7 PROCEEDI OF THE CALIFORNIA ACADEMY OF SCIENCES (Fourth Series) CALIFORNIA ACADEMY o& SCIENCES | FOUNDED 1853 SAN FRANCISCO, CALIFORNIA Copyright © 2005 by the California Academy of Sciences All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without permission in writing from the publisher. SCIENTIFIC PUBLICATIONS Alan E. Leviton, Ph.D., Editor Gary C. Williams, Ph.D., Associate Editor Michele L. Aldrich, Ph.D., Consulting Editor COVER IMAGE The Staff of the California Academy of Sciences November 1923 Photograph of the professional staff of the California Academy of Sciences taken on the steps of North American Hall by H.W. Steward. 7op row (left to right): Edward P. van Duzee, Curator, Entomology; Frank R. Cole, Curator, Dipterology; James O. Martin, Assistant, Entomology; Frank Tose, Chief Taxidermist (Exhibits); Merle C. Israelsky, Assistant Curator, Invertebrate Paleontology. Middle row (left to right): Joseph R. Slevin, Assistant Curator, Herpetology; Kate E. Phelps, Assistant, Botany; Constance W. Campbell, Office Assistant; Mary E. McLellan, Assistant Curator, Ornithology and Mammalogy; Joseph Mailliard, Curator, Ornithology and Mammalogy. Front row (left to right): Winthrop W. Sargeant, Secretary to Board of Trustees; Barton Warren Evermann, Director and Executive Curator (Ichthyologist); Susie (Susan) M. Peers, Secretary to the Director; Alice Eastwood, Curator, Botany. Not in photograph: Frank M. Anderson, Honorary Curator, Invertebrate Paleontology; Roy E. Dickerson, Honorary Curator, Invertebrate Paleontology; Walter K. Fisher, Curator, Invertebrate Zoology; Edward W. Gifford, Assistant, Ornithology and Mammalogy; Harvey M. Hall, Librarian; G Dallas Hanna, Curator, Invertebrate Paleontology; Alvin Seale, Superintendent, Steinhart Aquarium; John Van Denburgh, Curator, Herpetology. Photo from the Archives/Special Collections, California Academy of Sciences. ISSN 0068-547X The Proceedings of the California rteademy of Secenceé accepts manuscripts for publication in the Natural Sciences that stress systematics and evolutionary biology and paleontology, biogeography, and biodiversity, also relat- ed areas in the Earth Sciences, such as biostratigraphy, regional and global tectonics as they relate to biogeography, and paleoclimatology, and topics in astrobiology, anthropology, as well as the history of science as they relate to insti- tutions of natural history, to individuals, and to activities, such as expeditions and explorations, in the natural sciences. Priority consideration will be given to manuscripts submitted by Academy staff and others who are affiliated with the research programs of the institution and/or manuscripts that include reference in part or in whole to Academy research collections or to Academy-sponsored expeditions. Others who do not meet the above criteria may submit manuscripts for consideration for publication on a space-available basis. Manuscripts accepted for publication are subject to page charges; charges may be waived on a case-by-case basis. Note: All manuscripts submitted for publication in any of the Academy's scientific publication series (Proceedings, Occasional Papers, Memoirs) are subject to peer review. Peer review includes both internal and exter- nal review, internal review by at least one Academy scientist whose interests parallel those of the submission, and external review, ordinarily by two individuals who are recognized experts in the field. Published by the California Academy of Sciences 875 Howard Street, San Francisco, California 94103 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Volume 56, No. 18, pp. 189-199, 5 figs. August 26, 2005 Two New Species of Porostome Nudibranchs (Family Dendrodorididae) from the Coasts of California (USA) and Baja California (Mexico) Sandra V. Millen ! and Hans Bertsch 2-3 1 Department of Zoology, University of British Columbia, 6270 University Blvd., Vancouver, B.C., Canada, V6T 124; Email: millen@zoology.ubc.ca; 2? 192 Imperial Beach Bivd., #4, Imperial Beach,CA 91932, USA; Email: hansmarvida@sbcglobal.net. A new species of porostome nudibranch, Dendrodoris stohleri, with a yellow to cream body color and irregular black spots, is described from the northern Gulf of California, Mexico. Its discrete and intense spotting, which does not appear on the sole of the foot, is distinctive. A second species, Dendrodoris behrensi, is described from specimens collected from southern California and the Pacific side of the north- ern Baja California peninsula, Mexico. It is white with a smooth, elongate body and large brown irregularly distributed spots and pale rhinophores and gills. KEY worbDs: Opisthobranchia, Porostomata, Dendrodoris, new nudibranch species. A number of recent papers have enlarged our knowledge of the anatomical (Valdés et al. 1996; Brodie et al. 1997) and histological (Wagele et al. 1999; Brodie 2001) aspects of the Family Den- drodorididae. Despite these studies, there remains a dichotomy in the terms assigned to the anatom- ical parts of these animals. In this paper, we have followed recent terminology with the more tra- ditional terms given in brackets. Investigations have also been made to clarify either by molecular (Thollesson 2000; Wollscheid-Lengeling et al. 2001; Valdés 2003) or cladistic means (Valdés and Gosliner 1999; Valdés 2002) the position of the family in relation to other nudibranchs. It is cur- rently believed that the family Dendrodorididae is nested within the cryptobranch dorids (Valdés and Gosliner 1999; Wollscheid-Lengeling et al. 2001; Valdés 2002, 2003) and there is support for the monophyly of the Porostomata with the radula-less dorids Dendrodorididae and Phyllidiidae forming sister clades (Valdés and Gosliner 1999; Thollesson 2000; Valdés 2002, 2003). In the family Dendrodorididae, nudibranchs lack a radula and are often variable in color, which makes them difficult to identify. There are seven species in two genera, Doriopsilla and Dendro- doris that are currently recognized as occurring along the Pacific coast of North America. At one time, the two genera were merged (Steinberg 1961), but they are now separated by a large number of morphological features, some recognized earlier (Eliot 1906; Pruvot-Fol 1930, 1954) and others more recently (see Valdés and Gosliner 1999; Gosliner et al. 1999; Brodie 2001). The genus Doriopsilla is characterized by having synapomorphies of an eccentric anus (to the left), a flat, non- tubular prostate gland, and buccal ganglia that are dorsal to the esophagus but close to the central nerve ring. Like many cryptobranch dorid nudibranchs, they usually have small rounded tubercles and spicules on the notum, one duct to their bursa copulatrix and eversible spines on the penis. Five 3 Research Associate, California Academy of Sciences. 189 190 PROCEEDINGS OF THE CALIFORNIA ACADEY OF SCIENCES Fourth Series, Volume 56, No. 18 species along the Pacific coast of North America belong to this genus, Doriopsilla albopunctata (Cooper, 1863), D. gemela Gosliner et al., 1999, D. janaina Marcus and Marcus, 1967, D. nigro- maculata (Cockerell in Cockerell and Eliot, 1905) (= D. rowena Marcus and Marcus, 1967, accord- ing to Valdés and Behrens 1998; Valdés and Ortea 1998) and D. spaldingi Valdés and Behrens, 1998. It has been postulated that D. janaina is possibly a junior synonym of D. areolata Bergh, 1880 (see Valdés and Ortea 1997) although Valdés and Gosliner (1999) retain them separately. The genus Dendrodoris is characterized by synapomorphies of a soft body, lacking spicules even when tubercles are present, and a posterior mouth, located within a deep notch of the foot. Internally, there are paired ptyaline (oral) glands with a muscular duct and small esophageal glands. The central nervous system usually has ganglionic tubercles, and the buccal ganglia are far poste- rior on the esophagus, just in front of the esophageal (salivary) glands. The reproductive system usually has two ducts to the bursa copulatrix and the spines on the vas deferens usually have long bases (see Valdés and Gosliner 1999). There is a common but problematic species in this genus, commonly known as Dendrodoris krebsii (Mérch, 1863), found along the Pacific coast from the Gulf of California to Panama and the Galapagos that has recently been assigned to the Indo-Pacific species D. fumata Riippell and Leuckart, 1930 (Behrens 2004). However, a reassessment of D. fumata suggests that the black color variety with a red rim is in fact a separate species, Dendrodoris arborescens (Collingwood, 1881) based on differing larval forms (Brodie and Calado, in press). Since on the Pacific coast, specimens from the outer coast of Baja California and the Gulf of California are black with a red rim, and specimens further south, in Jalisco, Mexico and the Galapagos are variable from grey to red, further investigation must be made before settling upon a name for the Pacific northeast species. Recently described by Behrens and Valdés (2004) is a deep- water California species, D. azineae, which has a white body and dark purple-tipped gills and rhinophores. Two undescribed species belonging to the genus Dendrodoris are described in this paper, one with a yellow or cream body and small black spots, from the Gulf of California, and the other with a white body and large brown blotches from the Pacific coast of southern California and Baja California. Subclass Opisthobranchia Order Nudibranchia Porostomata Bergh, 1891 Family Dendrodorididae O’ Donoghue, 1924 Genus Dendrodoris Ehrenberg, 1831 Type Species: Dendrodoris lugubris Ehrenberg, 1831 [by subsequent designation by Gray, 1847]. Dendrodoris stohleri Millen and Bertsch, sp. nov. Figs. 1A—B, 2A, 3, 4 A-C. Dendrodoris sp. Bertsch 1991:18. ETYMOLOGY.— This species is named in honor of our colleague, mentor and friend, Dr. Rudolf Stohler (1901—2000), for his numerous contributions to malacology. As founding editor of The Veliger, he supported and encouraged many young students (Bertsch 2000). We remember his eccentricities and his graciousness — a true gentleman and scholar! MATERIAL EXAMINED.— Ho .otyre: CASIZ 166884, 10 mm. preserved length, 2-3 m. depth, Herra- dura, Bahia de los Angeles, Baja California, Mexico, 18 February 1999, S. Millen. PARATYPES: eight speci- mens, CASIZ 166887, intertidal, Isla Coronado, 27 April 1986, H. Bertsch, 1 dissected. Two specimens, CASIZ 069301, Punta la Gringa, Bahia de los Angeles, 27 April 1986, H. Bertsch, one dissected. Four spec- MILLEN AND BERTSCH: NEW SPECIES OF POROSTOME NUDIBRANCHS 19] imens, CASIZ 073507, Puerto Don Juan, Bahia de los Angeles, Baja California, Mexico, 5 October 1984, Gosliner, Bertsch and Ferreira. Two specimens, CASIZ 73505, 10-12 meters, Isla Coronado, Bahia de los Angeles, Baja California, Mexico, 4 October 1984, R. Van Syoe and D. Mulliner. Three specimens, CASIZ 166885, 2-3 m. depth, Herradura, Bahia de los Angeles, Mexico, 18 February 1999, S. Millen. One specimen, CASIZ 166886, 8 m., Cuevitas, Bahia de los Angeles, Mexico, 19 February 1999, S. Millen, dissected. EXTERNAL MORPHOLOGY.— This species has a low, elongate-oval body with a moderately sized, slightly crenulate margin (Figs. 1 A—B). The maximum size of a preserved animal measured 24 mm with a mantle margin 3 mm wide. The notum is smooth. There are a few fine striations in the mantle margin, approximately 3 per mm, dividing towards the edge. The straight rhinophores have 8—10 lamellae sloping posteriorly from an anterior ridge, to meet posteriorly in a V. The tip is elevated and flattened. The rhinophore opening is raised slightly and smooth. The low branchial pocket is smooth. There are 6—7 posteriorly directed tripinnate gills. Ventrally the foot is bilabiate as far posterior as the gonopore with both the labium and foot notched in the centre where the pore- like mouth opening is located. There are two small, triangular, unnotched tentacles fused to the upper labium of the foot. The foot has a small flange and a bluntly rounded posterior, which trails slightly when the animal is crawling. The ground color is pale cream to dark golden yellow. Small irregular black spots are scattered on the dorsum but rarely on the mantle margin. The rhinophores are slightly darker than the ground color and may have a few black spots. The gills are the same as the ground color and may also have a few spots. The ventral surface is the same as the ground color. A few animals had one or two spots on the sides, but never on the foot or the underside of the mantle. ANATOMY.— The notum is thick and soft with many lacunae just above the basal layer. It is strengthened by vertical connective tissue strands, but has no spicules. The viscera are within a col- orless envelope. The digestive tract is illustrated in Figure 2 A. The buccal bulb (oral tube) is mus- FiGURE 1. Dendrodoris stohleri sp. nov. (166885). A. Dark color form. B. Light color form. Both photos by Mike Miller. Specimens from Bahia de los Angeles, Mexico, Feb. 1999. C. Dendrodoris behrensi sp. nov. (082088). Photo by Mark Chamberlain, specimen from San Clemente Is., California, USA, Feb., 1988. D. Dendrodoris behrensi sp. nov. (070838). Photo by Gary McDonald. Specimen from Malibu, California, USA, Oct. 1971. 192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 18 cular, short and wide. Posteriorly it 1s attached to the body wall by lateral muscle bundles, three on each side. Ventrally, the long duct from the bilobed ptyaline (oral) glands enters the buccal bulb at its junction with the pharynx. The ptyaline duct widens and branches into two before it enters each irregularly rounded, somewhat flattened, light colored gland. The narrow, tubular, pharynx is high- ly muscular. It passes through the nerve ring and bends, to the left and right in a reverse S-shape in contracted specimens, in a C-shape in relaxed specimens. It constricts at its junction with the esophagus. Just anterior to this junction, on the dorsal surface, lie two round buccal ganglia. Posterior to this junction, the esophagus has two lateral, oval, esophageal (salivary) glands. Next to each gland are two slender retractor muscles that run half way to the first bend of the esophagus and insert on the esophagus. At this point, the esophagus widens to at least twice the diameter of the pharynx, becomes glandular and is capable of distending when filled with food. In its distend- ed state, it is smooth. When contracted its surface is somewhat irregular. The esophagus bends in an inverted S-shape to the left, then right, back to the midline. The esophagus is over twice as long as the pharynx (2.4:1). Posteriorly there is an internal, muscular sphincter controlling food flow into a thinner walled stomach, which can be tubular and the same diameter as the esophagus or expanded like a balloon when full of food. The remainder of the stomach is embedded in the con- fluent digestive glands. The intestine emerges near the centre of the digestive glands and turns sharply to the right. There is a small, round, anterior pyloric gland (caecum). The intestine lies over the digestive glands and under the renal-pericardial complex. It narrows to the anus emptying on a small, smooth, medial papilla, which is posteriorly placed in the gill circlet. The heart (Fig. 2A) forms a broad triangular auricle with a large anterior ventricle. A promi- nent renal syrinx, on the posteri- or right side, drains the pericar- dial sac. A medial aorta runs anteriorly from the ventricle to the region of the central nervous system. Anterior to the digestive gland, to the left of the aorta, is a moderately large, flat, irregular blood gland, which lies within the bend of the esophagus and posterior to the central nervous system. The central nervous system (Fig. 2A) has fused cerebro-pleu- ral ganglia. These ganglia are separated from each other by a very slight mid-line depression. FIGURE 2. Dendrodoris stohleri sp. nov. (166886). A. Digestive tract They have large nerve cells, drawn by camera lucida. Scale bar = 1 mm. B. Dendrodoris behrensi sp. nov. which nevertheless form a_ (!7!659). Digestive tract drawn by camera lucida. Scale bar = 1 mm. Key: ao=aorta, an=anus, bb=buccal bulb, bg=blood gland, cns = central nervous : system, dg=digestive gland, eg=esophageal gland, es=esophagus, hg=her- The stout rhinophoral N€lVES maphroditic gland, ht=heart, i=intestine, ph=pharynx, ptg=ptyaline gland, each have a slightly larger gan- _ pyg=pyloric gland, rs=renal sac, st=stomach. smooth, non-tuberculate surface. MILLEN AND BERTSCH: NEW SPECIES OF POROSTOME NUDIBRANCHS 193 glion at their bases. The eyes are large, almost sessile, with clear lenses. The small otocysts with otoconia are located just posterior and ventral to the eyes. The cerebro-buccal connectives are long and separate, the round buccal ganglia lie dorso-laterally on the posterior portion of the pharynx. There is a narrow, posterior, visceral commissure.The smaller, round pedal ganglia are ventral to the cerebral part of the cerebro-pleurals and the pharynx. They are connected by a short, broad pedal commissure and a narrow- am nye er, posterior, parapedal commis- sure. The large hermaphroditic gland (ovotestis) is separate from the digestive glands and anterior to them. The reproductive system (Fig. 3) has a long pre-ampullary duct that widens into a tubular ampulla with one loop. The short post-ampulla duct bifurcates into a long oviduct and the prostatic portion of the vas deferens. The prostate is wide and long. It loops back on itself, completely enveloping the bursa copulatrix, narrows and continues as a high- ly convoluted tubular duct. The FIGURE 3. Dendrodoris stohleri sp. nov. (166887). Reproductive system narrower muscular. deferent 4rawn by camera lucida. Scale bar= 1 mm. Key: am=ampulla, be=bursa cop- ulatrix, dd=deferent duct, fgm=female gland mass, ga=genital atrium, (ejaculatory) duct of the Vas def- nd=nidamental duct, od=oviduct, p=penis, pr=prostate, ud=uterine duct, erens is coiled distally, widening — yo~yagina. at the penial sac before it termi- nates in the common genital atrium. The penis is armed with many rows of large, almost straight chitinous spines from 18—28 um in height with small bases (Figs. 4A—C). There are up to 22 spines in each row. The common atrium is elongate and plicate internally. The long, tubular vaginal duct is internally villous and wider distally. It narrows gradually and convolutes slightly before opening into the large, rounded bursa copulatrix. A separate, long, narrow uterine duct connects the bursa copulatrix to the female gland mass close to the entrance of the oviduct. A short, wide duct from the small, oval seminal receptacle joins the uterine duct near its entrance to the gland mass. The bursa copulatrix is approximately 4.5 times larger than the seminal receptacle. The female gland mass consists of an anterior, granular, yellow portion dorsal to the entrances of the oviduct and uterine ducts and a highly convoluted, firm, white mucus gland with a slightly differentiated lobe at the proximal end of the long nidamental duct. The nidamental duct adheres closely to the atrium and opens posterior to it. There is no vestibular gland. A protruding flap separates the genital open- ings. They are located high on the right side of the animal, one quarter of the way back from the anterior mantle margin. NATURAL HISTORY.— The known range of Dendrodoris stohleri is within Bahia de los Ange- les (29°0'N, 113°30'W), in the Gulf of California, Mexico. It is restricted to the islands and outer shore locations. This species occurs extremely rarely at the frequently sampled and readily acces- sible shore sites (within 10 km of the town of Bahia de los Angeles) Cuevitas and Punta la Gringa. During the 10-year period from 1992-200I, one of us (HB) regularly surveyed the opisthobranch fauna of these sites. Of 6,095 specimens observed, only | was D. stohleri. However, over 30 spec- 194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 18 Penial spines from middle of penis. Scale bar = 30 um. C. Penial spines from proximal end of penis Scale bar = 30 pm. D. D. behrensi sp. nov. (171658). Penial spines from distal end of penis. Scale bar = 10 um. imens have been observed by HB at the islands and reefs between Puerto Don Juan, Isla Calabeza, Isla Coronado, and at the intertidal zone of the “‘old turtle farm” at Isla Coronado. This distributional pattern within Bahia de los Angeles clearly illustrates the 2 major opistho- branch communities within this bay as reported by Bertsch et al. 1998 (see their Fig. 1, map show- ing the different regions). This species has been found in February, April, October and November, with mature animals in April. They are found intertidally and subtidally under rocks, to a depth of 12 m. Dendrodoris stohleri appears to feed on an encrusting yellow demosponge. Discussion.— Dendrodoris stohleri can be distinguished from all others in the genus by its smooth body with distinct black spots, not blotches, on a yellow or cream background, light gills and rhinophores and no dark spots on the foot. Dendrodoris sadoensis Baba, 1993 is dull yellow with larger black blotches and black spots are also found on the underside. Internal differences are that the penis is unarmed, the bursae are close to each other in size, the proximal prostate is not enlarged and the vagina is threadlike. The reproductive system of D. stohleri lacks a vestibular gland and has a wide, massive prostate followed by a convoluted narrower prostatic section, then a shorter, muscular region. The bursa copulatrix is unusually large compared to the seminal receptacle. In these respects D. stoh- MILLEN AND BERTSCH: NEW SPECIES OF POROSTOME NUDIBRANCHS 195 leri is similar to D. tuberculosa Quoy and Gaimard, 1832 and D. limbata (Cuvier, 1804). It can be clearly distinguished externally from D. tuberculosa by its smaller size, higher oval shape, smooth dorsum and yellow or cream ground color. Internal differences are that D. tuberculosa has irregu- lar oral glands, an interdigitated hermaphroditic gland, no distal post-ampullar duct, a more tubu- lar prostate and longer bases on the penial hooks. Light colored tentacles and rhinophores and smaller spots, never ventral, distinguish D. stoh- leri externally from the yellow and spotted variety of D. Jimbata, which is usually dark brown with a yellow rim, although internally they are similar. A cladistic analysis, which includes 9 species in the genus Dendrodoris separates these 9 species by 13 characters (Valdés and Gosliner 1999). Using the Dendrodoris portion of this tree, rooted on D. coronata, results in the same topography as presented in Figure 23 in Valdés and Gosliner (1999:353). The addition of D. stohleri indicates it is a sister species to D. limbata. When an allzero ancestor is used as the outgroup, the topogra- phy changes slightly. Dendrodoris coronata remains basal and D. /imbata once more pairs with D. stohleri followed by D. grandiflora and D. fumata which are basal to the remaining species. In common, D. stohleri and D. limbata have a flattened proximal portion of the prostate and penial spines, which have short bases. A distal post-ampullary duct, present in D. stohleri, is absent in D. limbata. Dendrodoris behrensi Millen and Bertsch, sp. nov. Figs. 1C—D, 2B, 4D, 5. Dendrodoris sp. Lee and Brophy, 1969:220; Behrens, 1980:101, fig. 141. Dendrodoris species b: McDonald and Nybakken, 1981:54—S5, fig. 58; McDonald, 1983:171. Dendrodoris sp. 3: Behrens, 1991:72, fig. 132. Dendrodoris sp. 4: Rudman, 2003. Sea Slug Forum fact sheet. ETyMOLOGy.— This species is named for our friend and colleague David W. Behrens, whose books, articles and photographs have significantly contributed to our knowledge of opisthobranchs. MATERIAL EXAMINED.— Ho.ortyPe: 16 mm. preserved length, CASIZ 069303, Pyramid Point, San Clemente Is., intertidal, 14 Feb 1988, R. Van Syoc. PARATYPES: one specimen, CASIZ 071367, Castle Rock, San Miguel Island, 25—40 ft., 24 October 1986, R. Van Syoc. Two specimens, CASIZ 171658, Pyramid Point, San Clemente Is., intertidal, 14 February 1988, R. Van Syoc, one partially dissected. One specimen, CASIZ 068309, Pt. Pinos, Pacific Grove, July 1932, GE. MacGinitie, MacFarland collection. One specimen, dissect- ed, CASIZ 69299, Abalone cove, Palos Verdes, under rock, 15 ft., 29 July 1982, P.A. Haaker. One specimen, CASIZ 069300, Jim Gatewood, California, no locality. One specimen, CASIZ 070838, Dume Rocks, Pt. Dume, Malibu, 9 Oct 1971, S. Anderson, McDonald collection. One specimen, CASIZ 071019, Santa Monica Bay, 25 fathoms, 7 July 1971, S. Anderson, McDonald collection. One specimen. CASIZ 082088, n. end of West cove, San Clemente Is., intertidal, 17 February 1988, R. Van Syoc. One specimen, CASIZ 068310, Pacific Grove, large tide pool, 27 July 1929, F.M. MacFarland. One specimen, CASIZ 070836, dissected, 10 feet, California, no locality, S. Anderson, 20 October 1971, McDonald collection. One specimen, CASIZ 072006, '2 m s.w. of Pescacero Pt., the Pinnacles, Monterey county, 17 October 1975, Mark Silberstein, McDonald collection. One specimen, CASIZ 068311, Pacific Grove, Point Pinos, large tide pool, 21 July 1921, F.M. MacFarland. Three specimens, CASIZ 171659, 1 dissected, Cabo Falsa, Bahia de San Quintin, Baja California, Mexico, 22 May 2001, Jeff Goddard. One specimen, CASIZ 171660, dissected, Cabo Falsa, Bahia de San Quintin, Baja California, Mexico, 27 May 2001, Jeff Goddard. EXTERNAL MORPHOLOGY.— The body is slender and elongate with a narrow mantle margin, which undulates slightly (Figure 1C—D). The total length is up to 20 mm. The dorsum is smooth and without tubercles or spicules. There are some radiating connective tissue strands in the mantle margin. The rhinophores are straight, with 7-13 long sloping lamellae. The rhinophoral and branchial sheaths are slightly raised and smooth. There are 3—6 bipinnate or weakly tripinnate gills 196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 18 arranged in a circlet broken by the mid-posterior anus. The ground color is semi-translucent white or pale cream. There are round dark reddish brown or chocolate brown spots on the notum but not the mantle rim. These vary in size, but the larger spots tend to be clustered in four areas in the mid- line. These are, in front of the rhinophores, behind the rhinophores, the middle of the body and in front of the gills. Sometimes the anterior-most cluster is absent. Smaller spots are scattered towards the edges of the dorsum. Rhinophores and gills are white or cream; the gills may have a very few, small, brown spots near the tips. The ventral surface of the foot has no spots, but small spots may be present on the sides of the body and the dorsal surface of the trailing posterior foot. The head is small, with a pore-like mouth opening located within the notch of the foot. The short, ungrooved tentacles are attached to the foot and body wall. The foot is bilabiate with only the upper labium notched in the centre line; the main portion of the foot is entire, deeply wrinkled near the mouth. The double edge continues to the region of the reproductive openings. The foot flange is small and undulating. The posterior end is widely rounded. ANATOMY.— The notum is soft, without spicules and with fine connective tissue strands. The basal layer is thick and just above it is a layer with small lacunae. The viscera are within a color- less envelope. The digestive tract is illustrated in Figure 2B. The buccal bulb (oral tube) is conical and muscular. Posteriorly it is attached to the body wall by three slender muscles on each side. Ventrally, a long duct from the bilobed ptyaline (oral) glands enters the buccal bulb at its junction with the pharynx. This ptyaline duct branches into two within each gland, which are close togeth- er anteriorly. In a juvenile specimen, both glands were clearly separate. Posteriorly, each gland has a triangular white patch of larger round cells. The narrow, tubular, gradually widening pharynx is highly muscular. It bends in a reverse S-shape and sometimes loops before it passes through the nerve ring. Posterior to the nerve ring, the pharynx bends in a reverse C- or S-shape and constricts at its junction with the esophagus. Just anterior to this junction lie the two round, dorsal buccal gan- glia. Posterior to the junction are two small, round, lateral esophageal (salivary) glands. Next to each gland are two slender retractor muscles, which lie on either side of the anterior portion of the esophagus. After their insertion on the esophagus, the esophagus gradually widens, curves, and becomes irregular in texture. The esophagus is longer than the pharynx (1.4:1). Posteriorly, a sphincter separates the esophagus from the short, exposed section of the stomach. The remainder of the stomach is embedded in the confluent digestive glands. The intestine emerges from the digestive glands left of the midline and bends to the right. There is a short stalked, round, anterior pyloric gland (caecum). The narrow intestine curves medially to run posteriorly down the midline where it exits on the smooth anal papilla, located posteriorly in the gill circlet. The heart (Fig. 2B) forms a broad triangular auricle with a large, rounded anterior ventricle. A prominent renal syrinx, on the posterior right side, drains the pericardial sac. A medial aorta runs anteriorly from the ventricle to the region of the central nervous system. A small white blood gland is located to the right of the esophagus, with ducts attached to the esophagus at its bend. In the central nervous system (Fig. 2B), the elongate cerebro-pleural ganglia are fused but can be distinguished by a furrow separating the anterior cerebral ganglia from the posterior pleural gan- glia. There is a distinct mid-line furrow separating the two cerebro-pleural ganglia. The dorsal sur- face is smooth although large cells can be seen. The stout rhinophoral nerves each have a large gan- glion at their bases. The eyes are moderate in size and almost sessile. The cerebro-buccal connec- tives are long and thin, within the same connective tissue, extending to two round buccal ganglia, side by side on the dorsal posterior portion of the pharynx. The large, round pedal ganglia are ven- tral to the cerebro-pleural ganglia and the pharynx. They are connected by a very short commissure and have smaller nerve cells. The hermaphroditic gland (ovotestis) is spread over the digestive glands. The white female fol- MILLEN AND BERTSCH: NEW SPECIES OF POROSTOME NUDIBRANCHS 197 licles form clusters with granular, creamy-yellow male follicles between the clusters. The repro- ductive system (Fig. 5) has a long pre-ampullary duct, which widens abruptly into a bean- shaped ampulla. The ampulla has a distinct post ampullary duct, which bifurcates into a long oviduct and a vas deferens. The prostatic portion of the vas defer- ens is tubular and not greatly enlarged. The narrow, muscular deferent (ejaculatory) duct of the vas deferens is shorter than the prostatic portion. It widens slightly for half of its length and then enters the common atrium FIGURE 5. Dendrodoris behrensi sp. nov. (070836). Reproductive system with a short conical penis. The crayar by camera ducidas Scale bare 1 mm. Key: amampuls, be=bursa cop- +, eee : ulatrix, dd=deferent duct, fgm=female gland mass, ga=genital atrium, penis tip Is armed with 17-18 nd=nidamental duct, od=oviduct, p=penis, pr=prostate, ud=uterine duct, rows, 5—6 per row, of small, vg=vagina. recurved spines 5.4—7.5 um in height (Fig. 4D). The remainder of the vas deferens appears to have no spines. The elongate geni- tal atrium is glandular, striated and papillate internally. Between the male and female ducts is a muscular section with attached muscle strands from the junction of the prostatic and ejaculatory portion of the vas deferens. Other strands run to the seminal receptacle. The long, tubular vaginal duct is threadlike and muscular throughout its length. It enters a round or oval bursa copulatrix. The uterine duct is distinct but is attached to the vagina for a short distance, then it separates and con- volutes. The oval seminal receptacle has a moderately long duct, which joins the uterine duct near its junction with the oviduct. The oviduct and uterine duct together enter the female gland mass. The bursa copulatrix varies from equal in size to the seminal receptacle to twice its size. The female gland mass is large, elongate, and oval in shape. It consists of a large, granular pinkish or cream- colored section into which the combined uterine and oviduct enters ventrally and a highly convo- luted, firm, white mucus gland. There is a soft area at the base of the long nidamental duct. There is no vestibular gland. The nidamental duct adheres to the atrium and opens posterior to it. The reproductive openings usually appear as one small, round depression located high on the right side approximately one third of the way back from the anterior mantle margin, but the genital atrium can form an expanded papilla with the female gland mass opening posteriorly at its base. NATURAL HISTORY.— This species was first collected from the Great Tide Pool, in Pacific Grove, California, in 1921 and again in 1929 by Frank Mace MacFarland. The known range of Dendrodoris behrensi is from Point Pinos, Monterey Bay, California (approximately 37°27'N; 121°55’'W), to the San Benitos Islands, Baja California, Mexico (approximately 28°18'N; 115°34’'W). This species has been found in February, May, July and October, intertidally and sub- tidally to a depth of 50 m. The prey sponge of this species is unknown. The spawn and larval devel- opment have been studied by Goddard (2005a, b) and it is known to have ametamorphic direct development. Discussion.— The smooth dorsum, clustered, irregular chocolate brown spots on an off-white 198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 18 background and pale rhinophores and gills found in this species are distinctive. Other species with similar coloration are D. albobrunnea Allan, 1933 and D. elongata Baba, 1936. Both of these species can be distinguished by their flatter, more elongate bodies, presence of small tubercles, reg- ular brown spotting and leach-like locomotion. Internally, D. elongata is not known, but D. albo- brunnea has more rows of penial spines, which have long bases and a vestibular gland is present. Dendrodoris coronata Kay and Young, 1969 can be distinguished by small scattered pustules and a transverse raised ridge located anterior to the gills, which has large crown-like pustules. Internally it has no penial hooks. When D. behrensi is added to the cladograms along with D. stohleri, it lies just basal to the sibling species D. limbata and D. stohleri. It differs in that it has a smaller, tubu- lar prostate and fewer rows of penial hooks but the hooks of all three species have short bases. A recently described, smooth, white species from California, D. azinae Behrens and Valdés, 2004 can be distinguished by its dark purple-tipped gills and rhinophores and lack of brown spots on the body. ACKNOWLEDGMENTS We thank those who helped us collect, or gave us specimens or photos, enabling us to under- stand the zoogeography and anatomy of these two new species. Assistance with Dendrodoris stoh- leri is gratefully acknowledged to Brian Coleman, Carolina Espinoza, Antonio Ferreira, Terrence Gosliner, Alan Grant, Michael D. Miller, David K. Mulliner, Antonio Resendiz, Tom Smith, and Robert van Syoc. For Dendrodoris behrensi we thank Shane Anderson, Mark Chamberlain, Jeff Goddard, P.A. Haaker, Frank Mace MacFarland, George E. MacGinitie, Gary McDonald, and Mark Silberstein. This research was partly funded by the Department of Zoology, University of British Columbia, to Sandra Millen. We especially thank the graciousness and inspiration of Rudolf Stohler and David Behrens, without whom this manuscript would not have been possible. REFERENCES BEHRENS, D.W. 1980. Pacific Coast Nudibranchs, A Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers, Los Osos, California, USA. 112 pp. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs, A Guide to the Opisthobranchs Alaska to Baja California, 2d ed. Sea Challengers, Monterey, California, USA. 107 pp. BEHRENS, D.W. 2004. Pacific Coast Nudibranchs — Supplement II: New species to the Pacific Coast and new information on the oldies. Proceedings of the California Academy of Sciences 55:11—54. BEHRENS, D.W., AND A.VALDEs. 2004. A new species of Dendrodoris (Mollusca: Nudibranchia: Dendrodorididae) from the Pacific Coast of North America. Proceedings of the California Academy of Sciences 55:408-413. Bertscu, H. 1991. Seasonal, geographic and bathymetric distribution of opisthobranchs at Bahia de los Ange- les, Baja California, Mexico. Western Society of Malacologists, Annual Report 23:18. Bertscu, H. 2000. Dr. Rudolf Stohler: Some personal remembrances. The Veliger 43(4):ii. Bertscu, H., M.D. MILLER, AND A. GRANT. 1998. Notes on opisthobranch community structures at Bahia de los Angeles, Baja California, Mexico. Opisthobranch Newsletter 24:35-36. BropiE, GD. 2001. Some comparative histological aspects of the dendrodorid genera Doriopsilla and Dendrodoris (Opisthobranchia: Nudibanchia). Bollettino Malacologico, Roma 37:99-104. BropiE, GD., AND G CALADO. (In press.) Dendrodoris arborescens (Collingwood, 1881) (Mollusca: Nudibranchia): larval characteristics reveal a masked porostome. Records of the Western Australian Museum. Supplement. World Malacology Conference Opisthobranch Symposium. MILLEN AND BERTSCH: NEW SPECIES OF POROSTOME NUDIBRANCHS 199 Bropie, G.D., R.C. WILLAN, AND J.D. COLLINS. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia: Dendrodorididae) in the Indo-West Pacific Region. Journal of Molluscan Studies 63:407-423. Euior, C.N.E. 1906. The genus Doriopsilla Bergh. Journal of Conchology 11:366—367. Gray, J.E. 1847. A list of the genera of recent Mollusca, their synonyma and types. Proceedings of the Zoological Society of London, pt. 15:129-219. GODDARD, J.H.R. 2005a. Developmental mode in benthic opisthobranch molluscs from the northeast Pacific Ocean: feeding in a sea of plenty. Canadian Journal of Zoology 82:1954-1968. GODDARD, J.H.R. 2005b. Ametamorphic direct development in Dendrodoris behrensi (Nudibranchia: Dendrodorididae), with a review of developmental mode in the family. Proceedings of the California Academy of Sciences, ser. 4, 56:201—211. GOSLINER, T.M., M.C. SCHAEFER, AND S.V. MILLEN. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodorididae) from the Pacific Coast of North America, including a comparison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42:201—210. LEE, R.S., AND P. BRopHy. 1969. Additional bathymetric and locality data for some opisthobranchs and an octopus from Santa Barbara County, California. The Veliger 12:220-221. McDonaLp, G. 1983. A Review of the “Nudibranchs of the California Coast.” Malacologia 24:114-276. McDonaLp, G. AND J. NYBAKKEN. 1981. Guide to the Nudibranchs of California. Including most species found from Alaska to Oregon. American Malacologists, Inc., Melbourne, Florida. 72 pp. PRUVOT-FOL, A. 1930. Du genre Dendrodoris Ehrenberg et de ses rapports avec le genre Doriopsis Pease et avec queques autres. Note sur la taxonomie des nudibranchs. Bulletin du Museum d ‘Histoire Naturelle, Paris, ser. 2, 2(3):291-297. PRUVOT-FOL, A. 1954. Mollusques Opisthobranches. Faune de France, vol. 58. 460 pp. RUDMAN, W.B. 2003. Dendrodoris sp. 4. In: Sea Slug Forum. Australian Museum, Sydney, Australia. . STEINBERG, J. 1961. Notes on the Opisthobranchs of the West Coast of North America. The Veliger 4:57-63. THOLLESSON, M. 2000. Increasing fidelity in parsimony analysis of dorid nudibranchs by differential weigh- ing, or a tale of two genes. Molecular Phylogenetics and Evolution 16:161—172. VaLpes, A. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnaean Society 136:535—636. Vatpes, A. 2003. Preliminary molecular phylogeny of the radula-less dorids (Gastropoda:Opisthobranchia) based on 16S mtDNA sequence data. Journal of Molluscan Studies 69:75—-80. VALDES, iN AND D.W. BEHRENS. 1998. A new species of Doriopsilla (Mollusca, Nudibranchia, Dendrodorididae) from the Pacific coast of North America. Proceedings of the California Academy of Sciences, ser. 4, 50:307-314. VALDES, A., AND T.M. GOSLINER. 1999. Phylogeny of the radula-less dorids (Mollusca, Nudibranchia), with the description of a new genus and a new family. Zoologica Scripta 28:3 15-360. VALDES, A., AND J. ORTEA 1997. Review of the genus Doriopsilla Bergh, 1880 (Gastropoda. Nudibranchia) in the Atlantic Ocean. The Veliger 40:240-254. Vatpes, A., J. OrTEA, C. AviLa, AND M. BALLESTEROS. 1996. Review of the genus Dendrodoris Ehrenberg, 1831 (Gastropoda: Nudibranchia) in the Atlantic Ocean. Journal of Molluscan Studies 62:1-31. WAGELE, H., G. BRODIE, AND A. KLUSSMANN-KOLB. 1999. Histological investigations on Dendrodoris nigra (Stimpson, 1855) (Gastropoda, Nudibranchia, Dendrodorididae). Molluscan Research 20:79-94. WOLLSCHEID-LENGELING, E., J. BooRE, W. BROWN, AND H. WAGELE. 2001. The phylogeny of Nudibranchia (Opisthobranchia, Gastropoda, Mollusca) reconstructed by three molecular markers. Organisms Diversity and Evolution 1(4):241-256. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Volume 56, No. 19, pp. 201-211, 1 fig., 2 tables. August 26, 2005 Ametamorphic Direct Development in Dendrodoris behrensi (Nudibranchia: Dendrodorididae), with a Review of Developmental Mode in the Family Jeffrey H. R. Goddard Marine Science Institute, University of California, Santa Barbara, California 93106, USA; Email: goddard@lifesci.ucsb.edu Dendrodoris behrensi Millen and Bertsch, 2005 has ametamorphic direct develop- ment. The embryos passed through a vestigial veliger stage lacking a shell, opercu- lum, larval retractor muscle and pedal sensory cilia. After an embryonic period of 38 days (16—19°C) they hatched as juveniles averaging 512 um in dorsum length. Newly hatched juveniles possessed eyespots and a lattice of spicules on the ventral surface of the dorsum, and after an additional four days, they possessed rudimentary rhino- phores. Only three other examples of direct development have been noted from nudi- branchs from the northeast Pacific Ocean, all from the Family Dendrodorididae. Based on a survey of the literature, mode of development was determined for 26 species and forms of dendrodoridids worldwide. Fifty-four percent of these taxa have planktotrophic development, 4% lecithotrophic development, and 42% direct development. Direct development is significantly more prevalent in the Dendrodorididae than reported for opisthobranchs worldwide by Hadfield and Miller (1987). Three hypotheses are presented to explain this: (1) direct development is adaptive in overcoming size constraints on post-metamorphic juveniles stemming from their lack of a radula and suctorial mode of feeding on sponges. (2) Direct development is prevalent because small adult size, which is generally correlated with direct development in marine invertebrates, has been selected for in many den- drodoridids. (3) Direct development is an adaptation against high larval mortality in some regions. Limited evidence tends to support hypotheses 2 and 3, but with some interesting developmental exceptions, not hypothesis 1. KEY WORDS: Ametamorphic, direct development, dorid nudibranch, northeast Pacific Ocean, porostome Direct development, in which juveniles, rather than larvae, hatch from egg coverings, is rare in shallow-water, benthic opisthobranchs from the northeast Pacific Ocean (Goddard 2005). Among the nudibranchs, the porostome dorids Dendrodoris nigromaculata (Cockerell in Cockerell and Eliot, 1905) and Doriopsilla spaldingi Valdes and Behrens, 1998 have been noted to hatch from their egg masses as juveniles (Lance 1982; Behrens 1998; Valdés and Behrens 1998). Additionally, direct development has been reported for a yellow-gilled porostome from Bahia de los Angeles in the Gulf of California (Mulliner 1972; J. Lance, pers. commun., 2005). As present- ly understood, this form is considered Doriopsilla gemela Gosliner et al., 1999, which was described from Bahia de los Angeles, the west coast of Baja California and California (S. Millen, pers. commun., 2005). However, Doriopsilla gemela produces free-swimming larvae (Gosliner et al., 1999; pers. observ.; J. Lance, pers. commun., 2005). Given the rarity of variable developmen- tal mode, or poecilogony (Bouchet 1989; Chia et al. 1996), the form referred to by Mulliner (1972) 201 202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 19 may be an undescribed, cryptic species. All three of the above taxa with direct development occur south of Point Conception, the boundary between the Oregonian and Californian biogeographic provinces, although Doriopsilla spaldingi has also been collected from 64 m depth in central California (Behrens 2004). Development has not been described in detail for any of them. The pres- ent paper describes ametamorphic direct development in Dendrodoris behrensi Millen and Bertsch, 2005, a porostome known from central Baja California to Monterey, California (Millen and Bertsch 2005). This is the first detailed account of direct development in a nudibranch from the northeast Pacific Ocean. METHODS Four specimens of Dendrodoris behrensi, 12 to 22 mm long, were collected on 27 May 2001 from the interstices of a low intertidal oyster reef on the east shore of Bahia Falsa near San Quintin, Baja California, Mexico. These were transported back to the laboratory and held in a 100 ml jar of coarsely filtered seawater on a flow-through seawater table at 16—19°C until they laid egg masses. Three newly laid egg masses were examined using an Olympus compound microscope equipped with an ocular micrometer and 35-mm camera, and the diameters of a random sample of ten zygotes were measured from two of these. Each egg mass was then transferred to a separate vial. The water in all of these vials was changed at least once daily, and the egg masses examined every few days until the juveniles hatched. The live juveniles were then examined, measured and pho- tographed using the compound microscope. Juveniles were not reared more than eight days after hatching. After obtaining the above egg masses, the adult slugs were relaxed in 7.5% MgCl2 and then fixed in 70% ethanol. These adults were subsequently deposited as paratypes of this newly described species in the California Academy of Sciences (CASIZ 171659, 171660) (Millen and Bertsch 2005). Statistical analyses (see section on developmental mode in the Dendrodorididae) were con- ducted using JMP 4.0.4 (SAS Institute). RESULTS Dendrodoris behrensi laid white egg ribbons, up to 1.9 mm high, in a loose, open spiral of one turn. The mean diameter of the uncleaved eggs was 181.4 um (SD = 9.82 um, n = 10) in one egg mass and 187.5 um (SD = 6.75 um, n = 10) in another. Each egg mass contained approximately 100 white eggs, deposited one, or rarely two, per capsule. The walls of the egg capsules were 10- 20 um thick, and in some capsules, up to 30 um thick (Fig. la). The jelly matrix surrounding the egg capsules was slightly milky in appearance and tougher than that typically observed in the egg masses of nudibranchs from the northeastern Pacific (pers. observ.). The embryos were observed at irregular intervals until late in development, and therefore an exact chronology of the major stages of embryonic development cannot be presented. However, certain stages and events were noted on the following dates: blastulae 3 days after egg mass depo- sition (Fig. la); gastrulation in process on days 6 and 7; clear anal cells and the rudimentary lobes of the metapodium and velum on day 12. On day 18 the embryonic veligers had small cilia on the velum and were moving slightly; these also had very early eyespots. By day 24 the embryos had well-developed eyespots with lenses and were rotating slowly in their capsules (Fig. 1b). After 28 days, the mantle was semi-transparent and contained refractile glands, which presumably serve a defensive function after hatching. On day 31, the velum on most of the embryos was in the process of being resorbed, and this was complete by day 33. By this time the body had a more juvenile GODDARD: AMETAMORPHIC DIRECT DEVELOPMENT IN A NUDIBRANCH 203 velum SS “I 3 » PAY : hindgut “6 foot ETE : rhinophore spicules eyespot r. cg kidney dorsum foot spicules D FiGuRE 1. Stages in the development of Dendrodoris behrensi. (A) Blastulae, three days after oviposition. (B) Embryonic veligers, right lateral views, day 24. (C) Day 38, hatching juvenile, right lateral view. (D) Juvenile four days after hatching, dorsal view. All images are bright field photomicrographs of live specimens; scale bars = 100 «1m. Abbreviations used in figure: bm, buccal mass; ec, egg capsule; dg, digestive gland; pp mgl, propodial mucus gland; r. cg, right cerebral ganglion; r. pg, right pedal ganglion. shape, with a distinct dorsum and large foot. Slightly curved, spindle shaped spicules were observed in the dorsum on day 36, and 2 days later, on day 38, the embryos hatched as juvenile slugs. A shell, operculum and larval retractor muscle were not observed at any stage of develop- ment, and the velar cilia remained small compared to the locomotive cilia found on the velum of embryos hatching as swimming veliger larvae. The foot of the embryos also never developed the long, stiff sensory cilia observed on developing planktotrophic veligers (Thompson 1967; pers. observ. ). At hatching, the juveniles measured 465-550 um along the dorsum (mean = 512 um, n = 3) and had distinct eyespots, rhinophore rudiments and a small protruding tail (Fig. 1c). The ventral surface of the dorsum had a lattice of spicules 5 um in diameter. Outside the hepatic region the dor- sum was studded with defensive glands up to 12 um in diameter. Anteriorly, the end of the dorsum was not fully differentiated and lacked a distinct edge. The buccal mass, cerebral and pedal gan- glia, and propodial mucus gland were visible through the body wall. Posteriorly, the digestive gland and a posterior, refractile, translucent yellow organ (presumably a kidney) were also visible through both the dorsum and the lateral body wall (Fig.1c). Four days after hatching, the juveniles were 570 um long, and the dorsum was oval-shaped, slightly wider anteriorly and generally more adult-like in appearance (Fig. 1d). Rhinophores, 35- 40 um high, had developed, and the foot and dorsum measured up to 100 and 230 um wide, respec- tively. Eight days after hatching, the juveniles were about 600 um long. 204 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 19 DISCUSSION Dendrodoris behrensi has ametamorphic development, as described by Bonar (1978). Veliger structures such as the shell, operculum, larval retractor muscle, and pedal sensory cilia are all lack- ing, and the velum and its locomotive cilia are vestigial. Its development is similar to that described by Thompson (1967) for Cadlina laevis (Linnaeus, 1767). No shell was detected in the develop- ment of D. behrensi, however, and the development of the spicules in the dorsum was accelerated compared to those in C. /aevis, which develop after hatching. Although none of the other examples of direct development in nudibranchs from the northeast Pacific Ocean (see Introduction) has been described in detail, enough information exists to catego- rize further the development of two of them. Mulliner (1972) noted the lack of a shell in the embry- onic development of the yellow porostome from Bahia de los Angeles, indicating that this form has ametamorphic direct development. The illustration by Lance (1982), although not specifically labeled, clearly depicts a post-metamorphic Dendrodoris nigromaculata hatching at a length of 500 um with notal spicules and a distinct eyespot. An empty egg capsule, with no sign of a veliger shell, is also shown, suggesting that this species also has ametamorphic development. Not enough infor- mation is available on the development of Doriopsilla spaldingi to categorize its development as ametamorphic or capsular metamorphic (Goddard 2005, Table 1). The hatching juveniles of Dendrodoris behrensi are similar to those of D. miniata (Alder and Hancock 1864) in their possession of a spiculate dorsum (Thompson 1975), but their embryonic development differs in that the latter develops a coiled shell (Thompson 1975; Rose 1985). All known examples of direct development in nudibranchs from the northeast Pacific Ocean are from the family Dendrodorididae, and of the species of Dendrodoris from this region for which mode of development is known, both have ametamorphic development (Goddard 2005). These findings suggest that the non-feeding modes of development, especially direct development, might be more prevalent in the Dendrodorididae than in other Opisthobranchia, an estimated two-thirds of which produce planktotrophic larvae (Hadfield and Miller 1987). Patterns of developmental mode in the family are therefore examined below. Patterns of developmental mode in the Dendrodorididae Data on mode of development of the Dendrodorididae, gathered from the literature, are sum- marized in Table 1. Of the 28 species and forms listed, mode of development could be determined or inferred for 26. The egg diameters reported by Valdés et al. (1996) and Marcus and Marcus (1967) for Dendrodoris warta and Doriopsilla janaina, respectively, are enough above the 105 um minimum diameter known for nudibranch species with either lecithotrophic or direct development (see Hadfield and Miller 1987) that developmental mode can not be reliably inferred for them based on egg size alone. Two species in Table 1, Dendrodoris krebsii and Doriopsilla areolata, appear to have more than one type of development, suggesting that each may actually represent a complex of cryptic species, or that the taxonomy of the forms studied is simply problematic. As noted by Millen and Bertsch (2005), the latter appears to be the case for at least the specimens from the eastern Pacific previously referred to as Dendrodoris krebsii. According to Millen and Bertsch (2005) some of these specimens may actually be Dendrodoris arborescens, a possibility consistent with the deter- minations of developmental mode in Table 1. With regard to Doriopsilla areolata, three subspecies were recognized by Valdés and Ortea (1997), two of which have large egg diameters and direct development (Table 1). Valdés and Ortea (1997) did not specify the subspecies from which they observed relatively small eggs (and for which planktotrophic development is inferred in Table 1). GODDARD: AMETAMORPHIC DIRECT DEVELOPMENT IN A NUDIBRANCH 205 Based on the geographic distributions in Valdés and Ortea (1997, Figure 5) it presumably was Doriopsilla areolata areolata. However, according to Ballesteros and Ortea (1980) this subspecies has large eggs. If this discrepancy in egg diameters is accurate, then Doriopsilla areolata areolata as presently understood may actually consist of at least two cryptic taxa. Doriopsilla gemela was originally reported as having lecithotrophic larval development, from eggs 240 um in diameter (range: 120 to 300 um), based on observations of larvae from flat egg masses laid by adults collected 7 km north of the tip of Point Loma, San Diego (Schaefer 1997; Gosliner et al. 1999:207). However, the larva pictured in Figure 5 of Schaefer (1997) appears to have a small foot lacking a propodium, and based on the egg mass measurements reported by Gosliner et al. (1999:207), the eggs/embryos in their Figure 4B measure approximately 100 um in diameter. This egg size, and especially the lack of a propodium, indicate planktotrophic develop- ment (Bonar 1978: Hadfield and Miller 1987). Moreover, the 180 um range in egg diameter, if accurate, is much larger than any known from a NE Pacific opisthobranch (Goddard 2005) and could only indicate poecilogony, which is extremely rare, or a species complex. For these reasons, the egg sizes and lecithotrophic development reported by Gosliner et al. (1999) require confirma- tion, and I here classify its development as planktotrophic, based on my own recent observations and the older observations of J. Lance, both from flat egg masses laid by adults collected at Point Loma (see Table 1). Of the 26 species and forms in Table | for which mode of development was determined, 11, or 42% have direct development, 4% are lecithotrophic, and 54% are planktotrophic (Table 2). Direct development is significantly more prevalent in the Dendrodorididae than in opisthobranchs worldwide (Table 2; 2x2 contingency table analysis, log-likelihood ratio %* = 8.937, p = 0.003, lumping lecithotrophic and planktotrophic modes together as “indirect” development with a free- living larval stage). Moreover, at least five of the 11 taxa with direct development have ametamor- phic development (Table 1), presumably the most evolutionarily derived mode of development (Hadfield and Miller 1987). Three hypotheses, not mutually exclusive, might explain the high prevalence of direct devel- opment in the Dendrodorididae. These are presented below, each followed by a brief discussion of the pertinent evidence. (Hypothesis 1) Direct development has been selected for in the Dendrodorididae as a means of overcoming size constraints on post-metamorphic juveniles stemming from their lack of a radu- la and suctorial mode of feeding on sponges. It may be difficult to commence this mode of feed- ing, at least on some prey species, at the post-metamorphic sizes (typically well under 500 tm) characteristic of many indirectly developing opisthobranchs. Hadfield and Miller (1987, Figure 13) found that direct development, especially ametamorphic development, can result in post-metamor- phic sizes double that of planktotrophy and lecithotrophy. They concluded that the presence of a larval shell sets a maximum size limit of about 500 im in newly metamorphosed opisthobranch juveniles. Based on the hatching sizes of the directly developing species in Table | (which range from 300 to 800 um, with a mean of 474 um), this hypothesis does not appear to be supported. However, the sample size (n= 6) is small, and to reject this hypothesis, the hatching sizes of the directly developing species should really be compared to the post-metamorphic sizes of their planktotrophic and lecithotrophic congeners. To my knowledge, these data do not exist. If the hypothesis linking direct development to feeding-related constraints on juvenile body size is eventually borne out, then: (a) it obviously does not apply to all members of the family, depending perhaps on dietary composition. (b) We might expect to find a similarly high prevalence of direct development in the Phyllididae, a closely related family also characterized by the evolu- tionary loss of the radula and suctorial feeding on sponges (Valdés 2003). (c) We might expect 206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 19 TABLE |. Comparative data on embryonic development in the Dendrodorididae. Shell or juve- Mode of Max Reference nile length at develop- adult size hatching (um) ment (mm) Species Egg Embryonic Temp. diameter period (°C) (um) (days) Dendrodoris angolensis Valdés 64 - - & Ortea, 1996 D. arborescens (Collingwood, 121-159 6-9.2 22-27 1881) D. behrensi Millen & Bertsch, 181-187 38 16-19 2005 D. coronata Kay & Young, 55 6 27-29 1969 Distormale 105 6-7 22-26 D. form 2¢ 186 18-19 22-23 D. elongata Baba, 1936 165 - - D. fumata (Riippell & Leuckart, 1831) Gray form 100-151 5.5-17 16-30 Orange/red form 85-129 - - D. grandiflora (Rapp, 1827) 80 - - D. krebsii (Morch, 1863) From Florida and Caribbean 146-205 21-23 10 coast of Panama From Caribbean Sea, Brazil 66-114 9-10 10 and Pacific coast of Panama D. limbata (Cuvier, 1804) 270-300 >17 - D. nigra (Stimpson, 1855) 60-93 4-9.6 22-29 D. nigromaculata (Cockerell - - - in Cockerell and Eliot 1905) D. rubra (Kelaart, 1858) var. 308 7 - nigromaculata (Eliot, 1913) D. senegalensis Bouchet, 1975 < 11 Qh - - D. warta Marcus & Gallagher, 120 - - 1976 Doriopsilla albopunctata 108 18-191 14-15 (Cooper, 1863) D. areolata Bergh, 1880 106 - - D. areolata areolata Bergh, 209-266 - - 1880 D. areolata nigrolineata 195 >35 10 Meyer, 1977 D. gemela Gosliner, Schaefer & Millen, 1999 From Point Loma 80 - - From Point Loma! 111 8.5 15-21 From 7kmN of PointLoma 240 31 14 D. janaina Marcus & Marcus, 97-140 >22 10 1967 D. miniata (Alder & Hancock, 215-228 16-17 22-23 1864) D. pharpa Marcus, 1961! 203-234 14-116) 23225 D. spaldingi Valdés & - Behrens, 1998 Yellow porostome from Bahia ~260 de los Angeles, Baja California™ - (P) 45+ Valdés et al. 1996 144-153 P 78 Rose 1985b; Brodie et al. 1997; Brodie and Calado in press 512 AM 22 ‘This study - (P) 40 Johnson and Boucher 1983 150 P 138 Rose 1985; Orr 1981 440 AM - Rose 1985 - (L)4 75 Johnson and Boucher 1983 Gohar & Soliman. 1967; Soliman 1991: Brodie et al. 1997; Brodie & 157-220 P 97 Calado (in press) - (P) 38 Brodie et al. 1997 - (P) 32+ Valdés et al. 1996 410-460 CM 70° Clark & Goetzfried 1978; DeFreese & Clark 1983; Gonsalves-Jackson 2004 210 (P) 70 Bandel 1976; Valdés et al. 1996; Gonsalves-Jackson 2004 - cmt 53+ Tchang-Si 1931; Bonar 1978; Valdes et al. 1996 114-135 P 64 Ostergaard 1950; Kay & Young 1969; Johnson & Boucher 1983; Rose 1985b; Brodie et al. 1997; Brodie & Calado in press 500 (AM) 10 Lance 1982 165 (P) 70 Baba 1949; Amio 1963 : (P) 44+ = Valdés et al. 1996 - - 71+ Marcus & Gallagher 1976 195 P 60 Goddard 2005; personal observations = (P) 37+ Valdés & Ortea 1997 - (D) 38+ Ballesteros & Ortea 1980; Valdés & Ortea 1997 - (D) 30 Gonsalves-Jackson 2004 162 1v - J. Lance, pers. commun. 2005 173-184 iP - Personal observations - Lk 40 Shaefer 1997; Gosliner et al. 1999 - - 25 Marcus & Marcus, 1967: Gonsalves- Jackson 2004 360 AM 28 Thompson 1975; Rose 1985 300 CM 19 Clark and Goetzfried 1978; Eyster & Stancyk 1981; DeFreese & Clark 1983 - D 85 Valdés and Behrens 1998; Behrens 1998 800 (AM) 35 J. Lance, pers. commun., 2005; Mulliner 1972 GODDARD: AMETAMORPHIC DIRECT DEVELOPMENT IN A NUDIBRANCH 207 Notes to accompany Table 1 Mode of development: P, planktotrophic; L, lecithotrophic; CM, capsular metamorphic; AM, ametamorphic; D, direct (capsular metamorphic or ametamorphic not specified). Inferred modes of development are in parentheses and are based on morphological criteria, comparisons with congeners, and the egg size distributions reported by Hadfield and Miller (1987) for the major modes of development (see Goddard 2005). Values for egg diameter and shell or juvenile length at hatching are Means or ranges in means. 2 Values are maximum sizes of specimens examined by the authors of the references cited for each species. A plus sign (+) indicates a value for a preserved specimen. 5 Rose (1985) used the name Dendrodoris gemmacea (Alder and Hancock, 1864) for this planktotrophic species. However, the gemmacea-denisoni-gunnamatta species complex is in need of revision, and Rose did not deposit voucher specimens or photographs by which the identity of his specimens might be confirmed (B. Rudman, T. Gosliner, pers. com- mun., 2005). © Rose (1985) used the name Dendrodoris denisoni (Angas, 1864) for specimens of this directly developing form from Pilot Beach, an outer coast site in New South Wales (B. Rudman, pers. commun., 2005). The same comments in note b apply to this form. 4] consider lecithotrophic development most likely based on the egg size distributions reported by Hadfield and Miller (1987, Figure 1) for the major modes of development. Development in this species could actually be direct or (much less likely) planktotrophic. © Marcus and Marcus (1967:96) recorded 170 mm for a specimen from Biscayne Bay, Florida. f Hadfield and Switzer-Dunlap (1984) classified the development of this species as ametamorphic. As discussed by Bonar (1978) and Rose (1985), its development is intermediate between the capsular metamorphic and ametamorphic extremes of direct development, in that it possess “all of the larval structures, but in a partially reduced state” (Rose 1985). £ This value is probably inaccurate, as it is smaller than any other egg size known from the Opisthobranchia (see Hadfield and Miller 1987; Goddard 2005). h As stated by Valdés et al. (1996), 110 mm was the mean length of the egg capsules, not the eggs. i Gosliner et al. (1999) reported an anomalously long embryonic period of 31 days at 14°C. J The adults that laid the egg masses from which these data were collected have been deposited in the California Academy of Sciences (CASIZ catalogue numbers 172864-172865, and 172867) K Lecithotrophic development in D. gemela requires confirmation (see text). ! Clark and Goetzfried (1978) reported egg sizes and direct development in both Doriopsilla pharpa and D. sp., both from Florida. Given that the value they reported for D. sp. falls within the range reported for D. pharpa, and that the latter is the only species of Doriopsilla known from Florida (Valdés and Ortea 1997), I have assumed that their D. sp. is the same as D. pharpa. ™ This form is closest to D. gemela (see Introduction). TABLE 2. Percentage of developmental types in the Dendrodorididae, com- pared with the percentages for opisthobranchs worldwide (the latter from Hadfield and Miller 1987). Percent mode of development (number of species) Dendrodorididae Worldwide (from Mode of development (present study) Hadfield and Miller 1987)* Planktotrophic 54 (14) 68 (159) Lecithotrophic 4(1) 16 (37) Direct 42 (11) 16 (37) Total 100 (26) 100 (233) * These proportions and numbers have been recalculated, omitting all species also listed in Table 1 of the present study. 208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 19 other adaptations in the porostomes for escaping the constraints imposed by a larval shell and indi- rect development on post-metamorphic body size. (d) We might expect post-metamorphic juveniles of some of the indirectly developing porostomes to be at the upper size limits known for plank- totrophic and lecithotrophic opisthobranchs. Regarding prediction (b), to my knowledge, information on development has been reported for only three phyllidids, one of which is known to be planktotrophic (Ros 1981; Johnson and Boucher 1983; Soliman 1991). More data are therefore needed to characterize mode of development in the Phyllididae and evaluate this prediction. Regarding prediction (c), the planktotrophic veliger larvae of Dendrodoris arborescens, unlike most other nudibranchs, hatch without an operculum and have a large cephalopedal region which can not be withdrawn into the shell (Rose 1985b; Brodie and Calado 2005). Moreover, the shell is cast early in the pelagic phase (Brodie and Calado 2005). This larval morphology and development is strikingly similar to that of the aegirid dorids Aegires punctilucens (D’Orbigney, 1837) and A. albopunctata MacFarland, 1905 (Thiriot-Quiévreux 1977; Goddard 2001). After casting their shells larval A. punctilucens grow and develop a juvenile-like body with rhinophores, large foot and a highly spiculate dorsum (Thiriot-Quiévreux 1977). After settlement and loss of the velum, post-metamorphic juveniles are about 575 um long (Thiriot-Quiévreux 1977, Figure le), a size comparable to that attained by many ametamorphically developing opisthobranchs (Hadfield and Miller 1987). This two-stage metamorphosis, in which the larval shell is cast long before the loss of the velum, therefore may represent another adaptation for escaping the constraints on juvenile size imposed by a larval shell, and it appears to have evolved independently in Dendrodoris arborescens. Data to evaluate prediction (d) are lacking, but it is noteworthy that the shell size of the hatch- ing planktotrophic veligers of the gray form of Dendrodoris fumata, Dendrodoris kresbsii from the eastern Pacific and Doriopsilla albopunctata range from 195 to 220 um (Table 1), which is greater than the hatching shell size of most planktotrophic nudibranchs (Hadfield and Miller, Figures 1A and 8; Goddard 2005, Figures 2 and 3). Prediction (d) would be supported if the newly settled juve- niles of these species were found to be large compared to the juveniles of other planktotrophic opisthobranchs. (Hypothesis 2) Direct development is prevalent because small adult size, which is correlated with brooding and direct development in marine invertebrates (Strathmann and Strathmann 1982), has been selected for in many dendrodoridids. Species of dendrodoridids with direct development have a mean maximum adult size of 39.0 mm (n = 10), and are significantly smaller than the 15 species with indirect development, which have a mean maximum adult size of 61.9 mm of (Table 1, Wilcoxon two-sample test, normal approximation, Z = -2.277, p = 0.023), and this result is even more pronounced comparing the adult sizes of only those species with ametamorphic development to those with indirect development (Mean = 23.75 mm, n= 4 vs. 61.9 mm, n = 15, Wilcoxon two- sample test, normal approximation, Z = -2.852, p = 0.004). This hypothesis is, therefore, support- ed, but says nothing about the factors selecting for small adult size in the first place. (Hypothesis 3) Direct development has been selected for in some dendrodoridids as an adap- tation against high larval mortality (from low or poor quality food, predation, or transport away from favorable settlement sites) (see Goddard 2005). Most of the species listed in Table 1 occur in tropical to warm temperate waters, which, depending on the influence of the subtropical oceanic gyres, are often oligotrophic, less productive, and have a different phytoplankton composition com- pared to colder waters (Berger 1989; Mann and Lazier 1991). This hypothesis should, therefore, be investigated further, but it requires a biogeographic analysis, which is beyond the scope of this paper. GODDARD: AMETAMORPHIC DIRECT DEVELOPMENT IN A NUDIBRANCH 209 Regardless of the selective pressures that may have driven the evolution of direct development in the Dendrodorididae, once lost, larval feeding structures are not often regained (Strathmann 1978, 1993) and lineages constrained to non-feeding modes of development may then speciate or go extinct at geographic and temporal scales different from those for lineages with a larval stage or other potential for long range dispersal (e.g., Jablonski and Lutz 1983; Jablonski 1986; Palumbi 1994: Wares and Cummingham 2001). Given the prevalence of direct development in the Dendrodorididae, phylogenetic and phylogeographic analyses of this family should therefore strive to include species with this mode of development. ACKNOWLEDGMENTS I thank Sandra Millen, Bill Rudman, Angel Valdés and Richard Willan for reviewing Table | and (or) providing additional references. Jim Lance graciously shared data on the adult size and development of yellow-gilled porostomes from Bahia de los Angeles and San Diego. Terry Gosliner and Maria Schaefer shared images of the veliger larvae of Doriopsilla gemela. | also thank Terry Gosliner and an anonymous reviewer for their comments and suggestions on the man- uscript. LITERATURE CITED Amio, M. 1963. A comparative embryology of marine gastropods, with ecological considerations. The Journal of the Shimonoseki University of Fisheries 12:15—144. BaBa, K. 1949. Opisthobranchia of Sagami Bay collected by His Majesty The Emperor of Japan. Wwanami Shoten, Tokyo, Japan. 194 pp. BALLESTEROS, M., AND J.A. ORTEA. 1980. Contribuci6n al conocimiento de los Dendrodorididae (Moluscos, Opisthobranquios, Doridaceos) del litoral Ibérico. I. Publicaciones del Departmento de Zoologia, Universidad de Barcelona, Facultad de Biologia 5:25—37. BANDAL, K. 1976. Egg masses of 27 Caribbean opisthobranchs from Santa Marta, Columbia. Studies on Neotropical Fauna and Environment 11:87-118. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs. Sea Challengers, Monterey, California, USA. 107 pp. BEHRENS, D.W. 1998. Doriopsilla spaldingi Valdés & Behrens, 1998 [online]. Available from [accessed 21 March 2005]. BEHRENS, D.W. 2004. Pacific coast nudibranchs, Supplement I, New species to the Pacific coast and new information on the oldies. Proceedings of the California Academy of Sciences, ser. 4, 55:11—54. BerGER, W.H. 1989. Global maps of ocean productivity. Pages 429-455 in W.H. Berger, V.S. Smetacek, and G. Wefer, eds., Productivity of the Oceans: Present and Past. John Wiley & Sons, New York, New York, USA. Bonar, D.B. 1978. Morphogenesis at metamorphosis in opisthobranch molluscs. Pages 177-196 in F.S. Chia and M.E. Rice, eds., Settlement and Metamorphosis of Marine Invertebrate Larvae Elsevier/North- Holland Biomedical Press, New York, New York, USA. BOUuCHET, P. A. 1989. Review of poecilogony in gastropods. Journal of Molluscan Studies 55:67—78. Bropig, G.D., R.C. WILLAN, AND J.D. CoLLins. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia: Dendrodorididae) in the Indo-West Pacific region. Journal of Molluscan Studies 63:407-423. Bropiz, GD., AND G. CaLaApo. (In press). Dendrodoris arborescens (Collingwood, 1881) (Mollusca: Nudibranchia): larval characteristics reveal a masked porostome. Records of the Western Australian Museum. Supplement. World Malacology Conference Opisthobranch Symposium. CuI, F.-S., G. GIBSON, AND P.-Y. QUIAN. 1946. Poecilogony as a reproductive strategy of marine invertebrates. Oceanologica Acta 19:203-208. CLARK, K.B., AND A. GOETZFRIED. 1978. Zoogeographic influences on development patterns of North Atlantic 210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 19 Ascoglossa and Nudibranchia, with a discussion of factors affecting egg size and number. Journal of Molluscan Studies 44:283—294. DEFREESE, D.E., AND K.B. CLARK. 1983. Analysis of reproductive energetics of Florida Opisthobranchia (Mollusca: Gastropoda). /nternational Journal of Invertebrate Reproduction 6:\—10. EystTer, L.S., AND S.E. STANCYK. 1981. Reproduction, growth and trophic interactions of Doriopsilla pharpa Marcus in South Carolina. Bulletin of Marine Science 31:72-82. GODDARD, J.H.R. 2001. The early veliger larvae of Aegires albopunctatus (Nudibranchia: Aegiridae), with morphological comparisons to members of the Notaspidea. The Veliger 44:398—400. GODDARD, J.H.R. 2005 (for 2004). Developmental mode in benthic opisthobranch molluscs from the north- east Pacific Ocean: feeding in a sea of plenty. Canadian Journal of Zoology 82:1954-1968. GouAR, H.A.F., AND G.N. SOLIMAN. 1967. The biology and development of Dendrodoris (=Doridopsis) fuma- ta (Riippell and Leuckart) (Gastropoda, Nudibranchia). Publications of the Marine Biological Station, Al- Ghardagqa, Red Sea 14:31—54. GONSALVES-JACKSON, D.C. 2004. Opisthobranch Mollusks Across the Isthmus of Panama: Systematics and Biogeographic Distribution of Developmental Types. Ph.D. Dissertation, Florida Institute of Technology, Melbourne, Florida, USA. 378 pp. GOSLINER, T.M., M.C. SCHAEFER, AND S.V. MILLEN. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodorididae) from the Pacific Coast of North America, including a comparison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42:201—210. HADFIELD, M.G., AND M. SwITZER-DUNLAP. 1984. Opisthobranchs. Pages 209-350 in A.S. Tompa, N.H. Verdonk and J.A.M. van den Biggelaar, eds., The Mollusca, vol. 7, Reproduction. Academic Press, New York, New York, USA. HADFIELD, M.G, AND S.E. MILLER. 1987. On developmental patterns of opisthobranchs. American Malacological Bulletin 5:197—214. JABLONSKI, D. 1986. Larval ecology and macroevolution in marine invertebrates. Bulletin of Marine Science 39:565-587. JABLONSKI, D., AND R.A. Lutz. 1983. Larval ecology of marine benthic invertebrates: paleobiological impli- cations. Biological Reviews 58:21-89. JOHNSON, S., AND L.M. BOUCHER. 1983. Notes on some Opisthobranchia (Mollusca: Gastropoda) from the Marshall Islands, including 57 new records. Pacific Science 37:251-291. Kay, E.A., AND D.K. YOUNG. 1969. The Doridacea (Opisthobranchia; Mollusca) of the Hawaiian Islands. Pacific Science 23:172—231. LANCE, J.R. 1982. Dendrodoris nigromaculata. {illustration only] Opisthobranch Newsletter 14:29. MANN, K.H., AND J.R.N. LAZIER. 1991 Dynamics of Marine Ecosystems. Blackwell Science, Cambridge, Massachusetts, USA. 466 pp. Marcus, Ev., AND S.B. GALLAGHER. 1976. A new species of Dendrodoris from Florida. The Veliger 18:353-356. Marcus, Ev., AND ER. Marcus. 1967. Tropical American opisthobranchs. Studies in Tropical Oceanography 6:3-137. MILLEN, S.V., AND H. BERTSCH. 2005. Two new species of porostome nudibranchs (Family Dendrodorididae) from the coasts of California (USA) and Baja California (Mexico). Proceedings of the California Academy of Sciences, ser. 4, 56(19):189-199, 5 figs. MULLINER, D.K. 1972. Breeding habits and life cycles of three species of nudibranchs from the eastern Pacific. The Festivus 3:1—5. Orr, J. 1981. Hong Kong Nudibranchs. The Urban Council, Hong Kong, China. 82 pp. OSTERGAARD, J.M. 1950. Spawning and development of some Hawaiian marine gastropods. Pacific Science 4:75-115. PALUMBI, S.R. 1994. Genetic divergence, reproductive isolation, and marine speciation. Annual Review of Ecology and Systematics 25:547—572. Ros, J. 1981. Desarrolo y estrategias biondmicas en los opisthobranquios. Oecologia aquatica 5:147—183. Rose, R.A. 1985. The spawn and development of twenty-nine New South Wales opisthobranchs (Mollusca: Gastropoda). Proceedings of the Linnean Society of New South Wales 108:23-36. GODDARD: AMETAMORPHIC DIRECT DEVELOPMENT IN A NUDIBRANCH 211 Rose, R.A. 1985b. The spawn and embryonic development of color variants of Dendrodoris nigra (Mollusca: Nudibranchia). Jowrnal of the Malacological Society of Australia 7:75-88. SCHAEFER, M.C. 1997. Systematics and Biological Studies of Doriopsilla (Nudibranchia: Dendrodorididae) from California. Masters Thesis. San Francisco State University, San Francisco, California, USA. 56 pp. SOLIMAN, G.N. 1991. A comparative review of the spawning, development and metamorphosis of prosobranch and opisthobranch gastropods with special reference to those from the northwestern Red Sea. Malacologia 32:257-271. STRATHMANN, R.R. 1978. The evolution and loss of feeding larval stages of marine invertebrates. Evolution 32:894-906. STRATHMANN, R.R. 1993. Hypotheses on the origins of marine larvae. Annual Review of Ecology and Systematics 24:89-117. STRATHMANN, R.R., AND M.F. STRATHMANN. 1982. The relationship between adult size and brooding in marine invertebrates. American Naturalist 119:91—101. TCHANG-SI 1931. Un nouveau cas de condensation embryogénique chez un nudibranche (Doridopsis limbata Cuvier). Comptes Rendus Hebdomadaires des Seances de l’Academie des Sciences, Paris 192:302-304. THIRIOT-QUIEVREUX, C. 1977. Véligere planctotrophe du doridien Aegires punctilucens (D’Orbigny) (Mollusca: Nudibranchia: Notodorididae): description et métamorphose. Journal of Experimental Marine Biology and Ecology 26:177-190. THOMPSON, T.E. 1967. Direct development in the nudibranch Cadlina laevis, with a discussion of develop- mental processes in Opisthobranchia. Journal of the Marine Biological Association of the United Kingdom 47:1—22. THompsoNn, T.E. 1975. Dorid nudibranchs from eastern Australia (Gastropods, Opisthobranchia). Journal of Zoology, London 176:477-517. VALDES, A. 2003. Preliminary molecular phylogeny of the radula-less dorids (Gastrpoda: Opisthobranchia), based on 16s mtDNA sequence data. Journal of Molluscan Studies 69:75—80. VALDES, A., AND D.W. BEHRENS. 1998. A new species of Doriopsilla (Mollusca, Nudibranchia, Dendrodorididae) from the Pacific coast of North America. Proceedings of the California Academy of Sciences, ser. 4, 50:307-314. VaLpes, A., J. OrTEA, C. AVILA, AND M. BALLESTEROS. 1996. Review of the genus Dendrodoris Ehrenberg, 1831 (Gastropoda: Nudibranchia) in the Atlantic Ocean. Journal of Molluscan Studies 62:\—31. VALDES, A., AND J. ORTEA. 1997. Review of the genus Doriopsilla Bergh, 1880 (Gastropoda: Nudibranchia) in the Atlantic Ocean. The Veliger 40:240-254. Wares, J.P., AND C.W. CUMMINGHAM. 2001. Phylogeography and historical ecology of the North Atlantic intertidal. Evolution 55:2455-2469. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Volume 56, No. 20, pp. 213-273, 40 figs. 2 tables, Appendix August 26, 2005 Review of Acanthodoris Gray, 1850 with a Phylogenetic Analysis of Onchidorididae Alder and Hancock, 1845 (Mollusca, Nudibranchia) Shireen J. Fahey ! and Angel Valdés 2 ! California Academy of Sciences 875 Howard Street San Francisco, California 94103, USA Email: sfahey@calacademy.org; * Natural History Museum of Los Angeles County 900 Exposition Boulevard, Los Angeles, California 90007, USA; Email: avaldes@nhm.org The phylogenetic relationships of Acanthodoris Gray, 1850 within the Onchidorididae are examined based upon morphological characters. The present phylogenetic analysis supports the monophyly of Acanthodoris. Most of the described species of Acanthodoris and a newly described species from South Africa are examined. Species included in the present analysis are: Acanthodoris brunnea, A. falklandica, A. hudsoni, A. lutea, A. nanaimoensis, A. nanega, A. pilosa, A. pina, A. rhodoceras and A. serpentinotus. The new species has some external similarities to Acanthodoris pilosa (Abildgaard in Miiller, 1789) from the East Coast of North America and to Acanthodoris lutea MacFarland, 1925 from the West Coast of North America. Differences in the reproductive and radular morphology distinguish this new species. Reproductive differences in the new species include a shorter deferent duct, vaginal duct and receptaculum seminis duct than found in other externally similar species. Radula differences include multifid jaw rodlets and 6 or 7 elongate, pointed outer lateral teeth in the new species. This is in contrast to the triangular hook-like jaw structures and 5 or 6 flattened, triangular outer lateral plates in A. lutea. Acanthoris pilosa has no jaw rods and 3 flat outer lateral plates. Phylogenetic analyses place the new species in a relatively basal position to the remaining Acanthodoris species and closely related to Acanthodoris falklandica Eliot, 1907 from South America and Acanthodoris nanega Burn, 1969 from South Australia. The species from the West Coast of North America and the North Atlantic, i.e., A. pilosa, form a derived clade, separate from the Southern Hemisphere species. The phylogenetic relationships among the Onchidorididae are presented. Morphological characters of several species from the currently recognized family group are examined. Taxa included in the analysis are: Adalaria jannae Millen, 1987, Adalaria loveni (Alder and Hancock, 1862), Adalaria proxima (Alder and Hancock, 1854), Calycidoris guentheri Abraham, 1876, Diaphorodoris luteocincta Iredale and O’ Donoghue, 1923, Diaphorodoris lirulatocauda Millen, 1985, Diaphorodoris mitsuii (Baba, 1938), Diaphorodoris papillata Portmann and Sandmeier, 1960, Onchidoris bilamellata (Linnaeus, 1767) Onchidoris muricata (Miiller, 1776) and the Acanthodoris species listed above. Characters for the phylogenetic analysis are taken from the literature and from dissection of specimens when available. Characters were polarized using Bathydoris clavigera Thiele, 1912, Akiodoris salacia Millen, 2005 and Cadlina luteomarginata MacFarland, 1966. The phylogeny supports the monophyly of Adalaria and Onchidoris both situated in a basal clade to Acanthodoris. In addition, Calycidoris is basally situated to the clade containing the monophyletic Diaphorodoris, sister taxa to the clade of Goniodoris plus Okenia. Poi) 214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 The Onchidorididae have traditionally been placed within the Phanerobranchia (Anadoridoidea, Families: Corambidae, Goniodorididae, Polyceridae, Gymnodorididae, Aegiridae and Onchidorididae) (Rudman and Willan 1998). Although it was previously thought that the Phanerobranchia formed a monophyletic clade, recent studies have concluded that it is a para- phyletic group (Wagele and Willan 2000; Valdés 2002; Valles 2002; Fahey and Gosliner 2004). Until recently, the following genera had been placed within the Onchidorididae: Onchidoris, Adalaria, Arctadalaria, Acanthodoris, Doridunculus, Prodoridunculus, Actodoris, Calycidoris, Akiodoris (as Aciodoris) and Diaphorodoris (Millen 1987; Rudman and Willan 1998). However, the most recent phylogeny that includes the Onchidorididae (Millen and Martynov 2005) shows that the traditional Onchidorididae is a paraphyletic group that includes Onchidoris and no other members of the traditional Goniodorididae. Millen and Martynov’s analysis places the remaining Onchidorididae in a separate clade that they name the Akiodorididae. Their analysis also shows that there is one radular character supporting a clade of dorids that includes Acanthodoris, along with three other Onchidorididae genera. Previous work that included a detailed examination of Acanthodoris (Williams and Gosliner 1979), the group examined for the present study, included a review of all Acanthodoris described to date. The present study reviews the literature of all described Acanthodoris species and corroborates the descriptions and illustrations by dissection of additional specimens when available. Species of other Onchidorididae are examined and compared with the literature to provide morphological characters for the phylogenetic analysis. MATERIAL AND METHODS Eleven species of Acanthodoris were included in the analysis. Type material and additional non-type material were obtained from the following sources: The California Academy of Sciences (CAS), the Natural History Museum of Los Angeles County (LACM), the National Museum of Natural History, Smithsonian Institution (USNM), the South African Museum (A) and the Museo Nacional de Ciencias Naturales, Madrid (MNCM). Specimens were drawn under a dissecting microscope using a camera lucida then dissected by dorsal incision. The internal anatomy was drawn as described and then examined either by dissecting and compound or scanning electron microscopes (SEM). External features were examined directly when specimens were available, by photographs, or by literature review. Special attention was given to the reproductive anatomy, as some of these features were infrequently (or cursorily) described in the literature. Table 1 shows the list of character states derived from dissections and from the literature reviewed for the present study. Specimens from each group of Onchidorididae are included: Adalaria, Calycidoris, Diaphorodoris and Onchidoris. Members of the family group Goniodorididae are included for comparison (Goniodoris, Okenia). SPECIES DESCRIPTIONS Family Onchidorididae Gray, 1827 [= Acanthodoridinae P. Fischer, 1881 = Pseudodorididae Eliot, 1910 (n.a.) = Ancylodorididae Thiele, 1926 = Lamellidorididae Pruvot-Fol, 1933 = Villiersiidae Abbott, 1974 (n.a.) = Calycidorididae Roginskaya, 1972] D1AGNosis.— Williams and Gosliner (1979): a moderately depressed body with a simple man- tle edging the foot, elongate rhinophores and soft elongate dorsal papillae. The branchial leaves FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 DAS retract into separate cavities that are interconnected, labial cuticle with rodlets, rachidian absent, two main lateral teeth with a varying number of denticles and a varying number of outer laterals. Genus Acanthodoris Gray, 1850 TYPE SPECIES: Acanthodoris pilosa (Abildgaard in Miiller, 1789) by monotypy. [| = Lamellidoris Alder and Hancock, 1855.] DraGnosis.— Bergh (1880) and O’ Donoghue (1921) provided concise diagnoses of this genus and these are summarized as follows: Body soft depressed; notum thickly covered with short villi: margin of the rhinophore aperture lobed; few branchial plumes, tripinnate, arranged in a circle; head wide, veliform; tentacles short, lobiform. Armature of the labial disc of minute hooks below with projecting thickenings of the cuticle. Naked rachis. Narrow radula. First pleural tooth very large, hooked, few external lateral teeth. Armed penis, long vagina. Diagnosis based on characters examined for the present phylogenetic analysis: Narrow radula with no rachidian tooth; first lateral tooth large and hooked; labial disk with thickened cuticle; jaw rodlets blunt or pointed, single or multiple apices; triaulic reproductive system. Acanthodoris pilosa (Abildgaard in Miiller, 1789) Figs. LA—B, 2-3. Doris pilosa Abildgaard in Miller, 1789:7. Doris stellata Gmelin, 1791:3107. Doris nigricans Fleming, 1820:618. Doris flemingi Forbes, 1838:3. Doris sublaevis Thompson, 1840:87. Doris similis Alder and Hancock, 1842:32. Doris subquadrata Alder and Hancock, 1845:313. Doris sparsa Alder and Hancock, 1846:293. Doris fusca Lovén, 1846:4. Doris rocinella Leach, 1847:268. Doris quadrangulata Jeffreys, 1869:93. Doris bifida Verrill, 1870:406. Doris pilosa stellata Sauvage, 1873:30. Acanthodoris citrina Verrill, 1879:313. Acanthodoris ornata Verrill, 1879:313. Acanthodoris stellata Verrill, 1879:313. Acanthodoris pilosa var. albescens Bergh, 1880:246. Acanthodoris pilosa var. purpurea Bergh, 1880:247. Acanthodoris pilosa var pallida Bergh, 1905:97. Acanthodoris pilosa var. novzealandiae Bergh, 1905:94. MATERIAL EXAMINED.— Neortyre: CASIZ 118891 Dale County, Wales. Intertidal. 1 specimen, 21 mm dissected. August 1992, D. Geiger. HoLoTYPE of Acanthodoris ornata USNM 842118 Eastport Maine, USA. Intertidal. 10 mm. August 1872, Verrill. OTHER MATERIAL: Acanthodoris pilosa CASIZ 074711 Washington County, Maine, USA. No depth data. 2 specimens, 12, 15 mm. July 1977. T. Gosliner. MNCN 15.05/21439 (Onchidoris sparsa) Artedo, Oviedo, Spain. No depth data. | specimen, 10 mm. April 1993, G. Rodriguez. LACM 153980 Cutter Rock, Ketchikan, Alaska. No depth data. 1 specimen, 7 mm dissected, June 1987, S. Millen. DIsTRIBUTION.— North Atlantic (Europe and North America [SeaSlug Forum, accessed 2005]), North Pacific (Aleutian Islands south to Central California [Behrens 1991]). 216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 EXTERNAL MORPHOLOGY.— Numerous descriptions of the external morphology of this species have been provided in the literature (See in particular Abraham 1877; Bergh 1880; O’Donoghue 1921; Pruvot-Fol 1954; Thompson and Brown 1984). See Fig. 1A for a photo of the living animal and Fig. 1B for the original drawing by Abildgaard (1789) See Fig. 3F for a drawing of the ventral anterior surface of a specimen from Wales. DIGESTIVE SYSTEM.— Acanthodoris pilosa shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 3E). The radular sac protrudes noticeably from the bulb, under the esophagus. There is a prominent spherical buccal pump with a longitudinal crease. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have multi-tipped, rounded apices (Fig. 2D). The radular formula is 29 x 3.1.0.1.3 (CASIZ 118891), with all teeth similar to the descriptions of Bergh (1880), O’ Donoghue (1921) and Thompson and Brown (1984). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the three outer lateral teeth are flattened plates with a distinct ridge along the inner edge (Figs. 2A—C). The large innermost tooth is flattened with a pointed hook. The inner margin of each tooth has a large, thickened heel that blends into a large shoulder. The shoulder merges with a prominent ridge that has no distinct denticles, but some irregular thickenings or rounded knobs near the end of the ridge. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate and tubular (Fig. 3C). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a thin, elongate tube. It coils and twists, then narrows into the deferent duct, which folds back once on itself before entering the penial sheath. The penial sheath is elongate and wider than the deferent duct and terminates into a com- mon genital atrium. There are penial hooks at the tip of the penis (Fig. 3D). The vaginal duct 1s short and thinner than the deferent duct. At the distal end, the vagina is much thinner than the penial sheath. The proximal end widens before terminating into the large round bursa copulatrix. From the bursa a long duct connects to the smaller ovoid seminal receptacle. A short uterine duct leads from the base of the bursa into the female gland mass. CENTRAL NERVOUS SYSTEM.— As with other species of Onchidorididae, the cerebral and pleu- ral ganglia are fused together (Fig. 3B). The two pedal ganglia are located below the cerebro-pleu- ral complex and are joined by an elongate commissure. The buccal ganglia are placed under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two rela- tively short nerves. The eyes are stalked at the cerebro-pleural juncture. There are four cerebral nerves leading from each cerebral ganglion including the rhinophoral ganglia, and three large pleu- ral nerves and four small pedal nerves leading from the right and left pleural ganglia. There is a separate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. REMARKS.— The specimens we examined of A. pilosa closely match the descriptions of Bergh FiGure 1. Photos of living animals. A. Acanthodoris pilosa (Abildgaard in Miiller, 1789). Photo by T. Gosliner. B. Acanthodoris pilosa original drawing by Abildgaard (1789). C. Acanthodoris brunnea MacFarland, 1905. Photo by T. Gosliner. D. Acanthodoris hudsoni MacFarland, 1905. Photo by T. Gosliner. E. Acanthodoris lutea MacFarland, 1925. Photo by T. Gosliner. F. Acanthodoris nanaimoensis O'Donoghue, 1921. Photo by W. Lee. G. Acanthodoris pina Marcus and Marcus, 1967. Photo by A. Valdés. H. Acanthodoris rhodoceras Cockerell in Cockerell and Eliot, 1905. Photo by R. Ames. I. Acanthodoris serpentinotus Williams and Gosliner, 1979. Photo by T. Gosliner. J. Acanthodoris planca Fahey and Valdés, 2005. Photo by T. Gosliner. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 17 218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 wai : = eal : br FIGURE 2. Acanthodoris pilosa CASIZ 118891. Radular morphology. A First lateral teeth. Scale = 300 pm. B. Close up of denticles. Scale = 50 um. C. Outer lateral teeth. Scale = 50 um. D. Jaw rodlets. Scale = 50 um. (1880) and subsequent authors (O’ Donoghue 1921, Pruvot-Fol 1954, Thompson and Brown 1984). Only the reproductive system of the animal we examined differed slightly from Pruvot-Fol’s illus- tration. In her drawing, the vaginal duct is narrower and the oviduct is shorter than we found. Picton in Rudman (accessed 2004) reviewed the identity of Acanthodoris pilosa and the appar- ent historical misidentification of the animal currently recognized as A. pilosa. The original description and illustrations of this species (Abildgaard 1789) (see Fig. 1B) do not match the ani- mal now known as A. pilosa, (and recognized as such for over 100 years). That is to say, the ani- mal illustrated by Abildgaard appears to not have a gill. Although the original drawing of A. pilosa does not resemble the presently recognized species and since the type material is lost (could not be located in any of the major natural history muse- ums), Article 75 of the ICZN allows for the designation of a neotype. Thus, we hereby designate Acanthodoris pilosa CASIZ 118891 from Dale County, Wales as the neotype. This designation sta- bilizes the name A. pilosa with contemporary usage. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 29 ot Z FiGURE 3. Acanthodoris pilosa CASIZ 118891. A. Digestive system. B. Cerebro-pleural ganglia. C. Reproductive mor- phology with detail of oviduct, prostate and ampulla morphology. D. Detail of penial spines. E. Buccal bulb. F. Anterior ventral surface. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bg=buccal ganglion, blg=blood gland, bp=buccal pump, cl—3=cerebral ganglia, cg=cerebral ganglion, dd=deferent duct, e=esophagus, fg=female gland mass, ht-heart, od=oviduct, ot=oral tentacle, pg=pedal ganglion, pl—4=pedal nerves, pc=pedal commissure, pg+plg=pleural gan- glion, pll—3=pleural nerves, ppc=parapedal commissure, pr=prostate, r=rhinonophoral nerves, rs=radular sac, sg=salivary gland, sr=seminal receptacle. Scale bars = 1 mm. 220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 i) FIGURE 4. Acanthodoris brunnea CASIZ 105621. Radular morphology. A First lateral teeth. Scale = 200 pm. B. Close up of denticles. Scale = 50 um. C. Jaw rodlets. Scale = 30 um. Acanthodoris brunnea A MacFarland, 1905 Figs. 1C, 4—5. Acanthodoris brunnea MacFarland, 1905:52. MATERIAL EXAMINED.— CASIZ 105621 Monterey Bay, Monterey County, California. 50 m depth. 2 specimens, 14 mm, 15 mm dissected. May 1996, C. Mah. CASIZ 000443 San Juan Passage, Puget Sound, Washington. 146 m depth. 1 specimen, 15 mm. July 1925. No collector information. FIGURE 5. Acanthodoris brunnea CASIZ 105621. A. Anterior ventral sur- DISTRIBUTION.— West coast _ face. B. Reproductive morphology. C. Buccal bulb. Abbreviations: am=ampul- of North America. from Santa — !a, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, Monica. southern California e=esophagus, fg=female gland mass, pn=penial sheath, pr=prostate, rs=radula ; > sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bars = 1 mm. USA to Vancouver Island, Canada (MacFarland 1905; O’Donoghue 1921; Behrens 1991, and present study). EXTERNAL MORPHOLOGY.— MacFarland (MacFarland 1905; MacFarland 1926) gave thor- ough descriptions of the external morphology of this species. The specimens examined for the pres- ent study matched his descriptions and thus there is no additional information to present. See Fig. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 DOM 1 for a photo of the living animal and Fig. 5A for a drawing of the ventral anterior surface of a spec- imen from California. DIGESTIVE SYSTEM.— Acanthodoris brunnea shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus 1s short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. SC). The radular sac protrudes substantially from the bulb, under the esophagus. There is a large, spherical buccal pump with a longitudinal crease as described by MacFarland (1905). The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have multi-tipped, pointed apices (Fig. 4C). The radular formula (CASIZ 105621) is 29 x 4.1.0.1.4, with all teeth as described by MacFarland (1905). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the four outer lateral teeth are simply pointed plates (Figs. 4A and B). The large first tooth is pointed with 13 to 19 denticles on the inner border of the hook. The inner margin of each tooth has a large, thickened heel that narrows into a nearly straight-edged shoulder. The shoulder merges with a prominent inner edge from which the denticles protrude. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate (Fig. 5B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is an elongate, twisted tube. It coils twice before narrow- ing into the deferent duct, which also coils twice. The penial sheath is elongate and wider than the deferent duct and terminates into a common genital atrium. No penial hooks were observed. The vaginal duct is long and much wider than the deferent duct. The vagina was not examined internal- ly and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina is slightly wider than the penial sheath. The proximal end narrows and terminates into the round bursa copulatrix. From the bursa a short duct connects to the smaller pyriform seminal receptacle. The uterine duct leads from the seminal receptacle into the female gland mass. REMARKS.— The specimens we examined of A. brunnea match MacFarland’s descriptions and illustrations (1905, 1925) except for the absence of penial hooks in the specimens we dissect- ed. Williams and Gosliner (1979) noted contradictory information regarding the presence or absence of penial armature in Acanthodoris. These authors observed the difficulty in discerning the armature. Although we did perform microscopic examination of our specimens, in no instance did we observe penial spines in this species. Morphological characters for the present phylogenetic analysis are taken from our dissections and the original literature. Acanthodoris falklandica Eliot, 1907 Figs. 6-7. Acanthodoris falklandica Eliot, 1907:358. MATERIAL EXAMINED.— LACM 153981. Puerto Mott, Chile. 7 m depth. 3 specimens, 6—7 mm dis- sected. January 1995, S. Millen. DIsTRIBUTION.— Falkland Islands (Eliot 1907; Odhner 1926), southern Chile (Schrédl 1996; Schrédl 1997, and present study). EXTERNAL MORPHOLOGY.— Eliot 1907:358, Marcus 1959:60, and Odhner 1926:46 all described the external morphology of this species. The specimens examined for the present study matched their descriptions and thus there is no additional information to present. See Fig. 7A for a drawing of the ventral anterior surface of a specimen from Chile. DIGESTIVE SYSTEM.— Acanthodoris falklandica shares the same general digestive anatomy as 22D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 6. Acanthodoris falklandica LACM 153981. Radular morphology. A First lateral teeth. Scale = 50 um. B. Close up of denticles. Scale = 20 um. C. Outer lateral teeth. Scale = 10 um. D. Jaw rodlets. Scale = 20 um. other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 7C). The radular sac protrudes sub- stantially from an angular exten- sion of the bulb, under the esoph- agus. There is a large, spherical buccal pump with a longitudinal crease as described by Eliot FIGURE 7. Acanthodoris falklandica LACM 153981. A. Anterior ventral (1907). There is a thick plate surface. B. Reproductive morphology. C. Buccal bulb. Abbreviations: a am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=def- with . rodlets at the top of the erent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial opening. The rodlets have multi- sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, tipped, pointed apices (Fig. 6D). v=vagina. Scale bars = 0.5 mm. The radular formula is FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 223 23x2.1.0.1.2 (LACM 153981). There is no rachidian tooth and the large lateral teeth are similar in form while the two outer lateral teeth are simply flat plates with slightly scalloped edges (Figs. 6A and B). The large first tooth has a thickened heel that graduates into a ridge. There are 4-5 denti- cles on the ridge. The tip of each tooth ends in a pointed hook. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is short and tubular (Fig. 7B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is an elongate, thickly coiled tube. It nar- rows into the deferent duct, which loops once, then widens into the penial sheath. The penial sheath is wide, elongate and terminates into a common genital atrium. The penis does not contain penial hooks. The vaginal duct is as wide as the deferent duct. The vagina was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina is large and as wide as the penial sheath. At the proximal end, the vaginal duct narrows before terminating into the large, round bursa copulatrix. From the bursa a long duct connects to the smaller round seminal receptacle. The uterine duct leaves the base of the bursa and enters the female gland mass. REMARKS.— The specimens we examined of 4. falklandica nearly match Eliot’s (1907), Odhner’s (1926) and Marcus’ (1959) descriptions and illustrations of the radula. The specimen we examined has a formula of 23 x 5.1.0.1.5. Eliot’s description of the new species included a radular formula of 33 x 7.1.0.1.7 with up to seven pointed, flattened outer plates. Odhner described a radu- lar formula of 30 = 4.1.0.1.4 and illustrated four pointed outer lateral teeth. Marcus described and illustrated a radular formula of 4—7.1.0.1.4—7. He illustrated the outer lateral teeth as pointed and elongate. The radula formula of the specimens we examined fall within the range described by Marcus (1959). Morphological characters for the present phylogenetic analysis are taken from our dissections and the original literature. Acanthodoris hudsoni MacFarland, 1905 Figs. 1D, 8-9 Acanthodoris hudsoni MacFarland, 1905:51. MATERIAL EXAMINED.— CASIZ 070785 Monastery Beach, Monterey County, California. 8-10 m depth. 2 specimens, 13 mm, 14 mm dissected. August 1978, G McDonald. LACM 71-86 Seal Rocks State Park, Lincoln County, Oregon. Intertidal. 4 specimens, 20-40 mm. August 1971, G. Sphon. LACM 71-87 Neptune State Park, Lane County, Oregon. Intertidal. 6 specimens, 6-35 mm. August 1971, G. Sphon. DISTRIBUTION.— West coast of North America, from San Diego, southern California, USA to Vancouver Island, Canada (MacFarland 1905; O’Donoghue 1921; Behrens 1991, and present study). EXTERNAL MORPHOLOGY.— MacFarland (1905, 1926) and O’Donoghue (1921) gave thor- ough descriptions of the external morphology of this species. The specimens examined for the pres- ent study matched these descriptions and thus there is no additional information to present. See Fig. 1D for a photo of the living animal and Fig. 9A for a drawing of the ventral anterior surface of a specimen from California. DIGESTIVE SYSTEM.— Acanthodoris hudsoni shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is pear-shaped, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 9C). The radular sac protrudes substantially from an angular extension of the bulb, under the esophagus. There is a large, spherical buccal pump 224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 8. Acanthodoris hudsoni CASIZ 070785. Radular morphology. A First lateral teeth. Scale = 100 um. B. Close up of denticles. Scale = 30 um. C. Outer lateral teeth. Scale = 20 um. D. Jaw rodlets. Scale = 50 um. with a longitudinal crease as described by MacFarland (1905). The labial disk frames the open- ing to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have multi-tipped, pointed apices (Fig. 8D). The radular formula is 24 x 5.1.0.1.5 (CASIZ 070785), with all teeth as described by MacFarland. That is, there is no rachidian tooth and the large lat- eral teeth are similar in form while the two outer lateral teeth are simply pointed hooks (Figs. 8A-C). The large first tooth is pointed with 10 denticles on the inner border of the hook. The inner margin of each tooth has a FIGURE 9. Acanthodoris hudsoni CASIZ 070785 A. Anterior ventral sur- face. B. Reproductive morphology. C. Buccal bulb. Abbreviations: am=ampul- la, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagi- na. Scale bars = 1 mm. large, thickened heel that graduates into a ridge. The ridge merges into a shoulder from which the denticles protrude. The ridge continues to the point of the tooth. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 225 REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate (Fig. 9B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is an elongate, coiled tube. It narrows into the deferent duct, which coils once, then widens into the penial sheath. The penial sheath is wide, elongate and terminates into a common genital atrium. We could not confirm the presence of penial hooks. The vaginal duct is wider than the deferent duct. The vagina was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina is not as wide as the penial sheath. The middle portion of the vaginal duct expands substantially and then narrows for the last one-third length before terminating into the round bursa copulatrix. From the bursa a duct connects to the smaller round seminal receptacle. The uterine duct leaves the base of the bursa and enters the female gland mass. REMARKS.— The specimens we examined of A. hudsoni match MacFarland’s descriptions and illustrations (1905, 1925). Morphological characters for the present phylogenetic analysis are taken from our dissections and the original literature. Acanthodoris lutea MacFarland, 1925 Figs. 1E, 10-11. Acanthodoris lutea MacFarland, 1925:60. MATERIAL EXAMINED.— CASIZ 070677 Pigeon Point, San Mateo County, California. No depth data. 3 specimens, 20 mm, 25 mm, 28 mm, dissected. November 1975, G. McDonald. CASIZ 101570 Angel Island, San Francisco Bay, California. No depth data. 1 specimen, 20 mm, dissected. January 1976, Lindberg, Weitbrecht and Gray. LACM 70-74 Naples Reef, Santa Barbara County, California. 15 m depth. 1 specimen, 15 mm. October 1970, C. Swift, W. Stewart, D. Divine. LACM 71-1 Palos Verdes, Los Angeles, California. Intertidal. 2 specimens, 30 mm. January 1971, G. Sphon, E. Marcus, R. Roller, D. Cadien. LACM 73-55 Pirates Cove, California. No depth data. 1 specimen, 21 mm. November, 1973, G. McDonald. LACM 74-30 Palos Verdes, California. Intertidal. 1 specimen, 5 mm. February 1974, G. Sphon, G. Kennedy. LACM 140753 Zuma Beach, California. No depth data. 2 specimens, 22 mm, 25 mm. Spring, 1966. DISTRIBUTION.— West coast of the USA, Marin County, northern California to southern Baja California (Cape Colnett) (MacFarland 1925; Behrens 1991, and present study). EXTERNAL MORPHOLOGY.— MacFarland (1925) provided a thorough description of the exter- nal morphology of this species. The specimens examined for the present study matched his descrip- tions and thus there is no additional information to present. See Fig. 1E for a photo of the living animal and Fig. 11A for a drawing of the ventral anterior surface of a specimen from California. DIGESTIVE SYSTEM.— Acanthodoris lutea shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 11C). The radular sac protrudes from an angular extension of the bulb, under the esophagus. There is a large, spherical buccal pump with a longitu- dinal crease as described by MacFarland (1925). The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have rounded, scalloped apices (Fig. 10D). The radular formula is 34 x 7.1.0.1.7 (CASIZ 070677), with all teeth as described by MacFarland (1925). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the seven outer lateral teeth are pointed plates (Figs. 1|OA—C). The large first tooth is pointed with 2 to 3 rounded denticles on the inner bor- der of the hook. The inner margin of each tooth has a large, thickened heel that narrows into a wide, 226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 10. Acanthodoris lutea CASIZ 070677. Radular morphology. A First lateral teeth. Scale = 200 um. B. Outer lat- eral teeth. Scale = 20 um. C. Close up of denticles. Scale = 50 um. D. Jaw rodlets. Scale = 50 um. nearly straight-edged shoulder. A The shoulder merges with a prominent ridge from which the denticles protrude near the ante- f rior of the ridge. The ridge then J continues smoothly to the point / of the tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampulla is large and bulbous (Fig. 11B). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate rs is an elongate, thick, coiled tube. It narrows into the deferent duct. FiGurRE 11. Acanthodoris lutea CASIZ 070677 A. Anterior ventral surface. B. Reproductive morphology. C. Buccal bulb. Abbreviations: am=ampulla, 3 : : bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, widens into the penial sheath. e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, The penial sheath is tubular, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagi- elongate and terminates into a 4 Scale bars= 1 mm. which twists and coils, then FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 227 common genital atrium. The penis does not contain penial hooks. The vaginal duct is very long and slightly wider than the deferent duct. The vagina was not examined internally and thus the pres- ence of spines or hooks cannot be confirmed. At the distal end, the vagina does not widen from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct widens and then terminates into the round bursa copulatrix. At the base of the bursa a short duct connects to the slightly smaller pyriform seminal receptacle. The thin uterine duct leaves the base of the bursa and enters the female gland mass. REMARKS.— The specimens we examined of 4. /utea match MacFarland’s descriptions and illustrations (1905, 1925). The only difference between our description of the reproductive system and MacFarland’s is in regard to the vaginal duct. MacFarland noted a “glandular portion” of the duct at the proximal end. We observed a widening of the duct at this end; however, we cannot con- firm that it was glandular. Morphological characters for the present phylogenetic analysis are taken from our dissections and the original literature. Acanthodoris nanaimoensis O’ Donoghue, 1921 breswEI2=13: Acanthodoris nanaimoensis O’ Donoghue, 1921:172. Acanthodoris columbina MacFarland, 1926:94. MATERIAL EXAMINED.— CASIZ 076409 Salt Point, Sonoma County, California. 15 m depth. | spec- imen, 20 mm, dissected. October 1962, D. Sullivan. CASIZ 074558 Clayoquot Sound, Vancouver Island, Canada. No depth data. 4 specimens, 10-20 mm dissected. February 1991, G. MacGinitie. CASIZ 068331 Vancouver Island, Canada. No depth or collector data. 1 specimen, 25 mm dissected. LACM 72-103 Neptune National Park, Lincoln County, Oregon. Intertidal. 3 specimens, 10 mm, 20 mm, dissected, 20 mm. August 1972, G Kennedy and G. Sphon. LACM 72-108 Cape Arano State Park, Oregon. Intertidal. 1 specimen, 20 mm. August 1972, G Sphon and G. Kennedy. LACM 71-113 Whale Rock, Del Norte County, California. 10-15 m depth. 5 specimens, 13—30 mm. August 1971, C. Swift. LACM 71-109 Prisoner’s Rock, Trinidad, Humboldt County, California. 8-17 m depth. 1 specimen, 20 mm. July 1971, C. Swift. DISTRIBUTION.— West coast of North America, from Alaska to southern California (Santa Barbara) (MacFarland 1926; Behrens 1991, and present study). EXTERNAL MORPHOLOGY.— O’Donoghue (1921) and MacFarland (1926) gave thorough descriptions of the external morphology of this species. The specimens examined for the present study matched these descriptions and thus there is no additional information to present. See Figure 13A for a drawing of the ventral anterior surface of a specimen from Oregon. DIGESTIVE SYSTEM. — Acanthodoris nanaimoensis shares the same general digestive anato- my as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 13D). The knob-shaped radular sac pro- trudes from the bulb, under the esophagus. There is a large, spherical buccal pump with a longitu- dinal crease as found in other Acanthodoris species. The labial disk frames the opening to the buc- cal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the open- ing. The rodlets have blunt, deeply scalloped apices (Fig. 12D). The radular formula (LACM 72- 103) is 33 x 3.1.0.1.3, with all teeth as described by O’Donoghue (1921). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the five outer lateral teeth are small plates (Figs. 12A—C). The large first tooth is plate-shaped with a pointed hook that has 3-6 tiny pointed denticles on the inner border of the hook. The inner margin of each tooth has a large, 228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 ma) te FIGURE 12. Acanthodoris nanaimc wras , —— yensis LACM 72-103. Radular morphology. A First lateral teeth. Scale = 200 pm. B. Close up of denticles. Scale = 50 um. C. Outer lateral teeth. Scale = 50 um. D. Jaw rodlets. Scale = 50 pm. thickened heel that narrows into a wide, nearly straight-edged shoulder. The shoulder merges with a prominent inner edge from which the denticles protrude. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampulla is long and thin (Fig. 12B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the female gland mass. The prostate is an elongate, thick tube. It narrows into the long, thinner deferent duct, which twists and coils, then widens into the penial sheath. The penial sheath is tubular, elongate and terminates into the common genital atrium. The FIGURE 13. Acanthodoris nanaimoensis LACM 72-103. A. Anterior ventral surface. B. Reproductive morphology. C. Detail of reproductive system duct connections. D. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bars = limm:- penial sheath does not contain penial hooks. The vaginal duct is very long and wider than the def- FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 pipbg) erent duct. The vagina was not examined internally and thus the presence of spines or hooks can- not be confirmed. At the distal end, the vagina widens slightly from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large round bursa copulatrix. From the bursa a long duct connects to the smaller round seminal receptacle. The thin uterine duct leaves the base of the bursa and enters the female gland mass. REMARKS.— The specimens we examined of A. nanaimoensis match O’Donoghue’s (1921) description of the external morphology. However, our observations of the radular morphology dif- fer from O’Donoghue’s illustrations or descriptions. He reported 6—7 lateral teeth with six smaller outer teeth and we observed a formula of 31 x 3.1.0.1.3. O'Donoghue also found no signs of den- ticulation on the ridge of the large main tooth. We found the teeth to be denticulate, with two round- ed outermost denticles and 3—5 smaller, pointed denticles next to these. In addition, O’ Donoghue reported the labial armature as closely packed tiny triangular denticles. However we observed den- ticles having multifid tips as reported by MacFarland (1926) for Acanthodoris columbina. Thus, our specimens match the description and illustrations of the radular teeth as presented by MacFarland (1926). MacFarland (1926) reported that the specimens he examined were devoid of penial hooks and that the penis was very short and blunt. But O’ Donoghue (1921) reported that his specimens had a long, sub-conical penis, armed with minute hooks. Our specimens match MacFarland’s (1926) description of the reproductive organs. O’ Donoghue (1921) did not illustrate the reproductive sys- tem of his specimens, but only described the penial morphology. Steinberg (1963) synonymized Acanthodoris nanaimoensis and A. columbina stating that the variation noted between the specimens examined by O’Donoghue and MacFarland were not enough to justify two separate species. Our study suggests that the two species are synonymous, with A. nanaimoensis having priority. Morphological characters for the present phylogenetic analysis are taken from our dissections and the original literature. Acanthodoris nanega Burn, 1969 Figs. 14-15. Acanthodoris nanega Burn, 1969. MATERIAL EXAMINED.— CASIZ 100574 Victor Harbour, Granite Island, South Australia. 3.6 m depth. 2 specimens, 7-10 mm, 7 mm dissected. August 1994 (no collector identified). DISTRIBUTION.— South coast of Australia (Burn and present study). EXTERNAL MORPHOLOGY.— Burn (1969) described the external morphology of this species. The specimens examined for the present study matched his descriptions and thus there is no addi- tional information to present. Burn stated that the foot was bilabiate in front but the preserved spec- imens we examined did not display this morphology. See Fig. ISA for a drawing of the ventral anterior surface of a specimen from South Australia. DIGESTIVE SYSTEM. — Acanthodoris nanega shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 15C). The long radular sac protrudes substantially from the bulb, under the esophagus. There is a large buccal pump with a longitudinal crease and a pointed apex. The labial disk frames the opening to the buccal bulb and is lined with a thick cuti- cle. There is a thick plate with rodlets at the top of the opening. The rodlets have blunt, deeply scal- 230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 14. Acanthodoris nanega CASIZ 100574. Radular morphology. A First lateral teeth. Scale = 100 um. B. Jaw rodlets. Scale = 50 um. C. Close up of denticles. Scale = 50 um. D. Outer lateral teeth. Scale = 30 um. loped apices (Fig. 14B). The a radular formula is 27 x 5.1.0.1.5 (CASIZ 100574), with all teeth as described by Burn (1969). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the five outer lateral teeth are small, seed-shaped plates (Figs. 14A, C, D). The large first tooth is plate- shaped with a pointed hook that has 3—6 pointed denticles on the inner border of the hook. The inner margin of each tooth has a large, thickened heel that blends into a wide, nearly straight-edged shoulder. The shoulder merges FIGURE 15. Acanthodoris nanega CASIZ 100574. A. Anterior ventral sur- face. B. Reproductive morphology. C. Buccal bulb. Abbreviations: am=ampul- la, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagi- na. Scale bars = 1 mm. into a prominent inner edge from which the denticles protrude. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 733) REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate (Fig. 15B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is an elongate, serpentine tube. It does not narrow into the long deferent duct, which curves once before widening into the penis. The penial sheath is short and bulbous and terminates into the common genital atrium. The penial sheath does not con- tain penial hooks. The vaginal duct is very long and wider than the deferent duct. The vagina was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the dis- tal end, the vagina does not widen from the vaginal duct before it joins with the penis at the com- mon genital atrium. The proximal end of the vaginal duct terminates into the large round bursa cop- ulatrix. From the bursa a duct connects to the slightly smaller ovoid seminal receptacle. The uter- ine duct could not be seen. REMARKS.— The specimens we examined of A. nanega match the external description pro- vided by Burn (1969) except for the bilabiate anterior foot that Burn observed. The radular morphology of the specimens we examined matches the illustrations and descrip- tion by Burn. However, Burn did not examine the reproductive organs of his specimens, so the descriptions provided herein are used for the characters in the present phylogenetic analysis. Burn (1969) noted the similarities between A. nanega and A. globosa Abraham, 1877, from New Zealand. However, the only difference he mentioned to distinguish the two species was a broader lateral tooth base in A. nanega. It is possible that these two species are synonymous. Acanthodoris pina Ey. Marcus and Er. Marcus, 1967 Figs. 1G 16-17. Acanthodoris pina Marcus and Marcus, 1967:201. Acanthodoris stohleri Lance 1968:8, pl. 2, figs. 7-11. MATERIAL EXAMINED.— LACM 153100 Bahia de los Angeles, Baja California. 8.8 m. 1 specimen, 15 mm dissected. March 2000, M. Miller. LACM 140754 Punta Lobos, Sonora, Mexico. No depth data. 6 specimens, 6-30 mm. March 1975, F. and R. Poorman. LACM 67-17 Libertad, Sonora, Mexico. Intertidal. 1 specimen, 20 mm. March 1967, J. McLean. CASIZ 118701 Baja California Norte, Mexico. No depth data. 8 specimens, 5-20 mm. December 1964, W. Farmer. DISTRIBUTION.— Northwest Coast of Mexico, Baja California (Marcus and Marcus 1967, Lance 1968, and present study). EXTERNAL MORPHOLOGY.— Marcus and Marcus (1967) described the external morphology of this species. The specimens examined for the present study matched their descriptions and thus there is no additional information to present. See Fig. 1G for a photo of the living animal and Fig. 17A for a drawing of the ventral anterior surface of a specimen from Baja California. DIGESTIVE SYSTEM. — Acanthodoris pina shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 17D). The radular sac protrudes from the bulb, under the esophagus. There is a large buccal pump with a longitudinal crease as found in other Acanthodoris species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have blunt, deeply scalloped apices (Fig. 16D). The radular formula is 31 x 5.1.0.1.5 (LACM 153100), with all teeth as described by Marcus and Marcus (1967). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the five outer lateral teeth are small plates with a ridge DBP PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 ~*~ FIGURE 16. Acanthodoris pina LACM 153100. Radular morphology. A First lateral teeth. Scale = 200 tum. B. Close up of denticles. Scale = 50 um. C. Outer lateral teeth. Scale = 50 um. D. Jaw rodlets. Scale = 50 um. FIGURE 17. Acanthodoris pina LACM 153100 A. Anterior ventral surface. B. Reproductive morphology. C. Detail of reproductive system duct connections. D. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa_ copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bars = | mm. on the top (Figs. 16A—C). The large first tooth is plate-shaped with a pointed hook that has 3-4 blunt denticles on the inner border of the hook. The inner margin of each tooth has a large, thickened heel that blends into a wide, nearly straight-edged shoulder. The shoul- der merges into a prominent inner edge from which the denticles protrude. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is thick and elongate (Fig. 17B). It branches into the oviduct and the tubular prostate. The thin FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 233, oviduct enters the large female gland mass. The prostate is an elongate tube. It narrows into the long deferent duct, which loops and curves back on itself before widening into the penial sheath. The penial sheath is elongate and wide and terminates into the common genital atrium. There is a glandular portion of the penis located at the distal end. We did not observe penial hooks. The vagi- nal duct is very long, curved and wider than the deferent duct. The vagina was not examined inter- nally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina narrows slightly as compared to the vaginal duct before it joins with the penis at the common gen- ital atrium. The proximal end of the vaginal duct terminates into the round bursa copulatrix. From the bursa a duct connects to the much smaller round seminal receptacle. The thin uterine duct con- nects at the base of the bursa and enters the female gland mass. REMARKS.— The specimens we examined of 4. pina match Marcus and Marcus (1967) description of the external and radular morphology. The only difference between the specimens we examined and those examined by Marcus and Marcus is the presence of a glandular portion of the penis. Marcus and Marcus did not mention this glandular portion, but we observed it in the speci- mens we dissected. Morphological characters of the radular and reproductive systems used in the present phylogenetic analysis are taken from dissection of the specimens noted above. Lance (1968) described several specimens of Acanthodoris collected in Bahia San Luis Gonzaga, Baja California. He did not describe the reproductive system except to note the presence of penial spines. His publication, naming a new species, Acanthodoris stohleri, was preceeded by Marcus and Marcus’ (1967) publication of A. pina sp. nov. by only two months. We examined the type specimens of A. stohleri and concur with Keen’s (1971:828) synonymy of the two species. Acanthodoris rhodoceras Cockerell in Cockerell and Eliot, 1905 Figs. 1H, 18-19. Acanthodoris rhodoceras Cockerell in Cockerell and Eliot, 1905:38. Acanthodoris sp. MacGinitie and MacGinitie, 1949:363. MATERIAL EXAMINED.— CASIZ 068334 Orange County, California. 7 fathoms depth. 1 specimen, 18 mm dissected. May 1934, G. MacGinitie. CASIZ 072355 San Luis Obispo County, California. 6 m depth. 1 specimen, 20 mm. October 1985, D. Behrens. CASIZ 069078 Monterey County, California. No depth data. 1 specimen, 30 mm. April 1972, G. McDonald. CASIZ 169829 San Francisco Marina, California. | m depth. 1 specimen, 4 mm. April 2003, R. Ayres, C. Brown, M. Walton, S. Lattanzio. CASIZ 068322 Duxbury Reef, Marin County, California. No depth data. 2 specimens, 7-10 mm. November 1989, D. Contress. LACM 126371 Duxbury Reef, California. No depth data. 1 specimen, 15 mm. July 1986, R. Willan. LACM 140755 Corona del Mar, California. No depth data. 1 specimen, 15 mm. February 1963, no collector data. LACM 70- 78 Government Point, Santa Barbara County, California. 24 m depth. | specimen, 13 mm. December 1970, C. Swift. LACM 73-32 Kodiak Island, Alaska. Intertidal. | specimen, 12 mm. August 1973, J. McLean. LACM 73-23 Kenai Peninsula, Alaska, 5 fathoms. | specimen, 20 mm. August 1973, J. McLean. LACM 1971-362.6 Beaufort Sea, Arctic Ocean. 360 m. 4 specimens, 10-15 mm. September 1971, R/V Glacier. LACM 66-35 Vancouver, Canada. Intertidal. 1 specimen, 10 mm. June 1966, J. McLean. USNM 576629 San Francisco Bay, California. 46 fms. | specimen, 12 mm. October 1912, no collector data. USNM 791507 Point Loma, California. No depth data. 1 specimen, 8 mm. July 1956, J. Morrison. DISTRIBUTION.— West coast of North America, from Alaska to Baja California (Cockerell and Eliot 1905 and present study). EXTERNAL MORPHOLOGY.— Cockerell and Eliot (1905) and MacFarland (1925) described the external morphology of this species. The specimens examined for the present study matched their descriptions and thus there is no additional information to present. See Fig. 1H for a photo of the living animal and Fig. 19A for a drawing of the ventral anterior surface of a specimen from California. 234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 = > oe fH - | , , - ' y bh — i" " i be i ‘, ® . 7 ‘ oc a Be ° aS oe cai ti Sao 2 By araaE Pe Sen FIGURE 18. Acanthodoris rhodoceras CASIZ 068334. Radular morphology. A First lateral teeth. Scale = 200 um. B. Close up of denticles. Scale = 50 um. C. Outer lateral teeth. Scale = 30 um. D. Jaw rodlets. Scale = 50 um. DIGESTIVE SYSTEM. — Acanthodoris rhodoceras shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stom- ach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 19C). The radular sac protrudes from the bulb, under the esophagus. There is a large ovoid buccal pump with a longitudinal crease as found in other Acanthodoris species. The labial disk frames the opening to the buccal bulb and 1s lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have point- ed, deeply scalloped apices (Fig. 18D). The radular formula is 32 x 3.1.0.1.3 (CASIZ 068334), with all teeth as described by Cockerell and Eliot (1905). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the three or four outer lateral teeth are small plates (Figs. 18A—C). The large first tooth is rather flat with a long pointed hook that has up to 4 pointed denticles on the inner border of the hook. The inner margin of each tooth has a large, thickened heel that blends into a wide shoulder. The shoulder ends and there is a prominent ridge from which the denticles protrude. The ridge then continues to the tip of the hook, but has no further denticulation. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is sausage shaped and elongate (Fig. 19B). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate is an elongate tube. It narrows into FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 o3)5) the long deferent duct, which curves back on itself before widening into the penial sheath. The penial sheath is wide and ter- minates into the common genital atrium. The penis contains minute pointed hooks. The vagi- nal duct is very long and wider than the deferent duct. At the dis- tal end, the vagina widens slight- ly from the vaginal duct as it joins with the penis at the com- mon genital atrium. The proxi- mal end of the vaginal duct widens into a bulb before it ter- minates into the large round bursa copulatrix. From the bursa a duct connects to the smaller Figure 19. Acanthodoris rhodoceras CASIZ 068334. A. Anterior ventral ovoid seminal receptacle. The surface. Scale bar = 1 mm. B. Reproductive morphology. Scale bar = 1 mm. C. Buccal bulb. Scale bar = 1 mm. D. Sketch of penial spines.Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=def- ; erent duct, e=esophagus, fe=female gland mass, ot=oral tentacles, pn=penial REMARKS.— The specimens _ sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, we examined of A. rhodoceras v=vagina. Scale = 30 um. match Cockerell’s (1905) and MacFarland’s (1925) descriptions of the external, radular and reproductive morphology. Morphological characters of the radular and reproductive systems used in the present phylogenet- ic analysis are taken from the literature and corroborated with the dissection of the specimens noted above. long uterine duct connects at the base of the bursa. Acanthodoris serpentinotus Williams and Gosliner, 1979 Fig II. Acanthodoris serpentinotus Williams and Gosliner 1979:216. DISTRIBUTION..— Known only from San Felipe, Baja California, Mexico (Williams and Gosliner 1979). REMARKS.— This species was examined and described in detail by Williams and Gosliner (1979). Morphological characters for the present phylogenetic analysis were taken from the origi- nal literature and from re-examination of the type specimens held at the California Academy of Sciences. Other Acanthodoris species not included in the phylogenetic analyses Acanthodoris armata O’ Donoghue, 1927:4. LocaLity.— Vancouver Island, Canada REMARKS.— O’Donoghue (1927) described the external and radular morphology of this species. The type material could not be located at any of the major natural history museums. No further revision is possible. 236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 Acanthodoris atrogriseata O’ Donoghue, 1927:2. Locatity.— Vancouver Island, Canada REMARKS.— O’Donoghue (1927) described the external and radular morphology of this species. The type material could not be located at any of the major natural history museums. No further revision 1s possible. Acanthodoris caerulescens Bergh, 1880a:252 Locatiry.— Bering Sea, Alaska REMARKS.— Bergh (1880) described the external and radular anatomy of this species. The reproductive anatomy was not illustrated and only partially described. The type material could not be located at any of the major natural history museums, thus no further revision is possible. Acanthodoris globosa Abraham, 1877:262 Locatity.— New Zealand REMARKS.— Abraham (1877) briefly described the external and radular morphology of this species. Eliot (1907) examined and described the external and radular morphology of Abraham’s specimen. He speculated that A. globosa and A. metulifer may be synonymous. The type material could not be located at any of the major natural history museums. Acanthodoris metulifera Bergh, 1905:98 LocaLity.— Tasmania REMARKS.— Bergh (1905) briefly described the external and radular morphology of this species. The type material could not be located at any of the major natural history museums. Marcus and Marcus (1967:203) and Eliot (1907:356) speculated that this species may be a syn- onym of A. globosa Abraham, 1877. Certainly the descriptions of the external morphology are very similar, as are the descriptions and illustrations of the radular morphology. Without examination of the type material, we cannot formally synonymize the two species. A. mollicella Abraham, 1877:262 Loca.tity.— New Zealand REMARKS.— Abraham (1877) described the external morphology and briefly the radular mor- phology of this species. Eliot (1907) examined and described the external and radular morphology of Abraham’s specimen. Williams and Gosliner (1979) provided further anatomical details. Comparisons of the original descriptions and illustrations of 4. mollicella, A. metulifera, A. nanega, and A. globosa causes one to suspect that the four species may be synonymous. However, since the type material of 4. mollicella, A. metulifera, or A. globosa could not be located at any of the major natural history museums, no further revision of these species is possible. Acanthodoris uchidai Baba 1935:119 Locatity.— Akkeshi Bay, Hokkaido, Japan (Baba 1935). REMARKS.— Baba (1935) described the external and radular morphology of this species. However, no description of the reproductive anatomy exists for this species and we were unable to obtain the type material. Limited morphological characters were taken from the original literature for the present phylogenetic analysis. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 esi) Acanthodoris vatheleti Rochebrune and Mabille, 1891:11 Loca.ity.— Cape Horn REMARKS.— This species was only perfunctorily described by Rochebrune and Mabille (Rochebrune and Mabille 1891). Schrédl (2003) reported that 4. vatheleti may be conspecific with A. falklandica Eliot, 1907. However, the type specimen of 4. vatheleti cannot be located and thus no further revision is possible. NEW SPECIES Acanthodoris planca Fahey and Valdés sp. nov. Figs. 1J, 20-21. Acanthodoris sp. Gosliner, 1987:91 MATERIAL EXAMINED.— Holotype: CASIZ 171754 Cove Rock, False Bay, South Africa. No depth data. 20 mm. January 1991, M. Bursey. Paratype: ELM W1647 Igada-Gulu, False Bay, South Africa. No depth data. 18 mm dissected. February 1990, M. Bursey. DISTRIBUTION.— This species has only been recorded from South Africa (present study). EtyMoLoGy.— The specific name p/anca is taken from the Greek word meaning flat-footed. This Acanthodoris species has a prominent flat-footed appearance as it crawls. time FiGure 20. Acanthodoris planca CASIZ 171754. Radular morphology. A First lateral teeth. Scale = 100 tm. B. Close up of denticles. Scale = 50 um. C. Outer lateral teeth. Scale = 50 um. D. Jaw rodlets. Scale = 30 um. 238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 EXTERNAL MORPHOLOGY.— A The body shape of the living ani- mal is oblong and the foot extends beyond the mantle mar- gin (Fig. 1J). The dorsum is cov- ered with elongate papillae that appear short and conical when preserved. The papillae in the preserved specimens are longer at the mantle margin. The oral tentacles are short and blunt. The rhinophores are stout clubs. They angle towards the posterior and have 15—17 lamellae. The gill is broad and spreads to cover the posterior third of the animal. FIGURE 21. Acanthodoris planca CASIZ 171754. A. Anterior ventral sur- There are ten gill leaves that are face. Scale bar = | mm. B. Buccal bulb. C. Reproductive morphology. : aa? Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal bi- and tripinnate. The back- pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tenta- ground color of the dorsum is cles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=sem- pale brown or orange and there is _ inal receptacle, v=vagina. Scale bars = | mm. a darker orange patch of col- oration mid-dorsum. The foot is white with tiny brownish-orange dots. The rhinophores and gill leaves are the same color as the background color, but the rhinophore tips are white. See Fig. 21A for a drawing of the ventral anterior surface of the paratype. DIGESTIVE SYSTEM. — Acanthodoris planca shares the same general digestive anatomy as other Acanthodoris species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is a somewhat ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 21B). The radular sac protrudes from the bulb, under the esophagus. There is an ovoid buccal pump with a longitudinal crease as found in other Acanthodoris species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are elongate, blunt and have multi-apices (Fig. 20D). The radular formula is 32 x 6.1.0.1.6 (CASIZ 171754), with all teeth similar in shape to other Acanthodoris. That is, there is no rachidian tooth and the large lateral teeth are similar in form while the six to seven outer lateral teeth are elongate and have pointed tips (Figs. 20A—C). The large first tooth is plate-shaped with a pointed hook that has approximately 8 blunt denticles on a thickened shoulder along the edge of the hook. The inner margin of each tooth does not have the large, thickened heel that is found on some other Acanthodoris species. The thickened shoulder ends abruptly at the inner margin of the each tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is an elongate tube (Fig. 21C). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate is an elongate, narrow tube with two folds. It has a short constriction where it enters the short, straight deferent duct, which then widens slightly into the penial sheath. The penial sheath is short and terminates into the common genital atrium. No penial hooks were observed. The vaginal duct is very long and wider than the deferent duct. At the distal end, the vagina widens very slightly from the vaginal duct as it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large ovoid bursa FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 2BY. copulatrix. From the bursa a duct connects to the smaller ovoid seminal receptacle. The uterine duct is short and enters at the point where the bursa and seminal receptacle connect. REMARKS.— Acanthodoris planca is externally most similar to the description of A. citrina Verrill, 1879, later synonymized with 4. pilosa by Thompson and Brown 1984 and to A. lutea MacFarland, 1925. All three species are orange yellow or orange and have a large gill that extends the width of the body. All three species have rhinophores that are similar in color as the body. The dorsum of each of these species is densely covered with conical tubercles. Acanthodoris planca has ten branchial leaves and both Verrill (1879) and MacFarland (1925) reported nine for both A. pilosa and A. /utea. The foot color differs between the species. Acanthodoris planca has a white foot with tiny brownish-orange spots, while that of A. /utea is orange yellow. Verrill did not report the foot color for A. citrina but the foot color of A. pilosa (Doris sparsa) was reported by Alder and Hancock (1845) as colorless and by Bergh (1879) as whitish or yellowish. The radular morphology differs between these species. Acanthodoris planca has densely arranged, multifid jaw rodlets and 6—7 elongate, pointed outer lateral teeth. Acanthodoris lutea has rounded, scalloped rodlets and 5-6 flattened, triangular plates “with a slight basal thickening’(MacFarland 1925). Verrill (1879) did not report the radular morphology of A. citrina but Thompson and Brown (1984) provided scanning electron micrographs of A. pilosa. Acanthodoris pilosa has three outer lateral teeth that are flattened plates with rounded, multi-tipped labial rodlets. There are reproductive differences as well. Both the penis and vaginal duct of Acanthodoris planca are shorter and thinner than those found in either of the other two species. Acanthodoris planca has a much shorter deferent duct than found in either A. /utea or A. pilosa. The receptacu- lum seminis duct is much shorter in 4. planca than that of A. pilosa but similar in length to that of A. lutea. The combination of morphological characters identifies Acanthodoris planca as a previously undescribed species. The phylogenetic analysis performed for the present study shows A. planca to be most closely related to 4. falklandica and A. nanega. OTHER SPECIES EXAMINED FOR THE PRESENT PHYLOGENETIC ANALYSIS Genus Adalaria Bergh, 1878 TYPE SPECIES: Adalaria loveni (Alder and Hancock, 1854) Doris loveni Alder and Hancock, 1862:262 REMARKS.— Millen (1987) revised this genus. The characters used in the phylogenetic analy- sis of the present study are taken from this publication and corroborated with our own examina- tions of specimens. Adalaria jannae Millen, 1987 Figs. 22-23. Adalaria jannae Millen 1987:2696 Adalaria sp. Behrens 1980:100; Behrens 1991:50 Onchidoris muricata Roller and Long, 1969:427; McDonald, 1975:531; McDonald and Nybakken, 1978:111; Nybakken, 1978:135; McDonald and Nybakken, 1981:16 Onchidoris sp. Roller and Long 1969:427; McDonald and Nybakken 1980:44; McDonald and Nybakken 1980:16; McDonald 1983:124, 128 Onchidorididae sp. B Lemche in Just and Edmunds 1985:76 240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 22. Adalaria jannae CASIZ 142450. Radular morphology. A First lateral teeth. Scale = 50 um. B. Close up of outer lateral teeth. Scale = 20 tum. C. Close up of denticles. Scale = 20 um. D. Jaw rodlets. Scale = 10 um. MATERIAL EXAMINED.— CASIZ 142450 Whittier, Prince William Sound, Alaska. No depth data. 8 specimens, 4-6 mm, | mm and 5 mm, dissected. August 1999, J. Goddard. LACM 153983 Copper Cove, Vancouver Island, Canada. 1.5 m depth. | specimen, 6 mm. September 1985, S. Millen. DISTRIBUTION.— North Atlantic (Britain to France, and north across Norway, west to Iceland, Greenland and south to Massachusetts) and North Pacific (Bering Sea Alaska, south to California). (Millen 1987, Rudman 2004 and present study). EXTERNAL MORPHOLOGY.— Millen (1987) described the external morphology of this species. The specimens examined for the present study matched her description and thus there is no addi- tional information to present. See Fig. 23A for a drawing of the ventral anterior surface of a spec- imen from Alaska. DIGESTIVE SYSTEM. — Adalaria jannae shares the same general digestive anatomy as other Onchidorididae species (Fig. 3A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached; two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 23B). The radular sac protrudes from the bulb, under the esophagus. There is a large ovoid buccal bulb with a longitudinal crease as found in other FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 24] > Onchidorididae species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have blunt, deeply scalloped apices (Fig. 22D). The radular formula is 37 x 4.1.0.1.4 (CASIZ 142450) with all teeth as described by Millen. That is, there is no rachidian tooth and the large lateral teeth are similar in form while the 4 outer lateral teeth are elongate and multi-tipped (Figs. 22A—C). The large first tooth is beak- shaped with a flattened base and FIGURE 23 (right). Adalaria jannae CASIZ 142450. A. Anterior ventral a pointed hook that has 17 point- surface. Scale bar = 1 mm. B. Buccal bulb. Scale bar = 0.5 mm. C. ed denticles on the inner border Reproductive morphology. Abbreviations: am-ampulla, bb=buccal bulb, : : . bce=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, of the hook. The inner margin of fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, each tooth has a large heel that — rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bar blends into a wide shoulder. The =! mm. shoulder is on top of the promi- nent inner edge from which the denticles protrude. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is an elongate tube (Fig. 23C). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate is a curved, thickened tube. It does not narrow into the deferent duct, which is straight and widens into the penial sheath. The penial sheath is short and terminates into the common genital atrium. The penis does not contain penial hooks. The vaginal duct is very short. The vagina was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina widens enormously from the vaginal duct as it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large round bursa copulatrix. From the bursa a duct connects to the slightly smaller ovoid seminal receptacle. The uterine duct is very short and connects at the point where the receptaculum joins the vaginal duct. CENTRAL NERVOUS SYSTEM.— Millen described the central nervous system of this species as sim- ilar to other Onchidorididae. REMARKS.— Millen (1987) thoroughly described and illustrated the reproductive and radular mor- phology of this species. Our examination of specimens from Alaska concurs with these findings. The only difference we noted in our specimens 1s that the ampulla is longer than the illustration by Millen of specimens from British Columbia, Canada. Morphological characters for the present analyses are taken from the literature and corroborated by our own examinations. Adalaria proxima (Alder and Hancock, 1854) Figs. 24-25. Doris proxima Alder and Hancock, 1854:103. Adalaria albopapillosa Dall, 1871:137. 242 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 24. Adalaria proxima CASIZ 087252. Radular morphology. A. First and outer lateral teeth. Scale = 50 pm. B. Close up of first lateral tooth. Scale = 30 um. C. Jaw rodlets. Scale = 20 um. Adalaria pacifica Bergh, 1880:227. Adalaria virescens Bergh, 1880:83. MATERIAL EXAMINED.— CASIZ 087252 Isles of Shoals, York County, Maine. No depth data. 27 spec- imens, 4-6 mm, 5 mm, dissected. January 1979, T. Gosliner. CASIZ 087250 Rye Harbor, Rockingham County, New Hampshire. No depth data. 1 specimen, dissected. May 1978, T. Gosliner, E. Marcus and A. Kuzirian. CASIZ 070645 Otter Bay, Johnstone Strait, British Columbia. 13—20 m depth. 4 specimens, 12-15 mm, 15 mm, dissected. August 1976, P. Lambert. USNM 804963 Bay of Fundy, Canada. No depth data. | specimen, 10 mm. 1872. LACM 153982 Bamfield, Vancouver Island, Canada. 6 m depth. | specimen, 11 mm. September 1970, S. Millen. DISTRIBUTION.— North Atlantic (Britain to France, and north across Norway, west to Iceland, Greenland and south to Connecticut) and North Pacific (Bering Sea, Alaska, south to California). (Millen 1987, Rudman 2004 and present study). EXTERNAL MORPHOLOGY.— Bergh (1880), Alder and Hancock (1854), Thompson (1984) and FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 243 Millen (1987) described the external morphology of this species. The specimens exam- ined for the present study matched these descriptions and thus there is no additional information to present. See Fig. 25A for a drawing of the ventral anterior surface of a specimen from Maine. DIGESTIVE SYSTEM. — Adalaria proxima shares the same general digestive anato- my as other Onchidorididae species (Fig. 3A). The esopha- gus is short and connects directly to the stomach. The intestine makes a simple, wide FIGURE 25. Adalaria proxima CASIZ 087252. A. Anterior ventral surface. B. Reproductive morphology. C. Detail of reproductive system duct connections. D. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, bc=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral bulb is ovoid, with four large tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, muscles attached, two per side. _st=seminal receptacle, v=vagina. Scale bars = 1 mm. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 25D). The radular sac protrudes from the bulb, under the esophagus. There is a large ovoid buccal bulb with a longitudinal crease as found in other Onchidorididae species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are very small, blunt and have rounded tops (Fig. 24C). The radular formula is 43 x 9.1.1.1.9 (CASIZ 087252) with all teeth as described by Bergh (1880) and Thompson (1984). That is, there is a distinct rachidian tooth and the large lateral teeth are similar in form while the 9 outer lateral teeth have hooked tips (Figs. 24A and B). The large first tooth has a wide base and a long, point- ed hook. The inner margin of each tooth is flattened. A thickened ridge extends from the flat inner margin into the pointed tip of the tooth. There are no denticles on the inner border of the hook The outer edge of the tooth is thick and rounded. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is sausage shaped (Fig. 25B and C). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate is a thick, coiled tube. It does not narrow into the deferent duct, which coils once before curving into the penial sheath. The penial sheath is long and as wide as the deferent duct and it terminates into the common genital atrium. No penial armature was observed. The vaginal duct is short. At the distal end, the vagina widens enormous- ly from the vaginal duct as it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the small ovoid bursa copulatrix. The tiny seminal receptacle is sessile on the vaginal duct. The uterine duct is very short and connects at the vaginal duct. REMARKS.— Bergh (1880), Thompson (1958) and Millen (1987) described and/or illustrated the reproductive and radular morphology of this species. Our examination of specimens from the East and West Coasts of North America concur with these findings. The only difference we noted in our specimens is that the vaginal duct is longer than those illustrated by Millen from Great Britain and curve along the outside of the digestive gland. The buccal 244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 Canada. Morphological characters for the present analyses are taken from the literature and corrob- orated by our own examinations. Genus Calycidoris Abraham, 1876 TyPE SPECIES: Calycidoris guentheri Abraham, 1876 DraGnosis.— Abraham (1876) provided the following diagnosis for the genus: Body depressed, mantle ample extending over the head and the foot, and bearing soft conical papillae. Dorsal tentacles short and laminated, and retractile within sheathless cavities. Branchia simply laminate, forming a cup around the anus, together contained in a common mantle cavity, wall being not completely contractile over them. Oral tentacles represented by a fleshy, laterally extended veil. Odontophore narrow, bearing two bicuspid spines in each row, no central spine, spinous collar or under jaw. The present study confirms this diagnosis. Calycidoris guentheri Abraham, 1876:133 Figs. 26-27. Doris sibirica Aurivillius, 1887:372. FIGURE 26. Calycidoris guentheri CASIZ 086915. Radular morphology. A First lateral teeth. Scale = 300 um. B. Close up of first lateral tooth. Scale = 100 um. C. Close up of outer lateral teeth. Scale = 100 um. D. Jaw rodlets. Scale = 50 um. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 245 FIGURE 27. Calycidoris guentheri CASIZ 086915. A. Digestive system. B. Central nervous system. C. Reproductive morphology. D. Buccal bulb. E. Anterior ventral surface. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bg=buccal ganglion, blg=blood gland, bp=buccal pump, cl—3=cerebral ganglia, cg=cerebral ganglion, dd=deferent duct, e=esophagus, fg=female gland mass, ht=heart, od=oviduct, ot=oral tentacle, pg=pedal ganglion, pl—4=pedal nerves, pc—pedal commissure, pg+plg=pleural ganglion, pll—3=pleural nerves, ppc=parapedal commissure, pr=prostate, r=rhi- nonophoral nerves, rs=radular sac, sg=salivary gland, sr=seminal receptacle. Scale bars = 1 mm. sected. September 1973, unknown collector. CASIZ 086920 Arctic Ocean. 48 m depth. 1 specimen, 14 mm. August 1977, K. Frost. DisTRIBUTION.— Alaska, Arctic Ocean (Roginskaya 1972 and present study). EXTERNAL MORPHOLOGY.— Abraham (1876) and Roginskaya (1972) described the external 246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 morphology of this species. The specimens examined for the present study match their descriptions and thus there is no additional information to present. See Fig. 27E for a drawing of the ventral anterior surface of a specimen from Alaska. DIGESTIVE SYSTEM. — Calycidoris guentheri shares the same general digestive anatomy as other Onchidorididae species (Fig. 27A). The esophagus is short and connects directly to the stom- ach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two long salivary glands at the side of the oral tube, near the radular sac (Fig. 27D). The radular sac protrudes as an oval from the bulb, under the esophagus. There is a large ovoid buccal pump with a longitudinal crease as found in other Onchidorididae species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are elongate and have mostly blunt tips (Fig. 26D). The radular formula (CASIZ 086915) is 29 x 3.1.0.1.3 with all teeth as described by Roginskaya (1972). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the 3 outer lateral teeth are flattened plates (Figs. 26A-—C). The large first tooth has a wide base and a long, pointed hook that has no denticles on the inner border of the hook. The inner margin of each tooth does not have a flattened heel as found in Acanthodoris. There is a ridge that extends along the hooked portion of the tooth. The outer edge of the tooth is long and extends much longer than the inner edge. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is an elongate tube (Fig. 27C). It branches into the oviduct and the tubular prostate. The oviduct enters the large female gland mass. The prostate is a convoluted, thickened tube. It does not nar- row into the deferent duct, which is straight and wider, then it further widens into the penial sheath. The penial sheath is very wide and terminates into the common genital atrium. The penis does not contain penial hooks. The vaginal duct is very short. At the distal end, the vagina widens only slightly from the vaginal duct as it joins with the penis at the common genital atrium. The proxi- mal end of the vaginal duct terminates into the large round bursa copulatrix. From the bursa a thick and very elongate duct connects to the small round seminal receptacle. The uterine duct is long and connects at the base of the bursa. CENTRAL NERVOUS SYSTEM.— As with other species of Onchidorididae, the cerebral and pleu- ral ganglia are fused together (Fig. 27B). The two pedal ganglia are located below the cerebro-pleu- ral complex and are joined by an elongate commissure. The buccal ganglia are placed under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two rela- tively short nerves. The eyes are stalked at the cerebro-pleural juncture. There are four cerebral nerves leading from each cerebral ganglion including the rhinophoral ganglia, and three large pleu- ral nerves and four small pedal nerves leading from the right and left pleural ganglia. There is a separate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. REMARKS.— Phylogenetic characters for the present analyses are taken from the material examined herein, from Millen and Martynov (2005) and from Fahey and Gosliner (2004). Genus Diaphorodoris Iredale and O’ Donoghue, 1923 TyPE SPECIES: Diaphorodoris luteocincta (Sars, 1870) DIAGNosiIs.— Iredale and O’ Donoghue (1923) did not provide a diagnosis for the genus when they established this name in the List of British Nudibranchiate Molluscs. However, Pruvot-Fol (1954) gave a brief comparison of Diaphorodoris with Lamellidoris. She stated that Diaphorodoris differs from Lamellidoris in the body form, which is higher and narrower than the former. The oral FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 247 tentacles are not flattened but are two rounded bumps above the mouth. The tubercles of Diaphorodoris are fewer, long, conical and pointed. Diagnosis based on characters examined for the present phylogenetic analysis: Narrow radula with no rachidian tooth; first lateral tooth large and hooked with multiple pointed denticles; labial disk with thickened cuticle; one outer lateral tooth hooked; jaw rodlets pointed, single apex; triaulic reproductive system. Diaphorodoris luteocincta (Sars, 1870) Figs. 28-29. Doris luteocincta Sars 1870:191. Doris beaumonti Farran 1903:126. Diaphorodoris luteocincta var. alba Portmann and Sandmeier 1960:182. Diaphorodoris luteocincta var. reticulata Portmann and Sandmeier 1960:181. MATERIAL EXAMINED.— CASIZ 087443 Horta Harbor, Azores, Atlantic Ocean. 5—7 m depth. 7 spec- imens, 5—8 mm, 1, 7 mm dissected. November 1992, P. Wirtz. CASIZ 072580 Ilha San Miguel, Azores, Atlantic Ocean. 20 m depth. 6 specimens, 5—10 mm. 1988, T. Gosliner. DISTRIBUTION.— Mediterranean, north and eastern Atlantic (Sars 1870, Eliot 1910; Portmann 1960; Schmekel and Portmann 1982, and present study). EXTERNAL MORPHOLOGY.— Portmann and Sandmeier (1960), Schmekel and Portmann (1982) and Pruvot-Fol (1954) all described the external morphology of this species. The specimens exam- FIGURE 28. Diaphorodoris luteocinta CASIZ 087443. Radular morphology. A First lateral teeth. Scale = 20 um. B. Jaw rodlets. Scale = 10 um. C. Close up of denticles. Scale = 25 um. 248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 ined for the present study matched these descriptions and thus there is no additional information to present. See Fig. 29E for a drawing of the ventral anterior surface of a specimen from the Atlantic Ocean. DIGESTIVE SYSTEM. — Diaphorodoris luteocinta shares the same general digestive anatomy as other Diaphorodoris species (Fig. 29A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two very small sali- vary glands at the side of the oral tube, near the radular sac (Fig. 29C). The trapezoidal radular sac FIGURE 29. Diaphorodoris luteocinta CASIZ 087443. A. Digestive system. B. Cerebro-pleural ganglia. C. Buccal bulb. D. Reproductive morphology. E. Anterior ventral surface. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copula- trix, bg=buccal ganglion, blg=blood gland, bp=buccal pump, cl—3=cerebral ganglia, cg=cerebral ganglion, dd=deferent duct, e=esophagus, fg=female gland mass, ht=heart, od=oviduct, pg=pedal ganglion, pl—4=pedal nerves, pe=pedal com- missure, pg+plg=pleural ganglion, p!1—3=pleural nerves, ppc=parapedal commissure, pr=prostate, mrhinonophoral nerves, rs=radular sac, sg=salivary gland, srseminal receptacle. Scale bars = 1 mm. ; t } | | t FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 249 protrudes from the bulb, under the esophagus. There is a very large buccal pump with a longitudi- nal crease as found in other Diaphorodoris species. The labial disk frames the opening to the buc- cal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the open- ing. The rodlets have multi-tipped apices that are blunt (Fig. 28B). The radular formula (CASIZ 087443) is 28 x 1.1.0.1.1, with all teeth as described by the authors noted above. That is, there is no rachidian tooth and the large first lateral teeth are similar in form while the outer lateral tooth is a small plate with a blunt hook (Figs. 28A and C). The large first tooth is beak-shaped with a long pointed hook that has 13—16 pointed denticles on the inner border of the hook. There is no large, thickened heel or shoulder as found in species of Diaphorodoris. The denticles protrude from the edge of the tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate and tubular (Fig. 29D). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a thick elongate tube. It doesn’t narrow appreciably into the long deferent duct, which curves and twists before widening into the penial sheath. The penial sheath is short and bulbous and terminates into the common genital atrium. We did not observe penial hooks in the penis. The vaginal duct is elongate and as wide as the deferent duct. At the distal end, the vagina does not widen from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large rounded bursa copulatrix. From the bursa a duct connects to the slightly smaller ovoid seminal receptacle. The long uterine duct connects at the point where the seminal receptacle joins the bursa. CENTRAL NERVOUS SYSTEM.— As with other species of Onchidorididae, the cerebral and pleu- ral ganglia are fused together (Fig. 29B). The two pedal ganglia are located below the cerebro-pleu- ral complex and are joined by an elongate commissure. The buccal ganglia are placed under the esophagus, below the central nervous system. They are joined to the cerebral ganglia by two rela- tively short nerves. The eyes are stalked at the cerebro-pleural juncture. There are three cerebral nerves leading from each cerebral ganglion including the rhinophoral ganglia, three large pleural nerves, and three small pedal nerves leading from the right and left pleural ganglia. There is a sep- arate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. REMARKS.— Millen (1985) and Millen and Martynov (2005) described and presented mor- phological characters for phylogenetic analyses of this species. Characters for the present analyses are taken from these publications, from Fahey and Gosliner (2004) and from our dissections of specimens from the Azores. Diaphorodoris lirulatocauda Millen, 1985 Figs. 30-31. Diaphorodoris lirulatocauda Millen, 1985:84. MATERIAL EXAMINED.— CASIZ 076203 Monterey Bay, Monterey California. No depth data. 2 spec- imens, 3 mm, 6 mm dissected. August 1978, T. Gosliner. LACM 153984 Cutter Rock, Ketchikan, Alaska, | specimen, 3 mm, dissected. No depth data. June 1987, S. Millen. DISTRIBUTION.— West coast of North America from Alaska to San Luis Obispo County, California (Millen 1985, and present study). EXTERNAL MORPHOLOGY.— Millen (1985) thoroughly described the external and internal morphology of this species. The specimens examined for the present study matched her descrip- tion and thus there is no additional external information to present. See Fig. 31A for a drawing of the ventral anterior surface of a specimen from California. DIGESTIVE SYSTEM. — Diaphorodoris lirulatocauda shares the same general digestive anato- 250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 = oe a es nS ae FIGURE 30. Diaphorodoris lirulatocauda CASIZ 076203. Radular morphology. A First lateral teeth. Scale = 50 pm. B. Jaw rodlets. Scale = 10 um. C. Close up of outer lateral teeth. Scale = 20 ppm. my as other Diaphorodoris species (see description in Millen 1985). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 31C). The radular FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 251 sac protrudes from the bulb, under the esophagus. There is a large buccal pump with a longi- tudinal crease as found in other Diaphorodoris species. The labi- al disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets have blunt, scalloped apices, some which look like open flowers (Fig. 30B). The radular formula (CASIZ 076203) is 20x FIGURE 31. Diaphorodoris lirulatocauda CASIZ 076203. A. Anterior ven- 1.1.0.1.1, with all teeth as tral surface. B. Reproductive morphology. C. Buccal bulb. Abbreviations: described by Millen (198 5). That am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=def- erent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bars = 0.5 mm. is, there is no rachidian tooth and the large lateral teeth are similar in form while the outer lateral teeth are small plates with a curved pointed hook at the tip (Figs. 30A and C). The large first tooth is beak-shaped with a pointed hook that has 13-16 pointed denticles on the inner border of the hook. There is no large, thickened heel or shoulder as found in species of Acanthodoris. The den- ticles protrude from the edge of the tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is a thick sausage shape (Fig. 31B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a thin elongate tube. It does not narrow into the long deferent duct, which curves back on itself before widening into the penial sheath. The penial sheath is elongate and wide and terminates into the common genital atrium. Penial hooks were not observed. The vaginal duct is relatively short and wider than the deferent duct. The vagi- na was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina does not widen from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the round bursa cop- ulatrix. From the bursa a short duct connects to the slightly smaller ovoid seminal receptacle. The uterine duct could not be seen. REMARKS.— Our examination of specimens from California and Alaska match the description presented by Millen, except that we did not observe penial hooks. Characters for the present phy- logenetic analyses are taken from specimens examined herein and from Millen (1985). Diaphorodoris mitsuii (Baba, 1938) Figs. 32-33. Lamellidoridella mitsuii Baba, 1938:130. Diaphorodoris sp. Gosliner, 1987:92. MATERIAL EXAMINED.— CASIZ 069759 Barracuda Point, East side Pig Island [Tab Island], Madang, Papua New Guinea. 6 m depth. 1 specimen 5 mm, dissected. August 1989, T. Gosliner. DISTRIBUTION.— Japan (Baba 1938; Baba 1949), South Africa (Gosliner 1987), Australia (Willan and Coleman 1984), and Papua New Guinea (present study). JS) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES eis .¢ Fourth Series, Volume 56, No. 20 FIGURE 32. Diaphorodoris mitsuii CASIZ 069759. Radular morphology. A First lateral teeth. Scale = 30 um. B. Jaw rodlets. Scale = 10 um. C. Close up of denticles. Scale = 20 um. EXTERNAL MORPHOLOGY.— Baba (1938) and Millen (1985) described the external morpholo- gy of this species. The specimens examined for the present study matched these descriptions and thus there is no additional infor- mation to present. See Fig. 33. for a drawing of the ventral ante- rior surface of a specimen from Papua New Guinea. DIGESTIVE SYSTEM.— Diaphorodoris mitsuii shares the same general digestive anatomy as other Diaphorodoris species (Fig. 29A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the out- side of the digestive gland. The buccal bulb is ovoid, with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 33C). A FIGURE 33. Diaphorodoris mitsuii CASIZ 069759. A. Anterior ventral sur- face. Scale bar = 1 mm. B. Reproductive morphology. Scale bar = 0.5 mm. C. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copula- trix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=salivary glands, sr=seminal receptacle, v=vagina. Scale bar = 0.5 mm. The radular sac protrudes slightly from the bulb, under the esophagus. There is a large buccal pump ee FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 253 with a longitudinal crease as found in other Diaphorodoris species. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are pointed with a single apex (Fig. 32C). The radular formula (CASIZ 069759) is 20 x 1.1.0.1.1 with all teeth as described by Baba (1938). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the outer lateral tooth is a small plate with a pointed hook at the tip (Figs. 32A and B). The large first tooth is beak-shaped with a pointed hook that has 13—16 pointed denticles on the inner border of the hook. There is no large, thickened heel or shoulder as found in other species of Onchidorididae. The denticles protrude from the edge of the tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- FiGuRE 34. Diaphorodoris papillata CASIZ 099115. Radular morphology. A First lateral teeth. Scale = 20 tm. B. Close up of denticles. Scale = 20 um. C. Jaw rodlets. Scale = 10 um. 254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 oviduct enters the large female A ot gland mass. The prostate is a short tube. It does not narrow appreciably into the short defer- ent duct, which widens into the penial sheath. The penial sheath is short and wide and terminates into the common genital atrium. Penial armature was _ not observed. The vaginal duct is short and the same width as the deferent duct. At the distal end, the vagina does not widen from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large round bursa copu- latrix. From the bursa a duct con- nects to the ovoid seminal recep- FIGURE 35. Diaphorodoris papillata CASIZ 099115. A. Anterior ventral celia tnerne an , th surface. Scale bar = 1 mm. B. Reproductive morphology. Scale bar = 0.5 mm. BIS UNE US © same size as ME ©. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copu- bursa. The uterine duct connects |atrix, bp=buccal pump, dd=deferent duct, e=esophagus, fe=female gland at the base of the bursa. mass, ot=oral tentacles, pn=penial sheath, pr=prostate, rs=radula sac, sg=sali- REMARKS.— Baba (1938) VY@¥ glands, sr=seminal receptacle, v=vagina. Scale bar = 0.5 mm. and Millen (1985) described the external morphology of this species. Baba provided a discussion of the radular morphology. Characters for the present analyses are taken from this literature and from our dissections of specimens from New Guinea. Diaphorodoris papillata Portmann and Sandmeier, 1960 Figs. 34-35. Diaphorodoris papillata Portmann and Sandmeier, 1960:182 MATERIAL EXAMINED.— CASIZ 099115 Isla Tarifa, Strait of Gibraltar, Cadiz, Spain. 30 m depth. 2 specimens, 5 mm, 8 mm dissected. September 1994, T. Gosliner. DISTRIBUTION.— Western Mediterranean (Schmekel and Portmann, 1982 and present study). EXTERNAL MORPHOLOGY.— Several authors described the external morphology of this species (Portmann and Sandmeier 1960; Schmekel and Portmann 1982). The specimens examined for the present study matched these descriptions and thus there is no additional information to pres- ent. See Fig. 35A for a drawing of the ventral anterior surface of a specimen from Spain. DIGESTIVE SYSTEM.— Diaphorodoris papillata shares the same general digestive anatomy as other Diaphorodoris species (Fig. 29A). The esophagus is short and connects directly to the stom- ach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is almost elongate, with four large muscles attached, two per side. There are two small sali- vary glands at the side of the oral tube, near the radular sac (Fig. 35B). The radular sac barely pro- trudes from the bulb, under the esophagus. There is a large protruding buccal pump with a longi- tudinal crease as found in other Diaphorodoris species. The crop is the largest component of the buccal bulb. The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 255 There is a thick plate with rodlets at the top of the opening. The rodlets have pointed apices, some which look like spiked stars (Fig. 34C). The radular formula is 26 x 1.1.0.1.1 (CASIZ 099115), with all teeth as described by Schmekel and Portmann (1982). That is, there is no rachidian tooth and the large lateral teeth are similar in form while the outer lateral tooth is a small plate with a tiny pointed hook at the tip (Figs. 34A and B). The large first tooth is a concave hook with a point- ed hook that has a row 14-16 elongate denticles on the inner border of the hook. There is no large, thickened heel or shoulder as found in species of Acanthodoris. The denticles protrude from the edge of the tooth. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is sausage shaped (Fig. 35C). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a thin elongate tube. It does not narrow appreciably into the short deferent duct, which curves back on itself before widening into the penial sheath. The penial sheath is a wide, elongate tube and terminates into the common genital atrium. There were no penial hooks observed. The vaginal duct is relatively short and wider than the def- erent duct. The vagina was not examined internally and thus the presence of spines or hooks can- not be confirmed. At the distal end, the vagina does not widen from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large ovoid bursa copulatrix. From the bursa a long duct connects to the much smaller ovoid seminal receptacle. The long uterine duct connects at the base of the bursa. REMARKS.— Portmann and Sandmeier (1960) and Schmekel and Portmann (1982) illustrated the radular and reproductive morphology of this species. Our dissections corroborated their find- ings. Characters for the present analyses are taken from the literature and from our own examina- tion of this species. Genus Onchidoris Blainville, 1816 TyPE SPECIES: Onchidoris bilamellata (Linnaeus, 1767) Non Onchidoris (sic Onchidorus) de Blainville, 1816:96. Onchidorus, Onchidora Cuvier, 1830:52. Villiersia d’Orbigny, 1837:XI1. Oncidiodoris, Oncodoris Agassiz, 1846:417. Oicodespina (Okiodespina) Gistel, 1848:399. Proctaporia Morch, 1857:6 (Type species: Proctaporia fusca [Fabricius, 1780] Mérch = Doris pallida). Onchidora Desmarest, 1858:142. Onchidiorus Ferussac, 1882:XX VIII. Ancylodoris Dybowski, 1900:143. Atalodoris \redale and O’ Donoghue, 1923:221. DiaGnosis.— Schmekel and Portmann (1982) provided a diagnosis for this genus. In summa- ry, the body is oval and more broad than high. The notum is spiculose with dorsal papillae and without labial tentacles. The rhinophores are retractable into low sheaths. The gill is not retractable and is arranged in a circle around the anus. The radula is with or without a central tooth, with 2-3 lateral teeth. The jaw consists of a cuticle with weak jaw elements. Onchidoris bilamellata (Linnaeus, 1767) Figs. 36-37. Doris bilamellata Linnaeus, 1767:1083. Doris fusca Miiller, 1776:229. Doris elfortiania de Blainville, 1816:95. 256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 FIGURE 36. Onchidoris bilamellata CASIZ 068343. Radular morphology. A First lateral teeth. Scale = 300 tum. B. Close up of first lateral tooth. Scale = 100 um. C. Close up of outer lateral tooth. Scale = 100 um. D. Jaw rodlets. Scale = 20 um. Onchidoris leachii de Blainville, 1816:450. Doris liturata Moller, 1842:5. Doris liturata Beck, 1842:78. Lamellidoris bilamellata praecedentis Mérch, 1868:204. Lamellidoris n. sp. vel. var. praecedentis Mérch, 1868:204. Lamellidoris bilamellata var. liturata Bergh, 1878b:609 Lamellidoris bilamellata Eliot, 1910:156. Lamellidoris bilamellata var. pacifica Bergh, 1905:101. MATERIAL EXAMINED.— CASIZ 068343 Prince William Sound, Alaska. No depth data. 3 specimens, 18-20 mm, 1, 18 mm dissected. August 1965, S. Haven. CASIZ 081942 Woods Hole, Massachusetts. No depth data. 2 specimens, 10-12 mm. September 1906, F. MacFarland. CASIZ 068344 Charleston Yacht Harbor, Coos County, Oregon. No depth data. 3 specimens, 27-30 mm. February 1963, L. Andrews. CASIZ 060106 Sonoma County Coast, California. 50-115 fathoms. 1 specimen, 20 mm. February 1950, L. Miles. LACM 86-180 St. Andrews Bay, Fife, Scotland. Intertidal. 7 specimens, 7-12 mm. September 1986, J. McLean. USNM 574284 (Onchidoris liturata) Station 1182, Greenland. No depth data. 1 specimen, 14 mm. No collection date. J. Blake. DISTRIBUTION.— North Atlantic (Britain to France, and north across Norway, west to Iceland, Greenland and south to Connecticut) and North Pacific (Bering Sea, Alaska, south to California). FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 Dy) FIGURE 37. Onchidoris bilamellata CASIZ 068343. A. Digestive system. B. Cerebro-pleural ganglia. C. Reproductive morphology. D. Buccal bulb. E. Anterior ventral surface. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bg=buccal ganglion, blg=blood gland, bp=buccal pump, cl—3=cerebral ganglia, cg=cerebral ganglion, dd=deferent duct, e=esophagus, fg=female gland mass, ht=heart, od=oviduct, ot=oral tentacle, pg=pedal ganglion, pl—4=pedal nerves, pc=pedal commissure, pg+plg=pleural ganglion, pll1—3=pleural nerves, ppc=parapedal commissure, pr=prostate, r=rhi- nonophoral nerves, rs=radular sac, sg=salivary gland, sr=seminal receptacle. Scale bars = 1 mm. 258 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 EXTERNAL MORPHOLOGY.— The external morphology of this species has been described by several authors (Bergh 1878; Eliot 1910). The specimens examined for the present study matched these descriptions and thus there is no additional information to present. See Fig. 37E for a draw- ing of the ventral anterior surface of a specimen from Alaska. DIGESTIVE SYSTEM.— Onchidoris bilamellata shares the same general digestive anatomy as other Onchidorididae (Fig. 37A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal bulb is a flattened oval with four large muscles attached, two per side. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 37D). The radular sac protrudes from the bulb, under the esophagus. There is a large protruding buccal pump that has a fold (as found in other Onchidoris species) The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are elongate with single apices (Fig. 36D). The radular formula is 23 x 1.1.1.1 (CASIZ 068343), with all teeth as described by Thompson and Brown (1984). That is, there is a small vestigial rachidian tooth and the large lateral teeth are similar in form while the outer lateral tooth is a thin, curved hook (Figs. 36A-C). The large first tooth is plate-shaped with a long pointed hook that has no denticles on the inner border of the hook. There is a large, heel or shoulder as found in species of Acanthodoris and Onchidoris. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la is elongate (Fig. 37C). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a thin, curved elongate tube. It narrows slight- ly into the long deferent duct, which has one curve before it widens into the penial sheath. The penial sheath is short and wider than the deferent duct. It terminates into the common genital atri- um. The penis does not contain penial hooks. The vaginal duct is relatively short compared to other Onchidoris and wider than the deferent duct. The vagina was not examined internally and thus the presence of spines or hooks cannot be confirmed. At the distal end, the vagina slightly widens from the vaginal duct before it joins the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the large round bursa copulatrix. From the bursa a duct connects to the much smaller ovoid seminal receptacle. The short uterine duct connects below the point at which the receptaculum joins the oviduct. CENTRAL NERVOUS SYSTEM.— As with other Onchidorididae, the cerebral and pleural ganglia are fused together (Fig. 37B). The two pedal ganglia are located below the cerebro-pleural com- plex and are joined by an elongate commissure. The buccal ganglia are placed under the esopha- gus, below the central nervous system. They are joined to the cerebral ganglia by two relatively short nerves. The eyes are stalked at the cerebro-pleural juncture. There are four cerebral nerves leading from each cerebral ganglion including the rhinophoral ganglia, and three large pleural nerves and three small pedal nerves leading from the right and left pleural ganglia. There is a sep- arate abdominal ganglion on the right side of the visceral loop. Gastro-esophageal, rhinophoral and optical ganglia are present. REMARKS.— Characters for the present phylogenetic analyses are taken from our examination of material from Massachusetts, Alaska, Scotland and Greenland. Onchidoris muricata (Miiller, 1776) Figs. 38-39. Doris muricata Miller, 1776:229. Doris aspera Alder and Hancock, 1842:32. Doris diaphana Alder and Hancock, 1845:313. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 259 = or od FIGURE 38. Onchidoris muricata CASIZ 087251. Radular morphology. A First lateral teeth. Scale = 50 tum. B. Close up of denticles. Scale = 20 um. C. Close up of outer lateral tooth. Scale = 20 um. D. Jaw rodlets. Scale = 20 um. Doris pallida Agassiz, 1850:229. Lamellidoris hystricina Bergh, 1878b:605. Lamellidoris varians Bergh, 1878:613. Onchidoris muricata Nybakken and McDonald, 1980:44. Onchidoris sp. Goddard, 1984:152. (See Millen 1985 for a complete synonymy and discussion). MATERIAL EXAMINED.— CASIZ 087251 Isles of Shoals, York County, Maine. 8-15 m depth, 15 spec- imens, 8 mm dissected. January 1979, T. Gosliner. CASIZ 067705 Friday Harbor, Puget Sound, Washington. 0 m depth. 3 specimens, 3—5 mm. July 1950, P. Illg. USNM 574287 Provincetown, Massachusetts. No depth or collector data. 3 specimens, 15—20 mm. No collection date, J. Blake. USNM 804964 Bay of Fundy, Canada. No depth or collector data. 3 specimens, 6-12 mm. 1872. USNM 810779 Massachusetts Bay, Massachusetts. No depth or collector data. 1 specimen, 5 mm. 1878. USNM 810783 Bay of Fundy, Canada. No depth or col- lector data. 3 specimens, 5-10 mm. 1872. USNM 827473 (Lamellidoris aspera |sic]) Georges Bank, Massachusetts. 82 m depth. 1 specimen, 4 mm. November 1982, Battelle. USNM 827901 (Lamellidoris aspera |sic]) Georges Bank, Massachusetts. 82 m depth. 1 specimen, 4 mm. November 1982, Battelle. LACM 153985 Cutter Rock, Ketchikan County Alaska. No depth data. 1 specimen, 7 mm. June 1987, S. Millen. DISTRIBUTION.— North Atlantic (Britain to France, and north across Norway, west to Iceland, Greenland and south to Connecticut) and North Pacific (Bering Sea Alaska, south to California). (Rudman 2004 and present study). EXTERNAL MORPHOLOGY.— Several authors have described the external morphology of this 260 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 species, by most recently by A B Millen (1985). The specimens examined for the present study matched her description and thus there is no additional informa- tion to present. See Fig. 39A for a drawing of the ventral anterior surface of a specimen from Maine. DIGESTIVE SYSTEM.— On- chidoris muricata shares the same general digestive anatomy as other Onchidorididae (Fig. 37A). The esophagus is short and connects directly to the stomach. The intestine makes a simple, wide curve along the outside of the digestive gland. The buccal FIGURE 39. Onchidoris muricata CASIZ 087251. A. Anterior ventral sur- bulb is ovoid, with four large face. B. Reproductive morphology. C. Detail of reproductive system duct con- muscles attached, two per side. nections. D. Buccal bulb. Abbreviations: am=ampulla, bb=buccal bulb, be=bursa copulatrix, bp=buccal pump, dd=deferent duct, e=esophagus, fg=female gland mass, pn=penial sheath, pr=prostate, rs=radula sac, sg=sali- vary glands, sr=seminal receptacle, v=vagina. Scale bars = 1 mm. There are two small salivary glands at the side of the oral tube, near the radular sac (Fig. 37D). The radular sac protrudes noticeably from the bulb, under the esophagus. There is a promi- nent buccal bulb that has a longitudinal crease (as found in other Onchidoris species). The labial disk frames the opening to the buccal bulb and is lined with a thick cuticle. There is a thick plate with rodlets at the top of the opening. The rodlets are very small and rounded (Fig. 38D). The radu- lar formula is 32 x 1.1.1.1 (CASIZ 087251), with all teeth as described and illustrated by Millen (1985). That is, there is a large rachidian tooth that has parallel distinct edges and the first lateral teeth are all similar in form. The outer lateral tooth is a shallow plate with a thin hook (Figs. 38A—C). The large first tooth is has a long pointed hook that has up to 16 pointed denticles on the inner border of the hook. There is a curved heel or shoulder at the inner tooth edge that blends into the ridge from which the denticles protrude. The ridge extends to the tip of the hook devoid of den- ticles. At the base of the ridge there is a rounded protrusion. REPRODUCTIVE SYSTEM.— The hermaphroditic duct enters the ampulla terminally. The ampul- la resembles a curved sausage (Fig. 39B). It branches into the oviduct and the tubular prostate. The thin oviduct enters the large female gland mass. The prostate is a short tube. It narrows slightly into the short deferent duct, which has one curve before it widens into the penial sheath. The penial sheath is sausage shaped and only slightly wider than the deferent duct. It terminates into the com- mon genital atrium. The penis does not contain penial hooks. The vaginal duct is relatively short and wider than the deferent duct. At the distal end, the vagina slightly widens from the vaginal duct before it joins with the penis at the common genital atrium. The proximal end of the vaginal duct terminates into the small round bursa copulatrix. From the bursa a duct connects to the slightly smaller ovoid seminal receptacle. The uterine duct is short and connects at the point where the bursa and receptaculum join. REMARKS.— Millen (1985) reviewed this species and presented a thorough discussion of the synonymies. The material we examined for the present study closely matches the descriptions and ee —- FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 261 illustrations presented by Millen. Characters for the present phylogenetic analysis are taken from Millen and are corroborated with the specimens examined herein. Villiersia scutigera d’Orbigny, 1837:15 REMARKS.— Pruvot-Fol (1954) synonymized Villiersia with Lamellidoris. The description of Villiersia scutigera d’Orbigny, 1837 was limited to external characters. Pruvot-Fol (1954) wrote that because of this, the true identity of this species could not be con- firmed. Although the illustrations provided by d’Orbigny of the body shape, gill arrangement, notal papillae and ventral surface bear some resemblance to a few species of Onchidoris, the coloration of Villiersia scutigera most closely matches that of Onchidoris bouvieri as described by Schmekel and Portmann (1982). PHYLOGENETIC ANALYSIS Onchidorididae taxa included in the present analysis are Adalaria jannae Millen, 1987, A. loveni (Alder and Hancock, 1862), A. proxima (Alder and Hancock, 1854), Calycidoris guentheri Abraham, 1876, Diaphorodoris lirulatocauda Millen, 1985, D. luteocinta (Sars, 1870), D. mitsuii (Baba, 1938), D. papillata Portmann and Sandmeier, 1960), Goniodoris nodosa (Thompson and Brown, 1984) and Okenia mediterranea (Ihering, 1886). Characters were polarized with the out- group taxa Bathydoris clavigera Thiele, 1912, which, according to Valdés (2002), is a basally sit- uated species of Bathydoris, the closest sister clade to the Phanerobranchia and Cryptobranchia. Other outgroup taxa included in the present analysis are Cadlina luteomarginata MacFarland 1966, a species of the most basal clade of the Chromodorididae (Gosliner and Johnson 1999), and Akiodoris salacia Millen and Martynov, 2005, which according to Millen and Martynov (2005) is the most basally situated taxon in the sister clade to the remaining Onchidorididae. Character states were taken from both the literature and from examinations of museum material when available. Table 1 summarizes the information for the outgroup taxa and the Onchidorididae (other than Acanthodoris) included in the analysis. Morphological data were organized using MacClade, ver 3 (Maddison and Maddison 2000). Synapomorphies were examined using MacClade and the charac- ter-trace option, using the majority rule tree from PAUP analyses. CHARACTERS.— The following characters were considered for use in the analyses of Onchidorididae. The character states are indicated as follows: 0: the presumed plesiomorphic con- dition; 1,2: apomorphic condition. For character states that are not applicable, ‘“-” is used and for missing data, “?” is used. The distribution of plesiomorphic and apomorphic character states is pre- sented in Table 2. Character states for the taxa examined for the present study can be found in Table 2 (see Appendix A for the full data matrix). “Outgroup taxa” refers to the combination of Bathydoris clavigera, Cadlina luteomarginata and Akiodoris salacia. “Ingroup taxa” refers to all Acanthodoris plus Adalaria, Calycidoris, Diaphorodoris, Onchidoris, Goniodoris and Okenia. 1. Dorsal tubercle length: The dorsal tubercles are short (0) in Bathydoris, Adalaria, Diaphorodoris luteocinta, D. mitsuii, Goniodoris, Onchidoris, and Acanthodoris pilosa. In the remaining taxa, the tubercles are long (1). 2. Dorsal tubercle shape: The dorsal tubercles are pointed (0) in nearly all taxa included. But Adalaria species, Onchidoris bilamellata, Akiodoris, Cadlina and all Diaphorodoris except D. luteocincta have round- ed tubercles (1). 3. Dorsal tubercle distribution: The tubercles are sparsely distributed (0) in two species of Diaphorodoris, Goniodoris, Okenia, Acanthodoris falklandica, A. nanega, Bathydoris and Akiodoris. They are densely distributed (1) in the remaining taxa. 262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 TABLE |. Outgroup taxa and Onchidorididae species included in the present analysis, literature source and material examined. Species Literature Source Additional material examined (# specimens) Bathydoris clavigera Thiele, 1912 Marcus and Marcus 1962; Valdés 2002 Cadlina luteomarginata MacFarland, 1966 MacFarland 1966 Akiodoris salacia Millen and Martynov, 2005 Millen and Martynov 2005 Adalaria jannae Millen, 1987 Millen 1987 CASIZ 142450 (8) LACM 153983 (1) Adalaria loveni (Alder and Hancock, 1862) Thompson and Brown 1984, Millen 1987 Adalaria proxima (Alder and Hancock, 1854) Alder and Hancock 1854, CASIZ 087252 (27) Thompson 1984 CASIZ 087250 (1) CASIZ 070645 (4) LACM 153982 (1) USNM 804963 (1) Calycidoris giintheri Abraham, 1876 Abraham 1876, 1877; CASIZ 086915 (1) Roginskaya 1972; CASIZ 086920 (1) Valdés 2002 Diaphorodoris lirulatocauda Millen, 1985 Millen 1985 CASIZ 076203 (2) LACM 153984 (1) Diaphorodoris luteocinta (Sars,1870) Schmekel and Portmann 1982; CASIZ 072580 (6) Pruvot-Fol 1954; CASIZ 087443 (7) Valdés 2002 Diaphorodoris mitsuii Baba, 1938 Baba 1938 CASIZ 069759 (1) Diaphorodoris papillata Portmann &Sandmeier, 1960 Portmann &Sandmeier 1960 CASIZ 099115 (2) Goniodoris nodosa (Thompson and Brown, 1984) Thompson and Brown 1984 Okenia mediterranea (lhering, 1886) Schmekel and Portmann 1982; Valdés and Ortea 1995 4. Rhinophore length: In Bathydoris and most of the ingroup, the rhinophores are elongate (0). But in the remaining taxa, the rhinophores are short (1). 5. Rhinophore sheaths: In Bathydoris, Goniodoris and Okenia there are no rhinophore sheaths (0). In all other taxa the sheaths are not contractile (1). In Cadlina the sheaths are contractile (2). 6. Body Shape: In all taxa the body is ovoid (0) but in Cadlina the body is more elongate than ovoid (1). 7. Head shape: In all taxa, the head is veliform (0), but in Okenia and Cadlina the head is rounded (1). 8. Mantle margin: in Bathydoris and Okenia there is no mantle margin (0). But in Akiodoris, Diaphorodoris, and Cadlina the mantle margin is narrow (1) and in the remaining taxa the margin is wide (2). 9. Posterior mantle: In most taxa the posterior mantle is not bilobed (0), whereas in Goniodoris it is bilobed (1). 10. Gill protection: In Bathydoris and all species of Adalaria there is no single gill pocket (0). In Cadlina the gill is retractile into a pocket (1) and in the remaining taxa the gill is contractile but not retractile into an opening (2). 11. Gill notal sculpture: In Bathydoris, Cadlina, Goniodoris, Okenia and all species of Diaphorodoris ———— FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 263 TABLE 2. Characters and states examined for the phylogenetic analysis of Acanthodoris. 0 = presumed plesiomorphic; 1,2 = apomorphic conditions. Character States : elongate;1: short : pointed; 1: round : dense; 1: sparse Dorsal tubercles Dorsal tubercles Dorsal tubercles Rhinophores : elongate; 1: short Rhinophore sheaths : none; 1: non contractile; 2: contractile Body shape : ovoid; 1: elongate Head : veliform; 1: rounded none; 1: narrow; 2: wide : entire; 1: bilobed : no pocket; 1: retractile; 2: contractile : absent; 1: present : absent; 1: present : absent; 1: present : absent; 1: present : absent; 1: present : present; 1: absent : labial armature; 1: smooth, thin : single apex; 1: multi apices many; 1: few : first differs only; 1: more differ from outers : cusp with or w/o denticles; 1: beak shaped : both sides; 1: inner only; 2: absent many; |: few : elongate/hamate; 1: elongate/oval; 2: wide/rectangular : multiple; 1: one Mantle margin Posterior mantle Gill protection Gill notal sculpture Mid dorsal crest Caudal crest Buccal pump Peripheral muscles Rachidian Lip disk Jaw rodlets Number of teeth 1/2 row Number inner laterals Lateral teeth Denticulation Denticles Outer lateral teeth Outer lateral teeth Ampulla : elongate/narrow; 1: short/wide Receptaculum seminis absent; 1: present RS ducts : one duct; 1: two ducts RS duct insertion Oviduct connection Penial spines Vagina size Cerebro-pleural ganglia Body color Dorsal spots : on vagina; 1: on uterine duct; 2: base of bursa : into female gland mass; 1: into receptaculum : absent; 1: present : narrower than penial sheath; 1: same as penial sheath; 2: wider : separate; 1: fused : white/yellow; 1: other : absent; 1: present there are no notal structures such as tubercles within the gill ring (0). But in all remaining taxa, there are tuber- cles or other low structures within the gill circle (1). 12. Mid dorsal crest: In Goniodoris and Okenia there is a mid-dorsal crest (1) but in the remaining taxa, this crest is absent (0). 13. Caudal crest: In Diaphorodoris, Goniodoris and Okenia the anterior surface of the foot has a crest along the midline (1). This crest is missing on all other taxa (0). 14. Buccal Pump: The buccal pump is absent (0) in Bathydoris and Cadlina. But in the remaining taxa the pump is present (1). 15. Peripheral muscles: There is no band of longitudinal muscle on the pharynx (0) of Bathydoris, Akiodoris or Cadlina. The remaining taxa all have a band of muscle (1). 16. Rachidian: There is a rachidian tooth present in the outgroup taxa, species of Adalaria and Onchidoris (0). But the remaining taxa have no rachidian (1). 17. Lip disk: In most taxa, the lip disk contains armature (0), but in Akiodoris and species of Adalaria the lip disk is smooth and thin (1). 18. Jaw rodlets: In species with jaw rodlets, some have a single apex (0) such as Bathydoris, Calycidoris, 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 Diaphorodoris mitsuii, D. papillata, Goniodoris, Okenia and Onchidoris. In Acanthodoris the rodlets have multiple apices (1). 19. Number of teeth per half row: \n the outgroup taxa there are many teeth in each row (0), but in the remaining taxa there are few teeth (1). 20. Number of inner lateral teeth: In Akiodoris and Cadlina the first lateral tooth is the only tooth with that particular shape (0). In all other taxa, there is more than one inner lateral tooth that differs from the outer lateral teeth (1). 21. Lateral teeth shape: In most taxa examined, the innermost lateral tooth is a cusp with or without den- ticles (1). In the outgroup taxa the tooth is beak-shaped (0). 22. Tooth denticulation: In the outgroup taxa both sides of each tooth have denticles (0). In all remain- ing taxa except Calycidoris the teeth are denticulate on the inner side only. Calycidoris has no denticles (2). This character was omitted from the final analysis when interspecific variation was noted, seemingly depend- ent on the maturity of the specimen. 23. Denticle number: In the outgroup taxa, Acanthodoris hudsoni, Adalaria jannae, Goniodoris, Okenia and Diaphorodoris the denticles are numerous (0). However, in all remaining Acanthodoris species, Onchidoris and Akiodoris there are few denticles (1). 24. Outer lateral teeth shape: In those species having outer lateral teeth, there are elongate/hamate teeth (0) in Bathydoris and Cadlina. In all other species except Akiodoris there are elongate/oval teeth (1). In Akiodoris the outer lateral teeth are wide rectangles (2). 25. Outer later teeth number: In the outgroup taxa and most species examined, there are numerous outer lateral teeth (0). But in Diaphorodoris, Okenia, Acanthodoris serpentinotus and Onchidoris there is only one outer lateral tooth (1). 26. Ampulla length: In the outgroup taxa , Okenia and most Acanthodoris, the ampulla is elongate/nar- row (0). In Acanthodoris falklandica, A. serpentinotus and Onchidoris the remaining taxa it is short/wide (1). 27. Receptaculum seminis: In Bathydoris the receptaculum is absent (0). But in the remaining taxa the receptaculum is present (1). 28. Receptaculum duct: In all outgroup taxa and most other species included, the receptaculum is con- nected by a single duct (0). But in Adalaria and Onchidoris there are two ducts (1). 29. Receptaculum insertion: In Adalaria, Onchidoris, Akiodoris and Bathydoris the receptaculum duct inserts on the vaginal duct (0). In Goniodoris, Okenia and Cadlina the duct inserts on the uterine duct (1) and in the remaining taxa, the duct inserts at the base of the bursa (1). 30. Oviduct: In most taxa examined the oviduct connects to the female gland mass (0). In Adalaria and Onchidoris the oviduct connects to the receptaculum (1). 31. Penial spines: In Bathydoris, Adalaria and Onchidoris, penial spines are absent (0), but previous authors have observed spines in all other species (1). 32. Vagina size: In nearly all taxa examined, the vagina is narrower than the penial sheath (0). In Diaphorodoris mitsuii, Okenia and Onchidoris muricata the penial sheath and vagina are the same width (1). In Acanthodoris falklandica the vagina is wider than the penial sheath (2). 33. Cerebro-pleural ganglia: In Bathydoris and Adalaria the ganglia are separate (0). In all other taxa the ganglia are fused (1). 34. Body color: In nearly all taxa included in the analysis, the ground color is white or pale yellow (0). But in many Acanthodoris species the ground color is not white, but orange or brown or reddish (1). 35. Dorsum spots: In most taxa, dorsum pigment spots are absent (0). In Goniodoris, Okenia, Acanthodoris brunnea, A. lutea and A. pina spots are often found on the dorsum (1). RESULTS.— One most parsimonious tree was obtained with a length of 94 steps and had a con- sistency index (CI) of 0.45, a retention index (RI) of 0.68 and a homoplasy index (HI) of 0.55. The tree is shown in Fig. 40 with the character numbers and character reversals. The bold numbers indi- cate reversals. The trees indicated that Adalaria, Onchidoris, Diaphorodoris and Acanthodoris each form a monophyletic clade. Of the outgroup taxa included in the analysis, Bathydoris clavig- era is most basally situated, with Cadlina luteomarginata found as the sister species to the | i ! g FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 265 Onchidorididae plus Akiodorididae (Millen and Martynov 2005). Akiodoris salacia is sister taxon to the Onchidorididae. Within the ingroup taxa, the clade containing Adalaria and Onchidoris is the most basally sit- uated clade, with Onchidoris in the more basal position. Okenia mediterranea and Goniodoris nodosa form a clade that is basally situated to Diaphorodoris and Calycidoris guentheri is the most basally situated species of this clade. The clade containing all Acanthodoris species shows that the most basally situated species are Acanthodoris planca, A. nanega and A. falklandica, all from the Southern Hemisphere. The remaining species found in the Northern Hemisphere form a separate clade. Bremer support values (shown in Fig. 40, numbers below the branches) show poor support for most of the clades with the exception of the outgroups and the clade containing Adalaria plus Onchidoris and the clade of Goniodoris plus Okenia. Analyses were also performed using different outgroup taxa such as including Okenia and Goniodoris with Bathydoris, Cadlina and Akiodoris. Okenia and Goniodoris were also used as the only outgroup taxa, with Bathydoris, Cadlina and Akiodoris included in the ingroup. Results from both these trials produced the same single most parsimonious tree (tree not shown). Tree scores were the same in all trials. In the single tree that was produced by the outgroup trials, Bathydoris, Cadlina and Akiodoris group together in a derived sister clade to Adalaria and Onchidoris. The clade that contains these groups form the sister clade to Acanthodoris. Calycidoris 1s basally situated to both clades and Diaphorodoris forms a clade at the base of all these taxa. DISCUSSION Results from the phylogenetic analysis support the monophyly of Acanthodoris, Onchidoris, Adalaria and Diaphorodoris. Additionally, the placement of Adalaria and Onchidoris in a sister clade to Acanthodoris, Diaphorodoris, Goniodoris and Calycidoris agrees with the findings of Millen and Martynov (2005). Our findings also agree with Millen and Martynov’s results showing Cadlina in a basal position relative to Akiodoris. The synapomorphies for Acanthodoris are (18) jaw rodlets with multi-apices and (23) few den- ticles on the lateral teeth. Characters that unite members of the traditional Onchidoridae include (5) a contractile rhinophore sheath, (11) presence of notal sculpture within the gill circle, (14) presence of a buccal pump, (22) denticles on the inner side of the lateral teeth and (24) elongate/hamate outer lateral teeth. Our results support the findings of Millen (1985) with regard to the relationship among Diaphorodoris and Calycidoris, Acanthodoris and Akiodoris. Millen demonstrated that based on reproductive characters, these four genera are more closely aligned to each other than to Onchidoris. This same result is shown in our phylogenetic analysis (Fig. 40). Onchidoris is in a sis- ter clade to the clade containing Diaphorodoris, Calycidoris and Acanthodoris. In addition, Millen demonstrated that external and radular characters clearly separate Onchidoris and Diaphorodoris as separate genera. Our analysis also found radular characters to support the two genera (#25, outer lateral teeth and #17, lip disk). The results of the present analysis differ in some aspects from the phylogeny of the Nudibranchia reconstructed by molecular markers (Wollscheid-Lengeling et al. 2001). In that analysis, the tree constructed by combining three genes (16S, 18S and COI, p. 251) shows Goniodoris nodosa and Diaphorodoris luteocincta as basally situated to Cadlina luteomarginata. In our analysis (Fig. 40), Cadlina is more basal to Goniodoris and Diaphorodoris. However, in 266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 4 40,17,33 | (1) 1222 (2) 4,3,28,30,31 (>2) 222,26 25 4,2,3,10,222,35 18 11,13 8 : 2,18 (1) (1) 15,19,21,26 Dae 32 (1) 4 (1) 1 4 (1) 1,9 5,29,35 (>2) 7,8,25,26,32 2,32 34,35 1 D 102,16,292 OO (1) Y 23 52,11,14,22,24 (1) }34 ) (1) (1) (1) 24,2526 34 1,2,4,27,31.B3 (1) (>2) 18,23 (1) 26,322 17,20,23,24 » 3,5,6,7,8,10,18,20,29 Adalaria jannae Adalaria proxima Adalaria loveni Onchidoris bilamellata Onchidoris muricata Calycidoris guenthen Diaphorodoris lirulatocauda Diaphorodoris luteocinta Diaphorodoris papillata Diaphorodoris mitsuli Goniodoris nodosa Okenia mediterranea Acanthodoris brunnea Acanthodoris lutea Acanthodoris pina Acanthodoris hudsoni Acanthodoris pilosa Acanthodoris rhodoceras Acanthodoris serpentinotus Acanthodoris nanaimoensis Acanthodoris nanega Acanthodoris planca Acanthodoris falklandica Akiodoris salacia Cadlina luteomarginata Bathydoris clavigera FiGuRE 40. Phylogeny of Acanthodoris. Single most parsimonious tree, 94 steps. Numbers below the branches are Bremer decay values. Numbers shown above the branches are character numbers from Table 2. The bold numbers indicate reversals. The tree is derived from three outgroup taxa, Bathydoris, Cadlina and Akiodoris. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 267 their tree constructed from a single gene (CO1, maximum likelihood tree, p. 250) Cadlina and Diaphorodoris appear in sister clades, with Bathydoris clavigera in a clade basal to both. The placement of Barhydoris is consistent with our findings. In their tree derived from partial sequences of16S (maximum likelihood tree, p. 249), Acanthodoris and Onchidoris group together in a basal- ly situated clade to Cadlina with Diaphorodoris more derived than any of these groupings. Our phylogeny also shows Diaphorodoris as more derived than either Acanthodoris or Onchidoris. In the present analyses, as noted above, the selection of outgroups affected the placement of the major clades. If five outgroups were selected (Bathydoris, Akiodoris, Cadlina, Goniodoris and Okenia) a single most parsimonious tree resulted (tree not shown) with the same tree scores as obtained by using the three outgroups noted above. There was no difference in this second tree structure however, if only Goniodoris and Okenia were selected as outgroups. In the single tree produced, Okenia and Goniodoris are the most basally situated taxa and Diaphorodoris is a mono- phyletic clade basal to the remainder of taxa. Interestingly, Bathydoris, Cadlina and Akiodoris clus- ter in a clade more derived than the monophyletic clade of Acanthodoris. This tree agrees in some aspects with the tree derived from the partial sequence of 16S (Wollscheid-Lengeling et al. 2001). In that tree, Goniodoris is the most basally situated taxon and Diaphorodoris is basally situated to Cadlina, Onchidoris, Goniodoris and Acanthodoris. The placement of these taxa in Wollscheid- Lengeling et al.’s analysis agrees with our morphology based tree when we use either Goniodoris plus Okenia as the outgroups, or by using five outgroups as described above. The present phylogeny of Acanthodoris (Fig. 40) contains one conspicuous clade of Acanthodoris, including species found in the Northern Hemisphere. Basally situated to this clade are three species found exclusively in the Southern Hemisphere: Acanthodoris falklandica, A. plan- ca and A. nanega. This hypothesis suggests that Acanthodoris from the northern hemisphere have a common ancestor. It is also remarkable that species from the eastern Pacific are much more diverse than species from either the Atlantic or the southern hemisphere. The southern hemisphere species form a grade at the base of the more derived northern hemisphere clade. There are two most likely scenarios for this tree structure. One possibility is that there could have been a mass extinc- tion of Atlantic, western Pacific or Southern Hemisphere species. Another possibility is that the dis- persal scenario for Acanthodoris is similar to the findings of Fahey and Gosliner (2004) for Aegiridae. In that study, the distribution pattern indicated a possible dispersal scenario from the Antarctic, north along both coasts of Australia, then into the tropical Indo-Pacific. The topology of the present tree indicates that a similar scenario could have occurred for Acanthodoris. The differ- ence is that Acanthodoris could have dispersed from the Southern Hemisphere and then into the Eastern Pacific Ocean, where most Acanthodoris are found. However, as with the case of Aegiridae (Fahey and Gosliner 2004), it is likely that not all species of Onchidorididae have been described. It is possible that there may be other undescribed species that will enable a more thorough analysis of biogeographical relationships among this clade of nudibranchs. ACKNOWLEDGEMENTS We thank Sandra Millen for sharing her unpublished data on Adalaria, Akiodoris and Armodoris. Specimens were provided by the Smithsonian National Museum of Natural History, Museo Nacional de Ciencias Naturales, Madrid, Zoologisk Museum Kobenhavns Universitet, Copenhagen and Sandra Millen, University of British Columbia. Thanks to Hans Bertsch who thor- oughly researched the status of Acanthodoris stohleri syn. nov. And last but not least, thanks to Dr. Michele Aldrich who read proof and saved the authors from an embarrassment of “hidden” errors. 268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 This project was funded by the National Science Foundation PEET grant DEB 9978155 “Phylogenetic Systematics of Dorid Nudibranchs” to T. Gosliner. LITERATURE CITED ABILDGAARD, P.C. 1789. Nudibranchia. Page 7, pl. 8.5 in O. Miiller, ed., Zoologia Danica sev animalium Daniae et Norvegiae rariorum ac minus notorum descriptiones et historia. 3" edition. N. Molleri, Havniae, Denmark. ABRAHAM, P.S. 1876. Notes on some genera of nudibranchiate Mollusca, with notices of a new genus and some hitherto undescribed species in the collection of the British Museum. Annals and Magazine of Natural History, ser. 6, 7:132—146, 4 pls. ABRAHAM, P.S. 1877. Revision of the anthobranchiate nudibranchiate Mollusca, with descriptions or notices of forty-one hitherto undescribed species. Proceedings of the Zoological Society of London 1877:196-269, pls 27-30. AGassiz, L. 1846. Onchidorus de Blainville 1816. in H.S. Scudder, ed. Nomenclator Zoologicus. Government Printing Office, Washington, D.C., USA. 340 pp. Aaassiz, L. 1850. Notes in minutes of Nov. 7, 1849 meeting of Boston Society of Natural History drawings of Doris diademata, D. coronata, D. tenella, D. pallida, all new Agassiz species exhibited and Meliboea arbuscula reported from Gay Head, Mass. Proceedings of the Boston Society of Natural History 3:191. ALDER, J., AND A. HANCOCK. 1842. Descriptions of several new species of nudibranchous Mollusca found on the coast of Northumberland. Annals and Magazine of Natural History 9:31—36. ALDER, J., AND A. HANCOCK.1845. Notice of a new genus and several new species of nudibranchiate Mollusca. Annals and Magazine of Natural History 16:311—316. ALDER, J., AND A. HANCOCK. 1845. A Monograph of the British Nudibranchiate Mollusca: with Figures of All the Species, vol. 1, pt. 7. The Ray Society. London, UK. 54 pp., 8 pls. ALDER, J., AND A. HANCOCK. 1862. Descriptions of a new genus and some new species of naked Molluscs. Annals and Magazine of Natural History 10:261—265. AURIVILLIUS, C. 1887. Placophora och Gastropoda, Ofersicht 6fver de af Vega—expeditionen insamlade arkiska hafsmollusker. Kongelige Svenska Vetenskaps-Akademiens Handlingar 2:313-383. BaBA, K. 1935. The fauna of Akkeshi Bay Part 1 Opisthobranchia. Journal of the Faculty of Science Hokkaido University ( 6) Zoology 4:115—125, pls. 7-8. BaBa, K. 1938. Three new nudibranchs from Izu, Middle Japan. Annotationes Zoologicae Japonenses, Tokyo 17:130-133. BaBa, K. 1949. Opisthobranchs of Sagami Bay. lwanami Shoten, Tokyo. 194 pp. 50 pls. Beck, H. 1842. Doris literata. Page 78 in H. Moller, ed., Index Molluscorum Groenlandiae, Naturhistorie Tidsskrift, vol. 4(1). Kj6benhavn, Denmark. BEHRENS, D.W. 1980. Pacific Coast Nudibranchs. A Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers. Los Osos, California, USA. 112 pp. BEHRENS, D.W. 1991. Pacific Coast Nudibranchs: A Guide to the Opisthobranchs Alaska to Baja California. Sea Challengers. Monterey, California, USA. 107 pp. BERGH, R. 1878. Malacologische Untersuchungen. Pages 547-601, pls. 62-65 in C. Semper ed., Reisen im Archipel der Philippinen, Theil 2, Heft 13. Kreidel, Wiesbaden, Germany. BERGH, R. 1880. On the nudibranchiate gasteropod Mollusca of the North Pacific Ocean, with special refer- ence to those of Alaska, Part 2. Proceedings of the Academy of Natural Sciences, Philadelphia 40-127, pls. 9-16. BERGH, R. 1905. Die Opisthobranchiata der Siboga-Expedition. Pages 1-248, pls 1-20, in M. Weber. ed., Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch Gebied verzameld in Nederlandsch Oost-Indié 1899-1900 aan boord H. M. Siboga onder commando van Luitenant ter zee F° kl. G. F- Tydeman, vol. 50. Brill, Leiden, Netherlands. BLAINVILLE, H.D. de. 1816. Quartrieme mémoire sur les mollusques, de l’ordre des cyclobranches. Bulletin des Sciences par la Société Philomathique. Paris, ser. 3, (1—3):93-97. —— FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 269 Burn, R. 1969. A memorial report on the Tom Crawford Collection of Victorian Opisthobranchia. Journal of the Malacological Society of Australia 12:64—106. COCKERELL, T.D., AND C. ELior. 1905. Notes on a collection of Californian nudibranches. Journal of Malacology 12:31-53, pls. 7-8. Cuvier, GL. 1830. Les nudibranches (3). Le regne Animaux Nouvelle 2:50-58. D’ORBIGNY, A. 1837. Mémoire sur les especes et sur des genres nouveaux de |’ordre des nudibranches, observes sur les cotes de France. Magasin de Zoologie, 7 Class, 5:1—16, pls. 102-109. DALL, W. 1871. Descriptions of sixty new forms of molluscks from the West Coast of North America and the North Pacific Ocean, with notes on others already described. American Journal of Conchology 7:93—160. DesMAREST, E. 1858. Onchidorus. Page 142 in J. Chenu, ed., Encyclopédia Histoire Naturelle, vol. 4, Crustacés. Mollusques. Zoophytes. Marescq et cie, Paris. 22 vols. DyspowsklI, W. 1900. Beschreibung einer Hinterkiemer-Schnecke aus dem Baikal-See. Nachrichtshlatt der Deutschen Malakozoologischen Gesellschaft 32:143—152. Euiot, C. 1907. Nudibranchs from New Zealand and the Falkland Islands. Proceedings of the Malacological Society London 7:327-361. Eior, C. 1910. A Monograph of the British Nudibranchiate Mollusca. Part 8 (Supplement). The Ray Society. London. 198 pp. Fapricius, O. 1780. Fauna Groenlandica. 1.G. Rothe, Hafniae et Lipsiae, Denmark. 452 pp. FAHEY, S.J., AND T.M. GOSLINER. 2004. A phylogenetic analysis of the Aegiridae Fischer, 1883 (Mollusca, Nudibranchia, Phanerobranchia) with descriptions of eight new species and a reassessment of Phanerobranch relationships. Proceedings of the California Academy of Sciences, ser. 4, 55:613—689. FARRAN, GP. 1903. The nudibranchiate molluscs of Ballynakill and Bofin Harbours, Co. Galway. Report on the Sea and Inland Fisheries of Ireland for 1901, Part 2:123—132. FerussAc, A. 1922. Tableaux systématiques des animaux molluscques. Suivis d’un Prodrome Général I-XLVII:1-110. FLEMING, J. 1820. Doris nigricans. Brewster's Edinburgh Encyclopedia 14:618. ForBes, E. 1838. Malacologia Monensis, A Catalogue of the Mollusca Inhabiting the Isle of Man and the Neighboring Sea. J. Carfrae and Son, Edinburgh, Scotland. 63 pp. GisTEL. J. 1848. Villiersia D’Orbigny, A. 1837. Naturgeschichte des thierreichs fiir héhere Schulen. Hoffman’sche Verlags—Buchhandlung, Stuttgart, Germany. 216 pp., 32 pls. GMELIN, J. 1791. Jn C. Linnaeus. ed., Systema Naturae, ed. 3, 1(6): 3103-3107, 3147-3148. GODDARD, J. 1984. The opisthobranchs of Cape Arago, Oregon, with notes on their biology and a summary of benthic opithobranchs known from Oregon. The Veliger 27:143-163. GOSLINER, T.M. 1987. Nudibranchs of South Africa. Sea Challengers, Monterey, California, USA. 138 pp. GosLINER, T.M., AND R. JOHNSON. 1999. Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species. Zoological Journal of the Linnean Society 125:1—114. Gray, J.E. 1842-1857. Figures of Molluscous Animals, Selected from Various Authors. Longman, Brown, Green and Longmans, London, UK. Vol. 4, 123 pp; Vol. 5, 112 pls. HERRMANNSEN, A. 1852. Indicis generum Malacozoorum, supplementa et Corrigenda. Theodori Fischer, Cassellis, Germany. 140 pp. IREDALE, T., AND C. O’DoNoGHUE. 1923. List of British nudibranchiate Mollusca. Procedings of the Malacological Society of London 15:115—226. JAECKLE, W. 1984. The opisthobranch molluscs of Humboldt County, California. The Veliger 26:207-213. JEFFREYS, J. 1869. British Conchology or an Account of the Mollusca Which Now Inhabit the British Isles and the Surrounding Seas. J. van Voorst, London, UK. 5 Vols. Just, H., AND M. Epmunps. 1985. North Atlantic Nudibranchs (Mollusca) seen by Henning Lemche. Ophelia Supplement 2:170. Keen, A. M. 1971. Sea Shells of Tropical West America: Marine Mollusks from Baja California to Peru. Stanford University Press. Stanford, California, USA. 624 pp. Lance, J. 1968. New Panamic nudibranchs (Gastropoda; Opisthobranchia) from the Gulf of California. Transactions of the San Diego Society of Natural History 15:\—13. 270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 LeacH, W. 1847. The classification of the British Mollusca. Annals and Magazine of Natural History 20:267-273. LINNAEUS, C. 1767. Systema Naturae, 12 ed. Salvii, Holmiae, Sweden. 1384 pp. Loven, S. 1846. Index molluscorum litora Scandinaviae Occidentalia habitantium. Ofversigt af Kongl Vetenskaps Akademiens Férhandlinger Stockholm 3:135—160. MACFARLAND, F. 1905. Opisthobranchiate Mollusca from Monterey Bay, California, and vicinity. Bulletin of the United States Bureau of Fisheries 25:109-151, pls. 18-31. MACFARLAND, F. 1925. The Acanthodorididae of the California Coast. Nautilus 39:49—65, pls. 2-3. MACFARLAND, F. 1926. The Acanthodorididae of the Californian Coast. Nautilus 39:94—103, pls. 2-3. MACFARLAND, F. 1966. Studies of Opisthobranch Mollusks of the Pacific Coast of North America. Memoirs of the California Academy of Sciences, no. 16. California Academy of Sciences, San Francisco, California, USA. 546 pp. MACGINITIE, G., AND N. MACGINITIE. 1949. Natural History of Marine Animals. McGraw-Hill, New York, USA. 523 pp. ; MAppIson, W.P., AND D.R. MADDISON. 2000. MacClade. Sinauer Associates, Cambridge, Massachusetts, USA. Marcus, E. 1959. Lamellariacea und Opisthobranchia. Reports from the Lund University Chili Expedition 1948-49. Lunds Universitets Arsskrift 36:89-261. Marcus, E., AND E. MARcus. 1967. American Opisthobranch Mollusks. University of Miami Institute of Marine Sciences. Miami, Florida, USA. 256 pp. McDonaLp, G. 1975. Key D: Sacoglossa and Nudibranchia in R.I. Smith and J.T. Carlton eds., Light's Manual, Intertidal Invertebrates of the Central California Coast. University of California Press, Berkeley, California, USA. 716 pp. McDOonaLb, G. 1983. A review of the nudibranchs of the California coast. Malacologia 24:114—276. McDONALD, G., AND J. NYBAKKEN. 1978. Additional notes on the food of some California nudibranchs with a summary of known food habits of California species. The Veliger 21:110—119. McDONALD, G., AND J. NYBAKKEN. 1980. Guide to the Nudibranchs of California, Including Most Species Found from Alaska to Oregon. American Malacologists Inc., Melbourne, Florida, USA. 72 pp. MILLEN, S. 1985. The nudibranch genera Onchidoris and Diaphorodoris (Mollusca, Opisthobranchia) in the northeastern Pacific. The Veliger 28:80—93. MILLEN, S., AND A. MARTYNOY. 2005. Redescriptions of the nudibranch genera Akiodoris Bergh, 1879 and Armodoris Minichev, 1972 (Suborder Doridacea), with a new species of Akiodoris and a new family Akiodorididae. Proceedings of the California Academy of Sciences, ser. 4, 56(1):1—22. MILLEN, S.V. 1987. The nudibranch Adalaria, with a description of a new species from the northeastern Pacific. Canadian Journal of Zoology 65:2696-2702. MOLLER, H. 1842. Index Molluscorum Groenlandiae. Naturhistorisk Tidsskrift 1842:76—-94. Morcn, O. 1857. Pages 11-13, Part 4. Mollusks. Prodromus faunae molluscorum Grénlandiae. Fortegnelse over Grénlands Bléddyr in H.J. Rink, ed., Grénland geographisk og statistisk beskrevet, 1-28. Morcu, O. 1868. Faunula molluscorum Islandiae. Oversigt over Island Bléddyr. Videnskabelige Meddeleser fra Dansk Naturhistorisk 11—13:185—227. MULLER, O. 1776. Zoologiae Danicae, Prodromus seu animalium Daniae et Norvegiae ingenarum charac- teres, nomina, et Synonyma imprimis popularium. N. MOlleri et filii, Havniae, Denmark. 282 pp., 32 pls. NYBAKKEN, J. 1978. Abundance, diversity and temporal variability in a California intertidal nudibranch assemblage. Marine Biology 45:129-146. O’ DONOGHUE, C. 1921. Nudibranchiate Mollusca from the Vancouver Island Region. Transactions of the Royal Canadian Institute 13:147—209. O’DONOGHUE, C. 1927. Notes on the nudibranchiate Mollusca from the Vancouver Island Region. 5. Transactions of the Royal Canadian Institute 16:\—12, pl. 1. ODHNER, N. 1926. Die Opisthobranchien. Pages 1-100, Vol. 2 in N. Odhner, ed., Further Zoological Results of the Swedish Antarctic Expedition 1901—1903 under the direction of Dr. Otto Nordenskjold. Norstedt and Sdner, Stockholm, Sweden. OrTEA, J.. AND M. BALLESTEROS. 1982. Sobre algunos Onchidoris Blainville, 1816 (Mollusca, FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 Dail Opisthobranchia, Doridacea) del litoral ibérico. Investigacion Pesquera 46:239-254. PORTMANN, A., AND E. SANDMEIER. 1960. Zur kenntnis von Diaphorodoris (Gastropoda, Nudibranchia) und ihrer mediterranen Formen. Verhandelungen der Naturforschenden Gesellschaft in Basel 71:174-183. Pruvor-Fot, A. 1951. Etudes des nudibranches de la Méditerranée (2 Partie). Archives de Zoologie Expérimentale et Générale 88:1—80, pls. 1+. PruvoT-FoL, A. 1954. Mollusques Opisthobranches. Office Central de Faunistique. Paris, France. 460 pp. ROCHEBRUNE, A., AND J. MABILLE. 1891. Mollusques. Gauthier-Villars et fils, Paris, France. 143 pp. Roainskaya, I. 1972. Calycidoris guentheri (Gastopoda, Nudibranchia) taxonomy and geographical range. Zoological Journal, Institute of Oceanology, Moscow 6:913-918. ROLLER, R., AND S. LONG. 1969. An annotated list of opisthobranchs from San Luis Obispo County, California, USA. The Veliger 11:424—430. RupDMAN, W. 2004. Further species of the opisthobranch genus Okenia (Nudibranchia: Goniodorididae) from the Indo-West Pacific. Zootaxa 695:1—70. RUDMAN, W.B. (accessed 2004). SeaSlug Forum. Vol. 2003 http://www.seaslugforum.net RUDMAN, W.B., AND R. WILLAN. 1998. Opisthobranchia. Pages 915-942 in P.L. Beesley, G.J.B. Ross, A. Wells. eds., Mollusca: The Southern Synthesis. Fauna of Australia, vol. 5, pt. B. CSIRO Publishing, Melbourne, Australia. 1234 pp. Sars, G. 1870. Bidrag til Kundskab om Christianiafjordens Fauna 2. NYT Magazin for Naturividenskaberne 17:113—232. Nudibranchia 186—194. SAUVAGE, H. 1873. Catalogue des nudibranches des Cotes du Boulonnais, dressé, d’apres les notes de Bouchard-Chantereaux. Journal de Conchyliologie 3:25—36. SCHMEKEL, L. 1968. Ascoglossa, Notaspidea und Nudibranchia im Litoral des Golfes von Neapel. Revue Suisse de Zoologie 75:103-155. SCHMEKEL, L., AND A. PORTMANN. 1982. Opisthobranchia des Mittelmeeres. Nudibranchia und Saccoglossa. Springer-Verlag, Berlin, Germany. 410 pp. ScHRODL, M. 1996. Nudibranchia y Sacoglossa de Chile: Morfologia exterior y distribucion. Gayana Zoologia 60:17-62. ScHRODL, M. 1997. Range extension of Magellanic nudibranchs (Opisthobranchia) into the Peruvian faunal province. The Veliger 40:38-42. ScHRODL, M. 2003. Sea Slugs of Southern South America. Conch Books. Hackenheim, Germany. 165 pp. STEINBERG, J. 1963. Notes on the Opisthobranchs of the west coast of North America II]. Further nomencla- torial changes in the Order Nudibranchia. The Veliger 6:63—67. THompson, T.E. 1958. Observations on the radula of Adalaria proxima (A. & H.) (Gastropoda Opisthobranchia). Proceedings of the Malacological Society of London 33:49—56. THOMPSON, T.E., AND GH. Brown. 1984. Biology of Opisthobranch Molluscs, vol. 2. The Ray Society, London, UK. 280 pp. THompson, W. 1840. Contributions towards a knowledge of the Mollusca Nudibranchia and Mollusca Tunicata of Ireland, with descriptions of some apparently new species of Invertebrata. Annals and Magazine of Natural History 5:84-102. THompson, W. 1845. Additions to the fauna of Ireland including descriptions of some apparently new species of Invertebrata. Annals and Magazine of Natural History 15:308-322. VALDES, A. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society, London 136:535—636. VALLES, Y. 2002. Taxonomy and phylogeny of Kaloplocamus and Plocamopherus and their relationships with other phanerobranchs. Masters thesis. San Francisco State University. San Francisco, California, USA. 266 pp. VayssIERE, A. 1919. Recherches zoologiques et anatomiques sur les mollusques Opithobranches du Golfe de Marseille, 2me Supplément. Annales de la Museum Histoire Natural Marseille 17:53-92. VeERRILL, A. 1870. Contributions to zoology from the Museum of Yale College. No. 8. Descriptions of some New England Nudibranchiata. American Journal of the Sciences and Arts, ser. 2, 50:405—408. VERRILL, A. 1879. Notice of recent additions to the marine fauna of the east coast of North America. No. 4. American Journal of Science, ser. 3, 17:309-315. DD PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 20 WAGELE, H. AND R.C. WILLAN. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 130:83-181. WILLAN, R.C., AND N. COLEMAN. 1984. Nudibranchs of Australasia. N. Coleman, Springwood, Australia. 56 Pp. WILLIAMS, G.C. AND T.M. GOSLINER. 1979. Two new species of nudibranchiate molluscs from the west coast of North America, with a revision of the family Cuthonidae. Zoological Journal of the Linnean Society 67:203-223. WOLLSCHEID-LENGELING, E. AND J. BOORE, AND W. BROWN, AND H. WAGELE. 2001. The phylogeny of Nudibranchia (Opisthobranchia, Gastropoda, Mollusca) reconstructed by three molecular markers. Organisms Diversity & Evolution \:241—256. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. FAHEY AND VALDES: REVIEW OF ACANTHODORIS GRAY, 1850 Appendix A Data matrix for the taxa included in the phylogenetic analysis. Bisboohobobh-Hobhcbs-bclbebeb) poooosseoereernnonerseeees 1 —— Sal i | | | TT] DOO St tO) ish ooo obs HHNobbseespeHob6) ia) DOOR AHH falealalaaae iii Moco Se eat ——— ——— FROSSSOS SSS SSS SOS SS A4999 7 | , DOO NININNIN A RNNINAINANNAINNOOOOR lanl 1 Sari royal } FARDOSOOOS SOO SOOO S SO ONFO0 | { 1 ‘ | iu i \ } I i | i { 3 Ta an ad ead fel net cd ealicaeh {pa SRR RR RRR ARR AOR — 4 it —— AAA AAHOORMOODODODRSDORHDOClO [poet ea | {eet Hoo x5 BR OMRON Beet PPEPEPCEPECECHET Pere i S sacseuesesedasanansagiczs 015 bid 29995 5s Seren SmSoeS ss) Li ff ed ee | tet i ea i i tal & = nnososSsSoSoSes05e5000040 an ane Ea Sa SIS COR RAR RR ROCCO) +5 BEE REEEEEEEREREEREREERERE = eee fife) EEE Est | me poeo------bbbbbbpsiceclebs a5 saucersesceanscnansesanaa ajpppoopsopbooppb bobo) a ol Rs IS IS a a ISIE NEbossosesoHosspsoosesososoo- - i es SS) ea oleate wlopoococococ—pcSpSeoSSCSCCSCOSOSOCSCOS | Ww SAOSIN Tt DHOHO SOS S OSHS SAAS DORR Or) ] al weal mb AR OA OHS OH ORE HOHHAAoRAooH i NE aro HoH oHoobobbbboobHH| abo bo aHoHoeHaHoHHobobHoook4 = 1 i} ln! HSUPA cfd thea iS 1a) Sst teiwade dt = | a= Hetea eaten} s fae} iS | nig) yt iad ro) wo} | | {sie} hare [ont vie}_ is =\_} lo Rosters Atedt ae |) ais HGS] leloeie! Elo) | (el stlai_| (S| GBOD=| Gem sas Sa Geisis ic SSYe ave co YM sso godgicaaig in ser secgtgs~oyccOcoc a) Yost) SSDS Pikes \b\a =|S) Di SiE|=|OS ig) y|—j/E a Beis ecsciciaies alad|\s/s/0) Sessuuuvoge Free pla eo S/S | GROL SSE Cee eee seeeesesials eli>/O| jdlololo| BiG iEES Leflore | iia! 2 IO SY ooo OZ|d/00/5/O 00/0/00 |9/O/4/4 in) ie) 2-2 soookescssoscooso OE FMS ZisecOrrer or qooogc gC CCD CO OE), o RE RPCCCCSacccesesscssseosis See eH ie i es == |S EE] = SSSSSCSog=Sececceecececss|zo| a PSS gaa ai UC tt isan mee To 6 SOG GH KKK OF UU O OLE |g = Lt€CUADAA LO Ce tctctce¢eeccccooncy 2 elit tina a ~ of © va) oy cf ° o cect ein Semmes ae a EE ae SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 345 Chrosioderma namoroka Silva-Davila, sp. nov. Figs. 1A, 20; Map 1 Types.— Male holotype from Pare National de Namoroka, 16.9 km 317°NW Vilanandro, 16°24'24"S, 045°18'36"E, 100m, tropical dry forest, beating low vegetation, 12—16.xi.2002 (Fisher, Griswold et al.), deposited in CAS. ETYMOLOoGy.— The species name is a noun in apposition taken from the type locality. D1aGnosis.— Males are recognized by having a tegulum extending as long as one half cym- bial length and a ‘J’ shaped sclerotized process (Fig. 20A); additionally, all legs have dorsal black hairs surrounding the spine sockets (Fig. 1A). MALE (holotype).— Total length 6.9. Carapace and abdomen pale yellow; carapace partly coated with black hairs; legs with dark markings and black hairs surrounding spine sockets (Fig. 1A); abdomen with two guanine spots (Fig. 1A). Carapace 3.05 long, 2.55 wide; ocular area 0.48 long, 1.19 wide; OQA 0.67; OQP 0.73; diameter of eyes AM:AL:PM:PL, 0.25:0.17:0.17:0.17; AME-AME 0.21; AME-ALE 0.05; ALE-ALE 0.76; PME-PME 0.4; PME-PLE 0.13; PLE-PLE 0.92. Clypeal height 0.05. Sternum 1.58 long, 1.45 wide; labium 0.34 long, 0.52 wide; endites 0.8 long, 0.44 wide. Femur I 3.18 times carapace width. Spination: tibia I-II v7 pairs ; metatarsi I-II v2-2-0. Left legs I-II, 1V missing. Leg measurements: I IT III IV Palp Femur 8.1 7.8 4.5 6.5 1.8 Patella 1.6 1.6 Hail ilath 0.65 Tibia 9.9 9.3 4.4 6.9 0.93 Metatarsus 10.8 9.7 4.7 8.4 --- Tarsus Dal 1.8 1.0 1.4 Dep Total 325 30.2 1537, 24.3 5.58 Male palp as in Figures 20A—D. FEMALE.— Unknown MATERIAL EXAMINED.— MADAGASCAR: Mahajanga: HOLOTYPE above (CASENT 9017956). DISTRIBUTION.— Known from north western Madagascar (Map 1). Chrosioderma ranomafana Silva-Davila, sp. nov. Figs. 10, 12, 21-23; Map 1. Types.— Male holotype from Ranomafana, Radio Tower, 21°15.05’S, 047°24.4'E, 3700 ft, forest edge, malaise canopy trap, 27.1i-9.111.2003 (R. Harin'Hala), deposited in CAS. ETYMOLOGyY.— The species name is a noun in apposition taken from the type locality. D1aGnosis.— Males have a prolateral conductor arising on tegular apex (Figs. 10D, 21A—C, 22 A, 22C). Females have epigynal lateral lobes with small antero-lateral hood like projections (Figs. 12A, 23A); they differ from C. roaloha in having an elongate spermathecal head (Figs. 12B, 23B). MALE (holotype).— Total length 6.6. Carapace and abdomen pale yellow. Carapace 3.2 long, 2.75 wide; ocular area 0.5 long, 1.24 wide; OQA 0.68; OQP 0.74; diameter of eyes AM:AL:PM:PL, 0.21:0.18:0.15:0.18; AME-AME 0.25; AME-ALE 0.06; ALE-ALE 0.79; PME-PME 0.43; PME- PLE 0.13; PLE-PLE 0.96. Clypeal height 0.05. Sternum 1.74 long, 1.66 wide; labium 0.36 long, 0.62 wide; endites 0.96 long, 0.62 wide. Femur I 2.54 times carapace width. Legs I left and II miss- ing. Spination: tibia I v7 pairs, metatarsi I v2-2-0. Leg measurements: 346 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 I I I IV Palp Femur 7.0 - 4.1 5.9 1.8 Patella 1.6 - 1.3 1.3 0.7 Tibia 8.8 - 4.3 6.1 0.9 Metatarsus 9.3 - 4.5 8.0 --- Tarsus 1.9 - 1.0 1.4 2.4 Total 28.6 - 15.2 20a 5.8 RTA (Figs. 21A, 21C, 22A) elongate and sinuous, projecting into a small hook; cymbium with a short lobate retrolateral projection (Fig. 22B); tegulum (Figs. 21A—D; 22A—C) shorter than % cymbial length, nearly as wide as alveolus width; embolus originating mesally (Figs. 21B, 22 A-B); conductor short, arising on prolateral side of tegular apex (Figs. 10D, 21A—C, 22A, 22C). FEMALE (Ranomafana, Belle Vue).— Total length 9.5. Pale yellow as in male. Carapace 3.55 long, 3.15 wide; ocular area 0.46 long, 1.43 wide; OQA 0.75; OQP 0.86; diameter of eyes AM:AL:PM:PL, 0.24:0.17:0.17:0.17; AME-AME 0.29; AME-ALE 0.13; ALE-ALE 1.0; PME- PME 0.52; PME-PLE 0.17; PLE-PLE 1.16. Clypeal height 0.08. Sternum 2.0 long, 1.8 wide; labi- um 0.4 long, 0.68 wide; endites 1.0 long, 0.66 wide. Femur I 1.61 times carapace width. Spination: tibiae I-II v7 pairs, metatarsi I-I] v2-2-0. Leg measurements: SI II II IV Palp Femur Se) 5.8 3.8 5.1 1.53 Patella 1.7 1la7/ 192 1.3 0.73 Tibia 6.7 6.6 3333 4.8 1.13 Metatarsus 6.2 5.9 3.4 Sif --- Tarsus 1S 1.4 0.9 Mo? 2.08 Total 21.8 21.4 12.6 18.1 5.47 Epigynum (12A, 23A) with small antero-lateral hood-like projections. Vulva as in Figures IZ. 238. MATERIAL EXAMINED.— MabDAGaSscar: Fianarantsoa: Ranomafana, HOLOTYPE above (CASENT 1018736); Ranomafana, Belle Vue, 21°15.99'S, 047°25.21'E, 3340 ft, rainforest, malaise canopy trap, 14-21.1.2002 (R. Harin'Hala, CASENT 9005610), 1 2; Ranomafana, Radio Tower, 21°15.05’S, 047°24.43’E, 3700 ft, forest edge, open area, malaise trap (28.1-04.i1i.2002 (R. Harin'Hala, CASENT 9010722), 12; Madagascar (Hildebrandt, ZMB 7871), 192 6J. DISTRIBUTION.— Known from central Madagascar (Map 1). Chrosioderma roaloha Silva-Davila, sp. nov. Figs. 11, 24; Map 2. TypEs.— Female holotype from Parc National Ranomafana, Talatakely, 21°15’S, 047°25.26’E, 915—1000m, 30.x—20.x1.1998 (V.F. Lee and K. Ribardo), deposited in CAS. ETYMOLOGY.— The species name is from the Malagasy for “two heads” in reference to the bifid spermatheca head. DIAGNOsIS.— Females have epigynal lateral lobes with small antero-lateral hood like projec- tions (Fig. 11A, 24A); they differ from C. ranomafana in having the spermathecal head projecting into two asymmetrical branches (Figs. 11B, 24B). MALE.— Unknown. FEMALE (holotype).— Total length 9.1. Carapace and abdomen pale yellow. Carapace 3.9 long, 3.4 wide; ocular area 0.51 long, 1.49 wide; OQA 0.71; OQP 0.86; diameter of eyes AM:AL:PM:PL, 0.19:0.17:0.17:0.13; AME-AME 0.3; AME-ALE 0.14; ALE-ALE 0.98; PME- PME 0.52; PME-PLE 0.19; PLE-PLE 1.24. Clypeal height 0.05. Sternum 1.95 long, 1.75 wide; SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 347 labium 0.42 long, 0.7 wide; endites 1.06 long, 0.9 wide. Femur I 1.61 times carapace width. Spination: tibia I-II v7 pairs (+1, one side of right Ti I), metatarsi I-I]] v2-2-0. Leg measurements: I I TIT IV Palp Femur 5) / 5.8 3.6 4.9 ES Patella 1.8 1.8 3) 13: 0.4 Tibia 6.6 6.5 3.4 4.7 ei} Metatarsus 6.1 5.8 3.3) DO == Tarsus Le) 15 1.0 eZ Bol Total led 21.4 12.6 16.5 5.18 Epigynum (Figs. 11A, 24A) with small antero-lateral hood-like projections. Vulva as in Figures 11B, 24B. MATERIAL EXAMINED.— MADAGASCAR: Fianarantsoa: HOLOTYPE above (CASENT 9008465); Talatakely, 21°12’S, 047°27'E, 18.v.1992 (V. and B. Roth and Albert, CASENT 9008508), 1 juvenile ¢. DISTRIBUTION.— Known from central Madagascar (Map 2). Chrosioderma soalala Silva-Davila, sp. nov. Bigs) 8-9" 2527: Map 2. TypEs.— Male holotype from Pare National de Baie de Baly, 12.4 km 337° NNW Soalala, 16°0'36"S, 045°15'54”"E, 10 m, tropical dry forest, beating low vegetation, 26—30.x1.2002 (Fisher, Griswold et al.), deposited in CAS. ETyMOLOGyY.— The species name is a noun in apposition taken from the type locality. DiaGnosis.— Males are recognized by having a spherical tegulum and a large sclerotized process at the embolus base (Figs. 8A—C, 9, 25A—C, 26A—D). MALE (holotype).— Total length 5.65. Carapace and abdomen pale yellow. Carapace 2.85 long, 2.3 wide; ocular area 0.41 long, 1.03 wide; OQA 0.6; OQP 0.63; diameter of eyes AM:AL:PM:PL, 0.24:0.16:0.17:0.17; AME-AME 0.17; AME-ALE 0.00; ALE-ALE 0.86; PME- PME 0.29; PME-PLE 0.09; PLE-PLE 0.75. Clypeal height 0.03. Sternum 1.55 long, 1.28 wide; labium 0.32 long, 0.48 wide; endites 0.74 long, 0.42 wide. Femur I 2.83 times carapace width. Spination: tibia I-II v6 pairs ; metatarsi I-II v2-2-0. I left leg missing. Leg measurements: I I Il IV Palp Femur 6.5 6.2 3.6 5.4 LSS) Patella 1.6 ES) 0.9 ileil 0.55 Tibia ed Tell BP Sil 0.83 Metatarsus 8.8 He) 35) 6.6 --- Tarsus 1.9 ites) 0.8 ili 1.48 Total 24.6 23.6 12.0 19.3 4.4] Tegulum partly sclerotized (Figs. 25-26); embolus base with large proximal process; subtegu- lum projecting into a large ventral lobe (Figs. 8A—C, 9, 25A—C, 26A—D). FEMALE (Tombeau Vazimba).— Total length 8.0. Ornamentation pattern as in male. Carapace 3.45 long, 3.1 wide; ocular area 0.43 long, 1.23 wide; OQA 0.6; OQP 0.68; diameter of eyes AM:AL:PM:PL, 0.19:0.15:0.14:015; AME-AME 0.25; AME-ALE 0.09; ALE-ALE 0.81; PME- PME 0.38; PME-PLE 0.18; PLE-PLE 0.99. Clypeal height 0.05. Sternum 1.75 long, 1.69 wide; labium 0.32 long, 0.54 wide; endites 0.86 long, 0.52 wide. Femur I 1.74 times carapace width. Leg measurements: 348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 I II Ill IV Palp Femur 5.4 5.5 3.4 4.7 1.38 Patella ei lod EZ 13) 0.63 Tibia 5.8 5.9 2.9 4.2 1.0 Metatarsus 5.4 5.0 2.6 5.0 --- Tarsus 1.4 1S 0.9 Ell 1.53 Total 19.7 19.4 11.0 16.3 4.54 Epigynum (Fig. 27A) with flattened postero-lateral hood-like projections. Vulva as in Figure 7B: MATERIAL EXAMINED.— MADAGASCAR: Mahajanga: HOLOTYPE above (CASENT 9018006); Pare National de Baie de Baly, 12.4 km 337° NNW Soalala, 16°0’36"S, 045°15'54”E, 10 m, tropical dry forest, gen- eral collecting at night, 26—30.x1.2002 (Fisher, Griswold et al., CASENT 9018047), 1 &; Pare National Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, 19°8’31"S, 044°49'41”E, 50 m, tropical dry forest, general collecting ground spiders, 6—10.x1.2001 (B.L. Fisher et al. CASENT 9008803), 1 2; Forét de Tsimembo, 8.7 km 336° NNW Soatana, 19°1'17”"S, 044°26'26”E, 20 m, tropical dry forest, general collect- ing/beating, 21—25.xi.2001 (Fisher, Griswold et al, CASENT 9018047), 1 3; TOLIARA: Parc National de Kirindy Mite, 16.3 km 125°SE Belo sur Mer, 20°47'43”S, 044°8'49”E, 80m, tropical dry forest, general col- lecting/beating, 6—10.x11.2001 (B.L. Fisher et al., CASENT 9004644), 1 31 juvenile 3; Forét de Kirindy, 15.5 km 64° ENE Marofandilia, 20°2'42”S, 066°39'44”E, 100m, tropical dry forest, beating low vegetation, 28.xi—3.xii.2001 (B.L. Fisher et al., CASENT 9005038), 1 o 1 juvenile o. DISTRIBUTION.— Known from western Madagascar (Map 2). ACKNOWLEDGMENTS I am deeply indebted to Dr. C. Rollard (MNHN) and Dr. R. Jocqué (MRAC) for their kind hos- pitality and access to their unsorted collection of sparassid spiders. Also, I wish to thank Dr. J. Dunlop (ZMB) and Dr. P. Jager (Senckenberg Museum, Frankfurt) for lending additional speci- mens for this project. | am most thankful to Charles Griswold for his continuous support to my research work and time for reviewing drafts of this paper. I gratefully acknowledge Giovanni Maki for the illustrations, Michelle Koo and Xinping Wang for their assistance with the maps. This proj- ect was partially funded by the NSF grant DEB-0072713 to C. Griswold and B. Fisher and by the Frizzell and Planidium funds of CAS; additional support for examining various museum collec- tions was provided by the Schlinger Foundation. LITERATURE CITED PLATNICK, N.I. 2004. The World Spider Catalog, version 4.5. American Museum of Natural History, online at . Simon, E. 1897. Histoire naturelle des Araignées. Pages 1-192 and figs. 1-200 in Encyclopédie Roret, 24 ed., vol. 2, pt. 1. L. Mulo, Paris, France. Illustrations 349 350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 FIGURE 1. Chrosioderma, habitus, dorsal view. A. C. namoroka sp. nov., male holotype. B. Juvenile female (CASENT 9014911). ERS ea SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON FIGURE 2. Chrosioderma, juvenile carapace. A. Eyes, frontal view (CASENT 9014911). B—D. Juv. CASENT 9003610. B. Cheliceral teeth, ventral view, inset and white arrow to cheliceral glands. C. Right endite showing serrula, inset and white arrow to sieve plate. D. Close up of serrula, ventral view. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 FIGURE 3. Chrosioderma spp., tarsi-metatarsi. A-B. C. ranomafana sp. nov., female. A. Right pedipalp; arrow and inset to tarsal organ. B. Close up of palpal tarsal claw. C. C. soalala sp. nov., male, metatarsal membrane.D. C. roaloha sp. nov., female, metatarsal membrane. FIGURE 4. Chrosioderma spp., tarsi. A-B. C. havia, A. Female I left tarsus, arrow to tarsal organ. B. Close up of tarsal organ and trichobotrium. C. C ranomafana sp. nov., male | left tarsus, tarsal claws and claw tufts. D. C. ranomafana sp. nov., female II right tarsus. Tb = trichobotrium; TO = tarsal organ. 354 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 10um FIGURE 5. Chrosioderma ranomafana sp. nov. A. Male abdomen. B. Close up of epiandrous gland spigots. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON =) > AY ESS \3 . #7) FIGURE 6. Chrosioderma ranomafana sp. nov., male spinnerets. A. Overview. B. ALS. C. PMS. D. PLS. AC = aciniform gland spigots, ALS = anterior lateral spinneret, MAP = major ampullate gland spigots, mAP = minor ampullate gland spig- ots, nu =nubbin, PI = piriform gland spigots, PLS = posterior lateral spinneret, PMS = posterior median spinneret, tar = tartipore. 356 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 FIGURE 7. Chrosioderma roaloha sp. nov., female spinnerets. A. ALS. B. PMS. C. Close up of PMS. D. PLS. AC = acini- form gland spigots, CY = cylindrical gland spigots, MAP = major ampullate gland spigots, mAP = minor ampullate gland spigots, PI = piriform gland spigots, tar = tartipore. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 3) 0.5mm FIGURE 8. Chrosioderma soalala sp. nov., male CASENT 9018047. Left palp. A. Ventral view. B. Prolateral view. C. Retrolateral view. E = embolus, RTA = retrolateral tibial apophysis, ST = subtegulum, T = tegulum. Black arrows to embol- ic base. 358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 0.5mm Ficure 9 (left). Chrosioderma soalala sp. nov., male CASENT 9018047. Expanded right palp, prolateral and retrolat- eral views. An = anelli, E = embolus, P = petiole, RTA = retrolateral tibial apophysis, SC = cymbial scopula, ST = subtegu- lum, T = tegulum. Black arrows to embolic base. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON FIGURE 10. Chrosioderma ranomafana sp. nov., male CASENT 9010722. Expanded right palp. A. Prolateral view, black arrow to conductor base. B. Retrolateral view. C. Close up, retrolateral view. D. Ventral view, white arrow to conductor. An = anelli, BH = basal hematodocha, CY = cymbium, E = embolus, Eb = embolic base, Pp = pars pendula, ST = subtegulum, T = tegulum. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 360 min Fourth Series, Volume 56, No. 25 0.66 mm base of spermatheca, CD = copu- mum, ventral view. LL = epigynal lateral lobe, MS igy alia. A. Ep . nov., female genit ector, arrow to LL hood-like projection. B. Vulva, dorsal view. BS ynal median s g 5) FIGURE 11. Chrosioderma roaloha sp epi latory duct, FD spermathecal stalk. fertilization ducts, HS = head of spermatheca, St = SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON Ficure 12. Chrosioderma ranomafana sp. nov., female genitalia. A. Epigynum, ventral view. LL = epigynal lateral lobe, MS = epigynal median sector; arrows to LL hood-like projections. B. Vulva, dorsal view; arrow to spermathecal stalk. BS = base of spermatheca, CD = copulatory duct, FD = fertilization ducts, HS = head of spermatheca. 362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 ; Www 00'L Wu 00'F FIGURE 13. Chrosioderma albidum Simon, left male palp. A. Ventral view. B. Close up of ventral view, E=embolus, Pp=pars pendula, Tsp = tegular sclerotized process. C. Retrolateral view. D. Prolateral view. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 363 05mm Figure 14. Chrosioderma albidum Simon. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like pro- jections. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fer- tilization ducts, HS = head of spermatheca, MS = epigynal median sector. 364 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 FiGure 15. Chrosioderma analalava sp. nov. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like pro- jections. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fer- tilization ducts, HS = head of spermatheca, MS = epigynal median sector. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 365 wi 9'0 Ae) FiGurE 16. Chrosioderma havia sp. nov., left male palp. A. Ventral view, arrow to tegular sclerotized process. B. Retrolateral view. C. Prolateral view. D. Close up of prolateral view; arrow to subtegulum. Pp = pars pendula, ST = sub- tegulum. 366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 0.5mm 0.5mm FIGURE 17. Chrosioderma havia sp. nov. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like projec- tions. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fertil- ization ducts, HS = head of spermatheca, MS = epigynal median sector. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 367 Wu g'9 ML ub FIGURE 18. Chrosioderma mahavelona sp. nov., left male palp. A. Ventral view. B. Retrolateral view. C. Prolateral view, arrow to subtegulum. Eb = embolus base, T = tegulum. 368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 wu } FIGURE 19. Chrosioderma mipentinapentina sp. nov., left male palp. A. Ventral view. B. Close up, ventral; arrow to sub- tegulum, T = tegulum. C. Retrolateral view. D. Prolateral view. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 369 ww | ww) ww | FIGURE 20. Chrosioderma namoroka sp. novy., left male palp. A. Ventral view. B. Close up, ventral view, Tsp = tegular sclerotized process. C. Retrolateral view. D. Prolateral view, arrow to subtegulum. 370 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 uw | C FIGURE 21. Chrosioderma ranomafana sp. nov., holotype, left male palp. A. Ventral view. B. Close up of ventral view; arrow to subtegulum. C. Retrolateral view. D. Prolateral view. C = conductor, Eb = embolus base, T = tegulum. / ¥ SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON wy He. FIGURE 22. Chrosioderma ranomafana sp. nov., right male palp. A. Ventral view. B. Close up of retrolateral view. C. Prolateral view, arrow to conductor. D. Close up of cymbium showing position of tarsal organ, retrolateral view; inset to tarsal organ. C = conductor, Cym = cymbium, E = embolus, Eb = embolus base, ST = subtegulum, T = tegulum. 3D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 FIGURE 23. Chrosioderma ranomafana sp. nov. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like projections. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fertilization ducts, HS = head of spermatheca, MS = epigynal median sector. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON 3713 0.18 mm FIGURE 24. Chrosioderma roaloha sp. nov. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like pro- jections. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fer- tilization ducts, HS = head of spermatheca, MS = epigynal median sector. 374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 ww 90 wu 9°0 wu 9'0 FIGURE 25. Chrosioderma soalala sp. nov., holotype, left male palp. A. Ventral view. B. Prolateral view. C. Retrolateral view, arrow to embolus base. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON FIGURE 26. Chrosioderma soalala sp. nov. (CASENT 9005038), right male palp. A. Ventral view. B. Retrolateral view. C. Prolateral view. D. Close up of prolateral view. AN = anelli, E = emboius, Ep = embolic process, RTA = retrolateral tib- ial apophysis, ST = subtegulum, T = tegulum. 376 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 25 0.5mm FIGURE 27. Chrosioderma soalala sp. nov. Female genitalia. A. Epigynum, ventral view; arrows to LL hood-like pro- jections. B. Vulva, dorsal view; arrow to stalk of spermatheca. BS = base of spermatheca, CD = copulatory duct, FD = fer- tilization ducts, HS = head of spermatheca, MS = epigynal median sector. SILVA-DAVILA: REVISION OF THE SPIDER GENUS CHROSIODERMA SIMON ii ranomafana soalala namoroka mipentinapentina Juveniles mahavelona roaloha analalava havia albidum 100 200 300 Km FIGURE 28. Distribution range for Chrosioderma species. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Volume 56, No. 26, pp. 379-390, 13 figs. August 26, 2005 A New Genus and Species of Gorgonian Octocoral (Anthozoa: Plexauridae) from Antarctic Waters Gary C. Williams! and Pablo J. Lépez-Gonzalez* ! Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, California 94103 USA; Email: gwilliams@calacademy.org; 2 Departmamento de Fisiologia y Biologia Animal, Facultad de Biologia, Universidad de Sevilla, Reina Mercedes 6, 41012-Sevilla, Spain; Email: pjlopez@us.es A remarkable new genus and species of gorgonian octocoral is described from the Scotia Sea in the South Atlantic Ocean. Although familial placement is problematic due to shared characteristics particularly with regard to the sclerites, the new taxon is here assigned to the Plexauridae because of features consistent with that family pertaining to morphology of the axis and polyps. Perhaps the most striking attribute of the colonies are the modifications of the margin of the calyces, caused by symbi- otic polychaete worms. This association can produce finger-like, plate-like, or star- shaped processes. Additionally, arching processes from adjacent polyps can fuse at their apices to form arcades. Recent southern polar and subpolar investigations via R/V Polarstern and other endeavors have revealed a wealth of marine invertebrate taxa, many new to science. Included among these are new genera of octocorals such as Arntzia (Primnoidae), Gilibelemnon (Stachyptilidae), Sphaeralcyon (Alcyoniidae), Rosgorgia (Subergorgiidae), and one described herein. Consequently, these efforts have significantly increased our knowledge of the Antarctic benthos (Arntz, Gutt, and Klages 1997; Lopez-Gonzalez and Gili 2001; Lopez-Gonzalez, Gili, and Orejas 2002). Abbreviations used in the text are: BEIM (Biodiversidad y Ecologia de Invertebrados Marinos, University of Seville); CAS (California Academy of Sciences, San Francisco); SMF (Senckenberg Museum, Frankfurt); USNM (National Museum of Natural History, Smithsonian Institution, Washington, D.C.); ZIZMH (Zoologisches Institut und Zoologisches Museum, Hamburg). METHODS The material studied in this paper was collected on the Polarstern cruises ANT XV/3 (EASIZ- II, Ecology of the Antarctic Sea Ice Zone, 13 January to 26 March 1998), ANT XIX/3 (ANDEEP- 1, Antarctic benthic deep-sea Biodiversity: colonization history and recent community patterns, 23 January to 26 February 2002), ANT XIX/5 (LAMPOS, Latin American Polarstern Study, 3 April to 5 May 2002). All these programs were sponsored by the Alfred Wegener Institut fiir Polar-und Meeresforschung (Bremerhaven), under the auspices of the SCAR (Scientific Committee for Antarctic Research). Additional specimens were examined from the R/V Eltanin cruise of 1963. The colonies were collected using an Agassiz trawl or a bottom trawl at the Drake Passage and different localites of the Scotia Arc. The octocorals were fixed in buffered formalin (4% in sea water) then transferred to 70% ethanol. 379 380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 The underwater photos were part of a photographic transect carried out by Dieter Piepenburg (Kiel) during the Polarstern cruise ANT XV/3 (EASIZ-ID), stn 333, King George Island, 61°26.97'S 58° 07.87'W, 975—932 m depth, 19 March 1998. The total number of photographs taken at that station was 44, each photograph covering an area of approximately | m2. SYSTEMATIC ACCOUNT Family Plexauridae Gray, 1859 Gorgonian octocorals often form copiously branched fans, but can also be sparsely branched; rarely whip-like and unbranched. Axis brown or black, often tough, fibrous, and flexible; larger colonies often have axes that are woody and relatively soft in texture. Axis composed of a hollow, relatively wide, cross chambered central core, with a surrounding area of concentric layers of gor- gonin containing open spaces (loculi). These spaces may be filled with calcitic calcium carbonate, not in the form of sclerites. The holdfast region may contain aragonitic calcium carbonate. Polyps monomorphic, retractile directly into coenenchyme, or often the anthocodiae retract into conspicuous calyces. Anthocodiae often with crown and points, but with a distinct neck zone that contains few or no sclerites. In most cases, at least some of the sclerites of the anthocodiae, calyces, and coenenchyme, measure between 0.3 and 5.0 mm in length. Sclerites are mostly conspicuously tuberculated or thorny, with tubercles generally not arranged in regular whorls. Sclerite form highly diverse, including spindles, thornstars, thornscales, rooted leaves, stellate plates, rosettes, ovals, crutches, bifurcated rods, crescents, leaf spindles, leaf clubs, capstans, torches, double disks, and tuberulat- ed spheroids. The Plexauridae is a morphologically and taxonomically diverse family of approximately 38 genera, with widespread distribution in the Atlantic, Indian, Pacific, and Southern Oceans. Bayergorgia Williams and Lopez-Gonzalez, gen. nov. Type species: Bayergorgia vermidoma Williams and Lopez-Gonzales, sp. nov. DIAGNosis.— Colonies mostly unbranched. Axis proteinaceous with hollow cross-chambered core and loculated cortex. Anthocodiae retractile into prominent calyces. Margins of calyces often morphologically modified by symbiotic polychaetes. Polyps monomorphic and azooxanthellate. Coenenchymal sclerites are spindles < 0.9 mm in length and spiny rods < 0.4 mm long. Calyces with needle-like spindles < 1.2 mm long. Anthocodial sclerites present, neck zone sclerites absent. Sclerites colorless. ETYMOLOGy.— The new generic name is named for our colleague, Dr. Frederick M. Bayer, curator emeritus of the National Museum of Natural History, Smithsonian Institution; in combina- tion with the Greek word, gorgia (a commonly used suffix in octocorallian nomenclature). Dr. Bayer began preliminary work on this taxon in the 1980s, but the project was subsequently discon- tinued by him. Bayergorgia vermidoma Williams and Lopez-Gonzalez, sp. nov. Figs. 1-13. MATERIAL EXAMINED.— Hototyre: CAS #171740; Drake Passage, 61°26.08’S 58°06.2'W; 1047— 1227 m depth, 19 March 1998; Polarstern cruise ANT XV/3, stn 336, Agassiz trawl; single whole colony wet preserved, purple-violet in life. PARATYPES: CAS #1717141: same data as holotype; single fragmented colony wet preserved, purple-violet in life. ZIZMH #C11691:; Drake Passage, 61°26.08’S 58°06.2'W; 1047-1227 m WILLIAMS AND LOPEZ-GONZALEZ: NEW GORGONIAN OCTOCORAL 381 FIGURE 1. A. Polyps of Anthomuricea argentea; from Wright and Studer, 1889: plt. 23, fig. 1, scale not given; collect- ed from Patagonia (southern Atlantic), 256 m depth. B. Distal region of a living colony of Bayergorgia vermidoma gen. and Sp. nov.; portion shown approximately 65 mm in length; photograph of terminal portion of a colony (ANT-3337) by Martin Rauschert.. C. Jn situ photograph of Bayergorgia vermidoma gen. and sp. nov.; underwater photograph by Dieter Piepenburg, Polarstern Cruise ANT XV/3, Stn. 333, 61°26.8’S 58°08.1’W; 994 m depth; 19 March 1998; scale not given. depth; 19 March 1998; Polarstern cruise ANT XV/3, stn 336, Agassiz trawl; two colonies. SMF #(unavail- able at time of publication); Drake Passage; 61°26.08’S 58°06.2’W; 1047-1227 m depth; 19 March 1998; Polarstern cruise ANT XV/3, stn 336, Agassiz trawl; two colonies. OTHER MATERIAL: BEIM #ANT-1550; NW Elephant Island, 61°18.14’S 56°8.27'W, 281.1—288.6 m, 2 February 2002, Polarstern cruise ANT XIX/3, stn 61-1, bottom trawl; two colonies. BEIM #ANT-1622; NW Elephant Island, 61°18.14'S 56°8.27' W, 281.1—288.6 m, 2 February 2002; Polarstern cruise ANT XIX/3, stn 382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 61-1, bottom trawl; one colony. BEIM #ANT-1695; Shag Rocks, 53°23.59'S 42°41.78'W;, Polarstern cruise ANT XIX/5, stn 145; 223.6-307.1 m depth; 9 April 2002; three colonies. CAS #171742; Shag Rocks, 53°23.59'S 42°41.78' W; Polarstern cruise ANT XIX/5, stn 145; 223.6-307.1 m depth; 9 April 2002; one frag- mented colony. CAS # 171927; Shag Rocks, 53°23.59'S 42°41.78' W; Polarstern cruise ANT XIX/5, stn 145; 223.6-307.1 m depth; 9 April 2002; one partial colony. USNM #85223; South Atlantic Ocean, Antarctica, Antarctic Peninsula, South Shetland Islands, West of Elephant Island; 61°19'S 056°28’W; 403 m depth; 13 March 1964; coll. USARP, E/tanin R/V; one fragmented colony and one whole colony. USNM #85226; South Atlantic Ocean, Scotia Sea, South Georgia Island, off west tip of island; 54°29'S 039°22’ W; 659-686 m depth; 8 February 1966; coll. Eltanin R/V; one fragmented colony. USNM #85224; South Atlantic Ocean, Scotia Sea, South Georgia Island; 54°55’S 038°05’W; 595-677 m depth; 25 August 1963; coll. E/tanin R/V; one frag- mented colony. USNM #85225; South Atlantic Ocean, Scotia Sea, Antarctica, South Orkney Islands, Scotia Ridge, West of Islands; 60°22’S 046°50' W; 298-403 m depth; 15 April 1964; coll. USARP, Eltanin R/V; one fragmented colony and one whole colony. DESCRIPTION OF HOLOTYPE.— Axis: Axis smooth and continuous, brown, dark brown, or black in color, with a conspicuously hollow but cross-chambered central core and a surrounding cortex that is loculated and uncalcified. Axis mostly 1.5 mm in width, and 30 mm in length from the base of the holdfast to the proximal region of the polyp-bearing portion of the colony; axis at base of holdfast 3.0 mm wide. Growth form and size: Growth form is upright and unbranched. The colony is 145 mm in total length, and 10-12 mm wide in the polyp bearing region. The stalk, or region of the colony devoid of polyps between the holdfast and proximal region of the polypary, is 30 mm in length, whereas the polyp bearing portion of the colony is approximately 115 mm long. Polyps: Anthocodiae are retractile into prominent calyces. Sclerites are absent in the neck zones (introverts). The anthocodiae have eight intersceptal tracts of more-or-less longitudinally placed spindles below the tentacles, whereas a crown is absent. The calyces are cylindrical in shape with eight intersceptal tracts of spindles mostly longitudinally placed, or somewhat oblique and converging distally to form weak chevrons. The tentacles have somewhat flattened rods in the rachises, which are presumably absent from the pinnules. Sclerites: The coenenchyme contains two main kinds of sclerites. In the regions near the bases of the calyces, are needle-like spindles 0.60—0.90 mm long, that are mostly smooth in the midsec- tions with tuberculation toward the ends (Fig. 6, top row). In the region of the stem below the poly- pary and in the flat regions between the calyces are tuberculated rods 0.12—0.27 mm long, that vary in shape from relatively thin and elongate (Fig. 7) to blunt and more robust (Fig. 8), as in the axial sheath. Immediately below the superficial coenenchyme Is a very thin layer containing these short, blunt rods. Also present in the coenenchyme are sclerites of intermediate form and size 0.24—0.65 mm (Fig. 6, second and third rows). The calyces contain needles 0.6—1.2 mm in length, acutely pointed, with relatively sharp tubercles toward both ends (Fig. 9). These sclerites are either straight or somewhat curved and are longitudinally placed or in some polyps are oblique and converge dis- tally to form chevrons. The anthocodiae have eight intersceptal tracts below the bases of the tenta- cles that resemble points formed by spindles 0.21—0.65 mm long. These sclerites are more or less longitudinally placed. The sclerites are mostly straight, vary in the proportion of width vs. length, and are sparsely tubercated near the ends (Figs. 10-11). The transverse placement of sclerites to form a crown below the points is not evident. The rachises of the tentacles contain somewhat flat- tened rods and spindles 0.20—0.26 mm long, sparsely and shallowly tubercated on opposite mar- gins, most distinctly toward the ends of the sclerites (Fig. 12). These sclerites are not evident in the pinnules. Color: Color in life is brilliant magenta (red-purple) to violet-purple (Fig. 1), fading to a pale tan-white or grey-white in alcohol (Fig. 2). The vivid coloration is therefore most probably provid- WILLIAMS AND LOPEZ-GONZALEZ: NEW GORGONIAN OCTOCORAL 383 ed by alcohol-soluble pigments. Sclerites are colorless. DISTRIBUTION.— Antarctic Region: Archipelagos of the Scotia Sea, the northwestern Antarctic Peninsula, and Drake Passage (Fig. 13); depth range 223-1227 meters. Photographic transects of the seafloor were obtained during the expeditions ANT XV/3 in the King George area (Arntz and Gutt 1999). Each photograph covered approximately one square meter of the seabed. Bayergorgia vermidoma gen. and sp. nov. was present at the photographic transect Station 333 (Latitude: 61°26.97'S; Longitude 58°07.87' W; depth range 975-932 m). Three specimens of the new taxon appeared in three of forty-four photographs taken along the transect.(N. Teixid6, pers. commun.). Its abundance is therefore considered very low with a patchy pattern of distribution. EtyMoLoGy.— The specific epithet is derived from the Latin, vermis (worm) and domus (home or house), in reference to presence of commensal polychaetes that cause and inhabit the modified calyces of some polyps. REMARKS.—Two specimens (USNM 85229 and USNM 85230), both from Shag Rocks (Scotia Sea), are morphologically very similar to the new taxon with the exception that they show very sparse dichotomous or irregular branching. This material has not been included for examina- tion in the present study, but is planned for a subsequent assessment. DISCUSSION Familial Assignment: The new taxon shares general characteristics of the sclerites or axis with several gorgonian families — particularly the Acanthogorgiidae and the Plexauridae, therefore its proper placement as to family is problematic. The polyps of acanthogorgiid species are not retrac- tile and are not separable into functional anthocodiae and calyces, the tentacles contract over the oral disc, and the neck zones are covered with spindles commonly arranged in eight double rows forming inverted ‘V’s. On the other hand, the polyps of plexaurids are either completely retractile into the coenenchyme or are distinctly differentiated into anthocodiae and calyces, whereas the neck zones have few or no sclerites (Bayer 1981; Fabricius and Alderslade 2001). Characteristics of the polyps of the new taxon conform to those of the Plexauridae as outlined above. Symbiotic polychaetes: Perhaps the most remarkable aspect of the new taxon is the modifica- tion of the calyx due to a symbiotic association with polychaete worms. The margin of the calyx often forms finger-like (Fig.4C), star-shaped, or plate-like extensions (Fig. 4B). Occasionally, the modifications of adjacent polyps curve upwards and fuse at their tips forming arches or arcades (Fig. 4A). The worms that cause these modifications are presumably capable of moving around inside these extensions. In polyps with the most severely modified calyces, disk-like expansions result from the fusion of the prolongations and the filling in of the spaces between these prolonga- tions with coenenchymal material. Taxonomic identification of the worms as well as details of the association with the new taxon as host organism, are currently under investigation. Martin, Nufiez, Riera, and Gil (2002) record the presence of polychaetes of the genus Haplosyllis (Family Syllidae) with other gorgonians, namely the plexaurid genera Paramuricea and Villogorgia, and the acanthogorgiid genus Anthogorgia. These associations are described as symbiotic and either kleptoparasitic (stealing food from the host), or parasitic (feeding on the host), and at least partially mutualistic (cleaning the host). Related Taxon (Fig. 1A): A superficially similar plexaurid gorgonian, tentatively identified as Anthomuricea argentea Wright and Studer (189:103—104; pl. 23, fig. 1), which is also noted in Bayer (1981) and Grasshoff (1977), differs from Bayergorgia vermidoma gen. and sp. nov. in hav- 384 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 ing dichotomous branching, more copious branching, white coloration in life, infrequent to absent calyx modification from worm symbiosis, and a geographical restriction to the Patagonian Atlantic and Burdwood Bank (256 m depth). Recently collected material has allowed for a comparison with the new taxon and the potential for a redescription of Wright and Studer’s taxon, both of which are planned for future study. Grasshoff (1977) maintains that Anthomuricea is related to the more northern Atlantic genus Swiftia, with four or five species in the western Atlantic (Deichmann 1936: 186) and three species in the northern Atlantic, eastern Atlantic, and Mediterranean (Grasshoff 1977-62). ACKNOWLEDGMENTS We acknowledge the valuable assistance of the officers and crew of the Polarstern, and many colleagues on board during the EASIZ-I, ANDDEP-I and Lampos cruises. We take this opportu- nity to extend our thanks to the cruise leaders and steering committee, especially Wolf E. Arntz, Dieter Fiitterer, and Angelika Brandt, who kindly facilitated the work on board, and for the oppor- tunity to collaborate in these Antarctic programs. Special thanks are extended to some friends and colleagues for their valuable assistance on board, especially M. Isabel Alfonso, Nuria Te1xid6, Mercedes Conradi, Neus Vert and Estefania Rodriguez. Our thanks to Martin Rauschert for the photograph of a living colony (Fig. 1B) taken on board during the EASIZ-II cruise. Dieter Piepenburg and Nuria Teixido kindly provided the underwater photos and the information about the abundance of the new species in the photograph- ic transect carried out at the station 333 during the EASIZ-II cruise. Participation on these cruises was supported by the Spanish CICYT projects ANT 98-173-E, REN2001-4269-E/ANT, REN2001- 4929-E. We express our gratitude to Stephen D. Cairns (United States National Museum, Smithsonian Institution, Washington, D.C.) for the loan of specimens and unpublished study materials relating to this new taxon. LITERATURE CITED ARNTZ, W.E., J. GuTT, AND M. KLAGes. 1997. Antarctic marine biodiversity — An overview. Pages 3-14 in Battaglia B., J. Valencia, and D.W.H. Walton, eds., Antarctic Communities, Species, Structure and Survival. Cambridge University Press, Cambridge, UK. ARNTZ, W.E., AND J. GuTt. 1999. The expedition ANTARKTIS XV/3 (EASIZ II) of RV ‘Polarstern’ to the eastern Weddell Sea in 1998. Ber Polarforsch 301:1—229. BAYER. F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with diagnoses of new taxa. Proceedings of the Biological Society of Washington 94(3): 902-947. DEICHMANN, E. 1936. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum of Comparative Zoology at Harvard College 3(49):1—317. FABRICIUS, K. AND P. ALDERSLADE. 2001. Soft Corals and Sea Fans — A Comprehensive Guide to the Tropical Shallow-Water Genera of the Central-West Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville, Australia. 272 pp. GRASSHOFF, M. 1977. Die Gorgonarien des éstlichen Nordatlantik und des Mittelmeeres III. Die Familie Paramuriceidae (Cnidaria, Anthozoa). Meteor Forschungsergebnisse (D) 27:5—76. LOPEZ-GONZALEZ, P.J., AND J-M GILI. 2000. A new octocoral genus (Cnidaria: Anthozoa) from Antarctic waters. Polar Biology 23:452-458. LoPEZ-GONZALEZ, P.J., AND J-M GILI. 2001. Rosgorgia inexpectata, new genus and species of Subergorgiidae (Cnidaria, Octocorallia) from off the Antarctic Peninsula. Polar Biology 24:122-126. LOPEZ-GONZALEZ, P.J., AND J-M GILI, and C. OresAs. 2002. A new primnoid genus (Anthozoa: Octocorallia) from the Southern Ocean. Scientia Marina 66(4):383-397. WILLIAMS AND LOPEZ-GONZALEZ: NEW GORGONIAN OCTOCORAL 385 LOpEZ-GONZALEZ, P.J., AND G.C. WILLIAMS. 2002. A new genus and species of sea pen (Octocorallia: Pennatulacea: Stachyptilidae) from the Antarctic Peninsula. /nvertebrate Systematics 16:919—929. Martin, D., J. NUNEZ, R. RIERA, AND J. Git. 2002. On the associations between Haplosyllis (Polychaeta, Syllidae) and gorgonians (Cnidaria, Octocorallia), with the description of a new species. Biological Journal of the Linnean Society 77(4):455-477. WRIGHT, E.P., AND T. STUDER. 1889. Report on the Alcyonaria collected by H.M.S. Challenger during the years 1873-1876. Report of the Scientific Results of the Voyage of H.M.S. Challenger 1873-1876 31(1):1-314. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 FIGURE 2. Bayergorgia vermidoma gen. and sp. nov. A-B. Photographs of wet-preserved material. A. Non-Type speci- men (USNM 85225), scale bar = 20 mm. B. Paratype (CAS 171741), scale bar = 40 mm. C. Holotype (CAS 171740), scale bar = 30 mm. WILLIAMS AND LOPEZ-GONZALEZ: NEW GORGONIAN OCTOCORAL 387 FIGURE 3. Bayergorgia vermidoma gen. and sp. nov. A, scale bars MRS D. Scanning electron micrographs of unmodifed polyps, =1 mm showing calyces and anthocodiae. C. Single polyp showing two digitate modifications of the calyx on either side of the anthocodia. Scale bars = | mm. FIGURE 4. Bayergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of polyps. A. Two polyps with calicular prolongations fused together apically to form an arch. B. Single polyp with a broad and flattened calicular expansion. Entire retracted polyp. C. Single polyp with curved fingerlike calicular expansion. Scale bars = | mm. 388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 FIGURE 5 (upper left). Bavergorgia vermidoma gen. and sp. nov. AB. Scanning electron micrographs of two polyps with apically fused calicular modifications (also shown in figure 4A). Scale bars = 1 mm. FIGURE 6 (upper right). Bayergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of sclerites. Variation in coenenchymal sclerites from the polyp-bearing portion of a colony, scale bar = 0.10 mm. Inset shows tip of a single coenenchymal sclerite, scale bar = 0.02 mm. FIGURE 7 (lower left). Bayergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of sclerites. Coenenchymal sclerites. A. 0.24 mm. B. 0.19 mm. C. 0.17 mm. D. 0.24 mm. E. 0.28 mm. F. 0.19 mm. G. 0.26 mm. H. 0.27 mm. I. 0.23 mm. J. 0.20 mm. K. 0.38 mm. L. 0.19 mm. M. 0.17 mm. N. 0.24 mm. O. 0.24 mm. WILLIAMS AND LOPEZ-GONZALEZ: NEW GORGONIAN OCTOCORAL 389 FIGURE 8 (upper left). Bayvergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of sclerites. Sclerites from the axial sheath. A. 0.27 mm. B. 0.20 mm. C. 0.12 mm. D. 0.17 mm. E. 0.17 mm. F. 0.20 mm. G. 0.19 mm. H. 0.19 mm. I. 0.14 mm. J. 0.18 mm. K. 0.22 mm. FIGURE 9 (upper right). Bayergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of sclerites. Variation of sclerites from the calyx of a polyp, scale bar = 0.40 mm. Inset shows enlargements of the tips of two calicular scle- rites, scale bar = 0.15 mm. FiGurE 10 (lower left). Bayergorgia vermidoma gen. and sp. noy. Scanning electron micrographs of sclerites. Anthocodial sclerites. A. 0.26 mm. B. 0.38 mm. C. 0.22 mm. D. 0.21 mm. E. 0.52 mm. F. 0.40 mm. G. 0.48 mm. H. 0.22 mm. I. 0.11 mm. J. 0.36 mm. 390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. 26 FIGURE 12. Bayergorgia vermidoma gen. and sp. nov. Scanning electron micrographs of sclerites. Sclerites from FIGURE 11. Bayergorgia vermidoma gen. and sp. noy. the rachis of the tentacles; top scale bar = 0.15 mm, bottom Scanning electron micrographs of sclerites. Anthocodial scale bar = 0.20 mm. sclerites; scale bar = 0.20 mm. (¢) Atlantic Ocean Bouvet {* Island Tropic of Capricorn South South Georgia Sandwich ‘Islands | Falkland South _———_—_—__ islands ,- Orkney dp Scotia B\siands < : _ Antarctic ™™ 17 — > Bs. Peninsula ~ Ir South \Y-._ Weddell / Shetland /~ “\ Sea 7 Islands C —— Pacific Ocean é West Antarctica ~~ Patagonia L- * @ Sea Ss | +—3 South Pole Antarctic Circle aw, 180 \ FIGURE 13. Map of the Scotia Sea showing collecting stations for Bayergorgia vermidoma gen. and sp. noy. Arrow points to type locality. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. GONZALEZ ET AL.: NEW RECORDS FOR SCINCELLA VICTORIANA 391 Short Communications NEW RECORDS FOR SCINCELLA VICTORIANA (SHREVE, 1940) FROM THE CHIN HILLS, MyANMAR.— Heretofore Scincella victoriana was known from two specimens (Ouboter 1986) (MCZ 44738-44739) from the type locality of “Mt. Victoria, 2800 meters, Pakokku-Chin Hills, Burma” (Shreve 1940). While conducting herpetological surveys in 2001 and 2003 in the Chin Hills of western Myanmar, the California Academy of Sciences (CAS)/Myanmar Forest Department field team collected three additional specimens of S. victoriana. These were collected from the following localities (coordinates were taken using a Garmin 12 GPS, Datum WGS84): CAS 220628 (Figs. la—b), Myanmar, Chin State, Kan Pet Let Township, Nat Ma Taung National Park, 21°12'46.4"N, 94°00'54.7”E, ca. 2361 m, collected by Htun Win, Thin Thin, Awan Khwi Shein and Hla Tun on 4 March 2001; CAS 220629, Myanmar, Chin State, Kan Pet Let Township, Nat Ma Taung National Park, 21°13'19.7'N, 93°57'19.3”E, ca. 2676 m, collected by Htun Win, Thin Thin, Awan Khwi Shein and Hla Tun on 5 March 2001; and CAS 231488 Myanmar, Chin State, Falam Township, Laiva Forest Reserve, 20°50'35.5"N, 93°31'15.1"E, ca. 1954 m, collected by Htun Win, Kyi Soe Lwin and Awan Khwi Shein on 17 July 2003. 5 Ege 4 ope s , ¥ te ¢ gig FiGuRES la—b. Scincella victoriana (CAS 220628). a. Closeup of head to show details of pattern of dark and light mark- ings; b. Whole body. Photographs by Hla Tun. CAS 220628 and 220629 were collected on the slopes of Mount Victoria at a slightly lower elevation than the type locality. CAS 231488 extends the distribution of the species approximate- ly 183 km north of Mount Victoria, suggesting that the species ranges rather widely throughout the Chin Hills. The specimens were collected between 1102 and 1340 hours at air temperatures between 25.6—27.4°C. CAS 220628 and 220629 were collected in pine forest and CAS 231488 was collected in mixed hardwood and pine forest. Measurements for the paratype and new specimens are given in Table 1. Meristic characters that differ between the paratype and the new specimens are listed in Table 2. Scale counts for nuchals, supraoculars, loreals, supralabials, and midbody scale rows are as described by Shreve (1940). There are 7 supralabials and 6 infralabials on either side of all four specimens. The coloration in alcohol is similar to Shreve’s description except that CAS 220628 lacks gold spotting forming longitudinal lines on the back. We thank U Shwe Kyaw (Director General) and U Khin Maung Zaw (Director, Nature and Wildlife Conservation Division), Forest Department for their support of the Myanmar Herpetological Survey. José P. Rosado and James Hanken, Museum of Comparative Zoology, Harvard University, kindly provided us with the paratype of Scincella victoriana. Financial support for this project was provided by NSF grant DEB-9917861. 392 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, Short Communications TABLE. |. Mensural characters (in mm) for the paratype and the three new specimens of S. victoriana. MCZ 44739 CAS 220628 CAS 220629 CAS 231488 Snout to vent length 41.4 60.6 76.7 66.6 Axial to anterior insertion of hind limb length Pell 313 44.2 37.4 Hind limb length 14.6 17.4 23.9 Dill 4th finger length BP) 3.8 3] B07 4th toe length 4.8 6.9 8.0 7.3 Head length 7.8 10.0 ilcit 9.7 Head width 6.7 8.1 9.4 8.5 Snout to eye distance 3.7 4.5 53 4.7 Nares to orbit distance 2 DES 33 2.4 Horizontal eye diameter 2.6 2.9 3.0 2.6 Eye to ear distance 333 4.3 4.9 43 Palpebral disk horizontal diameter 1.2 1.4 1.5 1.5 TABLE 2. Meristic characters differing between the paratype and new specimens of S. victoriana. MVZ 44739 CAS 220628 CAS 220629 CAS 231488 Supraciliaries (right/left) 7/6 5/5 6/5 2/5 Dorsal scale rows 53 56 Sy 56 Right 4) finger subdigital lamellae 12 12 11 11 Right 4" toe subdigital lamellae 16 16 15 16 LITERATURE CITED OusotTeR, P.E. 1986. A revision of the genus Scincella (Reptilia, Sauria, Scincidae) of Asia, with some notes on its evolution. Zoologische Verhandelingen 229:1—66. SHREVE, B. 1940. Reptiles and amphibians from Burma with descriptions of three new skincs [sic.]. Proceedings of the New England Zodélogical Club 18:17-26. — Marlene Gonzalez!, Kyi Soe Lwin?, and Jens V. Vindum?. ! College of Arts and Sciences, University of San Francisco, 2130 Fulton Street, San Francisco, CA 94117; 7 Nature and Wildlife Conservation Division, Forest Department, Ministry of Forestry, Bayinmaung Road, West Gyogone, Insein, Yangon, Myanmar; > Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103; Email: jvindum@calacademy.org. A NOTE ON AMARANTHUS VIRIDIS IN THE CALIFORNIA FLORA.— In their annotated checklist of the native and naturalized plants of San Francisco County, Howell et al. (1958) reported sever- al new distribution records for California. One of these, Amaranthus gracilis Desf. ex Poir., was noted as a sidewalk weed in the Richmond District of San Francisco. A collection with several duplicates (Howell 32939 at CAS and DS) was made in 1957 to document the occurrence of this amaranth. Howell identified and annotated the specimens as 4. viridis L. and sent them to be ver- ified by J. Sauer, an authority on amaranths who was then working on several species of weedy plants in northern California (Tucker and Sauer 1958). Sauer confirmed the identification but noted that the name “4. viridis is a nomen confusum and must be rejected” (Howell et al. 1958). The next available name for the species was A. gracilis, with which name Sauer annotated Howell’s collec- DANIEL: AMARANTAUS VIRIDIS IN THE CALIFORNIA FLORA 393 tion at CAS in 1958. In the supplement to his 4 California Flora, Munz (1968) indicated that A. gracilis had been reported from San Francisco and that it resembles 4. deflexus L. except for its “wrinkled seeds.” Howell et al. (1958) had noted that these species could be distinguished by the wrinkled (vs. smooth) utricles of A. gracilis. In The Jepson Manual, Henrickson (1993) noted under 4. deflexus that plants “with wrinkled seeds have been called A. gracilis Desf.” All species of Amaranthus in California have generally smooth seeds (at least at magnifications to 50x) but there is variation in the ornamentation of the fruit (utricle) that closely envelops the seed, and it is undoubtedly this feature that both Munz (1968) and Henrickson (1993) intended. Recent treat- ments of Amaranthus (e.g., Allen and Akeroyd 1993; Mosyakin and Robertson 2003) use Linnaeus’ name, A. viridis, for this species and include A. gracilis as a synonym. In treating Amaranthus for a new taxonomic account of the plants in San Francisco County (including 8 species of Amaranthus; Daniel in preparation), | had the opportunity to compare var- ious published treatments of the genus with specimens collected in the county. Henrickson’s (1993) key works very well for most plants from San Francisco. However, his implications that the plants reported from California as A. gracilis (1.e., Howell’s collection) are similar to those of A. deflexus (except for the “wrinkled seeds”) and that these might represent the same taxon are inaccurate. These two species appear abundantly distinct (in fact, more so than several other species of ama- ranth in the California flora, e.g., A. hvbridus L. and A. powellii S. Wats; cf. Costea et al. 2001a). Although A. viridis was not treated as occurring in California in either The Jepson Manual or the Flora of North America, both it and A. deflexus have been collected in the state. These two species can be distinguished from one another by the characters in the following couplet: Mature utricles ovoid to ellipsoid, 2.5—3.2 mm long, smooth to somewhat wrinkled, inflated and much exceeding the size of the seed within; female flowers with 2 perianth segments (in San Francisco, elsewhere 2—3). A. deflexus Mature utricles subglobose, 1.3—1.7 mm long, prominently warty-roughened, not inflated and barely exceeding the size of the seed within; female flowers with 3-4 perianth segments. A. viridis They differ from most other species of Amaranthus in California by their indehiscent utricles. In other species the utricles either dehisce circumsessily (most species) or irregularly (4. spinosus L.); however, on occasion, species having utricles that usually dehisce may also possess fruits that remain indehiscent. Another species with indehiscent utricles, A. blitum L., has been noted to occur in California (Hrusa et al. 2002; Mosyakin and Robertson 2003). It was not treated by Hendrickson (1993) or Costea et al. (2001b) because it has only very recently been documented from the state. It appears to have been introduced with bedding plants raised outside of California and offered for sale at nurseries within the state (A. Sanders, pers. commun.). Its utricles resemble those of A. deflexus by their generally smooth surfaces; they differ from fruits of that species by their shape (subglobose to obovate) and lack of inflation (1.e., they do not appreciably exceed the size of the seed within). Amaranthus viridis always has conspicuously warty-roughened utricles (Fig. 1c). Variations in shape, size, and surface texture were observed among utricles of A. deflexus (Fig. la—b) in San Francisco County. The surfaces of utricles from some plants have wrinkles (e.g., Daniel 8792 from Bernal Heights) whereas others do not and are smooth (e.g., Cannon s.n. in 1895 from Post St.). This variation is likely responsible for some of the confusion between the two species in California. Neither taxon is believed to be indigenous to California, but their native ranges are not known with certainty. Amaranthus viridis is usually noted as native to South America but introduced into tropical and subtropical regions worldwide. Mosyakin and Robertson (2003) indicated that it is known from much of the eastern and southern United States to Arizona. Within California, A. viridis is still known only from San Francisco where several plants occurred in or along sidewalks. 394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 56, No. X It has not been relocated in the county since Howell’s collection, but it should be sought among amaranths that occur in disturbed habitats. Amaranthus deflexus is also believed to be native to South America and introduced into tropical or warm-temperate regions worldwide. Mosyakin and Robertson (2003) mapped its distribution in the eastern and southeastern United States, as well as on the Pacific Coast. It is ‘FIGURE 1. Perianth segments and utricles of Amaranthus. a. A. deflexus widespread in California, occur- (Cannon s.n. in 1895), with smooth surface and ovoid shape. b. 4. deflexus pi fae a : (Daniel 8792), with wrinkled surface and ellipsoid shape. c. A. viridis (Howell ring in disturbed habitats of west- 32939), with warty-roughened surface and subglobose shape. Drawn by Meg ern and central portions of the © Stalcup. State. a LITERATURE CITED ALLEN, P., AND J.R. AKEROYD. 1993. Amaranthus. Pages 130-132 in T.G. Tutin et al., eds., Flora Europaea, vol. 1, ed. 2. Cambridge University Press, Cambridge, England, UK. CosTEA, M., A. SANDERS, AND G. WAINES. 2001a. Preliminary results toward a revision of the Amaranthus hybridus species complex (Amaranthaceae). Sida 19:931—974. CosTEA, M., A. SANDERS, AND G. WAINES. 2001b. Notes on some little known Amaranthus taxa (Amaranthaceae) in the United States. Sida 19:975—992. HENRICKSON, J. 1993. Amaranthaceae. Pages 130-134 in J.C. Hickman, ed., The Jepson Manual: Higher Plants of California. University of California Press, Berkeley, USA. Howe Lt, J.T., P-H. RAVEN, AND P. RuBTzorr. 1958. A flora of San Francisco, California. The Wasmann Journal of Biology 16:1—157. Hrusa, F., B. ERTTER, A. SANDERS, G. LEPPIG, AND E. DEAN. 2002. Catalogue of non-native vascular plants occurring spontaneously in California beyond those addressed in The Jepson Manual—Part I. Madronio 49:61—91. MosyYAkIN, S.L., AND K.R. ROBERTSON. 2003. Amaranthus. Pages 410-455 in Flora of North America, vol. 4(1). Oxford University Press, New York, USA. Munz, P.A. 1968. Supplement to a Flora of California. Pages 1-224 in P.A. Munz and D.D. Keck, A California Flora (2nd printing). University of California Press, Berkeley, California, USA. Tucker, J.M., AND J.D. SAUER. 1958. Aberrant Amaranthus populations of the Sacramento-San Joaquin Delta, California. Madrofio 14:252-261. — Thomas F. Daniel. Department of Botany, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103; Email: tdaniel@calacademy.org. Copyright © 2005 by the California Academy of Sciences San Francisco, California, U.S.A. << ‘PSTaecve Daa’ fe? AP Thee an " T jeAl OSL. a 4 i in of aD | eis [ee gy tly ® evie -pe | Cage @ y ae. 7 =n > Ae Dimes qh), eed — g ¥ ‘ as Ps - » si => is o rte Ob bd y ' — i Viger : F - ws Oe wee ie , f 4 = a , 1 } 7 a c. = i, — } ! 7 -7T 5 ’ eal ae = . ss _ P ; 5 » Sa 7 5 as io pa a4 h ae ‘WiWrr ; mm & fin? *, = ors aX iq ye? Beh |) Sad r 2 af " ‘ ~ ce = = a > 7 asi, 7 e" wes Chae @ a7 j ‘ i i ; in é ne ii — e “ts nh ie ni os ( 1d, , ; rir DG cee ee a ni fal ey FA fiw ty ae ; r tai | i a ge, A me bends wk ec = he ye ove ieatiee YC Wiles rh ae ayes) ee oe ani i To » “yt \ ' im; te ‘Md Gua : es iy HG? ‘ Resteg Aig Po iy bt me - ; : : 7 7 oe coer - june eo) se! a ao) = er 83 — 7 aie - “ees, Oy), Ste aes | ss an eof a _. 3 7c a = eae a, ne mt 5 7 | o oe 2 = <> 7 a ae = aw * Ve =r i - , 5 = e Se eS am “ ie PA die, *. ee 3 a & « es ime @ Ps ee tee ad a hi oe pa a CALIFORNIA ACADEMY OF SCIENCES PROCEEDINGS SERIES INSTRUCTIONS TO AUTHORS Authors planning to submit papers for consideration for publication in the Academy’s Proceedings, Occasional Papers, or Memoir series must follow the directions given below in preparing their submissions. Under some circumstances, authors may not be able to comply with all the computer-based requirements for submission. Should this be the case, please contact the Editor or Associate Editor for guidance on how best to present the materials. The Scientific Publications Office of the Academy prepares all materials for publication using state-of-the-art, computer- assisted, page-description-language software. Final copy is sent to the printer for printing. The printer does not modify the files sent for printing. Therefore, it falls to the authors to check carefully page proof when it is returned for approval. Ordinarily, all communication with authors is done via email and galley and page proofs of manuscripts, including figures, are transmitted as attachments to email communications. Again, exceptions to this will be made in the event that an author is unable to communi- cate in this way. Authors are expected to provide digital copies of both manuscript text files and images, as well as a paper printout of their manuscript. Please note the following: TexT: Text can be in Microsoft Word, as a Word document file, WordPerfect, also as a WP document file, or, best of all, as an “rtf” (rich text format) file, which can be produced by most word processors. Authors who use non-standard fonts must include file copies of those fonts so that their symbols can be reproduced accurately. However, it is strongly recommended that the type style “New Times Roman” be used throughout and that the Symbols and Bookshelf Symbol 1-3 fonts be used for such items as 3, 2, 8. (Ge ImaGEs: Images should be in either JPG (JPEG), or TIF (TIFF) format. Resolution for grayscale images should be at least 600 ppi (1200 ppi if possible, especially for photomicrographs), and 300 ppi for color. All images should be sized so that none exceeds a maximum print size of 5.5"x7.875" (140 mm x 200 mm). TABLES: Our processing software allows for direct importation of tables. This reduces the chances for errors being introduced during the preparation of manuscripts for publication. However, in order to use this feature, tables must be prepared in Microsoft Excel or in Microsoft Word using Word’s table feature; do not prepare tables using tabs or space bars. Complex tables not pre- pared as described above will be returned to the author for revision. DiGiTaL FILES: IBM or MAC formatted disks will be accepted subject to the following conditions: (a) floppy disks must not exceed 1.4 mb and (b) zip disks, preferably IBM format, must not exceed 100mb. Authors are encouraged to submit their digital files on CD-ROM (CD-R formatted disks NOT CD-RW) inasmuch as these can be read by nearly all CD-ROM drives. FILE NAMING PROTOCOLS: To facilitate the handling of digital files submitted by authors, the following file-naming conven- tions are to be followed: text files should bear the author’s last name (in the case of multiple authors, only the first author’s name) followed by a space and a date in the format mmyy (e.g., 0603 for June 2003) to yield a file name such as Gosliner 0603.doc or Williams 0603.rtf. If an author has submitted two or more manuscripts and must distinguish between them, then the naming should include an additional numeral: Gosliner1 0603.doc for the first manuscript, Gosliner2 0603.doc (or .rtf) for the second. Figures should follow similar conventions, as follows: Gosliner F1 0603.tif, Gosliner F2 0603.tif, for figures in the first manu- script and, if more than one manuscript, then Goslinerl F1 0603.tif etc. for the figures associated with the first manuscript and Gosliner2 F1 0603.tif etc. for those with the second. Following these conventions will insure that figures submitted by one author are always maintained distinct from those submitted by another. Tables submitted as Excel files should follow the same naming conventions except the file type designation will be “.xls”: e.g., Gosliner T1 0603.xls. Please note that extraneous periods are omitted in file names. BIBLIOGRAPHY FORMAT: Three bibliographic styles are accommodated in the Academy’s scientific publications, one com- monly used in scientific journals publishing papers in systematic and evolutionary biology, a second used mainly in the geologi- cal literature, and lastly, the format most commonly used in the humanities by historians of science. On request, the author will be sent a style sheet that includes samples of the three formats. Authors are also encouraged to examine a copy of the latest pub- lished Proceedings. In all instances, however, authors should not abbreviate journal names but spell them out completely. For books, the reference must include the publisher and city of publication. It is recommended that the total number of pages in the book also be given. SUBSCRIPTIONS The Proceedings series of the California Academy of Sciences is available by exchange or subscription. For information on exchanges, please contact the Academy Librarian via regular mail addressed to the Librarian, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email addressed to amalley@calacademy.org. Subscription requests, including information on rates, should be addressed to Scientific Publications, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email to the Editor at scipubs@calacademy.org. Subscription price for 2005: $65 (US) plus $10 for mailing to U.S. and Canadian addresses and $18 to all others. The Occasional Papers and Memoirs are not available by subscription. Each volume is priced separately. Occasional Papers, Memoirs, and individual issues of the Proceedings are available for purchase through the Academy Stores (http://store.yahoo.com/casstore/bookstore.html). or from the Office of Scientific Publications. Credit card orders are available only through the Academy Stores. Sg COMMENTS Foal f Address editorial] correspondence or requests for pricing information to the Editor, Scientific Publications Office, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, U.S.A. or via email to the Editor, Scientific Publications, at scipubs@calacademy.org. TABLE OF CONTENTS SANDRA V. MILLEN AND HANS BERTSCH: Two New Species of Porostome Nudibranchs (Family Dendrodorididae) from the Coasts of California (USA) and Baja California (Mexico) JEFFREY H. R. GODDARD: Ametamorphic Direct Development in Dendrodoris behrensi (Nudibranchia: Dendrodorididae), with a Review of Developmental Mode in the Family SHIREEN J. FAHEY AND ANGEL VALDES: Review of Acanthodoris Gray, 1850 with a Phylogenetic Analysis of Onchidorididae Alder and Hancock, 1845 (Mollusca, Nudibranchia) Davip H. KAVANAUGH: The California Academy of Sciences Gulf of Guinea Expedition (2001). HI. A New Species of the Endemic Genus Straneoa Basilewsky, 1953, from Sao Tomé (Insecta: Coleoptera: Carabidae: Platynini) NorMAN D. PENNY: The California Academy of Sciences Gulf of Guinea Expedition (2001). IV. The Neuroptera of Sao Tomé and Principé Islands JoAo L. M. ARANHA FILHO, PETER W. FRITSCH, FRANK ALMEDA, AND ANGELA BORGES MARTINS: Two New Species of Symplocos Jacq. (Symplocaceae) from Minas Gerais, Brazil DARRELL UBICK: New Genera and Species of Cribellate Coelotine Spiders from California (Araneae: Amaurobiidae) DIANA SILVA-DAVILA: Revision of the Spider Genus Chrosioderma Simon (Araneae: Sparassidae) Gary C. WILLIAMS AND PABLO J. LOPEZ-GONZALEZ: A New Genus and Species of Gorgonian Octocoral (Anthozoa: Plexauridae) from Antarctic Waters SHORT COMMUNICATIONS MARLENE GONZALEZ, KYI SOE LWIN, AND JENS V. VINDUM: New records for Scincella victoriana (Shreve, 1940) from the Chin Hills, Myanmar THOMAS F. DANIEL: A Note on Amaranthus viridis in the California Flora SMITHSONIAN N INSTITUTION aT Hii