
Steinberg, 1963b: 70. MacFarland, 1966:
192-193, pl. 27, fig. 7; pl. 35, figs. 26-33.
Marcus & Marcus, 1967a: 80, 82, 87, 188,
189. Farmer, 1967: 341. Sphon & Lance,
1968: 79. Turner et al., 1969: 135. Thomp-
son, 1969: 763. Roller & Long, 1969: 427.
Long, 1969c: 232. Gosliner & Williams,
1970: 178. Bernard, 1970: 85. Bertsch et
al., 1972: 306. Gosliner & Williams, 1973b:
352. Abbott, 1974: 352, pl. 17, no. 4220.
Thompson, 1975: 487. Belcik, 1975: 276.
Lambert, 1976: 297-298. Bloom, 1976:
289 ff. Bertsch, 1977: 109. Nybakken,
1978: 134 ff. McDonald 8 Nybakken, 1978:
110; 112:
Archidoris montereyensis. Ricketts & Calvin,
1968: text fig. 86.
Type-specimens: U. S. Nat. Mus., no.
181282.
Rhinophores bear 10-16 lamellae and are
light yellow to dusky yellow on shaft and cla-
vus. Rhinophore sheaths low with a slightly
sinuous margin. Branchial plumes 8-10, tri-
pinnate, light yellow to dusky yellow, sprin-
kled with minute brownish flecks, rather whit-
ish on tips. General ground color varies from
light yellow to a more dusky yellow-tan or raw
umber, dorsum usually darker medially. Dor-
sum sprinkled with numerous minute red-
brown and brown to black dots, usually with
a major concentration just anterior of bran-
chial plumes. T.L.: 30 mm.
Radular formula of specimen examined
20(16-20.0.16-20), MacFarland (1966: 192)
reports 20(36-42.0.36-42), and Marcus
(1961: 19) reports 20-22(9-10.31.0.31.9-10),
while Bloom (1976: 293) gives the range 20-
22(36-42.0.36-42). Laterals (Fig. 77) strong-
ly hooked, but become long, thin and only
slightly curved near margin of radula.
Type-locality: Monterey Bay, California.
Range and habitat: Porcher Island, British
Columbia, Canada (Lambert, 1976) to Bahia
San Quintin, Baja California, Mexico (Farmer,
1967). Intertidal to 8 m, usually found in rocky
areas, rarely found on floating docks and pil-
ings in bays. Feeds upon the sponges: Ado-
cia gellindra, Halichondria panicea, and
Myxilla incrustans (Bloom, 1976; McDonald
& Nybakken, 1978).
Remarks: Marcus (1961: 20) states that
CALIFORNIA NUDIBRANCHS 181
Discodoris fulva O'Donoghue, 1924, is prob-
ably a young specimen of D. heathi.
Discodoris sandiegensis (Cooper, 1863)
Doris (Actinocyclus?) sandiegensis Cooper,
1863a: 204. Cooper, 1863b: 58.
Doris (? Actinocyclus) sandiegensis Cooper.
Carpenter, 1864: 608.
Doris sandiegensis Cooper. Carpenter, 1864:
609. Cooper, 1867: 14. Kelsey, 1907: 39.
Odhner, 1926a: 88.
Doridopsis sandiegensis (Cooper). Abraham,
1877: 240. O'Donoghue, 1922d: 143.
Diaulula sandiegiensis (Cooper). Bergh,
1878a: 567 (lapsus).
Diaulula sandiegensis (Cooper). Bergh,
1879a: 344. Bergh, 1879b: pl. 5, figs. 3-5.
Bergh, 1879c: pl. 5, figs. 3-5. Bergh,
1880a: 189-193. Bergh, 1880b: 40-44.
Bergh, 1891: 132. Bergh, 1892: 1097 (105).
Bergh, 1894: 172. MacFarland, 1897: 227,
245-257, pl. 20, figs. 24-31; pl. 21, figs.
33-44; pl. 22, figs. 45-46. MacFarland,
1905: 41. MacFarland, 1906: 122-123, pl.
18, figs. 22-24; pl. 23, fig. 5. Eliot, 1907:
356. Baily, 1907: 92. Berry, 1907: 35.
O'Donoghue, 1921: 159-161, pl. 1 (7), figs.
11-12. O'Donoghue, 1922a: 126. O'Don-
oghue & O'Donoghue, 1922: 137-138.
O'Donoghue, 1922b: 163-164. O’Dono-
ghue, 1922d: 143. O'Donoghue, 1924: 23,
29. O'Donoghue, 1926: 209. Odhner,
1926a: 88, 89. O'Donoghue, 1927a: 10.
O'Donoghue, 1927b: 83-84, pl. 1, figs. 14-
15. Fraser, 1932: 67. Ingram, 1935: 48, 49.
Hewatt, 1937: 200. Costello, 1938: 321,
323-327 ff, tabs. 1-4; pl. 1, figs. 1, 2, 8,
12, 14, 18; pl. 2, figs. 24-25, 36-37. He-
watt, 1946: 193, 198. Smith 8 Gordon,
1948: 181. LaRocque, 1953: 258. Baba,
1957: 9, 13, text fig. 6. Marcus, 1959: 53.
Marcus, 1961: 18-19, 57, pl. 3, figs. 59-
61. Steinberg, 1961: 58. Lance, 1961: 66.
Cook, 1962: 196. Paine, 1963a: 4. Stein-
berg, 1963b: 70. Farmer, 1963: 24. Wil-
lows, 1965: 707 ff. Lance, 1966: 69.
MacFarland, 1966: 190-192, pl. 27, fig. 6;
pL29 fig 15: ply 30 tig: 17; pl. 35, figs:
23-25. Hurst, 1967: 255 ff, text fig. 4a, pl.
27, fig. 6; pl. 35, fig. 40; fig. 24-17. Marcus
8 Marcus 1967a: 189, 238. Beeman,
1968b: 268-269. Sphon & Lance, 1968:
78. Roller & Long, 1969: 426. Turner et al.,
1969: 135. Bertsch, 1969: 231. Robilliard,
1969a: 290. Gosliner & Williams, 1970: 177.
Bernard, 1970: 85. Michel, 1970: 7. North,
1971: 57. Keen, 1971: 825. Holleman,
1972a: 60. Bertsch et al., 1972: 305-306.
Sphon, 1972a: 155. Goddard, 1973: 9.
Haderlie et al., 1974: tab. 4. Abbott, 1974:
353, pl. 17, no. 4234. Ferreira 8 Bertsch,
1975: 327. Schuler, 1975: 33. Belcik, 1975:
276. Bloom, 1976: 289 ff. Elvin, 1976: 194-
198. Lambert, 1976: 296. Michel, 1976: 48,
fig. 12. Thompson, 1976b: 33. Behrens 4
Tuel, 1977: 33, 35. O'Clair, 1977: 443. Ny-
bakken, 1978: 134 ff. McDonald & Nybak-
ken, 1978: 113, 116. Fuhrman et al., 1979:
291 ff.
Diaulula sandiegensis (Cooper) var. Bergh,
1879b: pl. 5, figs. 6-9. Bergh, 1879c: pl. 5,
figs. 6-9. Bergh, 1880a: 193-195. Bergh,
1880b: 44-46. Bergh, 1894: 173-175, pl.
6, figs. 2-5.
Doris (Diaulula) sandiegensis (Cooper). Or-
cutt, 1885: 548.
Diaululua sandiegensis (Cooper). Kelsey
1907: 39 (lapsus).
Peltodoris mauritiana Bergh. Baba, 1935a:
346. Baba, 1935b: 119.
Dialulu sandiegensis (Cooper). MacGinitie &
MacGinitie, 1949: 362, 379 (lapsus).
Dialula sandiegensis (Cooper). McLean, 1962:
111. Harris, 1973: 282, 287. Russo, 1979:
48. (lapsus).
Dilella sandiegagenesis. Hargens, 1977: 363
(lapsus).
Discodoris sandiegensis (Cooper). Bertsch,
1980: 224.
(Non) Diaulula sandiegensis var. pallida
Bergh, 1894: 172-175, (=Diaulula vestita
(Abraham, 1877)).
Type-specimens: not listed; Geological
Survey coll. (Cooper, 1863a).
Labial tentacles small and digitiform. Rhi-
nophores bear 12-30 lamellae, and are ochre
to raw umber. Branchial plumes 6-7, gray-
ish-white to dusky with whitish tips. General
ground color varies from almost white to
chocolate brown, but is most commonly pale
cream to raw umber; dorsum marked with
brown to almost black rings or occasionally
irregular blotches of various sizes, number,
and position. Most commonly these rings are
arranged in 2 longitudinal rows, one on either
side of median line, 3-4 in each row, but this
is highly variable, especially in specimens from
the northern part of the range. One specimen
which was collected lacked rings or blotches.
Specimens from slough or bay areas are typ-
ically darker than specimens from open coast
areas. T.L.: 25 mm, but specimens up to 150
182 MCDONALD
mm have been collected from Elkhorn Slough,
Monterey Co., California.
Radular formula of specimen examined
13(29-31.0.29-31), MacFarland (1966: 191)
reports 19-22(26-30.0.26-30), and Marcus
(1961: 19) reports 19-23(25-34.0.25-34).
Laterals (Fig. 74) hamate.
Type-locality: San Diego Bay, California.
Range and habitat: Unalaska, Alaska
(Bergh, 1894), to Cabo San Lucas, Baja Cal-
ifornia, Mexico (Lance, 1961); Japan (Baba,
1957). Intertidal to 37 m, commonly found in
rocky intertidal, occasional in sloughs and
sometimes found on pilings and docks in
bays. Bloom (1976: 195) reports that it is
found in association with and probably feeds
upon the sponges: Halichondria panicea,
Haliclona permollis, Myxilla incrustans, and
Petrosia dura. McDonald and Nybakken
(1978: 116) report it from Halichondria bow-
erbanki. Cook (1962: 196) states that in the
laboratory it was observed feeding upon Hal-
iclona sp.; Elvin (1976) reports that it feeds
upon Haliclona permollis.
Doridella Verrill, 1870
Body elliptical in outline, quite depressed
and flattened, dorsum entirely smooth and
extends well beyond foot. No postero-medi-
an notch in margin of dorsum such as occurs
in Corambe. Labial tentacles short and ta-
pered. Rhinophores retractile, smooth or with
a few vertical lamellae. Branchial plumes uni-
pinnate or ridge-like, arise posteriorly from
ventral surface of dorsum on either side of
median anus.
Radular formula n.1.0.1.n, laterals large,
with denticles below cusp; marginals smaller
and may bear small denticles.
Penis unarmed.
Type-species: Doridella obscura Verrill,
1870.
Doridella steinbergae (Lance, 1962)
Corambella sp. Steinberg, 1960: 49. Lance,
1961: 67.
Corambella steinbergae Lance, 1962b: 35-
38, pl. 1, text figs. 6-10. Steinberg, 1963b:
71. Sphon & Lance, 1968: 77. McBeth,
1968: 145-146. Gosliner, 1968: 147.
Thompson, 1976a: 36, pl. 4, fig. c.
Corambella bolini MacFarland, 1966: 133,
134-139, pl. 22, figs. 9-11; pl. 29, fig. 21;
pl. 32, figs. 1-12. Sphon & Lance, 1968:
77. Roller, 1970a: 371.
Doridella steinbergae (Lance). Franz, 1967:
75. Marcus & Marcus, 1967b: 208, 209.
Roller & Long, 1969: 427. Roller, 1970a:
371. Gosliner & Williams, 1970: 178. Ber-
nard, 1970: 84. Anderson, 1972: 19. Ab-
bott, 1974: 365. Thompson, 1976b: 48.
Seed, 1976: 1 ff. McDonald & Nybakken,
1978: 111. Bickell & Chia, 1979a: 291 ff.
Bickell & Chia, 1979b: 957.
Type-specimens: Calif. Acad. Sci., Paleo.
type coll. no. 12404.
Anterior margin of foot bluntly rounded or
only slightly emarginate. Labial tentacles short,
slender, and tapered distally. Rhinophores
smooth and tapered to blunt tips, translucent
grayish-white, retractile into rather high
sheaths with thin, entire margins. Branchial
plumes 2-4, translucent grayish-white; larg-
er, innermost plumes consist of a rather tri-
angular stalk which bears 3-4 small lamellae,
outermost plumes bear fewer or no lamellae.
General ground color translucent grayish-
white. Dorsum with approximately 7-10 bro-
ken, irregular, longitudinal, opaque white lines
which are borne upon ridges medially; mar-
ginally dorsum bears similar lines which are
shorter, more broken and more irregular. Be-
tween these lines are scattered small, irreg-
ular blotches of reddish-brown. In all, the col-
or pattern quite resembles the colonies of
Membranipora upon which the animal is most
often found. T.L.: 5 mm.
Radular formula of specimen examined
31(4-5.1.0.1.4-5), MacFarland (1966: 135)
reports 40-60(5.1.0.1.5). Laterals (Fig. 38f)
largest, bearing 3-8 denticles. Marginals (Fig.
38b-e) smaller with minute denticles, outer-
most marginals (Fig. 38a) smallest and non-
denticulate.
Type-locality: San Diego, San Diego Co.,
California.
Range and habitat: British Columbia, Can-
ada (50° N latitude) (Bernard, 1970), to Islas
Coronados, Baja California, Mexico (Lance,
1961). Intertidal and subtidal, almost always
found on the brown alga Macrocystis pyri-
fera, where it feeds upon an encrusting bryo-
zoan of the genus Membranipora (McBeth,
1968: 145); Lance (1962b: 38) states that it
feeds upon Membranipora serrilamella.
Doris Linnaeus, 1758
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum tuberculate.
Labial tentacles short and digitiform. Rhi-
nophores perfoliate, retractile into sheaths
with tuberculate margins. Branchial plumes
retractile, arranged in circle around anus.
CALIFORNIA NUDIBRANCHS 183
Radular formula 0.n.0.n.0, laterals hamate.
Penis acrembolic, no glans penis; vas def-
erens widened to a prostatic canal.
Type-species: Doris verrucosa Linnaeus,
1758:
Doris (s.l.) sp. MacFarland, 1966
Doris (s.l.) species MacFarland, 1966: 179-
181, pl. 25, figs. 1-6. Sphon & Lance, 1968:
79.
Material upon which MacFarland based his
description is at Calif. Acad. Sci.
Dorsum densely covered with numerous
small papillae, giving dorsum a velvety or vil-
lous appearance. Dorsal papillae each encir-
cled by a row of vertical spicules. Foot elon-
gate, elliptical and bilabiate anteriorly, upper
lip notched medially. Labial tentacles slender
and pointed distally. Rhinophores bear 8-18
lamellae, and are white to cream. Branchial
plumes 6, tripinnate, white or cream, tipped
with light brownish. General ground color off
white to cream tan, dorsum bears a number
of large, irregularly oval, light brown spots and
numerous flecks of same color; numerous
minute, opaque white flecks occur on margin
of dorsum, forming a diffuse band. T.L.: 25
mm.
Radular formula of specimen examined
16(22-23.0.22-23). Laterals (Fig. 73) are ha-
mate.
Type-locality: Newport Bay, Orange Co.,
California.
Range and habitat: Pescadero Point, Mon-
terey Co., California (personal observation),
to Newport Bay, Orange Co., California
(MacFarland, 1966). Intertidal to 6 m in rocky
areas, occasionally found upon the sponge
Dysidea amblia upon which it may feed. A
relatively rare species in California.
Remarks: This rather enigmatical species
in many respects (viz. radular teeth, dorsal
papillae, and coloration) resembles Discodor-
is sandiegensis, and may well be conspecific
with it. The coloration and radula are also
much like that of the widely distributed
species Jorunna tomentosa (Cuvier, 1804),
with which it may possibly be synonymous.
More extensive study is necessary to deter-
mine the position of Doris (s.l.) sp.
Doto Oken, 1815
Body limaciform and highly arched. Foot
narrow, linear, rounded anteriorly and ta-
pered posteriorly to short, blunt tail; anterior
foot corners rounded and not produced.
Cerata bulbous, tuberculate, rather club-
shaped, arranged in dorso-lateral series on
either side of dorsum; the larger, anterior
pairs bear a plume-like “gill'” on their inner
surface. Cerata lack cnidosacs and are de-
ciduous. Oral tentacles as such are absent;
in their place is a rounded frontal veil. Rhino-
phores smooth and bluntly tapered, retractile
into calyciform rhinophore sheaths.
Masticatory border of thin mandibles
smooth. Radula uniseriate, with numerous
teeth which are somewhat asymmetrical and
bear a few denticles.
Penis unarmed. Anus on right side, in line
of cerata.
Type-species: Doto coronata (Gmelin,
1791).
See ICZN (1964, Opinion 697) for valida-
tion of generic name and designation of type-
species.
Doto amyra Marcus, 1961
Doto amyra Marcus, 1961: 38, pl. 7, figs.
130-134. Steinberg, 1963a: 65. Steinberg,
1963b: 71. Marcus 8 Marcus, 1967a: 214,
238. Sphon & Lance, 1968: 79. Schmekel,
1968b: 6. Long, 1969c: 232. Gosliner 8
Williams, 1970: 178. Schmekel, 1970: 184.
Keen, 1971: 832. Thompson, 1972a: 75.
Nybakken, 1974: 371. Abbott, 1974: 371.
McDonald & Nybakken, 1978: 113.
Doto ganda Marcus, 1961: 39, 57, pl. 7, figs.
135-138. Farmer & Collier, 1963: 63.
Schmekel, 1970: 184. Abbott, 1974: 371.
Doto wara Marcus, 1961: 40-41, 57, pl. 8,
figs. 143-146. Schmekel, 1970: 184.
Thompson, 1972a: 75. Abbott, 1974: 371.
Doto amyra ? Marcus. Roller 8 Long, 1969:
427.
Doto gauda Marcus. Thompson, 1972a: 75
(lapsus).
Type-specimens: Dept. Zool., Univ. Sáo
Paulo, Brazil.
Frontal veil smooth, rounded, and slightly
laterally expanded, with an entire margin.
Rhinophore sheaths have rather smooth
margins. Rhinophores and sheaths both pig-
mented distally with numerous small, opaque
white dots. The 5-7 opposite pairs of cerata
bear numerous rather long, well spaced tu-
bercles which occur in 3-7 rings around each
ceras. General ground color translucent
cream-white. Cerata cores vary from cream
to pink, yellow, or light orange. T.L.: 8 mm.
Anus on a high papilla on right, in line of
184 MCDONALD
cerata, near second ceras. Genital aperture
on right side, just ventral of first ceras.
Radular formula of specimen examined
87(0.0.1.0.0), Marcus (1961: 38) reports about
70 teeth on radula. Rachidian teeth (Fig. 83)
asymmetrical, usually with 1-3 denticles on
either side of median denticle.
Type-locality: Monterey Bay, California.
Range and habitat: Dillon Beach, Marin
Co., California (Marcus, 1961), to Ensenada,
Baja California, Mexico (Farmer 8 Collier,
1963). A single specimen has been reported
from Puerto Penasco, Sonora, Mexico (Mar-
cus, 1967a: 214). Intertidal and subtidal, fre-
quently found on the hydroids: Aglaophenia
struthionides and Sertularia furcata upon
which it may feed; Marcus (1961: 39) found
one specimen on Obelia.
Doto columbiana O'Donoghue, 1921
Doto columbiana O'Donoghue, 1921: 204-
205, pl. 3 (9), fig. 33. O'Donoghue, 1926:
235. Odhner, 1936: 1119. Marcus, 1961:
36-38 ff, pl. 7 figs. 125-129. Steinberg,
1963b: 71. MacFarland, 1966: 288-289,
295. Gosliner & Williams, 1970: 178. Ber-
nard, 1970: 85. Schmekel, 1970: 184.
Marcus, 1972b: 303. Abbott, 1974: 371.
Belcik, 1975: 276. Lambert, 1976: 297,
298.
Idulia columbiana (O'Donoghue). O'Dono-
ghue, 1926: 235. LaRocque, 1953: 251.
Type-specimens: Mus. Dominion Biol.
Stat., Nanaimo, British Columbia, Canada.
Frontal veil rounded and slightly laterally
expanded, with a few small tubercles dorsally
and an entire margin. Rhinophore sheaths
have smooth margins. Rhinophores and
sheaths translucent grayish-white. The 5-7
opposite pairs of cerata bear numerous short,
flat tubercles which occur in 4-5 rings around
each ceras. General ground color translucent
grayish-white to pale grayish-yellow. Irregu-
lar, scattered lines of black occur on dorsum
and sides of body. Tubercles on cerata each
bear a black ring at their base. T.L.: 10 mm.
Anus on right, just anterior of second cer-
as. Genital aperture on right side, just ventral
of first ceras.
Marcus (1961: 37) gives the radular for-
mula 83(0.0.1.0.0), and MacFarland (1966:
288-289) reports 86-96(0.0.1.0.0). Rachidi-
an teeth (Fig. 84) asymmetrical, usually with
3-5 denticles on either side of median denti-
cle.
Type-locality: Nanoose Bay, Vancouver Is-
land, British Columbia, Canada.
Range and habitat: Pearse Island, British
Columbia, Canada (Lambert, 1976), to Dux-
bury Reef, Marin Co., California (Gosliner 8
Williams, 1970). Intertidal to 60 m, usually
found on hydroids.
Doto kya Marcus, 1961
Dota kya Marcus, 1961: 39-40, pl. 8, figs.
139-142. Steinberg, 1963b: 71. Haderlie,
1968: 333, 339. Schmekel, 1968b: 6. Had-
erlie, 1969: tabs. 1-2. Roller 8 Long, 1969:
427. Roller, 1970a: 372. Schmekel, 1970:
184. Thompson, 1972a: 75. Bertsch et al.,
1972: 306. Nybakken, 1974: 371. Abbott,
1974: 371:
Doto varians MacFarland, 1966: 288, 289-
295, pl. 42, figs. 1-8: pl. 44, figs. 8-17; pl.
48, figs. 9-13 (partim). Roller, 1970a: 372-
373. Bernard, 1970: 85. Schmekel, 1970:
184. Abbott, 1974: 371.
Type-specimens: Dept. Zool., Univ. Sáo
Paulo, Brazil.
Frontal veil smooth and slightly laterally
expanded, margin entire. Rhinophore sheaths
have scalloped margins which are slightly ex-
panded anteriorly. Rhinophores and sheaths
both bear numerous, small, opaque white
dots distally. The usually 8, opposite pairs of
cerata bear numerous ovoid tubercles which
occur in 4-5 rings around each ceras. Gen-
eral ground color translucent yellowish white.
Dorsum, sides of body, and cerata bear nu-
merous brownish-black spots and blotches.
Cerata cores usually brownish. T.L.: 8 mm.
Anus on right, just anterior of second cer-
as. Genital aperture on right side, ventral of
first ceras.
Radular formula of specimen examined
61(0.0.1.0.0), Marcus (1961: 40) reports
95(0.0.1.0.0). Rachidian teeth (Fig. 85) asym-
metrical, usually with 2-3 denticles on either
side of median denticle.
Type-locality: Point Pinos, Monterey Bay,
California.
Range and habitat: Duxbury Reef, Marin
Co., California (Gosliner 4 Williams, 1970), to
Shell Beach, San Luis Obispo Co., California
(Roller 8 Long, 1969). Intertidal and subtidal,
usually found upon the hydroid Aglaophenia
struthionides upon which it may feed.
Remarks: The species of the genus Doto
found along the California coast are present-
ly quite confused. Marcus (1961: 38-41)
CALIFORNIA NUDIBRANCHS 185
named four new species of Doto (D. amyra,
D. ganda, D. kya, and D. wara) from Califor-
nia. While further study is necessary, | rec-
ognize only D. amyra and D. kya and consid-
er D. ganda and D. wara synonyms of D.
amyra. MacFarland (1966: 289-295) named
Doto varians which appears to be, in part at
least, a junior synonym of D. kya, and also
possibly of D. amyra. A detailed study of the
California species of Doto is needed to clear-
ly define and differentiate the species of the
genus found in California. Preliminary evi-
dence from development and larval morphol-
ogy seems to indicate that there are more
valid species of Doto in California than are
accepted in this paper. However, no clearly
defined morphological differences have yet
been observed which will clearly distinguish
more than the three species herein recog-
nized.
Eubranchus Forbes, 1838
Body aeolidiform, rather high and narrow.
Foot narrow, linear, and elongate, tapered
posteriorly to tail; anterior foot corners usu-
ally rounded and not produced, but at least
one species (Е. misakiensis) has produced
foot corners. Cerata long, cylindrical, some-
what clavate, and rather irregularly inflated.
Oral tentacles cylindrical and tapered slightly
to blunt tips. Rhinophores non-retractile, long,
smooth, and tapered slightly to blunt tips.
Masticatory border of mandibles denticu-
late. Radula triseriate, rachidian teeth bear a
few denticles on either side of median cusp;
laterals rather thin, with a single, non-dentic-
ulate cusp.
Penis armed with a chitinous stylet in most
species. Anus acleioproct.
Type-species: Eubranchus tricolor Forbes,
1838.
See ICZN (1966b, Opinion 774), Eubran-
chus placed on Official List.
Eubranchus misakiensis Baba, 1960
Eubranchus misakiensis Baba, 1960b: 300-
301, pl. 34, figs. 2a-g. Hamatani, 1961:
353, 358-361, text figs. 5-8. Baba, 1964:
287. Burn, 1964: 14. Thompson, 1967: 9.
Edmunds & Kress, 1969: 907. Behrens,
1971b: 214-215. Baba, 1971: 63-64, 66,
pl. 6, figs. 1-6. Behrens & Tuel, 1977: 33,
35. Clark 8 Goetzfried, 1978: 290.
Type-specimens: Biol. Lab., Imperial Pal-
ace, Japan.
Foot corners produced into rather long,
tentaculiform processes. Oral tentacles col-
ored as rhinophores but usually bear only 0-
1 chocolate brown bands. Rhinophores
translucent yellowish-white, with a number of
small, chocolate brown dots which may be
concentrated into 2-3 bands. Cerata ar-
ranged in about 6 oblique rows dorso-later-
ally on either side of dorsum. General ground
color translucent yellowish-white. Dorsum,
sides of body, and cerata bear numerous
small, round, chocolate brown dots. Cerata
bear yellow to brown subapical ring, cores
bluish-green. T.L.: 5 mm.
Anus just anterior of third group of cerata,
slightly to right of midline. Genital aperture on
right side, just ventral and posterior of first
group of cerata. Penis unarmed.
Radular formula of specimen examined
46(0.1.1.1.0), Baba (1960b: 301) reports
40(0.1.1.1.0). Rachidian teeth (Fig. 105a) bear
3-4 denticles on either side of median cusp.
Laterals (Fig. 105b) rather thin plates with a
single, triangular cusp. Masticatory border of
mandibles bears a single row of about 15
denticles.
Type-locality: Misaki, Sagami Bay, Japan.
Range and habitat: Introduced into San
Francisco Bay, California (Behrens, 1971b);
originally described from Japan. Most often
on floating docks and pilings in bays, rather
rare in California.
Eubranchus olivaceus (O'Donoghue, 1922)
Galvina olivacea O'Donoghue, 1922b: 158-
160, 165, pl. 6, figs. 21-22.
Eubranchus olivaceus (O’Donoghue). O'Don-
oghue 8 O'Donoghue, 1922: 135, pl. 4, fig.
7. O'Donoghue, 1924: 25-26. O'Dono-
ghue, 1926: 230. LaRocque, 1953: 251.
Steinberg, 1963b: 72. Hurst, 1967: 255 ff,
textitigs 20; р! 32, fig: 26: pl. 38, fig. 51:
fig. 25-3. Edmunds & Kress, 1969: 907.
Bernard, 1970: 86. Nybakken, 1974: 371.
Abbott, 1974: 375. Lambert, 1976: 298.
Rivest & Harris, 1976: 146-147. Thomp-
son, 1976a: 79, text fig. 38g. McDonald &
Nybakken, 1978: 114.
Type-specimens: not listed.
Foot corners rounded, not produced. Oral
tentacles colored as, and shorter than rhi-
nophores. Rhinophores translucent yellow-
white with a subterminal band of brownish to
186 MCDONALD
olivaceous-green, and encrusted with white
dots which are more concentrated distally,
Cerata arranged in about 4-7 indistinct
groups on dorsum. First group contains about
4-6 cerata, posterior of first group, cerata
are nearly alternately arranged. General
ground color pale translucent yellow-green to
yellow-white. An irregular network of red-
dish-brown lines occurs on dorsum, along
with a faint, irregular, dorso-medial band of
olive green. A few opaque white dots occur
on head and dorsal surface of tail. Cerata
slightly frosted with white and bear an indis-
tinct, subterminal band of reddish-brown to
olivaceous-green, cores deep olivaceous
green. T.L.: 8 mm.
Radular formula of specimen examined
35(0.1.1.1.0), O'Donoghue (1922b: 159) re-
ports 32-33(0.1.1.1.0). Rachidian teeth (Fig.
106a) bear 5-6 denticles on either side of
median cusp. Laterals (Fig. 106b) rather thin
plates with a single triangular cusp. Masti-
catory border of mandibles finely denticulate.
Anus just anterior of third group of cerata,
slightly to right of midline. Genital aperture on
right side, postero-ventral of first group of
cerata.
Type-locality: Jesse Island, Vancouver Is-
land, British Columbia, Canada.
Range and habitat: Vancouver Island, Brit-
ish Columbia, Canada (O'Donoghue, 1922b),
to Asilomar Beach, Pacific Grove, Monterey
Co., California (Nybakken, 1974: personal
observation). Intertidal to 10 m. Rare in rocky
intertidal zone in California. O'Donoghue
(1922b: 159) states that it is usually found on
the hydroid Obelia longissima growing at the
base of Zostera marina. Although Sphon
(1972b: 64) gives the geographic range of E.
olivaceus as Bamfield, British Columbia,
Canada, to Bahía de los Angeles, Baja Cali-
fornia, Mexico, this is the range of Eubran-
chus rustyus (Robilliard, 1971a; Lance, 1961).
Since E. rustyus is not included in Sphon's
(1972b) list of nudibranchs of the west coast
of North America and it is reported elsewhere
in the literature to occur south of Asilomar
Beach, Monterey Co., California, it seems
certain that E. rustyus and E. olivaceus were
inadvertently combined into a single entry us-
ing the name of the latter and the range of
the former in Sphon’s list.
Remarks: Meyer (1971: 147-148) report-
ed E. olivaceus from Maine. However, Rivest
& Harris (1976: 146-147) suggest that she
collected Eubranchus exiguus (Alder 8 Han-
cock, 1848) rather than E. olivaceus.
Eubranchus rustyus (Marcus, 1961)
Capellinia rustya Marcus, 1961: 49-50, 57,
pl. 9, figs. 168-172. Lance, 1961: 68.
Steinberg, 1963b: 72. Paine, 1963a: 4.
Lance, 1966: 69. Sphon 8 Lance, 1968:
82. Roller & Long, 1969: 426. Roller, 1970a:
372. Abbott, 1974: 375.
Eubranchus occidentalis MacFarland, 1966:
323-326, pl. 62, fig. 7; pl. 65, figs. 19-25;
pl. 66, fig. 7. Roller, 1970a: 372.
Eubranchus rustyus (Marcus). Edmunds &
Kress, 1969: 907. Gosliner 8 Williams,
1970: 178. Robilliard, 1971a: 163, 165.
Keen, 1971: 837. Bertsch et al., 1972: 306.
Nybakken, 1974: 371. Keen & Coan, 1975:
45. Lambert, 1976: 298. McDonald & Ny-
bakken, 1978: 114.
Eubranchus (=Capellinia) rustyus (Marcus).
Robilliard, 1971a: 163-164.
Type-specimens: Dept. Zool., Univ. Sao
Paulo, Brazil.
Foot corners rounded, not produced. Oral
tentacles just over one half the length of the
rhinophores; colored as rhinophores with a
similar subapical band. Rhinophores a light
shade of body color with translucent whitish
tips and a subapical band of brownish to light
gray or greenish, and a few small, irregular
spots of the same color. Cerata may bear
tubercles, and are arranged in 4-6 groups on
either side of dorsum, with 2-6 cerata in each
group. General ground color varies from
grayish-white to light cream or yellowish to a
light brownish-yellow, numerous irregular
specks of brownish to light gray or greenish
over body and on cerata. Cerata each bear a
diffuse, subapical band of brownish to light
gray or greenish, and whitish tip; cores vary
from yellowish-green to brown. T.L.: 8 mm.
Anus just posterior of second group of ce-
rata, slightly to right of midline. Genital aper-
ture on right side, just ventral of first group
of cerata. Penis armed with a chitinous sty-
let.
Radular formula of specimen examined
59(0.1.1.1.0), Marcus (1961: 49) reports 50-
60(0.1.1.1.0), and MacFarland (1966: 324)
reports 58-62(0.1.1.1.0). Rachidian teeth (Fig.
107a) bear 3-5 denticles on either side of
median cusp. Laterals (Fig. 107b) are rather
thin plates with a single, triangular cusp.
Masticatory border of mandibles bears a sin-
gle row of about 10 transverse, ridge-like
denticles which may bear 12-20 small dentic-
ulations.
Type-locality: Monterey Bay, California.
CALIFORNIA NUDIBRANCHS 187
Range and habitat: Vancouver Island, Brit-
ish Columbia, Canada (Robilliard, 1971a), to
Punta Abreojos, Baja California, Mexico
(Keen, 1971); Bahia de los Angeles, Gulf of
California, Mexico (Lance, 1961). Intertidal
and subtidal, most common on floating docks
in bays, where it is usually found on the hy-
droids Obelia spp. Occasionally found in rocky
intertidal. It has been found on the hydroids:
Obelia sp., Plumularia lagenifera, and on Hy-
dractinia sp. at the base of the alga Cysto-
seira osmundacea (MacFarland, 1966; Mar-
cus, 1961; Robilliard, 1971a).
Fiona Alder & Hancock, 1855
Body aeolidiform, elongate, and broadest
medially. Foot rounded anteriorly and ta-
pered posteriorly to rather long tail, margins
thin and extend laterally well beyond body.
Anterior foot corners rounded, not produced.
Cerata cylindrical and tapered, the majority
bear a thin, sail-like expansion on proximal
three quarters of posterior surface; cnido-
sacs present. Oral tentacles cylindrical and
tapered to blunt tips. Rhinophores non-re-
tractile and smooth.
Masticatory border of mandibles denticu-
late. Radula uniseriate, with denticles on
either side of median cusp.
Penis unarmed. Anus acleioproct.
Type-species: Fiona pinnata (Eschscholtz,
1831).
Fiona pinnata (Eschscholtz, 1831)
Limax marinus Forskäl, 1775: 99. Cuvier,
1817: 25. O'Donoghue, 1922d: 145.
Lemche, 1964a: 37. (Non) Gunnerus, 1770:
170.
Doris fasiculata Gmelin in Linnaeus, 1791:
3104. (Non) Müller, 1776: 229, no. 2772.
Eolis fasciculata. Lamarck, 1819: 302.
Eolis leuconotus Hasselt in Ferussac, 1824a:
82.
Eolidia alba Hasselt in Férussac, 1824b: 239.
O'Donoghue, 1922d: 145.
Eolidia fasciculata. Bruguière 8 Lamarck,
1330: 115.
Eolidia pinnata Eschscholtz, 1831: 14, pl. 19,
fig. 1. Carpenter, 1857: 173. O'Donoghue,
1922d: 145.
Eolidia longicauda Quoy 8 Gaimard, 1832:
288-290, pl. 21, figs. 19-20. Finlay, 1927:
441.
Oithona nobilis Alder & Hancock in Forbes 4
Hanley, 1851a: 589. Alder 8 Hancock,
1851b: 291. Hancock, 1852: 74. Chenu,
1859: 413, fig. 3078. O'Donoghue, 1922d:
145. Pruvot-Fol, 1927: 45.
Fiona nobilis (Alder 8 Hancock). Adams &
Adams, 1854: 77, pl. 67, fig. 1. Alder 8
Hancock, 1855: 31, 32, 53, fam. 3, pl. 38a,
app. 23. Gosse, 1856: 100. Gray, 1857:
227. Bergh, 1859: 8-9. Jeffreys, 1869: 35,
pl. 2, fig. 2. Verrill, 1881: 300. Verrill, 1882a:
339. Verrill, 1882b: 551-552. Locard, 1886:
52. Tempere, 1900: 116. Vayssiere, 1901:
305. Grosvenor, 1903: 470. Pruvot-Fol,
1927: 45. White, 1938: 16.
Aeolis pinnata (Eschscholtz). Carpenter,
18572 313.
Fiona atlantica Bergh, 1858: 273-337, pls.
2-3. Bergh, 1859: 9-16, pl. 1, figs. 1-29;
pl. 2, figs. 30-53. Bergh, 1871: 1287-1288,
pl. 13, fig. 8. O'Donoghue, 1922d: 145.
Pruvot-Fol, 1927: 45.
Fiona longicauda (Quoy 8 Gaimard). Bergh,
1859: 16-18. Bergh, 1892: 1035 (43).
Bergh, 1894: 130. Simroth, 1895: 184.
Hymenaebolis elegantissima Costa, 1867: 29-
30, pl. 1, figs. 1-3. O'Donoghue, 1922d:
145. Iredale & O'Donoghue, 1923: 231.
Fiona pinnata (Eschscholtz). Bergh, 1859: 3.
Bergh, 1873a: 606-610, pl. 8, figs. 2-11;
pl. 9, fig. 13. Bergh, 1873b: 87, pl. 12, fig.
45. Bergh, 1892: 1035 (43). Bergh, 1894:
130. Iredale 8 O'Donoghue, 1923: 212-
213. O'Donoghue, 1926: 234. Finlay, 1927:
441. Pruvot-Fol, 1927: 45. O'Donoghue,
1929: 754. Winckworth, 1932: 237. John-
son, 1934: 155-156. Baba, 1937a: 197,
200. Baba, 1937b: 333. Graham, 1938:
300. White, 1938: 16, 18. Smith 8 Gordon,
1948: 181. Baba, 1949: 101, 176, pl. 43,
fig. 149. Tokioka, 1952: 13-14, text figs.
1-8. Baba & Hamatani, 1952: 9. La-
Rocque, 1953: 251. Graham, 1955: 153.
Haefelfinger, 1960: 341. Marcus, 1961: 50,
56, pl. 10, figs. 173-179. Lance, 1961: 68.
Steinberg, 1963b: 72. Bayer, 1963: 460-
465, figs. 5-7. Thompson, 1964: 277 ff.
Bennett, 1966: 46, pl. 11, fig. 2; pl. 12, figs.
1-2. Edmunds, 1966: 31 ff. Bieri, 1966:
166, 168. MacFarland, 1966: 355-358, pl.
68, figs. 23-28; pl. 70, figs. 11-11a, 12,
12-1’. 12-2’. Burn, 1966: 24. Burn, 1967a:
116-117, figs. 1-2. Marcus 8 Marcus,
1967a: 109, 124. Marcus 8 Marcus, 1967b:
216. Rees, 1967: 218. Sphon 4 Lance,
1968: 79. Schmekel, 1968a: 121, 147.
Schmekel, 1968b: 5. Roller 8 Long, 1969:
429. Gosliner & Williams, 1970: 178. Ed-
munds, 1970: 54, 55. Bernard, 1970: 86.
Schmekel, 1970: 136, 169-170, text figs.
188 MCDONALD
32-33. Keen, 1971: 837. Harris, 1971a: 84.
Schmekel, 1971: 123. Zeiller & Compton,
1971: 377. Holleman, 1972b: 142-146, text
fig. 5. McBeth, 1972a: 55. Salvini-Plawen,
1972: 388, 392. Abbott, 1974: 377. Bing-
ham 8 Albertson, 1974: 220, 223. Ed-
munds, 1975: 269. Ros, 1975: 340, 358.
Thompson & Brown, 1976: 192, fig. 104.
Thompson, 1976a: 35. Thompson, 1976b:
51, 67. Marcus, 1977: 14. McDonald & Ny-
bakken, 1978: 115. Ros, 1978a: 153 ff.
Ros, 1978b: 25 ff. Bertsch, 1979a: 58.
Eolis armoricana Hesse, 1872: 347.
Fiona marina var. pacifica Bergh, 1879b: 86-
88, pl. 1, figs. 7-8. Bergh, 1879c: 142-144,
pl. 1, figs. 7-8. Bergh, 1894: 130-132, pl.
1, figs. 13-15. O'Donoghue, 1922d: 145.
Marcus, 1959: 89. MacFarland, 1966: 355.
Fiona elegantissima (Costa). Tiberi, 1880:
236.
Cratena plicata Hutton, 1882: 166, pl. 6, fig.
1:
Fiona marina (Forskäl). Bergh, 1877c: 823-
824. Bergh, 1884a: 9-10, pl. 11, fig. 1.
Fischer, 1887: 541. Herdman & Clubb,
1889: 225, 235. Carus, 1889-1893: 215.
Bergh, 1892: 1035 (43). Herdman & Clubb,
1892: 146. Bergh, 1894: 130. Simroth,
1895: 165, 184. Herdman, 1896: 49. Herd-
man et al., 1896: 446. Tregelles, 1896: 223.
Bergh, 1898: 560-561. Cooke, 1899: 65.
Bergh, 1900: 239. Conchol. Soc., 1901: 25.
Casteel, 1904a: 325-405, pls. 21-35. Cas-
teel, 1904b: 505. Eliot, 1910: 5, 166. Eliot,
1913: 44-45. Sumner et al., 1913: 706.
Johnson, 1915: 168. Scott, 1922: 49.
O'Donoghue, 19224: 145-147. Finlay,
1927: 441. Pruvot-Fol, 1927: 45. Issel,
1928: 3-7. Russell, 1929: 219 ff, pl. 7.
Kropp, 1931: 120. White, 1938: 18. Mar-
cus, 1959: 93. Thompson, 1961: 237. Burn,
1962: 106. MacFarland, 1966: 355. Franz,
1975b: 250.
Fiona alba (Hasselt). Bergh, 1892: 1035 (43).
Simroth, 1895: 184.
Fiona ? alba (Hasselt). Bergh, 1894: 130.
Fiona primata (Eschscholtz). Simroth, 1895:
184 (lapsus).
Cratena ? spec. ? Simroth, 1895: 170-171,
pl. 20, figs. 11-22.
Cratena longicauda Heinke, 1897: 249, text
fig. 4.
Aeolis longicauda Quoy & Gaimard. O’Don-
oghue, 1922d: 145.
Dolicheolis longicauda (Quoy & Gaimard).
Finlay, 1927: 441.
Aeolis. Savilov, 1956: pl. fig. 5, text fig. 8.
Fiona. Bieri, 1970: 305.
Fione pinnata. Thiriot-Quiévreux, 1977: 178
(lapsus).
Type-specimens: not listed.
Oral tentacles about same length as rhi-
nophores and colored as body. Rhinophores
rather long with blunt tips, translucent gray-
ish-white to cream. Cerata borne upon thin,
marginal expansion of dorsum, with 4-6 or
more cerata in each transverse row. General
ground color translucent grayish-white; body
varies somewhat from pale cream to rather
pale raw umber, with a slight pinkish cast be-
tween and anterior of rhinophores. Cerata
colored as body, cores vary from brown to
raw umber; or bluish-purple in specimens
which have been feeding upon Velella vellel-
las О mms
Anus medial, slightly anterior of middle of
body. Genital aperture on right side, well for-
ward and ventral of area between oral ten-
tacles and rhinophores.
Radular formula of specimen examined
35(0.0.1.0.0), MacFarland (1966: 356) re-
ports 40(0.0.1.0.0), and Marcus (1961: 50)
reports 36-40(0.0.1.0.0). Rachidian teeth (Fig.
112) bear 5-7 small denticles on either side
of large, median cusp. Masticatory border of
mandibles bears one row of round to cres-
cent-shaped denticles.
Type-locality: Sitka, Alaska.
Range and habitat: Nearly cosmopolitan
(Lance, 1961), pelagic on floating objects, es-
pecially on logs with the barnacles Lepas spp.
Feeds upon the barnacles: Lepas anatifera,
Lepas anserifera, and Lepas sp., and upon
the chondrophorans: Porpita sp. and Velella
velella (Bergh, 1880b; Bieri, 1966; Burn,
1966; Eliot, 1910; Kropp, 1931; Marcus,
1961; McDonald & Nybakken, 1978; Rees,
1967; Savilov, 1956; Schmekel, 1968a;
Thompson & Brown, 1976). Kropp (1931:
121) also mentions that it feeds upon small
jellyfishes and crustaceans when it occurs in
the intertidal.
Hallaxa Eliot, 1909
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum may be
smooth or tuberculate. Labial tentacles short
and somewhat variable in form. Rhinophores
perfoliate, retractile into sheaths. Branchial
plumes unipinnate, retractile, arranged in a
circle around anus.
Labial disc bears spines. Radular formula
CALIFORNIA NUDIBRANCHS 189
n.1.1.1.n, rachidian teeth large and bear a
large cusp and a few smaller denticles below.
Marginals bear denticles below cusp.
Penis unarmed.
Type-species: Hallaxa decorata (Bergh,
1878).
Eliot (1909) proposed Hallaxa to replace
Halla Bergh (1878a), the latter being preoc-
cupied.
Hallaxa chani Gosliner & Williams, 1975
Hallaxa sp. McDonald, 1975a: 529, 540.
Hallaxa chani Gosliner & Williams, 1975: 396-
405, text figs. 1-10. Nybakken, 1978: 135.
McDonald & Nybakken, 1978: 110, 112.
Bertsch, 1980: 222. Jaeckle, 1981: 240.
Type-specimens: Calif. Acad. Sci., no. 674.
Dorsum covered with numerous tubercles
which are larger and more numerous near
margin. Foot relatively narrow and elongate,
anterior corners slightly extended laterally,
bluntly rounded posteriorly. Rhinophores bear
8-12 lamellae, and are pale yellow with
brownish-maroon distally. Branchial plumes
12-14, translucent yellowish-white with small
reddish-maroon spots near base of each
plume. General ground color light lemon yel-
low, varying from dull grayish-yellow in small
specimens to richer yellow in larger speci-
mens. Dorsum bears numerous reddish-
brown flecks of various sizes, and is some-
what darker medially; an irregular series of
larger, dark brown spots occurs medially, an-
terior of branchial plumes. Some specimens
bear a thin, marginal band of golden-brown
on dorsum. T.L.: 15 mm.
Radular formula of specimen examined
45(6-18.1.1.1.6-18), Gosliner & Williams
(1975: 401) report 35-36(7-14.1.1.1.7-14).
Rachidian teeth small and vestigial. Laterals
(Fig. 63a) each bear one large cusp and 3-5
small, blunt denticles. Marginals (Fig. 63b-e)
have 5-9 small denticles below cusp.
Type-locality: Duxbury Reef, Marin Co.,
California.
Range and habitat: Abalone Beach, Hum-
boldt Co., California (Jaeckle, 1981), to Shell
Beach, San Luis Obispo Co., California (Gos-
liner & Williams, 1975). Most individuals have
been found in rocky intertidal zone, but it is
also occasionally found on floating docks and
pilings in bays. Feeds upon the ascidian Di-
demnum carnulentum (McDonald & Nybak-
ken, 1978).
Hancockia Gosse, 1877
Body rather limaciform, somewhat elon-
gate, and compressed. Cerata digitate lobes,
arranged dorso-laterally on either side of dor-
sum. Oral tentacles as such are absent; fron-
tal veil prolonged into digitate lobes. Clavi of
rhinophores perfoliate and retractile into
sheaths on rhinophore shafts.
Labial disc armed with small rodlets. Mas-
ticatory border of mandibles denticulate.
Radula triseriate. Rachidian teeth bear a few
denticles on either side of median cusp. Lat-
erals somewhat rectangular plates with a sin-
gle cusp.
Anus located between first and second
cerata on right side. Penis unarmed.
Type-species: Hancockia eudactylota
Gosse, 1877.
Hancockia californica MacFarland, 1923
Hancockia californica MacFarland, 1923: 65-
104, pls. 1-6. O'Donoghue, 1926: 225.
Odhner, 1936: 1068 ff, text fig. 9. Smith &
Gordon, 1948: 181. Forrest, 1953: 234.
Marcus, 1957: 451, 454. Marcus, 1961:
33-34, 57, 58, pl. 7, figs. 119-120. Lance,
1961: 67. Steinberg, 1963b:71. deVries,
1963: 129. Ghiselin, 1965: 350. Mac-
Farland, 1966: 245, 246-254, pl. 38, figs.
7-9; pl. 43, figs. 45-50; pl. 49, fig. 7; pl.
50, fig. 5; pl. 52, figs. 7-7a; pl. 53, figs. 1-
6. Sphon 8 Lance, 1968: 82. Roller & Long,
1969: 427. Thompson, 1972a: 70, 72. Ny-
bakken, 1974: 371. Abbott, 1974: 369, fig.
4376.
Type-specimens: not listed.
Dorsum separated from foot by a well de-
fined, longitudinal groove. Foot narrow, line-
ar, bluntly rounded anteriorly with a narrow,
median notch, and slightly tapered posterior-
ly to bluntly rounded tail which bears a slight,
median notch. Anterior margin of body bears,
on either side, a broad, palmate, velar lobe
which bears 6-10 or more unequal, digitiform
processes. Clavi of rhinophores bear 6-8,
vertical lamellae, and a short blunt apex. Six
to nine irregularly spaced, rounded ridges oc-
cur along the thin margin of calyciform rhi-
nophore sheath. Rhinophores colored as
body. Cerata palmate distally, with 4-16 dig-
itiform processes arranged in a horseshoe
shape; the larger digitiform processes may
bear numerous, nodular tuberosities. Cerata
occur in 4—7 pairs, first pair opposite, those
on right side successively more posterior,
190 MCDONALD
such that posterior pairs are alternate. Gen-
eral ground color reddish-brown, varying from
light to dark in different specimens, occasion-
al specimens (especially young individuals)
may be greenish to greenish-brown. Red, red-
brown, and yellow blotches, which are formed
of minute dots, occur on body. Rhinophores
may bear scattered white spots. The convex,
dorsal sides of cerata bear oval patches of
white. T.L.: 15 mm.
Anus on right, between first and second
cerata. Genital aperture on right side, antero-
ventral of first ceras.
Radular formula of specimen examined
47(0.1.1.1.0), MacFarland (1966: 247) re-
ports 50-62(0.1.1.1.0). Rachidian teeth (Fig.
86a) bear 3-5 large denticles on either side
of median cusp. Laterals (Fig. 86b) rather flat,
rectangular plates, each with a single cusp.
Masticatory border of mandibles bears 20-
30 blunt denticulations.
Type-locality: Monterey Bay, California.
Range and habitat: Dillon Beach, Marin
Co., California (Lance, 1961), to Punta
Abreojos, Baja California, Mexico (Lance,
1961). Intertidal to 6 m, usually found in rocky
areas along open coast, frequently in the alga
Gigartina sp. Also found offshore on drifting
masses of the kelp Macrocystis pyrifera.
Hopkinsia MacFarland, 1905
Body basically doridiform, elongate-ellipti-
cal in outline, abruptly rounded anteriorly and
posteriorly, and quite depressed and flat-
tened dorso-ventrally. Dorsum densely cov-
ered with numerous very long, gently tapered
processes. Labial tentacles very broad, form-
ing a veil-like expansion. Rhinophores non-
retractile, perfoliate. Branchial plumes unipin-
nate, arranged in a semi-circle around anus.
Labial disc bears very short, thickened
rods. Radular formula 1.1.0.1.1. Laterals long
and hooked distally. Marginals flattened and
denticulate.
Penis armed with minute hooks.
Type-species: Hopkinsia rosacea Mac-
Farland, 1905.
Hopkinsia rosacea MacFarland, 1905
Hopkinsia rosacea MacFarland, 1905: 54.
Cockerell, 1905: 131. MacFarland, 1906:
149-151, pl. 21, figs. 97-103; pl. 31, figs.
24-25. Baily, 1907: 92. Berry, 1907: 35.
Kelsey, 1907: 41. O'Donoghue, 1926: 222.
O'Donoghue, 1927b: 100-101, pl. 2, figs.
57-59. Hewatt, 1937: 178 ff. Hewatt, 1938:
287. Costello, 1938: 320, 325, 327-330,
tabs. 1, 3, 5, pl. 1, fig. 20. Smith & Gordon,
1948: 180. Strain, 1949: 206-209. Good-
win 8 Fox, 1955: 1086. Marcus, 1961: 29-
30, pl. 6, figs. 103-106. Lance, 1961: 67.
Steinberg, 1963b: 71. Paine, 1963a: 4.
Paine, 1964: 385. MacFarland, 1966: 125-
126, pl. 21, figs. 2-3; pl. 31, figs. 32-36.
Sphon & Lance, 1968: 79. Roller 8 Long,
1969: 427. Lance, 1969: 37. Gosliner &
Williams, 1970: 178. McBeth, 1970: 28.
McBeth, 1971: 158-159. Harris, 1971a: 84.
McBeth, 1972a: 55. Bertsch et al., 1972:
307. Harris, 1973: 239, 240. Abbott, 1974:
364, pl. 17, fig. 4339. Schuler, 1975: 33.
Nybakken, 1978: 135. McDonald 8 Nybak-
ken, 1978: 111. Fuhrman et al., 1979: 292.
Russo, 1979: 44,
Hopkinsea rosacea MacFarland. O'Dono-
ghue, 1927a: 11 (lapsus).
Type-specimens: U. S. Nat. Mus., no.
181275;
Foot broad, truncated anteriorly with a deep
triangular notch, and abruptly rounded pos-
teriorly. Rhinophores bear 18-24 lamellae,
and are rose-pink on shaft, and somewhat
darker pink on clavus. Branchial plumes 7-
14, dark rose-pink. General ground color
everywhere rose-pink, dorsal processes same
color, or occasionally a lighter shade of same
color. T.L.: 20 mm.
Radular formula of specimen examined
22(1.1.0.1.1), MacFarland (1966: 125) and
Marcus (1961: 30) both report 16(1.1.0.1.1).
Laterals (Fig. 50a) flattened, very elongate
triangles, with a small, blunt hook apically.
Marginals (Fig. 50b) small and somewhat tri-
angular, usually with a single small cusp.
Type-locality: Monterey Bay, California.
Range and habitat: Coos Bay, Coos Co.,
Oregon (Steinberg, 1963b), to Ensenada, Baja
California, Mexico (Farmer & Collier, 1963).
Intertidal t0 6 m, usually in rocky areas.
McBeth (1971: 58) reports that it feeds upon
the rose pink bryozoan Eurystomella bilabia-
ta.
Hypselodoris Stimpson, 1855
Body basically doridiform, quite elongate,
somewhat compressed laterally; sides of
body quite high and lateral mantle margins
quite narrow, making body somewhat quad-
rilateral in cross section. Dorsum soft and
smooth, without tubercles or papillae. Labial
tentacles short and cylindro-conical. Rhino-
phores perfoliate, retractile into sheaths.
Branchial plumes retractile, usually unipin-
CALIFORNIA NUDIBRANCHS тэ
nate, arranged in а circle around anus. Most
species are brightly colored.
Labial armature minute rods which are
usually bifid at tip. Radula broad, without
rachidian tooth. Laterals have bifid cusps,
sometimes with denticles below cusp, espe-
cially on teeth nearest margin of radula.
Penis unarmed.
This genus is very closely related to Chro-
modoris Alder 8 Hancock, 1855 (q.v.).
Type-species: Hypselodoris obscura
(Stimpson, 1855).
Hypselodoris californiensis (Bergh, 1879)
Chromodoris californiensis Bergh, 1879b: 72,
109, 112-114. Bergh, 1979c: 128, 165,
168-170. Bergh, 1880a: 274, pl. 6 (14),
figs. 5-15. Bergh, 1880b: 125, pl. 6 (14),
figs. 5-15. Bergh, 1881b: 220. Bergh,
1884b: 649. Orcutt, 1885: 545. Bergh,
1890b: 940. Bergh, 1890c: 168, pl. 3, fig.
14. Bergh, 1891: 141. Bergh, 1892: 1110
(118). Bergh, 1898: 533. Bergh, 1905a: 71.
Bergh, 1905b: 156. Cockerell 8 Eliot, 1905:
36, 37-38. MacFarland, 1906: 129-130.
Kelsey, 1907: 37. Cockerell, 1908: 106. In-
gram, 1935: 48. Costello, 1947: 232. Pru-
vot-Fol, 1951b: 79. MacFarland, 1966: 154,
157-162 ff, pl. 24, figs. 1-3; pl. 34, figs.
12-23. Marcus 8 Marcus, 1967a: 176-178,
text figs. 30-33. Russell, 1968: 140-141.
Roller, 1970a: 371. Schmekel, 1970: 194.
North, 1971: 57. Schuler, 1975: 33. Bloom,
1976: 293, 294. Bertsch & Burn, 1979: 253
ff. (Non) Bergh, 1894: 181-182, pl. 7, figs.
23-28 (=Hypselodoris ghiselini Bertsch,
1978).
Chromodoris calensis Bergh, 1879a: 341.
Bergh, 1879e: 103. Bergh, 1879f: 3. Bergh,
1880a: 274, pl. 6 (14), figs. 5-15. Bergh,
1880b: 125, pl. 6 (14), figs. 5-15. Costello,
1947: 232. MacFarland, 1966: 157. Rus-
sell, 1968: 140-141.
Chromodoris universitatis Cockerell, 1901c:
79. Cockerell, 1902: 19-20. Cockerell 8
Eliot, 1905: 36, 37-38. MacFarland, 1906:
129-130. Kelsey, 1907: 37. Hilton, 1919:
34. Crozier, 1922: 304. Costello, 1947: 232.
Pruvot-Fol, 1951b: 90. MacFarland, 1966:
158, 163.
Chromodoris sp. Guernsey, 1912: 75, figs.
39C-D. O'Donoghue, 1927b: 78.
Glossodoris californiensis (Bergh). O'Dono-
ghue, 1926: 211. O'Donoghue, 1927a: 10.
O'Donoghue, 1927b: 90-91, pl. 2, figs. 38-
42. Cockerell, 1940: 504. Costello, 1947:
232. Pruvot-Fol, 1951b: 89-90. Pruvot-Fol,
1951c: 152. White, 1952: 114. Lance,
1961: 66. Steinberg, 1961: 62. Farmer 8
Collier, 1963: 62. Paine, 1963a: 4, 8. Stein-
berg, 1963b: 69. Lance, 1966: 69, 70, 72.
Farmer, 1967: 341. Sphon & Lance, 1968:
79. North, 1971: 57. Abbott, 1974: 355, fig.
4250.
Glossodoris (Chromodoris) californiensis
(Bergh). Smith & Gordon, 1948: 180.
Glossodoris calensis (Bergh). Pruvot-Fol,
1951b: 89.
Glossodoris universitatis (Cockerell). Pruvot-
Fol, 1951b: 90. Pruvot-Fol, 1951c: 152.
Hypselodoris californiensis (Bergh). Marcus
& Marcus, 1967a: 59, 176-178 (partim),
238, text fig. 30, (non) text fig. 31, (non)
pp. 176-178 (partim) (= Hypselodoris
agassizii (Bergh, 1894)). Roller & Long,
1969: 429. McBeth, 1970: 28. Roller,
1970a: 371. Long, 1970: 19. Farmer, 1971:
19. McBeth, 1971: 158. Sphon, 1971b:
214. Keen, 1971: 823, pl. 20, fig. 1, (non)
fig. 2335 (=H. agassizii). Sphon & Mulliner,
1972: 150. Harris, 1973: 240. Bertsch et
al., 1973: 287. Bertsch, 1973: 109. Marcus
8 Hughes, 1974: 520. Ferreira 8 Bertsch,
197578326, 327. Bertsch; 19/6a: 1158:
Bertsch, 1977: 114, text fig. 3M. Mc-
Donald & Nybakken, 1978: 112. Bertsch,
1978c: 241-244, figs. 27, 29-31, 61-64.
Fuhrman et al., 1979: 292. Bertsch & Burn,
1979: 255. Russo, 1979: 43. Bertsch,
1979b: 46, fig. 3M.
Chromodoris universitates (= Hypselodoris
californiensis). Harris, 1973: 240 (lapsus).
Type-specimens: not listed.
Anterior portion of dorsum widened and
somewhat expanded, forming a broad undu-
lating velum. Foot narrow and elongate,
rounded and bilabiate anteriorly, and tail
rather prolonged and rounded. Posterior
margin of dorsum bears 5-9 prominent,
hemispherical, glandular elevations ventrally,
each whitish with a central pore-like opening.
Rhinophores bear about 17-20 lamellae, cla-
vus bears a vertical ridge both anteriorly and
posteriorly; the low sheath has a smooth
margin. Rhinophores very dark blue. Bran-
chial plumes 9-12, usually unipinnate, very
dark blue. General ground color rich dark ul-
tramarine blue, margins of dorsum and foot
are bright cobalt blue. Dorsum bears numer-
ous round to oblong spots of bright yellow-
orange, frequently in lateral series on either
side of dorsum and more irregularly placed
192 MCDONALD
posterior of branchial plumes. Sides of body
bear about 10 round to oval spots of bright
yellow-orange. T.L.: 40 mm.
Radular formula of specimen examined
75(118.0.118), MacFarland (1966: 159) re-
ports 74(114.0.114), and Bertsch (1978c: 243)
reports 42-92(63-150.0.63-150), while
Bloom (1976: 293) reports the range 82-
92(98-132.0.98-132). Laterals (Fig. 64)
somewhat hooked and shallowly to deeply
bifurcated distally, thus forming 2 cusps; each
tooth bears 5-10 rather tuberculiform denti-
cles below the lower cusp.
Type-locality: Santa Catalina Island, Cali-
fornia.
Range and habitat: Monterey, Monterey
Co., California (MacFarland, 1966), to Bahía
Magdalena, Baja California, Mexico (Bertsch,
1978c) 24°11’N, 10955 W (La Paz), Baja
California, Mexico (Bergh, 1894); throughout
Gulf of California, Mexico. Rare north of Point
Conception, Santa Barbara Co., California.
Intertidal to 31 m, usually in rocky areas of
open coast, occasional on floating docks and
pilings in bays. Frequently found on the
sponge Dysidea amblia upon which it may
feed (McDonald & Nybakken, 1978). McBeth
(1971: 158) reports that it feeds upon the
sponges: Stelletta estrella and Haliclona sp.
Remarks: H. californiensis has been con-
fused in the literature with 2 other species,
Hypselodoris agassizii (Bergh, 1894) and
Hypselodoris ghiselini Bertsch, 1978 (see
Bertsch, 1978c for discussion).
Bertsch (1978c: 241) and Bertsch & Burn
(1979) consider Chromodoris glauca Bergh,
1879 a synonym of Hypselodoris californien-
sis. However, C. glauca was originally de-
scribed by Bergh (1879e: 107) as having a
double white line on the edge of the dorsum
(“Der ziemlich schmale Mantlerand mit ziem-
lich schmaler weisser Doppellinie.’’), and
Bergh (1879c: 168) said that in H. californien-
sis a “fine line seemed to border the margin
of the mantle-edge .. .,’’ while for Hypselo-
doris agassizii Bergh (1894: 182) mentions 2
narrow white lines on the edge of the dorsum
(... “an dem Ruckenrande selbst zwei
schmale, ringsum continuirliche weissliche
linien.''). С. glauca and H. agassizii are both
described as having numerous small yellow
flecks on the dorsum and side of the body,
while H. californiensis has fewer and larger
yellow spots. Bergh (1879e: 106) was appar-
ently not sure of the locality where C. glauca
was collected, as it was listed as: “Hab. М.
pacif. or. (California,’’ and as “Californien.”
Since California and Baja California were fre-
quently not clearly distinguished by early
collectors, C. glauca may well have been
collected in Baja California, where MH.
californiensis and H. agassizii are both known
to occur. Bertsch (1978b: 80) mentions a
similar situation when he rejects Bergh's list-
ing of Puget Sound, Washington as the type-
locality of Chromodoris dalli and proposes a
location south of Puertecitos, Baja California,
Mexico as the type-locality. Since the radula
and internal anatomy of C. glauca are un-
known, only the external coloration is avail-
able for comparing it to H. agassizii and H.
californiensis. From the above descriptions,
it seems certain that C. glauca much more
closely resembles H. agassizii than H. cali-
forniensis.
Laila MacFarland, 1905
Body doridiform, quite depressed dorso-
ventrally, rounded anteriorly and posteriorly.
A flattened subpallial ridge occurs on either
side of anterior end of body, just posterior
and dorsal of labial tentacles. Margin of dor-
sum bears numerous elongate, rather clavate
processes. Labial tentacles short and digiti-
form, grooved distally. Rhinophores retractile
into sheaths, perfoliate. Branchial plumes
non-retractile, tripinnate, arranged in circle
around anus.
Labial armature absent. Radular formula
n.2.1.2.n. Rachidian teeth spurious plates.
First laterals hooked, second laterals quite
large. Marginals smaller and flattened.
Penis armed with small hooks in irregular
longitudinal rows.
Type-species: Laila cockerelli MacFarland,
1905.
Laila cockerelli MacFarland, 1905
Laila cockerelli MacFarland, 1905: 47. Cock-
erell, 1905: 132. Cockerell 8 Eliot, 1905:
43-44. MacFarland, 1906: 134-145, pl. 19,
figs. 45-50; pl. 27, fig. 15. Berry, 1907: 35.
Kelsey, 1907: 41. Guernsey, 1912: 77, fig.
39A. Guernsey, 1913a: 88-92, text figs. 1-
2. Guernsey, 1913b: 137-157, figs. 1-5.
O'Donoghue, 1921: 163-165, pl. 2 (8), figs.
15-17. O'Donoghue, 1922b: 164. O'Don-
oghue & O'Donoghue, 1922: 138-139, pl.
4, fig. 8. O'Donoghue, 1924: 30. O'Dono-
ghue, 1926: 213. O'Donoghue, 1927a: 11.
O'Donoghue, 1927b: 99-100, pl. 2, figs.
54-56. Clayton, 1932: 6. Fraser, 1932: 67.
Hewatt, 1937: 178 ff. Costello, 1938: 320
ff, tabs. 1, 5. Smith & Gordon, 1948: 180.
CALIFORNIA NUDIBRANCHS 193
LaRocque, 1953: 257. Marcus, 1961: 21-
22, pl. 4, figs. 67-70. Lance, 1961: 66.
Paine, 1963a: 4. Steinberg, 1963b: 70.
MacFarland, 1966: 104-106, pl. 20, fig. 4;
pl. 29, fig. 1; pl. 31, figs. 6-12. Lance, 1966:
69. Sphon 8 Lance, 1968: 79. Roller 8
Long, 1969: 427. Lance, 1969: 37. Ro-
billiard, 1969a: 290. Gosliner & Williams,
1970: 178. Hertz, 1970: 6. Long, 1970: 19.
Keen, 1971: 827, pl. 21, fig. 3. Bertsch et
al., 1972: 307. Baker, 1972: 49. Sphon,
1972a: 155. Bertsch & Ferreira, 1974: 346.
Abbott, 1974: 359, pl. 17, fig. 4282. Had-
erlie et al., 1974: tab. 4. Belcik, 1975: 276.
Thompson, 1976a: 51, pl. 7, fig. d. Thomp-
son, 1976b: 77. Michel, 1976: 47, fig. 6.
Nybakken 1978: 135. Poorman 8 Poor-
man, 1978: 373. McDonald & Nybakken,
1978: 110, 112. Bertsch, 1980: 224.
Lialla. Clayton, 1932: 12 (lapsus).
Leilla cockerelli. Marcus, 1966: 58 (lapsus).
Type-specimens: U. S. Nat. Mus., no.
181290.
Dorsal processes usually arranged in rath-
er oblique rows of about 3-4, inner medial
processes generally longer than outer, lateral
processes. Anterior processes may occa-
sionally bear small tubercles distally. Pro-
cesses translucent white to yellowish-white,
tipped with red-orange. Foot linear; truncate,
bilabiate, and slightly emarginate anteriorly,
abruptly pointed posteriorly. Rhinophores
bear 9-14 lamellae, shaft whitish and clavus
red-orange. Rhinophore sheaths have smooth
edges. Branchial plumes 5, whitish, tipped
with red-orange flecks. General ground color
translucent grayish-white to yellowish-white.
In specimens from the northern part of range,
dorsum bears numerous small, randomly dis-
tributed, white tubercles medially; while in
specimens from the southern part of range
dorsum bears a median row of larger tuber-
cles which are tipped with red-orange.
Southern specimens are darker red-orange
than are northern specimens. In Monterey
Bay, California, specimens which bear tuber-
cles intermediate in size and number be-
tween the northern and southern forms are
occasionally found, some of the tubercles are
white and some red-orange. Tail may also be
tipped with red-orange distally. T.L.: 15 mm.
Penis armed with 10-12 slightly irregular,
longitudinal rows of minute, thorn-like hooks.
Radular formula of specimen examined
67(10.2.1.2.10), MacFarland (1966: 105) re-
ports 76-82(10-13.2.1.2.10-13), and Mar-
cus (1961: 21) reports 74-88(10-14.2.1.2.10-
14). Rachidian teeth very rudimentary, small,
flat rectangular, nearly colorless plates. First
laterals (Fig. 45a) long and thin with a distal
hook. Second laterals (Fig. 45b) large, quite
massive, with two distal cusps; below larger
cusp is a rounded prominence; lower part of
shaft bears a blunt projection. Marginals (Fig.
45c, d) quadrate in outline with a rather large,
broad cusp.
Type-locality: Monterey Bay, California.
Range and habitat: Mudge Island, Strait of
Georgia, British Columbia, Canada (O'Dono-
ghue, 1921), to Cabo San Lucas, Baja Cali-
fornia, Mexico (Lance, 1961); Bahía de los
Angeles, Baja California, Mexico (Keen,
1971). Intertidal to 34 m, occasional in rocky,
open coast areas, usually under rocks. Feeds
upon the bryozoan Hincksina velata (Mc-
Donald & Nybakken, 1978: 110).
Remarks: Since the northern and southern
forms, and individuals which are intermediate
between the two, occur in Monterey Bay,
California, it seems fairly certain that they are
a single species and not two species as has
occasionally been suggested, nor geographic
subspecies as suggested by Keen (1971:
827).
Melibe Rang, 1829
Body limaciform, somewhat laterally com-
pressed, and tapered posteriorly. Foot cor-
ners and oral tentacles as such absent. Head
separated from body by narrow neck and ex-
panded into an enormous oral hood. Margin
of hood bears cirri. Cerata spatulate, without
basal gills, and arranged in a longitudinal row
on either side of dorsum.
Radula absent, mandibles absent or rudi-
mentary.
Type-species: Melibe rosea Rang, 1829.
Melibe leonina (Gould, 1852)
Chioraera leonina Gould, 1852: 310-311, pl.
26, fig. 404. Adams 8 Adams, 1854: 71.
Carpenter, 1857: 210, 213, 313. Adams 8
Adams, 1858: 633, pl. 138, fig. 1. Chenu,
1859: 414, fig. 3083. Cooper, 1863b: 60.
Carpenter, 1864: 609. Cooper, 1867: 14.
O'Donoghue, 1921: 192. O'Donoghue,
1922а: 125. O'Donoghue & O'Donoghue,
1922: 134-135, pl. 4, fig. 8. O'Donoghue,
1922b: 165. O'Donoghue, 1922d: 148.
O'Donoghue, 1929: 716. Johnson, 1964:
54, 100. MacFarland, 1966: 280-287, pl.
194 MCDONALD
71, figs. 1-7; pl. 54, figs. 1-10. Schmekel,
1970: 178. North, 1971: 57.
Chioraera leontina Gould. Fewkes, 1889: 141.
Yates, 1890: 41. O'Donoghue, 1922d: 148.
(lapsus).
Melibe pellucida Bergh, 1904: 11-13, pl. 4,
figs. 33-34. O'Donoghue, 1922d: 148.
Odhner, 1936: 1117. Burn, 1960: 70.
MacFarland, 1966: 280.
Melibe leonina (Gould). Bergh, 1875b: 364.
Bergh, 1880d: 161. Fischer, 1887: 534, text
fig. 291. Bergh, 1890b: 882. Bergh, 1892:
1043 (51). Bergh, 1907: 95. Agersborg,
1919: 269. Agersborg, 1921a: 222-253,
text figs. 1-12. Agersborg, 1921b: 50-57.
O’Donoghue, 1922d: 150. Agersborg,
1922b: 439. Agersborg, 1923a: 507-592,
pls. 27-37. Agersborg, 1923b: 86-96, pls.
2-5. Agersborg, 1923c: 135. Agersborg,
1923d: 346. Agersborg, 1924: 216-229,
pls. 6-7. O'Donoghue, 1924: 24-25.
Agersborg, 1925: 167. Guberlet, 1928:
163-170. Fosbert, 1929: 135, text figs. a-
c. Fraser, 1932: 67. Chambers, 1934: 602,
604. Wismer & Swanson, 1935: 341.
Odhner, 1936: 1068, 1113, 1117, text fig.
8. Smith & Gordon, 1948: 181. LaRocque,
1953: 252. Forrest, 1953: 234. Graham,
1955: 153. Comfort, 1957: 226. Burn,
1960: 70. Eyerdam, 1960: 45. Lance, 1961:
67. deVries, 1963: 107 ff. Steinberg, 1963b:
71. Paine, 1963a: 4. Lance, 1966: 69, 78-
79. Hurst, 1967: 255 ff, text figs. 7a—b, pl.
27, fig. 8; fig. 24-16. Hurst, 1968: 151-
166, text figs. 1-8. Sphon & Lance, 1968:
80. Waidhofer, 1969: 295, 296. Roller &
Long, 1969: 427. Long, 1969c: 232. Gos-
liner & Williams, 1970: 179. Farmer, 1970:
78. Bernard, 1970: 85. Ghelardi, 1971: 411,
415. Ajeska, 1971: 13. Keen, 1971: 833,
pl. 21, fig. 6. Baker, 1972: 49. Thompson,
1972a: 75. Harris, 1973: 258 ff. Abbott,
1974: 370, fig. 4378. Birkeland, 1974: 218.
Ajeska & Nybakken, 1976: 19-26, figs. 4,
6-10. Lambert, 1976: 297. Thompson,
1976a: 28, text figs. 12b, 14a. Thompson,
1976b: 58. Poorman & Poorman, 1978:
373. McDonald & Nybakken, 1978: 113.
Bonar, 1978: 191.
Chioraera dalli Heath, 1917: 137-148, pls.
11-13. O'Donoghue, 1922d: 148. Mac-
Farland, 1966: 280 ff.
Melibe lenina. Bovard & Osterud, 1918: 134
(lapsus).
Melibe (Chioraera) leonina (Gould). O'Dono-
ghue, 1922d: 145-150.
Chicoraera leonina. O'Donoghue, 1924: 24
(lapsus).
Melibe dalli (Heath). Odhner 1936: 1117.
Type-specimens: not listed.
Foot narrow, linear; rounded anteriorly and
tapered slightly posteriorly to blunt tail; foot
separated from body by lateral groove. Oral
hood elliptical and arched dorsally. Margin of
hood bears two rows of outwardly directed
cirri, outermost longer, innermost half the
length of outermost series and alternates with
it. Within the previous series is an inwardly
directed series of shorter, more numerous
cirri, arranged at equal intervals. Rhino-
phores perfoliate with 5-6 lamellae; they are
located anteriorly on oral hood, shaft long and
cylindrical and bears a thin, triangular, sail-
like expansion on inner face; clavus retractile
into a campanulate sheath at tip of shaft.
Cerata laterally flattened, broadly ovate to
round in outline, and arise from a short, stout
stalk; the slightly truncate tips bear 2-6 small,
acute points. There are usually 5-6 cerata on
either side of dorsum. First pair is opposite,
and remaining pairs are more or less alter-
nate. They are readily dehiscent; the delicate
network of liver branches in the cerata may
easily be seen. General ground color, includ-
ing cerata, oral hood, and rhinophores is
translucent yellowish-brown, occasionally with
a slight blue-green tinge. Body irregularly
dotted with numerous small, bluish-white
dots. Liver branches usually light green to
brown. T.L.: 100 mm, but specimens up to
300 mm have been collected.
Anus on right, between and slightly ventral
of first and second cerata. Genital aperture
on right, antero-ventral of first ceras. Penis
unarmed.
Type-locality: Port Discovery, Puget Sound,
Washington.
Range and habitat: Dall Island, Alaska
(Heath, 1917), to Punta Abreojos, Baja Cali-
fornia, Mexico (Lance, 1966), and throughout
the Gulf of California, Mexico. Intertidal to 37
m, usually found on the alga Macrocystis
pyrifera, or on the eelgrass Zostera marina,
occasionally found on floating docks in bays.
The large oral hood is used to capture small
crustaceans, especially copepods which dwell
on the blades of Macrocystis. Feeds upon
copepods and amphipods (gammarids and
caprellids) (Ajeska & Nybakken, 1976; Hurst,
1968; MacGinitie & MacGinitie, 1949).
Remarks: This species is capable of swim-
CALIFORNIA NUDIBRANCHS 195
ming, and it releases a rather pungent fruity
odor.
Okenia Menke, 1830, ex Leuckart
in Bronn, Ms.
Body usually limaciform and elongated.
Lateral edge of dorsum bears a few elongate
dorsal processes. Labial tentacles as such are
absent. Rhinophores non-retractile, perfo-
liate (lamellae sometimes few and incom-
plete), and rather large. Branchial plumes uni-
to bipinnate, non-retractile, arranged around
anus.
Labial disc armed with small hooks. Rad-
ular formula 1.1.0.1.1., laterals hook shaped,
with or without denticles. Marginals smaller,
with a small cusp.
Penis armed.
Type-species: Okenia elegans Leuckart,
1828.
See ICZN (1974, Opinion 1014), Okenia
placed on Official List.
Okenia angelensis Lance, 1966
Okenia angelensis Lance, 1966: 76-78, text
figs. 9-12. Burn, 1967b: 55.
Sphon & Lance, 1968: 80. Roller 8 Long,
1969: 427. Keen, 1971: 829, pl. 21, fig. 2.
Abbott, 1974: 363. Schmekel, 1979: 357.
Cargoa angelensis (Lance). Vogel & Schultz,
1970: 389.
Type-specimens: Calif. Acad. Sci., no. 101.
Body somewhat compressed laterally, with
rather high, vertical sides. Dorsum bears a
few cylindrical, elongate, blunt tipped pro-
cesses, 6-9 of which occur dorso-laterally
along pallial ridge on either side; a pair of
processes occurs medially on dorsum be-
tween rhinophores and branchial plumes.
Foot elongate, rather narrow, and truncate
anteriorly; sides nearly parallel, and tail rath-
er blunt. Antero-lateral corners of head pro-
longed into a pair of rather triangular lobes.
Rhinophores long and slightly tapered to blunt
tips, they bear 1-3 incomplete lamellae on
posterior face, and are translucent grayish-
white with minute yellowish to white dots dis-
tally and minute brownish dots proximally.
Branchial plumes 5-7, uni- to bipinnate, ar-
ranged in an irregular semi-circle, they are
translucent grayish-white with a few flecks of
yellow and brown. General ground color
translucent grayish-white, with a few flecks
of yellow to white on dorsum and sides of
body, and with irregular patches of reddish-
brown dots on dorsum. Occasional speci-
mens may bear scatterings of bluish-green on
dorsum. T.L.: 5 mm.
Radular formula of specimen examined
18(1.1.0.1.1), Lance (1966: 77) reports
21(1.1.0.1.1). Laterals (Fig. 40b) hamate and
bear 14-30 denticles on margin of hook.
Marginals (Fig. 40a) oval to subquadrate with
a single, distal hook. .
Type-locality: Bahia de los Angeles, Baja
California, Mexico.
Range and habitat: San Francisco Bay,
California (Lance, 1966), to Mission Bay, San
Diego Co., California (Lance, 1966); Bahia de
los Angeles, Baja California, Mexico (Lance,
1966). Intertidal and subtidal, most common-
ly found on floating docks and pilings in bays.
Not uncommon, but easily overlooked, as it
is small and blends well with the substrate.
Okenia plana Baba, 1960
Okenia sp. Steinberg, 1960: 49.
Okenia plana Baba, 1960a: 80-81, pl. 7, figs.
2a-d. Steinberg, 1963a: 65. Steinberg,
1963b: 71. Burn, 1967b: 55. Gosliner &
Williams, 1970: 179. Abbott, 1974: 363.
Schmekel, 1979: 357.
Cargoa plana (Baba). Vogel & Schultz, 1970:
389.
Type-specimens: Dr. К. Baba’s Labora-
tory, Japan (Baba, personal communication).
Body rather doridiform, quite depressed
dorso-ventrally, rather equally rounded ante-
riorly and posteriorly. Dorsum bears a single
median, elongate, cylindrical, blunt tipped,
dorsal process just anterior of branchial
plumes; it is translucent grayish-white with a
number of irregular, chocolate-brown flecks
on proximal two thirds to three quarters,
some of which are concentrated into an irreg-
ular subapical band below the distal one
quarter to one third which is opaque white.
A row of 5 similarly shaped and colored dor-
sal processes occurs dorso-laterally on either
side, the first just anterior of rhinophores, the
next 3 in a line between rhinophores and
branchial plumes, and the last just posterior
of branchial plumes. Foot elliptical in outline;
head expanded into a semi-circular oral veil.
Rhinophores bear about 9-13 lamellae and
are translucent grayish-white, somewhat yel-
lowish distally, and bear a few irregular choc-
olate brown flecks proximally on shaft. Bran-
chial plumes 8-11, unipinnate, arranged in a
196 MCDONALD
semi-circle, translucent yellowish-white with
a number of irregular chocolate brown flecks.
General ground color translucent grayish-
white. Dorsum bears numerous irregular,
chocolate brown flecks. T.L.: 8 mm.
Radular formula of specimen examined
18(1.1.0.1.1), Baba (1960a: 81) reports
30(1.1.0.1.1). Laterals (Fig. 41a) hooked and
non-denticulate. Marginals (Fig. 41b) smaller,
somewhat quadrate with a rather large den-
ticle below which are 4-5 denticles. Labial
disc bears an incomplete ring of jagged
hooks.
Type-locality: Toba, Japan.
Range and habitat: San Francisco Bay,
California (Steinberg, 1963b); originally de-
scribed from Japan, from where it was prob-
ably introduced into California. Intertidal and
subtidal, usually on floating docks and pilings
in bays.
Onchidoris Blainville, 1816
Body doridiform, dorsum tuberculate or
papillate. Labial tentacles as such are ab-
sent; they are fused into an oral veil. Rhi-
nophores perfoliate and retractile into sheaths
with smooth margins. Branchial plumes re-
tractile and unipinnate.
Labial disc unarmed. Radular formula var-
les among the numerous species, 1.1.0.1.1,
Ул. OFZ hat 2. ete.
Penis unarmed.
Type-species: Onchidoris bilamellata (Lin-
naeus, 1767).
Onchidoris bilamellata (Linnaeus, 1767)
Doris bilamellatus Linnaeus, 1767: 1083.
Tonning, 1768, vide 1769: 484.
Doris fusca Müller, 1776: 229, no. 2768. Mül-
ler, +788: pl. 47, figs. 1-9. Cuvier, 1804:
449. Lamarck, 1819: 312. Blainville, 1819:
452. Rapp, 1827: 520. Bosc, 1830: 110:
Cuvier, 1834: 114. Gray, 1842: pl. 67, fig.
15. Adams & Adams, 1854: 51. Loyning,
1927: 250. Lemche, 1938: 1. (Non) Loven,
1846: 136 (? =Acanthodoris pilosa).
Doris verrucosa Pennant, 1777: 43, pl. 21,
fig. 23. Turton, 1807: 133. Pennant, 1812:
82, pl. 23, fig. 2. Fleming, 1828: 282. Thiele,
1931: 434, fig. 533. (Non) Linnaeus, 1768:
653.
Doris elfortiana Blainville, 1816: 95. Leach,
1852720, pl. 7, fiq..1.
Onchidorus leachii Blainville, 1816: 97.
Doris leachii Blainville, 1819: 450. Rapp,
1827: 520.
Onchidoris leachii Blainville. Blainville, 1825:
489. Blainville, 1827: pl. XLVI, fig. 8. Cu-
vier, 1834: 117.
Doris obvelata Bouchard-Chantereaux, 1835:
42. (Non) Müller, 1776.
Doris bilamellata Linnaeus. Cuvier, 1804: 448,
449. Johnston, 1838a: 54. Johnston,
1838b: pl. 2, fig. 8. Thompson, 1840: 86.
Hassall, 1842: 133. MacGillivray, 1843:
198. Thompson, 1844: 250. Reid, 1846:
377 ff, pl. 10, figs. 1-13, 15-21, 25. Alder,
1850: 105, 110. Forbes 8 Hanley, 1851:
567. Alder & Hancock, 1851c: fam. 1, pl.
1, fig. 13. Hancock & Embleton, 1852: 208
ff, pl. 11, fig. 2; pl. 12, fig: 5; ЕЛА ООВ:
pl. 17, figs. 6-7. Gosse, 1853: 12, 13-14,
62, 83, 232. Gosse, 1854: 107. Adams &
Adams, 1854: 51. Byerley, 1854: 44. Alder
& Hancock, 1855: 20, 31, 32, 43. Gosse,
1856: 105. Thompson, 1856: 272. Colling-
wood, 1859: 463, 470. E. Wright, 1859: 88.
Collingwood, 1860: 202. Norman, 1860:
7242. Collingwood, 1861: 114. Colling-
wood 8 Byerley, 1862: 189. Mcintosh,
1865: 390. Robertson, 1868: 205. Jef-
freys, 1869: 90. Gould, 1870: 228-229, pl.
21, figs. 305-309. Dall, 1870: 249. Verkru-
zen, 1872: 375. Sauvage, 1873: 25, 29.
Mcintosh, 1874: 432. McIntosh, 1875: 89.
Friele & Hansen, 1876: 71. Jeffreys, 1877:
337. Leslie & Herdman, 1881: 312. Locard,
1886: 28. Hertzenstein, 1885: 709. Hig-
gins, 1886: 25. Herdman, 1890b: 201.
Bergh, 1891: 103. Herdman & Clubb, 1892:
153, 159, 162. Lundbeck, 1893: 175.
Cooke, 1899: 64. Colgan, 1908: 106, 111.
Eliot, 1910: 16. Pelseneer, 1911: 53-55,
pl. 15, figs. 5-23. Chumley, 1918: 87, 169.
Pelseneer, 1923: 28-32, text figs. 1-4.
Loyning, 1927: 250. White, 1938: 14.
Lemche, 1938: 1.
Doris affinis Thompson, 1840: 85-86.
Thompson, 1856: 272. (Non) Gmelin in Lin-
naeus, 1791: 3106.
Doris liturata Beck in Möller, 1842: 78. Mörch,
1857: 6. Stimpson, 1862: 4. Bergh, 1878b:
607.
Doris vulgaris Leach, 1847: 268. Leach, 1852:
19.
Doris coronata Agassiz, 1850: 191. Gould,
1870: pl. 20, figs. 285, 286; pl. 21, figs.
297, 299.
Oncidoris (doris) bilamellata (Linnaeus). Hogg,
1868: pl. 10, fig. 43 (lapsus).
Lamellidoris n. sp. vel var. praecedentis ?
Morch, 1868: 204.
Lamellidoris bilamellata (Linnaeus). Mörch,
CALIFORNIA NUDIBRANCHS 197
1868: 203. Abraham, 1877: 198. Bergh,
1878b: 605, 606-609, pl. 64, fig. 13; pl.
64, figs. 1-5. Sars, 1878: 306-307 ff, pl.
XIII, fig. 5. Bergh, 1879a: 364. Bergh,
1880a: 210. Bergh, 1880b: 61. Herdman
1886: 268, 277. Fischer, 1887: 524, text
fig. 286. Garstang, 1889: 179. Garstang,
1890: 447. Bergh, 1890b: 985. Norman,
1890: 73. Bergh, 1892: 1152 (160). Herd-
man & Clubb, 1892: 132 ff. Bergh, 1894:
192. Garstang, 1894: 226. Bergh, 1894:
173. Herdman et al., 1896: 446. Tregelles,
1896: 220. Cooke, 1899: 66. Beaumont,
1900: 850. Nichols, 1900: 597. Allen &
Todd, 1900: 181, 212. Vayssiere, 1901:
297. Conchol. Soc., 1901: 26. Knight, 1901:
207. Johansen, 1902: 387. Mar. Biol. As-
soc., 1904: 284. Farran, 1904: 3. Cockerell
8 Eliot, 1905: 32. Bergh, 1905a: 101. Eliot,
1906c: 346. Woodland, 1907: 45, pl. 5, fig.
6. Odhner, 1907: 74. Walton, 1908: 239.
Elmhirst, 1908: 228. Farran, 1909: 17-18.
Eliot, 1910: 13, 28, 156, 346. Colgan, 1911:
25. Walton, 1913: 110. Farran, 1915: 10,
67. Johnson, 1915: 175. Renouf, 1916: 12-
15. Evans & Evans, 1917: 109. Bardarson,
1919: 73. Bardarson, 1920: 108. O'Dono-
ghue, 1921: 174-176, pl. 2 (8), fig. 21.
O’Donoghue, 1922a: 126-129, pl. 2.
O'Donoghue 8 O'Donoghue, 1922: 139, pl.
4, fig. 6. Odhner, 1922: 24-25. O'Dono-
ghue, 1924: 24. O'Donoghue, 1926: 221.
O'Donoghue, 1927a: 3, 10. Jutting, 1927:
LXXXVIII. Derjugin, 1928: 320. Fraser,
1932: 67. White, 1938: 19. Volodchenko,
1941: 60. Stock, 1952: 58. Pruvot-Fol,
1954: 296-297, figs. 116a-f. Swennen,
1959: 56, 58. Swennen, 1961: 197-199.
Zenkevitch, 1963: 112. Hadfield, 1963: 91.
Daro, 1969: 136 ff. Daro, 1970: 168. Voogt,
1973: 479 ff.
Lamellidoris liturata (Beck in Möller). Mörch,
1875: 125. Mörch, 1877: 436.
Lamellidoris bilamellata var. liturata Beck.
Bergh, 1878b: 609-613, pl. 64, figs. 14-
19; pl. 65, figs. 6-13.
Lamellidoris bilamellata var. pacifica Bergh,
1880a: 211-216, pl. 5, fig. 10; pl. 11, figs.
3-9. Bergh, 1880b: 62-67, pl. 5, fig. 10;
pl. 11, figs. 3-9. Bergh, 1894: 192-193.
Bergh, 1905a: 101.
Lamellidoris billamellata var. pacifica. Dall,
1884: 341 (lapsus).
Doris (Lamellidoris) bilamellata Linnaeus.
Vanoffen, 1897: 187, 193.
Onchidorus fusca (Müller). Iredale 8 O'Don-
oghue, 1923: 220. Mar. Biol. Assoc., 1931:
273. Johnson, 1934: 156. Renouf, 1934:
400. LaRocque, 1953: 254.
Onchidoris fusca (Müller). O'Donoghue, 1924:
24. Winckworth, 1932: 234. Steven, 1938:
60, 62. White, 1938: 14, 19. McMillan,
1944: 162. Barnes & Powell, 1951: 381.
Forrest, 1953: 233, text fig. 5d. Barnes &
Powell, 1954: 361-363, pl. 2. Williams,
1954: 106. Marcus, 1955: 129. Graham,
1955: 152. Mar. Biol. Assoc., 1957: 310.
Thompson, 1959: 240. Thompson, 1960b:
126-127. Miller, 1961: 111, 112. Thomp-
son, 1961: 234, 236, text fig. 1A. Tarasov,
1961: 8, 25, 54. Miller, 1962: 553-554, fig.
7. Roginskaya, 1962a: 88, 92, 105, fig. 1.3.
Hadfield, 1963: 88-93. Bruce et al., 1963:
203. Thompson, 1964: 279 ff. Miller, 1967:
9. Thompson, 1967: 9, 11, text fig. 7E. Ed-
munds, 1968b: 122 ff. Barrett, 1969: 69.
Hughes, 1970a: 609 ff. Hughes, 1970b: 81,
82, text figs. 15, 16. Potts, 1970: 269-292,
pls. 1-2, text figs. 1-8. Meyer, 1971: 139-
140. Holleman, 1972b: 142. Harris, 1973:
217 ff. Roginskaya, 1976: 23-26. Perron
& Turner, 1977: 182, 183. Eyerdam, 1977:
110. Chia & Koss, 1978: 118. Nybakken,
1978: 129, 143. Dehnel & Kong, 1979:
1843.
Onchidorus bilamellatus (Linnaeus). Odhner,
1926b: 26.
Onchidorus bilamellata (Linnaeus). Loyning,
1927: 262. Lemche, 1929: 17.
Onchidorus fuscus (Müller). O'Donoghue,
1927a: 3, 9-10.
Onchidoris bilamellata (Linnaeus). Lemche,
1938: 19-20 ff. Odhner, 1939: 40-41.
Lemche, 1941b: 20-22. Marcus, 1958: 27-
28, text figs. 44-45. Marcus, 1961: 27-28,
57, pl. 5, figs. 92-96. Steinberg, 1963b: 70.
Hurst, 1967:255 ff, pl. 28, fig. 9; pl. 37, fig.
45; fig. 24-15. Burn, 1968: 91. Robilliard,
1969a: 290. Roller, 1970a: 482. Mc-
Donald, 1970: 375. Connell, 1970: 54 ff.
Gosliner & Williams, 1970: 179. Bernard,
1970: 85. ¡Franz! 19702 172.1. Dayton;
1971: 383. Thompson & Bebbington, 1973:
148, pl. 8, fig. b. Birkeland, 1974: 218.
Rozsa, 1974: 7, 8. Clark, 1975: 45. Franz,
1975a: 81, 83. Belcik, 1975: 276. Lambert,
1976: 297. Thompson & Brown, 1976: 96,
fig. 49. Thompson, 1976a: 33, 36, 93, text
fig. 41e. Thompson, 1976b: 66 ff. Garlo,
1977: 23 ff. Crampton, 1977: 48 ff, text
figs. 1-17. Bleakney 8 Saunders, 1978:
82-85. McDonald 8 Nybakken, 1978: 111.
Mumaw, 1978: 74. Clark 8 Goetzfried,
198 MCDONALD
1978: 290. Todd, 1979a: 59. Todd, 1979b:
213.8. Todd, 19/9c::65:Tf.
Lamellidoris fusca (Müller). Baba, 1957: 9.
Abbott, 1974: 362.
Lamellidoris (Lamellidoris) fusca (Müller).
Baba, 1957: 9, 11, text fig. 2B.
Onchidoris (Lamellidoris) fusca (Múller). Had-
field, 1963: 86.
Type-specimens: not listed.
Body equally rounded anteriorly and pos-
teriorly, dorsum covered with numerous large,
round tubercles, which decrease in size near
margin. Foot rather broad, rounded to almost
truncate anteriorly, and somewhat tapered
posteriorly to rather bluntly rounded tail.
Crescent-shaped oral veil extends laterally
nearly to edge of dorsum. Rhinophores bear
14-20 lamellae and are encrusted with
brown except at translucent grayish-white
tips. Rhinophore sheaths low, with smooth
margins. Branchial plumes 16-32 or more,
unipinnate, arranged in 2 semi-circles just an-
terior of anus; they are encrusted with dark
brown. General ground color translucent
brownish-white, dorsum encrusted with irreg-
ular blotches of light rusty brown to dark
chocolate brown, occasionally in the form of
2-3 irregular longitudinal stripes. T.L.: 15 mm.
Radular formula of specimen examined
21(1.1.1.1.1), Marcus (1961: 27) reports 24-
30(1.1.1.1.1). Rachidian teeth (Fig. 35c) nar-
row, roughly rectangular plates which lack a
cusp or denticles. Laterals (Fig. 35b) large,
each with a long, tapered cusp. Marginals
(Fig. 35a) each bear a small cusp.
Type-locality: Oceano Norvegici.
Range and habitat: Kiska Island, Aleutian
Islands, Alaska (Bergh, 1894), to Morro Bay,
San Luis Obispo Co., California (McDonald,
1970); Japan (Baba, 1957), Greenland (Pru-
vot-Fol, 1954), Europe (Pruvot-Fol, 1954), At-
lantic coast of North America (Franz, 1975a).
Intertidal to 250 m, sporadically abundant on
pilings and floating docks in bays, usually in
association with the barnacles Balanus spp.
Feeds upon the barnacles: Balanus bala-
noides, Balanus crenatus, Balanus porcatus,
Elminius modestus, and Verruca stroemia
(Barnes & Powell, 1954; Barrett, 1969;
Bleakney & Saunders, 1978; Bruce et al.,
1963; Clark, 1975; McDonald 8 Nybakken,
1978; Meyer, 1971; Miller, 1961; Potts, 1970;
Swennen, 1961; Thompson 8 Brown, 1976;
Todd, 1979b). Juveniles have also been re-
ported to feed upon the bryozoans: Crypto-
sula and Umbonula (Thompson 8 Brown,
1976). Allan Fukuyama presented for identi-
fication two specimens of this species from
the stomach of a female oldsquaw duck
(Clangula hyemalis) collected at Homer, Alas-
ka; and 14 specimens from a Steller's eider
(Polysticta steller) collected at Nelson La-
goon, Alaska.
Remarks: Edmunds (1968b: 123) reports
that this species secretes acid of pH 1.
Lemche (1938: 20) discusses reasons for
maintaining the name O. bilamellata in pref-
erence to O. fusca.
Onchidoris hystricina (Bergh, 1878)
Lamellidoris hystricina Bergh, 1878b: 605,
614, pl. 68, figs. 17-23. Bergh, 1879a: 365.
Bergh, 1880a: 219-221. Bergh, 1880b: 70-
72. Bergh, 1890b: 985. Bergh, 1892: 1153
(161). Abbott, 1974: 362.
Onchidorus hystericina (Bergh). O'Dono-
ghue, 1926: 221 (lapsus).
Onchidoris hystricina (Bergh). Marcus, 1961:
28, 57, pl. 5, figs. 89-91. Paine, 1963a: 4.
Steinberg, 1963b: 71. Farmer, 1967: 342.
Sphon 8 Lance, 1968: 80. Roller & Long,
1969: 427. Gosliner & Williams, 1970: 179.
Keen, 1971: 838. Eyerdam, 1977: 110. Ny-
bakken, 1978: 135.
Onchidoris hystericina (Bergh). 1970: 85
(lapsus).
Type-specimens: not listed.
Body has rather parallel sides, and is bluntly
rounded anteriorly, and more acutely round-
ed posteriorly; dorsum thickly set with nu-
merous long, slender, white to yellowish-white
papillae which give animal a fuzzy appear-
ance. Foot truncate and bilabiate anteriorly
and tapered posteriorly to rounded tail. Oral
veil rather crescent-shaped. Rhinophores bear
7-20 lamellae, and are yellowish-white.
Branchial plumes 9-12, unipinnate, yellow-
ish-white, arranged in a circle around anus.
General ground color yellowish-white to
cream, dorsum and dorsal surface of tail may
bear small flecks of opaque white. T.L.: 8 mm.
Radular formula of specimens examined
27-32(1.1.1.1.1), Marcus (1961: 28) reports
32-34(1.1.1.1.1). Rachidian teeth (Fig. 36c)
narrow, roughly rectangular plates which lack
cusp or denticles. Laterals (Fig. 36b) large,
each bearing a cusp which bears 12-15 den-
ticles, Marcus (1961: 28) reports 6-11 den-
ticles on cusp. Marginals (Fig. 36a) some-
what quadrate and each bears a short cusp.
CALIFORNIA NUDIBRANCHS 199
Type-locality: Kiska Island, Aleutian Is-
lands, Alaska.
Range and habitat: Kiska Island, Aleutian
Islands, Alaska (Bergh, 1880a) to 12.9 km
south of Cabo Colonet, Baja California, Mex-
ico (Farmer, 1967). Intertidal to 30 m, spo-
radically abundant in rocky intertidal; occa-
sional on floating docks and pilings in bays.
May feed upon the bryozoan Tubulipora sp.
Remarks: The specimens which | have ex-
amined are in some ways intermediate be-
tween Onchidoris hystricina (Bergh) and
Onchidoris varians (Bergh, 1878). Bergh
(1880a: 217, 221) mentions 6-8 denticles on
the laterals of the former and about 20 for
the latter species, while the specimens | ex-
amined had 12-15 denticles on the laterals.
Bergh (1880a: 216) describes O. varians as
very similar to O. hystricina, the only real dif-
ferences being the number of denticles on the
laterals, and O. hystricina bears numerous
spicules in the dorsum, while O. varians has
very few spicules. Since the specimens which
| examined had numerous spicules in the dor-
sum, | feel that they more closely resemble
O. hystricina than O. varians. Further study
of these two species may well show them to
be conspecific.
Onchidoris sp.
Onchidoris spec. Roller 8 Long, 1969: 427.
Onchidoris muricata (Müller). Roller, 1970b:
482. McDonald, 1975a: 531, 540. Mc-
Donald 8 Nybakken, 1978: 111 (partim),
116. Nybakken, 1978: 135.
Body equally rounded anteriorly and pos-
teriorly, dorsum covered with numerous tu-
bercles which are somewhat constricted ba-
sally and bulbous apically, appearing rather
spherical. Foot rather broad, rounded, al-
most truncated anteriorly, broadly rounded
posteriorly. Oral veil crescent-shaped ex-
tending laterally nearly to edge of dorsum.
Rhinophores perfoliate with 9-15 lamellae,
lemon yellow; rhinophore sheaths with
smooth margins. Branchial plumes 10-13,
unipinnate, light to dark yellow, arranged in
incomplete circle around anus. General
ground color lemon yellow, tubercles slightly
darker than dorsum. T.L.: 5 mm.
Radular formula of specimens examined
34-39(5-6.1.0.1.5-6). Each lateral (Fig. 37f)
bears a long, curved hook with 11-21 small
denticles. Marginals (Fig. 37a-e) each have a
single cusp.
Range and habitat: Friday Harbor, Wash-
ington (Roller, 1970b), to Lion Rock, San Luis
Obispo Co., California (Roller, 1970b). Inter-
tidal to 10 m. Usually found upon the en-
crusting bryozoan Reginella mucronata which
it matches closely in color and upon which it
probably feeds (McDonald & Nybakken,
1978).
Remarks: Externally this species closely
resembles Adalaria proxima (Alder & Han-
cock, 1854) or Onchidoris muricata (Müller,
1776), but the denticulate laterals exclude it
from A. proxima and the lack of a rachidian
excludes it from both of the above. Further
study is needed to determine whether or not
it is a new species.
Phidiana Gray, 1850
Body aeolidiform, gradually tapered pos-
teriorly to the pointed tail. Anterior foot cor-
ners usually produced into elongate, tenta-
culiform processes, but may be rounded.
Cerata cylindrical and slightly clavate; ar-
ranged in regular rows. Rhinophores non-re-
tractile, may be smooth, papillate, verrucose,
annulate, or perfoliate.
Masticatory border of mandibles denticu-
late. Radula uniseriate, with denticles on
either side of median cusp.
Penis may be armed with a stylet or spines
or may be unarmed. Anus cleioproct.
Miller (1974) united a number of similar
genera (e.g. Hermissenda Bergh, 1878, Fa-
celina Alder & Hancock, 1855, Emarcusia
Roller, 1972, et al.) under the name Phidiana,
stating that the characters which had been
used to separate these genera overlap and
do not seem sufficient to distinguish separate
genera. | agree with Miller's definition of Phi-
diana.
Type-species: Phidiana patagonica (d’Or-
bigny, 1837).
Phidiana crassicornis (Eschscholtz, 1831)
Cavolina crassicornis Eschscholtz, 1831: 15,
fig. 1. Carpenter, 1857: 173. Trinchese,
1881: 31. O'Donoghue, 1922c: 76. O'Don-
oghue, 1922d: 133-135.
Facelina crassicornis (Eschscholtz). Gray,
1857: 224. Adams 8 Adams, 1858: 633.
Flabellina crassicornis (Eschscholtz). Car-
penter, 1857: 313.
Aeolis (Flabellina?) opalescens Cooper,
1863a: 205. O'Donoghue, 1922c: 75, 76.
Flabellina opalescens (Cooper). Cooper,
200 MCDONALD
1863b: 60. Carpenter, 1864: 609. Cooper,
1867: 14. Bergh, 1975a: 649. O'Dono-
ghue, 1922c: 75, 76. O'Donoghue, 1922d:
134.
Aeolis (? Flabellina) opalescens Cooper. Car-
penter, 1864: 608.
Eolis opalescens (Cooper). Dall, 1871: 137.
Hermissenda opalescens (Cooper). Bergh,
1879b: 81-85, pl. 1, figs. 9-12; pl. 2, figs.
1-6. Bergh, 1879c: 138-141, pl. 1, figs. 9-
12; pl. 2, figs. 1-6. Bergh, 1879d: 573.
Bergh, 1890a: 37. Bergh, 1892: 1030 (38).
Cockerell, 1901a: 86. Cockerell, 1901b:
122. Cockerell & Eliot, 1905: 50-51.
Guernsey, 1912: 78, fig. 39J. O'Donoghue,
1921: 201-204, pl. 3 (9), fig. 32. O'Dono-
ghue, 1922a: 125-126. O'Donoghue &
O'Donoghue, 1922: 136-137, pl. 3, fig. 3.
O'Donoghue, 1922b: 165. O'Donoghue,
1922c: 75-76. O'Donoghue, 1922d: 134.
Agersborg, 1922b: 425. Agersborg, 1923c:
134. O'Donoghue, 1924: 26, 31. Agers-
borg, 1925: 167. Costello, 1938: 321 ff,
tabs. 1, 3.5: pl. 1, fig: 19; pl: 2, figs: 33-
34, 42-43.
Aeolidia (Hermissenda) opalescens (Cooper).
Bergh, 1880a: 232. Bergh, 1880b: 83.
Facelina (Cavolina) crassicornis Eschscholtz.
Trinchese, 1881: 31.
Aeolis opalescens Cooper. Fischer, 1887:
559. Kelsey, 1907: 33.
Hermissenda crassicornis (Eschscholtz).
O'Donoghue, 1922a: 125-126. O'Dono-
ghue, 1922c: 76. O'Donoghue, 1922d:
133-135. Agersborg, 1923e: 349. O'Don-
oghue, 1924: 26, 31. Agersborg, 1925:
167-180, text figs. 1-22. O'Donoghue,
1926: 233. O'Donoghue, 1927b: 107-108,
pl. 3, figs. 74-76. Fraser, 1932: 67. In-
gram, 1935: 48, 49. MacGinitie, 1935: 740.
Hewatt, 1937: 178 ff. Volodchenko, 1941:
60. Worley & Worley, 1943: 367, pl. 3, figs.
7, 9-12. Hewatt, 1946: 191, 198. Smith 4
Gordon, 1948: 181. Pruvot-Fol, 1951a: 59,
fig. 36. LaRocque, 1953: 248. Marcus,
1961: 52-54, pl. 10, figs. 188-192. Lance,
1961: 68. Kohn, 1961: 292 ff. Lance,
1962c: 52. McLean, 1962: 111. Farmer 8
Collier, 1963: 63. Steinberg, 1963b: 72.
Paine, 1963a: 1, 4, pl. 1. Barth, 1964: 312.
Paine, 1964: 385. Farmer, 1964: 24. Bur-
gin, 1965: 205, 215, text figs. 1-9. Ghise-
lin, 1965: 336 ff. Paine, 1965: 604 ff.
MacFarland, 1966: 358-365, 368, pl. 55,
fig. 1; pl. 70, figs. 13-14; pl. 71, figs. 1-
14. Lance, 1966: 69, 79-80. Dennis, 1967a:
1441 ff. Dennis, 1967b: 259. Hurst, 1967:
255, text fig. 4a, pl. 31, fig. 22; fig. 24-3.
Eakin et al., 1967: 349 ff. Marcus 8 Mar-
cus, 1967a: 226-227, 238. Buchsbaum 4
Milne, 1967: pl. 64 (non pl. 62 =Antiopella
barbarensis). Mauzey et al., 1968: 607.
Beeman, 1968a: 96. Beeman, 1968b: 268.
Sphon & Lance, 1968: 79. Haderlie, 1968:
333 ff. DuShane 8 Sphon, 1968: 244.
Turner et al., 1969: 136, text fig. 52, ap-
pend. 1, 2. Haderlie, 1969: tabs. 1, 2. Roll-
er & Long, 1969: 427. Stensaas et al.,
1969: 510 ff. Robilliard, 1969a: 290. Wil-
liams & Gosliner, 1970: 33. Gosliner 8 Wil-
liams, 1970: 178. Grigg 8 Kiwala, 1970:
149, 151. Bernard, 1970: 85. Hughes,
1970b: 80. Michel, 1970: 7. Schmekel,
1970: 154. Fager, 1971: 243, 246. Robil-
liard, 1971c: 429. Crane, 1971: 57. Farmer,
1971: 19. Harris, 1971a: 80, 84-85, 88.
North, 1971: 57. Bertsch et al., 1972: 306-
307. Mulliner, 1972b: 2, fig. Holleman,
1972а: 60. Alkon & Fuortes, 1972: 631.
Baker, 1972: 45, 47. Blair & Seapy, 1972:
121 ff. Harris, 1973: 226 ff. Thompson,
1973: 167 ff, text fig. 9. Thompson 8 Beb-
bington, 1973: 148, 149, pl. 12, fig. c. God-
dard, 1973: 9. Alkon, 1973a: 444 ff. Alkon,
1973b: 185, 197. Alkon 8 Bak, 1973: 620-
634. Detwiler & Alkon, 1973: 618 ff. Wil-
lows, 1973b: 205. Gosliner & Williams,
1973b: 352. Nybakken, 1974: 371. Abbott,
1974: 379, fig. 4463. Miller, 1974: 43. Had-
erlie et al., 1974: tab. 4. Alkon, 1974a:
1083. Alkon, 1974b: 70, text figs. 1-2.
Birkeland, 1974: 211 ff. Chase, 1974a: 707.
Mariscal, 1974: 163. Murray 8 Lewis, 1974:
156. Zack, 1975a: 271-275. Zack, 1975b:
238 ff, text fig. 1. Belcik, 1975: 276. Alkon,
1975a: 46. Alkon, 1975c: 507. Detwiler 8
Fuortes, 1975: 107, 108. Harris, 1975: 539.
Masukawa, 1975: 359. Schuler, 1975: 33.
Alkon, 1976b: 341. Alkon, 1976c: 410 ff.
Detwiler, 1976: 691. Michel, 1976: 49, fig.
15. Anderson, 1976: 407 ff. Akaike 8 Al-
kon, 1977: 171. Defelice & Alkon, 1977b:
613. Alkon & Grossman, 1977: 171. Crow
& Alkon, 1978b: 191. Cooper, 1978: 8.
Schmidt et а/., 1978: 136a. McDonald &
Nybakken, 1978: 115. Bonar, 1978: 187.
Harrigan 8 Alkon, 1978a: 430 ff. Harrigan
& Alkon, 1978b: 299. Heldman 8 Alkon,
1978: 117 ff. Williams, 1978: 58. Crow &
Alkon, 1978a: 1239. Alkon & Grossman,
1978b: 1329 ff. Alkon, 1979b: 810-816.
Crow & Harrigan, 1979: 179 ff. Harrigan et
al., 1979: 1001. Grossman et al., 1979: 24
ff. Oakes, 1979: 28-29. Harris 8 Howe,
CALIFORNIA NUDIBRANCHS 201
1979: 145. Heldman et al., 1979: 153 ff.
Baba, 1979a: 17. Baba, 1979b: 7. Russo,
1979: 44, 46, 48. Crow et al., 1979: 181-
195. Susswein & Bennett, 1979: 523 ff.
Akaike & Alkon, 1980: 501 ff. Williams,
1980: 99 ff, text figs. 1a, 2-7. Crow & Al-
kon, 1980b: 412 ff. Stommell et al., 1980:
2104. Neary, 1980: 2166. Crow & Alkon,
1980a: 596. Lederhendler et al., 1980: 218
ff. Cockburn & Reid, 1980: 275 ff. Hodge
& Adelman, 1980: 220 ff.
Hermisisenda crassicornis (Eschscholtz).
O'Donoghue, 1927a: 11 (lapsus).
Cuthona (Hervia) emurai Baba, 1937b: 329-
331, text fig. 16A-D. Baba, 1937a: 199.
Marcus, 1958: 62. Baba 8 Hamatani, 1965:
109. Baba, 1979a: 17. Baba, 1979b: 7.
Hermissenda (Cavolina) crassicornis Esch-
scholtz. Pruvot-Fol, 1951a: 58.
Hervia emurai (Baba). Baba, 1957: 9.
Dondice emurai (Baba). Abe, 1964: 70, 89,
235, fig 125.
Shinanoeolis emurai (Baba). Baba & Hama-
tani, 1965: 109.
Hermissenda crassicornis (Eschscholtz in
Rathke). Keen, 1971: 840, pl. 2, fig. 6.
Hermissenda crassicornis (Eschscholtz).
Oakes, 1979: 27. Jerussi & Alkon, 1980:
596 (lapsus).
Phidiana crassicornus (Eschscholtz). Cooper,
1980: 284.
Type-specimens: not listed.
Foot rather broad, thickened, bilabiate, and
broadest anteriorly, tapered posteriorly to tail.
Margins of foot are thin and extend laterally
beyond body. Oral tentacles translucent
grayish-white. Rhinophores weakly perfo-
liate, with 8-24 lamellae, shaft translucent
grayish-white, clavus opalescent white to very
pale blue. Cerata arranged in 5-11 groups
dorsolaterally on either side of dorsum; first
2 groups separated medially by cardiac re-
gion, more posterior groups less well sepa-
rated medially. General ground color translu-
cent grayish-white. A median band of brilliant
cadmium orange occurs between oral tenta-
cles and passes between rhinophores, where
it is narrowest, to anterior of cardiac region.
An opalescent blue line occurs dorsally on
either oral tentacle, becoming broader and bi-
furcating proximally. Inner portion of blue line
extends between rhinophores, on either side
of orange line, and continues dorso-medially
to tip of tail. Outer portion of these blue lines
extends dorso-laterally to tip of tail, and is
interrupted by each group of cerata. Just
ventral of this last blue line is an orange line
on either side of head, extending from bases
of oral tentacles to first group of cerata. A
narrow, opalescent blue line extends poste-
riorly along dorsal margin of foot, from foot
corners to tip of tail. Cerata tipped with white
and each bears a subterminal band of cad-
mium orange which becomes lighter below.
Occasionally specimens may bear a bluish-
white band longitudinally on anterior surface
of each ceras. Cerata cores usually light burnt
umber to deep brown; occasionally speci-
mens may be found which have greenish-gray
cores. | have found these specimens on mud-
flats in pools with phoronids and presume
they feed upon the phoronid lophophores.
E25 mim:
Anus on right, dorso-laterally, between
second and third groups of cerata. Genital
aperture on right side, ventral of first group
of cerata. Penis unarmed.
Radular formula of specimen examined
25(0.0.1.0.0), MacFarland (1966: 361) re-
ports 22-25(0.0.1.0.0), and Marcus (1961: 53)
reports 28(0.0.1.0.0). Rachidian teeth (Fig.
108a) bear 3-6 rather long denticles on either
side of large median cusp which bears 6-15
minute, irregular serrulations on distal ventral
surface (Fig. 108b). Masticatory border of
mandibles has about 50 denticles.
Type-locality: Sitka, Alaska.
Range and habitat: Sitka, Alaska (Bergh,
1879b), to Punta Eugenia, Baja California,
Mexico (Lance, 1961); Isla Angel de la Guar-
da, Gulf of California, Mexico (Farmer & Col-
lier, 1963); Japan (Baba, 1937b); common in
northern Gulf of California, Mexico (Keen,
1971). Intertidal to 37 m. Common in rocky
intertidal and on floating docks and pilings in
bays, occasional on mudflats in bays. Feeds
upon various hydroids such as Obelia spp.
(Harris, 1973). Harris (1971a) reports that it
also eats small crustaceans and other nudi-
branchs. Birkeland (1974) reports that it feeds
upon the pennatulacean Ptilosarcus gurneyi,
the ascidian Aplidium californicum, and
stranded specimens of the scyphozoans
Phacellophora camtschatica and Cyanea
capillata. Harrigan & Alkon (1978) raised
specimens in the laboratory on Ciona intes-
tinalis, mantle muscles of Loligo pealii, and
Mytilus edulis. Cooper (1978) found that it
eats the polyps and gonophores of the hy-
droid Tubularia crocea. Oakes (1979) fed it
the hydroids Aglaophenia struthionides and
Sertularia furcata in the laboratory. Also oc-
casionally cannibalistic in captivity. This is
202 MCDONALD
probably the most common aeolid in Califor-
nia.
Remarks: The stomach of a buffalo sculpin
(Enophrys bison) which was captured near
the P. G. & E. outfall at Moss Landing, Mon-
terey Co., California, contained numerous
specimens of P. crassicornis, along with two
specimens of the sea anemone Anthopleura
elegantissima.
Additional references which mention Her-
missenda and are certainly referable to P.
crassicornis are: Alkon, 1975b; 1976a;
1979a; Alkon & Grossman, 1978a; Alkon et
al., 1978; Budelmann, 1976; DeFelice & Al-
kon, 1977a; Fuortes & O'Bryan, 1972; Ken-
nedy, 1967; Mellon, 1974; Zack, 1974a;
1974b.
Dr. K. Baba (1979 and personal communi-
cation) agrees that the form which has a
bluish-white band longitudinally on anterior
surface of each ceras is identical with Shi-
nanoeolis emurai (Baba, 1937), the latter
therefore being a junior subjective synonym
of P. crassicornis.
Phidiana hiltoni (O'Donoghue, 1927)
Facelina hiltoni O'Donoghue, 1927b: 104-
105, pl. 3, figs. 70-71. Marcus, 1958: 59.
Steinberg, 1961: 62.
Phidiana sp. Lance, 1961: 68.
Phidiana pugnax Lance, 1962a: 157-159, pl.
38, text figs. 4-8. Paine, 1963a: 4. Farmer
8 Collier, 1963: 63. Steinberg, 1963b: 72.
Sphon, 1966: 245. Sphon 4 Lance, 1968:
80. Roller 8 Long, 1969: 427. Roller,
1970a: 372. Keen, 1971: 840. North, 1971:
58. Robilliard, 1971c: 429. Baker, 1972: 46,
48. Nybakken, 1974: 371. Abbott, 1974:
379, text fig. 4465. Bertsch 8 Ferreira,
1974: 351. Fitch & Lavenberg, 1975: 115.
Thompson, 1976a: 33. Thompson, 1976b:
51, 66. McDonald & Nybakken, 1978: 115,
116. Russo, 1979: 46, 48. McCosker,
1980: 31.
Phidiana nigra MacFarland, 1966: 366-370,
pl. 62, figs. 1-3; pl. 70, figs. 15-16a; pl.
71, figs. 15-20. Sphon 4 Lance, 1968: 80.
Roller, 1970a: 372. Schmekel, 1970: 154.
Nybakken, 1978: 135. Childs, 1980: 53.
Type-specimens: not listed.
Foot relatively broad, and thickened, bila-
biate, and broadest anteriorly, and tapered
posteriorly to tail; foot extends laterally well
beyond body and is separated from it by a
longitudinal groove. Oral tentacles bluish-
white with a red-orange line on dorso-basal
portion, which extends over mouth to oppo-
site tentacle. Large specimens may also bear
a short line of same color mid-ventrally on
oral tentacles. Rhinophores perfoliate, with
14-22 lamellae and are рае orange-yellow
with rich orange-yellow on proximal lamellae.
Cerata cylindrical and slightly clavate, ar-
ranged in about 30 rows dorso-laterally.
About first 10 rows are anterior of cardiac
region and are separated by it medially. Re-
maining rows are posterior of cardiac region
and not well separated medially. General
ground color translucent grayish-white, with
a slight opalescent blue cast on dorsal sur-
face of foot. Large specimens may have an
area of suffused orange-yellow anteriorly on
head. Cerata rose pink distally, fading in in-
tensity proximally, and tipped with white or
gold; cores dark brown to almost black, rare-
ly very dark green. T.L.: 40 mm.
Anus on right just anterior of second group
of cerata. Genital aperture on right side, ven-
tral of posterior edge of first group of cerata.
Penis armed with minute, black chitinous
hook.
Radular formula of specimen examined
17(0.0.1.0.0), MacFarland (1966: 367) re-
ports 21(0.0.1.0.0). Rachidian teeth (Fig. 109)
bear 3-4 small denticles on either side, just
below cusp, and 4-6 larger denticles on either
side below the smaller denticles. Masticatory
border of mandibles bears 25-30 irregular,
blunt denticles.
Type-locality: Laguna Beach, Orange Co.,
California.
Range and habitat: Monterey Bay, Califor-
nia (MacFarland, 1966), to Puerto Rom-
piente, Baja California, Mexico (Farmer &
Collier, 1963); Gulf of Mexico (Childs, 1980).
Intertidal to 220 m, sporadically abundant in
rocky intertidal areas. McDonald & Nybakken
(1978: 116) report that it feeds upon the hy-
droid Hydractinia sp. Lance (1962a: 159)
states that it attacks other aeolids. Fitch 8
Lavenberg (1975: 115) state that they have
found only nudibranchs, including Phidiana
pugnax, in stomachs of the fish Chirolophis
nugator.
Remarks: O'Donoghue (1927b: 104-105)
described Facelina hiltoni from Laguna Beach,
California. Except for references to the origi-
nal description, this species has not been
mentioned in the literature since the original
description. O'Donoghue's description fails to
mention some important characters (e.g. lo-
cation of anus, penial stylet, etc.) but the in-
formation given seems sufficient to distin-
CALIFORNIA NUDIBRANCHS 203
guish the species. In comparing the
description of F. hiltoni with those of P. pug-
nax and its junior synonym P. nigra, | can
find no significant differences. The shape of
the radular teeth is certainly very similar
(O'Donoghue, 1922b: fig. 71; Lance, 1962a:
fig. 6; MacFarland, 1966: pl. 71, figs. 17-20).
The radular formulae are similar, P. pugnax
has 19 teeth and 6 or 7 denticles on either
side of the median denticle (Lance, 1962b:
158), P. nigra has 21 teeth and 7-10 denti-
cles on either side of the median denticle
(MacFarland, 1966: 367), and P. hiltoni has
19 teeth and 7-8 denticles on either side of
the median denticle (O'Donoghue, 1927b:
105). The masticatory border of the mandi-
bles is denticulate in all three cases and the
rhinophores are perfoliate. P. pugnax has 6
major groups of cerata with black cores
(Lance, 1962b: 157-158); P. nigra has 5
groups of cerata with brown to black cores
(MacFarland, 1966: 366-368); and P. hiltoni
has 6 oblique rows of cerata with chocolate
brown cores (O'Donoghue, 1927b: 104). The
type-locality for P. pugnax is Point Loma, San
Diego Co., California which is less 100 km
from that of P. hiltoni. Although Mac-
Farland's description of P. nigra was not
published until 1966, he mentions having col-
lected 10 specimens in 1932, showing that
P. pugnax was known on the California coast
at about the same time that P. hiltoni was
described. The type-locality of P. hiltoni is also
within the geographic range reported for P.
pugnax. The above evidence is sufficient to
establish the synonymy of P. pugnax with the
previously enigmatic P. hiltoni. The fact that
no other aeolid which even vaguely resem-
bles P. hiltonihas been reported from the Pa-
cific coast of North America lends additional
support to the synonomy of the two species.
Therefore, P. pugnax is a junior subjective
synonym of P. hiltoni, the latter being the
correct name by the law of priority.
Phidiana morroensis (Roller, 1972)
Emarcusia morroensis Roller, 1972: 420-423,
text figs. 19-28.
Type-specimens: Calif. Acad. Sci., no. 490.
Foot linear, slightly wider than body, rather
truncate and bilabiate anteriorly and tapered
posteriorly to tail. Oral tentacles long, cylin-
drical, and slightly tapered to rather bluntly
pointed tips, translucent grayish-white with
numerous opaque white dots on distal third.
Rhinophores smooth, pale orange on proxi-
mal two thirds and translucent grayish-white
with numerous opaque white dots on distal
third. Cerata arranged in 7-9 transverse rows
which occur on slightly elevated ridges on
either side of dorsum. Four or five rows oc-
cur anterior of cardiac region, and 3-4 rows
are posterior of cardiac region. General
ground color translucent grayish-white. Two
oval, light orange spots occur dorso-medially
on head, one just anterior and one just pos-
terior of rhinophores; in occasional speci-
mens these two spots may be connected by
a narrow, light orange line which passes be-
tween rhinophores. A narrow, light orange line
passes antero-laterally on either side of head,
from anterior-most cerata onto proximal two
thirds of oral tentacles. An irregular series of
white dots occurs medially from head to pos-
terior of cardiac region; an irregular, white line
extends medially from posterior of cardiac re-
gion to posterior-most cerata row. Cerata
translucent whitish, cores light ochre with
scattered, dark brownish-black spots which
are often concentrated in 3 bands, distal band
darkest. A distinct reddish-brown band often
occurs at bases of cerata; surface of cores
bears numerous, opaque white dots. T.L.: 10
mm.
Anus on right in first post-cardiac row of
cerata. Genital aperture on right side, ventral
of second and third cerata rows. Penis armed
with short, straight stylet.
Radular formula of specimen examined
24(0.0.1.0.0), Roller (1972: 422) reports 15-
22(0.0.1.0.0). Rachidian teeth (Fig. 110) bear
6-10 very long denticles on either side of
large, median cusp. Masticatory border of
mandibles bears 2 rows of denticles, inner
row with about 23 short denticles, and outer
row with about 13 very small denticles.
Type-locality: Morro Bay, San Luis Obispo
Co., California.
Range and habitat: Elkhorn Slough, Mon-
terey Co., California (Roller, 1972), to San
Diego, San Diego Co., California (Roller,
1972). Intertidal to 19 m, usually found in bays
on floating docks and pilings, in association
with hydroids upon which it probably feeds.
Phidiana stearnsi (Cockerell, 1901)
Facelina stearnsi Cockerell, 1901a: 86. Eliot,
1907: 331. O'Donoghue, 1926: 230:
O'Donoghue, 1927b: 105-107, pl. 3, figs.
72-73. Marcus, 1958: 58. Steinberg, 1961:
62.
Type-specimens: not listed.
204 MCDONALD
Foot rather narrow, bilabiate and rounded
anteriorly and tapered posteriorly to sharp,
pointed tail. Anterior foot corners bear a
groove on ventral margin. Oral tentacles
cream on distal third, scarlet-orange on me-
dial third, and maroon-pink on proximal third.
Rhinophores annulate with 10-13 annuli; tips
cream-yellow, distal half scarlet-orange,
shading to maroon-pink basally. Cerata cylin-
dro-conical, quite long, and tapered distally
to acute tips; arranged in 5-8 rows on slight-
ly elevated ridges dorso-laterally on either side
of dorsum. General ground color transcluent
pinkish-white. An irregular, longitudinal band
of brilliant vermilion to scarlet-orange occurs
on either side of head, from base of oral ten-
tacles to base of rhinophores, and a similar
band occurs dorso-medially on tail. Irregular
blotches of scarlet-orange to vermilion occur
on either side of dorsum, between groups of
cerata. Minute cream to white flecks occur
between and posterior of bases of rhino-
phores. Foot corners colored as oral tenta-
cles. Cerata translucent pinkish-maroon, each
with a subapical band of scarlet-orange to
vermilion below whitish tip which bears ma-
roon flecks. Opaque white occurs below the
subapical band and becomes more diffuse
proximally; cores dark maroon-brown to pale
olivaceous gray. T.L.: 15 mm.
Anus on right, just ventral and anterior of
second group of cerata. Genital aperture on
right side, just ventral of first group of cerata.
Radular formula of specimen examined
26(0.0.1.0.0), O'Donoghue (1927b: 106) re-
ports 21-23(0.0.1.0.0). Rachidian teeth (Fig.
111) bear 4-6 denticles on either side of me-
dian cusp. Masticatory border of mandibles
denticulate.
Type-locality: San Pedro, Los Angeles Co.,
California.
Range and habitat: Santa Barbara, Santa
Barbara Co., California (preserved specimen,
Calif. Acad. Sci.), to La Jolla, San Diego Co.,
California (personal observation). Intertidal in
rocky areas and occasionally on mudflats.
Quite uncommon.
Platydoris Bergh, 1877
Body doridiform, rather compressed dor-
so-laterally, equally rounded anteriorly and
posteriorly. Dorsum smooth or minutely
granular. Anterior margin of foot bilabiate.
Labial tentacles digitiform. Rhinophores per-
foliate and retractile. Branchial plumes 6 or
8, arranged in a circle around anus, retractile
into a sheath with valvular lobes.
Labial disc unarmed. Radular formula
0.n.0.n.0, laterals numerous and hamate.
Penis armed with short spines.
Type-species: Platydoris argo (Linnaeus,
1758).
Platydoris macfarlandi Hanna, 1951
Platydoris macfarlandi Hanna, 1951: 1-3, pl.
2, figs. 1-5. Lance, 1961: 66. Steinberg,
1963b: 70. Roller 8 Long, 1969: 429. Ab-
bott, 1974: 357.
Type-specimens: Calif. Acad. Sci., no.
9510.
Dorsum almost smooth, closely set with
very minute papillae, causing dorsum to ap-
pear velvety. Foot truncate anteriorly and ta-
pered posteriorly to a point. Labial tentacles
well developed. General ground color deep,
dark red. T.L.: 30 mm.
Radular formula of specimen examined
46(69-76.0.69-76). Laterals (Fig. 75) ha-
mate.
Type-locality: Pismo Beach, San Luis Ob-
sipo Co., California.
Range and habitat: Off Pismo Beach, San
Luis Obispo Co., California (Hanna, 1951);
subtidal to 157 m.
Remarks: This species is known only from
the original 3 specimens described by Hanna
(1951). No further specimens have been re-
ported and little is known about the species.
Polycera Cuvier, 1817
Body limaciform, highest in branchial re-
gion; with a dorso-lateral pallial ridge on either
side. Anterior margin of head somewhat ex-
panded, forming a frontal veil which bears
digitiform processes. Labial tentacles short
and lobiform. Rhinophores non-retractile and
perfoliate. Branchial plumes non-retractile,
arranged in a circle or semi-circle around
anus. Extra-branchial appendages present.
Mandibles bear a wing-like process. Rad-
ular formula n.2.0.2.n. Laterals hamate, sec-
ond lateral much larger than first. Marginals
smaller and flattened.
Penis armed with small hooks.
Type-species: Polycera quadrilineata (Mül-
ler, 1776).
Polycera atra MacFarland, 1905
Polycera atra MacFarland, 1905: 50-51.
MacFarland, 1906: 142-143, pl. 20, figs.
65-72; pl. 21, figs. 105, 111; pl. 29, fig. 22.
Berry, 1907: 35. Hilton, 1919: 34. O'Don-
CALIFORNIA NUDIBRANCHS 205
oghue, 1926: 217. Ingram, 1935: 48. Hew-
att, 1937: 200. Costello, 1938: tab. 1.
Odhner, 1941: 13, 17. Smith & Gordon,
1948: 180. Marcus, 1955: 159. Lance,
1961: 66. Steinberg, 1963b: 70. Paine,
1963a: 4. Collier & Farmer, 1964: 389.
Paine, 1964: 385. Paine, 1965: 604, 607.
MacFarland, 1966: 115-118, pl. 18, figs.
1-4; pl. 31, figs. 27-31. Marcus & Marcus,
1967a: 196-198. Sphon & Lance, 1968: 80.
Haderlie, 1968: 333 ff. Haderlie, 1969: tabs.
1-2. Roller & Long, 1969: 428. Turner et
al., 1969: 137. Long, 1969c: 232. Gosliner
& Williams, 1970: 179. Schmekel, 1970:
202. Robilliard, 1971b: 242. Keen, 1971:
827. Abbott, 1974: 359, pl. 17, fig. 4283.
Lewbel & Lance, 1975: 346. Thompson,
1976a: 35. Nybakken, 1978: 135. Haderlie
& Donat, 1978: 52, 60. McDonald & Ny-
bakken, 1978: 112. (Non) Bernard, 1970:
85 (=Polycera zosterae O'Donoghue,
1924).
Type-specimens: U. S. Nat. Mus., no.
181278.
Pallial ridges extend from posterior of rhi-
nophores to posterior of branchial plumes,
where they unite and continue postero-me-
dially as caudal crest, bearing a few small tu-
bercles. Each pallial ridge bears five or more
tubercles which are usually yellowish. Frontal
veil bears 4-8 tapered, pointed processes, the
medial being longest, and those located lat-
erally shorter; these processes are dusky
grayish-white with an orange-yellow band
medially. Foot relatively narrow. Rhinophores
bear 8-12 lamellae, and are translucent gray-
ish-white, with a slightly dusky gray hue, each
bears a subterminal band of orange-yellow
on clavus, and a large triangular spot of like
color on inner surface of shaft near base.
Branchial plumes 7-11, unipinnate, dusky
grayish, each with an orange spot distally on
outer face, a larger spot proximally and a pale
orange spot at base of inner face. There are
usually 2-4 extra-branchial appendages
borne upon the pallial ridge on either side of
branchial plumes, the appendages are yel-
low-orange with grayish-white tips. General
ground color pale grayish, with longitudinal
black lines of various widths on dorsum and
sides. Oblong spots of yellowish-orange oc-
cur everywhere between black lines, which
may branch somewhat. T.L.: 12 mm.
Radular formula of specimen examined
9(2-4.2.0.2.2-4), MacFarland (1966: 117) re-
ports 9-11(2-4.2.0.2.2-4). Laterals (Fig. 52a,
b) hamate, first laterals (Fig. 52a) smaller and
each bears a triangular expansion about mid-
way on shaft, below cusp. Second laterals
(Fig. 52b) larger and each has a rather trian-
gular expansion proximally on shaft. Margin-
als (Fig. 52c, d) roughly triangular, they de-
crease in size outwardly, and each bears a
longitudinal crest. Mandibles bear a thick-
ened, oval cutting surface.
Type-locality: Monterey Bay, California.
Range and habitat: Limantour Estero,
Marin Co., California (Gosliner & Williams,
1970), to Islas Coronados, Baja California,
Mexico (Lance, 1961). Intertidal to 18 m, oc-
casionally in rocky coastal areas, but more
common on floating docks and pilings in bays
where it feeds upon the bryozoan Bugula pa-
cifica (MacFarland, 1966; McDonald 8 Ny-
bakken, 1978); also reported to feed upon
the encrusting bryozoan Membranipora
membranacea and the gorgonian Lophogor-
gia chilensis (Lewbel 8 Lance, 1975; Mac-
Ginitie & MacGinitie, 1949).
Polycera hedgpethi Marcus, 1964
Polycera hedgpethi Marcus, 1964: 128-131,
text figs. 1-4. Collier 8 Farmer, 1964: 389.
Lance, 1966: 76. Marcus 8 Marcus, 1967a:
200. Sphon 8 Lance, 1968: 80. Roller 8
Long, 1969: 428. Robilliard, 1971b: 242.
Farmer, 1971:19. Keen, 1971: 827, fig.
2352. Gosliner & Williams, 1973b: 354. Ab-
bott, 1974: 359. Poorman 8 Poorman,
1978: 373. McDonald 8 Nybakken, 1978:
12:
Type-specimens: U. S. Nat. Mus., no.
575603.
Pallial ridges extend from posterior of fron-
tal veil to posterior of branchial plumes where
they unite and continue posteriorly as caudal
crest. Frontal veil bears 4-6 cylindrical, ta-
pered, pointed processes which are translu-
cent grayish-white proximally and distally,
with a narrow, subterminal band of gray-black
dots and a wider proximal band of yellow;
there are frequently gray-black dots scat-
tered throughout the length of the processes.
Foot relatively narrow, anterior corners
slightly produced. Rhinophores bear 8-12 la-
mellae, clavus tipped with white and bears a
subterminal band of yellow and numerous
closely set gray-black dots proximally, which
continue down onto shaft. Branchial plumes
7-9, tripinnate, translucent grayish-white,
densely covered with numerous gray-black
dots, and with a little yellow distally. There
are 2-3 cylindrical, bluntly pointed, extra-
206 MCDONALD
branchial appendages on either side of bran-
chial plumes; they converge somewhat pos-
teriorly, and are translucent grayish-white,
covered with various amounts of gray-black
dots, especially proximally, and in a more or
less distinct subterminal band, and with a
band of yellow proximal of gray-black subter-
minal band. General ground color translucent
grayish-white; dorsum, sides of body, head,
and dorsal surface of tail densely covered with
grayish-black dots, giving the animal an over-
all gray-black appearance. Pallial ridge and
caudal crest are nearly devoid of grayish-
black dots and may bear a few yellow dots.
Tip of tail usually yellow. T.L.: 15 mm.
Radular formula of specimen examined
14(3-4.2.0.2.3-4), Marcus (1964: 131) re-
ports 17(3-4.2.0.2.3-4). First laterals (Fig.
53a) hooked distally and each bears a low
tubercle near the base. Second laterals (Fig.
53b) slightly larger and each is hooked at tip
and bears a larger tubercle near base. Mar-
ginals (Fig. 53c-f) roughly rectangular. Mas-
ticatory border of mandibles smooth and
thickened.
Type-locality: Tomales Bay, Marin Co.,
California.
Range and habitat: Tomales Bay, Marin
Co., California (Marcus, 1964), to Mission
Bay, San Diego Co., California (Lance, 1966);
Bahía de los Angeles, Gulf of California,
Mexico (Lance, 1966). Intertidal to at least 5
m, usually on floating docks and pilings in
bays, where it feeds upon the bryozoan Bu-
gula pacifica (McDonald 8 Nybakken, 1978;
Robilliard, 1971b).
Remarks: Polycera gnupa Marcus 8 Mar-
cus (1967a: 198) is probably a synonym of
P. hedgpethi.
Polycera tricolor Robilliard, 1971
Polycera sp. Lance, 1969: 3.
Polycera tricolor Robilliard, 1971b: 235-243,
text figs. 1-10. Thompson, 1976b: 41.
Type-specimens: Calif. Acad. Sci., no. 447.
Pallial ridges extend from posterior of fron-
tal veil to extra-branchial appendages, pos-
terior of which they unite and continue pos-
tero-medially as caudal crest. Dorsum and
sides of body bear a few scattered, low tu-
bercles which are translucent grayish-white
to pale yellow. Frontal veil bears 8-11 cylin-
drical, tapered, pointed processes, the medi-
al processes longest and lateral processes
shorter; they are white tipped, with a yellow
band on distal one half to two thirds, and
black on base and continuing onto frontal veil.
Foot relatively narrow, anterior corners
somewhat produced. Rhinophores bear 10-
20 lamellae, clavus tipped with white with a
subterminal band of yellow and proximal of
this, a medial band of black. Branchial plumes
5-6, bi- and tripinnate, yellow on distal quar-
ter to half and pale to deep black on proximal
half to three quarters. There are 4-6 cylindri-
cal, tapered, extra-branchial appendages
borne on either side of branchial plumes; they
converge posteriorly and are white tipped with
a subterminal band of yellow on distal half to
two thirds and translucent grayish-white
proximally. General ground color translucent
grayish-white; a yellow line extends around
margin of foot, and may be discontinuous in
some specimens; a dorso-medial line of the
same color occurs on caudal crest. A narrow,
discontinuous line of yellow occurs along pal-
lial ridge; and a dorso-medial row of yellow
tubercles occurs on dorsum, and a few scat-
tered yellow tubercles occur elsewhere on
dorsum and sides of body. T.L.: 15 mm.
Radular formula of specimen examined
13(3-4.2.0.2.3-4), Robilliard (1971b: 240) re-
ports 9-16(3.2.0.2.3). First laterals (Fig. 54a)
are hooked and each bears a denticle near
base. Second laterals (Fig. 54b) are hooked
but lack any distinct denticles. Marginals (Fig.
54c-f) roughly rectangular. Masticatory bor-
der of mandibles smooth and thickened.
Type-locality: Bamfield, Barkley Sound,
British Columbia, Canada.
Range and habitat: Ucluelet, Vancouver Is-
land, British Columbia, Canada (Robilliard,
1971b), to La Jolla, San Diego Co., California
(Robilliard, 1971b). Subtidal to 60 m, usually
in rocky areas. Relatively rare in central Cal-
ifornia. Robilliard (1971b: 240) states that it
preys upon bryozoans.
Polycera zosterae O'Donoghue, 1924
Polycera zosterae O'Donoghue, 1924: 7-10,
pl. 1, figs. 5-9. O'Donoghue, 1926: 217.
Baba, 1937b: 291. Odhner, 1941: 17.
LaRocque, 1953: 258. Marcus, 1957: 434.
Steinberg, 1963b: 70. Collier 8 Farmer,
1964: 389. Bernard, 1970: 85. Robilliard,
1971b: 242. Gosliner & Williams, 1973a:
252-253. Abbott, 1974: 359. McDonald 8
Nybakken, 1978: 112.
Polycera atra MacFarland. Bernard, 1970: 85.
Type-specimens: not listed.
Pallial ridges extend from oral veil to bran-
chial plumes, and both bear a series of low
CALIFORNIA NUDIBRANCHS 207
tubercles which are translucent grayish-white,
encrusted with yellow and dark brown dots,
the yellow being more concentrated distally.
Similar tubercles occur elsewhere on dorsum
and sides of body and are largest and most
abundant just anterior of branchial plumes.
Frontal veil bears 5-6 anterior tubercles on
either side; these tubercles are grayish-white,
encrusted with yellow dots. Foot relatively
narrow. Rhinophores bear about 6 lamellae,
and are translucent brownish with encrusting
yellow and dark brown to black dots. Bran-
chial plumes 3-5, bi- and tripinnate, dusky
brownish-white, tipped with yellow dots and
a scattering of dark brown to black dots.
There are about 3-6 extra-branchial append-
ages on either side, postero-lateral of bran-
chial plumes, they are whitish with yellow
dots; the 3 posterior-most appear to arise
from a common base. A dorso-medial row of
tubercles passes from posterior of branchial
plumes to tail. General ground color translu-
cent grayish-white, with numerous small,
brown to black dots and a lesser number of
yellow dots over entire body, except ventral
surface of foot. T.L.: 10 mm.
Radular formula of specimen examined
13(5.2.0.2.5), O'Donoghue (1924: 8) reports
15(5-6.2.0.2.5-6). Laterals are hamate, first
laterals (Fig. 55a) smaller and each bears a
denticle about midway on shaft, and a small-
er denticle just below cusp. Second laterals
(Fig. 55b) larger and each bears a small den-
ticle near base. Marginals (Fig. 55c-g) rough-
ly quadrangular and decrease in size out-
ward. Mandibles bear a thickened cutting
surface.
Type-locality: Newcastle Island, Vancou-
ver Island, British Columbia, Canada.
Range and habitat: Shushartie Bay, Van-
couver Island, British Columbia, Canada
(Robilliard, 1971b), to Bodega Bay, Sonoma
Co., California (Gosliner & Williams, 1973a).
Intertidal and subtidal, in bays on floating
docks and pilings, and on eelgrass (Zostera
marina). This species occurs upon the bryo-
zoan Bowerbankia gracilis var. aggregata,
upon which it may feed. Robilliard (1971b:
242) reports that it feeds upon the bryozoan
Membranipora sp.
Precuthona Odhner, 1929
Body aeolidiform. Anterior foot corners
rounded, not produced. Cerata cylindrical,
linear, and tapered to blunt tips, arranged in
closely set transverse to slightly oblique dor-
so-lateral rows. Oral tentacles cylindrical and
tapered to blunt tips. Rhinophores non-re-
tractile, smooth, cylindrical, rather long, and
tapered distally.
Masticatory border of mandibles denticu-
late. Radula uniseriate, with a few denticles
on either side of median cusp.
Penis unarmed. Anus acleioproct.
Type-species: Precuthona peachii (Alder 8
Hancock, 1848).
Precuthona divae Marcus, 1961
Precuthona divae Marcus, 1961: 50-52, pl.
10, figs. 180-184. Steinberg, 1963b: 72.
Sphon 8 Lance, 1968: 80. Roller & Long,
1969: 428. Roller, 1970a: 372. Gosliner &
Williams, 1970: 179. Robilliard, 1971a: 164,
165. Bertsch et al., 1972: 307. Nybakken,
1974: 371. Abbott, 1974: 377, text fig.
4454. Harris et al., 1975: 264, 267. Chris-
tensen, 1977: 131. McDonald & Nybak-
ken, 1978: 114. Williams & Gosliner, 1979:
214.
Cuthona rosea MacFarland, 1966: 326-332,
р 59, figs. 1=2: р. 68, figs; 1=7: spl: 70;
figs. 9-10. Sphon & Lance, 1968: 80. Rol-
ler, 1970a: 372. Abbott, 1974: 377.
Cuthona divae (Marcus). Williams & Gosliner,
1979: 208, 210, 214, 215.
Type-specimens: Dept. Zool., Univ. Sáo
Paulo, Brazil.
Body slightly compressed dorso-ventrally,
sides of body distinct from margin of foot.
Foot elongate, widest and rounded anterior-
ly, tapered posteriorly to short, pointed tail.
Oral tentacles long, cylindrical, and tapered
to blunt tips, translucent cream to pinkish-
brown. A true frontal veil is absent, but head
broad and flattened ventrally and slightly ex-
tended laterally. Rhinophores colored as oral
tentacles. Cerata rows widely separated an-
tero-medially but converge postero-medially.
First 2-3 rows anterior of rhinophores and
consist of 3-4 cerata on either side. General
ground color translucent cream to brownish-
pink, cerata same color, tips encrusted with
opaque white, cores reddish-brown to pink
or burnt umber. T.L.: 15 mm.
Anus on right, just anterior of medial end
of eighth row of cerata. Genital aperture on
right side between fourth and fifth rows of
cerata.
Radular formula of specimen examined
23(0.0.1.0.0), MacFarland (1966: 327) re-
ports 28-32(0.0.1.0.0), and Marcus (1961: 51)
reports 21 (0.0.1.0.0). Rachidian teeth (Fig.
208 MCDONALD
115) each bear a large median cusp with 8-
10 (rarely as few as 3) smaller denticles on
either side. Masticatory border of mandibles
bears a single row of 10-15 blunt denticles.
Type-locality: Dillon Beach, Marin Co., Cal-
ifornia.
Range and habitat: San Juan Island, Puget
Sound, Washington (Robilliard, 1971a), to
Dume Rock, Paradise Cover, Malibu, Los
Angeles Co., California (personal observa-
tion). Intertidal to 18 m, occasional in rocky
intertidal areas. Feeds upon the hydroid Hy-
dractinia sp. (Christensen, 1977; Harris et al.,
1975; MacFarland, 1966) which it somewhat
resembles.
Remarks: This species very closely resem-
bles Precuthona peachii (Alder & Hancock,
1848) with which it may be conspecific.
Rostanga Bergh, 1879
Body doridiform, rather equally rounded
anteriorly and posteriorly; dorsum densely
covered with numerous minute papillae. La-
bial tentacles digitiform. Rhinophores perfo-
liate, retractile into sheaths. Branchial plumes
retractile, usually unipinnate, arranged in a
circle around anus.
Labial disc armed with small hooks. Rad-
ular formula 0.n.0.n.0. Inner laterals strongly
hooked, grading to outer laterals which are
thin and elongate.
Penis unarmed.
Type-species: Rostanga coccinea Forbes
in Alder & Hancock, 1848.
Rostanga pulchra MacFarland, 1905
Rostanga pulchra MacFarland, 1905: 40-41.
MacFarland, 1906: 119-122, pl. 18, figs.
18-21; pl. 24, fig. 8. Eliot, 1907: 330, 339-
341. Berry, 1907: 34. Guernsey, 1912: 77,
fig. 38A. Eliot, 1913: 20. O'Donoghue,
1922b: 152-154, 165, pl. 5, figs. 12-15.
O'Donoghue, 1924: 23, 29. O'Donoghue,
1926: 208. O'Donoghue, 1927b: 82-83, pl.
1, figs. 10-12. deLaubenfels, 1927: 266.
Boone, 1929: 38. Baba, 1933: 277. Baba,
1935a: 344, 345. Ingram, 1935: 48, 49.
Baba, 1937e: 3. Costello, 1938: 327 ff,
tabs. 1-5, pl. 1, figs. 13-15. Hewatt, 1946:
191, 193, 198. Smith & Gordon, 1948: 180.
Rigg & Miller, 1949: 344. LaRocque, 1953:
258. Marcus, 1958: 25. Marcus, 1959: 3,
7, 35-37 ff, figs. 65-68. Marcus, 1961: 15-
16, 57, pl. 3, figs. 46-49. Lance, 1961: 66.
Cooke, 1962: 194-196. Farmer & Collier,
1963: 62. Steinberg, 1963b: 70. Paine,
1963a: 4. MacFarland, 1966: 165-169,
170, pl. 25, fig. 7; pl. 29, figs. 7—10;pl.::35,
figs. 1-16. Lance, 1966: 69, 72. Hurst,
1967: 255 ff, text fig. 9, pl. 28, fig. 11; pl.
37, fig. 47; fig. 24-21. Sphon & Lance,
1968: 81. DuShane & Sphon, 1968: 244.
Miller, 1968: 134. Roller 8 Long, 1969: 428.
Long, 1969c: 232. Marcus 8 Marcus, 1969:
20-21. Fournier, 1969: 74. Burn, 1969: 82.
Robilliard, 1969a: 290. Gosliner 8 Wil-
liams, 1970: 179. Marcus & Marcus, 1970:
202-203. Long, 1970: 19. Bernard, 1970:
85. Keen, 1971: 821. Harris, 1971a: 82, 85.
Bertsch et al., 1972: 307-308. Baker,
1972: 49. Sphon, 1972a: 156. Harris, 1973:
239-240, 269. Navoni, 1973: 1334. Ander-
son, 1973: 121, 122. Abbott, 1974: 353,
pl. 17, fig. 4238. Haderlie, 1974: tab. 4.
Thompson, 1975: 489, 490. Lambert, 1976:
297, 299. Bloom, 1976: 293, 295. Behrens
& Tuel, 1977: 33, 35. Chia & Koss, 1978:
109-119, figs. 1-4. Hadfield, 1978: 167.
Chia, 1978: 283. Nybakken, 1978: 134 ff.
McDonald & Nybakken, 1978: 112. Bickell
& Chia, 1979a: 306, 310.
Rostangia pulchra. Hewatt, 1937: 178 ff (lap-
sus).
Type-specimens: U. S. Nat. Mus., no.
181292.
Dorsum thickly set with numerous small,
hispid papillae. Foot abruptly rounded and
deeply bilabiate anteriorly, and slightly ta-
pered posteriorly to short, bluntly pointed tail.
Labial tentacles digitiform, long, and slender.
Rhinophores bear 16-24 nearly vertical la-
mellae, shaft prolonged above clavus and
bluntly tipped. Rhinophore sheaths bear pa-
pillate margins. Rhinophores orange to scar-
let, lamellae darker shade of same color.
Branchial plumes 8-12, unipinnate, orange to
scarlet. General ground color orange to bright
red or scarlet, although occasional speci-
mens are light salmon pink. Dorsum sprin-
kled with minute dots of brown to black, the
amount of which is quite variable. T.L.: 10
mm.
Radular formula of specimen examined
70(0.75.0.75.0), MacFarland (1966: 167) re-
ports 68-80(81.0.81), and Marcus (1961: 16)
reports 80(75-90.0.75-90), while Bloom
(1976: 293) reports 65-80(39-90.0.39-90).
First laterals (Fig. 57f) strongly hooked and
bear 4-11 small denticles on hook. Succeed-
ing about 10 laterals (Fig. 57c-e) strongly
hooked. From about twelfth lateral outward,
the hook lengthens and becomes more slen-
CALIFORNIA NUDIBRANCHS 209
der and less curved. Outermost laterals (Fig.
57a) quite elongate, with 1-7 very long den-
ticles, appearing almost whisk-like.
Type-locality: Monterey Bay, California.
Range and habitat: Dundas Island, British
Columbia, Canada (Lamber, 1976), to Point
Loma, San Diego Co., California (Marcus,
1961); Bahia de los Angeles, Gulf of Califor-
nia, Mexico (Lance, 1966); Chiloé Island, Chile
(Marcus, 1961); Camarones Bay, Argentina
(Marcus & Marcus, 1969). Intertidal to 102
m, frequent in rocky intertidal areas. Feeds
upon the encrusting red sponges: Acarnus
erithacus, Esperiopsis originalis, Ophlita-
spongia pennata, Plocamia karykina, and
Plocamia lithophoenix (Chia & Koss, 1978;
Cook, 1962; deLaubenfels, 1927; Mac-
Farland, 1966).
Remarks: Baba (1933: 277; 1935a: 344)
synonymized Я. pulchra with Rostanga mus-
cula (Abraham, 1877), the latter being a syn-
onym of Rostanga arbutus (Angas, 1864).
However, Marcus (1958: 25; 1959: 36-37),
Marcus 8 Marcus (1969: 21), and Burn (1969:
82) give ample evidence, based on the radu-
la, for maintaining R. pulchra as a distinct
species.
Sclerodoris Eliot, 1904
Body doridiform, rather equally rounded
anteriorly and posteriorly; dorsum rough, with
tubercles, pits, depressions and reticulate
ridges developed to varying degrees. Spicu-
late sessile caryophyllidia and/or tubercles
with retractile papillae may be present. Rhi-
nophores perfoliate, retractile into crenulate
sheaths. Branchial plumes retractile, tripin-
nate and bushy.
Radular formula 0.п.0.п.0. Laterals ha-
mate, usually nondenticulate, outermost 3-4
teeth small and may be pectinate or serrated
distally.
Type-species: Sclerodoris tuberculata Eliot,
1904.
| use here Sclerodoris sensu Rudman
(1978). Bertsch (1981) gives ample evidence
for the placement of the following species in
this genus.
Sclerodoris tanya (Marcus, 1971)
Doris tanya Marcus, 1971: 357-362, text figs.
4-8. Sphon, 1973: 5. Marcus, 1973: 5.
Halgerda sp. Hertz, 1978: 90, fig. 1.
Sclerodoris tanya (Marcus). Bertsch, 1981:
217-220, figs. 2-7.
Type-specimens: Dept. Zool., Univ. Sáo
Paulo, Brazil (Marcus, personal communica-
tion).
Dorsum thickly set with numerous large, ir-
regular tubercles which bear smaller tuber-
cles. Foot rounded anteriorly and posteriorly,
unilabiate anteriorly. Labial tentacles rather
flattened and rounded distally. Rhinophores
bear about 40 lamellae, clavus yellowish-tan,
occasionally with a few darker tan markings.
Branchial plumes 7, multi-pinnate, colored as
dorsum, with a few tan to brownish flecks.
General ground color light yellowish-tan, a
number of darker tan to brown, irregularly
round spots occur on dorsum between tu-
bercles, smaller spots of similar color occur
on tubercles. Ventral surfaces of foot and of
mantle margin bear numerous irregular flecks
of brown. T.L.: 30 mm.
Radular formula of specimen examined
29(32-37.0.32-37), Marcus (1971: 358) re-
ports 23(0.33.0.33.0) and Bertsch (1981: 219)
reports 21-26(25-43.0.25-43). Laterals (Fig.
72) hamate.
Type-locality: Newport Bay, Orange Co.,
California.
Range and habitat: Newport Bay, Orange
Co., California (Marcus, 1971), to San Diego
Flood Control Channel, Mission Bay, San
Diego Co., California (personal observation);
also Gulf of California, Mexico (Bertsch,
1981). Intertidal to 5 m in bays.
Spurilla Bergh, 1864
Body aeolidiform, broadest anteriorly and
tapered posteriorly. Foot truncate anteriorly,
anterior foot corners somewhat produced into
triangular lobes. Cerata numerous, arranged
in numerous transverse or oblique rows, and
frequently decumbent; cnidosacs present.
Rhinophores non-retractile, perfoliate, lamel-
lae very oblique.
Masticatory border of mandibles smooth.
Radula uniseriate, rachidian teeth rather bi-
lobed and pectiniform, with the lateral denti-
cles largest on the medial portion of either
lobe.
Penis unarmed. Anus cleioproct.
The only character which seems to sepa-
rate Spurilla from Aeolidiella Bergh, 1867, is
the rhinophores, which are smooth in Aeoli-
diella and perfoliate in Spurilla. It should be
determined whether any other characters ex-
ist which sufficiently distinguish the two gen-
era. If none can be found, the shape of the
rhinophores alone is probably not sufficient
210 MCDONALD
to separate genera. If this should be the case,
they should be united under the oldest name,
Spurilla Bergh, 1864.
Type-species: Spurilla neapolitana (Delle
Chiaje, 1841).
Spurilla chromosoma Cockerell in
Cockerell & Eliot, 1905
Spurilla chromosoma Cockerell in Cockerell
8 Eliot, 1905: 32, 51-52. Marcus, 1961:
54-56, 57, pl. 10, figs. 196-199. Stein-
berg, 1961: 62. Lance, 1961: 68. Farmer 4
Collier, 1963: 63. Paine, 1963a: 4. Lance,
1966: 80. Marcus 8 Marcus, 1967a: 227
(partim). Sphon & Lance, 1968: 81. Du-
Shane & Sphon, 1968: 244. Burn, 1969:
97, 98. DuShane 8 Brennan, 1969: 361.
Bertsch, 1970: 16. Williams 8 Gosliner,
1970: 33. Bertsch & Smith, 1970: 19.
Farmer, 1971: 19. Keen, 1971: 839, pl. 22,
fig. 2. Harris, 1973: 265. Bertsch, 1973:
110. Abbott, 1974: 381. Michel, 1976: 46,
fig. 2. Poorman 8 Poorman, 1978: 373.
McDonald & Nybakken, 1978: 115.
Spurilla sp. Guernsey, 1912: 78, fig. 39K.
O'Donoghue, 1927b: 78. Steinberg, 1961:
62.
Eolidina chromosoma (Cockerell & Eliot).
O'Donoghue, 1926: 234.
Eolidina orientalis O'Donoghue, 1927b: 109-
110, pl. 3, fig. 78. Steinberg, 1961: 62.
Eolidina (Spurilla) chromosoma Cockerell 8
Eliot. O'Donoghue, 1927b: 109.
Type-specimens: not listed.
Foot broad, broadly rounded and bilabiate
anteriorly, and tapered posteriorly to rather
long, pointed tail; margins are thin and un-
dulating and extend laterally beyond body.
Foot corners produced into rather long, ten-
taculiform processes. Oral tentacles cylindri-
cal, tapered, quite long, and white. Rhino-
phores bear about 10-12 lamellae, clavus red
with white tip. Cerata arranged in about 13
crescentic, transverse rows. General ground
color translucent grayish-white. Head orange
dorsally. A row of opaque white blotches oc-
curs dorso-medially between cerata rows.
Anterior cerata have an orangish hue, tips are
white and cores are greenish-gray. T.L.: 20
mm.
Anus on right, just posterior of sixth row
of cerata. Genital aperture on right side, just
ventral of third and fourth rows of cerata.
Radular formula of specimen examined
24(0.0.1.0.0), Marcus (1961: 55) reports
21(0.0.1.0.0). Rachidian teeth (Fig. 97) pec-
tiniform with 24-32 denticles on either side
of median denticle.
Type-locality: San Pedro, Los Angeles Co.,
California.
Range and habitat: Purisima Point, Santa
Barbara Co., California (Sphon 8 Lance,
1968), to Tenacatita, Jalisco, Mexico (Keen,
1971). Intertidal to 18 m, usually found in
rocky intertidal areas. Harris (1973: 265) re-
ports that it feeds upon the sea anemone
Metridium senile.
Spurilla oliviae (MacFarland, 1966)
Aeolidiella oliviae MacFarland, 1966: 373-
377, pl. 62, figs. 4-6; pl. 72, figs. 9-14.
Sphon & Lance, 1968: 81. Roller, 1970a:
372. Harris, 1973: 221, 260, 265. Schuler,
1975: 33.
Spurilla oliviae (MacFarland). Sphon & Lance,
1968: 81. Roller & Long, 1969: 428. Roller,
1970a: 372. Gosliner & Williams, 1970: 179.
G. Williams, 1971: 215-216, text fig. 1. Ny-
bakken, 1974: 371. Abbott, 1974: 381, fig.
4477. McDonald & Nybakken, 1978: 115.
Spurilla chromosoma Cockerell & Eliot. Mar-
cus, 1961: 54-55 (partim). Nybakken,
1974: 371.
Type-specimens: type material at Calif.
Acad. Sci.
Foot rather broad, somewhat thickened and
bilabiate anteriorly and tapered posteriorly to
moderately long tail; margins are thin and ex-
tend laterally beyond body. Foot corners pro-
duced into rather long, tentaculiform pro-
cesses which bear a shallow groove. Oral
tentacles cylindrical, about twice as long as
rhinophores, and tapered to blunt tips; trans-
lucent grayish-white with encrusting white on
distal half. Rhinophores bear 10-14 oblique
lamellae which are orange-vermilion, shaft
rather short and translucent grayish-white.
Cerata decumbent and arranged in about 17
oblique rows, leaving dorsum free antero-
medially, posterior groups less separated
medially. General ground color translucent
grayish-white, becoming rather cream on
dorsum. A blotch of pale vermilion occurs on
head and another occurs in cardiac region.
Encrusting white occurs anteriorly on foot
margin. Cerata chrome orange with white tips,
cores raw umber. T.L.: 20 mm.
Anus right of cardiac region, between fifth
and sixth rows of cerata. Genital aperture on
right side, ventral of sixth and seventh rows
of cerata.
Radular formula of specimen examined
CALIFORNIA NUDIBRANCHS edn
27(0.0.1.0.0), MacFarland (1966: 374) re-
ports 24(0.0.1.0.0). Rachidian teeth (Fig. 98)
pectiniform with 20-35 denticles on either side
of median denticle.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Duxbury Reef, Marin
Co., California (Gosliner & Williams, 1970), to
Point Fermin, Palos Verdes Peninsula, Los
Angeles Co., California (Sphon, 1972b). Usu-
ally found under rocks in rocky intertidal zone.
Harris (1973: 265) reports that it feeds upon
the sea anemone Metridium senile.
Remarks: The range of variation in the col-
or of this species is such that it may prove to
be conspecific with Spurilla chromosoma.
Tenellia A. Costa, 1866
Body aeolidiform and rather narrow. An-
terior foot corners not produced. Cerata cy-
lindrical, slightly tapered distally, arranged in
groups of 1-4, laterally along either edge of
dorsum; cnidosacs present. Oral tentacles as
such absent, in their place a frontal veil. Rhi-
nophores non-retractile and smooth.
Masticatory border of mandibles finely
denticulate. Radula uniseriate, rachidian teeth
bear a few denticles on either side of median
cusp.
Penis armed with a chitinous stylet. Anus
acleioproct.
Type-species: Tenellia adspersa (Nord-
mann, 1845).
Tenellia adspersa (Nordmann, 1845)
Tergipes adspersus Nordmann, 1844: 270.
Nordmann, 1845: 498, pl. 1, figs. 4-5.
Nordmann, 1846: 110. Adams & Adams,
1854: 76. Gray, 1857: 229. Ostroumoff,
1893: 246. Sowinsky, 1904: 15, 144-145,
150, 310. Milachewitch, 1916: 132. Chukh-
chin, 1960: 111, text fig. 25. Gomoiu, 1961:
1251, fig. 4B. Gomoiu, 1966: 145, 146.
Turpaeva, 1972: 168-185. Lemche, 1973:
90. Roginskaya, 1974a: 139.
Terpiges lacinulatus Blainville. Schultze, 1849:
268, text figs. 1-11. Roginskaya, 1974a:
138. Bonar, 1978: 178.
Eolis ventilabrum Dalyell, 1853: 318, pl. 45,
fig. 28.
Embletonia pallida Alder & Hancock, 1854:
105. Byerley, 1854: 45. Alder & Hancock,
1855: 31, 32, 53, append. p. xii. Gosse,
1856: 100. Adams & Adams, 1858: 635.
Collingwood, 1859: 469. Collingwood,
1860: 196, 202. Collingwood, 1861: 114.
Sanford, 1861: fig. 1. Collingwood 8 Byer-
ley, 1862: 189. Meyer 8 Möbius, 1865: 17-
18, pl. Hensche, 1866: 105-106. Jeffreys,
1869: 36: Sars; 1878: pl. XVI; fig. 11.
Bergh, 1885: 34-37, pl. 2, figs. 14-19; pl.
3, figs. 11-13: р: 4, fig. 12; pl. 5, НО. 7.
Herdman, 1886: 276, 277. Higgins, 1886:
26. Bergh, 1890а: 33. Bergh, 1893: 1026
(34). Herdman & Clubb, 1892: 146. Herd-
man, 1896: 49. Herdman et al., 1896: 446.
Cooke, 1899: 65. Conchol. Soc., 1901: 25.
Odhner, 1907: 30, 81. Eliot, 1910: 1, 128-
129, 171, pl. 6, figs. 1-2. Colgan, 1913:
165, 167. Sumner et al., 1913: 706. Col-
gan, 1914: 181-182. Loyning, 1922: 65-
69, 94, text figs. 44-50, pl. 3, figs. 13a—b.
Iredale 8 O'Donoghue, 1923: 207. Naville,
1925: 812. Naville, 1926: 252-255, text
figs. 1-2. Hoffmann, 1926: 18. Poisson,
1927: 68-69. Cuenot, 1927: 263. Jutting,
1927: LXXXVIII, XCVI. Loyning, 1927: 262.
Winckworth, 1932: 237. Nicol, 1935: 215.
Engel, 1936: 107. Lemche, 1938: 39. White,
1938: 15. Rasmussen, 1944: 207, figs. 15-
20. Jutting, 1947: 64. Marcus & Marcus,
1955: 230, 238-242, pl. 38, figs. 23-31.
Marcus & Marcus, 1958: 94, 95. Swennen,
1959: 59. Jaeckel, 1961: 140, fig. 2. Swen-
nen, 1961: 217. Thompson, 1961: 236.
Coomans, 1962: 222. Coomans 4 de Con-
inck, 1962: 1. Hadfield, 1963: 93. Thomp-
son, 1964: 284 ff. Ghiselin, 1965: 351.
Thompson, 1967: 12, 14, 18. Lawinski,
1968: 410, figs. 1-6. Bebbington &
Thompson, 1968: 10. Tardy, 1970: 303.
Salvini-Plawen, 1972: 394. Suckow, 1972:
193-194, figs. 1, 3, 4. Rasmussen, 1973:
22, 25, 29, 269-270, 447, 449. Lemche,
1973: 90. Harris, 1973: 217 ff. Rogin-
skaya, 1974a: 138. Thompson, 1976a: 85,
86. Thompson, 1976b: 79. Bonar, 1978:
178. Harris et al., 1980: 70.
Embletonia hyalina ‘‘(Alder 8 Hancock).
Sanford, 1861: 152, 153.
Tenellia mediterranea Costa, 1866: 76, pl. 3,
fig. 7. Carus, 1889-1893: 211. Iredale &
O'Donoghue, 1923: 231. Tardy, 1970: 359.
Roginskaya, 1974a: 138.
Galvina adspersa (Nordmann). Bergh, 1868:
220.
Embletonia grayi Kent, 1869: 109, pl. 8. Gray,
1869: 247. Conchol. Soc., 1901: 25. Ire-
dale & O'Donoghue, 1923: 207.
Embletonia ? pallidus (Alder 8 Hancock).
Mörch, 1871: 184.
Eubranchus pallidus (Alder & Hancock). Jae-
ckel, 1952: 28.
212 MCDONALD
Embletonia mediterranea (Costa). Bergh,
1892: 1026 (34). Poisson, 1927: 68.
O'Donoghue, 1929: 747. Vannucci 8 Ho-
soe, 1953: 103-120, pls. 1-6. Hadfield,
1963: 92. Thompson, 1967: 9. Tardy, 1970:
303. Bonar, 1978: 178.
Tenellia ventilabrum (Dalyell). Pruvot-Fol,
1954: 413, text figs. 160a-d, 9-1. Stein-
berg, 1963a: 65. Schmekel, 1968a: 121,
149. Schonenberg, 1969: 252. Tardy, 1970:
303 ff. Schmekel, 1970: 136, 169, text fig.
30b. Schmekel, 1971: 126. Abbott, 1974:
372. Barletta & Melone 1976: 226, 233.
Bonar, 1978: 185, 190.
Tenellia pallida (Alder £ Hancock). Marcus,
1957: 467. Marcus 8 Marcus, 1958: 95.
Marcus 8 Marcus, 1960: 180-182, fig. 80.
Steinberg, 1963a: 65. Steinberg, 1963b:
72. Baba & Hamatani, 1963a: 337-338,
text fig. 1. Franz, 1968: 10. Marcus, 1970:
216. Marcus, 1972a: 314, fig. 8E. Rogin-
skaya, 1974a: 139. Franz, 1975b: 253.
Thompson & Brown, 1976: 180, fig. 97.
Marcus, 1977: 15. Clark 8 Goetzfried,
1978: 289. Bonar, 1978: 178. McDonald 4
Nybakken, 1978: 114. Eyster, 1979: 133
ff. Eyster, 1980: 582 ff. Williams, 1980: 113.
Embletonia sp. Steinberg, 1960: 49.
Tergipes. Gomoiu, 1961: 1251, text fig. 4B.
Tenellia sp. Steinberg, 1963a: 65.
Stiliger bellulus (d'Orbigny). Chukhchin,
1963a: 149. Chukhchin, 1963b: 197.
Chukhchin, 1963c: 218. Chukhchin, 1967:
102.
Tenellia adspersa (Nordmann). Turpaeva,
1969: 415. Roginskaya, 1970: 167-172,
text figs. 1-4. Maksimov et al., 1971: 902—
907. Roginskaya, 1974a: 139. Turpaeva 8
Simkina, 1975: 1149, 1152. Turpaeva &
Lebedeva, 1976: 1437, 1444. Cooper,
1978: 8. Williams 8 Gosliner, 1979: 215.
Brown, 1980: 251-252, text fig. 7B. Coo-
per, 1980: 284.
Type-specimens: not listed.
Foot narrow, rather linear, quite truncate
anteriorly and tapered posteriorly to short,
bluntly pointed tail. Circular frontal veil ex-
tends slightly laterally beyond body. Rhino-
phores rather short, translucent grayish-
white. Cerata cylindrical, only slightly tapered
to blunt tips, arranged dorso-laterally on either
side of dorsum in 5-6 groups of 1-2 cerata
each. General ground color translucent yel-
lowish-white, usually with a few small, black
flecks on dorsum. Cerata translucent yellow-
ish-white, occasionally tipped with orange and
occasionally with a pale, powder blue hue
distally; cores light yellowish to yellowish-
brown. T.L.: 5 mm.
Anus on right, between second and third
groups of cerata. Genital aperture on right
side, ventral of first and second groups of
cerata. Penis armed with chitinous stylet.
Radular formula of specimen examined
27(0.0.1.0.0), Eliot (1910: 129) reports
40(0.1.0). Rachidian teeth (Fig. 113) bear 5-
8 denticles on either side of median cusp.
Masticatory border of mandibles finely den-
ticulate.
Type-locality: Black Sea, U.S.S.R.
Range and habitat: San Francisco Bay,
California (Steinberg, 1963a), to Monterey
Harbor, Monterey Co., California (Steinberg,
1963b); Europe (Marcus & Marcus, 1960);
Japan (Baba 8 Hamatani, 1963a); Black Sea
(Nordmann, 1845); Cananeia, Brazil (Marcus
8 Marcus, 1960). Intertidal to 8 m, usually in
bays or estuaries on floating docks and pil-
ings. Feeds upon the hydroids: Bougainvillia
glorietta, Bougainvillia muscoides, Cordylo-
phora caspia, Cordylophora lacustris, Cory-
dendrium dispar, Gonothyraea loveni, Lao-
media gelatinosa, Laomedia longissima,
Laomedia loveni, Obelia sp., Obelia dichot-
oma, Perigonimus megas, Podocoryne sp.,
Protohydra leuckarti, and Psammohydra sp.
(Cooper, 1980; Marcus 8 Marcus, 1955; Na-
ville, 1926; Rasmussen, 1973; Salvini-Plaw-
en, 1972; Sanford, 1861; Schmekel, 1968;
Swennen, 1961; Thompson, 1964; Thomp-
son & Brown, 1976; Turpaeva, 1972). This
species is quite rare in central California.
Remarks: See ICZN (1977, Opinion 1084),
Tergipes adspersus placed on Official List.
Thordisa Bergh, 1877
Body doridiform, rather evenly rounded an-
teriorly and posteriorly; dorsum usually pa-
pillate or granular. Anterior margin of foot bi-
labiate. Labial tentacles short and digitiform.
Rhinophores perfoliate and retractile. Bran-
chial plumes retractile and arranged in circle
around anus.
Labial disc smooth. Radula bears numer-
ous laterals and a few marginals. Laterals
hamate, marginals bear numerous bristles
distally.
Penis may be armed or not.
Type-species: Thordisa maculigera Bergh,
1877.
CALIFORNIA NUDIBRANCHS 213
Thordisa bimaculata Lance, 1966
? Thordisa. Paine, 1963a: 4.
Aldisa sanguinea (Cooper). Farmer & Collier,
1963: 62.
Thordisa bimaculata Lance, 1966: 72-75,
figs. 1-8. Sphon & Lance, 1968: 81. Roller
& Long, 1969: 429. Abbott, 1974: 357.
Type-specimens: Calif. Acad. Sci., no. 100.
Dorsum thickly set with numerous, spicu-
late papillae of various sizes, which are larger
medially and smaller marginally; larger pa-
pillae may be constricted at base. Foot
rounded and bilabiate anteriorly, and slightly
tapered posteriorly to bluntly pointed tail. La-
bial tentacles relatively short and digitiform.
Rhinophores bear 14-16 lamellae which are
bright orange to dull yellow with a brownish
tinge, shaft colored as clavus but lacks
brownish tinge. Rhinophore sheaths have
papillate margins. Branchial plumes 6, bi- and
tripinnate, somewhat lighter in color than
dorsum and bear a few brown flecks on ra-
chis and are encrusted with white. General
ground color varies from bright orange to dull
yellowish. Two concentrations of brown dots
occur medially on dorsum, one just posterior
of rhinophores and the other just anterior of
branchial plumes. T.L.: 25 mm.
Penis armed with 7-10 longitudinally ori-
ented, large spines.
Radular formula of specimen examined
32(6.25.0.25.6), Lance (1966: 74-75) reports
32(8.34.0.34.8) and 31(6.29.0.29.6). Inner-
most laterals (Fig. 69f) small and hamate, the
laterals increase in size towards outermost
laterals (Fig. 69d). Marginals (Fig. 69a-c)
slightly hooked and bear numerous bristles
distally. Labial disc has a tessellated surface.
Type-locality: Windnsea Reef, La Jolla, San
Diego Co., California.
Range and habitat: Carmel, Monterey Co.,
California (Lance, 1966), to Isla de Natividad,
Baja California, Mexico (Lance, 1966). Inter-
tidal to 30 m, usually in rocky areas. This
species is rare, especially in the northern part
of its range.
Tochuina Odhner, 1963
Body limaciform, rather subquadrate,
somewhat depressed dorso-ventrally and ta-
pered posteriorly. The numerous branchial
appendages are plumose and arranged dor-
so-laterally along either margin of dorsum.
Dorsum tuberculate. Foot broad. Anterior
margin of head somewhat expanded, form-
ing a frontal veil which is tuberculate. Rhi-
nophores retractile, and bear plumose pro-
cesses distally.
Masticatory border of mandibles smooth.
Radular formula n.1.1.1.n. Rachidian teeth
bear a blunt cusp. Laterals short, blunt hooks;
marginals rather hamate.
Penis unarmed. Anus on right side, be-
neath margin of dorsum.
Type-species: Tochuina tetraquetra (Pal-
las, 1788).
Tochuina tetraquetra (Pallas, 1788)
Limax tetraquetra Pallas, 1788: 237, 239, pl.
5, fig. 22. Odhner, 1963: 50.
Doris tetraquetra (Pallas). Gmelin in Lin-
naeus, 1791: 3106.
Tritonia tetraquetra (Pallas). Bosc, 1830: 108.
Bergh, 1879b: 98-105, pl. 3, figs. 13-16;
pl. 4, figs. 5-12; pl. 5, figs. 1-2. Bergh,
1879c: 154-161, pl. 3, figs. 13-16; pl. 4,
figs. 5-12; pl. 5. figs. 1-2. Bergh, 1881b:
237. Bergh, 1884b: 701, 726-727. Bergh,
1892: 1068 (76). Cockerell & Eliot, 1905:
33. O'Donoghue, 1922b: 146-149, 165, pl.
5, figs. 1-5. Odhner, 1926a: 32. Odhner,
1936: 1080. Baba, 1937a: 197. Marcus,
1961: 32. Willows, 1965: 707 ff. Willows,
1968: 226. Willows 8 Dorsett, 1975: 118.
Willows, 1976: 329 ff.
Tritonia sp. Eliot, 1901: 163-165, text figs.
1-2.
Tritonia gigantea Bergh, 1904: 26-28, pl. 4,
figs. 29-32. Cockerell 8 Eliot, 1905: 33.
Odhner, 1926a: 32. Marcus, 1961: 32.
Odhner, 1963: 50. MacFarland, 1966: 241.
Sphaerostoma tetraquetra (Pallas). O'Dono-
ghue, 1924: 3. O'Donoghue, 1926: 204.
LaRocque, 1953: 261.
Tritoniopsis tetraquetra (Pallas). Odhner,
1926a: 32, 33. Odhner, 1936: 1080. Mar-
cus, 1961: 32. Odhner, 1963: 50. Stein-
berg, 1963b: 71.
Tritoniopsis gigantea (Bergh). Odhner, 1936:
1080.
Tritoniopsilla tetraquetra (Pallas). Baba,
1937b: 312.
Duvaucelia tetraquetra (Pallas). Baba, 1937a:
197, 199. Smith 8 Gordon, 1948: 180.
MacFarland, 1966: 207, 208-218 ff, pl. 30,
figs. 3-8; pl. 39, figs. 8-10; pl. 43, figs. 1-
9; pl. 44, fig. 1; pl. 45, figs. 1-5. Roller,
1970a: 372. Bernard, 1970: 85.
Tritoniopsis aurantia Mattox, 1955: 8-13, pls.
214 MCDONALD
4-5. Marcus, 1961: 32. Lance, 1961: 67.
Sphon & Lance, 1968: 81.
Tochuina tetraquetra (Pallas). Odhner, 1963:
50. Sphon & Lance, 1968: 81. Baba,
1969b: 134. Roller 8 Long, 1969: 429.
Roller, 1970a: 372. Thompson, 1971: 334—
335, text fig. 1. Wicksten & DeMartini, 1973:
195. Abbott, 1974: 369. Birkeland, 1974:
212. Belcik, 1975: 276. Lambert, 1976:
297. Thompson 4 Brown, 1976: 2. Thomp-
son, 1976a: 3. Thompson, 1976b: 6, 7, 91.
McDonald 8 Nybakken, 1978: 113.
Trochuina tetraquetra (Pallas). Baba, 1968a:
257-258, text figs. 1-5 (lapsus).
Tritonia (Tochuina) tetraquetra (Pallas). Ro-
billiard, 1969a: 290.
Type-specimens: not listed.
Dorsum bears numerous white tipped tu-
bercles. Foot nearly as broad as dorsum and
rather linear, bilabiate and rather truncate an-
teriorly, and slightly tapered posteriorly to
short bluntly rounded tail; margins project
laterally beyond sides of body. Upper surface
of foot well set off from sides of body, or-
ange-yellow to yellowish-brown, slightly light-
er than dorsum and bears white tipped tu-
bercles; sole of foot light salmon pink to
yellow, dorsal margin edged with a band of
white. Antero-dorsal surface expanded into
broad frontal veil which extends laterally as
thin, rounded lobes; dorsal surface bears a
few white tipped tubercles, and is a slightly
lighter shade of color of dorsum, ventral sur-
face a slightly darker shade of foot color. La-
bial tentacles are almost obscure, located at
outer angle of frontal veil, they are short and
externally grooved. Rhinophores bear 6-10
short, vertical, bi- to tripinnate plumose pro-
cesses which encircle shaft below tip, pos-
terior-most process adnate to shaft, others
free. The tapered rhinophore shaft termi-
nates distally in a blunt, cylindrical tip. Rhino-
phores retractile into high sheaths with thick
walls and thin, nearly smooth margins; outer
surface of sheath tuberculate, antero-lateral
margins cleft. A single, irregular series of
white, bi- and tripinnate plumose branchial
appendages occurs along undulating body
margins on either side, from the area of rhi-
nophores, posteriorly to tip of foot. General
ground color orange-yellow to brownish-yel-
low or almost gray-yellow. T.L.: 120 mm, but
specimens over 300 mm have been collect-
ed.
Anus on right side, close beneath dorso-
lateral margin, posterior of genital aperture.
Radular formula of specimen examined
64(162-212.1.1.1.162-212), MacFarland
(1966: 213) reports 50-94(225-312.1.225-
312). Rachidian teeth (Fig. 87e) slightly elon-
gated, with a blunt cusp posteriorly and
deeply notched anteriorly. Laterals (Fig. 87d)
somewhat elongated. Marginals (Fig. 87a-c)
somewhat hooked.
Type-locality: Kuril Islands, U.S.S.R.
Range and habitat: Unalaska, Alaska
(Bergh, 1879b), to Santa Catalina Island,
California (Mattox, 1955); northern Japan
(Baba, 1969b), to Kuril Islands. Subtidal to
363 m, usually in areas of rocky or mud and
cobble bottom. Feeds upon the alcyonacean
Gersemia rubiformis (Wicksten 8 DeMartini,
1973: 195), and the pennatulacean Ptilosar-
cus guerneyi (Birkeland, 1974: 212; Thomp-
son, 1971: 335).
Remarks: Bergh (1879c: 154) quotes Pal-
las as saying he obtained the animal from the
Kuril Islands, ‘‘where the inhabitants eat it,
raw or cooked, and where it is known by the
name of Tochni.”
Trapania Pruvot-Fol, 1931
Body limaciform, elongate, smooth, and
arched dorsally. Anterior foot corners ten-
taculiform. Labial tentacles rather long and
digitiform. Rhinophores non-retractile and
perfoliate; at base of each is a digitiform ex-
tra-rhinophoral appendage. Branchial plumes
non-retractile, arranged around anus. On
either side, lateral of branchial plumes, oc-
curs a digitiform extra-branchial appendage.
Labial disc armed with denticles. Radular
formula 0.1.0.1.0, laterals rather broad with
a denticulate margin.
Penis armed with spines.
Type-species: Trapania fusca (Lafont,
1874).
Trapania velox (Cockerell, 1901)
Thecacera velox Cockerell, 1901a: 87. Kel-
sey, 1907: 50. Cockerell, 1908: 106.
O'Donoghue, 1922d: 139-140. O'Dono-
ghue, 1926: 217. O'Donoghue, 1927a: 11.
MacFarland, 1931: 32.
Drepania velox (Cockerell). MacFarland,
1929: 487-496, pl. 35, figs. 1-15. Baba,
1935a: 337-338.
Drepanida velox (Cockerell). MacFarland,
1931232:
Trapania velox (Cockerell). Marcus, 1957:
443. Lance, 1961: 67. Steinberg, 1963b:
CALIFORNIA NUDIBRANCHS 215
71. Paine, 1963a: 4. Lance, 1966: 78.
MacFarland, 1966: 127-129, pl. 20, figs.
1-3; pl. 32, figs. 17-24. Sphon & Lance,
1968: 81. Roller 8 Long, 1969: 428. Kress,
1970: 115. Marcus, 1972b: 297. Abbott,
1974: 364, text fig. 4340.
Type-specimens: not listed.
Foot elongate, linear, and set off from body
by a narrow, thin ridge; anterior margin
broadly emarginate, bearing a series of small,
anteriorly directed, whitish papillae. Foot cor-
ners produced into long, tapered, blunt ten-
taculiform processes which are whitish and
are slightly grooved ventrally. Labial tenta-
cles cylindrical, long, and tapered to blunt tips;
with a broad, medial band of cadmium yellow
to orange, leaving tip and base white. Rhi-
nophores bear 10-12 lamellae; clavus cad-
mium yellow to orange on distal half and
whitish on proximal half and on shaft. On ex-
ternal base of rhinophore shaft is a horizon-
tal, cylindrical, bluntly pointed extra-rhino-
phoral appendage with a subterminal ring of
cadmium yellow to orange and a wide, oblique
band of dark brown just proximal of subter-
minal ring. Branchial plumes 3, uni- and bipin-
nate, whitish with cadmium yellow to orange
distally, and with a dark brown, rather oval
spot on outer basal surface of rachis of each
plume. A horizontal digitiform extra-branchial
appendage occurs immediately antero-later-
ally of branchial plumes on either side; distal
third of each is cadmium yellow to orange,
leaving extreme tip white, a dark brown line
extends along dorsal surface of proximal half
of each appendage. General ground color
translucent grayish-white. A narrow, median
dark brown line extends posteriorly from
frontal margin, between rhinophores to an-
terior of branchial plumes, where it joins the
two lines from the extra-branchial append-
ages. A similar line extends from posterior of
branchial plumes to tip of tail which bears a
subterminal band of yellow, the extreme tip
white. Another similar dorso-lateral line ex-
tends longitudinally on either side from just
posterior of rhinophores nearly to tip of tail,
being interrupted just ventral of branchial
plumes. All of these brown lines vary in width
and continuity. T.L.: 15 mm.
Penis armed with curved spines.
Radular formula of specimen examined
22(0.1.0.1.0), MacFarland (1966: 128) re-
ports 24(0.1.0.1.0). Laterals (Fig. 42) some-
what convex, bearing 8-24 irregular denti-
cles along one edge, usually with one denticle
considerably longer than the others.
Type-locality: La Jolla, San Diego Co., Cal-
ifornia.
Range and habitat: Hazard Canyon, San
Luis Obispo Co., California (Roller & Long,
1969), to San Diego, San Diego Co., Califor-
nia (Lance, 1961). Intertidal to 6 m, usually
found in rocky intertidal and subtidally on a
white sponge on pier pilings. Quite rare in
California.
Remarks: Cockerell (1901a: 87) mentions
that this species swims with an undulating
motion on the surface of the water.
Triopha Bergh, 1880
Body limaciform and elongate. An indis-
tinct dorso-lateral pallial ridge on either side
of dorsum bears a number of irregular (tu-
berculate or short-branched) processes.
Frontal margin somewhat expanded into a
veil-like process, the margin of which bears
processes similar to those on pallial ridge.
Labial tentacles short, blunt, and auriform,
usually with a longitudinal groove on distal
half. Rhinophores perfoliate, retractile into
rather prominent sheaths. Branchial plumes
non-retractile, usually tripinnate, arranged
around anus, usually five in number.
Mandibular plates triangular, with short,
closely-set rodlets. Radular formula n.n.2-
4.n.n. Rachidian plates spurious. Laterals
hooked; marginals quadrilateral.
Penis armed with minute hooks.
Type-species: Triopha catalinae (Cooper,
1863).
Triopha catalinae (Cooper, 1863)
Triopa catalinae Cooper, 1863b: 59. Carpen-
ter, 1864: 609. Cooper, 1870: 56. Abra-
ham, 1877: 230. Ferreira, 1977: 388.
Triopa carpenteri Stearns, 1873a: 78, fig. 2.
Stearns, 1873b: 209, fig. 2. Abraham,
1877: 230. Ferreira, 1977: 388.
Triopha carpenteri (Stearns). Bergh, 1880a:
262. Bergh, 1880b: 112. Tryon, 1883: 376.
Bergh, 1892: 1140 (146). Bergh, 1894: 184.
MacFarland, 1905: 48-49. Cockerell &
Eliot, 1905: 43. MacFarland, 1906: 135-
137, РГ. 19) 195-5155: pl. 21, 195. 108,
113; pl. 27, figs. 16-17. Berry, 1907: 35.
Cockerell, 1908: 107. Cockerell, 1915:
228-229. O'Donoghue, 1921: 167. O'Don-
oghue, 1922b: 164. O'Donoghue, 1922d:
136-138. O'Donoghue, 1926: 214, 238.
216 MCDONALD
O'Donoghue, 1927b: 96-97, pl. 2, figs. 45-
47. Hewatt, 1937: 200. Costello, 1938: 321
ff; tabs. 13, 5 plis: 11721; pl 2,
figs. 26, 31, 38-40. Hewatt, 1946: 195,
198. Smith & Gordon, 1948: 180. Baba,
1957: 8, 11, text figs. 1, 2A. Marcus, 1961:
22-23, 57, pl. 4, fig. 71. Thompson, 1961:
235. Steinberg, 1961: 60. Lance, 1961: 66.
McLean, 1962: 110. Paine, 1963a: 4.
Steinberg, 1963b: 70. Willows, 1965: 707
ff. MacFarland, 1966: 106-109, 115, pl. 19,
figs. 3-4; pl. 29, figs. 4-6; pl. 31, figs. 13-
18. Ghiselin, 1966: 333, 345. Hurst, 1967:
255 ff, pl. 28, fig. 12; fig. 24-9. Buchsbaum
& Milne, 1967: pl. 67. Sphon & Lance,
1968: 81. Roller & Long, 1969: 428. Had-
erlie, 1969: tab. 2. Turner et al., 1969: 137-
138. Long, 1969c: 232. Robilliard, 1969a:
290. Gosliner & Williams, 1970: 179.
McBeth, 1970: 28. Bernard, 1970: 85.
Thompson, 1971: 334. McBeth, 1971: 158,
159. Bertsch et al., 1972: 308. Sphon,
1972a: 156. McBeth, 1972a: 55 ff. Mc-
Beth, 1972b: 69. Thompson & Bebbing-
ton, 1973: 148, pl. 14, figs. a-b. Harris,
1973: 240, 281, 287. Abbott, 1974: 360,
pl. 17, 4295. Robilliard, 1974b: 990. Birke-
land, 1974: 218. Belcik, 1975: 276. Lam-
bert, 1976: 297. Thompson, 1976a: pl. 2,
fig. a. Thompson, 1976b: 37, 70. Nybak-
ken & Eastman, 1977: 279-289. Ferreira,
1977: 387-391. O'Clair, 1977: 443. Ny-
bakken, 1978: 134 ff. Haderlie & Donat,
1978: 60. McDonald & Nybakken, 1978:
111. Fuhrman et al., 1979: 292. Dickinson,
1979: 277 ff, text figs. 1A-B. Russo, 1979:
48.
Triopha modesta Bergh, 1880a: 261-266, pl.
14, figs. 17-20; pl. 15, figs. 1-11. Bergh,
1880b: 113-117, pl. 14, figs. 17-20; pl. 15,
figs. 1-11. Bergh, 1892: 1140 (146). Bergh,
1894: 184-187, pl. 7, fig. 29; pl. 8, figs. 2-
12; pl. 9, figs. 1-12. MacFarland, 1905: 49.
Cockerell & Eliot, 1905: 43. MacFarland,
1906: 137, 141. O'Donoghue, 1922d: 137-
138. O'Donoghue, 1926: 215. Thiele, 1931:
424. LaRocque, 1953: 258. Marcus, 1961:
22-23. MacFarland, 1966: 106, 109, 115.
Ferreira, 1977: 388, 390.
Triopa modesta. Fischer, 1887: 527.
Triopha catalinae (Cooper). Cockerell, 1915:
229. O'Donoghue, 1922d: 138. O'Dono-
ghue, 1926: 214. Smith & Gordon, 1948:
180. Marcus, 1961: 23, 56. Steinberg,
1961: 60. Lance, 1961: 66. Abbott, 1974:
360. Ferreira, 1977: 388, 389-396, figs. 1-
11, 16. Bertsch, 1977: 109.
Triopha scrippsiana Cockerell, 1915: 228-
229. O'Donoghue, 1922d: 137-138.
O'Donoghue, 1926: 215. Marcus, 1961: 23.
Steinberg, 1961: 60. Lance, 1961: 66. Ab-
bott, 1974: 360. Ferreira, 1977: 388, 390.
Triopha elioti O'Donoghue, 1921: 165-167.
Marcus, 1961: 23. Steinberg, 1961: 60.
Steinberg, 1963b: 70. MacFarland, 1966:
115. Ferreira, 1977: 388, 390.
Type-specimens: not listed, State Coll.
species 1002 (Cooper, 1863b).
Pallial ridge bears 4-9 slightly prolonged,
large orange tubercles which bear very small
branches or tubercles. Dorsum, dorsal sur-
face of tail, and sides of body bear a number
of irregular, orange tubercles and blotches of
various sizes. Margin of frontal veil bears 7-
16 or more irregularly lobed, orange process-
es. Foot rounded anteriorly and elongate with
nearly parallel margins extending to abruptly
pointed tail. Rhinophores bear 20-30 lamel-
lae, clavus inclined slightly posteriorly, or-
ange, shaft translucent grayish-white. Rhi-
nophore sheaths bear thin, smooth margins.
Branchial plumes 5, tripinnate, translucent
grayish-white with orange tips. General
ground color translucent grayish-white, very
large specimens may occasionally be quite
grayish. T.L.: 40 mm.
Radular formula of specimen examined
45(12-14.14.4.14.12-14), MacFarland (1966:
107) reports 33(9-14.9-18.4.9-18.9-14), and
Marcus (1961: 23) reports 29-33(9-20.9-
18.4.9-18.9-20). Rachidian plates (Fig. 46k-
|) very rudimentary and thin, innermost quad-
rangular, outer triangular. Laterals (Fig. 46h-
j) strongly hooked, marginals (Fig. 46a-g)
quadrilateral.
Type-locality: Santa Catalina Island, Cali-
fornia.
Range and habitat: Auke Bay, Alaska
(Robilliard, 1974b), to San Diego, San Diego
Co., California (Lance, 1961); Japan (Baba,
1957). Intertidal to 80 m. Common in rocky
intertidal and subtidal areas, occasional on
floating docks and pilings in bays. Feeds upon
the bryozoans: Bugula mollis, Caulibugula
ciliata, Cauloramphus spiniferum, Cellaria
mandibulata, Crisia occidentalis, Crisia ser-
rulata, Dendrobeania laxa, Filicrisia francis-
cana, Scrupocellaria californica, Scrupocel-
laria diegensis, and Tricellaria sp. (McBeth,
1971; Nybakken & Eastman, 1977).
Triopha maculata MacFarland, 1905
Triopha maculata MacFarland, 1905: 49.
Cockerell & Eliot, 1905: 43. MacFarland,
1906: 137-139, pl. 19, figs. 55a-59; pl. 21,
CALIFORNIA NUDIBRANCHS 217.
figs. 106-107; pl. 28, fig. 18. Berry, 1907:
35. Baily, 1907: 92. Cockerell, 1915: 229.
O'Donoghue, 1922b: 165. O’Donoghue,
1922d: 137-138. O'Donoghue, 1926: 214-
215. O'Donoghue, 1927b: 98-99, pl. 2, figs.
51-53. Hewatt, 1937: 200. Costello, 1938:
321 ff, tabs. 1-3, 5. Smith 8 Gordon, 1948:
180. Marcus, 1961: 23-24, pl. 4, figs. 72-
76. Steinberg, 1961: 61. Lance, 1961: 66.
Farmer 8 Collier, 1963: 62. Paine, 1963a:
4. Steinberg, 1963b: 70. Paine, 1964: 385.
Paine, 1965: 607. MacFarland, 1966: 109-
112, 115, pl. 19, figs. 5-6; pl. 31, figs. 19-
21. Farmer, 1967: 341. Sphon 8 Lance,
1968: 81. Roller 8 Long, 1969: 428. Gos-
liner & Williams, 1970: 180. North, 1971:
58. Bertsch et al., 1972: 308, tabs. 1-5.
Harris, 1973: 281. Abbott, 1974: 360, pl.
17, fig. 4296. Ferreira, 1977: 396-400, figs.
12-15, 17-18 (partim). Bertsch, 1977: 109.
Bloom & Bloom, 1977: 296. Nybakken,
1978: 134 ff. Haderlie & Donat, 1978: 60.
McDonald 8 Nybakken, 1978: 112. Fuhr-
man et al., 1979: 292. Russo, 1979: 48.
Triopha aurantiaca Cockerell, 1908: 107.
Cockerell, 1915: 229. O'Donoghue, 1922d:
137-138. O'Donoghue, 1926: 214. Fraser,
1932: 67. LaRocque, 1953: 258. Marcus,
1961: 115. Steinberg, 1961: 60-61. Lance,
1961: 66. MacFarland, 1966: 115. Ber-
nard, 1970: 85. Abbott, 1974: 360. Fer-
reira, 1977: 387, 388, 396.
Triopha sp. Cockerell 8 Eliot, 1905: 42-43.
Sphon & Lance, 1968: 81. Roller 8 Long,
1969: 428. Bertsch et al., 1972: 308, tabs.
2, 4, 5. Ferreira, 1977: 387, 396.
Type-specimens: U. S. Nat. Mus., no.
181276.
Pallial ridge bears 4-6 orange to vermilion,
slightly prolonged processes which are slightly
branched or bear tubercles distally. Margin of
frontal veil bears 10-12 stout processes
which bear a few tubercles distally and are
orange. Foot bluntly rounded anteriorly, elon-
gate with nearly parallel margins extending to
abruptly pointed tail. Rhinophores bear about
16-18 lamellae, clavus inclined slightly pos-
teriorly, orange; shaft slightly lighter orange.
Rhinophore sheaths bear thin, smooth mar-
gins. Branchial plumes 5-7, tripinnate, or-
ange, tipped with darker orange to vermilion.
General ground color varies from clear, light
orange in young specimens to darker orange
or yellow brown in larger specimens. In larger
specimens, dorsum bears numerous round to
oval, pale blue spots and more numerous
small, dark brown dots. T.L.: 15 mm.
Radular formula of specimen examined
14(8-10.1-5.4.1-5.8-10), MacFarland (1966:
110) reports 14(7-8.4-5.4.4-5.7-8), and
Marcus (1961: 24) reports 13-17(7-10.3-
5.4.3-5.7-10). Rachidian plates (Fig. 48i-j)
very rudimentary and thin. Laterals (Fig. 48f-
h) strongly hooked. Marginals (Fig. 48a-e)
quadrangular.
Type-locality: Monterey Bay, California.
Range and habitat: Bodega Bay, Sonoma
Co., California (Marcus, 1961), to Ensenada,
Baja California, Mexico (Farmer & Collier,
1963). Intertidal to 28 m, quite common in
rocky intertidal areas in central California.
Nybakken & Eastman (1977: 288) report that
it feeds upon the bryozoans: Bugula mollis,
Caulibugula ciliata, Cauloramphus spinife-
rum, Crisia occidentalis, Dendrobeania laxa,
Filicrisia franciscana, Membranipora mem-
branacea, Scrupocellaria californica, and Tri-
cellaria sp.
Triopha occidentalis (Fewkes, 1889)
Cabrilla occidentalis Fewkes, 1889: 140-141,
fig. O'Donoghue, 1926: 215. Steinberg,
1961: 60. Ferreira, 1977: 389.
Triopha grandis MacFarland, 1905: 50.
Cockerell & Eliot, 1905: 43. MacFarland,
1906: 139-141, pl. 19, figs. 60-64; pl. 28,
fig. 19. Cockerell, 1915: 229. O'Donoghue,
1922b: 165. O'Donoghue, 1922d: 137-138.
O’Donoghue, 1926: 214. O'Donoghue,
1927b: 97-98, pl. 2, figs. 48-50. Costello,
1938: 321, 325, tabs. 1—3. 5, pl. 1, fig. 7.
Smith & Gordon, 1948: 180. Lance, 1961:
66. Marcus, 1961: 23. Steinberg, 1963b:
70. MacFarland, 1966: 112-115, pl. 19,
figs. 1-2; pl. 31, figs. 22-26. Farmer, 1967:
341. Sphon & Lance, 1968: 81. Haderlie,
1968: 333, 338. Haderlie, 1969: tabs. 1, 2.
Roller & Long, 1969: 428. Gosliner & Wil-
liams, 1970: 180. Mulliner, 1972a: 38. Mul-
liner, 1972b: 3, figs. Abbott, 1974: 360, pl.
17, fig. 4297. Ferreira, 1977: 387, 388, 396,
397.
Triopha maculata MacFarland. Ferreira, 1977:
396-400, figs. 12-15, 17-18 (partim).
Type-specimens: not listed.
Pallial ridge bears 4-6 irregular, prolonged
processes which bear small branches or tu-
bercles, and are yellow-orange to vermilion.
Margin of frontal veil bears 7-12 tuberculate
or branched, yellow-orange to vermilion or
pale burnt sienna processes. Foot rounded
anteriorly, elongate with nearly parallel mar-
gins extending to abruptly pointed tail. Rhi-
nophores bear 15-20 lamellae, clavus in-
218 MCDONALD
clined slightly posteriorly and is yellow-or-
ange to vermilion, shaft slightly paler. Rhi-
nophore sheaths bear thin, smooth, slightly
flared margins. Branchial plumes 5-6, tri- and
quadripinnate, pale yellow-orange to almost
white with yellow-orange to vermilion tips.
General ground color varies from pale yellow-
ochre to dark yellow-brown, with yellow or-
ange being most frequent. Dorsum bears nu-
merous light powder blue spots which may
be almost white, or extremely faint in some
specimens: T.L.: 40 mm.
Radular formula of specimen examined
11(2-8.2-4.2.2-4.2-8), MacFarland (1966:
113) reports 18(8.7.2.7.8). Rachidian plates
(Fig. 47k-I) extremely rudimentary, thin, and
squarish. Laterals (Fig. 47i-j) strongly hooked.
Marginals (Fig. 47a—h) rather quadrilateral.
Type-locality: Prisoner's Harbor, Santa
Cruz Island, California.
Range and habitat: Limantour Estero,
Drakes Bay, Marin Co., California (Gosliner
& Williams, 1970), to San Quintín, Baja Cali-
fornia, Mexico (Farmer, 1967). Intertidal to 30
m, occasional on floating docks in bays, in
association with the alga Macrocystis pyri-
fera; also in offshore kelp (M. pyrifera and
Nereocystis luetkeana) beds.
Remarks: Ferreira (1977: 396) considered
Triopha grandis a junior synonym of Triopha
maculata. In reviewing his reasons for doing
so, | feel that the evidence which he presents
is insufficient to synonymize the two species.
Especially since | have seen juveniles and
adults of both species and feel that they are
reasonably easily separable. The correlation
between the number of hooks vs. number of
plates per half-row of radula (Ferreira, 1977:
399) does not seem sufficient reason to con-
sider the two species synonymous. Further
study is certainly necessary to resolve the
question.
Fewkes (1889: 139-141) described the new
genus and species Cabrilla occidentalis on the
basis of a single specimen from Santa Cruz
Island, California. Although a number of au-
thors (O'Donoghue, 1926; Steinberg, 1961;
Ferreira, 1977) have felt that the description
is very poor and therefore difficult to com-
pare to known species, it seems that the de-
scription corresponds very closely to that of
Triopha grandis MacFarland, 1905. The body
color of C. occidentalis is greenish-brown
covered with light green spots (Fewkes, 1889:
140) while that of T. grandis varies from pale
yellow-ochre to dark yellow-brown with light
blue spots (MacFarland, 1966: 113). C. oc-
cidentalis has 4 large processes on either
edge of the dorsum anterior of the branchial
plumes and 2 smaller ones posterior of the
branchial plumes, while 7. grandis has 4-6
processes on either side of the dorsum
(MacFarland, 1966: 112). The figure of C. oc-
cidentalis (Fewkes, 1889: 140) shows 7 or 8
velar processes, and T. grandis has 8-12
processes. Fewkes (1889: 140) states that
С. occidentalis has ‘‘lens-like bodies” on the
apex of the dorso-lateral processes, Mac-
Farland (1966: 112) mentions that T. grandis
has a dim white mass at the apex of each
appendage “probably the ‘lens’ of Fewkes.”’
The number of branchial plumes in C. occi-
dentalis is not mentioned by Fewkes; he says
simply: ‘The branchiae are stellate, bipin-
nate, consisting of primary arms and lateral
branches, of white color, transparent.” T.
grandis has 5-6 pale yellow-orange to white,
tri- to quadripinnate branchial plumes. The
bipinnate branchial plumes and their lateral
branches (of C. occidentalis) may be consid-
ered as tri- or quadripinnate plumes. The type-
locality of C. occidentalis is within the range
known for T. grandis, and C. occidentalis was
found on the anchor of a buoy, while T. gran-
dis is most often found on buoys, floating
docks, or in kelp holdfasts. These facts seem
to indicate that C. occidentalis is conspecific
with T. grandis, in which case the latter is a
junior subjective synonym of the former, the
proper name being Triopha occidentalis
(Fewkes, 1889) by the law of priority. Fur-
ther, specimens have been collected which
match the description of C. occidentalis much
more closely than that of T. grandis (i.e. they
are brownish with light green spots and the
rhinophores are brownish distally, as de-
scribed for C. occidentalis). It therefore seems
certain that C. occidentalis is a valid species,
and although some may consider it distinct
from T. grandis, | consider them conspecific.
Tritonia Cuvier, 1798
Body limaciform, rather subquadrilateral,
and somewhat depressed dorso-ventrally.
Branchial appendages plumose and ar-
ranged in a longitudinal series, dorso-laterally
on either side of dorsum. Dorsum smooth or
bears small tubercles. Foot broad. Antero-
dorsal surface expanded into broad frontal
veil which extends laterally and may be
rounded or bilobed and usually bears digiti-
form processes. Rhinophores retractile into
high sheaths, and bear a few plumose pro-
cesses distally.
CALIFORNIA NUDIBRANCHS 219
Masticatory border of mandibles denticu-
late. Radular formula n.1.1.1.n. Rachidian
teeth bear a median cusp and are broader
than in Tochuina. Laterals short, blunt hooks;
marginals hamate.
Penis unarmed. Anus on right side, be-
neath margin of dorsum.
Type-species: Tritonia hombergi Cuvier,
1803 (see ICZN, 1963, Opinion 668).
Tritonia diomedea Bergh, 1894
Tritonia diomedea Bergh, 1894: 146-150, pl.
2, figs. 10-11; pl. 3, figs. 6-10; pl. 4, figs.
1-5. Cockerell & Eliot, 1905: 33. Marcus,
1959: 68. Marcus, 1961: 57. Steinberg,
1963b: 71. Sakharov, 1966: 957. Dorsett,
1967: 140. Michelson, 1970: 108. Thomp-
son, 1971: 335-336, text fig. 2. Sanchis &
Castro, 1972: 181. Dorsett et al., 1973:
287. Thompson, 1976a: 23, pl. 5, fig. d.
Thompson, 1976b: 55, 59. Kempf & Wil-
lows, 1977: 261 ff. Getting, 1977: 325 ff.
Bonar, 1978: 178. McDonald & Nybakken,
1978: 113. Harrigan & Alkon, 1978b: 299.
Taghert & Willows, 1978: 253 ff. Audesirk
& Audesirk, 1979: 79 ff, text figs. 2A-F.
Dickinson, 1979: 278, text fig. 1D. Bulloch
8 Dorsett, 1979: 20. Strathmann & Leise,
1979: 524 ff. Lennard et al., 1980: 165 ff.
Getting et al., 1980: 151 ff.
Tritonia exsulans Bergh, 1894: 150-152, pl.
3, figs. 11-12; pl. 4, fig. 6. Cockerell 8 Eliot,
1905: 33. O’Donoghue, 1921: 152-154, pl.
1 (7), figs. 4-6. Baba, 1937b: 310-312, text
fig. E. Baba, 1957: 9. Marcus, 1959: 68.
Marcus, 1961: 32-33, 56, 57, pl. 6, figs.
115-118. Lance, 1961: 67. Steinberg,
1963b: 71. Willows, 1965: 707 ff. Hurst,
1967: 255 ff, text figs. 18a-d, pl. 31, fig.
24; pl. 37, fig. 48; fig. 24-14. Marcus &
Marcus, 1967a: 123. Sphon 8 Lance, 1968:
81. Willows, 1968: 242. Roller 8 Long,
1969: 428. Roller, 1970a: 372. Thompson,
1971: 336-337. Keen, 1971: 834. Bertsch
et al., 1972: 308. Holleman, 1972a: 60.
Goddard, 1973: 10. Gosliner 8 Williams,
1973b: 354. Abbott, 1974: 368. Willows 8
Dorsett, 1975: 118.
Tritonia diomedia Bergh. O'Donoghue, 1921:
151-152, pl. 1 (7), figs. 1-3. Volodchenko,
1955: 249, pl. 48, fig. 3. Veprintsev et al.,
1964: 352, 354. Borovyagin et al., 1965:
104 ff. Sakharov et al., 1965: 660 ff. Sa-
kharov, 1965: 365. Borovyagin 8 Sakha-
rov, 1965a: 642 ff. Borovyagin & Sakharov,
1965b: 458, 463. Gerasimov 8 Magura,
1965: 360, 363. Antonov et al., 1965: 1200.
Manokhina, 1966: 93 ff. Turpayev et al.,
1967: 619, 621. Dorsett, 1967: 140. Sa-
kharov 8 Turpaev, 1968: 305. Borovyagin
8 Sakharov, 1968: 3 ff. Kostyuk, 1968:
146. Bezruchko et al., 1969: 1107 ff.
Bezruchko et al., 1970: 1073 ff. Bezruchko
et al., 1971: 460 ff. Minichev, 1971: 282,
284. Willows, 1971: 69. Abraham 8 Wil-
lows, 1971: 271, 272. Junge, 1972: 975.
Krasts 8 Veprintsev, 1972: 289-290. Ad-
zhimolaev et al., 1972: 79 ff. McCaman et
al., 1973: 129. Willows et al., 1973a: 207.
Willows et al., 1973b: 255 ff. Willows et al.,
1973c: 461. Faugier & Willows, 1973: 244.
Adzhimolaev et al., 1973a: 60. Adzhimo-
laev et al., 1973b: 129. Getting & Willows,
1973: 424. Hoyle 8 Willows, 1973: 239,
240. Willows, 1973b: 200. Weinreich et al.,
1973: 970 ff. Adzhimolaev et al., 1973c:
131 ff. Getting, 1974: 846. Chase, 1974a:
707. Chase, 1974b: 721. Getting 8 Wil-
lows, 1974: 858. Grieshaber et al., 1974:
168. Willows 8 Dorsett, 1975: 117 ff. Zack,
1975b: 239. Chase, 1975: 37. Getting,
1975: 128. Dorsett, 1975: 292, 309. Par-
tridge, 1975: 161 ff. Willows, 1976: 329 ff.
Getting 1976: 271 ff. Hoyle, 1976: 33, fig.
2. Kuz'min et al., 1976: 231. Thompson,
1977: 465 ff. Tomosky-Sykes et al., 1977:
99. Willows, 1978: 155 ff. Audesirk, 1978a:
259 ff. Audesirk, 1978b: 541 ff. Audesirk,
1979: 71 ff. Audesirk, McCaman 8 Wil-
lows, 1979: 87 ff. Gakhova et al., 1979:
313 ff. McCaman, Ono 8 McCaman, 1979:
1111 ff (lapsus).
Sphaerostoma diomedia (Bergh). O’Dono-
ghue, 1924: 3. LaRocque, 1953: 261 (/ap-
sus).
Sphaerostoma exsulans (Bergh). O'Dono-
ghue, 1924: 3. O'Donoghue, 1926: 204.
LaRocque, 1953: 261.
Sphaerostoma diomedea (Bergh). O'Dono-
ghue, 1926: 204.
Duvaucelia diomedea (Bergh). Odhner, 1926a:
35. MacFarland, 1966: 241 ff. Bernard,
1970: 85.
Duvaucelia exsulans (Bergh) . Odhner, 1926a:
35. Baba, 1937a: 199. Smith & Gordon,
1948: 180. MacFarland, 1966: 212, 223,
226-235 ff, pl. 30, figs. 9-10; pl. 39, fig. 7;
pl. 43, figs. 20-26; pl. 44, figs. 3-4. Roller,
1970a: 372. Bernard, 1970: 85.
Duvaucelia (Duvaucelia) exsulans (Bergh).
Baba, 1937b: 310-312, text fig. 9. Baba,
1937d: 391. Baba, 1957: 9.
Duvaucelia (Duvaucelia) diomedea (Bergh).
Baba, 1937d: 391.
220 MCDONALD
Duvaucelia ( Duvaucelia) septemtrionalis
Baba, 1937d: 391-392, text figs. 1А-Е.
Duvaucelia gilberti MacFarland, 1966: 223,
224, 235-243, pl. 30, figs. 1-2; pl. 43, figs.
27-36; pl. 44, fig. 5; pl. 45, fig. 6. Roller,
1970a: 372.
Tritonia gilberti (MacFarland). Willows, 1967a:
570, 571. Willows, 1967b: 796. Willows &
Hoyle, 1967: 327. Willows, 1968: 219, 242.
Mauzey et al., 1968: 609. Willows & Hoyle,
1968: 443. Willows & Hoyle, 1969: 1549.
Dorsett et al., 1969: 711. Robilliard, 1969a:
290. Roller, 1970a: 372. Konishi, 1971: 60.
Rose, 1971: 185. Harris, 1973: 228 ff.
Birkeland, 1974: 212.
Tritonia primorjensis Minichev, 1971: 282-
284, text figs. 1-5.
Tritonia (Duvaucelia) diomedea (Bergh). Ab-
bott, 1974: 368.
Tritonia (Duvaucelia) gilberti MacFarland. Ab-
bott, 1974: 369.
Tritonia esculans. Willows, 1976: 349 (lap-
sus).
Tritonia diamedia. Audesirk, 1978b: 541 ff
(lapsus).
Type-specimens: not listed.
Dorsum covered with numerous, very small
tubercles. Foot somewhat broader than dor-
sum and rather linear, rounded and bilabiate
anteriorly and slightly tapered posteriorly to
short, rather blunt tail. Foot margins but
slightly set off from body. Margin of slightly
bilobed frontal veil bears about 10-30 light
pink to almost white, digitiform processes;
lateral processes somewhat longer than me-
dial processes, lateral-most process on either
side bears a ventral longitudinal groove.
Rhinophores bear about 20 short, vertical,
bipinnate, plumose, very pale yellow to
brownish processes which encircle shaft be-
low tip, posterior processes adnate to shaft,
others free distally. The slightly tapered rhi-
nophore shaft terminates in a white, blunt tip.
Rhinophore sheaths bear thin, nearly smooth,
white margins. A single, irregular series of
very pale yellow to brownish, bi- to tripinnate
plumose branchial appendages occurs dor-
so-laterally on either side of dorsum, from just
posterior of rhinophores to tip of foot. Gen-
eral ground color pinkish, dorsum a deep
rose-pink, sides of body and frontal veil light-
er. A narrow white line borders edge of foot;
a similar line occurs on anterior edge of fron-
tal veil. An indistinct, broken line of white may
occur on dorso-lateral body margins, be-
tween branchial appendages. A white line oc-
curs vertically on inner surface of rhinophore
sheath and continues down onto frontal veil.
TES 150 мм.
Anus on right, close beneath dorso-lateral
margin, posterior of genital aperture.
Radula formula of specimen examined
69(87-95.1.1.1.87-95), MacFarland (1966:
229) reports 44-55(64-82.1.1.1.64-82), Baba
(1937b: 312) reports 65(75-95.1.1.1.75-95),
and O'Donoghue (1921: 153) reports 45-
52(60-64.1.1.1.60-64). Rachidian teeth (Fig.
88f) tricuspidate with median cusp longer than
lateral cusps. Marginals (Fig. 88a-d) hooked.
Type-locality: Shumagin Bay, Aleutian Is-
lands, Alaska.
Range and habitat: Shumagin Bay, Aleu-
tian Islands, Alaska (Bergh, 1894), to Bay of
Panama, Panama (Sphon, 1972b); Japan
(Baba, 1957); Okhotsk Sea (Volodchenko,
1955); Florida (Marcus, 1961). Intertidal to
656 m. Not uncommonly taken in bottom
trawls on mud-sand bottom in central Cali-
fornia. Feeds upon the pennatulaceans: Pti-
losarcus gurneyi and Virgullaria sp. (Audesirk
8 Audesirk, 1979; Birkeland, 1974; Thomp-
son, 1971; Willows, 1978).
Remarks: Thompson (1971: 333) reviewed
the family Tritoniidae from the North Ameri-
can Pacific coast and found that the only
possible character for separating Tritonia di-
omedea and Tritonia exsulans is the shape
of the penis. The difference in shape, which
Thompson observed, could easily be attrib-
uted to differences in preservation technique.
Since all other observable characters fail to
differentiate the two species, and since they
occur in the same area, they are here consid-
ered conspecific and the name Tritonia di-
omedea is used since it has priority by being
described on the pages immediately preced-
ing the description of Tritonia exsulans. Tri-
tonia palmeri Cooper, 1863, while possibly
synonymous with T. diomedea, has been de-
clared anomen dubium by Thompson (1971:
334) because the original description is so
poor. Minichev (1971) described Tritonia pri-
morjensis from the northern part of the Sea
of Japan, stating that it may be differentiated
from T. diomedea most of all by its narrower
radula and by the lobed edge of the rhino-
phore cavities and the spiny jaws. The radu-
lar formula for T. primorjensis is 50-75(89-
100.1.1.1.89-100) (Minichev, 1971: 282) and
the total body length is 8-18 cm (Minichev,
1971: 282), while Thompson (1971: 336)
gives the radula formula 65(89.1.89) for a 16
CALIFORNIA NUDIBRANCHS 221
cm specimen of T. diomedea. Minichev (1971:
283) states that the lobed edge of the rhino-
phore cavities distinguishes Т. primorjensis
from T. diomedea. However, MacFarland
(1966: 235) states that 7. gilberti (a junior
syonym of T. diomedea) has irregularly cren-
ulate rhinophore sheaths. The masticatory
border of the mandibles of T. primorjensis
bears plates which grade into spines (Mini-
chev, 1971: 282) while the mandibles of 7.
exsulans bear large conical denticles grading
to plate-like markings (MacFarland, 1966:
229). It is apparent that Minichev (1971)
compared his specimens only with the de-
scription of T. diomedea in Bergh (1894) and
not with any of the more recent descriptions
of specimens of T. diomedea. From the above
discussion it is obvious that T. primorjensis
is a junior subjective synonym of T. diome-
dea. In addition, Soviet workers have contin-
ued to use the name T. diomedea and have
not used 7. primorjensis at all. Baba (1937d)
described Duvaucelia (Duvaucelia) septem-
trionalis from the Sea of Okhotsk. He distin-
guished it from T. diomedea by its larger rad-
ular formula, 80(100-115.1.1.1.100-115), and
larger number of branchial plumes (29-30).
However, Bergh (1894: 148) gives the radu-
lar formula 73(140-150.1.1.1.140-150) in the
original description of T. diomedea. Thomp-
son (1971: 335) mentions 7-8 major gills and
20-24 smaller plumes on either side of the
dorsum. It is therefore obvious that T. sep-
temtrionalis does not differ significantly from
T. diomedea and is therefore a junior subjec-
tive synonym of T. diomedea.
Additional references which mention Tri-
tonia sp., and are referable to T. diomedea
are: Dorsett, 1976; Field & MacMillan, 1973;
Gulrajani & Roberge, 1978; Harth et al., 1975;
Kennedy et al., 1969; Lloyd, 1979; Mano-
khina 8 Kuz'mina, 1972; Martin & Manning,
1974; McCaman, McCaman, & Stetzler,
1979; Mellon, 1974; Pentreath, 1977; Pent-
reath & Barry, 1977; Plant, 1979; Salanki 8
Willows, 1978; Veprintsev et al., 1966; Ve-
printsev 8 D'yakanova, 1967; Willows, 1973a.
Tritonia festiva (Stearns, 1873)
Lateribranchiaea festiva Stearns, 1873a: 77-
78, text fig. 1. O'Donoghue, 1926: 218.
Costello, 1938: 321, 324, tabs. 1, 5.
Lateribranchiaia festiva Stearns. Stearns,
1873b: 209, text fig. 1 (lapsus).
Tritonia reticulata Bergh, 1881b: 239-250, pl.
8, figs. 7-20; pl. 9, figs. 1-12; pl. 10, figs.
1-10. Bergh, 1884b: 727. Baba, 1937b:
310. MacFarland, 1966: 241.
Sphaerostoma undulata O'Donoghue, 1924:
3-6, pl. 1, figs. 1-4. O'Donoghue, 1926:
205. LaRocque, 1953: 261. Steinberg,
1961: 62. Marcus, 1961: 57. MacFarland,
1966: 241.
Duvaucelia reticulata (Bergh). Odhner, 1927a:
34.
Duvaucelia undulata (O'Donoghue). Odhner,
1936: 1078. Baba, 1940: 107.
Duvaucelia (Duvaucelia) undulata var. muro-
ranica Baba, 1940: 106-107, text figs. 6A-
С.
Tritonia undulata (O'Donoghue). Marcus,
1959: 69.
Lateribranchia festiva Stearns. Steinberg,
1961: 61 (lapsus).
Sphaerostoma festiva (Stearns). Steinberg,
1961: 61.
Tritonia festiva (Stearns). Marcus, 1961: 31-
32, 57, pl. 6, figs. 109-114. Steinberg,
1961: 61-62. Lance, 1961: 67. Paine,
1963a: 4. Steinberg, 1963b: 71. Willows,
1965: 707 ff. Willows, 1968: 226. Baba,
1968b: 258-259, figs. 1-2. Sphon 8 Lance,
1968: 81. Baba, 1969a: 132-134, fig. 1.
Bertsch, 1969: 231. Roller 8 Long, 1969:
428. Long, 1969c: 232. Robilliard, 1969a:
290. Roller, 1970a: 372. Gosliner 8 Wil-
liams, 1970: 180. Thompson, 1971: 337-
338, text fig. 3. North, 1971: 58. Bertsch
et al., 1972: 308. Sphon, 1972a: 156.
Gomez, 1973: 163-165, text fig. 1. Harris,
1973: 305. Thompson, 1973: 167 ff, text
fig. 8, pl. 2, fig. F. Nybakken, 1974: 371.
Abbott, 1974: 368, text fig. 4366. Birke-
land, 1974: 211 ff. Lewbel 8 Lance, 1975:
346. Belcik, 1975: 276. Willows & Dorsett,
1975: 118. Lambert, 1976: 297, 299. Mich-
el, 1976: 47, fig. 5. Thompson, 1976a: 17,
text fig. 11b. Thompson, 1976b: 71. Ny-
bakken, 1978: 135. McDonald & Nybak-
ken, 1978: 110, 113. Robilliard & Barr,
1978: 153. Bertsch, 1980: 224.
Duvaucelia festiva (Stearns). MacFarland,
1966: 212, 218-226 ff, pl. 39, figs. 1-6; pl.
43, figs. 10-19; pl. 44, fig. 2; pl. 45, figs.
7-8. Grigg 4 Kiwala, 1970: 151. Bernard,
1970: 85.
Type-specimens: not listed.
Dorsum smooth, lacking tubercles. Foot
somewhat broader than dorsum, rather line-
ar; rounded and slightly bilabiate anteriorly
and tapered posteriorly to rather blunt tail.
222 McDONALD
Margin of slightly bilobed frontal veil bears
about 7-12 long, tapered, white processes;
lateral processes somewhat longer than me-
dial processes; lateral-most process on either
side bears a ventral longitudinal groove.
Rhinophores bear about 8-10 vertical, uni- to
bipinnate, plumose, yellowish processes
which encircle shaft below tip, posterior-most
process adnate to shaft, others free, at least
distally. Tapered rhinophore shaft terminates
distally in blunt, white tip. Rhinophore sheaths
bear thin, undulating, white margins. A sin-
gle, rather irregular, undulating series of about
11-15, yellowish, bi- to tripinnate, plumose
branchial appendages occurs dorso-laterally
on either side of dorsum, from just posterior
of rhinophores, posteriorly almost to tail.
General ground color varies from pure white
to cream, light cadmium yellow, or occasion-
ally, burnt sienna. A series of white, reticulate
lines and loops occurs on dorsum. An opaque
white line occurs along edge of foot; similar
lines occur on anterior margin of frontal veil
and transversely on head between rhino-
phores up onto rhinophore sheaths. T.L.: 20
mm.
Anus on right, close beneath dorso-lateral
margin, posterior of genital aperture, and
ventral of fifth or sixth branchial appendage.
Radular formula of specimen examined
32(16-34.1.1.1.16-34), MacFarland (1966:
221) reports 57(49.1.49), and Marcus (1961:
31) reports 37(33.1.1.1.33). Rachidian teeth
(Fig. 89d) tricuspidate with median cusp long-
er than lateral cusps. Laterals (Fig. 89c)
broadly triangular with a slight hook distally.
Marginals (Fig. 89a, b) compressed hooks.
Masticatory border of mandibles bears 3-9
rows of denticles.
Type-locality: Point Pinos, Monterey Co.,
California.
Range and habitat: Port Dick, Kenai Pen-
insula, Alaska (Robilliard & Barr, 1978), to Is-
las Coronados, Baja California, Mexico
(Lance, 1961); Japan (Baba, 1969a). Not un-
common in low rocky intertidal to 10 m under
rocks and ledges, often in areas of heavy surf.
Feeds upon the pennatulacean Ptilosarcus
gurneyi, the gorgonian Lophogorgia chilen-
sis, and the alcyonarian Clavularia sp. (Birke-
land, 1974; Gomez, 1973; McDonald 8 Ny-
bakken, 1978).
Remarks: Thompson (1971: 337) reports
that this species is capable of swimming, and
Birkeland (1974: 224) states that it escapes
from its asteroid predators Crossaster pap-
posus and Solaster dawsoni by swimming off
the substratum.
ACKNOWLEDGMENTS
Many people have assisted in various as-
pects of this paper and in various degrees.
First, Richard A. Roller suggested that this
study would be useful, and provided much
discussion and many reprints during the early
stages of the investigation.
Doris Baron of Moss Landing Marine Lab-
oratories library was most helpful in obtain-
ing much of the literature. Susan Harris of
the Hopkins Marine Station library was most
gracious in allowing access to the F. M.
MacrFarland Memorial Library. The staff of the
interlibrary loan department, University of
California, Santa Cruz was extremely helpful
in obtaining much of the older and more ob-
scure literature, and | should especially like
to thank Joan Hodgson who did much more
than any researcher has any right to expect,
by obtaining many articles for which she was
given only the most meager and sometimes
questionable citations; without her friendly
assistance and dedication the synonymies
would be much less complete than they now
are. Dr. |. $. Roginskaya was very kind in
providing much of the Soviet literature which
was not available in this country. Carole M.
Hertz and William Old provided copies of ar-
ticles from “The Festivus” and “New York
Shell Club Notes” respectively.
Dustin Chivers and James T. Carlton of
California Academy of Sciences, Invertebrate
Zoology Department provided preserved
specimens of some rare species and also
helped in obtaining some of the literature. The
staff of Pacific Bio-Marine: Dr. Rim Fay, Pat
Brophy, Marion Patton, and especially R.
Shane Anderson provided numerous speci-
mens of rare species or species from south-
ern California, which greatly added to the
completeness of coverage of this paper. Stu-
dents, staff, and divers at Moss Landing Ma-
rine Laboratories provided specimens of many
species. Dr. Kikutaro Baba provided speci-
mens of Eubranchus misakiensis, and an-
swered a number of taxonomic questions. Dr.
Edward Lyke, Dr. Terrence Gosliner, John
Cooper, and Mark Silberstein provided spec-
imens of Okenia plana, Cuthona abronia, Te-
nellia adspersa, and Dendrodoris sp. b re-
spectively.
CALIFORNIA NUDIBRANCHS 223
Mark Pierson translated the Russian de-
scription of Tritonia primorjensis.
Lynn McMasters drew figures 2, 4, 6, 7,
10, and 15.
Dr. James W. Nybakken was most helpful
and encouraging as my major professor when
this paper was originally written as a mas-
ter's thesis, and was instrumental in its pub-
lication.
Rebecca Daughters typed the text and part
of the references, Carmen Germaine and
Barbara O’Connell typed portions of the ref-
erences and Appendix.
| would especially like to thank my wife An-
drea, who as friend, fiancee, and finally wife,
accompanied me on many dives and intertid-
al field trips (many of which were in the cold,
early morning), collected many specimens,
helped in collecting food data, did much
proofreading, assisted in the literature search,
provided encouragement and stimulating dis-
cussion, and helped in many other aspects
of preparing this paper.
Many other people helped in a variety of
ways, and to them | also express my thanks.
DEDICATION
This review is dedicated to Richard A. Rol-
ler, who has taken time and has had interest
in a budding biologist who was frustrated
trying to identify his tidepool finds, who has
given encouragement and advice in the
methods of studying opisthobranchs, and
who has always been available to discuss
taxonomic questions, lend reprints, and gen-
erate interest in nudibranchs.
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APPENDIX: MATERIAL EXAMINED
This list includes most of the material ex-
amined during this study; some material ex-
amined alive and returned to the field is not
included. Data concerning each species are
listed in the following order: accession num-
ber, locality of collection, date of collection,
collector, and number of specimens. Acces-
sion numbers beginning with an M and fol-
lowed by four groups of digits separated by
hyphens (e.g. M71-8-31-2) indicate speci-
mens in my personal collection. Accession
numbers beginning with an M and followed
by four digits (e.g. M0135) indicate speci-
mens in the Moss Landing Marine Labora-
tories. Specimens preceded by CASIZ are in
the California Academy of Sciences, Depart-
ment of Invertebrate Zoology. Specimens
preceded by LACM are in the Los Angeles
County Museum of Natural History. The as-
terisks indicate the lots from which radulae
were taken.
Acanthodoris brunnea
M71-8-31-2, Santa Monica Bay, Los An-
geles Co., Calif., 31 Aug. 1971, S. Anderson,
1. M71-8-31-7, Santa Monica Bay, Los An-
geles Co., Calif., 31 Aug. 1971, S. Anderson,
1. M71-8-31-8, Santa Monica Bay, Los An-
geles Co., Calif., 31 Aug. 1971, $. Anderson,
1. M71-10-7-1, Santa Monica Bay, Los An-
geles Co., Calif., 7 Oct. 1971, S. Anderson,
1. *M71-10-7-4, Santa Monica Bay, Los An-
geles Co., Calif., 7 Oct. 1971, S. Anderson,
1. M72-3-22-3, Monterey Breakwater, Mon-
terey Co., Calif., 22 Mar. 1972, G. McDonald,
1. M72-12-9-1, Monterey Bay, Calif., 9 Dec.
1972, G. McDonald, 2. M74-3-25-1, Monte-
rey Bay, Calif., 25 Mar. 1974, G. McDonald,
JE
Acanthodoris hudsoni
M70-10-16-1, Asilomar, Monterey Co.,
Calif., 16 Oct. 1970, G. McDonald, 1. M69-6-
3-1, $ of Spooner’s Cove, San Luis Obispo
Co., Calif., 3 June 1969, G. McDonald, 1.
M72-1-15-1, Asilomar, Monterey Co., Calif.,
15 Jan. 1972, G. McDonald, 1. M75-10-4-3,
262 MCDONALD
Monastery Beach, Monterey Co., Calif., 4
Oct. 1975, Andrea McDonald, 1. M76-1-17-
1, Monastery Beach, Carmel, Monterey Co.,
Calif., 17 Jan. 1976, G. McDonald, 1. M78-8-
24-3, Monastery Beach, Carmel, Monterey
Co., Calif., 24 Aug. 1978, G. McDonald, 2.
M0135, Asilomar, Monterey Co., Calif., 16
Oct. 1970, G. McDonald, 1.
Acanthodoris lutea
*M69-9-14-2, Morro Bay, San Luis Obispo
Co., Calif., 14 Sept. 1969, G. McDonald, 5.
M70-12-12-1, Santa Cruz Harbor, Santa Cruz
Co., Calif., 12 Dec. 1970, S. Anderson, 1.
M71-1-25-7, Sunset Palisades, San Luis
Obispo Co., Calif. 25 Jan. 1971, G. Mc-
Donald, 2. M71-1-26-2, Hazard Can., San
Luis Obispo Co., Calif., 26 Jan. 1971, G.
McDonald, 2. M71-9-21-1, Pt. Fermin, Los
Angeles Co., Calif., 21 Sept. 1971, S. Ander-
son, 1. M71-12-28-2, Sunset Palisades, San
Luis Obispo Co., Calif., 28 Dec. 1971, G.
McDonald, 2. M71-12-31-3, Shell Beach, San
Luis Obispo Co., Сай., 31 Dec. 1971, С.
McDonald, 1. M75-11-3-5, Pigeon Pt., San
Mateo Co., Calif., 3 Nov. 1975, G. McDonald,
3. M75-12-1-2, Pigeon Pt., San Mateo Co.,
Calif., 1 Dec. 1975, Andrea McDonald, 2.
M77-12-9-1, Terrace Pt., Santa Cruz Co.,
Calif., 9 Dec. 1977, G. McDonald, 1. M0018,
Pt. Fermin, Los Angeles Co., Calif., 21 Sept.
1971, S. Anderson, 1. M0134, Santa Cruz
Harbor, Santa Cruz Co., Calif., 12 Dec. 1970,
S. Anderson, 1.
Acanthodoris nanaimoensis
M71-3-6-1, Pigeon Pt., San Mateo Co.,
Calif., 6 Mar. 1971, invert. class, 1. M71-6-
24-1, Bodega Bay, Sonoma Co., Calif., 24
June 1971, J. Nybakken, 2. M71-7-25-5,
Morro Bay, San Luis Obispo Co., Calif., 25
July 1971, G. McDonald, 1. *M72-9-0-1, San
Francisco Bay, Calif., Sept. 1972, G. Huey,
1. M75-11-3-1, Pigeon Pt., San Mateo Co.,
Calif., 3 Nov. 1975, G. McDonald, 5. M75-12-
1-1, Pigeon Pt., San Mateo Co., Calif., 1 Dec.
1975, G. & A. McDonald, 9. M76-12-20-1,
Scott Cr., Santa Cruz Co., Calif., 20 Dec.
1976, A. K. McDonald, 1. M77-11-11-3, Scott
Cr., Santa Cruz Co., Calif., 11 Nov. 1977, G.
McDonald, 1.
Acanthodoris pilosa
M70-11-27-1, Morro Bay, San Luis Obispo
Co., Calif., 27 Nov. 1970, G. McDonald, 1.
M70-11-8-4, Morro Bay, San Luis Obispo
Co., Calif. 8 Nov. 1970, G. McDonald, 1.
*M71-10-23-1, Elkhorn Slough, Monterey Co.,
Calif., 23 Oct. 1971, G. McDonald, 1. M75-6-
27-1, Moss Landing, Monterey Co., Calif., 27
June 1975, В. Christiansen. M0143, Monte-
rey Bay, Calif., 7 May 1971, class, 1. M76-
12-19-1, Pescadero Pt., San Mateo Co., Cal-
if., 19 Dec. 1976, G. McDonald, 4. M77-11-
11-4, Scott Cr., Santa Cruz Co., Calif., 11
Nov. 1977, G. McDonald, 1.
Acanthodoris rhodoceras
M71-10-17-9, Morro Bay, San Luis Obispo
Co., Calif., 17 Oct: 1971, G. McDonald:
M71-11-2-1, S of Spooner's Cove, San Luis
Obispo Co. Calif., 2 Nov. 1971, G. Mc-
Donald, 1. M71-11-22-4, Morro Bay, San Luis
Obispo Co., Calif., 22 Nov. 1971, R. Roller,
1. *М71-12-28-1, Sunset Palisades, San Luis
Obispo Co., Calif., 28 Dec. 1971, G. Mc-
Donald, 6. M71-12-31-2, Shell Beach, San
Luis Obispo Co., Calif., 31 Dec. 1971, G.
McDonald, 3. M72-4-8-1, Elkhorn Slough,
Monterey Co., Calif., 8 Apr. 1972, G. Mc-
Donald, 1. M75-6-25-3, La Jolla, San Diego
Co., Calif., 25 June 1975, G. McDonald, 1.
M0127, Dillon Beach, Marin Co., Calif., 9 July
1971, J. Nybakken, 1.
Aegires albopunctatus
M69-9-14-1, Morro Bay, San Luis Obispo
Co., Calif., 14 Sept. 1969, G. McDonald, 2.
M70-6-5-4, Pirate's Cove, San Luis Obispo
Co., Calif., 5 June 1970, G. McDonald, 1.
M70-10-14-4, Asilomar, Monterey Co., Calif.,
14 Oct. 1970, G. McDonald, 1. M71-2-15-1,
Monterey Harbor, Monterey Co., Calif., 15
Feb. 1971, G. McDonald, 3. M71-2-24-8,
Morro Bay, San Luis Obispo Co., Calif., 24
Feb. 1971, G. McDonald, 1. M71-7-25-3,
Morro Bay, San Luis Obispo Co., Calif., 25
July 1971, G. McDonald, 1. M71-11-4-5, Car-
mel Pt., Monterey Co., Calif., 4 Nov. 1971,
G. McDonald, 1. M72-1-5-2, Malibu, Los An-
geles, Co., Calif., 5 Jan. 1972, S. Anderson,
4. M72-1-16-3, Monterey Breakwater, Mon-
terey Co., Calif., 16 Jan. 1972, R. Hilaski, 1.
M72-3-22-1, Monterey Breakwater, Monte-
rey Co., Calif., 22 Mar. 1972, G. McDonald,
4. M72-5-17-9, Carmel Pt., Monterey Co.,
Calif., 17 May 1972, G. McDonald, 1. *M72-
7-9-2, Shell Beach, San Luis Obispo Co.,
Calif., 9 July 1972, G. McDonald, 20. M73-6-
5-16, Asilomar, Monterey Co., Calif., 5 June
1973, G. McDonald, 1. M75-10-26-3, Morro
Bay, San Luis Obispo Co., Calif., 26 Oct.
1975, G. McDonald, 1. M77-12-10-11, Asilo-
mar, Monterey Co., Calif., 10 Dec. 1977, G.
McDonald, 1.
CALIFORNIA NUDIBRANCHS 263
Aeolidia papillosa
M70-10-22-2, Moss Landing, Monterey
Co., Calif., 22 Oct. 1970, $. Anderson, 1.
M70-11-14-1, Elkhorn Slough, Monterey Co.,
Calif. 14 Nov. 1970, G. McDonald, 1. M71-
1-26-12, Morro Bay, San Luis Obispo Co.,
Calif., 26 Jan. 1971, G. McDonald, 3. M71-1-
32-1, Monterey Harbor, Monterey Co., Calif.,
Jan. 1971, S. Anderson, 1. M71-2-15-3,
Monterey Harbor, Monterey Co., Calif., 15
Feb. 1971, S. Anderson, 1. M71-7-7-3, Mor-
ro Bay, San Luis Obispo Co., Calif., 7 July
1971, G. McDonald, 1. M71-8-8-2, Asilomar,
Monterey Co., Calif., 8 Aug. 1971, G. Mc-
Donald, 1. M71-10-17-11, Morro Bay, San
Luis Obispo Co., Calif., 17 Oct. 1971, G.
McDonald, 2. *M71-10-22-3, Monterey Har-
bor, Monterey Co., Calif., 22 Oct. 1971, G.
McDonald, 5. M71-11-1-4, Hazard Can., San
Luis Obispo Co., Calif., 1 Nov. 1971, G.
McDonald, 1. M71-11-17-8, Elkhorn Slough,
Monterey Co., Calif., 17 Nov. 1971, G.
McDonald, 1. M71-11-22-5, Morro Bay, San
Luis Obispo Co., Calif., 22 Nov. 1971, G.
McDonald, 1. M74-3-13-1, Monterey Bay,
Calif. 13 Mar. 1974, G. McDonald, 1. M75-
10-26-5, Morro Bay, San Luis Obispo Co.,
Calif. 26 Oct. 1975, G. McDonald, 2. M75-
11-4-2, Fanshell Beach, Monterey Co., Calif.,
4 Nov. 1975, G. McDonald, 2.
Aldisa sanguinea
M71-6-13-2, Monterey Breakwater, Mon-
terey Co., Calif., 13 June 1971, S. Anderson,
1. M71-6-13-3, Monterey Breakwater, Mon-
terey Co., Calif., 13 June 1971, S. Anderson,
1. M71-6-13-5, Monterey Breakwater, Mon-
terey Co., Calif., 13 June 1971, S. Anderson,
1. M71-8-15-1, Port San Luis, San Luis Obis-
po Co., Calif., 15 Aug. 1971, G. McDonald,
1. M71-12-2-8, Carmel Pt., Monterey Co.,
Calif., 2 Dec. 1971, G. McDonald, 1. M71-12-
28-5, Dume Rock, Los Angeles Co., Calif., 28
Dec. 1971, $. Anderson, 2. M72-3-19-2, As-
ilomar, Monterey Co., Calif. 19 Mar. 1972,
G. McDonald, 1. M72-4-17-1, Asilomar, Mon-
terey Co., Calif., 17 Apr. 1972, G. McDonald,
1. *M72-4-26-3, Monterey Breakwater, Mon-
terey Co., Calif., 26 Apr. 1972, G. McDonald,
2. M72-5-14-1, Rocky Pt., Monterey Co.,
Calif. 14 Мау 1972, G. McDonald, 1. M72-
5-17-8, Carmel Pt., Monterey Co., Calif., 17
May 1972, G. McDonald, 2. M72-7-18-1,
Spooner's Cove, San Luis Obispo Co., Calif.,
18 July 1972, G. McDonald, 1. M72-12-18-1,
Asilomar, Monterey Co., Calif., 18 Dec. 1972,
G. McDonald, 1. M72-12-20-1, Asilomar,
Monterey Co., Calif., 20 Dec. 1972, G.
McDonald, 2. M73-1-0-1, Los Angeles Har-
bor, Los Angeles Co., Calif., Jan. 1973, R.
Fay, 1. M73-4-9-1, Carmel Pt., Monterey Co.,
Calif., 9 Apr. 1973, G. McDonald, 3. M67-11-
17-1, Hazard Can., San Luis Obispo Co.,
Calif., 17 Nov. 1967, R. Roller, 1. M71-1-8-8,
White's Pt., Los Angeles Co., Calif., 8 Jan.
1971, R. Roller, 2. M73-4-9-8, Carmel Pt.,
Monterey Co., Calif., 9 Apr. 1973, G. Mc-
Donald, 2. M73-5-5-10, Asilomar, Monterey
Co., Calif., 5 May 1973, G. McDonald, 1.
M75-11-2-4, Carmel Pt., Monterey Co., Cal-
if., 2 Nov. 1975, G. McDonald, 1.
Ancula lentiginosa
*M71-10-22-6, Monterey Harbor, Monte-
rey Co., Calif., 22 Oct. 1971, G. McDonald,
5. M71-11-4-2, Asilomar, Monterey Co., Cal-
Н., 4 Nov. 1971, G. McDonald, 1. M71-11-17-
1, Elkhorn Slough, Monterey Co., Calif., 17
Nov. 1971, G. McDonald, 2. M71-11-22-6,
Morro Bay, San Luis Obispo Co., Calif., 22
Nov. 1971, G. McDonald, 2. M71-12-4-2, Elk-
horn Slough, Monterey Co., Calif., 4 Dec.
1971, G. McDonald, 3.
Ancula pacifica
M70-6-5-1, Pirate's Cove, San Luis Obis-
po Co., Calif., 5 June 1970, G. McDonald, 1.
M70-6-28-3, Pirate's Cove, San Luis Obispo
Co., Calif. 28 June 1970, G. McDonald, 1.
M70-7-18-2, Shell Beach, San Luis Obispo
Co., Calif. 18 July 1970, G. McDonald, 2.
M70-7-18-3, Shell Beach, San Luis Obispo
Co., Calif., 18 July 1970, G. McDonald, 3.
M70-10-14-2, Asilomar, Monterey Co., Calif.,
14 Oct. 1970, G. McDonald, 3. M71-1-26-23,
Morro Bay, San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 1. M71-2-8-2, Asi-
lomar, Monterey Co., Calif., 8 Feb. 1971, G.
McDonald, 1. M71-11-1-2, Hazard Can., San
Luis Obispo Co., Calif., 1 Nov. 1971, G.
McDonald, 2. M73-5-5-2, Asilomar, Monte-
rey Co., Calif., 5 May 1973, G. McDonald, 1.
M73-5-6-2, Pt. Pinos, Monterey Co., Calif., 6
Мау 1973, G. McDonald, 2. M73-6-5-13, As-
ilomar, Monterey Co., Calif., 5 June 1973, G.
McDonald, 1. *M75-11-2-3, Carmel Pt., Mon-
terey Co., Calif., 2 Nov. 1975, G. McDonald,
5. M78-6-22-9, Carmel Pt., Monterey Co.,
Calif. 22 June 1978, G. McDonald, 1.
Anisodoris nobilis
M70-12-12-7, Santa Cruz Harbor, Santa
Cruz Co., Calif. 12 Dec. 1970, S. Anderson,
1. M70-12-12-8, Pigeon Pt., San Mateo Co.,
Calif., 12 Dec. 1970, G. McDonald, 1. M70-
264 MCDONALD
12-12-10 Santa Cruz Harbor, Santa Cruz Co.,
Calif., 12 Dec. 1970, S. Anderson, 1. M71-1-
25-8, Pirate's Cove, San Luis Obispo Co.,
Calif., 25 Jan. 1971, G. McDonald, 2. *M71-
1-26-1, Hazard Can., San Luis Obispo Co.,
Calif., 26 Jan. 1971, G. McDonald, 5. M71-1-
26-19, Hazard Can., San Luis Obispo Co.,
Calif., 26 Jan. 1971, G. McDonald, 1. M71-5-
26-3, Asilomar, Monterey Co., Calif., 26 May
1971, G. McDonald, 1. M71-11-2-7, Spoon-
er's Cove, San Luis Obispo Co., Calif., 2 Nov.
1971, G. McDonald, 1. M72-1-16-2, Monte-
rey Breakwater, Monterey Co., Calif., 16 Jan.
1972, R. Hilaski, 1. M73-6-5-2, Asilomar,
Monterey Co., Calif., 5 June 1973, G. Mc-
Donald, 1. M75-10-5-1, Carmel Pt., Monterey
Co., Calif., 5 Oct. 1975, G. McDonald, 4.
M0012, Anacapa Is., Calif., Oct. 1968, P.
Brophy, 1. M0015, Asilomar, Monterey Co.,
Calif., 3 Nov. 1971, J. Nybakken, 1. M0167,
Pt. Pinos, Monterey Co., Calif., 25 Apr. 1967,
V. Mansfield, 1. M0169, Moss Beach, Mon-
terey Co., Calif., 16 Oct. 1966, J. Nybakken,
de
Antiopella barbarensis
M71-1-26-10, Hazard Can., San Luis Obis-
po Co., Calif., 26 Jan. 1971, G. McDonald, 1.
M71-2-1-5, Port San Luis, San Luis Obispo
Co., Calif., 1 Feb. 1971, G. McDonald, 1.
M71-8-8-4, Asilomar, Monterey Co., Calif., 8
Aug. 1971, G. McDonald, 1. M71-12-30-4,
Hazard Can., San Luis Obispo Co., Calif., 30
Dec. 1971, G. McDonald, 3. M69-10-18-1,
San Juan Is., Wash., 18 Oct. 1969, G. Ro-
billiard, 6.
Archidoris montereyensis
M71-1-25-6, Pirate's Cove, San Luis Obis-
po Co., Calif., 25 Jan. 1971, G. McDonald, 1.
M71-1-26-20, Мото Bay, San Luis Obispo
Co., Calif., 26 Jan. 1971, G. McDonald, 2.
M71-11-1-1, Hazard Can., San Luis Obispo
Co., Calif., 1 Nov. 1971, G. McDonald, 1.
M71-11-16-2, Elkhorn Slough, Monterey Co.,
Calif., 16 Nov. 1971, G. McDonald, 1. M71-
11-17-5, Elkhorn Slough, Monterey Co., Cal-
if., 17 Nov. 1971, G. McDonald, 1. M71-12-
4-1, Elkhorn Slough, Monterey Co., Calif., 4
Dec. 1971, G. McDonald, 1. M72-1-16-8, ЕК-
horn Slough, Monterey Co., Calif., 16 Jan.
1972, G. McDonald, 3. M72-11-1-6, Moss
Landing, Monterey Co., Calif., 1 Nov. 1972,
G. McDonald, 4. M75-8-6-1, Elkhorn Slough,
Monterey Co., Calif., 6 Aug. 1975, B. Antrim,
1. M75-8-8-1, Pacific Grove, Monterey Co.,
Calif. 8 Aug. 1975, М. E. Anderson, 1.
M0139, Elkhorn Slough, Monterey Co., Cal-
if., Nov. 1970, G. McDonald, 1. M0176, Car-
mel Pt., Monterey Co., Calif., 15 Oct. 1970,
J. Nybakken, 1.
Archidoris odhneri
M71-1-27-2, Sunset Palisades, San Luis
Obispo Co., Calif., 27 Jan. 1971, G. Mc-
Donald, 1. *M71-7-3-2, Sunset Palisades, San
Luis Obispo Co., Calif., 3 July 1971, G.
McDonald, 2. M71-7-3-3, Sunset Palisades,
San Luis Obispo Co., Calif., 3 July 1971, P.
Clark, 1. M72-7-16-1, Pirate's Cove, San Luis
Obispo Co., Calif., 16 July 1972, G. Mc-
Donald, 5. M75-9-28-3, Monastery Beach,
Monterey Co., СаШ., 28 Sept. 1975, A.
McDonald, 2. M0237, Monastery Beach,
Monterey Co., Calif., 25 Nov. 1970, E. Stark,
1. M0523, Hopkin's Reef, Monterey Co., Cal-
Н., 8 Dec. 1974, D. Hunt, 2.
Armina californica
M71-4-10-4, Monterey Bay, Calif., 10 Apr.
1971, а. McDonald, 1. M71-5-4-1, Monterey
Bay, Calif., 4 May 1971, С. McDonald, 1.
M71-10-0-1, Zuma Beach, Los Angeles Co.,
Calif., Oct. 1971, S. Anderson, 1. M71-11-
18-1, Monterey Bay, Calif., 18 Nov. 1971, D.
Varoujean, 1. M71-11-15-1, Santa Monica
Bay, Los Angeles Co., Calif., 26 Nov. 1971,
S. Anderson, 3. M72-1-7-4, Santa Monica
Bay, Los Angeles Co., Calif., 7 Jan. 1972, S.
Anderson, 2. M69-8-27-2, Friday Harbor,
Wash., 27 Aug. 1969, R. Roller, 1. M73-4-
16-1, Monterey Bay, Calif., 16 Apr. 1973, G.
McDonald, 1. M0022, Año Nuevo Pt., San
Mateo Co., Calif., 3 Aug. 1971, D. Ventresca,
1. M0321, Monterey Bay, Calif., 8 Mar. 1972,
С. Kukowski, 1. M0322, Monterey Bay, Cal-
if., 8 Mar. 1972, G. Kukowski, 1.
Atagema alba
M69-7-31-1, Sunset Cliffs, San Diego Co.,
Calif., 31 July 1969, R. Roller, 1. *M73-10-0-
1, Escondido Can., San Diego Co., Calif., Oct.
1973, M. Patton, 1. CASIZ, radula of holo-
type of Petelodoris spongicola.
Babakina festiva
M71-10-0-9, Los Angeles Co., Calif., Oct.
1971, $. Anderson, 1. *M71-11-0-4, Malibu,
Los Angeles Co., Calif., Nov. 1971, S. An-
derson, 1. M75-1-25-2, Matanchen, Nayarit,
Mexico, 25 Jan. 1975, C. Boone, 4.
Cadlina flavomaculata
M69-10-26-8, Pirate's Cove, San Luis
Obispo Co., Calif., 26 Oct. 1969, G. Mc-
Donald, 2. M71-1-25-4, Sunset Palisades,
CALIFORNIA NUDIBRANCHS 265
San Luis Obispo Co., Calif., 25 Jan. 1971, G.
McDonald, 1. M71-12-28-4, Dume Rock, Los
Angeles Co., Calif., 28 Dec. 1971, S. Ander-
son, 2. M71-12-31-4, Shell Beach, San Luis
Obispo Co., Calif. 31 Dec. 1971, G. Mc-
Donald, 2. M73-6-5-7, Asilomar, Monterey
Co., Calif., 5 June 1973, G. McDonald, 2.
M75-10-5-6, Carmel Pt., Monterey Co., Cal-
Н., 5 Oct. 1975, G. McDonald, 2. M75-11-2-
10, Carmel Pt., Monterey Co., Calif., 2 Nov.
1975, G. McDonald, 1. MO318, Asilomar,
Monterey Co., Calif., 15 May 1972, J. Nybak-
ken, 1.
Cadlina marginata
M70-2-5-3, Field Ranch, San Luis Obispo
Co., Calif., 5 Feb. 1970, G. McDonald, 2.
M70-10-17-2, Pacific Grove, Monterey Co.,
Calif., 17 Oct. 1970, С. McDonald, 1. M70-
12-12-6, Pigeon Pt., San Mateo Co., Calif.,
12 Dec. 1970, G. McDonald, 2. M71-1-26-3,
Hazard Can., San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 1. M71-12-29-1,
Pirate's Cove, San Luis Obispo Co., Calif.,
29 Dec. 1971, G. McDonald, 2. M72-5-14-2,
Rocky Pt., Monterey Co., Calif., 14 May 1972,
J. Nybakken, 1. *M73-4-9-7, Carmel Pt.,
Monterey Co., Calif., 9 Apr. 1973, G. Mc-
Donald, 1. M75-9-28-1, Monastery Beach,
Monterey Co., Calif., 28 Sept. 1975, A.
McDonald, 1. M75-10-4-1, Monastery Beach,
Monterey Co., Calif., 4 Oct. 1975, A. Mc-
Donald, 5. M75-11-2-1, Carmel Pt., Monterey
Co., Calif., 2 Nov. 1975, G. McDonald, 3.
M0131, Monastery Beach, Monterey Co.,
Calif., 18 Nov. 1970, E. Stark, 1. M0164,
Carmel Pt., Monterey Co., Calif., 15 Oct.
1970, J. Nybakken, 1. MO189, Pt. Pinos,
Monterey Co., Calif., 27 Apr. 1967, V. Mans-
field, 2. M0311, Rocky Pt., Monterey Co.,
Calif., 14 May 1972, J. Nybakken, 2.
Cadlina modesta
M69-10-26-6, Pirate's Cove, San Luis
Obispo Co., Calif., 26 Oct. 1969, G. Mc-
Donald, 1. M70-10-17-3, Pacific Grove, Mon-
terey Co., Calif., 17 Oct. 1970, G. McDonald,
2. M71-1-26-4, Hazard Can., San Luis Obis-
po Co., Calif., 26 Jan. 1971, G. McDonald, 6.
M71-11-2-3, Spooner's Cove, San Luis Obis-
po Co., Calif., 2 Nov. 1971, G. McDonald, 1.
M71-12-29-2, Pirate's Cove, San Luis Obis-
po Co., Calif., 29 Dec. 1971, G. McDonald,
3. M73-6-5-6, Asilomar, Monterey Co., Calif.,
5 June 1973, G. McDonald, 1. M75-10-5-5,
Carmel Pt., Monterey Co., Calif., 5 Oct. 1975,
G. McDonald, 2. M75-11-2-8, Carmel Pt.,
Monterey Co., Calif., 2 Nov. 1975, G. Mc-
Donald, 4. M0174, Asilomar, Monterey Co.,
Calif., 10 Nov. 1970, G. McDonald, 1. MO219,
Carmel Pt., Monterey Co., Calif., 15 Oct.
1970, J. Nybakken, 1.
Cadlina sparsa
M71-12-27-1, Malibu, Los Angeles Co.,
Calif., 27 Dec. 1971, $. Anderson, 1. M72-3-
22-2, Monterey Breakwater, Monterey Co.,
Calif. 22 Mar. 1972, G. McDonald, 1. M72-
10-22-1, Asilomar, Monterey Co., Calif., 22
Oct. 1972, J. Nybakken, 2. *M73-6-5-5, Asi-
lomar, Monterey Co., Calif., 5 June 1973, G.
McDonald, 1. M75-6-25-2, La Jolla, San Die-
go Co., Calif., 25 June 1975, G. McDonald,
1. M77-11-11-1, Scott Cr., Santa Cruz Co.,
Calif., 11 Nov. 1977, G. McDonald, 1.
Cerberilla mosslandica
M71-11-10-1, Monterey Bay, Calif., 21 Aug.
1971, 1; 10Nov.. 1971, 1; 3 Feb: 197271.
M72-8-23-6, Monterey Bay, Сай., 23 Aug.
1972, G. McDonald, 2. M72-8-23-9, Monte-
rey Bay, Calif., 23 Aug. 1972, G. McDonald,
2. M0323, Monterey Bay, Calif., 10 May 1972,
G. McDonald, 7. M0481, Monterey Bay, Cal-
if., 1971-1972, Sea Grant, approx. 100.
M0482, Monterey Bay, Calif., 3 Feb. 1972,
Sea Grant, 1, paratype. M0483, Monterey
Bay, Calif., 20 Aug. 1971, Sea Grant, 1,
paratype. CASIZ no. 596, Monterey Bay,
Calif., 10 May 1972, Sea Grant, 1. holotype.
CASIZ no. 597, Monterey Bay, Calif., Sea
Grant, 3, paratypes. LACM 1725, 1726,
Monterey Bay, Calif., Sea Grant, 3, para-
types.
Chromodoris macfarlandi
*M70-7-9-1, Sunset Palisades, San Luis
Obispo Co., Calif, 9 July 1970, G. Mc-
Donald, 1. M70-7-9-2, Sunset Palisades, San
Luis Obispo Co. Сай., 9 July 1970, G.
McDonald, 1. M70-7-26-1, Sunset Palisades,
San Luis Obispo Co., Calif., 26 July 1970, G.
McDonald, 1. M71-12-2-13, La Jolla, San
Diego Co., Calif., 2 Dec. 1971, S. Anderson,
1. M75-6-25-7, La Jolla, San Diego Co., Cal-
Н., 25 June 1975, G. McDonald, 1. M78-8-17-
5, Monterey Breakwater, Monterey Co., Cal-
Н., 17 Aug. 1978, А. К. McDonald, 1.
Chromodoris porterae
M71-8-2-1, Malibu, Los Angeles Co., Cal-
if., 2 Aug. 1971, $. Anderson, 1. M71-8-2-5,
Malibu, Los Angeles Co., Calif., 2 Aug. 1971,
S. Anderson, 1. M71-8-2-6, Malibu, Los An-
geles Co., Calif., 2 Aug. 1971, S. Anderson,
266 MCDONALD
1. M71-9-14-3, Malibu, Los Angeles Co., Cal-
if., 14 Sept. 1971, $. Anderson, 1. *M71-10-
0-5, Malibu, Los Angeles Co., Calif., Oct.
1971, S. Anderson, 4. M71-11-23-2, Malibu,
Los Angeles Co., Calif., 23 Nov. 1971, S. An-
derson, 1. M72-3-2-1, Malibu, Los Angeles
Co., Calif., 2 Mar. 1972, S. Anderson, 1. M71-
11-23-5, Malibu, Los Angeles Co., Calif., 23
Nov. 1971, S. Anderson, 1.
Conualevia alba
*М71-9-21-2, Pt. Fermin, Los Angeles Co.,
Calif., 21 Sept. 1971, S. Anderson, 2. M71-
1-8-7, White's Pt., Los Angeles Co., Calif., 8
Jan. 1971, 2. M71-10-20-4, Los Angeles Co.,
Calif., 20 Oct. 1971, S. Anderson, 2. M71-
10-22-8, Los Angeles Co., Calif., 22 Oct.
1971, S. Anderson, 1. M75-1-23-8, Cruz de
Huanacaxtle, Nayarit, Mexico, 23 Jan. 1975,
G. McDonald, 15. M77-11-12-2, Carmel Pt.,
Monterey Co., Calif., 12 Nov. 1977, R. Slutz,
1. MO363, Carmel Pt., Monterey Co., Calif.,
9 Apr. 1973, G. McDonald, 1.
Corambe pacifica
M70-10-7-3, Monterey Bay, Calif., 7 Oct.
1970, G. McDonald, 12. M70-11-27-2, Morro
Bay, San Luis Obispo Co., Calif., 27 Nov.
1970, G. McDonald, 2. M71-1-26-21, Morro
Bay, San Luis Obispo Co., Calif., 26 Jan.
1971, G. McDonald, 1. M71-10-17-10, Morro
Bay, San Luis Obispo Co., Calif., 17 Oct.
1971, G. McDonald, 12. *M71-11-22-3, Mor-
ro Bay, San Luis Obispo Co., Calif., 22 Nov.
1971, G. McDonald, 4. M69-11-15-1, Morro
Bay, San Luis Obispo Co., Calif., 15 Nov.
1969, R. Roller, 2. M73-6-21-1, Pirate's Cove,
San Luis Obispo Co., Calif., 21 June 1973,
G. McDonald, 1. M75-10-14-4, Monterey Bay,
Calif., 14 Oct. 1975, G. McDonald, 1. M75-
10-26-10, Morro Bay, San Luis Obispo Co.,
Calif., 26 Oct. 1975, 7. M78-8-17-6, Monte-
rey Breakwater, Monterey Co., Calif., 17 Aug.
1978, G. McDonald, 12.
Coryphella cooperi
*M70-11-16-3, Elkhorn Slough, Monterey
Co., Calif., 16 Nov. 1970, G. McDonald, 1.
M70-11-16-8, Elkhorn Slough, Monterey Co.,
Calif., 16 Nov. 1970, G. McDonald, 1. M70-
12-7-1, Elkhorn Slough, Monterey Co., Calif.,
7 Dec. 1970, G. McDonald, 1. M76-11-12-1,
Elkhorn Slough, Monterey Co., Calif., 12 Nov.
1976, J. Cooper, 3. M76-11-20-1, Elkhorn
Slough, Monterey Co., Calif., 20 Nov. 1976,
G. McDonald, 1.
Coryphella iodinea
M70-6-13-1, Port San Luis, San Luis Obis-
po Co., Calif., 13 June 1970, G. McDonald,
1. M71-2-1-2, Port San Luis, San Luis Obis-
po Co., Calif., 1 Feb. 1971, G. McDonald, 1.
M71-2-1-3, Port San Luis, San Luis Obispo
Co., Calif., 1 Feb. 1971, G. McDonald, 1.
M71-2-2-1, Port San Luis, San Luis Obispo
Co., Calif., 2 Feb. 1971, G. McDonald, 1.
M71-2-2-2, Port San Luis, San Luis Obispo
Co., Calif., 2 Feb. 1971, G. McDonald, 1.
M71-2-2-4, Port San Luis, San Luis Obispo
Co., Calif., 2 Feb. 1971, G. McDonald, 1.
M71-7-3-1, Sunset Palisades, San Luis Obis-
po Co., Calif., 3 July 1971, G. McDonald, 1.
M71-8-18-3, Santa Monica Bay, Los Angeles
Co., Calif., 18 Aug. 1971, В. $. Anderson, 2.
M71-8-31-5, Santa Monica Bay, Los Angeles
Co., Calif., 31 Aug. 1971, R. S. Anderson, 2.
M71-11-26-3, Santa Monica Bay, Los Ange-
les Co., Calif., 26 Nov. 1971, R. S. Anderson,
1. *M72-7-9-1, Shell Beach, San Luis Obispo
Co., Calif., 9 July 1972, G. McDonald, 5. M75-
6-26-3, La Jolla, San Diego Co., Calif., 26
June 1975, M. U. Purdue, 1. M75-11-23-1,
Pacific Grove, Monterey Co., Calif., 23 Nov.
1975, А. К. McDonald, 1. M0186, Del Monte
Beach, Monterey Co., Calif., 1 Apr. 1967, V.
Mansfield, 1. MO362, Carmel Pt., Monterey
Co., Calif., 9 Apr. 1973, G. McDonald, 1.
Coryphella pricei
*M71-11-0-1, Zuma Beach, Los Angeles
Co., Calif., Nov. 1971, S. Anderson, 1.
Coryphella trilineata
M70-2-26-2, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 1.
M70-11-8-1, Morro Bay, San Luis Obispo
Co., Calif., 8 Nov. 1970, G. McDonald, 2.
M70-11-13-3, Elkhorn Slough, Monterey Co.,
Calif., 13 Nov. 1970, G. McDonald, 1. M70-
11-16-2, Elkhorn Slough, Monterey Co., Cal-
if., 16 Nov. 1970, G. McDonald, 1. M70-11-
16-6, Elkhorn Slough, Monterey Co., Calif.,
16 Nov. 1970, G. McDonald, 5. M71-10-17-
6, Morro Bay, San Luis Obispo Co., Calif.
17 Oct. 1971, G. McDonald, 1. M71-11-17-7,
Elkhorn Slough, Monterey Co., Calif., 17 Nov.
1971, G. McDonald, 2. M71-12-2-5, Carmel
Pt., Monterey Co., Calif., 2 Dec. 1971, G.
McDonald, 1. M72-10-18-9, Elkhorn Slough,
Monterey Co., Calif., 18 Oct. 1972, G. Mc-
Donald, 4. M72-11-1-2, Elkhorn Slough,
Monterey Co., Calif., 1 Nov. 1972, G. Mc-
Donald, 3. M75-10-21-3, Elkhorn Slough,
CALIFORNIA NUDIBRANCHS 267
Monterey Co., Calif., 21 Oct. 1975, G. Mc-
Donald, 6. M0197, Elkhorn Slough, Monterey
Co., Calif., 13 Nov. 1970, G. McDonald, 1.
M0198, Elkhorn Slough, Monterey Co., Cal-
Н., 13 Nov. 1970, G. McDonald, 1. M0207,
Carmel Pt., Monterey Co., Calif., 15 Oct.
1970, J. Nybakken, 1.
Coryphella sp.
*M72-10-18-2, Elkhorn Slough, Monterey
Co., Calif., 18 Oct. 1972, G. McDonald, 15.
M72-10-18-7, Elkhorn Slough, Monterey Co.,
Calif. 18 Oct. 1972, G. McDonald, 2. M72-
10-19-1, Elkhorn Slough, Monterey Co., Cal-
Н., 19 Oct. 1972, G. McDonald, 5. M72-10-
19-2, Elkhorn Slough, Monterey Co., Calif.,
19 Oct. 1972, G. McDonald, 10.
Crimora coneja
M75-6-25-1, La Jolla, San Diego Co., Cal-
if., 25 June 1975, G. McDonald, 2. *Prepared
radula of specimen, Pt. Loma, San Diego Co.,
Calif., 25 Apr. 1973, R. Roller, 1.
Cumanotus beaumonti
M70-10-28-1, Elkhorn Slough, Monterey
Co., Calif., 28 Oct. 1970, G. McDonald, 1.
M70-11-13-1, Elkhorn Slough, Monterey Co.,
Calif., 13 Nov. 1970, G. McDonald, 1. M70-
11-13-2, Elkhorn Slough, Monterey Co., Cal-
if., 13 Nov. 1970, G. McDonald, 6. M70-11-
16-1, Elkhorn Slough, Monterey Co., Calif.,
16 Nov. 1970, G. McDonald, 3. *M70-11-16-
7, Elkhorn Slough, Monterey Co., Calif., 16
Nov. 1970, G. McDonald, 5. M72-10-18-1,
Elkhorn Slough, Monterey Co., Calif., 18 Oct.
1972, G. McDonald, 30. M72-10-18-5, Elk-
horn Slough, Monterey Co., Calif., 18 Oct.
1972, G. McDonald, 7. M72-10-18-6, Elkhorn
Slough, Monterey Co., Calif., 18 Oct. 1972,
G. McDonald, 6. M72-11-1-3, Elkhorn Slough,
Monterey Co., Calif., 1 Nov. 1972, G. Mc-
Donald, 8. M75-10-21-1, Elkhorn Slough,
Monterey Co., Calif., 21 Oct. 1975, G. Mc-
Donald, 9. M75-10-21-5, Elkhorn Slough,
Monterey Co., Calif., 21 Oct. 1975, G. Mc-
Donald, 18. M0201, Elkhorn Slough, Monte-
rey Co., Calif., 13 Nov. 1970, G. McDonald,
6. M0203, Elkhorn Slough, Monterey Co.,
Calif., 28 Oct. 1970, G. McDonald, 1.
Cuthona abronia
*M67-9-6-1, Duxbury Reef, Marin Co., Cal-
if., 6 Aug. 1967, T. Gosliner, 10. M72-8-9-2,
Asilomar, Monterey Co., Calif., 9 Aug. 1972,
G. McDonald, 1. M73-3-0-4, Los Angeles Co.,
Calif., Mar. 1973, R. Fay, 1. M73-5-6-6, Pt.
Pinos, Monterey Co., Calif., 6 May 1973, G.
McDonald, 4. M73-6-2-2, Pescadero Pt.,
Monterey Co., Calif., 2 June 1973, G. Mc-
Donald, 1. M78-7-21-5, Pt. Pinos, Monterey
Co., Calif., 21 July 1978, G. McDonald, 8.
Cuthona albocrusta
M71-5-25-3, Asilomar, Monterey Co., Cal-
Н., 25 May 1971, G. McDonald, 1. *M71-10-
22-4, Monterey Harbor, Monterey Co., Calif.,
22 Oct. 1971, G. McDonald, 2. M72-10-18-4,
Elkhorn Slough, Monterey Co., Calif., 18 Oct.
1972, G. McDonald, 2. M73-6-5-19, Asilo-
mar, Monterey Co., Calif., 5 June 1973, G.
McDonald 1. M75-10-26-12, Morro Bay, San
Luis Obispo Co., Calif., 26 Oct. 1975, G.
McDonald, 2. M76-11-20-2, Elkhorn Slough,
Monterey Co., Calif., 20 Nov. 1976, G.
McDonald, 4. M78-7-21-7, Pt. Pinos, Mon-
terey Co., Calif., 21 July 1978, G. McDonald,
de
Cuthona columbiana
M70-11-16-4, Elkhorn Slough, Monterey
Co., Calif., 16 Nov. 1970, G. McDonald, 1.
M71-1-13-1, Elkhorn Slough, Monterey Co.,
Calif., 13 Jan. 1971, G. McDonald, 9. *M71-
1-13-2, Elkhorn Slough, Monterey Co., Calif.,
13 Jan. 1971, G. McDonald, 15. M71-5-25-5,
Asilomar, Monterey Co., Calif., 25 May 1971,
G. McDonald, 1. M71-6-26-3, Asilomar, Mon-
terey Co., Calif., 26 June 1971, G. McDonald,
1. M71-12-2-7, Carmel Pt., Monterey Co.,
Calif., 2 Dec. 1971, G. McDonald, 1. M72-5-
15-9, Asilomar, Monterey Co., Calif., 15 May
1972, G. McDonald, 1. M72-10-22-4, Asilo-
mar, Monterey Co., Calif., 22 Oct. 1972, G.
McDonald, 1. M72-11-1-1, Elkhorn Slough,
Monterey Co., Calif., 1 Nov. 1972, G. Mc-
Donald, 1. M73-5-5-6, Asilomar, Monterey
Co., Calif., 5 May 1973, J. Nybakken, 1. M73-
6-5-17, Asilomar, Monterey Co., Calif., 5 June
1973, M. Silberstein, 1. M76-11-20-3, Elk-
horn Slough, Monterey Co., Calif., 20 Nov.
1976, G. McDonald, 3. M78-7-21-6, Pt. Pi-
nos, Monterey Co., Calif., 21 July 1978, G.
McDonald, 1.
Cuthona flavovulta
M70-10-14-10, Asilomar, Monterey Co.,
Calif. 14 Oct. 1970, G. McDonald, 1. *M71-
1-26-22, Hazard Can., San Luis Obispo Co.,
Calif., 26 Jan. 1971, G. McDonald, 2. M71-5-
25-2, Asilomar, Monterey Co., Calif., 25 May
1971, G. McDonald, 1. M71-8-8-6, Asilomar,
Monterey Co., Calif., 8 Aug. 1971, G. Mc-
268 MCDONALD
Donald, 1. M71-10-5-3, Asilomar, Monterey
Co., Calif., 5 Oct. 1971, G. McDonald, 1. М71-
10-22-5, Monterey Harbor, Monterey Co.,
Calif., 22 Oct. 1971, G. McDonald, 1. M73-6-
5-18, Asilomar, Monterey Co., Calif., 5 June
1973, G. McDonald, 2. MO313, Asilomar,
Monterey Co., Calif., 15 May 1972, G. Mc-
Donald, 1.
Cuthona fulgens
*M71-5-25-4, Asilomar, Monterey Co., Cal-
if., 25 May 1971, G. McDonald, 1. M73-5-5-
7, Asilomar, Monterey Co., Calif., 5 May 1973,
G. McDonald, 3. M73-6-5-20, Asilomar, Mon-
terey Co., Calif., 5 June 1973, G. McDonald,
6. M78-7-21-8, Pt. Pinos, Monterey Co., Cal-
if., 21 July 1978, G. McDonald, 1. MO366,
Asilomar, Monterey Co., Calif., 5 June 1973,
G. McDonald, 1.
Cuthona lagunae
M70-7-18-4, Shell Beach, San Luis Obispo
Co., Calif. 18 July 1970, G. McDonald, 1.
M71-1-26-14, Hazard Can., San Luis Obispo
Co., Calif., 26 Jan. 1971, G. McDonald, 5.
M71-5-25-1, Asilomar, Monterey Co., Calif.,
25 May 1971, G. McDonald, 4. M71-5-26-1,
Asilomar, Monterey Co., Calif., 26 May 1971,
G. McDonald, 6. M71-6-26-4, Asilomar, Mon-
terey Co., Calif., 26 June 1971, G. McDonald,
1. M71-7-25-7, Morro Bay, San Luis Obispo
Co., Calif., 25 July 1971, G. McDonald, 1.
M71-8-8-7, Asilomar, Monterey Co., Calif., 8
Aug. 1971, G. McDonald, 1. M72-3-19-1, As-
ilomar, Monterey Co., Calif., 19 Mar. 1972,
G. McDonald, 1. *M72-5-15-8, Asilomar,
Monterey Co., Calif., 15 May 1972, G. Mc-
Donald, 4. M68-8-8-1, Hazard Can., San Luis
Obispo Co., Calif., 8 Aug. 1968, R. A. Roller,
2. M73-5-5-8, Asilomar, Monterey Co., Calif.,
5 May 1973, G. McDonald, 4. M73-5-6-5, Pt.
Pinos, Monterey Co., Calif., 6 May 1973, G.
McDonald, 1. M73-6-5-4, Asilomar, Monte-
rey Co., Calif., 5 June 1973, G. McDonald, 5.
M0192, Asilomar, Monterey Co., Calif., 14
Oct. 1970, R. Ajeska, 1. M0312, Asilomar,
Monterey Co., Calif., 15 May 1972, G. Mc-
Donald, 2.
Cuthona virens
*CASIZ, radula of holotype, 1.
Cuthona sp.
*M72-9-20-1, Palo Alto, Santa Clara Co.,
Calif., 20 Sept. 1972, M. Silberstein, 5. M72-
9-20-2, Palo Alto, Santa Clara Co., Calif., 20
Sept. 1972, M. Silberstein, 2.
Dendrodoris albopunctata
M70-10-17-4, Pacific Grove, Monterey Co.,
Calif., 17 Oct. 1970, G. McDonald, 1. M70-
12-12-4, Pigeon Pt., San Mateo Co., Calif.,
12 Dec. 1970, G. McDonald, 1. M71-6-29-1,
Pirate's Cove, San Luis Obispo Co., Calif.,
29 June 1971, G. McDonald, 1. M71-8-2-2,
Malibu, Los Angeles Co., Calif., 2 Aug. 1971,
В. $. Anderson, 1. M71-9-24-1, Malibu, Los
Angeles Co., Calif., 24 Sept. 1971, R. S. An-
derson, 1. M72-1-16-1, Monterey Breakwa-
ter, Monterey Co., Calif., 16 Jan. 1972, R.
Hilaski, 2. M72-12-16-2, Shell Beach, San
Luis Obispo Co., Calif., 16 Dec. 1972, G.
McDonald, 1. M73-4-9-2, Carmel Pt., Mon-
terey Co., Calif., 9 Apr. 1973, G. McDonald,
2. M73-6-5-3, Asilomar, Monterey Co., Calif.,
5 June 1973, G. McDonald, 6. M75-6-24-1,
La Jolla, San Diego Co., Calif., 24 June 1975,
G. McDonald, 3. M75-6-26-5, La Jolla, San
Diego Co., Calif., 26 June 1975, G. Mc-
Donald, 1. M75-8-8-4, Pacific Grove, Monte-
rey Co., Calif., 8 Aug. 1975, M. E. Anderson,
1. M75-10-17-2, Pescadero Pt., Monterey
Co., Calif., 17 Oct. 1975, M. Silberstein, 1.
M75-11-2-17, Carmel Pt., Monterey Co., Cal-
if., 2 Nov. 1975, G. McDonald, 3. M0132, Pa-
cific Grove, Monterey Co., Calif., 17 Oct.
1970, G. McDonald, 1. M0141, Carmel Pt.,
Monterey Co., Calif., 15 Oct. 1970, J. W. Ny-
bakken, 1. M0166, Pt. Pinos, Monterey Co.,
Calif., 26 Apr. 1967, V. Mansfield, 1. M0168,
Pigeon Pt., San Mateo Co., Calif., 30 Oct.
1966, J. W. Nybakken, 1.
Dendrodoris nigromaculata
M75-6-26-4, La Jolla, San Diego Co., Cal-
if., 26 June 1975, G. McDonald, 2.
Dendrodoris sp. a.
M72-7-16-2, Pirate s Cove, San Luis Obis-
po Co., Calif., 16 July 1972, G. McDonald, 1.
M73-11-12-1, Pirate's Cove, San Luis Obis-
po Co., Calif., 12 Nov. 1973, G. McDonald,
1. M72-9-23-1, Elkhorn Slough, Monterey
Co., Calif., 23 Sept. 1972, G. McDonald, 1.
M75-0-0-1, 17 Mile Dr., Monterey Co., Calif.,
1975, J. Harvey, 1.
Dendrodoris sp. b.
M71-7-7-6, Santa Monica Bay, Los Ange-
les Co., Calif., 7 July 1971, S. Anderson, 1.
M71-10-20-2, Los Angeles Co., Calif., 20 Oct.
1971, S. Anderson, 1. M71-10-9-1, Malibu,
Los Angeles Co., Calif., 9 Oct. 1971, S. An-
derson, 2. M75-10-17-1, Pescadero Pt.,
Monterey Co., Calif., 17 Oct. 1975, M. Sil-
berstein, 1.
CALIFORNIA NUDIBRANCHS 269
Dendronotus albus
M70-4-3-1, Pirate's Cove, San Luis Obis-
po Co., Calif., 3 Apr. 1970, G. McDonald, 1.
M70-6-5-2, Pirate's Cove, San Luis Obispo
Co., Calif., 5 June 1970, G. McDonald, 1.
M70-6-5-3, Pirate's Cove, San Luis Obispo
Co., Calif., 5 June 1970, G. McDonald, 1.
*M70-7-18-1, Shell Beach, San Luis Obispo
Co., Calif., 18 July 1970, G. McDonald, 2.
M71-5-26-4, Asilomar, Monterey Co., Calif.,
26 May 1971, G. McDonald, 5. M72-5-15-4,
Asilomar, Monterey Co., Calif., 15 May 1972,
G. McDonald, 1. M72-7-11-1, Pirate's Cove,
San Luis Obispo Co., Calif., 11 July 1972, G.
McDonald, 3. M73-5-6-3, Pt. Pinos, Monte-
rey Co., Calif., 6 May 1973, G. McDonald, 2.
M73-6-5-8, Asilomar, Monterey Co., Calif., 5
June 1973, 2. M78-7-21-3, Pt. Pinos, Mon-
terey Co., Calif., 21 July 1978, G. McDonald,
2. M0214, Asilomar, Monterey Co., Calif., 25
May 1971, G. McDonald, 2.
Dendronotus diversicolor
*M0506, Asilomar, Monterey Co., Calif., 14
Nov. 1974, J. Nybakken, 1. M76-8-2-3, Otter
Bay, British Columbia, Canada, 2 Aug. 1976,
P. Lambert, 1.
Dendronotus frondosus
M70-2-26-1, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 2.
M70-2-26-4, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 1.
M70-3-29-1, Morro Bay, San Luis Obispo
Co., Calif., 29 Mar. 1970, G. McDonald, 1.
M70-9-21-1, Monterey Bay, Calif., 21 Sept.
1970, S. Anderson, 3. *M70-10-7-1, Monte-
rey Bay, Calif., 7 Oct. 1970, G. McDonald, 4.
M71-11-17-9, Elkorn Slough, Monterey Co.,
Calif., 17 Nov. 1971, G. McDonald, 1. M75-
10-14-1, Monterey Bay, Calif., 14 Oct. 1975,
G. McDonald, 3. M75-10-23-1, Monterey Bay,
Calif., 23 Oct. 1975, P. Slattery, 20. M78-7-
21-20, Elkhorn Slough, Monterey Co., Calif.,
21 July 1978, G. McDonald, 1. M0213, Mon-
terey Bay, Calif., 7 Oct. 1970, G. McDonald,
4. M0215, Monterey Bay, Calif., 21 Sept.
1970, S. Anderson, 15. M0499, Monterey
Bay, Calif., 8 Oct. 1974, G. McDonald, 1.
Dendronotus iris
M71-2-15-4, Monterey Harbor, Monterey
Co., Calif., 15 Feb. 1971, G. McDonald, 1.
M71-5-23-1, Monterey Breakwater, Monte-
rey Co., Calif., 23 May 1971, G. McDonald,
1. M71-5-23-2, Monterey Breakwater, Mon-
terey Co., Calif., 23 May 1971, P. Clark, 1.
M71-5-23-3, Monterey Breakwater, Monte-
rey Co., Calif., 23 May 1971, G. McDonald,
2. M71-5-23-4, Monterey Breakwater, Mon-
terey Co., Calif., 23 May 1971, P. Clark, 1.
M70-8-18-1, Morro Bay, San Luis Obispo
Co., Calif., 18 Aug. 1970, R. Roller, 1. M72-
1-7-1, off Topanga Can., Los Angeles Co.,
Calif., 7 Jan. 1972, S. Anderson, 2. M72-3-
22-5, Monterey Breakwater, Monterey Co.,
Calif., 22 Mar. 1972, G. McDonald, 2. M72-
3-22-6, Monterey Breakwater, Monterey Co.,
Calif., 22 Mar. 1972, G. McDonald, 2. M73-
3-0-2, Los Angeles Co., Calif., Mar. 1973, В.
Fay, 2. M78-8-4-1, Monterey Breakwater,
Monterey Co., Calif., 4 Aug. 1978, G. Mc-
Donald, 4. M69-8-0-3, Friday Harbor, Wash.,
Aug. 1969, R. Roller, 1. MO221, Monterey
Breakwater, Monterey Co., Calif., 23 May
1971, G. McDonald, 1.
Dendronotus subramosus
M71-7-25-6, Morro Bay, San Luis Obispo
Co., Calif., 25 July 1971, G. McDonald, 1.
M71-8-8-3, Asilomar, Monterey Co., Calif., 8
Aug. 1971, G. McDonald, 3. M71-11-1-5,
Hazard Can., San Luis Obispo Co., Calif., 1
Nov. 1971, G. McDonald, 1. M71-11-4-3, As-
ilomar, Monterey Co., Calif., 4 Nov. 1971, D.
Shonman, 2. M72-5-15-3, Asilomar, Monte-
rey Co., Calif., 15 May 1972, G. McDonald,
1. M72-5-17-7, Carmel Pt., Monterey Co.,
Calif., 17 May 1972, G. McDonald, 1. M75-
8-8-2, Pt. Pinos, Monterey Co., Calif., 8 Aug.
1975, M. E. Anderson, 1. M75-11-2-7, Car-
mel Pt., Monterey Co., Calif., 2 Nov. 1975,
С. McDonald, 1. M0210 Asilomar, Monterey
Co., Calif., 25 May 1971, G. McDonald, 2.
M0314, Asilomar, Monterey Co., Calif., 15
May 1972, G. McDonald, 1.
Dirona albolineata
M70-12-12-2, Pigeon Pt., San Mateo Co.,
Calif., 12 Dec. 1970, G. McDonald, 1. M71-
5-25-7, Asilomar, Monterey Co., Calif., 25
Мау 1971, G. McDonald, 1. *M71-6-26-1,
Asilomar, Monterey Co., Calif., 26 June 1971,
G. McDonald, 1. M71-10-5-2, Asilomar, Mon-
terey Co., Calif., 5 Oct. 1971, G. McDonald,
1. M68-7-11-1, Shell Beach, San Luis Obispo
Co., Calif., 11 July 1968, R. A. Roller, 3. M72-
5-15-2, Asilomar, Monterey Co., Calif., 15
May 1972, G. McDonald, 2. M75-11-3-4, Pi-
geon Pt., San Mateo Co., Calif., 3 Nov. 1975,
G. McDonald, 1.
Dirona picta
M70-2-26-6, Morro Bay, San Luis Obispo
Co., Calif. 26 Feb. 1970, G. McDonald, 1.
M70-12-11-1, Asilomar, Monterey Co., Calif.,
270 MCDONALD
11 Dec. 1970, G. McDonald, 1. M71-1-26-5,
Morro Bay, San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 2. M71-1-26-17,
Hazard Can., San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 1. M71-2-8-1, Asi-
lomar, Monterey Co., Calif., 8 Feb. 1971, G.
McDonald, 3. M71-2-24-4, Morro Bay, San
Luis Obispo Co., Calif., 24 Feb. 1971, G.
McDonald, 2. M71-2-24-5, Morro Bay, San
Luis Obispo Co., Calif., 24 Feb. 1971, G.
McDonald, 1. *M71-5-26-5, Asilomar, Моп-
terey Co., Calif., 26 May 1971, G. McDonald,
3. M71-7-25-4, Morro Bay, San Luis Obispo
Co., Calif., 25 July 1971, G. McDonald, 1.
M71-10-7-3, Marina del Rey, Los Angeles
Co., Calif., 7 Oct. 1971, R. S. Anderson, 3.
M71-12-1-2, Asilomar, Monterey Co., Calif.,
1 Dec. 1971, D. Shonman, 1. M71-12-30-3,
Hazard Can., San Luis Obispo Co., Calif., 30
Dec. 1971, G. McDonald, 1. M72-5-17-3,
Carmel Pt., Monterey Co., Calif., 17 May
1972, G. McDonald, 1. M75-11-2-15, Carmel
Pt., Monterey Co., Calif., 2 Nov. 1975, G.
McDonald, 2. M78-7-21-11, Pt. Pinos, Mon-
terey Co., Calif., 21 July 1978, A. K. Mc-
Donald, 2. M0220, Asilomar, Monterey Co.,
Calif., 11 Nov. 1970, В. $. Anderson, 1.
Discodoris heathi
M70-6-28-2, Pirate's Cove, San Luis Obis-
po Co., Calif., 28 June 1970, G. McDonald,
1. M70-10-16-3, Asilomar, Monterey Co.,
Calif., 16 Oct. 1970, G. McDonald, 1. M70-
12-12-11, Pigeon Pt., San Mateo Co., Calif.,
12 Dec. 1970, J. Hansen, 1. M71-1-26-7,
Hazard Can., San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 1. M71-2-1-6, Port
San Luis, San Luis Obispo Co., Calif., 1 Feb.
1971, G. McDonald, 1. M71-11-2-2, Spoon-
er's Cove, San Luis Obispo Co., Calif., 2 Nov.
1971, G. McDonald, 1. M71-11-2-9, Spoon-
er's Cove, San Luis Obispo Co., Calif., 2 Nov.
1971, G. McDonald, 1. *M71-12-29-4, Pi-
rate's Cove, San Luis Obispo Co., Calif., 29
Dec. 1971, G. McDonald, 1. M75-11-2-9,
Carmel Pt., Monterey Co., Calif., 2 Nov. 1975,
G. McDonald, 5. M77-12-10-15, Asilomar,
Monterey Co., Calif., 10 Dec. 1977, G.
McDonald, 5. M0165, Pt. Pinos, Monterey
Co., Calif., 26 Apr. 1967, V. Mansfield, 1.
M0486, Carmel Pt., Monterey Co., Calif., 22
June 1974, M. E. Anderson, 1.
Discodoris sandiegensis
M69-9-3-10, San Diego, San Diego Co.,
Calif., 3 Sept. 1969, G. McDonald, 1. M69-
10-26-4, Pirate's Cove, San Luis Obispo Co.,
Calif., 26 Oct. 1969, G. McDonald, 2. M70-6-
28-5, Pirate s Cove, San Luis Obispo Co.,
Calif., 28 June 1970, G. McDonald, 1. M70-
10-14-1, Asilomar, Monterey Co. Calif., 14
Oct. 1970, G. McDonald, 1. M70-10-17-1,
Pacific Grove, Monterey Co., Calif., 17 Oct.
1970, G. McDonald, 2. M70-12-12-5, Pigeon
Pt., San Mateo Co., Calif., 12 Dec. 1970, G.
McDonald, 1. M71-1-25-3, Pirate's Cove, San
Luis Obispo Co., Calif., 25 Jan. 1971, G.
McDonald, 1. M71-11-2-4, Spooner's Cove,
San Luis Obispo Co., Calif., 2 Nov. 1971, G.
McDonald, 1. M71-11-2-8, Spooner's Cove,
San Luis Obispo Co., Calif., 2 Nov. 1971, G.
McDonald, 1. *M71-12-31-1, Shell Beach,
San Luis Obispo Co., Calif., 31 Dec. 1971,
G. McDonald, 5. M72-1-16-4, Monterey
Breakwater, Monterey Co., Calif., 16 Jan.
1972, R. Hilaski, 1. M72-1-16-9, Elkhorn
Slough, Monterey Co., Calif., 16 Jan. 1972,
G. McDonald, 4. M73-6-5-1, Asilomar, Mon-
terey Co., Calif., 5 June 1973, G. McDonald,
2. M75-7-18-1, Elkhorn Slough, Monterey
Co., Calif., 18 July 1975, G. McDonald, 1.
M75-10-21-4, Elkhorn Slough, Monterey Co.,
Calif., 21 Oct. 1975, О. Ituarte, 1. M75-11-2-
2, Carmel Pt., Monterey Co., Calif., 2 Nov.
1975, G. McDonald, 3. M75-11-4-1, Pesca-
dero Pt., Monterey Co., Calif., 4 Nov. 1975,
G. McDonald, 2. MO160, Del Monte Beach,
Monterey Co., Calif., 1 Apr. 1967, V. Mans-
field, 1. M0487, Monterey Breakwater, Mon-
terey Co., Calif., 8 July 1974, M. E. Ander-
son, 1.
Doridella steinbergae
*M71-7-7-4, Morro Bay, San Luis Obispo
Co., Calif., 7 July 1971, G. McDonald, 6. M73-
6-21-2, Pirate's Cove, San Luis Obispo Co.,
Calif., 21 June 1973, G. McDonald, 40. M75-
10-14-3, Monterey Bay, Calif., 14 Oct. 1975,
G. McDonald, 20.
Doris (s.l.) sp.
M71-9-14-1, Malibu, Los Angeles Co., Cal-
if., 14 Sept. 1971, В. $. Anderson, 1. M71-
9-14-2, Malibu, Los Angeles Co., Calif., 14
Sept. 1971, R. S. Anderson, 1. *M71-12-2-
12, La Jolla, San Diego Co., Calif., 2 Dec.
1971, В. $. Anderson, 4. M75-6-26-1, La Jol-
la, San Diego Co., Calif. 26 June 1975, G.
McDonald, 1. M75-6-26-6, Pescadero Pt.,
Monterey Co., Calif., 26 June 1975, M. Sil-
berstein, 1.
Doto amyra
*M72-5-15-5, Asilomar, Monterey Co., Cal-
if., 15 Мау 1972, G. McDonald, 3. M73-5-5-
9, Asilomar, Monterey Co., Calif., 5 May 1973,
CALIFORNIA NUDIBRANCHS 271
С. McDonald, 4. M73-6-5-12, Asilomar, Моп-
terey Co., Calif., 5 June 1973, G. McDonald,
4. M78-6-22-4, Carmel Pt., Monterey Co.,
Calif., 22 June 1978, G. McDonald, 5. MO315,
Asilomar, Monterey Co., Calif., 15 May 1972,
G. McDonald, 2.
Doto kya
*M71-10-17-2, Morro Bay, San Luis Obis-
po Co., Calif., 17 Oct. 1971, G. McDonald, 1.
M73-5-6-7, Pt. Pinos, Monterey Co., Calif., 6
May 1973, G. McDonald, 1. M73-6-5-15, As-
ilomar, Monterey Co., Calif., 5 June 1973, С.
McDonald, 6. M75-10-26-2, Morro Bay, San
Luis Obispo Co., Calif., 26 Oct. 1975, G.
McDonald, 12.
Eubranchus misakiensis
*Mukaishima, Inland Sea of Seto, Japan,
22 Mar. 1962, K. Baba, 2.
Eubranchus olivaceus
*M73-5-5-4, Asilomar, Monterey Co., Cal-
if., 5 May 1973, G. McDonald, 1.
Eubranchus rustyus
M70-11-16-5, Elkhorn Slough, Monterey
Co., Calif. 16 Nov. 1970, G. McDonald, 1.
M71-1-26-8, Morro Bay, San Luis Obispo
Co., Calif., 26 Jan. 1971, G. McDonald, 3.
M71-6-26-5, Asilomar, Monterey Co., Calif.,
26 June 1971, G. McDonald, 1. M71-10-17-
3, Morro Bay, San Luis Obispo Co., Calif.,
17 Oct. 1971, G. McDonald, 1. *M73-5-5-1,
Asilomar, Monterey Co., Calif., 5 May 1973,
G. McDonald, 7. M73-5-5-2, Asilomar, Mon-
terey Co., Calif., 5 May 1973, G. McDonald,
2. M73-5-6-1, Pt. Pinos, Monterey Co., Calif.,
6 May 1973, G. McDonald, 8. M75-10-19-1,
Monterey Breakwater, Monterey Co., Calif.
19 Oct. 1975, A. K. McDonald, 2. M75-10-
26-8, Morro Bay, San Luis Obispo Co., Cal-
if., 26 Oct. 1975, G. McDonald, 4. M78-6-22-
10, Carmel Pt., Monterey Co., Calif., 22 June
1978, G. McDonald, 1. MO316, Asilomar,
Monterey Co., Calif., 15 May 1972, J. W. Ny-
bakken, 1.
Fiona pinnata
M70-9-21-2, Monterey Bay, Calif., 21 Sept.
1970, R. S. Anderson, 2. M70-10-5-5, Mon-
terey Bay, Calif., 5 Oct. 1970, G. McDonald,
4. M70-10-5-6, Monterey Bay, Calif., 5 Oct.
1970, G. McDonald, 11. M70-11-1-1, Mon-
terey Bay, Calif., 1 Nov. 1970, R. S. Ander-
son, 6. *M70-11-1-3, Monterey Bay, Calif., 1
Nov. 1970, R. S. Anderson, 9. M70-11-1-4,
Monterey Bay, Calif., 1 Nov. 1970, R. S. An-
derson, 3. M72-8-9-1, Monterey Bay, Calif.,
9 Aug. 1972, G. McDonald, 14. M0031, Mon-
terey Bay, Calif., Sept. 1971, 2. M0193,
Monterey Bay, Calif., 1 Nov. 1970, R. S. An-
derson, 7. M0194, Monterey Bay, Calif., 8
Nov. 1970, В. $. Anderson, 7. M0195, Mon-
terey Bay, Calif., 12 Sept. 1970, R. S. An-
derson, 9. M0199, Monterey Bay, Calif., 5
Oct. 1970, G. McDonald, 9.
Hallaxa chani
M71-12-30-2, Hazard Can., San Luis Obis-
po Co., Calif., 30 Dec. 1971, G. McDonald,
2. M71-12-31-6, Shell Beach, San Luis Obis-
po Co., Calif., 31 Dec. 1971, G. McDonald,
2. M72-1-15-2, Asilomar, Monterey Co., Cal-
Н., 15 Jan. 1972, G. McDonald, 1. *M72-10-
22-2, Asilomar, Monterey Co., Calif., 22 Oct.
1972, G. McDonald, 1. M75-10-26-4, Morro
Bay, San Luis Obispo Co., Calif., 26 Oct.
1975, G. McDonald, 3. M77-11-11-2, Scott
Cr., Santa Cruz Co., Calif., 11 Nov. 1977, G.
McDonald, 1.
Hancockia californica
M70-9-21-3, Monterey Bay, Calif., 21 Sept.
1970, R. S. Anderson, 3. M70-10-14-3, Asi-
lomar, Monterey Co., Calif., 14 Oct. 1970, G.
McDonald, 1. M71-5-25-6, Asilomar, Monte-
rey Co., Calif., 25 May 1971, G. McDonald,
1. M71-6-26-2, Asilomar, Monterey Co., Cal-
if., 26 June 1971, С. McDonald, 1. M71-7-25-
2, Morro Bay, San Luis Obispo Co., Calif.,
25 July 1971, G. McDonald, 12. *M71-8-8-1,
Asilomar, Monterey Co., Calif., 8 Aug. 1971,
G. McDonald, 25. M71-7-25-10, Morro Bay,
San Luis Obispo Co., Calif., 25 July 1971, G.
McDonald, 3. M72-8-9-3, Asilomar, Monte-
rey Co., Calif., 9 Aug. 1972, G. McDonald, 2.
M73-6-5-10, Asilomar, Monterey Co., Calif.,
5 June 1973, G. McDonald, 3. M75-8-8-3,
Pacific Grove, Monterey Co., Calif., 8 Aug.
1975, M. E. Anderson, 1. M76-11-21-1, Asi-
lomar, Monterey Co., Calif., 21 Nov. 1976, G.
McDonald, 1. M78-7-21-1, Pt. Pinos, Mon-
terey Co., Calif., 21 July 1978, G. McDonald,
6. M0206, Monterey Bay, Calif., 12 Sept.
1970, R. S. Anderson, 5.
Hopkinsia rosacea
M70-6-28-1, Pirate's Cove, San Luis Obis-
po Co., Calif., 28 June 1970, G. McDonald,
1. M70-6-28-4, Pirate's Cove, San Luis Obis-
ро Co., Calif., 28 June 1970, G. McDonald,
1. M71-1-25-1, Sunset Palisades, San Luis
Obispo Co., Calif., 25 Jan. 1971, G. Mc-
Donald, 1. M71-1-26-8, Hazard Can., San
Luis Obispo Co., Calif., 26 Jan. 1971, G.
McDonald, 1. M71-11-2-10, Spooner's Cove,
272 MCDONALD
San Luis Obispo Co., Calif., 2 Nov. 1971, G.
McDonald, 1. M71-11-2-11, Spooner's Cove,
San Luis Obispo Co., Calif., 2 Nov. 1971, G.
McDonald, 1. M72-5-17-2, Carmel Pt., Mon-
terey Co., Calif., 17 May 1972, G. McDonald,
2. M73-4-9-5, Carmel Pt., Monterey Co., Cal-
if., 9 Apr. 1973, G. McDonald, 3. M73-6-1-1,
Pacific Grove, Monterey Co., Calif., 1 June
1973, G. McDonald, 4. M75-10-5-2, Carmel
Pt., Monterey Co., Calif., 5 Oct. 1975, G.
McDonald, 10. MO136, Asilomar, Monterey
Co., Calif., 11 Nov. 1970, G. McDonald, 1.
M0188, Pacific Grove, Monterey Co., Calif.,
16 Oct. 1966, J. W. Nybakken, 1. MO202, Pt.
Pinos, Monterey Co., Calif., 27 Apr. 1967, V.
Mansfield, 1.
Hypselodoris californiensis
M71-8-16-1, Malibu, Los Angeles Co., Cal-
if., 16 Aug. 1971, В. $. Anderson, 1. M71-8-
2-3, Malibu, Los Angeles Co., Calif., 2 Aug.
1971, В. $. Anderson, 1. M71-8-2-4, Malibu,
Los Angeles Co., Calif., 2 Aug. 1971. В. $.
Anderson, 1. *М71-10-0-2, Malibu, Los Ап-
geles Co., Calif., Oct. 1971, R. S. Anderson,
1. M71-10-0-3, Malibu, Los Angeles Co., Cal-
if., Oct. 1971, В. $. Anderson, 1. M71-10-0-
4, Malibu, Los Angeles Co., Calif., Oct. 1971,
В. $. Anderson, 1. M71-11-23-1, Malibu, Los
Angeles Co., Calif., 23 Nov. 1971, В. $. An-
derson, 1. M71-11-23-4, Malibu, Los Angeles
Co., Calif., 23 Nov. 1971, В. $. Anderson, 1.
Laila cockerelli
M70-2-5-2, Pecho, San Luis Obispo Co.,
Calif., 5 Feb. 1970, G. McDonald, 2. *M71-5-
25-8, Asilomar, Monterey Co., Calif., 25 May
1971, G. McDonald, 1. M71-6-10-1, Monte-
rey Breakwater, Monterey Co., Calif., 10 June
1971, О. В. Lewis, 1. M71-10-5-1, Asilomar,
Monterey Co., Calif., 5 Oct. 1971, G. Mc-
Donald, 2. M71-11-3-1, Carmel Pt., Monterey
Co., Calif., 3 Nov. 1971, G. McDonald, 5.
M71-11-4-4, Carmel Pt., Monterey Co., Cal-
if., 4 Nov. 1971, G. McDonald, 1. M71-12-2-
11, Carmel Pt., Monterey Co., Calif., 2 Dec.
1971, G. McDonald, 2. M69-7-30-1, Pt. Loma,
San Diego Co., Calif., 30 July 1969, R. A.
Roller, 1. M72-1-16-7, Monterey Breakwater,
Monterey Co., Calif., 16 Jan. 1972, R. Hilas-
ki, 1. M72-2-14-1, Asilomar, Monterey Co.,
Calif., 14 Feb. 1972, G. McDonald, 1. M72-
5-15-6, Asilomar, Monterey Co., Calif., 15
May 1972, G. McDonald, 1. M72-5-17-6,
Carmel Pt., Monterey Co., Calif., 17 May
1972, G. McDonald, 2. M73-4-9-4, Carmel Pt.,
Monterey Co., Calif., 9 Apr. 1973, G. Mc-
Donald, 1. M73-5-6-4, Pt. Pinos, Monterey
Co., Calif., 6 May 1973, G. McDonald, 1.
M73-6-1-3, Pacific Grove, Monterey Co.,
Calif., 1 June 1973, G. McDonald, 2. M75-6-
25-4, La Jolla, San Diego Co., Calif., 25 June
1975, G. McDonald, 1. M75-10-5-3, Carmel
Pt., Monterey Co., Calif., 5 Oct. 1975, G.
McDonald, 1. M75-11-2-14, Carmel Pt.,
Monterey Co., Calif., 2 Nov. 1975, G. Mc-
Donald, 8. M0212, Carmel Pt., Monterey Co.,
Calif., 15 Oct. 1970, Е. Stark; 1:
Melibe leonina
M69-10-3-1, Morro Bay, San Luis Obispo
Co., Calif., 3 Oct. 1969, G. McDonald, 3. M70-
10-7-2, Monterey Bay, Calif., 7 Oct. 1970, G.
McDonald, 1. M70-12-2-2, Monterey Bay,
Calif., 2 Dec. 1970, G. McDonald, 2. M75-10-
14-5, Monterey Bay, Calif., 14 Oct. 1975, J.
Harvey, 1. MO218, Monterey Bay, Calif., 7
Oct. 1970, G. McDonald, 1. M0496, Elkhorn
Slough, Monterey Co., Calif., Sept. 1974, C.
Keusink, 1.
Okenia angelensis
M70-11-8-2, Morro Bay, San Luis Obispo
Co., Calif., 8 Nov. 1970, G. McDonald, 1.
M70-11-27-3, Morro Bay, San Luis Obispo
Co., Calif., 27 Nov. 1970, G. McDonald, 2.
M71-1-26-13, Morro Bay, San Luis Obispo
Co., Calif., 26 Jan. 1971, G. McDonald, 3.
M71-2-24-2, Morro Bay, San Luis Obispo
Co., Calif., 24 Feb. 1971, G. McDonald, 2.
M71-2-24-7, Morro Bay, San Luis Obispo
Co., Calif., 24 Feb. 1971, G. McDonald, 3.
*M71-10-17-7, Morro Bay, San Luis Obispo
Co., Calif., 17 Oct. 1971, G. McDonald, 13.
M71-10-22-7, Monterey Harbor, Monterey
Co., Calif., 22 Oct. 1971, G. McDonald, 7.
M72-10-29-1, Morro Bay, San Luis Obispo
Co., Calif., 29 Oct. 1972, G. McDonald, 4.
M69-11-23-1, Morro Bay, San Luis Obispo
Co., Calif., 23 Nov. 1969, В. A. Roller, 1. M69-
10-12-4, Morro Bay, San Luis Obispo Co.,
Calif., 12 Oct. 1969, R. A. Roller, 4. M75-10-
26-7, Morro Bay, San Luis Obispo Co., Cal-
if., 26 Oct. 1975, G. McDonald, 10. M0025,
Monterey Harbor, Monterey Co., Calif., 21
Oct. 1971, G. McDonald, 14. M0171, Morro
Bay, San Luis Obispo Co., Calif., 8 Nov. 1970,
G. McDonald, 1.
Okenia plana
*M72-11-0-1, Pt. Richmond, San Francis-
co Bay, Calif., Nov. 1972, E. Lyke, 2.
Onchidoris bilamellata
*M70-10-22-1, Moss Landing, Monterey
Co., Calif., 22 Oct. 1970, R. S. Anderson, 10.
M70-10-22-3, Moss Landing, Monterey Co.,
CALIFORNIA NUDIBRANCHS 273
Calif., 22 Oct. 1970, В. $. Anderson, 11. M70-
10-22-5, Moss Landing, Monterey Co., Calif.,
22 Oct. 1970, R. S. Anderson, 12. M71-2-15-
2, Monterey Harbor, Monterey Co., Calif., 15
Feb. 1971, G. McDonald, 1. M71-11-17-3,
Elkhorn Slough, Monterey Co., Calif., 17 Nov.
1971, G. McDonald, 2. M72-2-10-1, Elkhorn
Slough, Monterey Co., Calif., 10 Feb. 1972,
G. McDonald, 7. M65-4-3-1, England, 3 Apr.
1965, T. E. Thompson, 1. M75-7-18-2, Elk-
horn Slough, Monterey Co., Calif., 18 July
1975, G. McDonald, 1. MO129, Moss Land-
ing, Monterey Co., Calif., 22 Oct. 1970, R. S.
Anderson, 14. M0172, Moss Landing, Mon-
terey Co., Calif., 22 Oct. 1970, R. S. Ander-
son, 6. M0173, Moss Landing, Monterey Co.,
Calif., 22 Oct. 1970, R. S. Anderson, 20.
Onchidoris hystricina
M70-7-18-5, Shell Beach, San Luis Obispo
Co., Calif., 18 July 1970, G. McDonald, 3.
M70-7-18-6, Shell Beach, San Luis Obispo
Co., Calif., 18 July 1970, G. McDonald, 3.
*M71-6-28-2, Morro Bay, San Luis Obispo
Co., Calif., 28 June 1971, В. A. Roller, 3. M71-
7-7-2, Morro Bay, San Luis Obispo Co., Cal-
if., 7 July 1971, G. McDonald, 1. M71-10-17-
5, Morro Bay, San Luis Obispo Co., Calif.,
17 Oct. 1971, G. McDonald, 2. M75-10-26-9,
Morro Bay, San Luis Obispo Co., Calif., 26
Oct. 1975, G. McDonald, 1. M78-8-4-3, Mon-
terey Breakwater, Monterey Co., Calif., 4
Aug. 1978, G. McDonald, 2. M0126, Asilo-
mar, Monterey Co., Calif., 14 Oct. 1970, J.
W. Nybakken, 1.
Onchidoris sp.
*M71-1-26-15, Hazard Can., San Luis
Obispo Co., Calif., 26 Jan. 1971, G. Mc-
Donald, 3. M69-8-24-2, Friday Harbor, Wash.,
24 Aug. 1969, R. A. Roller, 1. M71-12-30-1,
Hazard Can., San Luis Obispo Co., Calif., 30
Dec. 1971, G. McDonald, 2. M75-0-0-1,
Monastery Beach, Monterey Co., Calif., 1975,
A. К. McDonald, 3. M0505, Monastery Beach,
Monterey Co., Calif., 18 Jan. 1975, A. K.
McDonald, 5.
Phidiana crassicornis
M69-10-26-9, Pirate's Cove, San Luis
Obispo Co., Calif. 26 Oct. 1969, G. Mc-
Donald, 3. M69-10-26-11, Pirate's Cove, San
Luis Obispo Co., Calif., 26 Oct. 1969, G.
McDonald, 1. M70-3-29-2, Morro Bay, San
Luis Obispo Co., Calif., 29 Mar. 1970, G.
McDonald, 1. M70-10-17-5, Elkhorn Slough,
Monterey Co., Calif., 17 Oct. 1970, G. Mc-
Donald, 1. M70-10-28-3, Elkhorn Slough,
Monterey Co., Calif., 28 Oct. 1970, G. Mc-
Donald, 1. M71-10-22-1, Monterey Harbor,
Monterey Co., Calif., 22 Oct. 1971, G. Mc-
Donald, 2. M71-11-17-6, Elkhorn Slough,
Monterey Co., Calif., 17 Nov. 1971, G.
McDonald, 1. M71-11-29-1, Morro Bay, San
Luis Obispo Co., Calif., 29 Nov. 1971, G.
McDonald, 1. M72-10-18-3, Elkhorn Slough,
Monterey Co., Calif., 18 Oct. 1972, G. Mc-
Donald, 5. M72-10-18-8, Elkhorn Slough,
Monterey Co., Calif., 18 Oct. 1972, G. Mc-
Donald, 3. M75-10-5-7, Carmel Pt., Monterey
Co., Calif., 5 Oct. 1975, G. McDonald, 1. M75-
10-26-6, Morro Bay, San Luis Obispo Co.,
Calif., 26 Oct. 1975, G. McDonald, 1. M75-
11-2-16, Carmel Pt., Monterey Co., Calif., 2
Nov. 1975, G. McDonald, 1. M76-7-0-1, E.
Redonda Island, B. C., Canada, July 1976,
M. E. Anderson, 6. M0191, Pt. Pinos, Mon-
terey Co., Calif., 26 Apr. 1967, V. Mansfield,
1. M0204, Elkhorn Slough, Monterey Co.,
Calif., 17 Oct. 1970, G. McDonald, 2.
Phidiana hiltoni
M69-10-26-7, Pirate's Cove, San Luis
Obispo Co., Calif. 26 Oct. 1969, G. Mc-
Donald, 1. *M71-1-25-2, Sunset Palisades,
San Luis Obispo Co., Calif., 25 Jan. 1971, G.
McDonald, 6. M71-1-26-6, Hazard Can., San
Luis Obispo Co., Calif., 26 Jan. 1971, G.
McDonald, 6. M71-12-2-10, Carmel Pt.,
Monterey Co., Calif., 2 Dec. 1971, G. Mc-
Donald, 1. M71-12-28-3, Sunset Palisades,
San Luis Obispo Co., Calif., 28 Dec. 1971,
G. McDonald, 3. M72-5-17-4, Carmel Pt.,
Monterey Co., Calif., 17 May 1972, ©. Mc-
Donald, 1. M75-10-4-2, Monastery Beach,
Monterey Co., Calif., 4 Oct. 1975, A. K.
McDonald, 1. M75-11-2-11, Carmel Pt.,
Monterey Co., Calif., 2 Nov. 1975, G. Mc-
Donald, 2. M78-7-21-12, Pt. Pinos, Monterey
Co., Calif., 21 July 1978, G. McDonald, 1.
M0187, Carmel Pt., Monterey Co., Calif., 26
May 1967, R. Schinoke, 1. MO310, Carmel
Pt., Monterey Co., Calif., 17 May 1972, G.
McDonald, 1.
Phidiana morroensis
M71-10-17-4, Morro Bay, San Luis Obispo
Со., Calif., 17 Oct. 1971, G. McDonald, 1.
M71-11-17-2, Elkhorn Slough, Monterey Co.,
Calif., 17 Nov. 1971, G. McDonald, 5. M71-
12-4-3, Elkhorn Slough, Monterey Co., Calif.,
4 Dec. 1971, G. McDonald, 5. *M72-9-21-1,
Monterey Bay, Calif., 21 Sept. 1972, J. Oliv-
er, 3. M72-11-1-5, Elkhorn Slough, Monterey
Co., Calif., 1 Nov. 1972, G. McDonald, 3.
274 MCDONALD
Phidiana stearnsi
M75-6-0-1, La Jolla, San Diego Co., Calif.,
June 1975, J. R. Lance, 1. *CASIZ, Santa
Barbara, Santa Barbara Co., Calif., Aug.
1966, J. E. Steinberg, 1.
Platydoris macfarlandi
"CASIZ 9511, Pismo Beach, San Luis
Obispo Co., Calif., 23 Oct. 1950, W. E. Ripley
et al., 1, paratype. CASIZ 9512, Pismo Beach,
San Luis Obispo Co., Calif., 23 Oct. 1950, W.
E. Ripley et al., 1, paratype.
Polycera atra
M70-2-26-7, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 1.
M70-3-31-1, Morro Bay, San Luis Obispo
Co., Calif., 31 Mar. 1970, G. McDonald, 1.
M70-10-22-4, Monterey Harbor, Monterey
Co., Calif., 22 Oct. 1970, G. McDonald, 1.
M70-10-28-2, Elkhorn Slough, Monterey Co.,
Calif. 28 Oct. 1970, G. McDonald, 1. M71-1-
26-16, Morro Bay, San Luis Obispo Co., Cal-
if., 26 Jan. 1971, G. McDonald, 1. M71-7-25-
9, Morro Bay, San Luis Obispo Co., Calif.,
25 July 1971, G. McDonald, 1. M71-10-17-8,
Morro Bay, San Luis Obispo Co., Calif., 17
Oct. 1971, G. McDonald, 2. M71-11-22-1,
Morro Bay, San Luis Obispo Co., Calif., 22
Nov. 1971, G. McDonald, 2. *M72-3-12-1,
Morro Bay, San Luis Obispo Co., Calif., 12
Mar., 1972, G. McDonald, 2. M72-11-1-4,
Elkhorn Slough, Monterey Co., Calif., 1 Nov.
1972, G. McDonald, 1. M75-10-21-2, Elkhorn
Slough, Monterey Co., Calif., 21 Oct. 1975,
D. Ituarte, 2. M75-10-26-1, Morro Bay, San
Luis Obispo Co., Calif., 26 Oct. 1975, G.
McDonald, 2. M75-11-11-1, Elkhorn Slough,
Monterey Co., Calif., 11 Nov. 1975, B. S. An-
trim, 4. MO209, Elkhorn Slough, Monterey
Co., Calif., 28 Oct. 1970, G. McDonald, 1.
Polycera hedgpethi
M70-11-8-3, Morro Bay, San Luis Obispo
Co., Calif., 8 Nov. 1970, G. McDonald, 1.
M71-6-28-1, Morro Bay, San Luis Obispo
Co., Calif., 28 June 1971, R. A. Roller, 2.
*M71-7-7-5, Morro Bay, San Luis Obispo Co.,
Calif., 7 July 1971, G. McDonald, 2. M71-9-
20-1, Redondo Beach, Los Angeles Co., Cal-
if., 20 Sept. 1971, В. S. Anderson, 1. M71-
11-22-2, Morro Bay, San Luis Obispo Co.,
Calif. 22 Nov. 1971, G. McDonald, 1. M71-
1-26-26, Morro Bay, San Luis Obispo Co.,
Calif., 26 Jan. 1971, В. A. Roller, 1.
Polycera tricolor
*M71-8-31-3, Santa Monica Bay, Los An-
geles Co., Calif., 31 Aug. 1971, R. S. Ander-
son, 1. M71-8-31-4, Santa Monica Bay, Los
Angeles Co., Calif., 31 Aug. 1971, В. $. An-
derson, 1. M71-11-26-1, Santa Monica Bay,
Los Angeles Co., Calif., 26 Nov. 1971, В. $.
Anderson, 1. M71-11-26-2, Santa Monica
Bay, Los Angeles Co., Calif., 26 Nov. 1971,
В. $. Anderson, 1.
Polycera zosterae
*Prepared slide of radula, Friday Harbor,
Wash., 23 Apr. 1973, R. A. Roller, 1.
Precuthona divae
M71-8-8-5, Asilomar, Monterey Co., Calif.,
8 Aug. 1971, G. McDonald, 1. M71-11-2-6,
Spooner's Cove, San Luis Obispo Co., Calif.,
2 Nov. 1971, G. McDonald, 3. M71-11-4-1,
Asilomar, Monterey Co., Calif., 4 Nov. 1971,
D. Shonman, 1. M72-1-5-1, Dume Rock, Los
Angeles Co., Calif., 5 Jan. 1972, R. S. An-
derson, 1. M72-5-17-5, Carmel Pt., Monterey
Co., Calif., 17 May 1972, G. McDonald, 2.
M76-8-5-1, Lasqueti Island, B. C., Canada, 5
Aug. 1976, М. E. Anderson, 5. M0196, Asi-
lomar, Monterey Co., Calif., 11 Dec. 1970, J.
W. Nybakken, 1.
Rostanga pulchra
M70-4-3-5, Pirate's Cove, San Luis Obis-
po Co., Calif., 3 Apr. 1970, G. McDonald, 1.
M70-11-11-1, Asilomar, Monterey Co., Calif.,
11 Nov. 1970, G. McDonald, 1. M71-1-25-5,
Pirate's Cove, San Luis Obispo Co., Calif.,
25 Jan. 1971, G. McDonald, 1. M71-1-26-9,
Hazard Can., San Luis Obispo Co., Calif., 26
Jan. 1971, G. McDonald, 3. M71-1-27-1, Shell
Beach, San Luis Obispo Co., Calif., 27 Jan.
1971, G. McDonald, 1. M71-6-13-1, Monte-
rey Breakwater, Monterey Co., Calif., 13 June
1971, G. McDonald, 1. M71-11-1-3, Hazard
Can., San Luis Obispo Co., Calif., 1 Nov.
1971, G. McDonald, 1. M71-12-2-9, Carmel
Pt., Monterey Co., Calif., 2 Dec. 1971, G.
McDonald, 2. M71-12-28-6, Dume Rock, Los
Angeles Co., Calif., 28 Dec. 1971, R. S. An-
derson, 2. M71-12-29-3, Pirate's Cove, San
Luis Obispo Co., Calif., 29 Dec. 1971, С.
McDonald, 2. M71-12-31-5, Shell Beach, San
Luis Obispo Co., Calif., 31 Dec. 1971, G.
McDonald, 2. M73-1-16-1, Asilomar, Monte-
rey Co., Calif., 16 Jan. 1973, G. McDonald,
1. M73-6-5-9, Asilomar, Monterey Co., Calif.,
5 June 1973, G. McDonald, 3. M75-11-3-3,
Pigeon Pt., San Mateo Co., Calif., 3 Nov.
1975, G. McDonald, 1. M76-2-21-1, Monas-
tery Beach, Monterey Co., Calif., 21 Feb.
1976, A. K. McDonald, 3. M0163, Pt. Pinos,
Monterey Co., Calif., 27 Apr. 1967, V. Mans-
CALIFORNIA NUDIBRANCHS 275
field, 1. M0308, Rocky Pt., Monterey Co.,
Calif., 14 May 1972, J. W. Nybakken, 1.
M0309, Rocky Pt., Monterey Co., Calif., 14
May 1972, G. McDonald, 2.
Sclerodoris tanya
M74-3-27-1, Mission Bay, San Diego Co.,
Calif., 27 Mar. 1974, J. Patton, 1. *M73-9-0-
1, Mission Bay, San Diego Co., Calif., Sept.
1973, M. Patton, 1.
Spurilla chromosoma
*M73-4-18-2, Puertecitos, Baja Calif., Mex.,
18 Apr. 1973, M. Silberstein, 1. M75-1-26-3,
Punta Mita, Nayarit, Mex., 26 Jan. 1975, G.
McDonald, 1.
Spurilla olivae
M70-3-3-1, San Simeon, San Luis Obispo
Co., Calif., 3 Mar. 1970, P. Clark, 1. M71-1-
26-11, Morro Bay, San Luis Obispo Co., Cal-
if., 26 Jan. 1971, G. McDonald, 1. M71-5-26-
2, Asilomar, Monterey Co., Calif., 26 May
1971, В. 5. Anderson, 1. M71-6-10-2, Моп-
{егеу Breakwater, Monterey Co., Calif., 10
June 1971, G. McDonald, 1. *M71-9-14-4,
Malibu, Los Angeles Co., Calif., 14 Sept.
1971, R. S. Anderson, 2. M71-10-22-2, Mon-
terey Harbor, Monterey Co., Calif., 22 Oct.
1971, G. McDonald, 1. M71-11-22-10, Mali-
bu, Los Angeles Co., Calif., 22 Nov. 1971, В.
S. Anderson, 1. M73-4-7-1, Asilomar, Mon-
terey Co., Calif., 7 Apr. 1973, G. McDonald,
1. M73-4-9-3, Carmel Pt., Monterey Co., Cal-
if., 9 Apr. 1973, G. McDonald, 1. M0024,
Monterey Harbor, Monterey Co., Calif., 21
Oct. 1970, G. McDonald, 1. M0217, Monte-
rey Breakwater, Monterey Co., Calif., 18 May
1971, э/ Расе, 1.
Tenellia adspersa
*M0580, Elkhorn Slough, Monterey Co.,
Calif., 3 June 1977, J. W. Cooper, 5.
Thordisa bimaculata
M74-4-3-1, Escondido Can., San Diego
Co., Calif., 3 Apr. 1974, M. Patton, 1. *M75-
6-26-2, La Jolla, San Diego Co., Calif., 26
June 1975, G. McDonald, 1.
Tochuina tetraquetra
M76-7-30-1, Roffey Island, B. C., Can., 30
July 1976, P. Lambert, 1. M0472, Monterey
Bay, Calif., 10 Mar. 1974, G. McDonald, 1.
M0500, Monterey Bay, Calif., 8 Oct. 1974, G.
McDonald, 1.
Trapania velox
M71-2-1-1, Port San Luis, San Luis Obis-
po Co., Calif., 1 Feb. 1971, G. McDonald, 5.
*M71-2-1-4, Port San Luis, San Luis Obispo
Co., Calif., 1 Feb. 1971, G. McDonald, 5.
M71-2-2-3, Port San Luis, San Luis Obispo
Co., Calif., 2 Feb. 1971, G. McDonald, 7.
Triopha catalinae
M69-10-12-1, Port San Luis, San Luis
Obispo Co., Calif., 12 Oct. 1969, G. Mc-
Donald, 1. M69-10-12-2, Port San Luis, San
Luis Obispo Co., Calif., 12 Oct. 1969, G.
McDonald, 1. M70-1-6-7, Sunset Palisades,
San Luis Obispo Co., Calif., 6 Jan. 1970, G.
McDonald, 1. M70-10-14-7, Asilomar, Mon-
terey Co., Calif., 14 Oct. 1970, G. McDonald,
1. M71-5-25-9, Asilomar, Monterey Co., Cal-
if., 25 May 1971, G. McDonald, 1. M71-7-7-
1, Morro Bay, San Luis Obispo Co., Calif., 7
July 1971, G. McDonald, 1. M71-12-2-6, Car-
mel Pt., Monterey Co., Calif., 2 Dec. 1971, С.
McDonald, 1. *M72-1-16-6, Monterey Break-
water, Monterey Co., Calif., 16 Jan. 1972, R.
Hilaski, 1. M72-4-26-4, Monterey Breakwa-
ter, Monterey Co., Calif., 26 Apr. 1972, P.
Clark, 1. M72-5-17-1, Carmel Pt., San Luis
Obispo Co., Calif., 17 May 1972, G. Mc-
Donald, 1. M73-6-5-14, Asilomar, Monterey
Co., Calif., 5 June 1973, G. McDonald, 1.
M75-11-2-12, Carmel Pt., Monterey Co., Cal-
if., 2 Nov. 1975, G. McDonald, 2. M0161, Pi-
geon Pt., San Mateo Co., Calif., 30 Oct. 1966,
J. W. Nybakken, 1. M0162, Carmel Pt., Mon-
terey Co., Calif. 26 May 1967, R. Schinoke,
1. M0211, Asilomar, Monterey Co., Calif., 14
Oct. 1970, G. McDonald, 1.
Triopha occidentalis
M70-2-26-3, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 1.
M70-2-26-8, Morro Bay, San Luis Obispo
Co., Calif., 26 Feb. 1970, G. McDonald, 1.
M70-10-14-6, Asilomar, Monterey Co., Calif.,
14 Oct. 1970, G. McDonald, 1. *M71-7-25-8,
Morro Bay, San Luis Obispo Co., Calif., 25
July 1971, G. McDonald, 1. M72-1-9-1, Mon-
terey Breakwater, Monterey Co., Calif., 9 Jan.
1972, R. Hilaski, 1. M73-11-11-1, Morro Bay,
San Luis Obispo Co., Calif., 11 Nov. 1973,
G. McDonald, 4. M73-11-11-4, Morro Bay,
San Luis Obispo Co., Calif., 11 Nov. 1973,
G. McDonald, 2. M75-6-25-5, La Jolla, San
Diego Co., Calif., 25 June 1975, G. Mc-
Donald, 1. M0170, Monterey Bay, Calif., 20
Dec. 1970, G. McDonald, 1.
Triopha maculata
M69-10-26-2, Pirate's Cove, San Luis
Obispo, Calif., 26 Oct. 1969, G. McDonald,
4. *M69-10-26-5, Pirate's Cove, San Luis
276 MCDONALD
Obispo, Calif., 26 Oct. 1969, G. McDonald,
8. M70-7-18-8, Shell Beach, San Luis Obispo
Co., Calif., 18 July 1970, G. McDonald, 3.
M70-10-14-5, Asilomar, Monterey Co., Calif.,
14 Oct. 1970, G. McDonald, 1. M70-10-16-2,
Asilomar, Monterey Co., Calif., 16 Oct. 1970,
G. McDonald, 1. M71-2-24-6, Morro Bay, San
Luis Obispo Co., Calif., 25 Feb. 1971, G.
McDonald, 1. M72-1-16-5, Monterey Break-
water, Monterey Co., Calif., 16 Jan. 1972, R.
Hilaski, 1. M73-6-5-11, Asilomar, Monterey
Co., Calif., 5 June 1973, G. McDonald, 1.
M75-11-2-13, Carmel Pt., Monterey Co., Cal-
if., 2 Nov. 1975, G. McDonald, 2. M75-11-4-
3, Fanshell Beach, Monterey Co., Calif., 4
Nov. 1975, G. McDonald, 3. M75-10-5-4,
Carmel Pt., Monterey Co., Calif., 5 Oct. 1975,
С. McDonald, 1. M0175, Carmel Pt., Monte-
rey Co., Calif., 15 Oct. 1970, J. W. Nybak-
ken, 1.
Tritonia diomedea
M71-4-10-1, Monterey Bay, Calif., 10 Apr.
1971, G. McDonald, 1. M71-4-10-2, Monte-
rey Bay, Calif., 10 Apr. 1971, G. McDonald,
1. M71-4-10-3, Monterey Bay, Calif., 10 Apr.
1971, G. McDonald, 1. M71-8-32-1, Santa
Monica Bay, Los Angeles Co., Calif., Aug.
1971, P. Brophy, 1. M71-10-7-2, Marina del
Rey, Los Angeles Co., Calif., 7 Oct. 1971, В.
S. Anderson, 1. M72-1-7-2, Santa Monica
Bay, Los Angeles Co., Calif., 7 Jan. 1972, R.
S. Anderson, 1. M74-3-0-3, Monterey Bay,
Calif., Mar. 1974, D. Rold, 1. M76-1-16-1,
Monterey Bay, Calif., 16 Jan. 1976, B. Alford,
1. M0223, Monterey Bay, Calif., 5 Nov. 1970,
L. Talent, 1. M0471, Monterey Bay, Calif., 13
Mar. 1974, G. McDonald, 1. M0494, Monte-
rey Bay, Calif., 14 Aug. 1974, G. McDonald,
1.
Tritonia festiva
M69-10-26-3, Pirate's Cove, San Luis
Obispo Co., Calif., 26 Oct. 1969, G. Mc-
Donald, 8. M69-10-26-12, Pirate's Cove, San
Luis Obispo Co., Calif., 26 Oct. 1969, G.
McDonald, 1. M70-12-12-3, Pigeon Pt., San
Mateo Co., Calif, 12 Dec. 1970, G. Mc-
Donald, 1. M71-11-4-6, Carmel Pt., Monterey
Co., Calif., 4 Nov. 1971, G. McDonald, 1.
M73-6-2-1, Pescadero Pt., Monterey Co.,
Calif., 2 June 1973, G. McDonald, 1. M75-
11-2-5, Carmel Pt., Monterey Co., Calif., 2
Nov. 1975, G. McDonald, 1. M75-11-3-2, Pi-
geon Pt., San Mateo Co., Calif., 3 Nov. 1975,
G. McDonald, 7. M75-12-2-1, Pigeon Pt., San
Mateo Co., Calif., 2 Dec. 1975, А. К. Mc-
Donald, 4. MO190, Pt. Pinos, Monterey Co.,
Calif., 26 Apr. 1967, V. Mansfield, 1. MO200,
Pigeon Pt., San Mateo Co., Calif., 12 Dec.
1970, G. McDonald, 1. MO208, Asilomar,
Monterey Co., Calif., 27 Apr. 1971, J. W. Ny-
bakken, 1. MO216, Pirate's Cove, San Luis
Obispo Co., Calif., 26 Oct. 1969, G. Mc-
Donald, 1. M0364, Carmel Pt., Monterey Co.,
Calif., 4 May 1973, G. McDonald, 1.
MALACOLOGIA, 1983, 24(1-2): 277-288
FACTORS REGULATING THE DISTRIBUTION OF FRESH-WATER
SNAILS (GASTROPODA) IN NORWAY"
Jan Okland
Section of Limnology, Department of Marine Biology and Limnology,
University of Oslo, P.O. Box 1027, Blindern, Oslo 3, Norway
ABSTRACT
Environmental factors and snail faunas were investigated in about 1,000 lakes. Abundance
of species was estimated according to a time-catch method.
Single factor analyses showed that 18 species tolerated total hardness values down to 1°dH,
and 13 species down to 0.5°dH. The number of species tended to decrease with a decrease
in pH. The drop in species number was particularly noticeable at about pH 6.0 and no species
occurred in lakes with a pH below 5.2. Total hardness, pH, macro-vegetation in the water, and
substratum showed a high correlation with the snail fauna and were considered primary factors
of direct importance to the snails.
Bivariate analyses showed that both total hardness and pH affected the snail fauna per se.
Lakes without gastropods tended to be more acid than those where snails were found, re-
gardless of the level of total hardness. Furthermore, it appears that the snails' tolerance to low
pH increases with increasing total hardness.
Stepwise multiple regression analyses demonstrated that total hardness and water vegeta-
tion were the two most significant independent variables that accounted for the number of
species found and total time-catch abundance. Total hardness, water vegetation and pH were
listed as steps 1, 2, 3, respectively, in the analyses for low total hardness lakes (<1°dH).
The data suggest that acidification of lakes will adversely affect snail species. This implies
that snails can be used as “early warning” organisms of acidification. Since snails are impor-
tant fish-food items, their disappearance will cause a reduction in fish production.
Key words: acidification; distribution; ecology; environment; fresh-water snails; hydrogen-
ion concentration; Norway; pH.
INTRODUCTION
Field investigations on fresh-water snails
have recently been focused upon because
they are sensitive to low pH and consequent-
ly are affected by acid precipitation. Field-
work for the present study was begun in
1953, long before problems connected with
acid rain became evident. Inspiration for this
investigation came from studies of the habi-
tats of fresh-water Mollusca in Britain by
Boycott (1936) and those on the distribution
patterns of fresh-water snails in South Swe-
den by Hubendick (1947). The main purpose
of this study was to elucidate the importance
of various environmental factors for the geo-
graphical and microgeographical distribution
of fresh-water snails in Norway.
Some of the results from this investigation
have already been published (J. Vkland, 1969,
1979a, b; J. Okland 8 К. A. O@kland, 1979).
Most of the material collected is still being
studied and these results will be published
later. The present contribution considers
mainly the snail fauna as a unit. It also in-
cludes much of the background material used
in two short synopses dealing with the fresh-
water fauna and acidification problems (J.
Okland, 1980b; J. Vkland & К. A. Gkland,
1980). The possible effects of acidification are
discussed.
MATERIALS AND METHODS
About 1,500 lakes, rivers, ponds, etc. were
investigated from 1953 to 1973. The distri-
bution of these localities is shown in Fig. 1.
Only lakes, of which there were about 1,000,
will be considered here. Two thirds of the lo-
' Only slightly updated since presentation at the Seventh International Malacological Congress, 31 August-7 September,
1980.
(277)
278 OKLAND
LOCALITIES AM
INVESTIGATED
FIG. 1. Map of Norway indicating the distribution
of approximately 1,500 fresh-water localities where
environmental parameters were recorded and bot-
tom fauna investigated. Area S comprises south-
eastern Norway.
calities (including approx. 600 lakes) are lo-
cated in the southeastern part of Norway.
Within this fairly restricted area environmen-
tal conditions are extremely diverse.
Each locality was usually studied once,
during the summer (1 June-30 September).
In each lake only one habitat was investigat-
ed and defined as a stretch of shore about
200 m long. This habitat was described by
ten environmental parameters and surveyed
down to a depth of 1.5 m. The major collect-
ing device among vegetation and on soft bot-
tom was a sieve mounted on a rod about 1.8
m in length. Stones, branches and other items
were also picked up by hand and inspected
for snails.
The abundance of the snails was estimat-
ed according to a time-catch method, i.e.
number of individuals collected per half-hour.
The level of chemical environmental factors
refers to the summer values from surface
water.
RESULTS
Single Factor Analyses
Of the 27 species of fresh-water snails in
Norway, 18 species tolerated total hardness
(“calcium” content) down to 1°dH, which
equals 10 mg ‘‘CaO’’/I, and 13 species tol-
erated values down to 0.5°dH (Fig. 2). Toler-
ance is defined as the presence of a species
in a lake with a recorded value for a particu-
lar environmental parameter, or presence in
lakes with both the lower and higher values
for that parameter. From Fig. 2 it appears
that the snail fauna has no problems existing
in a lake so long as the total hardness is
above approximately 1°dH.
Fig. 3 indicates the number of species of
snails which tolerate a given pH. Twenty
species were found at pH 7.0. There was a
decline in the number of species in lakes with
a pH below 7.0 and a pronounced decrease
in the number at about pH 6.0 and below.
Fig. 4 shows the correlation between en-
vironmental parameters and number of
species of snails present. For those param-
eters exhibiting discontinuous variation ab-
breviations have sometimes been used to de-
scribe the category (see the caption). Details
on the discrimination between categories will
be given later in connection with a further
treatment of these data. The data in Fig. 4 are
self-explanatory. Therefore, only a few com-
FRESH-WATER SNAILS IN NORWAY 279
NUMBER OF SPECIES
0 à
0 05 10. MS 29 OS
TOTAL HARDNESS (°dH)
FIG. 2. Species tolerance to total hardness, range
0-2.5°dH (1°ан = 10 mg ‘‘CaO’’/I) of snails. Ma-
terial: 959 lakes investigated.
ments will be made. With regard to geology,
lakes which belong to category A have from
two to twelve species with an average of
about six species, while those that belong to
category D have a maximum of four species.
Under the parameters with continuous vari-
ation it was found that at different elevations
above sea level lowland lakes could have few
or many species, while high-altitude lakes al-
ways had few.
The correlation between environmental pa-
rameters and total time-catch abundance of
snails is shown in Fig. 5. The bar graphs rep-
resent the number of individuals collected
during a half-hour period. Among those lakes
which are influenced by unaltered Cambro-
Silurian rocks (IIA), there was а fairly large
number of the lakes investigated in which
more than 150 individuals were collected.
However, in those lakes influenced by
strongly altered Cambro-Silurian rocks (C), or
by Precambrian rock (D), the number of lakes
with such a high population density was neg-
ligible.
Pearson correlation matrices were made for
ten environmental parameters and number of
species of snails (Table 1) and total time-catch
abundance of snails (Table 2). In order to in-
clude parameters with discontinuous varia-
tion (Such as geology) into the calculations,
categories of such parameters were given
numbers from one and upwards, ranking the
categories from favourable to unfavourable
for the snails using the frequency deviation
method described by J. Gkland (1969, 1979a).
T el
20+ / \20
| | | J
if
m | | oo. 4
L | a
L a]
15 e 415
un [ | A
Ww | ! / 4
ie NE и |
un |
we | | 1
S10 410
a | ; Al
WwW | |
(ea) 1 |
> Г | 1
=) & | | >|
Z | |
L |
5H eo | 45
E el | 4
P BER: 4
Ir ! =]
F =
ил nement 0
4.0 5.0 6.0 70
HYDROGEN - ION CONCENTRATION (рн)
FIG. 3. Species tolerance to pH range 4.0 to 7.0
of snails from the same lakes as in Fig. 2.
Correlation coefficients for such parameters
versus snail parameters were therefore all
negative.
The bottom row of each of the two parts
in Tables 1-2 shows how the snail parame-
ters are correlated with the various environ-
mental parameters. The correlation coeffi-
cients describing correlation between
environmental factors with continuous varia-
tion (pH, total hardness (=log*dH), elevation,
and temperature give some idea of the types
of lakes which were studied.
Table 3 compares Pearson and Kendall
correlation coefficients. Since the material is
large (594 lakes) significant values are often
obtained. Therefore major attention should be
focused on the magnitude of the coefficients
and not on significance/non-significance. To-
tal hardness, macro-vegetation in the water,
pH, and substratum have high correlation
coefficients and are primary factors. High
correlations are also found between the snail
fauna and two secondary factors, ¡.e. geolo-
gy and terrestrial vegetation.
Bivariate Analyses
Cursory examination of Fig. 6 which rep-
resents low-calcium lakes in southeastern
280
OKLAND
ENVIRONMENTAL PARAMETER
II. GEOLOGY
r= -0.66
А. UNALTERED C-S ROCKS.
B. MARINE DEPOSITS
С. STRONGLY ALT. C-S.R
D. PRE-EOCAMB. R., ЕТС
00
4
%e
|
|
Ш. TERREST.
VEGETATION
r= -0.48
100
A. CULT. FIELDS, PAST. L
В. CULT.+ CONIF. FOREST
C. CONIFEROUS FOREST
D
. SUBALP.+ALPINE AREAS à
[У MACROVEG,
IN THE WATER
r= - 061
RICH (QUANT.+ QUAL) '
RICH (QUANT.)
SPHAGNUM lo
POOR VEGETATION
4
“lo
00
4
GYTTJA, CLAY
DY - GYTTJA
DY
STONES, SAND
10
BOTH SMALL AND MED ,
SMALL WAVE ACTION o
MEDIUM WAVE ACTION ””
HEAVY WAVE ACTION
VI. WAVE
EXPOSURE
r=-030|
CLEAR, COLOURLESS |
SLIGHTLY BR.-YELL. y,
STRONGLY BR.-YELL
TURBID WATER
. 143 tipaw te
Se | ted
Tr,
A AT в ‹
A IA
п wma ::
Sram .
XII.
TOTAL
TOTAL TIME-| SEEN
HARD -
NESS
r= 074
CATCH
ABUND
r= 0.78
|
150 +
LAKES % | 4 |
INVESTI- | ais
GATED 0
FIG. 4. Correlations of ten environmental parameters and number of species of snails found in 594 lakes
in southeastern Norway. The upper part of the figure represents six parameters with non-continuous
variation (geology, etc.) and the lower part four parameters with continuous variation (elevation, etc.). The
frequency distribution of investigated lakes according to number of species present is presented in the
lower left and the total time-catch abundance correlated with number of species is found in the bottom
right corner. The Pearson correlation coefficient (r) is indicated for each of the 11 sets of correlations.
Abbreviations for categories representing parameters with non-continuous variation: For: geology: A. Un-
altered Cambro-Silurian rocks, C. Strongly altered Cambro-Silurian rocks, D. Precambrian rocks, and
Permian plutonic and effusive rocks of the Oslo region. For terrestrial vegetation: A. Cultivated fields,
pasture lands, B. Both cultivated fields and coniferous forests. For water colour: B. Slightly brownish-
yellowish water, C. Strongly brownish-yellowish water.
FRESH-WATER SNAILS IN NORWAY 281
= TOTAL Е: SE ABUND.
150 310,0 150 310,0 150 310,0 150 310
ENVIRONMENTAL PARAMETER
10
an
A UNALTERED CAMB.-SIL. ROCKS
Il GEOLOGY B. MARINE DEPOSITS à
г--053 | © STRONGLY ALT CAMB.-SIL.R. "
D PRE-EOCAMB. ROCKS, ETC
CULTIVATED FIELDS, PAST. L. 'Y
CULT+CONIF. FOREST
CONIFEROUS FOREST
SUBALPINE + ALPINE AREAS
Ш. TERRESTRIAL
VEGETATION
г= - 040
onu»
A.RICH (QUANT.+ QUAL.) 4
B. RICH (QUANT.)
| С. SPHAGNUM
О. POOR VEGETATION
IV. MACROVEG.
IN THE WATER
r= - 053
| A.GYTTJA, CLAY
V. SUBSTRATUM | B.DY-GYTTJA
С
D
ET DY
f= 052 . STONES, SAND
.BOTH SMALL AND MEDIUM
VI. WAVE À
| B. SMALL WAVE ACTION
EXPOSURE | С. MEDIUM WAVE ACTION
D
Г= - 0.26 | 5 HEAVY WAVE ACTION
A.CLEAR, COLOURLESS WATER
B. SLIGHTLY BROWNISH-YELL.
C. STRONGLY BROWNISH -YELL.
D. TURBID WATER
NUMBER
N = 594
OF
ES lo SPECIES
INVE [=
САТЕО 0 r= 0.78
FIG. 5. Correlations of ten lake parameters with total time-catch abundance of snails (number of individ-
uals collected per half-hour). The parameters and calculations are the same as those described in Fig. 4.
Norway shows that most of the lakes in which Furthermore, lakes in the pH range 6.0-7.0
snails were found were in the pH range 6.0 without gastropods tended to have a lower
to 7.0 and had a total hardness above 0.25” total hardness than those in which snails were
dH. Lakes without gastropods tended to be present.
more acidic than those in which snails were Study of data from all parts of Norway re-
present, regardless of the level of total hard- vealed that nine of the eleven most common-
ness. ly found species increased their tolerance to
QT
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284 OKLAND
TABLE 3. Pearson and Kendall correlation coefficients from one-factor analyses of environmental param-
eters correlated with snail parameters (number of species and time-catch abundance). Data from 594 lakes
in southeastern Norway. Kendall values are put in parentheses. Pearson correlations are significant + 0
at 0.001 level if |r| > 0.126 (symbol @), at 0.05 level if |r| > 0.067 (assuming normal distributions). Kendall
correlations significant + O at 0.001 level if |t| > 0.11 (symbol 6), at 0.05 level if |t| > 0.05. All tests are
one-tailed.
A A a A A AA AAA AAA EE EEE A A
Total time-catch
Number of species abundance
Pearson Kendall Pearson Kendall
Environmental parameter r t r t
High Correlations
A. PRIMARY FACTORS
(IN THE WATER)
Total hardness 0.740 (0.560) 0.610 (0.490)
Macro-vegetation -0.610 (0.490) 0.530 (0.450)
pH 0.580 (0.470) 0.450 (0.410)
Substratum -0.52@ (—0.40@) -0.42@ (—0.34@)
B. SECONDARY FACTORS
(OUTSIDE THE WATER)
Geology -0.66@ (— 0.51) -0.53@ (—0.43@)
Terrestrial vegetation -0.48@ (—0.39@) -0.40@ (—0.33@)
Low Correlations
Wave exposure -0.30@ (—0.22@) —0.26@ (—0.23@)
Elevation -0.28@ (-0.21@) —0.24@ (—0.16@)
No or negligible
Correlations
Water temperature 0.11 (0.05) 0.11 (0.02)
Water colour DAA (051) — 0.03 (-0.06)
low pH as the total hardness level increased. DISCUSSION
This pattern, as well as tendencies described
in connection with Fig. 6, is shown in Fig. 7.
Multiple Regression Analyses
Multiple regression analyses may be used
to show the relative importance of environ-
mental factors to explain variability in the
data. Using stepwise multiple regression
analyses, where the number of species and
total time-catch abundance, respectively,
were dependent variables and the ten envi-
ronmental parameters independent vari-
ables, ranking lists of the various environ-
mental factors were obtained. Table 4 gives
results for number of species and Table 5 for
time-catch abundance. An increase in r re-
flects improvement of the model by the ad-
dition of more parameters. Total hardness
and macro-vegetation in the water had the
greatest explicatory value both for “all lakes”
and “low total hardness lakes’ in both ta-
bles. pH appears in Step 3 in the low calcium
groups, but is insignificant for “all lakes.”
Calcium concentration and pH level are im-
portant factors for determining presence and
well-being of snails (Boycott, 1936; Huben-
dick, 1947; Aho, 1966, 1978; J. ОКапа, 1969,
1979a, b; J. Okland 8 К. A. Okland, 1979).
The decline in species number with decreas-
ing values for total hardness (Fig. 2) and pH
(Fig. 3) is, therefore, in agreement with pre-
vious information.
The concept “species tolerance” was used
in relation to values of total hardness and pH
of lakes. If a given species was collected from
a lake with a particular parameter value, it
was considered to tolerate this value or con-
ditions associated with this value. When the
species was not found in lakes where the pa-
rameter was below a given value, and a suf-
ficiently large number of lakes below this val-
ue were investigated with negative results, we
assumed that this condition was not tolerat-
ed by the species. In those cases where the
species was recorded in only one or a few
lakes having the lowest value of a particular
FRESH-WATER SNAILS IN NORWAY 285
or о
ОЗ
01+ [0] OLAKE WITH
GASTROPODA
OLAKE WITHOUT
Г GASTROPODA
TOTAL HARDNESS (*dH)
0 ER re e
40 50 60 70 80 90
HYDROGEN - ION CONCENTRATION (pH)
FIG.6. Presence/absence of snails in low calcium
lakes of southeastern Norway with a given pH and
total hardness.
parameter, these lakes were considered to
be marginal habitats for the species. The
curves in Figs. 2-3 represent such marginal
habitats.
The bivariate analyses in Fig. 7 allow us to
make some general statements about the
dependence of the snail fauna on pH and cal-
cium concentration. Since this figure is based
on a large amount of data for abundance of
widespread species from all parts of Norway,
it includes data from Fig. 6 which only rep-
resents presence/absence data from a re-
stricted part of the calcium spectrum and lim-
ited to southeastern Norway.
We note that at a given value of calcium
the number of species and abundance in-
crease with increasing pH.
At a given pH value the number of species,
and abundance, increase with increasing cal-
cium content.
It may, therefore, be concluded that both
pH and calcium concentration are important
factors, per se, in spite of the fact that they
are correlated (Table 1).
Marginal habitats for the snails are repre-
sented by dots bordering the white area. It is
evident that these dots form a curved line
which extends further to the left—towards
more acid water—when calcium concentra-
tion is medium or high, and further down-
wards—towards lower calcium concentra-
tion—when pH is medium or high. This
implies that in those habitats where pH is
VERY ACIDIC ACIDIC SLIGHTLY ACIDIC
<
NO
SPECIES
PRESENT
INCREASING Ca
INCREASING pH
INCREASING NUMBER OF SPECIES
ABUNDANCE
FIG. 7. Schematic diagram of lakes with mea-
sured values for pH and calcium concentration
containing widespread species of snails (N = 9),
small mussels (N = 10), and the crustacean Gam-
marus lacustris. Approximately 1,000 lakes inves-
tigated.
minimal for a given species, total hardness is
above minimum—-and in those habitats where
total hardness is minimal, pH is above mini-
mum. The species accordingly does not tol-
erate more than one minimal factor at the
same time. If one factor approaches the tol-
erance limit of the species, other factors have
to be favourable.
The single factor analyses in Table 3 places
those factors showing a high correlation with
the snail fauna into the following groups: Pri-
mary factors, those directly influencing the
snails (total hardness, macro-vegetation in the
water, pH, and substratum), and Secondary
factors, those which affect the species indi-
rectly through other factors for example: ge-
ology influences total hardness. Following a
model for direct/indirect action of environ-
mental factors on the snail fauna (J. Okland,
1979a) the effect of a given factor was con-
sidered a direct one when its action did not
necessarily involve a step through any of the
other factors.
It also appears from Table 3 that wave ex-
posure and elevation above sea level had low
correlations with the snail fauna, while no or
negligible correlations were found for water
temperature and water colour.
Correlations may reflect causality either di-
rectly between the given environmental fac-
286 OKLAND
TABLE 4. Stepwise multiple regression analyses of number of species of snails (dependent variable) and
ten lake parameters (independent variables) from 594 lakes in southeastern Norway (''all lakes’’), and from
low total hardness lakes (N = 403). The parameters are: (1) elevation above sea level, (2) water tempera-
ture, (3) geology, (4) terrestrial vegetation, (5) macro-vegetation in the water, (6) substratum, (7) wave
exposure, (8) water colour, (9) total hardness (log°dH), and (10) hydrogen-ion concentration (pH). Sequence
and selection of environmental parameters are listed according to decreasing prediction value.
Percent of
variability
r accounted for
ALL LAKES Step 1. Log°dH* 0.74 54.7
Step 2. Macro-vegetation 0.79 62.0
Step 3. Geology 0.79 63.0
Step 4. Substratum 0.80 63.7
Step 5. pH 0.80 64.1
Step 6. Terr. vegetation 0.80 64.6
Step 7. Water temp. 0.81 65.0
Step 8. Wave exposure 0.81 65.2
Step 9. Elevation 0.81 65.4
LOW TOTAL HARDNESS Step 1. Log’dH* 0.53 27.9
LAKES (=1°dH) Step 2. Macro-vegetation 0.63 39.1
Step 3. pH 0.66 43.0
Step 4. Geology 0.67 44.7
Step 5. Substratum 0.68 45.7
Step 6. Water temp. 0.68 46.2
Step 7. Elevation 0.69 47.7
Step 8. Wave exposure 0.69 48.1
* Total hardness (‘‘calcium’’).
tor and the snails or indirectly through other
factors which are influenced by the environ-
mental factor in question. Causality can only
be proven, however, by experimentation or
corroborated by knowledge of the ecology
and physiology of the species.
It is of special importance to verify causal-
ity for the primary factors, which had high
TABLE 5.
correlations with the snail fauna. Low values
of total hardness may be critical, since snails
need calcium for their shells. Limnic organ-
isms (fish) are affected by low pH through
changes in the ion balance, creating lower
concentrations of Na and Cl in the blood
plasma with secondary effects at the cellular
level (Leivestad et al., 1976; Leivestad &
Stepwise multiple regression analysis of total time-catch abundance of snails (dependent vari-
able) and ten lake parameters (independent variables) of 594 lakes in southeastern Norway (‘all lakes’’),
and low total hardness lakes (N = 403). Same parameters etc. as indicated in Table 4.
ALL LAKES Step 1. Log’dH*
Step 2. Macro-vegetation
Step 3. Terr. vegetation
Step 4. Water colour
Step 5. Substratum
LOW TOTAL HARDNESS
LAKES (<1°dH)
Step 1. Log"dH*
Step 3. pH
Step 4. Geology
Step 5. Elevation
Step 6. Water temp.
* Total hardness (‘‘calcium’’).
Step 2. Macro-vegetation
Percent of
variability
r accounted for
0.61 37483
0.66 44.1
0.67 44.5
0.67 44.8
0.67 45.1
0.40 157
0.47 21.9
0.51 25.9
0.52 27.2
0.53 27.8
0.54 29.3
FRESH-WATER SNAILS IN NORWAY 287
Muniz, 1976). Much work on the physiology
of the snails seems to be needed in order to
understand how low calcium concentration
and low pH affect the snail fauna. Dussart &
Kay (1980) point out that the precise way in
which fresh-water gastropods respond to
their chemical environment at a cellular level
has yet to be discovered.
The importance of the macro-vegetation is
probably connected to different ecological
mechanisms. A rich growth of macro-vege-
tation creates sheltered conditions and re-
duces wave action. It also provides organic
matter to feed on, including attached algae,
and suitable substratum to crawl upon. Dif-
ferent types of macro-vegetation reflect dif-
ferent trophic states which in indirect ways
(through chemical factors, etc.) may affect the
snail fauna.
Substratum also had a high correlation with
the snail fauna. Gyttja and dy-gyttja repre-
sent more productive lakes. Algae growing
on these types of sediment may serve as food
for several species, a few also using the or-
ganic material in the substratum per se. A
loose dy sediment is obviously a disadvan-
tage for all species which to some extent are
in need of a substratum to crawl upon.
From the stepwise regression analyses
(Tables 4 and 5), it appears that regarding
“all lakes’’ most of the variability in the data
can be explained by two environmental fac-
tors, total hardness and macro-vegetation in
the water. If only low-calcium lakes are con-
sidered (bottom part of the tables), these
same two factors top the list, but pH should
not be disregarded. These three factors to-
gether account for much of the variation in
the data. The increase in importance of pH in
the groups of low total hardness lakes is rea-
sonable, since the effect of pH is connected
with acid water which almost always has a
hardness value below 1°dH.
The sequence of the first three environ-
mental factors in the low total hardness lakes
in Tables 4-5 (total hardness, macro-vege-
tation, PH) suggests that a change in pH will
affect the snail populations. In a situation of
acidification, a slight increase in total hard-
ness has been observed in some lakes. If
macrovegetation should change, it would be
towards adverse conditions for the snails. A
slight increase in total hardness is, however,
not sufficient to maintain snail populations
unaffected during acidification since the num-
ber of species tolerating given values for pH
decreases drastically with decreasing pH (Fig.
3). Since we have seen that pH affects the
snail fauna per se and not through total hard-
ness with which it is correlated we conclude
that decreasing pH may cause snails to dis-
appear.
Thousands of lakes both in Europe and in
North America have been acidified during the
last decades mainly by the impact of acid
precipitation (Drablos & Tollan, 1980). In Nor-
way more than one thousand lakes have lost
their fish populations (Sevaldrud et al., 1980).
Only low-calcium lakes become acidified,
particularly those with total hardness below
0.5°dH, but sometimes lakes with values up
to 1°dH (Henriksen, 1979; J. Okland, 1980a;
J. Vkland 8 К. A. Okland, 1980). Since 13 of
the Norwegian species of fresh-water snails
tolerate hardness values down to 0.5°dH, at
least one-half of the Norwegian species may
be influenced, if acidification continues.
Snails form an important part of the diet of
fresh-water fish. They are more sensitive to
low pH than fish, which means that they will
disappear before the fish become extinct. It
has been suggested that the disappearance
of snails will reduce fish production (J. @k-
land 8 К. A. Okland, 1980). Snails may also
be used as biological indicators or “early
warning’ organisms for monitoring acidifica-
tion (J. Vkland, 1980b).
ACKNOWLEDGEMENTS
The work has been sponsored by the Nor-
wegian Research Council for Science and the
Humanities (1953-1976), the Directorate for
Wildlife and Freshwater Fish (1965-1980),
and the Norwegian interdisciplinary research
project, “Acid Precipitation—Effects on For-
est and Fish” (the SNSF-project) (1977-
1979). Cand. real. Vidar Berteig at the Nor-
wegian Computing Center has assisted in
calculations and given statistical advice.
Cand. real. Karen Anna Okland at the Section
of Limnology, University of Oslo, has partici-
pated in fieldwork and laboratory activities.
We also thank Dr. May Haugstad for her as-
sistance in the preparation of the final ver-
sion of this manuscript. This is SNSF-contri-
bution FA 60/80.
REFERENCES CITED
AHO, J., 1966, Ecological basis of the distribution
of the littoral freshwater molluscs in the vicinity
288 OKLAND
of Tampere, South Finland. Annales Zoologici
Fennici, 3: 287-322.
AHO, J., 1978, Freshwater snail populations and
the equilibrium theory of island biogeography. И.
Relative importance of chemical and spatial vari-
ables. Annales Zoologici Fennici, 15: 155-164.
BOYCOTT, A. E., 1936, The habitats of fresh-water
Mollusca in Britain. Journal of Animal Ecology,
5: 116-186.
DRABLOS, D. & TOLLAN, A., eds., 1980, Ecolog-
ical impact of acid precipitation. Proceedings of
an international conference, Sandefjord, Nor-
way, March 11-14, 1980. SNSF project, Nor-
way, 383 p.
DUSSART, G. 8 KAY, R., 1980, Relationships be-
tween water chemistry and respiration rate in
several populations of Lymnaea peregra Müller
(Gastropoda: Mollusca). Hydrobiologia, 69: 57-
65.
HENRIKSEN, A., 1979, A simple approach for
identifying and measuring acidification of fresh-
water. Nature, 278: 542-545.
HUBENDICK, B., 1947, Die Verbreitungsverhált-
nisse der limnischen Gastropoden in Süd-
schweden. Zoologiska Bidrag fran Uppsala, 24:
419-559.
LEIVESTAD, H., HENDREY, G., MUNIZ, I. P. &
SNEKVIK, E., 1976, Effects of acid precipitation
on freshwater organisms. In: BRÆKKE, Е. H.,
ed., Impact of acid precipitation on forest and
freshwater ecosystems in Norway. SNSF proj-
ect FR6/76, Oslo-Ás, Norway, p. 87-111.
LEIVESTAD, H. & MUNIZ, 1. P., 1976, Fish kill at
low pH in a Norwegian river. Nature, 259: 391-
392.
OKLAND, J., 1969, Distribution and ecology of the
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@KLAND, J., 1979a, Distribution of environmental
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Norway: use of European Invertebrate Survey
principles. Malacologia, 18: 211-222.
OKLAND, J., 1979b, Kalkinnhold, surhetsgrad (pH)
og snegler i norske innsjoer. Abstract in English:
Total hardness, hydrogen-ion concentration (PH)
and Gastropoda in Norwegian lakes, with re-
marks on acidification of watercourses. Fauna
(Oslo), 32: 96-111.
OKLAND, J., 1980a, Acidification in 50 Norwegian
lakes. Nordic Hydrology, 11: 25-32.
@KLAND, J., 19806, Environment and snails (Gas-
tropoda): studies of 1,000 lakes in Norway. In
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impact of acid precipitation. Proceedings of an
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March 11-14, 1980. SNSF project, Oslo-As,
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Ás, Norway, p. 326-327.
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POSTSCRIPT
TOTAL HARDNESS, CALCIUM AND MAGNESIUM
Values for total hardness represent calcium plus magnesium. For 514 lakes, calcium and
magnesium were also measured separately. Based on this material, average concentration of
calcium and magnesium in Norwegian lakes can be calculated for given values of total hard-
ness. Some examples are given below.
Given values for total hardness
°dH “Cao:
(1°dH = 10 mg ''СаО’/) mg/l
0.2 2
0.4 4
0.6 6
0.8 8
1.0 10
5.0 50
10.0 100
20.0 200
Calculated empirically
Ca
mg/l mg/l
lee 0.2
2.2 0.4
32 0.7
4.2 0.9
5.2 122
30.8 3.0
62.4 5.6
MALACOLOGIA, 1983, 24(1-2): 289-297
THE PREDATORY BEHAVIOUR OF MARISA CORNUARIETIS ON
EGGS AND NEONATES OF BIOMPHALARIA GLABRATA,
THE SNAIL HOST OF SCHISTOSOMA MANSONI
Aragua Cedeno-Leön? & J. D. Thomas
School of Biological Sciences, University of Sussex, Falmer,
Brighton BN1 9QG, Sussex, England
ABSTRACT
The predatory behaviour of Marisa cornuarietis (L.) was evaluated as an agent for the control
of Biomphalaria glabrata (Say). The results show that predation by Marisa is influenced both
by endogenous and exogenous factors. Although some Marisa preyed deliberately on Biom-
phalaria eggs and juveniles, a minority did not do so during the experiment. There is evidence
that Marisa predation tends to increase with experience, age and onset of sexual maturity.
Mature females ingest significantly more eggs than mature males. Predation is also enhanced
by the presence of plant food in flow systems but depressed by heterotypic conditioning. The
possible cause of these effects is discussed. The above considerations, and the fact that the
rate of predation by Marisa does not increase with egg density, shows that Marisa is not a
good predator. The conclusion that Marisa can only be effective as a predator if it is present
at high densities, of about 1 per 150-300 cm?, is supported by field observations.
INTRODUCTION
The feeding behaviour of freshwater snails
that serve as hosts of human schistosomes,
and those proposed as agents for their con-
trol, has received little detailed attention (Fer-
guson, 1977; Brown, 1980). Whilst it is gen-
erally assumed that freshwater pulmonate
and prosobranch snails feed mainly on living
or dead plant material as well as small ani-
mals associated with epilithic and epiphytic
algae (World Health Organization, 1965), re-
cent detailed studies (Ndifon, 1979; Ndifon 8
Ukoli, 1980) have shown that animal materi-
al, including protozoa, gastrotrichs, cope-
pods, ostracods, rotifers, nematodes and
even chironomid larvae, may become domi-
nant in the diets of the pulmonate Bulinus
globosus (Morelet), at certain times at least.
There is also evidence that snail species, such
as Marisa cornuarietis (L.), Helisoma duryi
(Wetherby), Physa acuta (Drap.) and Poma-
cea spp., are predators, as eggs and juve-
niles of pulmonate snails feature in their diets
(Oliver-Gonzalez et al., 1956; Frandsen 4
Madsen, 1979; Paulinyi 8 Paulini, 1972; Fer-
guson, 1977). Before their potential as con-
trol agents can be properly evaluated, it is
important that their predatory behaviour be
better understood.
The present paper concerns the predatory
behaviour of Marisa cornuarietis and at-
tempts to answer the following questions.
Firstly, does Marisa deliberately feed on the
eggs and juveniles of the snail hosts? Ac-
cording to Oliver-Gonzalez et al. (1956) and
Chernin et al. (1956) the ingestion of egg
masses is incidental while Marisa is feeding
on plant material on which they have been
deposited. Demian 4 Lutfy (1965, 1966) take
the opposite view and claim that predation
by Marisa is intentional, although no quanti-
tative evidence is provided. Secondly, is Ma-
risa an obligate omnivore, or does it cease to
feed on eggs when plant food is present in
abundance? Thirdly, to what extent is pre-
dation by Marisa dependent on media con-
ditioning, previous experience, age and sex?
Fourthly, does Marisa show any of the attri-
butes of an efficient predator?
' Based on part of a dissertation submitted for a Ph.D. degree from the University of Sussex by A. Cedeño-León (1975).
2 Present address: Universidad Central de Venezuela, Facultad de Ciencias, Instituto de Zoologia Tropical, Apartado
47058, Caracas 1041-A, Venezuela.
(289)
290 CEDENO-LEON & THOMAS
MATERIALS AND METHODS
The methods used for culturing the albino,
Venezuelan strain of Biomphalaria glabrata
(Say), and the Puerto Rican strain of Marisa
cornuarietis, have been described by Thom-
as (1973) and Cedeño-León (1975). Both
species were maintained in 40 aquaria con-
taining aerated, filtered tap-water maintained
at a temperature of 26 + 1°C and a photo-
period of 12 hr light and 12 hr dark. They
were fed lettuce daily. As Marisa tends to
pollute its environment, the tap-water was
replaced by means of a ‘drip-feed’ (8 mi min ')
and an overflow.
The experiments described below were de-
signed to answer the questions raised above.
1. The influence of experience, and location
of egg masses, on predation by Marisa
In the first treatment six juvenile Marisa
12 + 2 mm in shell diameter, which had pre-
viously coexisted with Biomphalaria for three
months, were placed, together with an equal
number of adult Biomphalaria (14 + 1 mm),
in plastic buckets containing 1.2 | of Stan-
dard Snail Water (SSW2) (Thomas et al.,
1975). The second treatment resembled the
first in all respects, except that the Marisa
had not previously encountered eggs, juve-
niles or adults of Biomphalaria. The snails in
the first and second treatments were desig-
nated experienced and non-experienced
predators respectively. As a control, six adult
Biomphalaria in the same size range as those
used in the treatments were placed in 600 ml
of SSW2 in a plastic bucket. Each treatment
and control were replicated six times.
The SSW2 was changed every three days.
An excess of lettuce was provided as food.
Uneaten remains were removed each day be-
fore adding new portions. The number of egg
masses laid on different kinds of substrates,
the total number of egg masses and the
number of juvenile Biomphalaria present were
counted at three day intervals for a period of
15 days.
2. The influence of egg deposition substrate
and media conditioning on predation by Ma-
risa
If egg masses are only consumed acciden-
tally during feeding, then obviously they
should be safe from attack if laid on non-food
substrates. To test this, 14 plastic buckets,
each containing 10 adult Biomphalaria fed on
an excess of lettuce, were left for 4 days in
an environmental unit at a temperature of
26 + 1°C. During this period they deposited
between 12 and 20 egg masses in each of
the containers. The snails were then re-
moved, and the number of egg masses in
each container recorded. The water was
carefully decanted and replaced by 600 ml of
fresh SSW2. Six non-experienced juvenile
Marisa (14-17 mm shell diameter) were in-
troduced into each of ten buckets and lettuce
was added in excess. Four buckets without
snails were used as controls. The media were
changed every third day.
The number of Biomphalaria egg masses
and juveniles present in each container was
counted every day for seven days.
3. Influence of experience and Biomphalaria
egg density on egg predation by juvenile
Marisa
In these experiments the functional re-
sponse (Solomon, 1949; Holling, 1959a, b)
was investigated by measuring the extent of
predation on varying numbers of Biompha-
laria eggs by individually isolated, experi-
enced and non-experienced juvenile Marisa
(shell diameter 17-23 mm).
To familiarise Marisa with Biomphalaria
eggs, two groups of 20 juvenile Marisa were
maintained separately at 26 + 1°C in the aer-
ated flow-through aquaria previously de-
scribed. Approximately 500 Biomphalaria egg
masses, deposited on floating transparent
plastic sheets, were introduced into each
aquarium. Initially, the snails were deprived
of food and remained on the sediment, mak-
ing no attempt to prey on the egg masses.
From the fourth day onwards they were pro-
vided with lettuce below the level deemed to
be in excess. As a result, all the lettuce was
eaten by the end of the day. After this change
in feeding regime, the rate of consumption of
egg masses increased from zero to a stable
level of 1.81 + 0.19 egg masses per Marisa
per day. The snails were maintained under
these conditions for eight weeks.
After this treatment, the experienced Ma-
risa were placed individually, with a 1 cm di-
ameter lettuce disc, in buckets containing ap-
proximately 50, 100, 150, 200 and 250 Biom-
phalaria eggs in 200 ml SSW2. Each
treatment was replicated four times. A par-
allel experiment was set up with non-experi-
enced snails. The experimental snails were
maintained under the standard conditions al-
ready described. The number of eggs remain-
ing uneaten was counted after 24 hr.
MARISA PREYING ON BIOMPHALARIA 291
e EXPERIENCED M.C
o NON-EXPERIENCED M.C 30
= CONTROL B.G
: 25 у
Sp ONM.C SHELL UNATTACHED
£ 0 =
RO SC 8 I E
= DAYS zer
= 5 a
m а
es <
<<] = )
= 10 Fon B.6. SHELL = 10
2 =
(Un,
CO
5 =
= =
= =
=) S 0
[æ) —
Os EA er es
= DAYS = DAYS
© 25 as 625
2 i 2
2 ON LETTUCE = 29 LON CONTAINER WALLS т
Е E
S 15 =
eS ud
2 E
= ==
m =
5 =>
=
5
0
ON EIA E A Soe (deals ea E a TE
DAYS DAYS
FIG. 1. The cumulative mean number (X + S.E.) of egg masses laid by six Biomphalaria on various
substrates in three treatments (control: Biomphalaria on their own, B. glabrata with experienced Marisa,
and B. glabrata with non-experienced Marisa). Key: B. glabrata = B.g; M. cornuarietis = M.c.).
292 CEDENO-LEON & THOMAS
FIG. 2. The mean number (X + S.E.) of Biom-
phalaria egg masses counted each day in the three
treatments described in Fig. 1. (Key as in Fig. 1.)
4. Influence of previous experience, and
variation in Biomphalaria egg density, on egg
predation by adult Marisa
A procedure similar to the one described
for juvenile Marisa was used for adults (34—
40 mm shell diameter), except that the treat-
ments consisted of 100, 200, 300, 400 and
500 Biomphalaria eggs and the snails were
provided with two 1 cm diameter lettuce discs
each.
RESULTS
1. The influence of experience and location
of egg masses on predation by Marisa
It was found that Biomphalaria deposited
egg masses on all possible substrates,
namely lettuce leaves, shells of their conspe-
cifics, Marisa shells and the walls of the con-
tainers. From day nine onwards many egg
masses were found loose on the bottom of
the containers, but as Biomphalaria cannot
oviposit without attaching the egg masses to
a substrate, it can be assumed that they had
become detached after oviposition. As the
egg masses do not attach very firmly to let-
tuce leaves, it is probable that most, if not
all, of the detached egg masses originated
from the lettuce. When the excess lettuce
leaves were removed each day, the portions
of the leaf containing the egg masses were
left in the container.
The cumulative mean number of detached
egg masses and those found on the various
substrates are given separately for the two
treatments and the control in Fig. 1. The data
which were subjected to a three-way analy-
sis of variance revealed the following trends.
DIV LAB RA TA
NON-EXPERIENCED M.C
FIG. 3. The mean number (X + S.E.) of neonate
B. glabrata present in the three treatments de-
scribed in Fig. 1 after day 9 of the experiment. (Key
as in Fig. 1.)
There were statistically significant tenden-
cies for the mean number of eggs deposited
by Biomphalaria on all four substrates to in-
crease progressively until the ninth day.
Thereafter the number of egg masses count-
ed on the Marisa shells and container walls
continued to increase, whereas the mean
numbers on lettuce and Biomphalaria shells
declined. This decline was due, in part, to the
Biomphalaria eggs becoming detached, as
Fig. 1 shows that the mean number of unat-
tached eggs counted on the floor of the con-
tainer increased throughout the period of ob-
servation, in both the control and the two
treatments. Lettuce is clearly the preferred
oviposition site and the substrates may be
ranked in order of preference, as follows:
lettuce > container wall > Biomphalaria
shells > Marisa shells. The number of egg
masses found was always significantly lower
on the treatments containing Marisa than in
the controls (P < 0.001), but there were no
significant differences between the mean
number of egg masses in treatments contain-
ing experienced and non-experienced Ma-
risa.
Fig. 2 shows the mean number of Biom-
phalaria eggs found in the two treatments and
the control. These values are based on the
total number of eggs counted in each repli-
cate at each observation time, irrespective of
the substrate on which they were laid. It can
be seen that there was a significant tendency
(Р < 0.01) for the number of eggs to decline
in the control after the twelfth day. There were
significantly more Biomphalaria egg masses
in the controls than in the treatments con-
taining Marisa (P < 0.001). However, there
MARISA PREYING ON BIOMPHALARIA 293
kl
WN
m
o
(=>)
un
MEAN №, OF EGG MASSES CONSUMED / MARISA / DAY ( e—e )
DAYS
FIG. 4. The mean daily number (X + S.E.) and mean cumulative percentage (X
100 |
T
a
=
D
5
50-7 >
cn
uy
mn
”n
=
с (o)
©
uy
Be
0
5 6 7
+ S.E.) of В. glabrata
egg masses consumed by individual Marisa over a period of seven days.
were no significant differences at any time
between the values in the treatments con-
taining experienced and non-experienced
Marisa.
Fig. 3 shows the mean number of neonate
Biomphalaria found in the control and the two
treatments, from the time they appeared in
the control until the end of the experiment.
Significantly more neonates were found in the
controls than in either of the two treatments
on both days 12 and 15 (P < 0.01). There
were fewer neonates in the treatment con-
taining experienced Marisa than in that con-
taining their non-experienced conspecifics on
days 12 and 15; but the differences were only
statistically significant on day 12 (P < 0.05).
2. The influence of egg deposition, substrate
and media conditioning on predation by Ma-
risa
The mean rate of consumption of egg
masses per day by Marisa were higher on
days 1, 4 and 7 (the days after the media
were changed) than on other days. The val-
ues on day 4 were significantly higher (P <
0.05) than on other days. Fig. 4 also shows
that there was a progressive increase in the
percentage of eggs eaten, and by the end of
the experiment 90.5% + 5.3 had been con-
sumed. As might be expected, there were
significantly more juveniles in the control than
in the treatment containing Marisa (P <
0.001) from the fourth day onwards (Fig. 5).
3. Influence of experience, and Biomphalaria
egg density, on egg predation by juvenile
Marisa
Fig. 6 shows that the number of eggs eat-
en per day does not change significantly with
increase in egg density. Although experi-
enced Marisa ate more Biomphalaria eggs
than their non-experienced conspecifics (with
one exception), the differences were only sta-
tistically significant (P < 0.05) in the 50 egg
treatment. The high standard errors indicate
the high level of individual variation in preda-
tory activity. In fact, several of the Marisa did
not prey on egg masses.
294 CEDENO-LEON & THOMAS
20
>)
e CONTROL BUCKETS т
o BUCKETS WITH MARISA
.GLABRATA / CONTAINER
S
100
B
=
>
250
= 0
= CRE (aE A Ze ee e a |
0 ] 2 3 4 5 6 7
DAYS
FIG. 5. The mean number (X + S.E.) of juvenile
B. glabrata per container, counted each day over
a seven day period.
4. Influence of previous experience and vari-
ation in Biomphalaria egg density on egg pre-
dation by adult Marisa
Fig. 7 shows that the number of eggs eat-
en per day by adult Marisa did not vary sig-
nificantly with egg density. Neither could any
significant differences be detected between
the number of egg masses consumed by ex-
perienced and non-experienced Marisa, al-
though, as with juveniles, there was a ten-
dency (with one exception) for experienced
EI EXPERIENCED SWAIL
O NON-EXPERIENCED SNAILS
АК!
EGGS CONSUMED PER
OF
MEAN NO.
FIG. 6. The mean number (X + S.E.) of egg
masses consumed per juvenile Marisa per day in
the various egg density treatments.
EXPERTENCED SNATLS
СЗ NON-EXPERIENCED SNAILS
/ MARISA / DAY
OF EGGS CONSUMED
MEAN NO,
100 200 300 400 500
№, OF EGGS
FIG. 7. The mean number (X + S.E.) of egg
masses consumed per adult Marisa per day in the
various egg density treatments.
snails to eat more egg masses than their non-
experienced conspecifics. The standard errors
indicate that, as with juveniles, there was a
great deal of individual variability in predatory
activity.
Table 1 shows that mature female Marisa
ate more Biomphalaria eggs than mature
males. A two-way analysis of variance indi-
cates that the differences between the sexes
were highly significant (P < 0.001). However,
there were no significant differences attrib-
utable to experience of egg eating, or to in-
teractions between the sex differences and
experience.
DISCUSSION
The results of the first 15 day experiment
show that significantly fewer Biomphalaria
eggs were invariably found in the treatment
containing Marisa than in controls. These dif-
ferences may have been caused directly by
Marisa preying on the eggs, or indirectly by
Marisa causing a decline in egg production
by Biomphalaria as a result of competition.
Although Cedeño-León (1975) showed that
conditioning of the media and competition for
resources by Marisa does impair the growth
and reproduction of Biomphalaria, the results
of experiments 2-4 in the present investiga-
tion show unequivocally that Marisa preys
significantly on both eggs and juveniles of
Biomphalaria. These results confirm those of
Oliver-Gonzalez et al. (1956), Chernin et al.
(1956), Demian & Lutfy (1965, 1966), Huben-
MARISA PREYING ON BIOMPHALARIA 295
TABLE 1.
Response of female and male Marisa cornuarietis to an increasing density of Biomphalaria
glabrata eggs, measured as number of eggs consumed for M. cornuarietis per day (X + S.E.).
Experienced snails
Non-experienced snails
Female Male
40.91 + 15.35 7.88 + 6.56
39 A0EE NAN д.14 =: 2182
dick (1966), Msangi & Kihaule (1972) and
strengthen the suspicion of Jobin (1970) that
the decline in Biomphalaria populations in
nature, following the introduction of Marisa,
was largely due to predation.
Biomphalaria has a catholic taste for ovi-
position sites (Ferguson, 1977). In this inves-
tigation these snails deposited their egg
masses on the following substrates in order
of preference: the plant food (lettuce) > walls
of the container > Biomphalaria shells >
Marisa shells. Although predation pressure is
likely to be greater on the egg masses de-
posited on the lettuce, the results show that
Marisa will also prey on egg masses depos-
ited on non-food substrates. This observa-
tion supports the contention of Demian &
Lutfy (1965, 1966) that predation by Marisa
is intentional, not accidental, as claimed by
Chernin et al. (1956) and Oliver-Gonzalez et
al. (1956).
However, the present results demonstrate
that the predatory behaviour of Marisa is an
extremely plastic phenomenon that may be
influenced by environmental factors, as well
as by genotype, age and the physiological
state of the snail. The experiments involving
plant food shed some light on the predatory
activity of Marisa. The first experiment shows
that the presence of plant food does not de-
ter Marisa from preying on egg masses or
juveniles of Biomphalaria. On the contrary,
the experiments in flow aquaria demonstrate
that Marisa will only prey on egg masses
when plant food is present. The increased
predatory activity in the presence of plant
food may be caused by kairomones, diffusing
from the lettuce, acting as arousal factors.
Alternatively, increased activity following the
ingestion of lettuce could be the cause. Fur-
ther experiments are needed to test these
hypotheses. The behaviour of Marisa differs
in this respect from that of Pomacea, which
continues to feed on Biomphalaria eggs and
juveniles even in the absence of plant food
(Paulinyi & Paulini, 1972).
In contrast to the effects of lettuce, condi-
tioning of the SSW2 by snails feeding on let-
tuce is followed by a decline in the predatory
activity of Marisa. The reason is unknown,
but it is possibly caused by the accumulation
of certain chemical species such as H* or
ammonia (as NH,* or free NH,) or to a de-
crease in the concentration of Ca?*. There is
experimental evidence that such changes do
occur in media conditioned by Biomphalaria
and Marisa (Thomas, Goldsworthy 8 Benja-
min, 1974; Cedeño-León, 1975). These
chemical changes may act by lowering the
metabolic activity of the Marisa, or by creat-
ing background “noise” thus making the de-
tection of egg masses difficult. Demian 4 Lut-
fy (1966) have postulated that both egg
masses and juveniles release attractants, but
there is no experimental evidence for their
existence.
There is also a great deal of variability in
the responses shown by Marisa to Biom-
phalaria eggs. This is reflected in the high
standard errors of the mean values. There is
some evidence that predatory behaviour
might be an individual characteristic, as a mi-
nority of the snails never preyed on egg
masses. Further experiments are needed to
ascertain whether this is a consistent, genet-
ically determined trait. There is also the pos-
sibility that the variability in response is due
to individual snails having learnt to eat egg
masses, while others have not. Complex
manifestations of learning have recently been
demonstrated and partially analysed in gas-
tropod molluscs (Davis & Gillette, 1978;
Chang & Gelperin, 1978). The higher level of
predation by experienced Marisa provides
some support for this hypothesis. However,
the differences between experienced and non-
experienced Marisa were only statistically
significant in one case. The lack of statisti-
cally significant differences in other cases may
have been due to the high variances.
Predatory activity tends to increase with
age, and is significantly higher in sexually
mature females than males. It is plausible to
attribute this change to the need for higher
quality food when the snails enter the repro-
ductive phase. The differences between the
296 CEDEÑO-LEÓN & THOMAS
sexes might be explained by the higher costs
of reproduction in the females. There is
therefore a greater need for females to select
food rich in the galactogen and protein re-
quired for egg formation. There is some evi-
dence that other snails may supplement their
plant diet with protein rich food, particularly
when they mature. Thus Bovbjerg (1968)
found that although lymnaeid snails feed pri-
marily on plant food, they will also feed on
carrion when available. Animal food also fig-
ures prominently in the diets of snail species
such as Physa acuta, Helisoma duryi and
Pomacea sp. (Ferguson, 1977), which have
been proposed as agents for biological con-
trol of the snail hosts. It has been shown un-
der experimental conditions, both in the field
(Eisenberg, 1966, 1970), and in the labora-
tory (Frank, 1963; and El Emam & Madsen,
in prep.), that improved food quality is fol-
lowed by an increase in the growth and na-
tality of pulmonate snails. Generally, higher
protein food appears essential for maximiz-
ing egg production. Further detailed work on
the feeding habits and dietary requirements
of aquatic snails is clearly needed.
In view of the probabilitistic nature of pre-
dation by Marisa, it is necessary to question
whether it has the attributes needed as a
predator for successful biological control. A
good predator should respond to an increase
in prey density in three major ways. Firstly,
individual predators should ingest propor-
tionately more prey as density increases, as
described in the type Ш functional response
of Holling (1959a, b). Secondly, they should
be capable of a numerical response by ag-
gregating in the vicinity of the high prey den-
sities. Thirdly, their reproductive response
should be sufficiently strong to enable them
to control the prey species in a density de-
pendent manner.
Unfortunately, Marisa does not appear to
satisfy these requirements. Thus the preda-
tory activity of the individual does not in-
crease over the range of egg densities used.
It is possible that a functional response could
arise if smaller numbers of egg masses were
used. Secondly, it is unlikely that Marisa could
show a numerical response because Biom-
phalaria deposits its eggs over a wide range
of substrates. Thirdly, the active space within
which the egg masses can be discerned is
likely to be very small. Finally, it seems im-
probable from the information given by Fer-
guson (1977) that Marisa can show a suffi-
ciently strong reproductive response to
increasing numbers of Biomphalaria. De-
tailed demographic studies are needed to
verify this last assumption.
It can be concluded that to optimize on the
predatory activity of Marisa for biological
control it is necessary to introduce them at
high densities. There have, in fact, been cases
where Marisa has been used successfully. For
example, Jobin (1970) and Demian 8 Kamel
(1978) introduced them at densities of one
per 163 cm?, and one per 225 cm? respec-
tively. These densities are very similar to
those used in the present experiments,
namely one per 283 cm”. However, it should
be remembered that Marisa can also induce
other detrimental effects on Biomphalaria, by
acting as a competitor for resources, and by
producing allelopathic factors. Possible
methods of maximising these attributes will
be considered in subsequent papers.
ACKNOWLEDGEMENTS
We are indebted to Consejo de Desarrollo
Cientifico y Humanistico de la Universidad
Central de Venezuela (A.C.L.) and the UNDP/
World Bank/WHO Special Programme for
Research and Training in Tropical Disesases
(J.D.T.) for financial support, Dr. Jesús Al-
berto León for his valuable suggestions, to
Dr. Brian Charlesworth for help with statis-
tics, and Professor John Maynard Smith for
providing facilities.
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brata (Say), the snail host of schistosomiasis:
The search for factors in their media conditioned
by snails which inhibit their growth and repro-
duction. Journal of Applied Ecology, 12: 421-
436.
WORLD HEALTH ORGANIZATION, 1965, Snail
control in the prevention of bilharziasis. WHO
Monograph Series No. 50, Geneva.
MALACOLOGIA, 1983, 24(1-2): 298-311
CONCHIOLIN LAYERS AMONG THE UNIONIDAE AND MARGARITIFERIDAE
(BIVALVIA): MICROSTRUCTURAL CHARACTERISTICS
AND TAXONOMIC IMPLICATIONS
Pieter W. Kat
Department of Earth and Planetary Sciences, The Johns Hopkins University,
Baltimore, Maryland 21218, U.S.A.
ABSTRACT
The purposes of this paper are to determine the function of the conchiolin layers deposited
within the shell of the freshwater bivalve families Unionidae and Margaritiferidae, whether they
contain species-specific characteristics, and whether the microstructure of these layers sup-
ports inclusion of the margaritiferids within the family Unionidae, as proposed by Davis & Fuller
(1981). Scanning electron microscope observations were conducted on 23 unionids and three
margaritiferids. In addition, fossil unionids from the Eocene and Late Jurassic of Wyoming
were examined to determine whether the microstructure of such layers is preserved and can
be used as a taxonomic character among fossils.
The results indicate that unionids possess two types of conchiolin layers within the shell:
thin, homogeneous and thick, more complex layers. The latter attain their highest degree of
complexity within the Unionidae: Ambleminae, especially Elliptio. Unionid complex layers can
be divided into three distinct regions. Margaritiferids, on the other hand, possess only one type
of layer, also divisible into three distinct regions, which closely resembles the periostracum in
ultrastructure.
Species examined possess highly individualistic conchiolin layer characteristics, with the
exception of some members of the recently radiating Elliptio, among which overlaps in both
reticulate region characteristics and inter-population variability occur. Placement of the mar-
garitiferids within the family Unionidae is considered incompatible with the differences observed
among the taxa.
The conchiolin layers prevent rapid shell dissolution in acidic or poorly buffered waters.
Bivalves collected from habitats in which dissolution is severe often show significantly greater
numbers of conchiolin layers per millimeter of shell thickness than do conspecific populations
from habitats where dissolution is less severe. This indicates that these bivalves exercise
control over the frequency of layer deposition, but the mechanism that actuates formation of
layers other than damage-response layers remains obscure.
Unionid fossils from the Eocene of Wyoming have remarkably well preserved conchiolin
layers. Overall characteristics of these layers are highly consistent with those that occur among
Recent taxa examined. When preserved, this feature should allow discrimination of relation-
ships among fossil taxa, and estimation of some environmental parameters, such as water
acidity.
Key words: conchiolin layers; microstructure; Unionidae; Margaritiferidae; taxonomy.
INTRODUCTION
The occurrence of conchiolin layers within
the shells of freshwater bivalves of the fami-
lies Unionidae and Margaritiferidae has been
documented for some time (e.g. Gray, 1833;
Tolstikova, 1974), and recent studies sug-
gest that possession of such conchiolin lay-
ers increases resistance of these bivalves to
shell dissolution after the periostracum has
worn away (Tevesz 8 Carter, 1980; Kat,
1982). Examination of specimens of Elliptio
complanata indicated that two types of con-
chiolin layers are present within the shell of
this species: thin, undifferentiated and thick,
differentiated layers which can be subdivided
into three regions. | proposed (Kat, 1982) that
the microstructure of the thick conchiolin lay-
ers, especially characteristics of the central
reticulate region, might be useful as a taxo-
nomic character at the species level. The pur-
pose of this paper is to test that hypothesis
with a number of Atlantic Slope unionids rep-
resenting two subfamilies, the Anodontinae
and the Ambleminae (sensu Davis & Fuller,
1981).
(298)
CONCHIOLIN LAYERS OF THE UNIONACEA 299
In addition, Tolstikova (1974) indicated a
substantial difference between the Unionidae
and the Margaritiferidae with respect to both
shell and conchiolin layer microstructure. Da-
vis & Fuller (1981) recently included North
American margaritiferids within the family
Unionidae, which seems to contradict Tolsti-
kova's evidence. Conchiolin layer microstruc-
ture of three margaritiferids is compared to
that of a number of unionids to determine
whether the differences observed by Tolsti-
kova (1974) are compatible with subfamilial
status of the margaritiferids.
Taxonomic problems among the Uniona-
cea are especially pronounced among fossil
forms. Loss of all soft-part characteristics, as
well as evidence of ecophenotypic plasticity
has led to much taxonomic uncertainity
among these fossil taxa (Haas, 1969). If mi-
crostructure of conchiolin layers is pre-
served, discrimination among fossil species
should be simplified. Three fossils, one from
the Late Jurassic and two from the Eocene
of Wyoming, were examined to study the
taxonomic value of conchiolin layer micro-
structure in fossils.
METHODS
The classification and collection locations
of the specimens used in this study are listed
in the Appendix. All shells were embedded in
clear plastic, radially sectioned with a circular
rock saw, polished with carborundum grit, and
etched for 5 seconds with 5% HCI in prepa-
ration for scanning electron microscopy
(SEM). During SEM, the specimens were tilt-
ed to reveal clearly the microstructure of the
reticulate portions of the conchiolin layers; the
relative thickness of the uppermost homo-
geneous region of the conchiolin layers thus
is slightly distorted.
Variability within a species and within a
population of a species was studied by ex-
amining seven populations of Elliptio com-
planata and four individuals within each of
four populations of this species (Ellenwood,
French Lake, Norwich, Bull Run). Е. com-
planata was chosen for this survey of vari-
ability because К is one of the most variable
unionid species in terms of shell shape, soft-
part characteristics such as stomach anato-
my and siphonal papillation, and molecular
genetics of the Atlantic Slope unionid assem-
blage (Davis et al., 1981; Kat 8 Davis, in
press). Variability of conchiolin layer charac-
teristics occurring in this species thus might
similarly represent an upper limit to that oc-
curring in other species.
RESULTS
The results of this study indicate the exis-
tence of two separate groups within the bi-
valves examined; as indicated by Tolstikova
(1974), the conchiolin layers among the Mar-
garitiferidae have a different structural orga-
nization from those in the Unionidae.
A) Margaritiferidae (Margaritifera margariti-
fera, Fig. 1C; M. falcata, Fig. 1E; Cumberlan-
dia monodonta, Fig. 1F)
The shell of the margaritiferids contains
only one type of layer, a thick (35 to 80 um)
conchiolin band (Fig. 1A) which is composed
of three distinct regions (Fig. 1C). An outer-
most homogeneous region of approximately
10 to 16 um in thickness surmounts a vacu-
olated region approximately 10 to 20 um
thick. The third region is composed of rather
widely spaced organic lamellae, between
which fit blocks of subprismatic shell materi-
al. This innermost region is the thickest of the
three and varies from about 15 to 40 um. The
general appearance of the conchiolin layers
of the margaritiferids is very similar to that of
the periostracum among these bivalves (Tol-
stikova, 1974; see also Tevesz 8 Carter,
1980).
Among the Margaritiferidae examined,
Margaritifera margaritifera (Fig. 1C) has the
thickest layers, which show the best defini-
tion of the three regions. Cumberlandia mon-
odonta (Fig. 1F) also has good definition of
each region, but the layers are only about half
as thick as those deposited in the shell of M.
margaritifera. Finally, M. falcata (Fig. 1E) has
layers equally thick to those of C. monodon-
ta, but the vacuolated region generally is
poorly developed in most individuals, and the
lamellae seem more randomly placed.
B) Unionidae
Anodontinae (Anodonta gibbosa, Fig. 2A;
А. с. cataracta, Fig. 2B; A. с. fragilis, Fig.
2C; A. implicata, Fig. 2D; Strophitus un-
dulatus, Fig. 2E; Alasmidonta undulata,
Fig. 2F)
Ambleminae: Lampsilini (Lampsilis radiata,
Fig. 2G, H; L. splendida, Fig. 21; L. sp.,
Fig. 2J; L. teres, Fig. ЗА; L. dolabraefor-
300 KAT
— = A : ~ >.
' y. a ae 1e EN eee.
== a er III! A И. Ke
мч МУ A
=
=
Site nr Sag as À
7 Ух
e -
it~ So 4
ae Se
7 - - >. y à or rt e ;
ASI PI EE WA a ee
eee er u > ESE TA A A
FIG. 1. A. Shell of Margaritifera falcata illustrating the exclusive presence of thick conchiolin layers in the
shell. The arrow indicates the direction to the outside of the shell; scale bar = 100 um. B. Shell of Elliptio
complanata illustrating the presence of thick (a) and thin (b) conchiolin layers characteristic of the unionids.
The arrow indicates the direction to the outside of the shell; scale bar = 50 um. C. Thick conchiolin layer
of Margaritifera margaritifera illustrating the three regions present in each layer: upper homogeneous (a),
CONCHIOLIN LAYERS OF THE UNIONACEA 301
mis, Fig. 3B; Ligumia nasuta, Fig. 3C;
Villosa delumbis, Fig. 3D)
Ambleminae: Pleurobemini (Elliptio hope-
tonensis, Fig. 3E; E. shephardiana, Fig.
3F; E. spinosa, Fig. 3G; E. lanceolata,
Fig. ЗН; E. fisheriana, Fig. 31; E. follicu-
lata, Fig. 3J; E. dariensis, Fig. 4A; E.
arctata, Fig. 4B; E. complanata, Fig.
4C-l)
The conchiolin layers of the Anodontinae
are among the simplest observed in this
study. The shell contains two types of layers;
simple, thin (2.5-5 um), homogeneous layers
and slightly more complex layers composed
of at most three partially defined regions: an
outermost homogeneous region (about 3 um
thick), a reticulate region composed of rather
poorly defined chambers, and an innermost,
thin, homogeneous sheet. The reticulate re-
gion in particular is not very well developed;
for example, in Anodonta gibbosa (Fig. 2A)
thin lamellae form only partial chambers, and
in Strophitus undulatus (Fig. 2E) the cham-
bers are small and highly irregular in shape.
The reticulate region varies in thickness from
about 7 um in A. gibbosa to about 5 um in
S. undulatus.
The Ambleminae form a cohesive group
characterized by possession of two types of
layers: thin, homogeneous (2-5 um) and thick,
complex (5-18 um). Thick and thin layers
commonly alternate, but thin shells, such as
those of Lampsilis splendida, Elliptio fisher-
jana, and Ligumia nasuta, frequently have
only one thick layer among several thin ones.
Formation of these two layers seems rather
similar; thick layers could result from elabo-
ration of thin layers. The dimensions of the
homogeneous upper portions of the thick
layers are certainly similar to those of the thin
layers, and thin layers sometimes have an in-
complete reticulate region.
In the Ambleminae thick conchiolin layers
consist of three regions: an outermost, ho-
mogeneous portion, which varies in thick-
ness from about 2 to about 8 um; a central,
reticulate portion (ranging in thickness from
about 3 to about 14 um) composed of cham-
bers of various shapes formed by sheet-like
lamellae; and a lowermost, thin homoge-
neous region. The reticulate region contains
calcareous material within the chambers
formed by the lamellae (‘irregularly shaped
polyhedra” according to Tolstikova, 1974),
which largely dissolved when the specimens
were etched with HCI, but remains visible, for
example, in Fig. 4H. The reticulate region of
the conchiolin layers seems best developed
in the genus Elliptio. The various features of
a thick conchiolin layer characteristic of the
Unionidae are illustrated in Fig. 5.
The fossil specimen of Late Jurassic age
had a highly altered shell in which neither
conchiolin layer microstructure nor shell mi-
crostructure was preserved. In contrast, both
Eocene specimens (Fig. 1G, H) were excep-
tionally well preserved in that they seemed to
retain the original conchiolin and contained
both thin, undifferentiated and thick, differ-
entiated layers. These conchiolin layers are
similar to those of Recent Ambleminae, es-
pecially the Lampsilini, but are quite plesio-
morphic in that the lamellae and chambers
are poorly developed, and the entire reticu-
late region presents a disorganized appear-
ance.
Based on the observed variability of con-
chiolin layer microstructure, it is possible to
define a set of characters that determine ple-
siomorphic and apomorphic conditions among
the Unionidae (Table 1). Highly plesiomorphic
characters appear mainly among the Ano-
dontinae; the Lampsilini and some Pleuro-
bemini have some plesiomorphic characters;
and most other Pleurobemini have apo-
morphic characters. This division agrees in a
general fashion with the previously proposed
times of origin of these taxa: The Anodonti-
nae appeared during the Upper Cretaceous,
the Lampsilini appeared during the Oligo-
cene, and the Pleurobemini appeared during
the Pleistocene (Haas, 1969; Davis et al.,
1981).
Tolstikova (1974) noted no differences
among the microstructures of Unio tumidus
os
central vacuolated (b), and lowermost lamellar (c). Scale bar = 20 um. D. Thick conchiolin layer of Elliptio
waccamawensis illustrating the presence of three regions: uppermost homogeneous (a), central lamellar
(b), and lowermost thin homogeneous (c). Scale bar = 10 um. E. Thick conchiolin layer of Margaritifera
falcata. Scale bar = 15 um. F. Thick conchiolin layer of Cumberlandia monodonta. Scale bar = 15 um. G.
Thick conchiolin layer of a Pseudelliptio from the Eocene of Wyoming. Scale bar = 7 um. H. Thick con-
chiolin layer of a Pseudelliptio from the Eocene of Wyoming. Scale bar = 5 um.
302 KAT
FIG. 2. A. Thick conchiolin layer of Anodonta gibbosa. Scale bar = 4 um. B. Thick conchiolin layer of
Anodonta cataracta cataracta. Scale bar = 4 ит. С. Thick conchiolin layer of Anodonta cataracta fragilis.
Scale bar = 6 ит. D. Thick conchiolin layer of Anodonta implicata. Scale bar = 5 um. E. Thick conchiolin
layer of Strophitus undulatus. Scale bar = 3 um. F. Thick conchiolin layer of Alasmidonta undulata. Scale
bar — 5 um. G. Thick conchiolin layer of Lampsilis radiata (Locality 1). Scale bar — 4 um. H. Thick con-
chiolin layer of Lampsilis radiata (Locality 2). Scale bar — 3 um. I. Thick conchiolin layer of Lampsilis
Splendida. Scale bar = 3 um. J. Thick conchiolin layer of Lampsilis sp. Scale bar = 4 um.
CONCHIOLIN LAYERS OF THE UNIONACEA 303
FIG.3. A. Thick conchiolin layer of Lampsilis teres. Scale bar = 3 um. B. Thick conchiolin layer of Lamp-
silis dolabraeformis. Scale bar = 7 um. C. Thick conchiolin layer of Ligumia nasuta. Scale bar = 5 um. D.
Thick conchiolin layer of Villosa delumbis. Scale bar = 5 ит. E. Thick conchiolin layer of Elliptio hopeto-
nensis. Scale bar = 3 ит. Е. Thick conchiolin layer of Elliptio shephardiana. Scale bar = 3 ит. G. Thick
conchiolin layer of Elliptio spinosa. Scale bar = 7 ит. Н. Thick conchiolin layer of Elliptio lanceolata. Scale
bar = 7 um. 1. Thick conchiolin layer of Elliptio fisheriana. Scale bar = 5 um. J. Thick conchiolin layer of
Elliptio folliculata. Scale bar = 5 um.
304 KAT
FIG.4. А. Thick conchiolin layer of Elliptio dariensis. Scale bar = 5 um. B. Thick conchiolin layer of Elliptio
arctata. Scale bar = 5 ит. С. Thick conchiolin layer of Elliptio complanata (Locality 1). Scale bar = 7 um.
D. Thick conchiolin layer of Elliptio complanata (Locality 2). Scale bar = 5 um. E. Thick conchiolin layer of
Elliptio complanata (Locality 3). Scale bar = 6 um. F. Thick conchiolin layer of Elliptio complanata (Locality
4). Scale bar = 7 um. G. Thick conchiolin layer of Elliptio complanata (Locality 5). Scale bar = 7 um. H.
Thick conchiolin layer of Elliptio complanata (Locality 6). Scale bar = 6 um. I. Thick conchiolin layer of
Elliptio complanata (Locality 7). Scale bar = 7 um. J. Presence of many conchiolin layers in shells from
CONCHIOLIN LAYERS OF THE UNIONACEA 305
FIG. 5. Semi-diagrammatic representation of a
unionid thick conchiolin layer. Illustrated are the
upper (or outermost) homogeneous region (UH), the
central reticulate region (CR), and the lower (or in-
nermost) homogeneous region (LH). The reticulate
region is composed of chambers (C), which are
separated by lamellae (L). The back walls of the
chambers are illustrated on the right; each cham-
ber was filled with calcareous material which dis-
solved during HCl etching. The arrow at the top
indicates the direction to the outside of the shell.
from different provinces in the U.S.S.R., but
such differences are evident when popula-
tions of the phenotypically plastic and genet-
ically variable Elliptio complanata are com-
pared. In eastern Canada, for example,
populations from the Sydney and Nine Mile
Rivers and from French Lake have variously
spaced, approximately vertical lamellae (with
respect to the homogeneous layer) within the
reticulate portion of the conchiolin layer (Fig.
4C, D, E), while the populations from Placide
and Ellenwood Lakes possess curved lamel-
lae (Fig. 4F, G). In contrast, Bull Run (Virgin-
ia) and Norwich Creek (Maryland) popula-
tions are characterized by lamellae that are
interwoven in a variably complex network (Fig.
4H, 1). Individuals in a population, however,
resemble each other closely.
Resemblance in conchiolin layer micro-
structure can occur among different species
of Elliptio. For example, E. arctata is similar
to E. dariensis (Fig. 4A, B). Despite such sim-
ilarities, pronounced differences usually oc-
cur. For example, the lanceolate forms E.
lanceolata, E. folliculata, E. fisheriana, E.
shephardiana, and E. arctata are clearly dif-
ferent (Fig. 3H, J, I, F, and 4B, respectively).
Variability among populations of Elliptio
—
TABLE 1. Plesiomorphic and amorphic charac-
teristics of the reticulate portion of unionid con-
chiolin layers.
A. Plesiomorphic characters.
1. Absence of lamellae, or presence of coarse
lamellae which vary in width or length.
2. Absence of chambers, presence of poorly
defined chambers formed by partial lamel-
lae, or presence of variously sized cham-
bers.
3. Reticulate portion only slightly thicker than
the upper homogeneous portion of the con-
chiolin layer.
B. Apomorphic characters.
1. Presence of well-developed lamellae of rath-
er uniform length and appearance.
2. Presence of well-defined chambers of gen-
erally equal dimensions.
3. Reticulate portion considerably thicker than
the upper homogeneous portion of the con-
chiolin layer.
complanata and overlap of characteristics
among species of this genus contrasts with
conservatism among the Lampsilini and An-
odontinae. For example, L. sp., an unde-
scribed species from Lake Waccamaw, North
Carolina, is different in reticulate region char-
acteristics from both L. radiata and L. splen-
dida, the species found in the drainages
around Lake Waccamaw and from which L.
sp. presumably was derived (Fig. 2G, H, 1, J)
(Kat, in press a).
Despite the high degree of plesiomorphy
among the Anodontinae, it is possible to dis-
criminate among anodontine species. For ex-
ample, Anodonta c. cataracta differs in retic-
ulate region characteristics from A. c. fragilis,
hypothetically a closely related anodontine
from Nova Scotia (Fig. 2B, C) (Clarke 8 Rick,
1963). A. implicata (Fig. 2D) is an exception
to the plesiomorphic trend among the ano-
dontines; while the reticulate portion still pre-
sents a somewhat disorganized picture, the
lamellae are thin and the chambers complete.
DISCUSSION
Functional Significance
The presence of thick conchiolin layers in
bivalve shells usually has been associated
habitats in which dissolution is severe. Specimen is Elliptio complanata from Locality 5. The arrow indi-
cates the direction to the outside of the shell; scale bar = 50 um.
306
(e.g. by Lewy & Samtleben, 1979) with resis-
tance to both predation by boring gastropods
and shell dissolution. In the limited case of
the unionids and margaritiferids the function
of the shell layers has been proposed to fall
in the latter category; after the periostracum
has been worn away from the umbonal re-
gion (which is the oldest part of the bivalve
shell), these successive organic layers might
retard shell dissolution sufficiently to permit
compensatory shell deposition under most
conditions (Kat, 1982). For instance, as a re-
sult of possession of such layers, unionids
can survive with minimal shell damage in
habitats where Corbicula experience con-
siderable mortality due to excessive shell dis-
solution because their shells lack conchiolin
layers (Kat, 1982). In addition, when the
unionid shell is damaged, similar conchiolin
layers are deposited to seal off the affected
area (Beedham, 1965; Tevesz 8 Carter,
1980; personal observations). These dam-
age-response layers can be distinguished
from other conchiolin layers by the presence
of an underlying prismatic layer (Tevesz 4
Carter, 1980).
With the exception of damage-response
layers, it is uncertain what actuates forma-
tion of these conchiolin layers. Layers are de-
posited mainly in small patches in the region
of the umbo, and usually end abruptly (Fig.
1B). There is a suggested relationship be-
tween frequency of conchiolin layers within
the shell and water acidity, although confir-
mation of this trend requires measurement of
inter-population variability in conchiolin layer
abundance with both the averages and stan-
dard deviations of environmental parameters
such as water pH and hardness. Neverthe-
less, bivalves from habitats in which shell dis-
solution is extensive (such as Lake Wacca-
maw, North Carolina; various mesotrophic
and oligotrophic lakes in Nova Scotia; and
certain small creeks in Georgia and Florida)
often have more bands per millimeter of shell
(Fig. 4J) than do conspecifics in habitats
where minimal shell dissolution occurs (Fig.
1B); it thus appears that these bivalves ex-
ercise some degree of control over frequency
of layer deposition. There is no evidence that
these layers correspond to growth stops as
proposed by Tolstikova (1974).
Russell-Hunter et al. (1981) found no rela-
tionships between total organic content and
water hardness in several populations of
freshwater limpets from habitats in which
hardness varied by an order of magnitude.
KAT
Their methods, however, are open to some
criticism: the variance of environmental pa-
rameters such as water pH and hardness ap-
parently was not taken into account, limpets
of various size classes were lumped (large
limpets could have had eroded shells), and
tissues were not extracted from the shells
prior to determination of organic content. It
is not known whether any freshwater gastro-
pods deposit conchiolin layers within their
shells to counter dissolution. Various other
relationships among shell calcium and water
hardness were presented by Russell-Hunter
et al. (1981); it is apparent that freshwater
molluscs have a variety of responses in terms
of shell components and their relationships
to environmental parameters.
Taxonomic Significance
Species-level Discrimination
SEM examinations of conchiolin layer mi-
crostructure reveal that features of the retic-
ulate region, in particular, can often be used
to discriminate among species within a ge-
nus. Exceptions to this trend occur among
some species of the genus Elliptio, which vary
considerably in characteristics among geo-
graphic subgroups and some overlap of
characteristics among species. Conchiolin
layer discriminants among anodontine species
could be fewer and/or more equivocal be-
cause the Anodontinae examined here usu-
ally lack a well-defined reticulate region.
The overlap of characteristics among some
species of Elliptio and the variability among
geographic subgroups of E. complanata are
not surprising. The genus apparently is
undergoing a Recent radiation, and levels of
molecular genetic resemblance among some
species are not different from those that
characterize different populations of a wide-
ranging species. For example, species within
the E. complanata ‘‘group’’ defined by Davis
et al. (1981) are genetically cohesive with
identity values (Nei, 1972) ranging from 0.90
to 0.99. Populations of morphologically de-
fined E. complanata from various locations,
on the other hand, show genetic identity val-
ues ranging from 0.82 to 0.99 (Kat 8 Davis,
in press), and | propose elsewhere (Kat, in
press b) that Е. companata should be re-
garded as a highly polytypic species present-
ly distributed as a Rassenkreis, or ring
species, around the Appalachian mountain
chain. This pattern of close interspecific and
variable intraspecific resemblances generally
CONCHIOLIN LAYERS OF THE UNIONACEA 307
has been considered to be indicative of rapid
speciation and little time since divergence of
species within a genus (Avise et al., 1975;
Avise, 1976; Davis et al., 1981; Kat, in press
a). Morphological characters such as con-
chiolin layer microstructure therefore could
show overall similarity among species unless
those genetic changes that accompanied
speciation directly or indirectly (through pleio-
tropy) affected loci that regulate pattern and
process of conchiolin layer deposition or un-
less such differences accumulated since di-
vergence. The latter process seems to be re-
sponsible for accumulated differences among
widely-separated (and presumably reproduc-
tively isolated) populations of E. complanata,
but it is not possible to determine which pro-
cess is responsible for observed differences
between species which are well separated
genetically. For example, E. folliculata and E.
fisheriana are distantly related (I = 0.64; Da-
vis et al., 1981) and exhibit considerable dif-
ferences in conchiolin layer microstructure
(Fig. 31, J), but it is not possible to determine
by which pathway (directly associated with a
punctuated process of speciation or as a re-
sult of divergence over time) the differences
arose.
Three species within Elliptio (E. shephar-
diana, E. hopetonensis, E. spinosa; Fig. 3E,
F, G) show considerable plesiomorphy in
conchiolin layer microstructure when com-
pared with other members of the genus ex-
amined here. All three species are endemic
to the ancient Altamaha River drainage
(Johnson, 1970) and could represent a group
of species that diverged from ancestral Ellip-
tio early in time and thus retained some ple-
siomorphic characters.
Members of the Anodontinae and Amble-
minae: Lampsilini do not show such variabil-
ity. Both groups are characterized by rather
plesiomorphic conchiolin layers, but there are
some exceptions to this overall trend. Lamp-
silis dolabraeformis (Fig. 3B), for instance,
has the most apomorphic conchiolin layers
among the lampsilines examined. This species
can be characterized as “advanced” with re-
spect to other features as well; for example,
the mantle flap and marsupium show highly
derived conditions. Anodonta implicata (Fig.
2D) is an exception to the plesiomorphic trend
among the anodontines; the lamellae in this
species are thin and complete, and the cham-
bers are quite regular. | have proposed sep-
aration of A. implicata from the subgenus Py-
ganodon (which includes all other Anodonta
TABLE 2. Characteristics of the conchiolin layers
which differentiate the Unionidae and Margeritifer-
idae.
Unionidae
1. Presence of two types of conchiolin layers
within the shell: thin, undifferentiated, and
thick, differentiated.
2. Thick, differentiated layers can always be di-
vided into three or more or less distinct lay-
ers: an uppermost homogeneous portion; a
central reticulate portion, and a lowermost thin
homogeneous portion.
3. Chambers of various sizes are present within
the reticulate portion of the conchiolin bands,
which are filled with unconsolidated, chalky
shell material.
Margaritiferidae
1. Presence of one type of conchiolin layer with-
in the shell: thick, differentiated.
2. Differentiated layers can be divided into three
layers: an uppermost homogeneous portion,
a central vacuolated portion, and a lower-
most lamellar portion. The appearance of the
conchiolin layers is similar to that of the peri-
ostracum.
3. Chambers of various sizes are formed within
the lamellar portion, which are filled with
blocks of subprismatic shell material.
examined here) because of electrophoretic
and soft-part characteristic differences (Kat,
1983); the differences observed in conchiolin
layer microstructure support this separation.
| also described a substantial difference be-
tween A. c. cataracta and A. c. fragilis; the
latter “subspecies’ differs considerably from
the former in electrophoretic and soft-part
characters (in fact, A. c. fragilis shows close
affinity to the European A. cygnea with re-
spect to stomach morphology); conchiolin
layer characteristics again positively corre-
late with other discriminants (Fig. 2B, C).
Taxonomic concepts based on conchiolin
layer similarities, however, sometimes dis-
agree with relationships suggested by elec-
trophoresis: for example, Lampsilis sp. seems
more closely related to L. splendida than to
L. radiata with respect to overall conchiolin
layer microstructure (Fig. 2H, 1, J) while the
reverse relationship is suggested by electro-
phoretic and soft-part similarities (Kat, in
press a).
Higher-order Taxonomy
Two very distinct categories can be de-
fined on the basis of characteristics of the
308
conchiolin layers among the species exam-
ined (Table 1). The Unionidae show varying
degress of elaboration of the conchiolin lay-
ers, which appear to be the most complex
among the Ambleminae. Also, all unionids
possess thin, undifferentiated bands, which
could be indicative of plesiomorphy. The mar-
garitiferids show very different, periostra-
cum-like bands, which are unlike any encoun-
tered among the unionids examined.
Tolstikova (1974) was equally convinced of
the differences between unionids and mar-
garitiferids with respect to this character and
described some additional significant differ-
ences in shell microstructure.
| propose two hypotheses to explain these
differences:
(1) Recent margaritiferids show conchiolin
layers that are as highly derived from an an-
cestral state as those of the Pleurobemini are;
in other words, the fact that these layers are
now very different does not mean that they
could not once have been very similar. This
hypothesis is rejected for two reasons. First,
while the conchiolin layers of the Pleurobe-
mini can be regarded as apomorphic, the si-
multaneous presence of thin, undifferentiated
layers within the shell, as well as the pres-
ence of features that link differentiated and
undifferentiated bands among the unionids,
points to common ancestry. This is support-
ed by close resemblance of conchiolin layer
microstructure between Recent species and
Eocene taxa. A similar argument cannot be
constructed for the margaritiferids; they lack
all but differentiated layers, and these layers
exhibit few features in common with those of
the unionids. Second, conchiolin layers of the
margartiferids resemble the periostracum in
ultrastructure, while those of the unionids do
not.
(2) Recent margaritiferids have conchiolin
layers that are derived from a very different
ancestral condition; in other words, they
arose in a lineage different from that of the
Unionidae. This hypothesis is most compati-
ble with the data.
It appears that there have been two dis-
tinctly different pathways taken by different
unionacean lineages towards the solution of
the common problem of shell dissolution. An-
cestral unionids likely laid down simple to
slightly elaborated conchiolin layers, similar
to those encountered among Recent ano-
dontines. Elaboration of such bands is en-
KAT
countered among Recent Lampsilis and Ellip-
tio. Ancestral margaritiferids likely formed
layers similar to the periostracum. Recent
margaritiferids continue to have this ances-
tral condition. This hypothesis should be
testable in the fossil record.
The classification of Davis & Fuller (1981),
which relegates the margaritiferids to
subfamilial status within the Unionidae, is
therefore rejected; the margaritiferids have
conchiolin layer microstructural differences
suggestive of an early divergence from the
unionid lineage, and on the basis of this char-
acter | propose restoration of familial status.
On the other hand, similarities among the
Lampsilini and Pleurobemini with respect to
conchiolin layer microstructure should con-
stitute additional evidence to support includ-
ing them in the same subfamily (Ambleminae)
as proposed by Davis & Fuller (1981).
Placement of the margaritiferids within the
Unionidae was based on molecular genetic
and some morphologic data (Davis & Fuller,
1981). First, genetic similarity was found to
be higher than expected if the taxa belonged
to different families (comparative data were
derived from comparisons of different gastro-
pod families), especially because fossil evi-
dence indicated that the divergence could
have begun before the Cretaceous (Haas,
1969). With respect to electrophoretically de-
termined genetic distances, remaining simi-
larities among distantly related taxa generally
are found among slowly evolving loci (Sarich,
1977). Divergence times based on genetic
distances that include such loci must include
a large margin of error, because rates of dif-
ferentiation for such loci are unknown, be-
cause it is unknown to what extent natural
selection maintains similarities among such
loci, and because it is not intuitively obvious
why genetic distances between distantly re-
lated taxa should continue to increase in a
regular fashion. In fact, the strict applicability
of the molecular clock has recently been
questioned: sea urchin species pairs sepa-
rated by the Isthmus of Panama reveal radi-
cally different genetic distances, even though
they have presumably been isolated for ex-
actly the same amount of time (Lessios, 1979,
1981). Some taxa thus might diverge more
rapidly than others. Cluster-ordination analy-
sis of immunoelectrophoretic distances also
indicates high levels of similarity between the
Ambleminae and Margaritifera (Davis 8 Full-
er, 1981). In sum, the combined electropho-
retic and immunoelectrophoretic data indi-
CONCHIOLIN LAYERS OF THE UNIONACEA 309
cate that the taxa proabably are not of
polyphyletic origin, but the distinct differ-
ences observed between the Margaritiferidae
and the Unionidae in conchiolin layer micro-
structure must place the taxa in different
families, which | propose shared a common
ancestor before the Cretaceous (Fig. 6).
Second, similarities exhibited by the mar-
garitiferids to the unionid morphological
groundplan, including the glochidial larval
type, could imply similarity of response to
similar selective pressures. Margaritiferids and
unionids share identical habitats, and para-
sitize identical hosts (fishes); it is thus entire-
ly likely that, given a similar ancestral bivalve
groundplan, the taxa now resemble each
other in a general fashion. It must be noted
that margaritiferid glochidia are much smaller
than those found among the unionids, and
that careful comparative observations have
not been made to support contentions of their
similarity. An initial examination of soft-part
characteristics of margaritiferids and union-
ids reveals significant differences between the
taxa. Siphonal papillae among the margariti-
ferids are both muscular and arborescent, a
condition not encountered among unionids
thus far examined (although some Amblemini
apparently possess arborescent papillae; see
Davis & Fuller, 1981); margaritiferids pos-
sess no true septa or water tubes in their
lamellae; and examination of characteristics
of the margaritiferid stomach reveals that it
is simple in structure, resembles those of the
anodontines, but differs from the unionids
thus far examined in possession of a sorting
pouch beneath the minor typhlosole fold (Kat,
1983; personal observations).
Fossils
According to Haas (1969), loss of all soft-
part characters, evidence of a high degree of
phenotypic plasticity of unionacean shell
shape, and gaps in knowledge of Recent
forms contribute to render classification of
fossil forms an extremely difficult endeavor.
In addition, fossil specimens often are frag-
mentary, or preserved only as casts or molds,
which further reduces the amount of infor-
mation that can be deduced from them. Pres-
ervation of microstructural characteristics of
the conchiolin layers within shells that have
undergone little replacement, or replacement
with little deformation of the original micro-
structure of the shell, would allow identifica-
tion of species as well as degree of pheno-
UNIONIDAE
MARGARITIFERIDAE
AMBLEMINAE
ANODONTINAE
FIG. 6. Relationships among North American
Unionacea based on conchiolin layer microstruc-
ture. In the diagram, a generalized unionacean
ancestor A gives rise to descendant groups B and
C, present during the Cretaceous (K). Group B is
characterized by deposition of conchiolin layers
which resemble the periostracum in ultrastructure,
and is ancestral to the Margaritiferidae. Group C
is characterized by deposition of simple, non-dif-
ferentiated layers within the shell, and is ancestral
to the Unionidae. This group then diverges (D) into
the Ambleminae, which has differentiation of con-
chiolin layers into separate portions, and the Ano-
dontinae, which largely retain simple layers.
typic plasticity shown by such species, even
from fragmentary material. Such information
is important in that it allows assessment of
fossil species diversity, determination of re-
lationships between fossil and Recent taxa,
and because the number of layers deposited
in the shell seem related to water acidity,
some environmental reconstruction. Unfor-
tunately, preservation of conchiolin layer mi-
crostructure requires rather exceptional con-
ditions, which might only seldom be met;
permineralization which faithfully replicates
the conchiolin layer could be extremely rare.
The Eocene fossils studied were exception-
ally well preserved, and appear to have re-
tained the original components of the con-
chiolin layers (Fig. 1G, H). In contrast, a fossil
specimen from the Late Jurassic had been
extensively altered. Preservation of taxo-
nomically valuable characters might thus be
limited to specimens of Tertiary or younger
age. Nevertheless, good preservation of con-
chiolin layer microstructure observed in this
preliminary study of fossil forms is extremely
encouraging, and should provide a useful way
to discriminate among previously problemat-
ical fossil taxa.
310
SUMMARY
Thick conchiolin layers within the shells of
the Unionidae and the Margaritiferidae seem
to serve a common purpose: prevention of
rapid shell dissolution in the region of the
umbo once the protective periostracum has
worn away. Most species examined, with the
exception of some recently diverging taxa
within the genus Elliptio, seem to possess
highly individualistic characteristics of espe-
cially the reticulate regions of the layers.
Placement of the margaritiferids within the
Unionidae as proposed by Davis & Fuller
(1981) is considered incompatible with the
significant differences observed in the micro-
structure of the conchiolin layers of these
taxa; there seem to have been two distinctly
different pathways taken by the different lin-
eages toward the solution of the common
problem of shell dissolution. The margaritife-
rids deposit layers that resemble the perios-
tracum, while the unionids lay down simple
and variably complex chambered conchiolin
layers, which do not resemble the periostra-
cum.
Preservation of conchiolin layer micro-
structure among Eocene taxa suggests the
existence of a powerful tool to discriminate
among fossil forms, and should facilitate fos-
sil classification. Such discriminatory ability
should additionally allow estimates of fossil
assemblage diversity, relationships among
fossil and Recent taxa, and reconstruction of
some environmental parameters such as
water pH.
ACKNOWLEDGEMENTS
| thank Drs. Alan Walker and Pat Shipman,
and especially Jenny Rose for SEM assis-
tance; their patience during tedious SEM ex-
aminations of shell structures was remark-
able. | am further indebted to Gene Meyer for
assistance during many field trips, to Al Leech
for guidance during specimen preparation, to
Dr. George M. Davis for insightful comments
which improved the manuscript, and to the
anonymous reviewers for their suggested re-
visions. This study was funded, in part, by a
Biomedical Research Support Grant.
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APPENDIX. Classification and collection
localities of the species studied
Recent Taxa
Unionidae
Anodontinae
Anodonta gibbosa Say, 1824
Ocmulgee River, Ben Hill Co., Georgia
Anodonta cataracta cataracta Say, 1817
Pickering Creek, Chester Co., Pennsyl-
vania
Anodonta cataracta fragilis Lamarck, 1819
Nowlans Lake, Digby Co., Nova Scotia,
Canada
Anodonta implicata Say, 1829
Fletcher Lake, Halifax Co., Nova Scotia,
Canada
Strophitus undulatus (Say, 1817)
Norwich Creek, Talbot Co., Maryland
Alasmidonta undulata (Say, 1817)
Norwich Creek, Talbot Co., Maryland
Ambleminae
Lampsilini
Lampsilis radiata (Gmelin, 1791)
1. Shubenacadie Grand Lake, Halifax Co.,
Nova Scotia, Canada
2. Newville Lake, Cumberland Co., Nova
Scotia, Canada
Lampsilis splendida (Lea, 1838)
Ocmulgee River, Ben Hill Co., Georgia
Lampsilis dolabraeformis (Lea, 1838)
Ocmulgee River, Ben Hill Co., Georgia
Lampsilis teres (Say, 1834)
Withlacoochee River, Lacoochee, Pasco
Co., Florida
Lampsilis sp.
Lake Waccamaw, Columbus Co., North
Carolina
Ligumia nasuta (Say, 1817)
Norwich Creek, Talbot Co., Maryland
Villosa delumbis (Conrad, 1834)
Fountain Mill, Pulaski Co., Georgia
Pleurobemini
Sune complanata (Lightfoot, 1786)
. Sydney River, Cape Breton Co., Nova
Scotia, Canada
2. French Lake, Sunbury Co., New
Brunswick, Canada
3. Nine Mile River, Halifax Co., Nova
Scotia, Canada
4. Ellenwood Lake, Yarmouth Co., Nova
Scotia, Canada
5. Placide Lake, Digby Co., Nova Scotia,
Canada
6. Norwich Creek, Talbot Co., Maryland
7. Bull Run, Prince William Co., Virginia
Elliptio fisheriana Ortmann, 1919
Norwich Creek, Talbot Co., Maryland
Elliptio waccamawensis (Lea, 1863)
Lake Waccamaw, Columbus Co., North
Carolina
Elliptio hopetonensis (Lea, 1838)
Ocmulgee River, Ben Hill Co., Georgia
Elliptio spinosa (Lea, 1836)
Ocmulgee River, Ben Hill Co., Georgia
Elliptio shephardiana (Lea, 1834)
Ocmulgee River, Ben Hill Co., Georgia
Elliptio lanceolata (Lea, 1820)
Fountain Mill, Pulaski Co., Georgia
Elliptio dariensis (Lea, 1842)
Bowens Mill, Ben Hill Co., Georgia
Elliptio arctata (Conrad, 1834)
Mosquito Creek, Gadsden Co., Florida
Elliptio folliculata (Lea, 1858)
Lake Waccamaw, Columbus Co., North
Carolina
Margaritiferidae
Margaritifera margaritifera (Linnaeus, 1758)
Maccan River, Cumberland Co., Nova
Scotia, Canada
Margaritifera falcata (Gould, 1850)
Deschutes River, Deschutes Co., Oregon
Cumberlandia monodonta (Say, 1829)
Clinch River, Hancock Co., Tennessee
Fossil Taxa
Genus?
Jurassic: Lower Morrison Formation, East
Como Bluff, Albany Co., Wyoming.
Johns Hopkins University M-COMO-
CLM-1
Pseudelliptio sp.
Eocene: Willwood Formation, Elk Creek,
Big Horn Co., Wyoming. Johns Hop-
kins University W-KC-PSE-1
MALACOLOGIA, 1983, 24(1-2): 312-322
INDEX TO TAXA IN VOLUME 24
An asterisk (*) denotes a new taxon
Aaeolidia, 140
abietina, Abietinaria, 176
Abietinaria, 172, 173, 176
abronia, Cratena, 164
abronia, Cuthona, 127, 132, 222, 267
abronia, Trinchesia, 164
Acanthodoris, 116-136, 261, 262
Acarnus, 170, 209
Acleioprocta, 118, 120
Acroloxus, 72
acronicus, Gyraulus, 1-101
acronicus, Planorbis, 45
Actinia, 141
Actinocyclidae, 119
Actinocyclus, 181
Actinothoe, 141
acuta, Physa, 289, 296
Adalaria, 199
adhaerens, Mycale, 146, 149
Adocia, 180
adspersa, Galvina, 211
adspersa, Tenellia, 126, 132, 211, 212, 222, 275
adspersus, Tergipes, 211, 212
Aegires, 119, 121, 129, 137, 138, 262
Aegiretidae, 119
Aegirus, 137
aeglifinus, Melanogrammus, 141
Aeolidia, 115-141, 200, 262
Aeolidida, 140
Aeolidiella, 115, 120, 131, 140-142, 209, 210
Aeolidoidea, 115, 118, 120
Aeolis, 138-200
Aeolodia, 140
Aerolidia, 140
affinis, Doris, 196
affinis, Flabellina, 160
Afrogyrorbis, 75, 80, 93
Afrogyrus, 80, 88, 93
agassizii, Hypselodoris, 192
agennes, Myxilla, 146
aggregata, Bowerbankia, 207
Aglaophenia, 176, 177, 179, 184, 201
agraulus, Gyraulus, 36
Aiptasia, 141
akkeshiensis, Dirona, 178
alabastrina, Doris, 117
Alasmidonta, 299, 302, 311
alba, Atagema, 122, 130, 150, 264
alba, Conualevia, 121, 130, 157, 266
alba, Eolidia, 187
alba, Fiona, 188
alba, Glossodoridiformia, 150, 151
albescens, Acanthodoris, 135, 136
albidus, Gyraulus, 67, 72, 73
albina, Eolis, 140
albocrusta, Cratena, 165
albocrusta, Cuthona, 127, 132, 165, 267
albocrusta, Trinchesia, 165
albolineata, Dirona, 125, 131, 178, 179, 269
albopunctata, Aegires, 137, 268
albopunctata, Dendrodoris, 122, 169-171
albopunctata, Doriopsilla, 138, 170
albopunctata, Doriopsis, 170
albopunctata, Doris, 138, 169
albopunctatus, Aegires, 121, 129, 137, 138, 262
albopunctatus, Aegirus, 137
albopunctatus, Cregires, 137
albopunctatus, Dendrodoris, 170
albus, Anisus, 33
albus, Dendronotus, 125, 131, 172, 269
albus, Gyraulus, 1-101
albus, Planorbis, 27, 28, 99, 100
Alcyonidium, 136
Aldisa, 119, 122, 129, 142, 143, 213, 263
Aldisidae, 119
alpha, Catriona, 166
alpha, Cuthona, 166, 167
alpicola, Paratrichocladius, 93
Ambleminae, 298, 299, 301, 307-309, 311
amblia, Dysidea, 157, 183, 192
amphispicula, Syringella, 149
Amphitridea, 174
Amphitrite, 173
amphora, Abietinaria, 172
Amphorina, 166, 167
amplificatus, Gyraulus, 100
amyra, Doto, 126, 131, 183, 185, 270
Anadoridacea, 118, 119
anatifera, Lepas, 188
Ancula, 119, 121, 129, 143-145, 263
Anculinae, 119
Anemonia, 141
angelensis, Cargoa, 195
angelensis, Okenia, 121, 195, 272
angulata, Caillaudia, 100
angulata, Gyraulus, 55
Anisodoris, 116-148, 180, 263
Anisus, 10-102
Anodonta, 299, 301, 302, 305, 307, 311
Anodontinae, 298, 299, 301, 305, 307, 309, 311
anserifera, Lepas, 188
Anthea, 141
Anthopleura, 141, 202
Antiopella, 117, 120, 131, 146, 147, 200, 264
Aplidium, 201
Aplysilla, 153, 154
arborescens, Dendronotus, 173, 174
arborescens, Doris, 173
arborescens, Tritonia, 173, 174
arbutas, Rostanga, 209
Archidorididae, 119
Archidoris, 119-149, 180, 264
arctata, Elliptio, 304, 305, 311
Arctocorisa, 93
argentea, Sertularia, 176
argentea, Thuiaria, 172
(312)
INDEX 313
Armiger, 10-102
Armina, 120, 129, 131, 149, 150, 264
Arminidae, 120
Arminoidea, 118, 120
armoricana, Eolis, 188
artemisia, Anthopleura, 141
ascani, Tritonia, 173
Asteronotidae, 119
Asteronotus, 141
Atagema, 119, 122, 130, 150, 151, 264
atlantica, Fiona, 187
atra, Polycera, 121, 129, 204, 206, 274
aurantia, Tritoniopsis, 213
aurantiaca, Triopha, 217
aurelia, Paramaecium, 24
aureocincta, Antiopella, 146, 147
auricularia, Radix, 60
Austrodoris, 149
Babaina, 152
Babakina, 120, 125, 132, 151, 152, 264
balanoides, Balanus, 198
Balanus, 198
balli, Anthopleura, 141
banyulensis, Aldisa, 143
barbarensis, Aeolis, 146
barbarensis, Antiopella, 117, 125, 131, 146, 147,
200, 264
barbarensis, Janolus, 146
Barentsia, 144
Bathyomphalus, 39, 74-102
beaumonti, Coryphella, 163, 267
beaumonti, Cumanotus, 115, 126, 132, 163, 164
bellulus, Stiliger, 212
bicarinatus, Gyraulus, 75
bifida, Doris, 135
bilabiata, Eurystomella, 190
bilamellata, Doris, 196, 197
bilamellata, Lamellidoris, 196, 197
bilamellata, Onchidoris, 122, 129, 196-198
bilamellata, Onchidorus, 197
bilamellata, Oncidoris, 196, 272
bilamellatus, Doris, 196
billamellata, Lamellidoris, 197
bimaculata, Thordisa, 123, 130, 142, 213, 275
Biomphalaria, 11, 14, 289-297
biwaensis, Anisus, 69
biwaensis, Gyraulus, 67-73, 75, 100
biwaensis, Planorbis, 69, 71
bodoensis, Doris, 138
bodoensis, Eolidia, 138
boettgerianus, Acroloxus, 72
boholiensis, Discodoris, 179, 180
bolini, Corambella, 182
borealis, Gyraulus, 46
borealis, Heterocope, 93
borealis, Planorbis, 45
Botryllus, 176
Bougainvillia, 176, 212
Bowerbankia, 207
bowerbanki, Halichondria, 149, 182
brepha, Hymendesmia, 143
brongersmai, Gyraulus, 94, 97
brunnea, Acanthodoris, 123, 128, 131, 133, 261
Bugula, 147, 205, 206, 216, 217
Bulinus, 10, 11, 22, 79, 289
Cabrilla, 217, 218
Cadlinidae, 119
Cadlina, 115-133, 152-154, 264, 265
Caillaudia, 1, 55, 76-102
calensis, Chromodoris, 191
calensis, Glossodoris, 191
californica, Armina, 129, 131, 149, 150, 264
californica, Bugula, 147
californica, Corynactis, 141
californica, Hancockia, 124, 131, 189, 271
californica, Pleurophyllidea, 150
californica, Pleurophyllidia, 149, 150
californica, Scrupocellaria, 216, 217
californicum, Aplidium, 201
californiensis, Chromodoris, 191
californiensis, Glossodoris, 191
californiensis, Hypselodoris, 117, 122, 129, 191,
192, 272
callarius, Gadus, 141
Callopora, 136
Campaspe, 174
camtschatica, Phacellophora, 201
Capellinia, 186
capillata, Cyanea, 201
Cardlina, 152
Cargoa, 195
carinata, Arctocorisa, 93
carinata, Atagema, 150
carinata, Lacuna, 178
carinatus, Planorbis, 9, 92
Carinogyraulus, 1, 67-102
Carinulorbis, 75
carnea, Chromodoris, 156
carnulentum, Didemnum, 189
carpenteri, Triopha, 215
caspia, Cordylophora, 212
catalinae, Triopha, 122, 129, 215, 216, 275
cataracta, Anodonta, 299, 302, 305, 307, 311
Catriona, 164, 166, 168
Caulibugula, 216, 217
Cauloramphus, 216, 217
Cavolina, 199-201
celata, Cliona, 170
Cellaria, 216
Celleporella, 179
Ceratophallus, 75
Cerberilla, 120, 129, 131, 155, 265
Cereus, 141
cervina, Doris, 173
cervina, Tritonia, 173
chani, Hallaxa, 123, 129, 189, 271
chilensis, Lophogorgia, 205, 222
chinensis, Gyraulus, 1-103
chinensis, Planorbis, 56
Chioraera, 193, 194
Chirolophis, 202
Chironomidae, 289
Choanomphalodes, 1, 42, 73-102
Choanomphalus, 66-79, 99, 100
314
Chromodorididae, 119
Chromodoridinae, 156
Chromodoris, 116-130, 155-157, 191, 192, 265
chromosoma, Eolidina, 210
chromosoma, Spurilla, 126, 131, 210, 275
ciliata, Caulibugula, 216, 217
cincta, Diadumene, 141
Ciona, 201
circumstriatus, Gyraulus, 22, 81
citrina, Acanthodoris, 135
Clangula, 198
Clavularia, 220
Cleioprocta, 118, 120
Cliona, 170
coccinea, Rostanga, 208
coccinea, Stomphia, 141
cockerelli, Laila, 121, 129, 192, 272
cockerelli, Leilla, 193
cocoachroma, Cuthona, 129, 165
coeruleopictus, Janolus, 117, 147
columbiana, Amphorina, 166, 167
columbiana, Armina, 150
columbiana, Catriona, 166
columbiana, Cratena, 166
columbiana, Cuthona, 127, 132, 166, 167, 267
columbiana, Doto, 126, 131, 184
columbiana, Galvina, 166
columbiana, Idulia, 184
columbina, Acanthodoris, 133, 134
commissuralis, Obelia, 176
complanata, Elliptio, 298-301, 304-307, 311
complanatus, Hippeutis, 11, 41
compressus, Planorbis, 97
concinna, Porella, 136
coneja, Crimora, 121, 129, 163, 267
connollyi, Gyraulus, 53, 55, 79, 100
contortus, Bathyomphalus, 92
Conualevia, 119, 120, 130, 157, 266
Conualeviidae, 119
conventus, Pisidium, 94
convexiusculus, Gyraulus, 1, 55-62, 102
convexiusculus, Planorbis, 55
cooperi, Aldisa, 143
cooperi, Coryphella, 126, 132, 159, 162, 266
Copepoda, 289
Corambe, 116, 119, 129, 158, 182, 266
Corambella, 182
Corambidae, 119
Corbicula, 306
Cordylophora, 212
coriacea, Tealia, 141
cornuarietis, Marisa, 289-297
coronata, Doris, 196
coronata, Doto, 183
Corydendrium, 212
Corymorpha, 147
Corynactis, 141
Coryne, 176
Coryphella, 120, 126, 132, 158-163, 266
Coryphellidae, 120, 161
costulatus, Gyraulus, 11, 19, 20, 23, 53-55, 100
costulatus, Planorbis, 54, 55, 100
couchi, Aiptasia, 141
Craniella, 149
INDEX
crassicornis, Cavolina, 199-201
crassicornis, Facelina, 199, 200
crassicornis, Hermissenda, 146, 200, 201
crassicornis, Phidiana, 117, 125, 132, 199, 202, 273
crassicornis, Tealia, 141
crassicornus, Phidiana, 201
Cratena, 164-168, 188
Cregires, 137
crenatus, Balanus, 198
crenophilus, Gyraulus, 19, 67, 68, 100
Crimora, 119, 121, 129, 162, 163, 267
Crisia, 216, 217
crista, Armiger, 42, 43, 94
crista, Gyraulus, 16, 42-44, 79-100
crista, Nautilus, 44, 100
cristata, Ancula, 144
cristata, Antiopella, 146
crocea, Tubularia, 115, 159, 162, 164-166, 176,
201
Crossaster, 222
crosseanus, Planorbis, 27
Cryptobranchia, 119
Cryptosula, 136, 198
Cumanotinae, 120
Cumanotus, 115, 120, 132, 162-164, 267
Cumberlandia, 299, 301, 311
cupressina, Sertularia, 176
Cuthona, 120, 127, 129, 132, 164-169, 201, 207,
222, 267, 268
Cuthonidae, 118, 120
Cuthoninae, 120
cuverii, Eolis, 138
cuvieri, Aeolis, 139
cuvieri, Eolidia, 139
cuvieri, Eolis, 138
cuvierii, Eolidia, 138
cuvierii, Eolis, 139
Cyanea, 201
cygnea, Anodonta, 307
Cystoseira, 187
dalli, Chioraera, 194
dalli, Chromodoris, 192
dalli, Dendronotus, 175
dalli, Melibe, 194
dariensis, Elliptio, 301, 304, 305, 311
dawsoni, Solaster, 222
decorata, Hallaxa, 189
deflectus, Gyraulus, 33
deformis, Gyraulus, 27, 45
deformis, Planorbis, 27, 33
delumbis, Villosa, 301, 303, 311
Dendrobeania, 216, 217
Dendrodorididae, 120
Dendrodoris, 120, 122, 169-171, 222, 268
Dendrontidae, 120
Dendronotoidea, 118, 120
Dendronotus, 116-131, 171-177, 269
Dendrophis, 90
devians, Gyraulus, 48, 49
devians, Planorbis, 48
Diadumene, 141
Dialula, 181
diamedia, Tritonia, 220
INDEX 315
dianthus, Metridium, 141 elegantissima, Hymenaeolis, 187
Diaulula, 179-181 elfortiana, Doris, 196
Diaululae, 180 elioti, Triopha, 216
Diaululua, 181 Elliptio, 298-301, 303-308, 310, 311
dichotoma, Obelia, 212 Elminius, 198
Dicodoris, 180 Emarcusia, 199, 203
Didemnum, 189 Embletonia, 211, 212
diegensis, Scrupocellaria, 216 emurai, Cuthona, 201
digueti, Armina, 150 emurai, Dondice, 201
Dilella, 181 emurai, Hervia, 201
diomedea, Duvaucelia, 219-221 emurai, Shinanoeolis, 201, 202
diomedea, Sphaerostoma, 219 Eolida, 138
diomedea, Tritonia, 117, 124, 131, 219, 220, 276 Eolidia, 138, 139, 187
diomedia, Sphaerostoma, 219 Eolidina, 141, 210
diomedia, Tritonia, 219 Eolis, 138-141, 187, 188, 200, 211
Diplosoma, 160 Epiactis, 141
Dirona, 116-131, 177-179, 269 equina, Actinia, 141
Dironidae, 120 erithacus, Acarnus, 170, 208
Discodorides, 180 esculans, Tritonia, 220
Discodorididae, 119 Esperiopsis, 209
Discodoridinae, 119 essingtonensis, Gyraulus, 82
Discodoris, 115-145, 179-183, 270 estrella, Estrella, 192
Disculifer, 84, 89 Euarminacea, 120
dispar, Corydendrium, 212 Eubranchidae, 118, 120
distans, Hydrallmania, 173 Eubranchinae, 120
divae, Cuthona, 207 Eubranchus, 116-132, 185, 186, 211, 222, 271
divae, Precuthona, 129, 132, 207, 274 eudactylota, Hancockia, 189
diversicolor, Dendronotus, 125, 131, 172, 269 Eudendrium, 160, 162
dolabraeformis, Lampsilis, 299, 303, 307, 311 Eudoridacea, 115, 118, 119, 180
Dolicheolis, 188 Eueolidacea, 120
Dondice, 201 “eugyne, Gyraulus, 1, *64-102
Doridella, 119, 129, 182, 270 euphraticus, Gyraulus, 15-103
Dorididae, 119 Eurystomella, 190
Doridoidea, 119 exiguus, Eubranchus, 186
Doridopsis, 170, 171, 181 eximia, Syncoryne, 167, 176
Doriopsilla, 138, 169-171 exsulans, Duvaucelia, 219
Doriopsis, 169-171 exsulans, Sphaerostoma, 219
Doris, 117-213, 270 exsulans, Tritonia, 219-221
dortmanna, Lobelia, 93
Dotidae, 120 Facelina, 199, 200, 202, 203
Doto, 116, 120, 126, 131, 183-185, 270, 271 facrici, Amphitridea, 173
Draco, 95 falcata, Margaritifera, 299-301, 311
Drepania, 214 farinacea, Aeolis, 139
Drepanotrema, 83 farinacea, Eolis, 139
dumerili, Callopora, 136 fasciculata, Eolidia, 187
dura, Petrosia, 182 fasciculata, Eolis, 187
duryi, Helisoma, 289, 296 fasiculata, Doris, 187
Duvaucelia, 213, 219-221 felina, Dendronotus, 174
Dynamena, 176 felina, Tealia, 141
Dysidea, 157, 183, 192 felina, Tritonia, 173
festiva, Babaina, 152
echinata, Hydractinia, 176 festiva, Babakina, 125, 132, 151, 152, 264
Echinostoma, 2 festiva, Lateribranchia, 221
edulis, Mytilus, 201 festiva, Lateribranchiaea, 221
ehrenbergi, Gyraulus, 23, 52-99 festiva, Lateribranchiaia, 221
ehrenbergi, Planorbis, 52 festiva, Sphaerostoma, 221
Eichhornia, 62 festiva, Tritonia, 124, 131, 221, 276
eleanor, Leucosolenia, 138 feuerborni, Gyraulus, 95
Electra, 136 Ficulina, 170
elegans, Dendronotus, 174 Filicrisia, 216, 217
elegans, Okenia, 195 fimbriatus, Pachycerianthus, 177
elegans, Sagartia, 141 Fiona, 120, 126, 132, 187, 188, 271
elegantissima, Anthopleura, 141, 202 Fione, 188
elegantissima, Fiona, 188 Fionidae, 120
316 INDEX
firma, Lissodendoryx, 146
fischeri, Coryphella, 161
fisheri, Coryphella, 161
fisheriana, Elliptio, 301, 303, 305, 307, 311
flabellata, Obelia, 176
Flabellina, 159, 160, 199, 200
Flabellinidae, 118
Flabellinopsis, 159, 160
flavomaculata, Cadlina, 124, 129, 152, 264
flavomaculata, Cardlina, 152
flavovulta, Cratena, 167
flavovulta, Cuthona, 127, 132, 167, 267
flemingii, Doris, 134
Flustrellidra, 136
folliculata, Elliptio, 301, 303, 305, 307, 311
fontinalis, Gyraulus, 19, 67, 68, 100
fragilis, Andonta, 299, 302, 305, 307, 311
francis, Filicrisia, 216, 217
frondosa, Amphitrite, 173
frondosa, Dendronotus, 175
frondosa, Doris, 173
frondosus, Dendronotus, 125, 131, 172-175, 269
fulgens, Cratena, 167
fulgens, Cuthona, 127, 132, 167, 268
fulgens, Trinchesia, 167
fulva, Dendrodoris, 170
fulva, Discodoris, 181
fulva, Doridopsis, 170
fulva, Doriopsilla, 170
fulva, Doriopsis, 169, 170
furcata, Sertularia, 184, 201
fusca, Acanthodoris, 136
fusca, Doris, 134, 196
fusca, Lamellidoris, 198
fusca, Onchidoris, 197, 198
fusca, Onchidorus, 197
fusca, Trapania, 214
fuscus, Antiopella, 146
fuscus, Janolus, 146, 147
fuscus, Onchidorus, 197
Gadus, 141
Galvina, 166, 185, 211
ganda, Doto, 183, 185
Gastropoda, 1-113
Gastrotricha, 289
gauda, Doto, 183
gelatinosa, Laomedia, 212
gelatinosum, Alcyonidium, 136
gellindra, Adocia, 180
Gellius, 157
geniculata, Laomedea, 176
Gersemia, 214
ghiselini, Hypselodoris, 192
gibberum, Holopedium, 93
gibbosa, Ancula, 143
gibbosa, Anodonta, 299, 301, 302, 311
gigantea, Tritonia, 213
giganteus, Dendronotus, 176
Gigartina, 190
gilberti, Duvaucelia, 220
gilberti, Tritonia, 220, 221
glabrata, Biomphalaria, 11, 14, 289-297
glacialis, Aplysilla, 153, 154
glauca, Chromodoris, 117, 192
globosus, Bulinus, 289
glorietta, Bougainvillia, 176, 212
Glossodoridiformia, 150, 151
Glossodoris, 156, 157, 191
gnupa, Polycera, 206
Goniodorididae, 119
Gonothyraea, 212
gracilis, Bowerbankia, 207
grandis, Triopha, 217, 218
grayi, Embletonia, 211
gredleri, Gyraulus, 46, 67, 72
gredleri, Planorbis, 45
gurneyi, Ptilosarcus, 150, 201, 214, 220, 222
Gyraulus, 1-113
Gyrorbis, 66, 67
Halgerda, 209
Halichondria, 146, 149, 154, 180, 182
Haliclona, 146, 157, 182, 192
Haliplanella, 169
Halla, 189
Hallaxa, 119-129, 188, 189, 271
Hancockia, 116-131, 189, 271
Hancockiidae, 120
heathi, Discodoris, 123, 131, 180, 181, 270
hebraicus, Gyraulus, 51, 52
hebraicus, Planorbis, 51
hedgpethi, Polycera, 121, 129, 205, 206, 274
helicinus, Margarites, 178
Helisoma, 81, 289, 296
herculea, Aeolidia, 140
hercules, Aeolidia, 140
Hermissenda, 146, 199-202
Hervia, 168, 201
Heterocope, 93
Heteroprocta, 119
hibernicum, Pisidum, 94
hiemantium, Gyraulus, 60, 61
hiemantium, Planorbis, 61
Higginsia, 154
hiltoni, Facelina, 202, 203
hiltoni, Phidiana, 126, 132, 202, 203, 273
Hincksina, 193
Hippeutis, 10, 11, 41, 97
hirsutis, Gyraulus, 33
hirsutum, Alcyonidium, 136
hispida, Flustrellidra, 136
hispidus, Planorbis, 27, 99, 100
Holohepatica, 119
Holopedium, 93
hombergi, Tritonia, 219
hopetonensis, Elliptio, 301, 303, 307, 311
Hopkinsia, 119, 121, 129, 190, 271
Hopkinsiinae, 119
hudsoni, Acanthodoris, 123-136, 261
hyalina, Celleporella, 179
hyalina, Embletonia, 211
hyaloderma, Zygerphe, 146
Hydractinia, 176, 187, 202, 208
Hydrallmania, 173
hyemalis, Clangula, 198
Hymenaeolis, 187
Hymendesmia, 143
INDEX 317
Hypselodoris, 117-129, 156, 159, 190-192, 272 leachii, Onchidoris, 196
hystericina, Lamellidoris, 198 leachii, Onchidorus, 196
hystericina, Onchidorus, 198 Leilla, 193
hystricina, Onchidoris, 124, 129, 198, 199, 273 lenina, Melibe, 194
lentiginosa, Ancula, 121, 129, 143, 263
Idulia, 184 leonina, Chioraera, 193, 194
ilocanum, Echinostoma, 2 leonina, Melibe, 125, 193, 194, 272
imbricatus, Planorbis, 42 leontina, Chioraera, 194
implicata, Anodonta, 299, 301, 305, 307, 311 Lepas, 188
incrustans, Halichondria, 149 lesliana, Aeolis, 138
incrustans, Myxilla, 154, 180, 182 Leucilla, 137
indivisa, Tubularia, 141, 176 leuckarti, Protohydra, 212
intermixtus, Gyraulus, 20 leuconotus, Eolis, 187
intermixtus, Planorbis, 84, 91 Leucosolenia, 138
intestinalis, Ciona, 201 leucostomus, Anisus, 92
iodinea, Aeolis, 159 Lialla, 193
iodinea, Coryphella, 126, 132, 159-162, 266 Ligumia, 301, 303, 311
iodinea, Flabellina, 159, 160 lillieborgii, Pisidium, 94
iodinea, Flabellinopsis, 159, 160 Limax, 138, 187, 213
iodinea, Phidiana, 159 limbaughi, Cadlina, 115, 124, 129, 153
iris, Dendronotus, 125, 131, 176, 269 lineatus, Draco, 95
Isoetes, 93 Lissodendoryx, 146
lithophoenix, Plocamia, 209
Janolus, 117, 146, 147 liturata, Doris, 196
japonicus, Choanomphalus, 67, 68, 71, 100 liturata, Lamellidoris, 197
Jorunna, 183 Lobelia, 93
Juanella, 154 Loligo, 201
longicauda, Aeolis, 188
karykina, Plocamia, 209 longicauda, Cratena, 188
kobelti, Acroloxus, 72 longicauda, Dolicheolis, 188
kollikeri, Renilla, 150 longicauda, Eolidia, 187
kya, Doto, 131, 184, 185, 271 longicauda, Fiona, 187
longicirrha, Cerberilla, 155
lacinulatus, Terpiges, 211 longissima, Laomedia, 212
lactea, Tritonia, 173 longissima, Obelia, 186
lacteus, Dendronotus, 174 Lophogorgia, 205, 222
Lacuna, 178 loveni, Gonothyraea, 212
lacustris, Cordylophora, 212 loveni, Laomedia, 212
lacustris, Isoetes, 93 luciae, Diadumene, 141
laevis, Cadlina, 152-154 luciae, Haliplanella, 169
laevis, Doris, 134 luetkeana, Nereocystis, 218
laevis, Gyraulus, 3-44, 72-103 lugubris, Dendrodoris, 169
laevis, Planorbis, 34 lutea, Acanthodoris, 123-133, 262
lagenifera, Plumularia, 187 luteolus, Dendronotus, 174
lagunae, Catriona, 168 luteomarginata, Cadlina, 133, 153, 154
lagunae, Cuthona, 126, 132, 168, 268 lychnidicus, Gyraulus, 19, 67, 68, 100
lagunae, Hervia, 168 Lymnaeidae, 296
lagunae, Trinchesia, 168
Laila, 119, 121, 129, 192, 272
Lamellidoris, 196-198 macedonicus, Acroloxus, 72
Laminaria, 165, 167, 168 macedonicus, Gyraulus, 20
Lamorbis, 42, 74-102 macedonicus, Gyrorbis, 66
Lampsilini, 299, 301, 305, 307, 311 macedonicus, Planorbis, 66, 91
Lampsilis, 299, 301-303, 307, 308, 311 macfarlandi, Chromodoris, 122, 129, 156, 157, 265
lanceolata, Elliptio, 301, 303, 305, 311 macfarlandi, Glossodoris, 156
Laomedea, 176 macfarlandi, Platydoris, 122, 131, 204, 274
Laomedia, 212 macginnitiei, Mycale, 146
larynx, Tubularia, 167, 176 Macrocystis, 158, 182, 190, 194, 218
Lateribranchia, 221 maculata, Triopha, 118-129, 216-218, 275
Lateribranchiaea, 221 maculigera, Thordisa, 212
Lateribranchiaia, 221 magnifica, Chromodoris, 156
laticeps, Cumanotus, 163 major, Campaspe, 174
laxa, Dendrobeania, 216, 217 *malayensis, Gyraulus, 1, 19, 61, *62-103
leachii, Doris, 196 mandibulata, Cellaria, 216
318 INDEX
mansoni, Schistosoma, 289-297
marcusi, Conualevia, 157
mareoticus, Planorbis, 52
Margarites, 178
Margaritifera, 299-301, 308, 311
margaritifera, Margaritifera, 299, 300, 311
Margaritiferidae, 298-311
marginata, Cadlina, 129, 153, 154, 265
marginata, Doris, 154
marginatum, Metridium, 141
marginatus, Metridium, 141
marina, Fiona, 188
marina, Tubularia, 167
marina, Zostera, 186, 194, 207
marinus, Limax, 187
Marisa, 289-297
mauritiana, Peltodoris, 181
mauritianus, Gyraulus, 55, 95
mediterranea, Embletonia, 212
mediterranea, Tenellia, 211
megas, Perigonimus, 212
Melanogrammus, 141
Melibe, 116, 120, 125, 193, 194, 272
membranacea, Membranipora, 205, 217
Membranipora, 136, 159, 182, 205, 217
Metarminacea, 120
Metridium, 141, 210
Mexichromis, 156, 157
misakiensis, Eubranchus, 127, 132, 185, 222, 271
modesta, Cadlina, 124, 129, 154, 265
modesta, Triopha, 216
modestus, Elminius, 198
mollis, Bugula, 216, 217
monodonta, Cumberlandia, 299, 301, 311
Montereina, 145
montereyensis, Archidoris, 124, 130, 148, 149, 180,
264
montereyensis, Doris, 148
morroensis, Emarcusia, 203
morroensis, Phidiana, 127, 132, 203, 273
mosslandica, Cerberilla, 129, 131, 155, 265
mucronata, Reginella, 199
multiformis, Planorbis, 75
multiformis, Poecilospira, 75
muricata, Onchidoris, 199
murrayana, Aeolis, 138
muscoides, Bougainvillia, 212
muscula, Rostanga, 209
Mycale, 146, 149
Mytilus, 141, 201
Myxilla, 146, 149, 154, 180, 182
nana, Cuthona, 164
nanaimoensis, Acanthodoris, 123, 133, 136, 262
nanus, Planorbis, 51
nasuta, Ligumia, 301, 303, 311
natalensis, Afrogyrorbis, 75
nautileus, Turbo, 42
Nautilus, 44, 100
Nematoda, 289
Nereocystis, 218
nigra, Phidiana, 202, 203
nigricans, Doris, 134
nigromaculata, Dendrodoris, 122, 170, 268
nigromaculata, Doridopsis, 171
nigromaculata, Doriopsilla, 171
nigromaculata, Doris, 171
nivosa, Pseudodiamesa, 93
nobilis, Anisodoris, 124-148, 263
nobilis, Archidoris, 145
nobilis, Fiona, 187
nobilis, Montereina, 145
nobilis, Oithona, 187
Nonsuctoria, 119
nov. zealandiae, Acanthodoris, 136
Nudibranchia, 114-276
nugator, Chirolophis, 202
numidicus, Gyraulus, 36
nuttingi, Leucilla, 137
nyctea, Archidoris, 148
Obelia, 167-187, 201, 212
obscura, Doridella, 182
obtusalis, Eolis, 138
obvelata, Doris, 196
occidentalis, Cabrilla, 217, 218
occidentalis, Crisia, 216, 217
occidentalis, Eubranchus, 186
occidentalis, Triopha, 122, 129, 217, 275
odhneri, Archidoris, 124, 131, 149, 264
odhneri, Austrodoris, 149
Oithona, 187
Okenia, 119, 121, 129, 195, 222, 272
Okeniinae, 119
olivacea, Galvina, 185
olivaceus, Eubranchus, 127, 132, 185, 186, 271
oliviae, Aeolidiella, 210
oliviae, Spurilla, 126, 131, 275
Onchidorididae, 119
Onchidoris, 119-129, 196-199, 273
Onchidorus, 196, 197
Oncidoris, 196, 272
opalescens, Aeolis, 199, 200
opalescens, Eolis, 200
opalescens, Flabellina, 199, 200
opalescens, Hermissenda, 200
Ophlitaspongia, 143, 209
Opisthobranchia, 118
orientalis, Eolidina, 210
originalis, Esperiopsis, 209
ornata, Acanthodoris, 135
osmundacea, Cystoseira, 187
Ostracoda, 289
ovata, Radix, 94
Pachycerianthus, 177
Pachygnatha, 120
pacifica, Aeolidia, 140
pacifica, Ancula, 121, 129, 144, 263
pacifica, Bugula, 205, 206
pacifica, Corambe, 129, 158, 266
pacifica, Fiona, 188
pacifica, Lamellidoris, 197
paladilhi, Planorbis, 42
pallasiana, Cryptosyla, 136
pallida, Acanthodoris, 136
pallida, Diaulula, 181
pallida, Embletonia. 211
pallida, Tenellia, 212
pallidus, Embletonia, 211
pallidus, Eubranchus, 211
palma, Corymorpha, 147
palmeri, Tritonia, 117
panicea, Halichondria, 146, 149, 154, 180, 182
papatasii, Phlebotomus, 90
papillata, Crimora, 163
papillosa, Aaeolidia, 140
papillosa, Aeolidia, 115, 127, 131, 138-141, 263
papillosa, Aeolidida, 140
papillosa, Aeolidiella, 140
papillosa, Aeolis, 138
papillosa, Aeolodia, 140
papillosa, Aerolidia, 140
papillosa, Doris, 138
papillosa, Eolidia, 138
papillosa, Eolis, 138
papillosa, Tritonia, 138
papillosus, Limax, 138
pappilosa, Aeolidia, 140
papposus, Crossaster, 222
paradoxus, Gyraulus, 67
Paramaecium, 24
Paratrichocladius, 93
Paresperella, 146
parvus, Gyraulus, 10-44, 71, 89, 99, 103
parvus, Planorbis, 38, 39, 99
patagonica, Phidiana, 199
paupera, Chromodoris, 156
pauxillus, Anisus, 91
peachii, Precuthona, 208
pealii, Loligo, 201
pedunculatus, Cereus, 141
pellucida, Melibe, 194
Peltodoris, 181
pennata, Ophlitaspongia, 143, 209
peregra, Radix, 94
Perigonimus, 212
permollis, Haliclona, 146, 182
perstriatulus, Choanomphalus, 69, 71
perstriatulus, Gyraulus, 71
petechialis, Chromodoris, 156
Petelodoris, 151
Petrosia, 182
Phacellophora, 201
Phanerobranchia, 119
Phidiana, 116-132, 199, 201-203, 273, 274
Phidianidae, 120
Phidianinae, 120
Phlebotomus, 90
Physa, 289, 296
Physastra, 79, 80
picta, Dirona, 125, 131, 178, 179, 269
pictus, Dendrophis, 90
pilosa, Acanthodoris, 123-136, 262
pilosa, Doris, 134-136
pilosa, Electra, 136
pinnata, Aeolis, 187
pinnata, Eolidia, 187
pinnata, Fiona, 126, 132, 187, 271
pinnata, Fione, 188
piscinarum, Gyraulus, 51-99
INDEX
319
piscinarum, Planorbis, 52
Pisidium, 94
piunca, Coryphella, 161
pizoni, Diplosoma, 160
plana, Cargoa, 195
plana, Okenia, 121, 129, 195, 222, 272
Planorbarius, 10
Planorbidae, 1-113
Planorbis, 9-102
planorbis, Planorbis, 91, 92, 98
Planorbula, 79
Platydorididae, 119
Platydoridinae, 119
Platydoris, 119, 122, 131, 204, 274
Pleurobemini, 301, 307, 311
Pleurophyllidea, 150
Pleurophyllidia, 149, 150
Pleuroprocta, 118, 120
plicata, Cratena, 188
Plocamia, 209
plumata, Eolis, 139
Plumularia, 172, 187
Podocoryne, 212
Poecilospira, 75
Polycera, 116-129, 204-206, 274
Polyceratidae, 119
polyoum, Alcyonidium, 136
Polypylyis, 79
Polysticta, 198
Pomacea, 289, 295, 296
Porella, 136
Porodoridacea, 120
Porostomata, 120
porterae, Chromodoris, 123, 130, 156, 157, 265
porterae, Glossodoris, 157
porterae, Hypselodoris, 157
porterae, Mexichromis, 157
Precuthona, 120, 129, 132, 207, 208, 274
Precuthoninae, 120
presbensis, Gyraulus, 20
presbensis, Planorbis, 91
Prianos, 146
pricei, Coryphella, 132, 161, 266
primata, Fiona, 188
primorjensis, Tritonia, 220, 221, 223
proclivis, Gyraulus, 95
prolifera, Epiactis, 141
Promenetus, 20, 79, 88
Protohydra, 212
Protozoa, 289
proxima, Adalaria, 199
Psammohydra, 212
Pseudelliptio, 301, 311
Pseudodiamesa, 93
psila, Paresperella, 146
Ptilosarcus, 150, 201, 214, 220, 222
pugnax, Phidiana, 202, 203
pulchella, Dendronotus, 174
pulchella, Tritonia, 173
pulchellus, Dendronotus, 174
pulchra, Rostanga, 116, 122, 129, 208, 209, 274
pulchra, Rostangia, 208
pumila, Dynamena, 176
punctilucens, Aegires, 137
320
punctuolata, Anisodoris, 145, 180
pupillus, Margarites, 178
purpurascens, Dendronotus, 175
purpurea, Acanthodoris, 135, 136
purpureus, Dendronotus, 174, 175
pusilla, Campaspe, 174
Pyganodon, 307
pyrifera, Macrocystis, 158, 182, 190, 194, 218
quadrangulata, Acanthodoris, 136
quadrangulata, Doris, 135
quadrilineata, Polycera, 204
quadrimaculata, Atagema, 151
radiata, Lampsilis, 299, 302, 305, 307, 311
Radix, 60, 82, 94
ramosa, Barentsia, 144
ramosum, Eudendrium, 160
Reginella, 199
Renilla, 150
reticulata, Doriopsilla, 170
reticulata, Doriopsis, 169
reticulata, Duvaucelia, 221
reticulata, Smittina, 136
reticulata, Tritonia, 221
reticulatus, Bulinus, 79
reynoldsii, Tritonia, 173, 174
reynoldsi, Tritonia, 173
rhodocera, Acanthodoris, 136
rhodoceras, Acanthodoris, 123, 128, 136, 262
riparius, Gyraulus, 3, 15, 16, 41-44, 79-101
riparius, Planorbis, 41, 42, 99
rocinela, Doris, 134
ronga, Catriona, 168
rosacea, Hopkinsia, 121, 129, 190, 271
rosea, Cuthona, 207
rosea, Eolis, 138
rossmaessleri, Gyraulus, 3, 14-52, 79-101
Rostanga, 116, 119, 129, 208, 209, 274
Rostangia, 208
Rostangidae, 119
Rotifera, 289
rowena, Doriopsilla, 171
rubiformis, Gersemia, 214
rufescens, Radix, 60
rufibranchialis, Coryphella, 159
rufus, Dendronotus, 175
rustya, Capellinia, 186
rustyus, Capellinia, 186
rustyus, Eubranchus, 127, 132, 186, 271
rutila, Cratena, 168
sabulicola, Coryphella, 160, 161
Sagartia, 141, 142
Sagartiogeton, 141
sandiegagenesis, Dilella, 181
sandiegensis, Actinocyclus, 181
sandiegensis, Dialula, 181
sandiegensis, Dialula, 180, 181
sandiegensis, Diaululua, 181
sandiegensis, Discodoris, 115, 123, 131, 181, 183,
270
sandiegensis, Doridopsis, 181
INDEX
sandiegensis, Doris, 181
sanguinea, Aldisa, 122, 129, 142, 143, 263
sanguinea, Asteronotus, 142
sanguinea, Doris, 142, 143
sanguineus, Asteronotus, 142
sarasinorum, Anisus, 91
sarasinorum, Planorbis, 91
sargassicola, Corambe, 158
Schistosoma, 289-297
Schizoporella, 136
schlosseri, Botryllus, 176
Sclerodoris, 119, 123, 131, 209, 275
scrippsiana, Triopha, 216
Scrupocellaria, 216, 217
Segmentina, 10, 97
Selva, 118
senile, Metridium, 141, 210
septemtrionalis, Duvaucelia, 220, 221
septemtrionalis, Tritonia, 221
serotina, Aeolidia, 139, 140
serrilamella, Membranipora, 158, 182
serrulata, Crisia, 216
Sertularella, 173
Sertularia, 176, 184, 201
shephardiana, Elliptio, 301, 303, 305, 307, 311
Shinanoeolis, 201, 202
similis, Doris, 134
singularis, Planorbis, 81
Smittina, 136
soemmerringii, Aeolidiella, 141
Solaster, 222
spadix, Cratena, 166
sparsa, Cadlina, 123, 129, 154, 265
sparsa, Juanella, 154
Sphaerostoma, 213, 219
sphyrodeta, Actinothoe, 141
spiniferum, Cauloramphus, 216, 217
spinosa, Elliptio, 301, 303, 307, 311
spirillus, Anisus, 92
spirillus, Gyraulus, 1, 10-33, 58-76, 102
spirillus, Planorbis, 58
splendida, Lampsilis, 299, 301, 302, 305, 307, 311
spongicola, Petelodoris, 151
Spurilla, 120, 126, 131, 209, 210, 275
Spurillidae, 120
Stagnicola, 82
stankovici, Gyraulus, 68-70, 72, 73, 100
stearnsi, Facelina, 203
stearnsi, Phidiana, 126, 132, 203, 274
steinbergae, Corambella, 182
steinbergae, Doridella, 129, 182, 270
stella, Tealiopsis, 141
stellata, Acanthodoris, 135
stellata, Doris, 134, 135
stelleri, Polysticta, 198
Stelletta, 192
Stiliger, 212
stipitata, Stylissa, 149
Stomphia, 141
strauchianus, Anisus, 93
stroemia, Verruca, 198
Strophitus, 299, 301, 302, 311
struthionides, Aglaophenia, 177, 184, 201
Stylissa, 149
suberea, Ficulina, 170
Suberites, 170
sublaevis, Doris, 134
subquadrata, Acanthodoris, 135
subquadrata, Doris, 134, 136
subramosus, Dendronotus, 125, 131, 177, 269
Suctoria, 119
sulcata, Anemonia, 141
sulphurea, Ancula, 145
sumatranus, Gyraulus, 95
swinhoei, Radix, 60
Syncoryne, 167, 176
Syringella, 149
INDEX 321
Tritonia, 116-138, 173, 174, 213-223, 276
Tritoniidae, 102, 220
Tritoniopsilla, 213
Tritoniopsis, 213
trochiformis, Gyraulus, 75
troglodytes, Sagartia, 141
tropicus, Bulinus, 10
tryoni, Chromodoris, 156
tuberculata, Archidoris, 148, 149
tuberculata, Doris, 148
tuberculata, Sclerodoris, 209
Tubularia, 115, 141, 159, 162, 164-167, 176, 201
Tubulipora, 199
Turbo, 42
Umbonula, 198
takanosimensis, Aeolidiella, 115, 126, 131, 141, 142 undata, Sagartiogeton, 141
takanosimensis, Eolidina, 141
tanya, Doris, 209
tanya, Sclerodoris, 123, 131, 209, 275
Tealia, 141
Tealiopsis, 141
Tedania, 149
Tenellia, 120, 126, 132, 211, 212, 222, 275
tenellus, Gyrautus, 27
teres, Lampsilis, 299, 303, 311
Tergipes, 211, 213
Terpiges, 211
terraesacrae, Gyraulus, 63, 81, 95
Tethyidae, 120
tetraquetra, Doris, 213
tetraquetra, Duvaucelia, 213
tetraquetra, Limax, 213
tetraquetra, Sphaerostoma, 213
tetraquetra, Tochuina, 124, 131, 213, 214, 275
tetraquetra, Tritonia, 213, 214
tetraquetra, Tritoniopsilla, 213
tetraquetra, Tritoniopsis, 213
Thaumatoporella, 179
Thecacera, 214
thermalis, Gyraulis, 34-36
thermalis, Planorbis, 34
Thordisa, 119, 123, 130, 142, 212, 213, 275
Thuiaria, 172
tigrina, Armina, 149
Tochuina, 120, 124, 131, 213, 214, 219, 275
tokyoensis, Anisus, 61
tokyoensis, Gyraulus, 3, 9, 22, 60-62, 82-103
tomentosa, Doris, 134
tomentosa, Jorunna, 183
tondanensis, Gyraulus, 94, 97
Torquis, 1, 39, 74-103
Trapania, 116, 119, 120, 129, 214, 275
trapezoides, Gyraulus, 67, 68, 100
Tricellaria, 216, 217
tricolor, Eubranchus, 185
tricolor, Polycera, 121, 129, 206, 274
tricuspidata, Sertularella, 173
trillineata, Coryphella, 126, 132, 161, 266
Trinchesia, 164, 167, 168
Triopha, 116-129, 215-218, 275
Triophidae, 119
Triophinae, 119
undulata, Alasmidonta, 299, 302, 311
undulata, Duvaucelia, 221
undulata, Sphaerostoma, 221
undulata, Tritonia, 221
undulatus, Strophitus, 299, 301, 302, 311
unicornis, Schizoporella, 136
Unionidae, 298-311
universitates, Chromodoris, 191
universitatis, Chromodoris, 191
universitatis, Glossodoris, 191
vancouverensis, Armina, 150
vancouverensis, Pleurophyllidia, 150
varians, Doto, 184
varians, Onchidoris, 199
velata, Hincksina, 193
Velella, 188
velella, Velella, 188
velox, Drepania, 214
velox, Thecacera, 214
velox, Trapania, 120, 129, 214, 275
ventilabrum, Eolis, 211
ventilabrum, Tenellia, 212
venustus, Dendronotus, 175
vermigera, Doris, 138
Verruca, 198
verrucosa, Doris, 183, 196
vicina, Chromodoris, 156
vidua, Dendrodoris, 171
vidua, Doridopsis, 170, 171
Villosa, 301, 303, 311
virens, Cratena, 168
virens, Cuthona, 127, 132, 168, 268
virens, Trinchesia, 168
Virgullaria, 220
viridis, Corynactis, 141
vortex, Anisus, 92, 94
vorticulus, Anisus, 84, 85, 92
vulgaris, Doris, 196
waccamawensis, Elliptio, 301, 311
wara, Doto, 183, 185
xanthogrammica, Anthopleura, 141
Zephyrinidae, 120
322 INDEX
zetlandica, Aldisa, 142 Zostera, 186, 194, 207
zetlandica, Eolida, 138 zosterae, Polycera, 121, 129, 205, 206, 274
zilchianus, Gyraulus, 64 Zygerphe, 146
IL. 24 1983
MALACOLOGIA
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MALACOLOGIA, VOL. 24
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A. CEDENO-LEON & J. D. THOMAS
The predatory behaviour of Marisa cornuarietis on eggs and neonates of
Biomphalaria glabrata, the snail host of Schistosoma mansoni ........... 289
Р. W. КАТ
Conchiolin layers among the Unionidae and Margaritiferidae (Bivalvia): micro-
structural characteristics and taxonomic implications ................... 298
G. R. MCDONALD
A review of the nudibranchs of the California coast .................... 114
C. MEIER-BROOK
Taxonomic studies on Gyraulus (Gastropoda: Planorbidae).............. 1
J. VKLAND
Factors regulating the distribution of fresh-water snails (Gastropoda) in
A A ge ee м oR WRC 277
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CONTENTS
C. MEIER-BROOK
Taxonomic studies on Gyraulus (Gastropoda: Planorbidae) .............. м
С. В. MCDONALD F
A review of the nudibranchs of the California coast .................... 114
J. OKLAND
Factors regulating the distribution of fresh-water snails (Gastropoda) in
Ме ER De e ee En
A. CEDENO-LEON & J. D. THOMAS
The predatory behaviour of Marisa cornuarietis on ‚eggs and neonates of
Biomphalaria ‚the snail post ERROR EN so AN 2
P. W. KAT | RR Na SE EE $
Conchiolin layers ong tbe Unionidae and Marat ide (Bivalvia): micro-
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INDEX TO VOL. 24, NOS. 1-22... 44. ut: EN, Weine nn ide tee DU LL ИИЙ
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