A Ху“ a at A Y in RENE LA ar en pia Ar ER VOL EU PEN DO Nc \ . O +. > PR EN: EEE o A PSE te > wi АХ ыы ye VS ARA x PET PER toe рам ро АНТИ RCE F es TR AA 43, uw | > ERA г ee O es LORS ne | PA LAB arret г perry AA MEL sur , PET pc AA rd = РО ды" + FEA ATA я PA 1 ARTS © ne mI ви. Е имели тв ИН APRA EE 91 HARVARD UNIVERSITY € Library of the | Museum of Comparative Zoology Per mas A u u 7 \ av = A Y р L Г : À de. Fi a BA E are á u es \ és и > р | $ — | me в $ Fr съ У qa EA A TEA L , 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 ‘рахе!елий, IE NE D PIN ЕЕ Я A AAA A CR A A A A AAA EEE _ 0+9 ИСТ $'0 + 95 еб G2 Le + $55 05—02 el ¿SY + NZ N “ЦО = GOS | 265 CAL O’EF SE LVN OL EG +692 6-12 Gil “INF ‘1}U99 “‘иолое + Отт ÿe- LL zo +80 01-90 LL zp ‘d 3х3} aas ‘uen ‘B}SUO + GL'O 60-90 80 6 0-/'0 v 5'8 01-2 v snuedu + 00 + SEO 7'0-5'0 20 +03 e 2-81 1 се + с 81 15-91 2 UM ‘Id “Wwssos + Lo+s0 90-r0 corr ОСТ pl La FEEL v1-8 СЕ ээе$|\ "WSSOJ + 80 +07 92-91 zo + SEL 91058 LL 176 = 1701 71-9 6 "pl39] 'snased + eorgı Senet eoFrt 0.260 St Gerla 26 Ss! “ING ‘1}u89 “snaJed + €07 91 MOL r0 +60 61-90 LL La + гб 1-9 2 ‘HEEH 'S/ineB] + vOF6L вое! cOFE£EL 7 on 0 BL LeFre 91-6 6 ‘Yan ‘SA8E/ eorgı Geral eoFrzt 81-80 er er Fgel Conk Le (с эае1) snqe ‘ty yds ‘DiS EX эбиел PISE эбиел u DIS TX эбиел u saloads did:ysd oney (шш) чзбие! ¡ezo 1 "uanıp `1$049 ‘ON иебло Asoyejndoo ajeyy ‘Aaupiy ayeydas = 14 ‘\ds ‘eis N pue edoung ui snjnesAdH jo влэоелецэ ‘ег 37191 SoOrZt oc a | ОО 81-60 67 9'6 + 991 0078 el “eels ‘1x8 0-91 AA ВОО 1 81780 er Sv + sick 0071 LE [2101 t 8l 81-91 vl GC v (aqiu189s1pui) == ‘MIIJBPUIM y Zar Op! 9! steil v SL! ec st y wıeyusNg A + 871 d (ejqiu189s1pui) — иэбицещиом + 0? 60 | 1 с цэщепу + eoFr6l So vl rorFret ЧЕ 9 (equuiaosipul) > aasıN t GL Al! 01 & 160 v 8 | neyosi4 t 02. +0° +05 AN yel € Br SE OI € 99SU8/I910 t el! € 0C+6! T2! & gasysunz ЕН Ур + 6`0 Ce (equuiaosipul) == aasuspog t GO gel Во КО FOL 81-80 pl 958 = 6 Lh 92% 81 риенее!$ ‘uds “qlo ‘18 ‘95 DIS EX abueı HISZERX эбиел u PIS EE X эбиел u Ayyeoo7 xıyeındoo esing dıd:ysd oney (ww) yyBua] ¡ejo | "U1aAIP ‘}S01d ‘ON ueBio Aıoyeindoa ajey RN ‘papnjoul эле (+961) HOOIG-ISI8N шо езеа 'sngje snineJsAH jo suebio элцопрол4ан ‘| 3198V1 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 - 4x yd: u г ) Er HE 944 grrr h + FARES Я TT I Y. Y az ve 1 z VU y гг HART & Fr] T 353127 022 ЕН RE + E 1777 Г LA 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 50 "P3Pnjox3 sjenpiaipul развелий г "paxejajun ı eorgı 8 1-60 80 +61 ОСИ! 7 ez + 9'61 61-21 Z ¡¿UOS-MIYSBS SO+61 О! vol? LEO el ez + 0'61 ЗЕЕ OL ¿UOS-NANSNBILIIO | “SISUSOÁ NO] 0 +91 Sel 90 +92 Дб 8 0'€ + 0'pL 61-6 8 uesuny tO+6!1 wey lt ro +07 Scart G Or! 91-81 v Buolsey eorFrgı (6 ey e0+ ге DCE Sl rzF0or 81-8 Sl биолабиоц rorgı dell £O +07 0'e-p"1 ee VEZF 161 lei pS иеме | 170 + 0°< Ой 70 +05 Чел | G 9 FOZ 02-91 G ¿¡OY9-NQOJON eorrı ie tae COFLL (СИЕ G gLFor 91-21 G ¿UOS-BUUO 'SnyuIds rorsgı ear €0+ $71 Sicalit el ¿LF 63 GUS PE 8 {Buoy Buoy ‘s/suauiyo КО OL A! €0 + gel girl 2 Ge + £73l 91-01 G ‚риенечу а | 21-07 61 22-1 9 Sad 2-02 ‚тру AE RAI Ser ОИ Гас 8 КЕ + eel Sı-21 L губуу + ‘INEW ‘зп/позпихалиоэ QL Ganar 8! Diesel p S Zt 61-561 p ¡BLpuexa|y e0+6l ВЕР #0 + 92 82-61 el УЕ 9'91 61-71 81 олеЭ ‘5поцовлеш Ss! Ост v2 EEO У Or 91-31 p JIMBqueÁIg ‘иплешо$!а rorrı e: 2-80 70791 GG: OL rı Free GI-LL 9 ‚елАцил + eonjwny ‘5по!елдац 90 +61 62-€'1 COF LS 92-91 81 G'e + L'EL 81-6 91 UBA] $ ‘зпоцелцапа E эбиел DIS Ex эбиел u EAS эбиел u Ayyeoo) ‘эшеи saisads ‘SiAOï4 did:ysd oney (ww) yyBua] je}O 1 уебло Aıoyeindoa ajey епэцлэлмр ‘}S01d ‘ON ‘EISY 3 pue © ui sninelÁo) jo sueB10 anıyonpolday ‘9 319V1 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 ‘7961 'ADIPUSQNH » “ıandun *y001g-1919/N 'Ep—9Z e "1961 '93PNH ‘08-61 2 "99/61 ‘HOOIG-ISI8W ‘16-61 ı (+) + ¿ yt „+ й пецашоиеоц + + (+) + (+) + (+) + (+) sE sninesAy a ES > > в snreydwo/yjeg + > > ES + snsiuy + (+) + + iL qe SIQJOUR|q juasald juasqe разеци! "био!э „eu juasaid juasqe diy ‘4 di ureBe juasaid juasqe -qou,, }UP SIP Jeau peonpai (eus 1npe) xuyeindoo esing ‘\djnos :¡nonay OL ads je1ued MOIIOH pajenys Buiuexoiu} di siuag LL 6 aod sıuad / 8 ¿ y+ é 6 ¿ ¿ sn¡eydwoueoya (+) + (+) + (+) + + (+) (+) + (+) + sn mello + + 5 et + snjeydwohyjeg + 7 7 г+ + + snsiuy + + + i+ . + SIGIOUE/4 ueße adeus ‘1puiA9 apim = MOJEU aS00| paoeds MO} ybiy Ayoyed = asnyip ayeinpun jyBiens “1IpuAo -qnjo Ajasojo ES 5 SuaJajJap SEA Jaqunu yeyuawbid suibiew ÁSupiy yyeauys siu б }уэшэбиелле "NP ‘}SOid эвчеи | -э4 jo adeys "NP “ISO Id e с 9 y Е A A A A A AA к A me "биилпээо Alaaeı—(+) 'биллпээо Auen6ai— + aqu) au} UIUJIM siajoeJeyo Ul эбиецо Алецоцтюла jo чоцоалр pasoddns :suuun¡o9 ul зубы O] Ya} WOI4 “3QUY-SIQIOUE/H ay} JO esauab ul $аазоелецо “/ FIGVL 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. LITERATURE CITED AGOCSI, P. 8 PINTER, L., 1971, Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 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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 в at en ра AN A. ten. FREE ; Е т ВАК ИЕ HR A IL (A 14 RT / < ES 3 / A 16 18 | Ay > a | Page = 7 А | | / \ 7 N N | / Vi Lo € | | BEN a |5 23 | [7 19 20 22 24 Ех. br. app. we: rid. LA 6 Cer. Lat. pro. a Rh. Ги Г Fr. veil A 4 aS 9 F Br. app. Pal. rid. Rh. / rl A $ >> e A RE Sp =" veil A 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 A Se = —= TH Seta 7! ga Senn >| == an N n) S = Wann avy i as | - Er 00 ws >| a NUL = C4 YI — 34 b a5 op Se en x c в N с е + | В 38 т | L в RENT ON BY De dl S JV | = LE — | ‹ 39 40 | a ZEN T ER 43 : EN) % =) Don ak 007: ST ys | ( WgefS h wt Ik 46 IN 42 — : 3 a с д | b FA b 45 И ) | [| SS GEES ea h i J 47 ДД) a ab de ¢ | x 5 Ga = С Die aan. И 2 $ a re e [NW = 0071 55 = С diet Le > \ 1) Fa Я | = | q) QA ES WII Jlösel АМИ le ЛМ | с E NS eT Е 59 3 5 С я b + u ? : с Br | mm N € R NS PEN AMI | FÜ 7 a Pr Bu SN Y gee ie N | me À 27 | ES We N a A rig oy СЕ $ ere 60 d я [ » I % T rn e i ca SN SN > ( \ ) Al Ñ | | AT dou > o A IN: sid e f 63 64 lv: 65 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 N * y $ - | CE | т mM — 2 =? a b В д CPS 9 b и e t 9 67 5 b с 68 66 д b I \ ( \ P Po I a Te 2 Kim a \) ots d a he Е f 69 д h | J 70 b c 7(\\ b = d A ==> =— Le = ( u \ Г JE T ab = eo oy L + c 4 с . ob . ES Hop d ge | d ES — / д = AA ae: == = I (| \\ | | I A WAS \) Y TT a + o с 7 76 abc di 8 f b d 75 d e A e gf У N AA f À. К \ vn Yh, % Г | JL N, \ / МА \ | 4 » || / y \ N AN NM Mr = QU | ( I { | | a | | bec Je + a bc, ry PE a ) (|| | A b \ sul e f 9 Cc d J ) 78 79 80 e фон 4 gn T oy | Se N | 82 fi M =" | h y y 84 a b | Nh 7 p\ a o | m У Y Kh, es 86 Е" 83 Li || AN - BEN he y > & = 7 С ААУ C3 et I er : d e у 88 А ab cd® lg h i j k CI — a b “e Set > 90 И 87 i ng 91 a a ус 7 : g h 2 b 93 b d e f с КАМ Am. А ИИ NE nu ul NA Ц } ШИ И МА \ Pere 77 wi" a ha м \ qu UV N y N ae Y AN Ил $ 7 4 N % и М N DT. guet un dl NL” N y - 95 M T и il F 7 97 Е АЙ / ve À V4 à Sy y J y Y 94 96 98 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 N р Ay ; y ", T ] À >, i ) | E | à LL e 3 | | ai IN 2 B Ze а b b = Y) AN b 99 a b 101 a 103 | 100 102 Por } T = N | 106 /\ u rm E см | vai b | a b | \ = b Il | Mi N в. 104 105 Avr du de: a E b 107 108 + An м e if ! h 7 : | UA A | m. y, A: x | A 5 en A y" —^ № г a TN te N а U | pn ty I SA / uy \\ - ; Vi \ N / EA \ \ now) Vile И 14 \ y un ir | УМ т NW y NY mi MT Тот И | M | MAN LU À l Е le 225 \ | EN N т EDS M A Ves Sl 117 | | fio| |1 LS U ee \ | ; o 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. REFERENCES CITED ABBOTT, R. T., 1974, American seashells, the marine Mollusca of the Atlantic and Pacific coasts of North America. Ed. 2, Van Nostrand Reinhold, New York, 633 p., 24 pl. ABE, T., 1964, Opisthobranchia of Toyama Bay and adjacent waters. Hokuryu-kan, Tokyo, ix + 99 p., 36 pl. м ABILDGAARD, Р. С., 1789, In MULLER, О. F., Zoologica Danica. Ed. 3, 3: 1-71, pl. 81-120. ABOUL-ELA, |. A., 1959, On the food of nudi- branchs. Biological Bulletin, 117: 439-442. ABRAHAM, Е. D. & WILLOWS, A. O. D., 1971, Plasticity of a fixed action pattern in the sea slug Tritonia diomedia. Communications in Behavior- al Biology, (A) 6: 271-280. ABRAHAM, P. 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Translated by Sophia Botcharskaya. Allen & Unwin, London, 955 p. 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 8 cz al 68 — Sy — 98 Erz 0c — eg Sr BE saisads ‘ON 001 0) vO' GG) 9 Bl 02 vO' 90 — Le OL 1N0[09 001 Ве ZO 0) = er — Sl LS OL вЕ— 6 alnyesadula | 001 co ol 6, OL LL JAS 20 = 8 элпзодха элем 001 95 ce LE: Ol Gm 065 2 un]e1sqns 00"! 95° cv 35 bz — йе = 9 иоцезэбэл-олэеи\ 001 6€' LL 1e 10 G иоцезэбал |211S8119 | 001 2 ce — co — Y Ábojoan 00 1 Goi LL € uolNeAgIZ 001 Or С HP 607 001 | на OL 6 8 L 9 G y $ с | 11909 ain] ainsodxe wunje1sqns иоцезэабэл uonejaban ABojoey uonera|3 Hp 507 Hd -eladua | элем -OJ9BN — 121459491 = ‘CO+ = N (HP.L=) SANV1 SSSNGHVH 1VLOL MOT = ae LL 0€ — es" 19 — 8, — 99 — 82 — pl 85° LL заюэ4$ ‘ON Se 001 10 e0' EG 03 8l vl 60° vo - Sl OL INO|OD о 001 8d 80' 9 rt = las 99 — Sl #0` 6 alnyesadua | 001 LE 0€' 8d 93 Sl io US 8 элпзоаха SALM 001 09 er LG Go by — ve — L wnyelsans 001 Ai 95 93 09 - д 9 иоцезэбэл-олоеи\ 00° gr 99' eu — Br G иоцезэбал [81159118 | 001 LE cL’ 9g — y Абоюэ9 001 Де FAQ) = € uolenelg 001 OL’ С Hp 507 001 | Hd OL 6 8 L 9 G v £ С | 119109 ain} einsodxe шпзедзапс uone}sBan иоцезэбэл Абооээ uoij2A8|3 Hp 607 Hd -eJadula | элем -OJ9BN — |¡eusala] ‘v6S = М SIVIT TIV ER ‘KEMION ила}5еэцпо$ WO} SAAB] Ul SIIBUS JO Sa/Dads jo 1эашпи pue siajaweJed aye] из} 10} зээщеш чоцее.люэ UOSIESd ‘| FIGVL 282 283 FRESH-WATER SNAILS IN NORWAY OL — v0 — 80 — le — ve — 1 — ce — эк Or’ ge’ LL y9}29-aUuI 1 00'L €0'- 70° co 92 8L' 02 70` 90 — 6l’— OL INO|OD 001 8c - 20° e0'- ev — EL — lg — OL €L'— 6 aınyeledweL 001 co cr OL OL’ EJE AS A) 8 элп5оаха элем 001 95° ze’ Le 91 co 02° — L wnjeusang 001 9€' er Sl ve — Le — 9 uone}sßan-o0eyy 001 6€' LL Le - 10 — G uonejoßen 12118119 | 00‘! 211 ge — do v Ábojoan 001 CoS LL € цоцела!3 00! Or’ с Hp 607 001 | Hd OL 6 8 L 9 G У e С | 1nN0[09 ainje ainsodxe шпуедзапс ичоцезэбэл чоцезэбэл Ábojoay) uonernaj3 Hp 607 Hd -Jadua | элем -OI9EW 1214594191 ‘Ov = М (HP.L=) SIMYI SSSNGYVH IVLOL MOI €0 — LE 9 — ov — es — Or — es — ve - LQ’ GY’ LL y9}29-OUUI | 001 10 — €0° Lo OC 8L' pl 60' v0 — GL'— OL INO|OD 00! 8c — 80 —- 91° — by — Le — 95 — Gi v0’ 6 aunyesoduue | 00! 21 0€' 82 92° St’ LE — 6L'— 8 einsodxe SALEM 00! 09' er LS ge’ by — ve — L wnjeusang 00 1 cs 99° 97 0S — ОИ = 9 цчоцезэбэл-о.эеи\ 00`1 gr 99° СИ = Ad G uone}aben 1211S8118 | 001 LG el — 9g — У Абоюэ9 001 1e— 20° e uonenglg 001 OL rd Hp 607 00 1 | Hd OL 6 8 й 9 G v e с | 10009 ainje ainsodxa uwnje11sqns uonejeBaa ичоцезэбэл Абооэю чоцелаз Hp 607 Hd -Jadua | элем -OJO0BIN — 1214591191 `6у5 = NASA VIN ‘AEMION uJ3]sesymos шод sexe] ul SIIEUS jo HOUEPUNGE yojeo-au ¡e30] pue $лэ}эшелеа aye] ua] 40} S9911IJ811 UONEISAOD UOSIBS] ‘с FIIIVL 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 fresh-water snails (Gastropoda) of Norway. Mal- acologia, 9: 143-151. @KLAND, J., 1979a, Distribution of environmental factors and fresh-water snails (Gastropoda) in 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 DRABLOS, D. & TOLLAN, A., eds., Ecological impact of acid precipitation. Proceedings of an international conference, Sandefjord, Norway, March 11-14, 1980. SNSF project, Oslo-As, Norway, p. 322-328. OKLAND, J. 8 VKLAND, К. A., 1979, Use of fresh- water littoral fauna for environmental monitor- ing: aspects related to studies of 1,000 lakes in Norway. In: The Use of Ecological Variables in Environmental Monitoring. The National Swed- ish Environment Protection Board, Report PM 1151, р. 168-183. OKLAND, J. 4 VKLAND, К. A., 1980, pH level and food organisms for fish: studies of 1,000 lakes in Norway. In DRABLOS, D. & TOLLAN, A., eds., Ecological impact of acid precipitation. Proceed- ings of an international conference, Sandefjord, Norway March 11-14, 1980. SNSF project, Oslo- Ás, Norway, p. 326-327. SEVALDRUD, |. H., MUNIZ, 1. P. 8 KALVENES, S., 1980, Loss of fish populations in southern Norway. Dynamics and magnitude of the prob- lem. In DRABLOS, D. 8 TOLLAN, A., eds., Eco- logical impact of acid precipitation. Proceedings of an international conference, Sandefjord, Nor- way, March 11-14, 1980. SNSF project, Oslo- Ás, Norway, p. 350-351. 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. LITERATURE CITED BOVBJERG, R. V., 1968, Responses to food in lymnaeid snails. Physiological Zoology, 41:412- 423. BROWN, D. S., 1980, Freshwater snails in Africa and their medical importance. Taylor and Fran- cis, London, 487 p. CEDENO-LEON, A., 1975, Interactions between Biomphalaria glabrata (Say) and Marisa corn- uarietis (L.) under laboratory conditions. Ph.D. thesis, University of Sussex, England. CHANG, J. T. & GELPERIN, A., 1978, Learned modification of a molluscan feeding resonse pro- duced in the isolated nervous system. Society for Neuroscience Abstracts, 4: 189. CHERNIN, E., MICHELSON, E. H. & AUGUSTINE, D. L., 1956, Studies on the biological control of schistosome-bearing snails, I. The control of Australorbis glabratus populations by the snail Marisa cornuarietis, under laboratory conditions. American Journal of Tropical Medicine and Hy- giene, 5: 297-307. MARISA PREYING ON BIOMPHALARIA 297 DAVIS, W. J. & GILLETTE, R., 1978, Neural cor- relate of behavioural plasticity in command neu- rones of Pleurobranchaea. Science, 199: 801- 804. DEMIAN, E. $. 8 KAMEL, Е. G., 1978, Displace- ment of Bulinus truncatus by Marisa cornuarie- tis under semi-environmental conditions in Egypt. Proceedings of the International Conference on Schistosomiasis, p. 731-739. DEMIAN, Е. $. 8 LUTFY, В. G., 1965, Predatory activity of Marisa cornuarietis against Biom- phalaria alexandrina under laboratory condi- tions. Annals of Tropical Medicine and Parasi- tology, 59: 337-339. DEMIAN, E. S. 8 LUTFY, R. G., 1966, Factors af- fecting the predation of Marisa cornuarietis on Bulinus (B.) truncatus, Biomphalaria alexandrina and Lymnaea caillaudi. Oikos, 17: 212-230. EISENBERG, R. M., 1966, The regulation of den- sity in a natural population of the pond snail, Lymnaea elodes. Ecology, 47: 889-906. EISENBERG, R. M., 1970, The role of food in the regulation of the pond snail Lymnaea elodes. Ecology, 51: 680-684. EL EMAM, M. A. & MADSEN, H., 1983, The effect of temperature, darkness, starvation and var- ious food types on the growth and reproduction of Helisoma duryi: (Wetherby), Biomphalaria alexandrina (Ehrenberg) and Bulinus truncatus (Audouin) (Pulmonata: Planorbidae). Hydrobio- logía, 88: 265-275. FERGUSON, F. F., 1977, The role of biological agents in the control of schistosome-bearing snails. U.S. Department of Health, Education and Welfare, Atlanta, Georgia. FRANDSEN, F. & MADSEN, H., 1979, A review of Helisoma duryi in biological control. Acta Tropi- ca, 36: 67-84. FRANK, G. H., 1963, Some factors affecting the fecundity of Biomphalaria pfeifferi (Krauss) in glass aquaria. Bulletin World Health Organiza- tion, 29: 531-537. HOLLING, C. S., 1959a, The components of pre- dation as revealed by a study of small mammal predation of the european pine sawfly. Canadian Entomology, 91: 293-320. HOLLING, C. S., 1959b, Some characteristics of simple types of predation and parasitism. Ca- nadian Entomology, 91: 385-398. HUBENDICK, B., 1966, Some aspects of vector snail control. Malacologia, 5: 31-32. JOBIN, W. R., 1970, Population dynamics of aquatic snails in three farm ponds of Puerto Rico. American Journal of Tropical Medicine and Hy- giene, 19: 1038-1048. MSANGI, A. S. & KIHAULE, P. M., 1972, Pros- pects of biological control of schistosomiasis in East Africa. In: ANDERSON, C. & KILAMA, W. L. (eds.), Parasitoses on man and animals in Af- rica. Nairobi, East African Literature Bureau, p. 439-447. NDIFON, С. T., 1979, Studies on the feeding biol- ogy, anatomical variations and ecology of vec- tors of schistosomiasis and other freshwater snails in South Western Nigeria. Ph.D. thesis, University of Ibadan, Nigeria. NDIFON, G. T. & UKOLI, F. M. A., 1980, Studies on the dietary habits of naturally occurring pop- ulations of Bulinus (P.) globosus (Pulmonata, Planorbidae): the snail host of Schistosoma hae- matobium. The 3rd European multicolloguium of Parasitology, Cambridge, England, p. 90. OLIVER-GONZALEZ, J., BANMAN, P. M. & BE- NENSON, A. S., 1956, Effect of the snail Marisa cornuarietis on Australorbis glabratus in natural bodies of water in Puerto Rico. American Jour- nal of Tropical Medicine and Hygiene, 5: 290- 296. PAULINYI, H. М. & PAULINI, E., 1972, Laboratory observations on the biological control of Biom- phalaria glabrata by a species of Pomacea (Am- pullariidae). Bulletin of World Health Organiza- tion, 46: 243-247. SOLOMON, M. E., 1949, The natural control of animal populations. Journal of Animal Ecology, 18—35. THOMAS, J. D., 1973, Schistosomiasis and the control of molluscan hosts of human schisto- somes with particular reference to possible self- regulatory mechanisms. Advances in Parasitol- ogy, 11: 307-394. Academic Press, London and New York. THOMAS, J. D., GOLDSWORTHY, С. J. & BEN- JAMIN, M., 1974, Chemical conditioning of the environment by the freshwater pulmonate snail Biomphalaria, and its effect on growth and na- tality rates. Journal of Zoology, London, 172: 443-467. THOMAS, J. D., LOUGH, A. S. 8 LODGE, R. W., 1975, The chemical ecology of Biomphalaria gla- 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. REFERENCES CITED AVISE, J. C., 1976, Genetic differentiation during the speciation process. In AYALA, F. J. (ed.), KAT Molecular Evolution. Massachusetts, Sinauer Press, p. 106-122. AVISE, J. C., SMITH, J. J. 8 AYALA, Е. 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D., 1981, Interpopulation variation in calcareous and proteinaceous shell compo- nents in the stream limpet, Ferrissia rivularis. Malacologia, 20: 255-266. SARICH, V. M., 1977, Rates, sample sizes, and the neutrality hypothesis for electrophoresis in evolutionary studies. Nature, 265: 24-28. TEVESZ, M. J. 8 CARTER, J. C., 1980, Environ- mental relationships of shell form and structure of Unionacean bivalves. In RHOADS, D. C. 4 LUTZ, R. A. (EDS.), Skeletal Growth of Aquatic Organisms. Plenum Press, New York, p. 295- 322. TOLSTIKOVA, N. V., 1974, Microstructural char- acteristics of freshwater bivalves (Unionidae). Paleontological Journal [translation of Paleon- tologicheskii Zhurnal], 8: 55-60. 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 ernational Journal of Malacology Revista Internacional de Malacologia Journal International de Malacologie Международный Журнал Малакологии Internationale Malakologische Zeitschrift Publication date Vol. 23, No. 2—28 February 1983 E. 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