NH PRIL 22, 2004 * VOLUME 55 * NUMBERS 1-12 & SHORT NOTES 7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES CALIFORNIA. ACADEMY SCIENCES ESTABLISHED 1853 SAN FRANCISCO, CALIFORNIA Copyright © 2004 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 Painting by E. Helgason (1964) of Academy’s schooner Academy (U.S. register #202055), the former U.S. Navy schooner Earnest, decked out for its 17-month voyage to the Galapagos Archipelago with new paint and new rigging. The schooner had been purchased from the Navy in May 1905 for $1000 plus an additional $50 for a new set of sails and binnacle. It was further refurbished and equipped for the Galapagos expedition at a cost of $4,550. The vessel measured 85 feet in length, 23.5 feet in width, and had a gross weight of 114 tons. The expedition departed from San Francisco 28 June 1905 and returned 29 November 1906. Its 10-man crew included Rollo Beck, expedition leader and ornithologist; Edward W. Gifford, ornithologist; Joseph S. Hunter, ornithologist; Ernest S. King, assis- tant herpetologist; Frederick T. Nelson, mate; Washington H. Ochsner, geologist and conchologist; J.J. Parker, navigator; Joseph R. Slevin, herpetologist; Alban Stewart, botanist; and Francis X. Williams, entomologist. ISSN 0068—574X Published by the California Academy of Sciences Golden Gate Park, San Francisco, California 94118 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Sit HSON/4 ry Volume 55, No. 1, pp. 1-10, 5 figs. April 22, 2004 MAY 0.2 2004 | ~~ LibRARIES Two New Species of Dorid Nudibranchs (Gastropoda, Opisthobranchia) from the Indian Ocean Terrence M. Gosliner! and David W. Behrens? ! Department of Invertebrate Zoology and Geology. California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA. Email: tgosliner@ calacademy.org; 2 Department of Invertebrate Zoology and Geology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA. Email: dave @ seachallengers.com. Two new species of cryptobranch dorid nudibranchs are described from the Indian Ocean. Chromodoris cazae sp. nov. is known from the United Arab Emirates, Gulf of Oman. This species is characterized by having a smooth white body with several irregu- lar patches of deep red-maroon found scattered on the mid-dorsal region. Each of these patches has a number of smaller irregular orange spots within it. Aldisa andersoni sp. nov. is found from Sri Lanka. This species has a series of rows of low tubercles on the blue dorsum. Black pigmentation demarcates the rows of tubercles. The species also has an opaque yellow saddle posterior to the rhinophores, and other yellow marks on the dorsum. Other species having similar coloration and that also appear to mimic phyllidid nudibranchs are discussed. Both species are distinguished from other described taxa based on differences in body coloration and external and internal morphology and anatomy. During the past several years, the described opisthobranch taxa of the western Pacific has increased dramatically (Brunckhorst 1993; Gosliner 1993, 1994; Gosliner and Draheim 1996; Gosliner and Behrens 1997; 1998a and b, 2000; Johnson and Gosliner 1998; Gosliner and Fahey 1998; Gosliner and Johnson 1999; Carlson and Hoff 2000; Fahey and Gosliner 1999a and b; Fahey and Gosliner 2001, 2003; Hamatani 2001; Yonow 2001; Dorgan et al. 2002; Ortiz and Gosliner 2003; Smith and Gosliner 2003; Gosliner and Smith 2003). The Indian Ocean, however, has a rich- ly diverse but reasonably unknown opisthobranch gastropod fauna (Gosliner 1994; Gosliner and Behrens 2000; Valdés, Mollo, and Ortea 1999: Yonow 1984, 1994, Yonow et al. 2002). Increasing awareness of differences in opisthobranch morphology by sharp-eyed, dedicated recreational divers has produced a number of interesting finds. Two such new discoveries are described here. The family Chromodorididae comprises one of the most colorful and morphologically diverse groups of dorid nudibranchs. More than 600 species have been described to date belonging to the 16 genera within the Chromodorididae. The morphological characteristics and the systematic rela- tionships of members of the family have been comprehensively discussed in recent years (Rudman 1984, 1986, 1987, 1995; Baba 1994, 1995; Hamatani 1995; Gosliner and Behrens 1998a, 2000; Johnson and Gosliner 1998; Gosliner and Johnson 1999; Schrédl 1999; Valdés et al. 1999). The Chromodoris species described here broadens the basis for the genus and adds new records to the fauna of a little studied area of the Gulf of Oman. Millen and Gosliner (1985) defined the diagnostic characters of the genus Aldisa, listing eleven species. The genus is characterized by having elongate denticulate teeth, penial spines, con- ical tubercles and reduced oral tentacles. Several additional species have been added since then (Ortea and Ballesetros 1989; Elwood et al. 2000; Perrone 2001). Perrone (2001) listed twenty 1 D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 1 species of Aldisa worldwide. He overlooked two species described the year before (Elwood et al. 2000). All but three of these species are white, yellowish or red in color. Perrone (2001) included in his list several undescribed species and the specimen shown in Debelius (1996:213 as Chromodoris sp.) as a junior synonym to Aldisa erwinkoehleri. We were fortunate to acquire the specimen (pictured in Debelius) from its collector, Dr. Charles Anderson, and describe it here as a new species. To paraphrase Dr. Bill Rudman (Sea Slug Forum, March 13, 2002), we too “used to think of Aldisa as a genus of red dorids which mimicked sponges, but now it seems they include a group of nodulose species which mimic phyllidiids.” This paper describes the anatomy and systematic relationships of two new species, one Chromodoris, from the United Arab Emirates and the one Aldisa mentioned above, from Sri Lanka. Specimens examined in this study are housed in the Department of Invertebrate Zoology and Geology of the California Academy of Sciences (CASIZ). SPECIES DESCRIPTIONS Chromodorididae Bergh, 1891 Chromodoris Alder and Hancock, 1855 Chromodoris cazae Gosliner and Behrens, sp. nov. (Figs. 1A, 2-3) Chromodoris sp. Coleman, 2001:76. MATERIAL EXAMINED.— HOLOTYPE (CASIZ 168299). Rashid Wreck, Khawr Fakkan, United Arab Emirates, 12 m, December 1, 1999. PARATYPE (CASIZ 168300). Jumeirah Reef, Khawr Fakken, United Arab Emirates, 12m, December 23, 2000. DISTRIBUTION.— Known only from the Gulf of Oman, United Arab Emirates. ETYMOLOGY.— Chromodoris cazae is named in recognition of Carole Harris, Dubai, United Arab Emirates, who was the first to bring this species to our attention and was instrumental in col- lecting the type specimens. The name cazae refers to Carole’s childhood nickname (Caz). EXTERNAL MORPHOLOGY.— The living animals are 40-70 mm in length. The body is oval and high, the notal surface is smooth (Fig. 1A), but with low pigmented elevations. The anus is situat- ed within the ring formed by the gill branches. The perfoliate rhinophores have 18-26 lamellae. The branchial plume is made up of nine unipinnate branches. The background color of the body is translucent white. There are several irregular patches of deep red-maroon found scattered on the mid-dorsal region. Each of these patches has a number of smaller irregular orange spots within it. The number of orange spots varies greatly from zero to sixteen or more. Some specimens, as indi- cated in photographs of specimens not collected, have orange spots outside of the maroon patches, within the white regions. Each colored patch and spot is slightly elevated or raised from the other- wise smooth notal surface. The margin of the mantle has an irregular series of similar patches. Some specimens have a thin blue line along the edge of the mantle. In some specimens the rhinophoral sheaths have a small maroon patch on the posterior side. The gills and rhinophores are uniformly white, with opaque white specks and lines. Ventrally (Fig. 2B), the foot is about a quar- ter of the width of the notum. The genital aperture is located on the right side in the anterior third of the body. The triangular, conical oral tentacles are well separated from each other. MANTLE GLANDS.— The mantle glands (Fig. 2A) are continuous around the lateral margins of the mantle and are absent from anterior and posterior portions of the notum. The glands consist of simple spherical bodies that are well separated from each other. BUCCAL ARMATURE.— The buccal mass is composed of an oval to rounded buccal bulb and GOSLINER AND BEHRENS: NEW NUDIBRANCHS FROM THE INDIAN OCEAN 3 k ie a FiGuRE 1. Living animals. A, Chromodoris cazae sp. nov., specimen from Khawr Fakken, United Arab Emirates, photograph by Carole Harris. B. Aldisa andersoni sp. noy., specimen from Sri Lanka, holotype (CASIZ 167461), photograph by Charles Anderson. the short, glandular oral tube (Fig. 2C). The muscular portion is about two-thirds the length of the oral tube. A pair of elongate salivary glands extends posteriorly from the junction of the posterior portion of the buccal bulb and the esophagus. At the anterior end of the muscular portion of the buccal mass there is a chitinous labial cuticle, which bears numerous jaw rodlets. The rodlets (Fig. 3A) have a short base and a curved apex that may be undivided or, more commonly, a short bifid 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 1 apex. The radular formula of the paratype is 33 x 42.1.42. The rachidian row of teeth is well- developed (Fig. 3B). The rachid- ian teeth are short, triangular and devoid of denticles. The inner- most lateral teeth (Fig. 3B) have a broad triangular cusp. On the inner side of the primary cusp is a single rounded inner denticle. There are 4-5 small, acutely pointed denticles on the outer side of the primary cusp. The subsequent inner lateral teeth lack the inner denticle and pos- sess 5—6 outer denticles. The middle lateral teeth (Fig. 3C) are elongate and sharply curved with 5-6 acutely pointed denticles on the outer side of the primary cusp. The three to four outermost teeth (Fig. 3D) are more elongate with a broad base. They have 3-4 rounded denticles on the outer side of the small primary FIGURE 2. Chromodoris cazae (CASIZ 168300). A. Schematic distribution of mantle glands. B. Ventral view of anterior border of the foot, scale bar = 1.0 mm. C. Buccal mass, scale bar = 1.5 mm. D. Reproductive system; scale bar = Cusp. 0.3mm. Abbreviations: am, ampulla; bc, bursa copulatrix; bm, buccal mass; e, REPRODUCTIVE SYSTEM.— _ esophagus; ej, ejaculatory duct; fgm, female gland mass; 0, oral tube; ot, oral tentacle; p, penis; pr, prostate; sg, salivary gland; rs, receptaculum seminis; vg, The reproductive system is tri- vestibular gland. aulic (Fig. 2D). The ampulla is relatively short and curved. From its proximal end it bifurcates into the short, thick, slightly convoluted prostate and the short oviduct that enters the female gland mass. From the prostatic portion of the vas deferens, the duct narrows somewhat and changes texture as it becomes the long and convoluted ejaculatory segment. More proximally to the genital atrium, the ejaculatory duct narrows and again abruptly expands into the short muscular penial sac. The thick vaginal duct opens into the genital atrium adjacent to the penis and a short, globular vestibular gland. Proximally, the vaginal duct connects directly with the spherical bursa copulatrix. The curved, pyriform receptaculum seminis enters the vagina immedi- ately below the bursa copulatrix. Just below the entrance of the receptaculum seminis, the vagina is joined by the short, narrow uterine duct, which enters the female gland mass. The female gland mass is large and completely developed. DisCUSSION.— Chromodoris cazae is unique in its white color with patches of maroon with smaller orange spots inside the patches. A few other species have similar colors. Chromodoris kitae Gosliner, 1994, has purple spots extending from the edge of the notum and covering the entire dor- sal surface. There is a ring of orange patches along the submarginal area of the mantle. The gills and rhinophores are grayish white with opaque white spots. This species is known only from Madagascar. Chromodoris naiki Valdés, Mollo, and Ortea, 1999, from India, also has a white body but with purple and orange markings. In this case, the orange markings are not restricted to a sub- GOSLINER AND BEHRENS: NEW NUDIBRANCHS FROM THE INDIAN OCEAN 5 Jaw rodlets, scale bar = 2 um. B. Inner lateral teeth, scale bar = 10 um. C. lateral teeth from the central por- tion of half-row, scale bar = 10 um. D. Outer lateral teeth, scale bar = 10 um. marginal band and are found on the notum as well. This differs from the color pattern of C. cazae, where the orange spots are much smaller than the purplish ones. Additionally, C. naiki bears small opaque white spots on the notum that are absent in both C. kitae and C. cazae. Internally, the three species differ as well. In C. naiki, a rachidian row of radular teeth is absent, whereas in C. cazae and C. kitae a row of triangular rachidian teeth is present. In C. cazae, the rachidian teeth are far more prominent than in C. kitae. In the reproductive system of C. naiki and C. kitae, the elongate and coiled receptaculum seminis enters directly into the base of the bursa copulatrix at the distal end of the vagina. In C. cazae, the receptaculum is pyriform with a shorter duct and enters the vagina in a position that is more proximal to the genital atrium. Aldisidae Bergh, 1891 Aldisa Bergh, 1878 Aldisa andersoni Gosliner and Behrens, sp. nov. (Figs. 1B, 4-5) Chromodoris sp. Debelius, 1996:213 (bottom of page). Aldisa erwinkoehleri Perrone, 2001: misidentification of specimen in Debelius, 1996. MATERIAL EXAMINED.— HoLotyPeE (CASIZ 167461). Pigeon Island, Trincomal, Sri Lanka, March 7, 1995, 8 m deep, Charles Anderson. Photographs of two additional specimens taken on the same dive the holotype was collected, taken by Dr. Charles Anderson. 6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 1 DISTRIBUTION.— Known only from Pigeon Island, Trinco- mal, Sri Lanka. ETYMOLOGY.— Aldisa an- dersoni is named in recognition of Charles Anderson, Malé, Maldives, who was the first to bring this species to our attention and was instrumental in collect- ing the holotype. EXTERNAL MORPHOLOGY.— The living animals are 20-30 mm in length. The body is oval and the notal surface covered with a series of low conical tubercles (Fig. 1B). The tubercles form distinct ridges running lon- gitudinally along the center of the notum, and perpendicular lat- erally along the margin of the notum. Rows of these tubercles are situated in regions of opaque blue pigment. These regions are separated by areas of black. There is an opaque, bright yellow saddle across the notum behind the rhinophores. This saddle Ficure 4. Aldisa andersoni sp. nov., holotype (CASIZ 167461) A. Ventral extends right to the edge of the _ view of the anterior border of the foot, scale bar = 1.0 mm. B. Buccal mass, mantle. A few bright yellow dorsal view, scale bar = 0.6mm. C. Buccal mass, ventral view, scale bar = 0.5 mm. D. Reproductive system; scale bar = 2.3 mm. E. Penial hooks, scale bar = 35 um. Abbreviations: al, albumen gland; am, ampulla; be, bursa copulatrix; gill pocket. Some specimens eng, central nervous system; e, esophagus: ej, ejaculatory duct; fgm, female have yellow patches posterior to — gland mass; og, oral gland; p, penis; pr, prostate; ra, radular sac; rm, retractor the rhinophores. The anus is muscle; rs, receptaculum seminis; sg, salivary glands; v, vagina. patches may partially encircle the positioned dorso-ventrally. The perfoliate rhinophores have 17 lamellae. The gill is made up of seven tripinnate branches. The rhinophores and gill are the same blue as the notum. Each rhinophoral pocket has two equally-sized tubercles on either side. The gill sheath has a single row of twelve tubercles along its margin. The anterior portion of the foot (Fig. 4A) is smooth lacking a notch and the oral tentacles are rudimen- tary, typical of Aldisa. BUCCAL ARMATURE.— The buccal mass is composed of an oval to rounded buccal bulb and the glandular oral tube (Fig. 4B—C). The esophagus opens on the dorsal side of the buccal mass, where two large, darkly pigmented salivary glands attach. Below the posterior end of the oral tube portion of the buccal mass lies a large granular oral gland mass (Fig 4C). The radular formula is impossible to determine given the elongate intertwining radular teeth (Fig. 5A). There is no trace of a rachidian row of teeth. The teeth are all extremely narrow and elongate with a broad, triangu- lar base. The inner lateral teeth (Fig. 5C) are much longer than the outer ones. The inner teeth bear 7-8 sharp denticles with a wide groove extending from the apex to about the middle of each den- GOSLINER AND BEHRENS: NEW NUDIBRANCHS FROM THE INDIAN OCEAN 7 FiGureE 5. Aldisa andersoni sp. nov., holotype (CASIZ 167461). Scanning electron micrographs. A. Lateral teeth, scale bar = 10 um. B. Close up of inner lateral teeth from the central portion of half-row, scale bar = 1 pm. C. Inner lateral teeth, scale bar = 2 um. D. Close up of inner lateral teeth, scale bar = 2 um. ticle (Fig. 5B, D). A series of approximately ten smaller, simply upturned denticles are present along the distal portion of the tooth below the grooved denticles. REPRODUCTIVE SYSTEM.— The reproductive system is triaulic (Fig. 4D). The ampulla is tubu- lar and thick, winding back on itself before narrowing where the distal portion emerges at the prostate and oviduct. The oviduct connects with the female gland mass beneath the prostate. The prostate is granular and tubular, convoluted, distally narrowing and expanding again into the short muscular ejaculatory portion. The distal portion of the deferent duct exits at a separate genital atri- um. The penial bulb is armed with approximately 6 rows of minute hooks (Fig 4E). The hooks have a wide base tapering to a long curved cusp. The vaginal duct is about the same length as the defer- ent duct, and opens into the bursa copulatrix. The bursa copulatrix is rounded and connects with the seminal receptacle by a short duct. Near the seminal receptacle this duct is joined by the uter- ine duct, which enters the female gland mass. The pyriform receptaculum seminis is equal in size to the bursa copulatrix. Discussion.— The revision of the genus Aldisa by Millen and Gosliner (1985) diagnosed the genus with the following characteristics: elongate radular teeth with multiple denticulations, a dor- sum with low conical tubercles (except Aldisa pikokai Bertsch and Johnson, 1982, which lacks tubercles), a ring of tubercles around the rhinophoral and branchial pockets, bi- and tri-pinnate gills, penial spines in most species, absence of oral tentacles and the anterior portion of the foot smooth with no notch. Aldisa andersoni exhibits all the characteristics of the genus Aldisa noted above. Elwood, et al. (2000) added a previously unnoticed character when describing Aldisa albatrossae and A. williamsi: the presence of a pair of unique hamate radular teeth, which are much wider and more robust than the thin denticulate ones. 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 1 Aldisa andersoni is the fourth described tropical Indo-Pacific species exhibiting bright blue coloration with black and/or yellow markings. It is similar, in this regard, to Aldisa albatrossae, A. williamsi and A. erwinkoehleri Perrone, 2001, but it differs in a number of characteristics. All other species, except Aldisa barlettai Ortea and Ballesteros, 1989, are species exhibiting subdued white, green, tan, or red coloration making them cryptic on their sponge prey. In Aldisa andersoni, the low conical tubercles are clearly arranged in rows or ridges. Tubercles on the notum of Aldisa albatrossae, A. williamsi and A. erwinkoehleri are randomly distributed. The black coloration on A. williamsi and A. erwinkoehleri forms concentric rings, and in A. alba- trossae it forms a rectangle on the dorsum with a T-shape mark anterior to the rhinophores. In A. andersoni, the black pigmentation is found uniformly over the mantle separating the rows of tuber- cles. The opaque yellow pigmentation on Aldisa andersoni is quite similar to that found on A. alba- trossae and A. erwinkoehleri. There are, however, consistent differences that clearly separate them. A. williamsi has no yellow pigmentation. The yellow pigmentation covers the tubercles in A. erwinkoehleri and in A. andersoni, but not in A. albatrossae, where it occurs between the tubercles. The yellow color markings on A. andersoni are far more extensive than in the other species, with a yellow saddle extending to the very edge of the mantle that is not isolated into smaller broken patches, as in A. erwinkoehleri. The rhinophores differ in color among all four species. In Aldisa albatrossae, they are white- grey; in A. williamsi, pale brown; in A. erwinkoehleri, black; whereas in A. andersoni they are the same blue as the body. The arrangement of tubercles around the rhinophoral pockets also differs among the four species. Aldisa albatrossae has four similarly sized equally separated tubercles; A. erwinkoehleri has two large and two small tubercles, whereas A. andersoni and A. williamsi have two equally sized tubercles, which are separated on either side of the pocket. Differences exist also in the tubercles around the gill sheath. A. albatrossae has a double row of tubercles; A. williamsi has a single row of eight tubercles; A. erwinkoehleri a single series tubercles; and A. andersoni has a single series of twelve tubercles around the sheath. Internally, the radular teeth vary in one significant regard. In Aldisa albatrossae and A. williamsi, there is a pair of smooth, hamate teeth in addition to the elongate teeth typical of species of Aldisa. These hamate teeth are absent in A. andersoni and were not described either in A. erwinkoehleri. A massive granular gland on the ventral surface of the buccal mass 1s present in A. andersoni. Re-examination of type specimens of A. williamsi and A. albatrossae revealed the pres- ence of a similar gland, but it is proportionately much smaller in these two species than that found in A. andersoni. A few differences were found in the reproductive systems of the four species. The penial spines of Aldisa andersoni are similar in shape to those found in A. williamsi, but they differ from those of A. albatrossae. Perrone (2001) reports no armature at all in the penial bulb of A. erwinkoehleri. In the description of A. erwinkoehleri, the prostatic portion of the vas deferens was not differentiated, as it is in the other three species. It is not certain whether these discrepancies are due to morphological differences, but they are more likely the result of incomplete description in the case of A. erwinkoehleri. The bursa copulatrix is relatively small in all four species, being approximately equal to the receptaculum seminis in size. ACKNOWLEDGMENTS We thank Carole Harris, Leon Betts, and Charles Anderson for collection of type material and their continued keen enthusiasm for discovering and documenting new species from the Indo- Pacific tropics. GOSLINER AND BEHRENS: NEW NUDIBRANCHS FROM THE INDIAN OCEAN 9 This paper has been supported by the National Science Foundation, through the PEET grant DEB-9978155 “Phylogenetic systematics of dorid nudibranchs.” LITERATURE CITED Basa, K. 1994. Descriptions of four new, rare, or unrecorded species of Hypselodoris (Nudibranchia: Chromodorididae) from Japan. Venus 53(3):175—187. Basa, K. 1995. Anatomical and taxonomical review of four blue patterned species of Hypselodoris (Nudibranchia: Chromodorididae) from Japan. Venus 54 (1):1-15. BRUNCHHORST, D.J. 1993. The systematics and phylogeny of phyllidiid nudibranchs (Doridoidea). Records of the Australian Museum, Supplement 16:1—107. CARLSON, C.H., AND P.J. Horr. 2000. Three new Pacific species of Halgerda (Opisthobranchia: Nudibranchia: Doridoidea). The Veliger 43(2):154-163. COLEMAN, NEVILLE. 2001. 1001 Nudibranchs — Catalogue of Indo-Pacific Sea Slugs. Neville Coleman’s Underwater Geographic Pty Ltd., Queensland. 144 pp. DEBELIUS, HELMuT. 1996. Nudibranchs and Sea Snails — Indo-Pacific Field Guide. IKAN — Unterwasser- archive, Frankfurt. 321 pp. DorGAN, K.M., A. VALDES, AND T.M. GOSLINER. 2002. Phylogenetic systematics of the genus Platydoris (Mollusca, Nudibranchia, Doridoidea) with descriptions of six new species. Zoologica Scripta 31: 271-319. ELwoop, H.R., A. VALDES, AND T.M. GOSLINER. 2000. Two new species of Aldisa Bergh, 1878 (Mollusca, Nudibranchia) from the tropical Indo-Pacific. Proceedings of the California Academy of Sciences 52(14): 171-181. FAHEY, S., AND T.M. GOSLINER. 1999a. description of three new species of Halgerda from the western Indian Ocean, with a redescription of Halgerda formosa, Bergh, 1880. Proceedings of the California Academy of Sciences 51(8):365-383. FAHEY, S., AND T.M. GosLiner. 1999b. Preliminary phylogeny of Halgerda (Nudibranchia: Halgerdidae) from the Tropical Indo-Pacific with descriptions of three new species. Proceedings of the California Academy of Sciences 51(11):425-448. FAHEY, S., AND T.M. GOsLINER. 2001. On the genus Halgerda (Nudibranchia: Halgerdidae) from Western Australia with descriptions of four new species. Bollettino Malacologico 37(5—8):55—76. FAHEY, S., AND T.M. GOSLINER. 2003. Mistaken identities: On the Discodorididae genera Hoplodoris and Carminodoris Bergh (Opisthobranchia, Nudibranchia). Proceedings of the California Academy of Sciences 54(10):169-208. GOsLINER, T.M. 1994. New species of Chromodoris and Noumea (Nudibranchia: Chromodorididae) from the western Indian Ocean and southern Africa. Proceedings of the California Academy of Sciences 48(12):239-252. GOSLINER, T.M., AND D.W. BEHRENS. 1997. Description of four new species of phanerobranch dorids from the Indo-Pacific, with a redescription of Gymnodoris aurita (Gould, 1852). Proceedings of the California Academy of Sciences 49(9):287-308. GOSLINER, T.M., AND D.W. BEHRENS. 1998a. Five new species of Chromodoris (Mollusca: Nudibranchia: Chromodorididae) from the tropical Indo-West Pacific. Proceedings of the California Academy of Sciences 50:139-165. GOSLINER, T.M., AND D.W. BEHRENS. 1998b. Two new discodorid nudibranchs from the western Pacific with a redescription of Doris luteola Kelaart, 1858. Proceedings of the California Academy of Sciences 50(11): 279-293. GOSLINER, T.M., AND D.W. BEHRENS. 2000. Two new species of Chromodorididae (Mollusca: Nudibranchia) from the tropical Indo-Pacific, with a redescription of Hypselodoris dollfusi (Pruvot-Fol, 1993). Proceed- ings of the California Academy of Sciences 52(10):111—124. GOSLINER, T.M., AND S.J. FAHEY. 1998. Description of a new species of Halgerda, with a redescription of Halgerda elegans Bergh, 1905. Proceedings of the California Academy of Sciences 50(15):346-359. 10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 1 GOSLINER, T.M., AND R.F. 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. GOSLINER, T.M., AND V.G. SMITH. 2003. Systematic review and phylogenetic analysis of the nudibranch genus Melibe (Opisthobranchia: Dendronotacea) with descriptions of three species. Proceedings of the California Academy of Sciences 54(18):302-355. Hamatanl, I. 1995. Two species of Chromodorididae (Nudibranchia), one newly recorded and one newly established, from middle Japan. Venus 54(2):101—107. HAMATANI, I. 2001. Two new species of Goniodorididae (Opisthobranchia; Nudibranchia) with a new genus from Kuroshima Island, Okinawa, Japan. Venus 60(3):151—156. JOHNSON, R.F., AND T.M. GOSLINER. 1998. The genus Pectenodoris (Nudibranchia: Chromodorididae) from the Indo-Pacific, with the description of a new species. Proceedings of the California Academy of Sciences 50(12):295—306. MILLEN, S., AND T.M. GOSLINER. 1985. Four new species of dorid nudibranchs belonging to the genus Aldisa (Mollusca: Opisthobranchia), with a revision of the genus. Zoological Journal of the Linnean Society 84:195-233. OrTEA, J., AND M. BALLESTEROS. 1989. Descripcidn de una espectacular especie del genero Aldisa Bergh, 1878 (Mollusca, Opisthobranchia) dedicada a la memoria Dr. Giorgio Barletta. Bollettino Malacologico 24(9-12):155—160. Ortiz, D.M., AND T.M. GOSLINER. 2003. A new species of Phyllodesmium Ehrenberg, 1931 (Mollusca, Nudibranchia) from the tropical Indo-Pacific. Proceedings of the California Academy of Sciences 54(9):161—-168. PERRONE, A.S. 2001. A new species of Nudibranchia of the genus Aldisa Bergh (Gastropoda, Opisthobranchia) from Thailand. Basteria 65:105—116. RUDMAN, W.R. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: a review of the genera. Zoological Journal of the Linnean Society 81(2—3):115—273. RUDMAN, W.R. 1986. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Noumea purpurea and Chromodoris decora colour groups. Zoological Journal of the Linnean Society. 86:309-353. RUDMAN, W.R. 1987. The Chromodorididae (Opisthobranchia, Mollusca) of the Indo-West Pacific: Chromodoris epicuria, C. aureopurpurea, C. annulata, C. coi and Risbecia tryoni colour groups. Zoologi- cal Journal of the Linnean Society 90(4):305—407. RUDMAN, W.R. 1995. The Chromodorididae (Opisthobranchia: Mollusca of the Indo-West Pacific: further species from New Caledonia and the Noumea romeri group. Molluscan Research 16:1-43. SCHRODL, M. 1999. Glossodoris charlottae, a new chromodorid nudibranch from the Red Sea (Gastropoda, Opisthobranchia). Vita Marina 46(3-4):89-94. SMITH, V.G., AND T.M. GOSLINER. 2003. A new species of Tritonia from Okinawa (Mollusca: Nudibranchia), and its association with a gorgonian octocoral. Proceedings of the California Academy of Sciences 54(16):255-278. VALDES, A., E. MOLLO, AND J. ORTEA. 1999. Two new species of Chromodoris (Mollusca, Nudibranchia, Chromodorididae) from Southern India, with a redescription of Chromodoris trimarginata (Winckworth, 1946). Proceedings of the California Academy of Sciences 51:461-472. Yonow, N. 1984. Doridacean nudibranchs from Sri Lanka, with descriptions of four new species. The Veliger 26(3):214-228. Yonow, N. 1994. Opisthobranchs from the Maldives Islands including descriptions of seven new species (Mollusca: Gastropoda). Revue Francaise d’Aquariologie 20(4):97-129. Yonow, N. 2001. Part 11. Doridacea of the families Chromodorididae and Hexabranchidae (Mollusca, Gastropoda, Opisthobranchia, Nudibranchia), including additional moluccan material. Jn Results of the Rumphius Biohistorical Expedition to Ambon (1990). Zoologische Meddelingen. Leiden 75:1—S0. Yonow, N., R.C. ANDERSON, AND S.G Buttress. 2002. Opisthobranch molluscs from the Chagos Archipelago, Central Indian Ocean. Journal of Natural History 36(7):831-882. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2, pp. 11-54, 4 pls., 15 figs. April 22, 2004 Pacific Coast Nudibranchs, Supplement IT New Species to the Pacific Coast and New Information on the Oldies David W. Behrens Department of Invertebrate Zoology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, U.S.A. Email: dave @ seachallengers.com. Pacific Coast Nudibranchs — A Guide to the Opisthobranchs Alaska to Baja California (Behrens 1991) presented taxonomic and biological information on 217 species of opisthobranch molluscs found within the geographic range of the title. Only a few species were excluded at that time, because of depth or rarity. Since its printing many new species have been discovered and the ranges of others found to occur with- in the coverage of the original work. Researchers have learned a great deal about morphological variation and the ecology of many species. The goal of this supple- ment is to bring current the biological, biogeographical, morphological and taxo- nomic information concerning those species found along this coastline by incorpo- rating both published material and confirmed observations of field workers and opishtobranch enthusiasts. The opisthobrach fauna of the Pacific Coast, from the Bering Strait of Alaska to Cabo San Lucas, Baja California Sur, Mexico, now totals some 252 documented species. This supplement is intended to update the contents of Pacific Coast Nudibranchs, second edi- tion, (Behrens 1991) by providing information on newly discovered species, unknown at the time of publication of that edition, and new occurrences, range extensions and ecological information, as well as, changes to and additions of species names published since 1991. This new information is compiled not only from the publication of recent scientific journal articles, but also from observations and collections shared with the author by opisthobranch researchers and enthusiasts up and down the Pacific Coast. Without this collaborative effort, accu- mulation of scientific knowledge in this area would certainly not be as extensive as it is today. Included here are 26 species new to the region covered by the 1991 edition. This supplement also includes species, previously undescribed, which now have names, several newly discovered for- eign introductions, and 23 additional undescribed species, bringing the total number of opistho- branch species from the Bering Strait, Alaska to the tip of Baja California, Mexico, to 252 species. Interest in opisthobranch mollusks has a broadening audience as indicated not only by the number of new publications featuring this taxonomic group, but by the media attention being given it. Nudibranch or sea slug is becoming a household word. The recent report of the introduction of a Philine from New Zealand waters to San Francisco Bay and its rapid population spread to other coastal bays and estuaries in California received tele- vision and newspaper coverage across the country. With the growth of the Internet has come its concomitant use by opisthobranch enthusiasts to share information. It all began with Bernard Picton and C. Morrow’s entire book entitled A Field II 1D PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Guide to the Nudibranchs of the British Isles presented on a Web Site. Since then, Nudibranch web sites have sprung up all over the World. On the West Coast of the United States, we have two web sites. “The Slug Site” features a collection of color photos from various regions around the world, whereas “The Opisthobranch” web site contains online editions of the current month’s Opisthobranch Newsletter, a bibliography of Opisthobranchia, a nudibranch bibliography com- piled by Gary McDonald with gopher search capabilities, a list of color illustrations of opistho- branchs on the net, a taxonomic and systematic list of Opisthobranchia, and the addresses of per- sons following this group of marine animals. Internationally, the Australian Museum’s — “Sea Slug Forum” has become the largest and most active site providing realtime communication among sea slug enthusiasts and scientists alike. Lastly, Erwin Koehler’s Opisthobranchs of the World is a list of every photograph of an opisthobranch on the world wide web. New information was accepted for this supplement up until September 30, 2003. ACKNOWLEDGEMENTS I would like to acknowledge the generous sharing of observations, photographs and collections which make the publication of this supplement possible. My sincere thanks to each of the follow- ing: Peter Ajtai Jerry Allen Jim Anderson Kelvin Barwick Clinton Bauder Bob Bayer Michael Behrens Hans Bertsch Robert Bolland Peter Brueggeman Don Cadien Yolanda Camacho-Garcia Orso Angulo-Campillo Juan Lucas Cervera Marc Chamberlain Roger Clark Kathy deWet Sarah Douglass Jack Engle E. Daniel Erikson Nora Foster Steve Gardner Donna and Charlie Gibbs Jeff Goddard Dale Marie Gonsalez Terry Gosliner Dan Gotshal] Constance Gramlick Alan Grant Herb Gruenhagen Pete Haaker Bernard Hanby Gordon Hendler Richard Hermann Ali Hermosillo-Gonzalez Steve Horvath Mary Ellen Hill Paul Hughes Derek Hutchinson Jun Imanoto Jerry Jacobs Anthony Keefe Darcy Kehler Kirsty Kaiser Erwin Koehler Alan Kuzirian Andy Lamb Jim Lance Lovell and Libby Langstroth Doug Mason Rosa Larralde Ridaura Megan Lilly Steve Long Carlos Ramos Mantecon Nishina Masayoshi Neal McDaniel Gary McDonald Duke McPherson Bill Merilees Jane Middleton Sandra Millen Kathy Ann Miller Mike Miller Phanor Montoya Kurtis Morin Alan Murray Ryan Murphy Kathy Noonan James Nybakken Bill Pence Bernard Picton Rita O’ Clair Dan Richards Jeff Rosenfeld Bill Rudman Maria Schaefer Caroline Schooley Marc Shargel Paul Sin Bob Sinclair Hugh Erle Smith George Spalding Cynthia Trowbridge Marli Wakeling Bruce Wight Angel Valdés Ron Veralde Special thanks to the referees who gave their valuable time to review the information present- ed here and offer their candid comments. I would also like to thank my wife, Diana for her patience with my obsession for opisthobranchs. BEHRENS: PACIFIC COAST NUDIBRANCHS 13 SPECIES NEW TO THE PACIFIC COAST The following species were not known by the author to occur between Alaska and the tip of the Baja California peninsula during the preparation of the second edition of Pacific Coast Nudibranchs in 1991. Some are new discoveries, some were published in obscure publications, and some where purposely excluded from the 1991 edition due to their depth of occurrence below that of the average SCUBA diver. The new records are presented here in phylogenetic order. Order Cephalaspidea Family Acteonidae Acteon sp. Gosliner (1996) reports an undescribed species of Acteon from the Santa Maria Basin, California. IDENTIFICATION.— The shell is white and 3 mm long with 22 finely, punc- tate spiral lines on the body whorl (Gosliner 1996) (Fig. 1). It has three whorls. This species has a large operculum. The species is reported to be similar to Acteon panamensis Dall, 1908, but different from A. traskii, which has rose colored bands on the shell (Gosliner 1996). RaDULA.— Each half row has more than 50 minute teeth, each with 16-18 denticles along the inner edge. a ec, ay ee S1ZE.— 1-3 mm in length. Son DHA GF oll RANGE.— Santa Maria Basin, off Purisima Point, California (Gosliner (After Gosliner 1996.) 1996). Family Hydatinidae Parvamplustrum sp. Gosliner (1996) reports an undescribed species of Parvamplustrum from the Santa Maria Basin, California. IDENTIFICATION.— A tiny species, the animal has a short, triangular foot (Gosliner 1996). It has bifid head tentacles. The shell is bulloid. RADULA.— Gizzard plates are absent and the jaws are unarmed polygonal rodlets (Gosliner 1996). The radular formula is 20 x 1.0.1. The lateral teeth are curved with broad denticles. S1zE.— Less than | mm in length. RANGE.— Santa Maria Basin, off Purisima Point, California (Gosliner 1996). Family Cylichnidae Cylichna diegensis (Dall, 1919) Gosliner (1996) reports Cylichna diegensis from the Santa Maria Basin, California. IDENTIFICATION.— The shell is bulloid (Fig. 2A) with fine spiral striae and brown periostrical lines at each end (Gosliner 1996). The body of the animal is brown in the preserved state. RADULA.— The jaws have multifid rodlets (Fig. 2B). The radular formula is 13 x 3-4.1.1.1.3-4 (Gosliner 1996). The rachidian teeth are bifid with 5—7 denticles (Fig. 2C). The inner lateral teeth have 9-11 denticles (Fig. 2D). The three gizzard plates are equal in size and shape (Gosliner 1996). 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 SIZE.— The shell is 3—7mm in length. RANGE.— Pt. Loma, San Diego, California (9-191 fathoms) to the Santa Maria Basin, off Point Sal, California (Gosliner 1996). A C D FIGURE 2. Cylichna diegensis (Dall, 1919). A. Drawing of shell. B. Jaw rodlets, C. Rachidian tooth, D. Lateral Tooth. (After Gosliner 1996.) Acteocina eximia (Baird, 1863) Gosliner (1996) discusses nomenclatural issues surrounding the subspecies originally assigned to Acteocina culcitella. He concludes that the sub- species, Acteocina culcitella eximia, is in fact a dis- tinct species in the California fauna. IDENTIFICATION.— Externally similar to Acteo- cina culcitella (Species #4 in Behrens 1991), but with a shorter spire and no columellar pleat (Gosliner 1996) (Fig. 3A). RADULA.— The radular formula is 19-28 x 1.0.1 (Gosliner 1996). The lateral teeth have a strong central A denticle, with 30—50 short denticles along the margin FIGURE 3. Acteocina eximia (Baird, 1863). A. (Fig. 3B). There are three gizzard plates, two large Drawing of shell. B. Lateral Tooth. (After Gosliner, and concave and one small one that is laterally com- !99®-) pressed and concave (Gosliner 1996). SIZE.— 4-10mm in length. NATURAL History.— Found in deep water, 22 m in its northern range to 60 m in the southern areas. RANGE.— Vancouver, British Columbia, to San Diego, California (Gosliner 1996). Family Retusidae Retusa zystrum Dall, 1919 Gosliner (1996) reports Retusa zystrum from the Santa Maria Basin, California. IDENTIFICATION.— The shell is bulloid with numerous characteristic longitudinal striae (Gosliner 1996). Buccal Morphology.— Jaws are absent. The gizzard plates are not equal in size, one being larger than the other two (Gosliner 1996). The plates are covered with tubercles, the largest being on the inner side. S1zE.— 1.5 mm in length. RANGE.— Described from San Diego and San Pedro, California. Now known from the Santa Maria Basin off Point San Luis and Purisima Point, California (Gosliner 1996). BEHRENS: PACIFIC COAST NUDIBRANCHS 15 Volvulella californica Dall, 1919 Volvulella californica Dall, 1919, is reported by Gosliner (1996) from the Santa Maria Basin, Califonia, just north of its type locality. IDENTIFICATION.— Shell smooth, without sculpture (Figure 4A). Widest in middle, tapered at both ends. Posterior project acute, elongate (Gosliner 1996). RADULA.— None NATURAL History.— Little is known. Specimens were collected at between 90-345 m depth. S1zE.— 1-3 mm in length. RANGE.— Santa Cruz, California, to Todos Santos Bay, Baja California (Abbott 1974). EtyMoLoGy.— Named for the State of California, its type locality. Volvulella panamica Dall, 1919 Volvulella panamica, on the other hand, was known only from Panama, and it too is noted because of its recent collection in the Santa Maria Basin (Gosliner, 1996). IDENTIFICATION.— Shell smooth, without sculpture, and with a prolonged posterior projection (Fig. 4B). Posterior end of outer lip originating at base of posterior projection (Gosliner 1996). RADULA.— None NATURAL History.— Little is known about the biology of | A B this species. Specimens were collected from 90-197 m depth. FIGuRE 4. Drawings of shells of S1zZE.— 2.5—5 mm in length. (left) Volvulella californica and (right) RANGE.— Santa Maria Basin, California (Gosliner 1996), Volvulella panamica. (After Gosliner to Panama (Abbott 1974). 1996.) ETYMOLOGyY.— Named for the type locality, Panama Bay, Panama. Family Philinidae A white species of Philine was first noted in San Francisco Bay when it was observed in trawl samples collected in conjunction with the on-board educational program offered by the Marine Science Institute of Redwood City, in the summer of 1992. It was identified by Gosliner (1995) as Philine auriformis Suter, 1909. Rudman (www.seaslugforum /philcali.htm) refutes this identifica- tion, stating the California species is quite different in internal anatomy from the New Zealand Philine auriformis. Since that time, similar white Philine have become abundant along the entire California coastline. As a result of careful study, a total of four species of white Philine have now been identified throughout the region (see discussions on the Sea Slug Forum — www.seaslugfo- rum/philcali.htm). Three more deepwater species are reported from the Santa Maria Basin, California, by Gosliner (1996). Philine aperta (Linnaeus, 1767) IDENTIFICATION.— The animal is white like P. auriformis, P. orientalis and P. japonicus. BuccAL MorPHOLOGY.— Gizzard plates same as P. orientalis and P. japonicus, two larger mirror imaged and a third smaller plate. All have small holes in the center. S1zE.— To 70 in length. RANGE.— Atlantic coast of South Africa. California (T. Gosliner, pers. commun.). 16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Philine auriformis Suter, 1909 IDENTIFICATION.— The animal (Plate 1A) is transparent white to yellow in color. The Hancock’s organs are reddish brown, and the darker visceral mass and gizzard plates are clearly visible through both the dorsal and ventral surfaces of the body (Gosliner 1995). This species also has a broad, shallow internal shell, with a spiral beaded sculpture. BuCcCAL MORPHOLOGY.— There are equally sized spindle shaped gizzard plates with grooves on the outer sur- face (Gosliner 1995) (Fig. 5B). The radular formula is 21 x 1.1.0.1.1. There are 30-50 fine denticles along the inner masticatory edge on the inner lateral tooth (Fig. 5A). The outer tooth is smaller but similar in morphology. NATURAL History.— This species is found on muddy — A B bottoms, intertidally to 15—20 ft. deep. Egg masses are large, ovoid, mucous masses attached to the substrate by a thread. They contain numerous spirals of egg capsules, each capsul containing two eggs (see Gosliner (1995) for details). This voracious carnivore feeds on the clam Gemma gemma and several species of Transennella. S1zE.— 15—30mm in length. RANGE.— South San Francisco Bay and Bodega Harbor, California (Gosliner 1995). Also reported from Dillon Beach (H.E. Smith, pers. commun.), Morro Bay (M. Behrens, pers. commun.) Elkhorn Slough (G. McDonald, pers. commun.), Mission Bay (D. Cadien, pers. commun.), and Coos Bay, Oregon (Jeff Goddard, pers. commun.), Barkley Sound, Vancouver Island, British Columbia (S. Millen, pers. commun.). FiGuRE 5. Philine auriformis Suter, 1909. A. Lateral radular tooth; B. Gizzard plate. Philine japonica Lischke, 1872 IDENTIFICATION.— The animal is white like P. aperta, P. auriformis, and P. orientalis. BuccAL MORPHOLOGY.— Gizzard plates same as P. aperta and P. orientalis, two larger mir- ror imaged and a third smaller plate. All have small holes in the center. SIZE.— To 50 mm in length. RANGE.— Japan and Tomales Bay, California (Gosliner, pers. commun.). Philine orientalis A. Adams, 1854 IDENTIFICATION.— The animal is white similar to P. aperta, P. auriformis, and P. japonicus. RADULA.— Gizzard plates are relatively large. Two are mir- ror images of one another. The third is smaller and symetrical in shape. Each plate has a pair of pin holes in the center (Fig. 6A). The radular formula is 1.0.1. The teeth are hook-like with a denticulated A. B. edge, like that of P. auriformis FiGcureE 6. Philine orientalis A. Adams, 1854. A. Gizzard plate. B. Radu- (Fig. 6B). lar dentition. NATURAL History.— Found on intertidal mudflats. Periodically in dense populations, dozens per square meter ( M. Chow, pers. commun.). BEHRENS: PACIFIC COAST NUDIBRANCHS 7 S1zE.— To 50 mm in length. RANGE.— Originally described from “eastern seas,” it has been reported from the Philippines by Watson (1886) and from Hong Kong by Morton and Chiu (1990). Bodega Bay and San Francisco Bay, California (M. Chow and T. Gosliner, pers. commun.). Philine polystrigma (Dall, 1908) Orignally named Clistaxis polystrigma Dall, 1908, Gosliner (1996) reports this species from the Santa Maria Basin, California and reassigns the species to the genus Philine. IDENTIFICATION.— Typical Philine body shape, the preserved specimens were light tan (Gosliner 1996). The anterior head shield is rounded at both ends. The posterior shield has a blunt lobe on either side (Figure 7A). RADULA.— The radular formula is 16x 1.0.1. The lateral teeth have 25-30 fine, triangular denticles (Gosliner 1996) (Fig. 7B). The three gizzard plates are long, oval and equal in size and shape (Fig. 7C). S1zE.— 0.5 to 3.0 mm in length. RANGE.— Described from South Coronado Island. Gosliner extends the range to the Santa Maria Basin, off Point San Luis and Point Sal, California. B C FiGure 7. Philine polystrigma (Dall, 1908). A. Drawing of animal. B. Lateal tooth. C. Gizzard Plate. (After Gosliner 1996.) Philine sp. 1 Gosliner (1996) reports two undescribed species from the Santa Maria Basin, California. IDENTIFICATION.— Philine sp. 1 is morphologically similar to P. polystrigma as discussed above (Gosliner 1996) (Fig. 8A). The shell has a characteristic pointed lobe (Fig. 8B). RADULA.— The radular formula is 15 x 1.1.0.1.1. The inner lateral teeth have 22—34 elongate denticles (Gosliner 1996) (Fig. 8C). The outer lateral teeth are simple and smooth. The three giz- zard plates are equal in size. They have a prominent ridge down the centerline (Gosliner 1996). NATURAL History.— Collected only on fine soft sediments in 90-155 m of water. Probably feeds on Foraminifera (Gosliner 1996). S1zeE.— 1-5 mm in length. RANGE.— Known from the Santa Maria Basin, off Point San Luis and Point Sal, California (Gosliner 1996). FiGure 8. Philine sp. 1. A. Drawing of animal. B. Lobe of shell. C. Inner Lateral tooth. (After Gosliner 1996.) 18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Philine sp. 2 Gosliner (1996) reports a second undescribed Philine from the Santa Maria Basin, California. IDENTIFICATION.— A tiny species, the anterior shield is short and rounded posteriorly, where- as the posterior shield is elongate, terminating in skirt-like lobes (Gosliner 1996). RADULA.— This species has a unique radular formula of 13 x 6.1.0.1.6 (Gosliner 1996). The inner lateral teeth are broad and hook-shaped and bear no denticles (Gosliner 1996). The outer lat- eral teeth are narrow hook-shaped and are also free of secondary denticles (Gosliner 1996). SIZE.— Approximately | mm in length. RANGE.— Known only from the Santa Maria Basin, off Point San Luis, California (Gosliner 1996). Family Aglajidae Aglaja sp. | A new, undescribed species of Aglaja is reported from Santa Maria Basin, California (Gosliner 1996). IDENTIFICATION.— The body is a uniform whitish brown. Both ends of the head shield are rounded. The posterior shield terminates in an elongate tail on the left side of the body and a short- er lobe on the right side (Gosliner 1996). BuccAL MorPHOLOGY.— There are no radula or gizzard plates (Gosliner 1996). S1zE.— 0.5—1 mm in length. RANGE.— Known only from the Santa Maria Basin, off Purisima Point and Point Sal, California (Gosliner 1996). Family Philinoglossidae Philinoglossa sp. Nine specimens of this undescribed species have been collected on a fine sand bottom in 65 ft. of water off San Diego (Cadien 1998). The specimens ranged in size from 1—2 mm in length, suggesting that this species may be quite common but overlooked due to its small size (Plate 1B). According to Cadien (1998), the species is ovate-elongate and lacks a head shield, parapodia, exter- nal gills, rhinophores, oral tentacles, attenuated foot corners, posterior mantle lobes surface eyes and an external shell. The color is tan, with darker reddish brown visceral mass showing through. Internally there is no shell, jaws or gizzard plates. The radula formula is 15 x 2.1.0.1.2. The lateral teeth are flat and appressed, the cusp simple, curved and acute (Plate 1C). The lateral tooth is larger, stronger and strong shouldered. The central cusp is curved with some small marginal denticles. Family Akeridae Akera sp. The benthic monitoring program carried out in southern California, referred to a Bight °98 EMAP project, has collected an unidentifiable species determined to belong to the genus Akera (Ron Veralde and Kelvin Barwick, pers. commun.). The specimen was collected in 40 m of water near the south western shore of Santa Catalina Island. As seen in Plate 1D, the body of the animal is cream colored. The internal organs can be seen through the external coiled shell, which has a depressed spire. BEHRENS: PACIFIC COAST NUDIBRANCHS 19 Order Anaspidea Family Aplysiidae Aplysia parvula Gilding in Morch, 1863 This species occurs worldwide in warm temperate waters and has recently been observed in southern California (Bruce Wright, pers. commun. ). IDENTIFICATION.— Typical sea hare body form. Mottled brown in color, the parapodia has a dark blue-black line along the edge of the parapodia (Plate 1E). The parapodia encloses the man- tle cavity. S1ZE.— To 50 mm in length. RANGE.— Worldwide tropical to sub-tropical. Along the Pacific Coast of North America from the Gulf of California to Galapagos Islands (Gosliner 1991), and San Clemente Island. Order Notaspidea Family Pleurobranchidae Pleurobranchus cf. areolatus (Moérch, 1863) Although originally thought to be a new species of Pleurobranchus, because it was so far out- side of the range of P. areolatus and because it lacked the characteristic white speckles on the rhinophores and white snow flakes over the notal tubercles, internal anatomical similarities suggest we should refer to it as a color variation of P. areolatus until further studies are conducted, suggest- ing otherwise. IDENTIFICATION.— A large pleurobranch, the notal surface is covered with large smooth tuber- cles (see Plate 1F). The largest tubercles are brown whereas smaller ones are golden brown. In juveniles, the border between the tubercles is white whereas in larger specimens is @ light tan to peach color. The rolled rhinophores are tan to orange. RADULA.— 90 x 230.0.230. S1zeE.— To 190 mm in length (Angulo-Campillo et al. 2001). RANGE.— The new occurrences outside the Gulf of California, include: Isla Revillagigedos, Mexico (K. Kaiser), San Benitos Island (E. Houcke), La Jolla (P. Hughes) and San Clemente Island (M. Tegner, K. Miller and C. Gramlick), San Diego County, Anacapa and Catalina Islands, California (J. Engle, E. Erikson, R. Herrmann and D. Richards). Previously, Atlantic coast, Florida and Caribbean to Canal Zone, Barbados and Curacao, and Gulf of California to Panama. Order Sacoglossa Family Stiligeridae Placida sp. Discovered by Jim Lance and Wes Farmer, the species is undescribed. IDENTIFICATION.— Aecolid-like in appearance, this sacoglossan has a cream colored head and green cerata covered with white specks and have white tips (Plate 1G). The long cephalic tentacles are also white at the tips. RADULA.— Unknown NATURAL History.— This species lives on intertidal mudflats and feeds on the green alga, Codium magnum. S1ZE.— To about 15 in length. RANGE.— To date known only from Bahia San Quintin, Baja California, Mexico. 20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Order Nudibranchia Suborder Doridacea (= Doridina) Family Gnathodoridoidea Bathydoris aioca Marcus and Marcus, 1962 Originally described from Guadalupe Island, Baja California, Mexico, I was unaware of any photographs of this species and, due to its depth of occurrence, I excluded it from the 1991 edition of Pacific Coast Nudibranchs. The species has now been dredged off the Oregon coast. IDENTIFICATION.— Valdés and Bertsch (2000) report that although the color of the living ani- mal is unknown, the dorsum is smooth and preserved specimens give no evidence of spots or other color markings (Plate 1H). The rhinophores are quite long and bear 45 lamellae. The gill consists of 18 bipinnate branchial leaves. RADULA.— The radular formula is 53-64 x 64—81.1.64—81. NATURAL History.— Dredged from depths of 2700-2850 m. S1zE.— To 71 mm in length. RANGE.— Known only from the type locality off Guadalupe Island, Mexico, and Oregon (Valdés and Bertsch 2000). Family Goniodorididae Okenia sp. 1 A new species of Okenia has been showing up regularily in southern California (Plate 11). It has been encountered at diver depths off Huntington Beach, Newport Beach, San Onofre and Redondo Beach on worm tubes on soft bottoms (Don Cadien, pers. commun.). Further south in San Diego, the species is found in association with the solitary tunicate, Mogula sp. (Ron Velarde and Megan Lilly, pers. commun.). Preliminary evaluations indicate that the species is undescribed but shares some anatomical features with Okenia aspersa (Alder and Hancock, 1845) from British waters (D. Cadien, pers. commun. ). Okenia sp. 2 A second undescribed species of Okenia is reported from the Santa Maria Basin, California (Gosliner 1996). IDENTIFICATION.— The notum has 3-4 papillae present along either lateral margin of the notum (Gosliner 1996). There are four gills arranged in a semi-circle (Gosliner 1996). RADULA.— The radula formula is 18 x 1.1.0.1.1. The inner lateral teeth are large, curved with 7-8 denticles on the inner side (Gosliner 1996). The outer lateral teeth are narrow and bicuspid. SIZE.— | mm in length. RANGE.— Known only from the Santa Maria Basin off Point Sal, California (Gosliner 1996). Family Onchidorididae Calycidoris guentheri Abraham, 1876 The distribution of this species has previously been given as Arctic Seas to Bering Strait in 1444 meters of water. Nora R. Foster and Rae Baxter were responsible for the first collections of this species in the northern Pacific, the southern most being from the central Bering Sea. It is a syn- BEHRENS: PACIFIC COAST NUDIBRANCHS DiI onym of Doris (Adalaria) sibirica Aurivillius, 1887 (= Calycidoris guentheri, fide T. Gosliner, pers. commun.). See complete description below. IDENTIFICATION.— Typically doridoid in shape, the body is large, slightly wider in front and depressed. It is densely covered with small conical spiculose tubercles which vary in size. The ground color is tan to dark brown, often with reddish pink to purple. The head is a narrow, thick, veil. It has 18—23 pinnate gills in a circle around a central anus, which contract into a sheath. RADULA.— The radular formula is 25-28 x 3.1.0.1.3. NATURAL HisTory.— Dredged from depths of 9-270 m, usually 9-60 m (Roginskaya 1972). S1zE.— To 32 mm in length. RANGE.— Widespread in the Arctic Ocean from the Barents Sea to the Chukchi Sea and Bering Strait (Roginskaya 1972); also found across the Beaufort Sea to Liverpool Bay, North WestTerritory (S. Millen, pers. commun.). Specimens are also reported from the Bering Sea (Goddard and Foster 2002). ETYMOLOGyY.— Named for Dr. Albert Giinther of the British Museum of Natural History. Family Discodorididae Diaulula greeleyi (MacFarland, 1909) Peltodoris nayarita Ortea and Llera, 1981, was described from the Nayarit coast of Mexico by Ortea and Llera (1981). Since then it has been collected within the coverage of Pacific Coast Nudibranchs, 2nd ed., at Punta Eugenia (Bertsch et al. 2000). Camacho-Garcia and Valdés (2003) report that this species is synonymous with Diaulula greeleyi (MacFarland, 1909). IDENTIFICATION.— An orange red dorid with brown blotches (Plate 2A). The dorsum is tuber- culate, bearing caryophyllidia. The rhinophores and gills pockets are elevated. RADULA.— The radular formula is 37 x 55.0.55. The lateral teeth are hamate, having a single cusp and lacking denticles. Size.— To 100 mm in length. RANGE.— Florida, Brazil, South Carolina, Isla Isabela, Nayarit, Mexico and Punta Eugenia, Baja California, Mexico, and the Pacific Coast of Costa Rica (Camacho-Garcia and Valdés 2003). Family Polyceratidae A recent, deepwater study in the Santa Maria Basin of Central California has produced a num- ber of previously undescribed species (Gosliner 1996). The following genus, new to the Pacific Coast of North America, was represented among specimens collected during that study. The new species has yet to be described. Holoplocamus sp. Among the new species collected in the Santa Maria Basin study (Gosliner 1996) was this primitive polyceratid. Coincidentally, Shrake (1977) reported the collection of five specimens taken in 600 ft of water at locations, also in the Santa Maria Basin (Platform Harvest and Platform Hildago), of a species identified at the time, as Triopella sp., another primitive polyceratid. Subsequent discussions with Don Cadien, who originally identified the specimens as Triopella, have led to the conclusion that those specimens were likely the same as Gosliner’s species report- ed here as Holoplocamus sp. IDENTIFICATION.— The body is elongate with 11—15 spiculate velar tentacles and 4—6 lateral appendages per side of the body (Fig. 9). The three gills are unipinate to bipinnate. The mouth is ED PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 flanked by a pair of elongate tentacles (Gosliner 1996). RADULA.— The radular formula is 30 x 7-8.2.1.2.7-8. See Gosliner (1996) for SEMs of the radular morphology. NATURAL HIstToRy.— Collected by trawl net on both hard and soft bottoms. SIZE.— To 0.5—1.5 mm. RANGE.— Know only from the Santa Maria Basin, Platform Harvest and Platform Hildago (Shrake 1977) and off Morro Bay and Purisima Point, San Luis Obispo FIGURE 9. Holoplocamus sp. Drawings of preserved speci- County, California (Gosliner 1996). men. Dorsal and ventral views. (After Gosliner 1996.) Colga pacifica Bergh, 1894 IDENTIFICATION.— This phanerobranch dorid looks very much like a Triopha. The notum has a pre-branchial notal processes. A main row of elongate processes runs down the middle of notum and usually a pair of rows of similar processes accompanied it along both sides. In some specimens these lateral rows are almost the same length as the middle row. Specimens vary from pure white with yel- low gills and rhinophores shown above, and as described originally by Bergh, to speci- mens that are completely orange (Plate 2B). RapuLa.— The radular formula IS FIGURE 10. Colga pacifica. Drawing of jaw and radula. (After x 5—6.1.1.1.1.1.5-6. Drawings of the jaw Martynoy and Baranets 2002) and radula are presented in Fig. 10. NATURAL HIstory.— This species feeds on various bryozoans. The specimens shown here were collected between 90-275 m deep. S1ZE.— To about 20 mm. RANGE.— Alaska to Shikotan Island (southern Kuril Islands); southwest Greenland. Family Dendrodorididae Doriopsilla spaldingi Valdés and Behrens, 1998 This recently discovered species has probably been passed over for years as a common yellow porostome (Doriopsilla albopunctata or Dendrodoris fulva) with a light margin. The collection of several specimens from the La Jolla area bearing a distinctive blue margin has drawn attention to the identity of this species (J. Lance, pers. commun.). IDENTIFICATION.— As mentioned above, this species looks strikingly similar to Doriopsilla albopunctata (= Dendrodoris fulva). Gills and rhinophores are yellow-orange and the notum has evenly distributed low tubercles. The margin is the key identifying feature; it is highly undulated and white to iridescent blue in color (Plate 2C). Close analysis by Jim Lance indicates that in iri- descent blue specimens, the tissue morphology is similar to that of the blue in the aeolid Hermissenda, being a concentration of coalesced sacs acting as vesicles that hold the color platelets together. BEHRENS: PACIFIC COAST NUDIBRANCHS Pop} RaADULA.— None NATURAL History.— Collected in depths of 200 feet in Scripp’s Canyon, to 80 feet, and in the Channel Islands. This species lays a flat coiled egg mass, which produces direct development or lecithotrophic larvae (J. Lance, pers. commun.). S1zE.— To 83 mm. RANGE.— South Coronado Island, Baja California, La Jolla Submarine Canyon and Pt. Loma, San Diego County, and San Miguel Island, California. It has also been observed at 210 ft. at The Pinnacles, Carmel, California (Clinton Bauder, pers. commun.). ETyMOLOGy.— Named for George E. Spalding III of Solana Beach, California, who discov- ered this species. Suborder Dendronotacea (= Dendronotina) Family Tritoniidae Tritonia pickensi Marcus and Marcus, 1967 IDENTIFICATION.— Similar in appearance to Tritonia myrakeenae, the body is translucent and bears an opaque white region mid-dorsally, which branches to extend to the base of each gill (Plate 2D). There are approximately 12 gill tufts. NATURAL History.— Found on the red gorgoniam, Lophogorgia chilensis, to depths of 65 ft or more. SIZE.— To about 25 mm. RANGE.— La Jolla Canyon, San Diego and California Channel Islands, California; Bahia de los Angeles to Cabo San Lucas, Baja California (Skoglund 2002), Puerto Vallarta, Mexico (Hermosillo-Gonzalez 2003) and Costa Rica (INBIO 2003). ETYMOLOGyY.— Named to honor Dr. Peter E. Pickens, University of Arizona, who collected extensively in the Gulf of California for the Marcus’. Family Scyllaeidae Crosslandia daedali Poorman and Mulliner, 1981 This species had been known only from the mainland of Mexico and the Gulf of California. IDENTIFICATION.— The species has lobed parapodia, the first of which is largest (Plate 2E). The edges of the lobes are irregularly notched and digitate and have numerous branchial tufts along the edges. The color varies from orange to green and red. There are fine brown lines along the body as well as some brown spots. The species also has a series of 5—6 bight blue spots along the sides and dorsally. NATURAL History.— On the mainland, the species was found on the alga Padina sp., where it grazes on small hydroids that live on the alga. S1zE.— To 25 mm in length. RANGE.— First reported on this coast from Punta Eugenia, Baja California (Bertsch et. al. 2000), the species is now known to range to Guaymas in the Gulf of California and south to Puerto Vallarta, Mexican mainland (Hermosillo-Gonzalez 2003). Family Dendronotidae Dendronotus sp. It is not known if this is the same as Dendronotus sp. 1, (Species #148), which is known from 24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Alaska and as far south as the California Channel Islands, as I have no specimens to make this determination. What is known is that this very large species, reaching over 7 inches, matches no other species recorded from along this coast. I have received several photographs of this species but as yet no specimens by which to set the record straight. Note the white specks on a red body as described for Dendronotus sp. 1, but the handsome specimen shown in Plate 2F has a wide patch of opaque white along its mid-dorsal line. IDENTIFICATION.— A large, red-bodied Dendronotus with white specks covering the entire body. Some specimens may have a wide patch of opaque white along its mid-dorsal line. RADULA.— Unknown. NATURAL History.— Nothing is known of this species’ habits. SIZE.— Large, to 180 mm in length. RANGE.— Seymour Inlet, Tredwell Bay, Vancouver area, British Columbia. Suborder Arminacea (= Arminina) Family Arminidae Armina cordellensis Gosliner and Behrens 1996 IDENTIFICATION.— Grey in color, the notum is granular with irregularly spaced, low, rounded tuberacles (Fig. 11). RADULA.— 46 X 42.1.69. The rachidian teeth are broad with a large triangular central cusp, flanked by 8—13 elongate denticles per side (Fig. 12). The lateral teeth are triangular with 7—9 short triangular denticles (Fig. 12). NATURAL History.— The type specimen was collected at a depth of 46-52 m. SIZE.— 60 mm. RANGE.— Known only from Cordell Banks, 20 km off Point Reyes, California. ETYMOLOGyY.— Named for the type locality, the Cordell Bank. FIGURE 11. Armina cordellensis Gosliner and Behrens, 1996. Draw- FIGURE 12. Armina cordellensis ing of preserved specimen. Gosliner and Behrens, 1996. Radular teeth. Janolus sp. IDENTIFICATION.— Similar in coloration to Dirona pellucida Volodchenko, 1941(= Dirona aurantica [Hurst, 1966]), the body being transparent to orange with white specks and a white line up each ceras and each rhinophore. The digestive gland is clearly seen within each ceras (Plate 3A), unlike D. pellucida, where the gland is absent. RADULA.— Unknown. NATURAL History.— Nothing is known of this species’ habits. SIZE.— 3040 mm. BEHRENS: PACIFIC COAST NUDIBRANCHS 25 RANGE.— Photographed in Quarry Bay, Nelson Island, British Columbia in 60 feet of water by Bernard P. Hanby (Andy Lamb, pers. commun.). Suborder Aeolidacea (= Aeolidina) Family Flabellinidae Flabellina bertschi Gosliner and Kuzirian 1990 Until recently, this species was known only from the Pacific coast of Panama and within the Gulf of California. Goddard and Schickel (2000) have reported the species from the outer coast of Baja California. IDENTIFICATION.— This species is typically flabellinid in shape. The body is translucent with an opaque white band on the notum (Plate 3B). The cerata are orange-red with white tips. The rhinophores are smooth. The oral tentacles and rhinophores are tipped with white. RADULA.— The radular formula is 24-31 x 1.1.1. The rachidian tooth has 6-8 denticles, the laterals 9-11 denticles. NATURAL History.— Little is known about this species. S1zE.— To 30 mm length. RANGE.— Known from Pacific coast of Panama (Gosliner 1994), Puerta Vallarta, Mexico (Hermosillo-Gonzalez 2003), Costa Rica INBIO 2003), Gulf of California (Gosliner and Kuzirian 1990), Punta Rosarita on the west coast of Baja California (Goddard and Schickle 2000), and Little Farnsworth, Catalina Island, California (E. Erikson, pers. commun.). Also from the Galapagos (Gosliner 1994). ETYMOLOGyY.— Named to honor marine biologist and friend, Hans Bertsch. Flabellina cf. islandica (Odhner, 1937) This species is possibly the same as Flabellina islandica, which 1s known from Iceland (Sandra Millen, pers. commun. ). IDENTIFICATION.— This species is typically flabellinid in shape, but slightly stouter and wider, an adaptation believed to be related to its soft bottom existence. The ground color is translucent pinkish cream with stubby red cerata (Plate 3C). Opaque white specks occur on the dorsal surface and cerata, the ends of the smooth rhinophores and the extremely long oral tentacles. RADULA.— The radular formula for specimens from Iceland is 15 x 2.1.1. The rachidian has 5-7 denticles, the laterals are smooth and there is a small outer lateral on the left side of some spec- imens. NATURAL History.— Shallow subtidal on mud bottoms in the spring and summer. S1ZE.— To 22 mm length. RANGE.— Known from southern British Columbia, Victoria Harbor (Vancouver Island) and Vancouver Harbor (mainland) (Neil McDaniel and Sandra Millen, pers. commun.). Similar animals have been found in the Sea of Japan (Sandra Millen, pers. commun.). Family Tergipedidae Cuthona sp. IDENTIFICATION.— Another transparent tergipedid with brown ceratal cores. Internal anatomy indicates it is a new species (J. Lance, pers. commun.). There are 8—9 rows of cerata, tipped with a white cnidosac. The tips of the rhinophores and cephalic tentacles are white encrusted. 26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 RADULA.— Unknown. NATURAL History.— Found on the hydroid, Tubularia sp., on boat docks (J. Lance, pers. commun. ). SIZE.— 5-9 mm in length. RANGE.— San Diego, California (J. Lance, pers. commun.). Family Aeolidiidae Aeolidia herculea Bergh, 1894 Aeolidia farallonensis Gosliner and Behrens, 1996 (synonym). Aeolidia grandis Volodchenko, 1941 (synonym). Gosliner and Behrens (1996) described two new deepwater species from northern California. One was Aeolidia farallonensis, described from several specimens collected from 510 and 1400 m deep off the Farallon Islands, Gulf of the Farallons, California. At the time the authors described A. farallonensis, A herculea was regarded as a synonym of Aeolidia papillosa (Linnaeus, 1781) (Marcus 1961). At the suggestion of colleague A.V. Martynov, the authors reviewed Bergh’s orig- inal description and found these species to be identical. IDENTIFICATION.— Similar in external morphology to Aeolidia papillosa (Linnaeus, 1761) (Species no. 212) except the rhinophores are rugose (Fig. 13A) rather than smooth, and it has fewer rows of cerata, but with nearly twice the number of cerata per row (Gosliner and Behrens 1996). Internal anatomies clearly differentiate these two species. The color of the living animal is unknown. RADULA.— 30 x 0.1.0. Each tooth bears 38—53 shallow triangular denticles (Fig. 13B). The jaws (Fig. 13C) are twice as long and more than twice as wide as those of a similar size A. papil- losa. The jaws of A. herculea are similar in size and shape to those described for A. farallonensis. NATURAL History.— Specimens have been collected from 510 and 1400 m deep. SIZE.— 25 mm. RANGE.— Originally described from Alaska. Gosliner and Behrens (1996) add the Farallon Islands, Gulf of Farallons, California. Additional collections have been recorded from Monterey Bay, 570-700 m deep (Jim Nybakken, pers. commun.). FicureE 13. Aeolidia herculea, Bergh, 1894. A. Rhinophore; B. Radular tooth; C. Jaw. (Drawings after Gosliner and Behrens 1996.) BEHRENS: PACIFIC COAST NUDIBRANCHS Di Plate 1 I A. Philine auriformis Suter, 1909, living animal, San Francisco Bay, California (T. Gosliner); B. Philinoglossa sp. Dorsal and ventral views of preserved specimens, San Diego, California (Kelvin Barwick); C. Philinoglossa sp. Radula; D. Akera sp., preserved specimen, dorsal and ventral view, Santa Catalina Island, California (Kelvin Barwick); E. Aplysia parvula, living animal San Clemente Island, California (Bruce Wright). F. Pleurobranchus cf. areolatus, living animal, La Jolla, California (David Behrens); G. Placida sp., living animal, Bahia San Quintin, Baja California, Mexico (Wes Farmer and Jim Lance); H. Bathydoris aioca, preserved specimen, Oregon (Hans Bertsch); I. Okenia sp., freshly preserved speci- men, Huntington Beach, California (Sarah Douglass). 28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Plate 2 A. Diaulula greeyeli (MacFarland, 1909), living animal, Punta Eugenia, Baja California, Mexico (Hans Bertsch); B. Colga pacifica, living animal, Kuril Islands, Alaska (Roger Clark); C. Doriopsilla spaldingi Valdés and Behrens 1998, liv- ing animal, San Diego, California (Mike Miller); D. Tritonia cf. pickensi, living animals, Channel Islands, California (Kathy deWet); E. Crosslandia daedali, living animal, Puerto Vallarta, Mexico (Ali Hermosillo-Gonzalez); F. Dendronotus sp., living animal, Seymour Inlet, Tredwell Bay, Vancouver, British Columbia (Bernard P. Hanby). BEHRENS: PACIFIC COAST NUDIBRANCHS 29 A. Janolus sp., living animal, Quarry Bay, Nelson Island, British Columbia (Bernard P. Hanby); B. Flabellina bertschi, living animal, Puerto Vallarta, Mexico (Ali Hermosillo-Gonzdlez). C. Flabellina cf. islandica, living animal, Victoria Harbor, Vancouver Island, BC, Canada (Neil McDaniel). D. Ancula gibbosa, living animals, Southern California (Bruce Wright); E. Ancula gibbosa, living animals, Vancouver, British Columbia area (Charlie Gibbs); F. Onchidoris bilamellata, mass feeding on barnacles in Monterey, California (Clinton Bauder); G. Onchidoris bilamellata, mass spawning in Monterey, California (Clinton Bauder). 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Plate 4 2 A. Taringa aivica, living specimen, Puerto Vallarta, Mexico (Ali Hermosillo-Gonzalez); B. Dendrodoris sp. 2., living animal (Clinton Bauder); C. Flabellina trophina, living dark color morph, Coronation Island, Alaska (Clinton Bauder); D. Anetarca armata, feeding on hydoids living on shell of living Decipifus californicus, Bahia de Los Angeles, Baja California, Mexico (James R. Lance); E. Decipifus californicus covered with Anetarca egg ribbon, Bahia de Los Angeles, Baja California, Mexico (James R. Lance); F. Cerberilla cf. pungoarena, living animal, Channel Islands, California (Kathy deWet). BEHRENS: PACIFIC COAST NUDIBRANCHS 31 NEw INFORMATION AND NOMENCLATURAL CHANGES The following is an update to the information presented in the second edition of Pacific Coast Nudibranchs (Behrens 1991). The numbers listing the species below correspond to the species identification numbers used in that text. Included in brackets are new names given species that were reported as undescribed in 1991, synonymies and nomenclatural changes. New information on color variation and possible taxonomic confusion, and extensions of the known ranges are also presented. A comprehensive study of the food preferences of nudibranchs worldwide was published by McDonald and Nybakken (1997) Referencing the prey species of the nudibranchs of this coast here would be too voluminous and the mere number of references would surely triple the length of this document. I refer you to the original document at — http://www. veliger.org/nudibranch_food.html. Order Cephalaspidea 1. Rictaxis punctocaelatus (Carpenter, 1864) Mikkelson (1996) reviewed the Cephalaspidea and proposed moving the Opisthobranch fam- ily Acteonidae from the Opisthobranchia, to the Gastropod subclass Heterobranchia. I do not sup- port this move here. The range of this species should be extended into the Gulf of California to Bahia San Carlos, Sonora, Mexico (Poorman and Poorman 1988). 5. Acteocina harpa (Dall, 1871) The range of this species is extended south to Rocas Alijos, Mexico (McLean and Coan 1996). 8. Cylichna alba (Brown, 1827) This little known species has been observed recently in Boundary Bay, Vancouver, British Columbia on the estuary’s mud flat, hidden in drying mats of drift eel grass (Bill Merilees, pers. commun.). This species has been found in the guts of western sandpipers. 11. Diaphana californica Dall, 1919 The range of this species is extended north to Kayostla Beach, Olympic Peninsula, Washington (Jeff Goddard et al. 1997). 21. Navanax inermis (Cooper, 1863) The range of this species is extended north to Bolinas Lagoon, Marin County, California, where adult specimens were collected in December 1992 during that year’s El Nifio event. 22. Navanax polyalphos (Gosliner and Willaims, 1972) The species is now reported as far south as Costa Rica (Camancho-Garcia, pers. commun.). 23. Philine bakeri Dall, 1919 [now: Philine alba Mattox, 1958] The correct designation for the photo of the Philine shown in Behrens, 1991, photo 23, has been bantered back and forth. Due to the size of the specimen (15 mm), it should be listed as Philine alba. 3) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Order Anaspidea Family Aplysiidae 24. Aplysia californica Cooper, 1863 The range of this species has been extended to Hachijo Island, Japan where several 70-cm specimens have been collected (Jun Imamoto and Nishina Masayoshi, pers. commun.) and to El Salvador (Hernandez C. 1992). Order Notospidea Family Pleurobranchidae 30. Berthella californica (Dall, 1900) A specimen closely matching Berthella californica was recently trawled from 400 feet in the Galapagos Islands (T. Gosliner, pers. commun.). This extension should be considered tentative, however. Martynov (1997) reports this species from Beringa Island, Peter the Great Bay, Sea of Russia. 32. Berthella strongi (MacFarland, 1966) The range of this species is extended north from Moss Beach, California, to Nanaimo, British Columbia (Mike Miller, pers. commun.), and south from Santa Cruz Island to Punta Rosarito, Baja California (Goddard and Schickel 2000). 33. Berthella engeli (Gardner, 1936) [now: Berthellina ilisima Marcus and Marcus, 1967] For sometime there has been uncertainty concerning the proper placement of the bright orange pleurobranchs on this coast and elsewhere. Berthella engeli and B. citrina have been used inter- changably throughout the literature. A recent study of all the smooth orange Berthella’s worldwide undertaken by Terrence Gosliner and Lucas Cervera concludes that the eastern Pacific species is Berthellina ilisima (Juan Lucas Cervera, pers. commun.). Order Sacoglossa A review of the natural history of the sacoglossans of the northeastern Pacific can be found in Trowbridge (2002). 36. Elysia hedgpethi Marcus, 1961 Hermosillo-Gonzalez (2003) reports this species from Puerto Vallarta, Mexico. 38. Ercolania boodleae (Baba, 1938) The range is extended into the Gulf of California at Puerto Penasco (Trowbridge 2002). 39. Placida dendritica (Alder and Hancock, 1843) The range of this species is extended north to Bertha Bay, Chichagof Island, Alaska, where it has been collected on Codium fragile (Rita O’Clair, pers. commun.), and south into Gulf of California at both Bahia de los Angeles, Baja California, and Bahia San Carlos, Sonora, Mexico (Trowbridge 2002), on Codium setchelli (Cynthia Trowbridge, pers. commun.). BEHRENS: PACIFIC COAST NUDIBRANCHS 33 41. Stiliger fuscovittatus Lance, 1962 Please note that the photo in the 1991 edition of Pacific Coast Nudibranchs shows a specimen on the algae, Microcladia. This algal species is not fed upon by Stiliger and, additionally, is rarely associated with it (Cynthia Trowbridge, pers. commun.). 42. Alderia modesta (Loven, 1844) This species was previously reported as far south as Newport Bay, California. It has now been reported from San Quintin, Baja California, Mexico (J. Lance, pers. commun.). The range is also extended north to Prince William Sound, Alaska (Goddard and Foster 2002). Krug (1998) reports that Alderia modesta practices poecilogony, or the production of eggs of different sizes that yield either planktotrophic veligers or lecithotrophic veligers, depending on their available yolk supply. Back crosses of these two forms yield the same ratio of reproductive types seen initially and give evidence of their conspecificity. Starved individuals, which had previ- ously spawned exclusively or primarily lecithotrophic larvae, switiched to production of plank- totrophic larvae after a short period of no food. Whether larvae are planktonic or not significantly affects the dispersal potential of a spawning event. 43. Aplysiopsis oliviae (MacFarland, 1966) [now: Hermaea oliviae (MacFarland, 1966)] The taxonomic placement of this species has changed several times. As I suspected (Behrens 1991), the morphology of the radular teeth places this species in Hermaea (Kathe Jensen, Sea Slug Forum, 7 November 2001). The range of this species is extended south to El Morro, Isla Cedros (Bertsch et al. 2000) and north to Seymour Inlet, British Columbia (Marli Wakeling, pers. commun.). 44. Aplysiopsis enteromorphae (Cockerell and Eliot, 1905) The color of the body in adults (15—25mm) is yellowish-white with patches of greenish black. Small specimens (<5—10mm in length) are much darker than their larger peers (Cynthia Trowbridge, pers. commun.). Trowbridge also points out that specimens feeding on Chaetomorpha are browner than those feeding on Cladophora, infact they shimmer with a golden-reddish reflectance. The range is extended south to Bahia San Quintin, Baja California (C. Trowbridge, per. comm.), and Puerto Vallarta, Mexico (Hermosillo-Gonzalez 2003). 45. Hermaea vancouverensis (O’ Donoghue, 1924) This species feeds on Isthmia nervosa in Oregon (C. Trowbridge, per. comm.) as was previ- ously documented by Williams and Gosliner (1973). 46. Olea hansineensis Agersborg, 1923 The range of this species is extended to Prince William Sound, Alaska (Goddard and Foster 2002), where it was found feeding upon the egg mass of the cephalaspidean, Melanochlamys diomedea. Specimens of this species may reach 13 mm (Chia and Skeel 1973). 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 Order Nudibranchia Suborder Doridacea (= Doridina) 47. Corambe pacifica MacFarland and O’Donoghue, 1929 The genus Corambe has been subjected to several nomenclautural reviews and changes in recent years. Martynov (1994) divided Corambe into several newly defined genera and assigned C. pacifica to the genus Gulbinia. More recently, Valdés and Bouchet (1998) have reexamined the value of the characters Martynov used to differentiate the new genera and have simplified the tax- onomy to three genera, reestablishing, this species as Corambe pacifica. The range of this species is extended to the south to Mismaloya, Puerto Vallarta, Mexico (Hermosillo-Gonzalez 2003). 48. Corambe sp. 1 [now: Loy thompsoni (Millen and Nybakken, 1991)] This species has now been described in the scientific literature and given the name Corambe thompsoni by Sandra Millen and Jim Nybakken (1991). The trivial name thompsoni is given in recognition of the British opisthobranch researcher Dr. Thomas Thompson. Martynov (1994) reas- signed the species to the genus Psammodoris. Valdés and Bouchet (1998), in their reorganization, placed it in the genus Loy Martynovy (1994). The radular teeth, which were not included in PCN Supplement I — Rabula (Behrens 1992), are shown here (Fig. 14). The radular formula for this species is 45-48 x 5—6.1.0.1.5-6. The lateral tooth has a rectangular base with a long, denticulate hook. Also FIGURE 14. Loy thompsoni (Millen and Nybakken, 1991). A. Radular tooth. B. Drawing of arrangement of mantle spicules. (After Millen and Nybakken 1991.) AS SSOIINGE shown is the arrangement of the spicules in the mantle tissue. The range of the species is extended to Port Valdez, Alaska (Nora Foster, pers. commun.). 49. Doridella steinbergae (Lance, 1962) [now: Corambe steinbergae (Lance, 1962)] Like other corambid species, Doridella steinbergae has also undergone some recent nomen- clatural revision. Martynov (1994) created the genus Suhinia for D. steinbergae. Swennen and Dekken (1995, p. 104), placed this species in a newly established genus, Paracorambe. Here again, Valdés and Bouchet (1998), in their reexamination of the value of the characters used to differen- tiate the eleven proposed genera of Martynov (1994) and Swennen and Dekker (1995), simplified the taxonomy, referred this species to the genus Corambe. The range of this species is extended north to Little Takli Island, Alaska (Goddard and Foster 2002). 50. Okenia angelensis Lance, 1966 The range of this species is extended south to Chile, South America (Fischer and Ortea 1996). 51. Okenia plana Baba, 1960 Rudman (1995), in his review of the Chromodorididae of New Caledonia and the Noumea BEHRENS: PACIFIC COAST NUDIBRANCHS 35 romeri color group, when describing Cadlinella hirsuta, he clarified some taxonomic confusion in this genus that resulted in the combination of a number of species, including Okenia plana Baba, 1960, under Doris eolida and its placement in the genus Hopkinsia. Gosliner (2004:144), howev- er, places this species in the genus Okenia. 52. Okenia vancouverensis O’ Donoghue, 1921 The range of this species is extended south to Sechelt Inlet, Vancouver, British Columbia (Donna Gibbs and Andy Lamb, pers. commun. ). 54. Ancula lentiginosa Farmer in Farmer and Sloan, 1964 The range of this species is extended south to Islote Mosquera, Isla Galapagos, Ecuador (Fischer and Ortea, 1996). It has been collected on several occasions (1995 and 2002) in 2300 ft of water off San Diego, California (Ron Velarde, pers. commun.). This disjunct occurrence is likely related to water temperature similarities between shallow Canadian and deep California waters. 55. Ancula pacifica MacFarland, 1905 [now: Ancula gibbosa (Risso, 1818)] This species has been synonymized with Ancula gibbosa (Risso, 1881) by Thompson and Brown (1984). The range of this species is extended north to Prince William Sound (Goddard and Foster 2002) and Beringa Island, Sea of Russia (Martynov 1997). On an ecological note, during the summer of 1996, hundreds of specimens were observed feeding on the entoproct, Barentsia, in Diablo Cove, San Luis Obispo, California (Mike Behrens, pers. commun.). In the Vancouver area, specimens of two color variations occur together. The color form shown in Plate 3D is typical of the color originally described for Ancula gibbosa and has not been record- ed from the southern part of the range on this coast. Plate 3E depicts the more typical southern color and the one shown in the 1991 edition of Pacific Coast Nudibranchs. 56. Trapania velox (Cockerell, 1901) The range of this reasonably uncommon species is extended north to Morro Bay, San Luis Obispo County, California, where it has be observed and collected commonly in deep holes in this coastal estuary (Michael Behrens and Duke McPherson, pers. commun. ). 57. Trapania sp. 1 [now: Trapania goslineri Millen and Bertsch, 2000] This beautiful little goniodorid was named to honor my friend and opisthobranch researcher, Terrence Gosliner, invertebrate zoologist and Provost of the California Academy of Sciences. With the description now published, we can now cite the radular formula of 37-41 x 1.0.1. The spawn is a narrow, white, upright ribbon, of 1 % coils, 5-6 mm in diameter (Millen and Bertsch 2000). Hermosillo-Gonzalez (2003) extends the range to Puerto Vallarta, Mexico. 58. Hopkinsia rosacea MacFarland, 1905 [now: Okenia rosacea (MacFarland, 1905)] In Gosliner’s (2004) review of the genus Okenia, he concluded that the genus Hopkinsia and its included species had to be referred to Okenia, which is the senior synonym. Gosliner expressed 36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 his regrets in having to take this action, but he believes it is justified based on recent phylogenetic analyses of the relevant morphological data (Gosliner, pers. commun.). 61. Acanthodoris lutea MacFarland, 1925 Publication of the northernmost collections of this species was overlooked during the prepara- tion of the second edition of Pacific Coast Nudibranchs. Goddard (1987) reports this species occur- rence as far north as Punta Gorda, Humboldt County, California. Goddard has now collected this species at North Cove, Cape Argo, Oregon, feeding on Alcyonidium sp. Subsequently, I have learned of this species occurrence in 60 feet of water at Indian Arm, near Vancouver, British Columbia (Andy Lamb, pers. commun.). 62. Acanthodoris nanaimoensis O’ Donoghue, 1921 The range of this species is extended north to Kachemak Bay, Alaska, near Homer, where it has been collected intertidally (Jane Middleton, pers. commun.). 64. Acanthodoris rhodoceras Cockerell and Eliot, 1905 The range of this species is extended south into the Gulf of California, to Bahia de los Ange- les (Hans Bertsch, pers. commun.) where it occurs in January, and north to the mouth of the Umpqua River, Oregon (Jeff Goddard, pers. commun.). 65. Adalaria jannae Millen, 1987 The range of this species is extended north to Prince William Sound (Goddard and Foster 2002) where it was found feeding on the bryozoan Membranpora, which was living on the kelp Laminaria. 66. Adalaria sp. 1 The range of this species is extended north to Prince William Sound (Goddard and Foster 2002). 70. Onchidoris bilamellata (Linnaeus, 1767) This barnacle-grazing dorid is not particularly common along the Pacific Coast, but when it is found, it is usually in large swarms. Clinton Bauder documented huge aggregations of this species in Carmel Bay, California, in December 2001 (Plate 3F). The presence of the nudibranch followed a massive settlement of the barnacle, Balanus crenatus. As the barnacle population declined, mass spawning by the nudibranchs occurred (Plate 3G). Similar observations of these large population explosions have been made by Bernard Picton (pers. commun.) in Galway, Ireland and by Jim Anderson (pers. commun.) off Scotland. 72. Aegires albopunctatus MacFarland, 1905 The range of this species is extended north to Mountain Point, Ketchikan, Alaska (Kurtis Morin and A. Murray, pers. commun.). BEHRENS: PACIFIC COAST NUDIBRANCHS 37 73. Crimora coneja Marcus, 1961 This is a rare species, previously reported to occur from Point Loma, San Diego, California, to Cape Arago, Oregon. Within this range it has only been found at two locations: rocky intertidal at Punta Gorda, California (Jeff Goddard pers. commun.) and subtidally within Morro Bay, California (Michael Behrens, pers. commun.). Goddard reports that it has been found in abundance only once, when 66 specimens were collected on a single low tide. In that area, Crimora feeds on the encrusting bryozoan, Hincksina minuscule. 74. Laila cockerelli MacFarland, 1905 [now: Limacia cockerelli (MacFarland, 1905)] The genus Laila, described by MacFarland in 1905, has been synonymized with the older Limacia Miiller, 1781 (Vallés et al. 2000). Limacia cockerelli has several color morphs and it has been speculated that these might repre- sent distinct species. The distinctive color form of Limacia with the large red blotches was thought to occur just in California as reported previously. Darcy Kehler (pers. commun.) photographed specimens exhibiting this color pattern at the very northernmost extreme of its range, in Bedwell Bay, Vancouver, British Columbia. Additionally, its range must now be extended north to Mountain Point, Ketchikan, Alaska (K. Morin and A. Murray, pers. commun. ). 75. Polycera alabe Collier and Farmer, 1964 The range of this species has been extended south to Puerto Vallarta, Mexico, Costa Rica (INBIO, 2003), and Chile (Schréd1 2003) and north, within Baja California to the Coronado Islands (Jerry Allen, pers. commun.), La Jolla Canyon (G. Spalding and M. Miller, pers. commun.), and Catalina, Anacapa, and Santa Barbara Islands (Erik Erikson, Dan Richards, and Kathy deWet, pers. commun.; also Lonhart and Tupen 2001). 77. Polycera hedgpethi Marcus, 1964 The range of this species 1s extended to Balboa, Panama (D. Cadien, pers. commun.) and to the Izu Pennisula and Suruga Bay, Japan (Okutani 2000). 78. Polycera tricolor Robillard, 1971 The range of this species is extended north to Hayden Passage, British Columbia (Jeff Rosenfeld, pers. commun.) and to Settler Cove, Ketchikan, Alaska (K. Morin and A. Murray, pers. commun.). 79. Polycera zosterae O’ Donoghue, 1924 [now: Palio dubia (M. Sars, 1829)] Polycera zosterae has been reassigned to the genus Palio by Rivest (1984). Goddard and Foster (2002) report P. zosterae as a junior synonym of the northern Sea of Japan species, Palio dubia (M. Sars 1829). Angulo-Campillo (2002) reports P. zosterae from La Paz, Baja California Sur, Mexico. Because this species is clearly temperate, further study is necessary to confirm the identity of the Mexican species. 80. Polycerella glandulosa Behrens and Gosliner, 1988 This species is now reported along the Pacific Coast of Central America to Costa Rica (INBIO 2003). 38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 81. Tambja eliora (Marcus and Marcus, 1967) Recently, members of the University of California at San Diego dive club, The Sea Deucers, have documented large numbers of this species as far north as Ensenada Bay. Individuals varied in size from 45-68 mm (Mike Miller, pers. commun.). The species is also reported from Puerto Vallarta (Hermosillo-Gonzalez 2003) and Costa Rica (INBIO 2003). 82. Tambja fusca Farmer, 1978 [now: Tambja abdere Farmer, 1978] Tambja fusca and T: abdere were both described by Farmer in 1978. These two species have been determined to be color variations of a single species; the name T: abdere takes precedence as it appears first in the publication. With the synonymy, it should be noted that the range of the species now extends south to Huatulco, Oaxaca, Mexico. Hermosillo-Gonzalez (2003) reports it from Puerto Vallarta, and INBIO (2003) reports the species from Costa Rica. 83. Triopha catalinae (Cooper, 1863) The range of this species is extended northwest to Slemya Island in the Aleutians (Goddard and Foster 2002). In a laboratory study, Geiger and Holyoak (1996) demonstrated that 7. catalinae prefers shad- ed area over lighted conditions even though its primary prey, erect bryozoans, occurs in both light and shaded environments. 5 84. Triopha maculata MacFarland, 1905 The range of this species is extended from Punta Cono to Punta Rosarito, Baja California (Goddard and Schickel 2000), and Beringia and Mednyi Islands, Sea of Russia (Martynov 1997). 85. Triopha sp. 1 [now: Triopha catalinae (Cooper, 1863)] The correct location of the collection that includes the specimen shown in the photograph (Behrens 1991) for this species is Boardman State Park, Oregon (Jeff Goddard, pers. commun.). Since his original collection, other specimens have been photographed by Steve Horvath and Marli Wakeling at Seymour Narrows, British Columbia, Canada. Based on an examination of the internal anatomy of specimens that were colleceted from Washington, British Columbia, and Alaska, Sandra Millen determined it to be a color morph of Triopha catalinae. 86. Cadlina flavomaculata MacFarland, 1905 The range of this species is extended to La Paz, Baja California Sur (Angulo-Campillo, 2002) and to Costa Rica (INBIO 2003). The specimens from Costa Rica may prove to be a new species (R. Johnson, pers. commun.). 90. Cadlina sparsa (Ohdner, 1921) Although this species was reported from the Juan Fernandez Island, far offshore of Chile, it has now been collected from the Chilean shoreline at Bahia de Coliumo, Chile (Schrédl 1997), and Argentina (Schrédl 2003). BEHRENS: PACIFIC COAST NUDIBRANCHS 39 91. Chromodoris galexorum Bertsch, 1978 Previously known only as far north as Guadalupe Island, Baja California, this species quite surprising showed up at Catalina Island, California (Jack Engle and Erik Erikson, pers. commun.). 92. Chromodoris macfarlandi Cockerell, 1901 Goddard (2000b) reports that this dorid feeds on both the pink sponge, Aplysilla glacialis and the deep red sponge, Aplysilla polyraphis in the La Jolla, California, portion of its range. 94. Glossodoris dalli (Bergh, 1879) Gosliner (1991), reports two specimens collected intertidally on Isla Fernandina and Isla Santa Cruz, Galapagos Archipelago. 99. Tyrinna evelinae (Marcus, 1958) The range of this species has been extended south to Costa Rica (Camacho-Garcia, pers. com- mun.) and to El Rubio, Peru (Schrédl and Millen 2001). 101. Jorunna pardus Behrens and Henderson, 1981 Since its original description, collection of numerous additional specimens has shown some degree of color variation. Individual specimens have been collected ranging from white with sparse black speckling to deep yellow with large round spots, the largest of which being dorso-medially positioned. A few specimens have been collected with no spots on the foot or tail. The degree of black pigment on the gills also varies from dense black to white tipped only slightly with black. 103. Aldisa albomarginata Millen, 1985 in Millen and Gosliner 1985 The range of this species is extended to Agamemnon Channel, Vancouver, British Columbia (D. Gibbs and A. Lamb, pers. commun.). 105. Aldisa sanguinea (Cooper, 1863) The northernmost range of this species has been extended by Jeff Goddard, who has collected three specimens at Otter Crest, Lincoln County, Oregon. 107. Rostanga pulchra MacFarland, 1905 Some clarification of the sponges Rostanga feeds upon should note that R. pulchra is chemi- cally attracted to and feeds on Ophlitaspongia pennata Labbe. Anderson (1971) adds Esperiopsis originalis and Plocamia karykina to the list of authenticated prey sponges. 108. Atagema alba (O’ Donoghue, 1927) This species was recently collected from a depth of 120 feet in Scripps Canyon, San Diego County, California. (Jim Lance, pers. commun.). 109. Sclerodoris tanya (Marcus, 1971) This species has been collected in Puerto Vallarta, Mexico (Hermosillo-Gonzalez, 2003). 40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 110. Archidoris montereyensis (Cooper, 1862) [now: Doris montereyensis Cooper, 1863] Bergh (1878) introduced the genus Archidoris based on Doris tuberculata, Doris montereyen- sis and others. Doris montereyensis has simple round tubercles, with spicules that do not protrude from the dorsal surface. For this reason, Valdés (2002) referred Archidoris to the synonymy of Doris Linnaeus, 1758. Aside from the obvious presence of this species’ preferred food, the encrusting sponges Haliclona sp. and Halichondria sp. in shaded habitats, a study reported by Geiger and Holyoak (1996) indicates that intertidally this dorid unquestionably prefers shaded over lighted conditions. Geiger and Holyoak’s study was conducted in the high intertidal and did not take into account the effects of current strength. Ryan Murphy (pers. commun.) suggests in his study, in an area of strong currents, that no preference was found between shaded and unshaded conditions. 111. Archidoris odhneri MacFarland, 1966 [now: Doris odhneri (MacFarland, 1966)| As reported above for Doris montereyensis, because Doris odhneri also has simple round tubercles, with spicules that do not protrude from the dorsal surface, it has been transferred to Doris Linnaeus, 1758 (Valdés 2002). The range of this species is extended north to the Bering Sea side of the Alaska Peninsula (Goddard and Foster 2002). 112. Anisodoris lentiginosa Millen 1982 [now: provisionally listed as cf. Peltodoris lentigi nosa (Millen 1982)] The placement of Anisodoris lentiginosa is problematic (see the discussion below for Peltodoris nobilis). Because Anisodoris lentiginosa 1s also not a caryophyllidia-bearing species, it is provisionally listed here as Peltodoris until a more complete analysis of this species is published. 113. Anisodoris nobilis (MacFarland, 1905) [now: Peltodoris nobilis (MacFarland, 1905)] Anisodoris nobilis was originally placed in the genus Montereina (MacFarland 1905) and later transferred to the genus Anisodoris. Valdés and Gosliner (2001), having conducted a careful review of caryophyllidia-bearing dorid nudibranchs, noted that the type species of Anisodoris was caryophyllidia-bearing and they, therefore, transferred it to Diaulula. Anisodoris nobilis, however, unlike the type species of the genus, has small rounded tubercles with no protruding spicules on the dorsal surface. Because it does not have caryophyllidia, this species recently has been trans- ferred to the genus Peltodoris Bergh, 1880 (Valdés 2002). 115. Diaulula sandiegensis (Cooper, 1863) The range of this species is extended northward to Kachemak Bay, Alaska (Jane Middleton, pers. commun.) and the central Aleutian Islands (Goddard and Foster 2002), and to Peter the Great Bay (Martynov 1997). In a study by Geiger and Holyoak (1996), this species prefers shaded over lighted conditions. Ryan Murphy’s findings mentioned above dispute this preference. Behrens and Valdés (2001) in their examination of the Doris (s.l.) species described the vari- ability in radular morphology of D. sandiegensis and presented evidence that the hamate teeth of Diaulala may have denticles in some specimens. 116. Diaulula sp. 1 [now: Diaulula sandiegensis (Cooper, 1863)] In the 1980 edition of Pacific Coast Nudibranchs, 1 referred to this species as Doris (s.1.) BEHRENS: PACIFIC COAST NUDIBRANCHS 4] species MacFarland (1966). In the 1991 edition, it was suggested that this species more properly belongs in the genus Diaulula. Behrens and Valdés (2001) examined the single specimen of Doris (s.l.) species referred to in MacFarland (1966) and other material collected from southern California that matched the external coloration of Doris (s./.) species. The internal anatomy was found to be identical to that of Diaulula sandiegensis. In that study, Doris odonoghuei Steinberg 1963 (= Doris echinata O’Donoghue, 1922) was also found to be a junior sysnomym of D. sandiegensis. 117. Discodoris sp. 1 [now: Diaulula aurila (Marcus and Marcus, 1967)] In their review of caryophyllidia-bearing dorids, Camacho-Garcia and Valdés (2003) placed Discodoris aurila in the genus Diaulula based on the presence of caryophyllidia. During their study, they were unaware of the specimen collected by T. Gosliner and shown in Behrens (1991). This specimen is identical of D. aurila (T. Gosliner, pers. commun. ). The range of this species is now extended north from Ft. Kobbe Beach, Canal Zone, Panama (Camacho-Garcia), to Punta Rosarito, Baja California, Mexico (Goddard and Schickel 2000). 119. Geitodoris heathi (MacFarland, 1905) The range of this species is extended north to Prince William Sound (Goddard and Foster 2002). 120. Taringa aivica timia Marcus and Marcus, 1967 [now: Taringa aivica Marcus and Marcus, 1967) The subspecies 7: a. timia appears to be a color morph of the nominal species 7? aivica, so this subspecific designation is dropped (Camacho-Garcia and Valdés 2003). The range is extended to Puerta Vallarta, Mexico (Hermosillo-Gonzalez 2003). Hermosillo-Gonzalez (2003) reports that the color varies widely from that reported in Behrens (1991) (see Plate 4A). 123. Thordisa rubescens Behrens and Henderson, 1981 Valdés (2002) provides a much more detailed description of the internal anatomy of this species than given by Behrens and Henderson (1981), adding information on the central nervous and reproductive systems, as well as SEMs of the radula, dorsal papillae and penial armature. The range of this species has been greatly extended to the south with the collection of a6 mm specimen at Punta Eugenia, Baja California by Hans Bertsch (pers. commun.). 124. Peltodoris sp. 1 [now: Peltodoris mullineri Millen and Bertsch, 2000] The descriptive material in the 1991 edition of Pacific Coast Nudibranchs is unchanged, but we now have the radular formula which is 23-24 x 42-63.0.42—63. Sandra Millen and Hans Bertsch named this species to honor their friend Dave Mulliner of San Diego, California. The range of this species is extended north to Santa Barbara and Anacapa Islands (Marc Shargel, pers. commun.), and south to Isla de Malpelo, Columbia (Kaiser and Bryce 2001). It has also been found in the Gulf of California at Bahia de los Angeles (Hans Bertsch and Allen Grant, pers. commun.). 42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 126. Platydoris macfarlandi Hanna, 1951 This rare species had only been collected once since its description in 1951 (Behrens and Henderson 1983). With the collection of two additional specimens, the range is extended south to Bahia San Cristobal, Baja California (Bertsch et al. 2000). 127. Dendrodoris fulva (MacFarland, 1905) [now: Doriopsilla albopunctata (Cooper, 1863)] After years of fielding questions concerning the identification of this species and difficulty dis- tinguishing Dendrodoris fulva from Doriopsilla albopuntata, a careful study of the internal anato- my, has shown the two to be synonymous (Gosliner, Schaefer, and Millen 1999). D. albopunctata, being the older of the two names, takes precedence. 128. Dendrodoris krebsii (Morch, 1863) [now: Dendrodoris fumata Riippell and Leuckart, 1830] The Caribbean and Eastern Pacific species referred to as Dendrodoris krebsii are actually dis- tinct species (Valdés, Ortea, and Ballestros 1996). Brodie, Willan and Collins (1997) refer to color photographs in Valdés et al. (1996) labeled as D. nigra (Stimpson, 1855), that the Pacific coast species should be referred to as D. fumata Riippell and Leuckart, 1830. Gosliner (1991) and Valdés et al. (1996) report this species from the Galapagos Islands, and Kaiser and Bryce (2001) at Isla de Malpelo, Columbia. 129. Dendrodoris nigromaculata (Cockerell in Cockerell and Eliot, 1905) This species is reported south to Costa Rica (INBIO 2003). 130. Dendrodoris sp. 1 [now: Doriopsilla gemela Gosliner, Schaefer, and Millen, 1999] The photograph of species #130 Dendrodoris sp. 1 was taken of an intertidal specimen collect- ed in San Luis Obispo, California. Since the publication of Behrens (1991), several “yellow gilled” porostomes have been discovered along this coast. Doriopsilla gemela, is the species shown in photograph #130, but is thought to be distinct from an identical looking deep water species found in La Jolla Canyon (Jim Lance, pers. commun.). IDENTIFICATION.— Identical in color to Doriopsilla albopunctata except that the branchial plume is orange. The body and gills are bright yellow to orange or orange-brown. The minute tubercles on the surface of the notum have small white specks. RADULA.— None NATURAL History.— Found intertidally (Bertsch 2002). The egg mass 1s distinctive and dif- fers from that of D. ablopunctata. The coil consists of about three whorls of yellow eggs which lie flat on the substrate, rather than on edge, as in D. ablopunctata. Bertsch (2002), in some detail, also describes its ecology and natural history in Bahia de los Angeles. S1zE.— To 40 mm in length. RANGE.— Gulf of California and along the Pacific coast of North America from Bahia Tortugas, Baja California Sur, Mexico, to Elkhorn Slough, Monterey County, California. ETYMOLOGy.— The name gemela comes from the Spanish for twin, as the species is so close in appearance to D. albopunctata. BEHRENS: PACIFIC COAST NUDIBRANCHS 43 131. Dendrodoris sp. 2 Two additional sightings of this still undescribed species have been made in 150 and 170 feet of water in La Jolla Canyon (George Spalding, pers. commun.). Additionally, a specimen was pho- tographed by Clinton Bauder in Carmel Bay, at 180 ft. on July 12, 2000 (Plate 4B). The specimen measured 30 mm in length. Bauder’s photograph is a much better representation of the living ani- mal than the moribund specimen figured in Behrens (1991). This species remains undescribed. 132. Dendrodoris sp. 3 This yet undescribed porostome species (currently under study by Sandra Millen and Hans Bertsch) is becoming increasingly more common along the mainland coast where it has been col- lected south to Bird Rock, La Jolla, California (Jim Lance, pers. commun.). Subtidal observations have also been made at Quast Rock at 60 feet (George Spalding, pers. commun.). Suborder Dendronotacea (= Dendronotina) 136. Tritonia diomedea Bergh, 1894 An unpublished doctoral thesis by James Murray (1966) presents fascinating new information on Tritonia diomedea. The following notes are extracted from from Murray (1966): Tritonia diomedea feeds on pennatulacean cnidarians — the sea pen Ptilosarcus gurneyi (unpublished), and the sea whips Virgularia (McDonald and Nybakken 1978), Stylatula elongata (Willows, pers. commun.) in Washington and British Columbian waters, and Acanthoptilum in Californian waters as deep as 200 m (Rim Fay, pers. commun.). 7: diomedea will also feed on the sea pansy, Renilla kéllikeri (G. Brown, R. Longley, pers. commun.), in the laboratory, although it is unknown if they do so in nature. It appears that geographically isolated groups of Tritonia will eat only one prey species, and this prey species varies among groups. Virgularia beds tend to be found in silty mud at greater depths (>25 m) than the sandy beds of Ptilosarcus (<20 m). Tritonia diomedea found among Virgularia tend to be smaller (0.5—20 cm) than those found among Ptilosarcus (8-30 cm). Also, those found among Ptilosarcus tend to be orange (like Ptilosarcus), with distinct tubercles on the dorsum, whereas those found among Virgularia tend to be redder in hue, from brick red to pale pink, and have a smoother dorsum (MacFarland 1966 for color illustration). There also appears to be variation in the number of embryos per egg mass cap- sule among geographically separated groups that is not related to prey items (C.E. Lee, unpub- lished). Tritonia diomedea lives among beds of these soft corals on flat or gently sloping soft bottoms of sand or silt. The silty surface of Virgularia beds (~30 m deep) is softer than the sandy surface of Ptilosarcus beds (4-18 m deep), presumably due to lower water flow rates in the deeper Virgularia beds that allow the finer silt to settle out of suspension. This correlates with the observation that 7. diomedea found among Virgularia can orient to lower flow rates than those found among Ptilosarcus (unpublished). 7: diomedea can grow to 30 cm in length and to a mass of 1700 g. 137. Tritonia festiva (Stearns, 1873) James Murray also reports that during the studies described above on Tritonia diomedea, he collected T. festiva up to 10 cm in length near Tofino, British Columbia. 44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 138. Tritonia myrakeenae Bertsch and Mozqueira, 1986 This species has been collected as far south as Costa Rica (INBIO, 2003). 139. Hancockia californica Macfarland, 1923 The range of this species is extended to Fort Bragg, California (C. Schooley and M. Ellen Hill, pers. commun.), inside the Gulf of California at Bahia de los Angeles (Hans Bertsch and Jim Lance, pers. commun.), and south to Puerto Vallarta (Hermosillo-Gonzalez 2003) and Costa Rica (INBIO 2003). 145. Dendronotus iris (Cooper, 1863) The range of this species is extended to Cabo San Lucas, Baja California (Jim Lance, pers. commun. ). The color of this species varies widely. Behrens (1980) described three distinct morphs, but later (Behrens 1991) he mentions only the red and white morphs. The color of this species varies tremendously from white through grey, orange and red. The color of the gill tufts is also variable — white, orange or purple, depending on the overall body color. The most distinctive character is the white line along the edge of the foot. This characteristic is found in every specimen. 148. Dendronotus sp. 1 La Jolla Shores, San Diego County can be added to locations at which this undescribed species has been photographed (Steve Gardner, pers. commun.). Also, see the description of a new Dendronotus in “Newly Discovered Species” section. 149. Doto amyra Marcus, 1961 Goddard (1996) provides an excellent study of the larval development of this species. In this paper, he points out that D. amyra, as currently accepted, may include at least two and possibly four distinct species. The following 1s a brief review of the distinctions Goddard reports for the four forms: Doto amyra Marcus, 1961 — Feeds on the hydroids Abietinaria sp., Garveia sp., Coryne sp., and Sertularia furcata and reaches a length of 14 mm. The larvae are lecithotrophic. The body and head are translucent white and occasionally have an irregular scattering of fine-grain subcutaneous brown to black pigment. The five to eight pairs of cerata have pale yellow to light pinkish-orange or orange-brown colored cores. The cerata possess four to seven rings of tuberacles and lack dark pigment. The rhinophores are translucent and contain opaque white grains. Doto form A — Feeds on small athecate hydroids and measures up to 7 mm. This form is sim- ilar in color to the previous, but with no dark pigment. It has five to six pairs of cerata, usually with orange to pinkish cores. Because of the presence of semi-translucent white glands in the tuberacles, they appear whiter that in the previous form. The larvae are planktotrophic. Doto form B — This form occurs on Aglaophenia. It differs from the above two in having dark pigment on the cerata, which are smooth or bear only low tuberacles. Specimens lack the opaque white grains in the rhinophores. It is similar to D. columbiana O’ Donoghue, 1921. Doto form C (from La Jolla, California) — Similar to Doto form A, above, it has planktotroph- ic larvae and feeds on short, unidentified hydroids. Hermosillo-Gonzalez (2003) extends the range of this species to Puerto Vallarta, Mexico. BEHRENS: PACIFIC COAST NUDIBRANCHS 45 152. Doto lancei Marcus and Marcus, 1967 The range of this species is extended northward from Mission Bay, California to Monterey Bay (Lovell and Libby Langstroth, pers. commun.). Lovell and Libby Langstroth’s specimen was dark in color, similar to the species referred to as Doto columbiana in Behrens (1980, Species #92). The range of this species is extended south to Puerto Vallarta, Mexico Hermosillo-Gonzalez (2003) and Costa Rica (INBIO, 2003). Suborder Arminacea (= Arminina) 156. Armina californica (Cooper, 1863) The range of this species is extended to the Aleutian Akutan Islands (Goddard and Foster 2002). 158. Dirona aurantia Hurst, 1966 [now: Dirona pellucia Volodchenko, 1941] Dirona aurantia was reported to be a junior synonym of D. pellucida (Martynov 1997). The species complete range now would include Charleston, Oregon (Jeff Goddard, pers. commun.) to Norton Sound, Alaska, across the Bering Sea and the Sea of Japan to Russia (Martynov 1997). It is found to 60 m. In Oregon, the individuals were found on the bryozoan Bugula pacifica, living on floating boat docks (Jeff Goddard, pers. commun. ). 159. Dirona picta MacFarland in Cockerell and Eliot, 1905 Previously known only to Charleston, Oregon, this species has now been collected on the south side of Cape Mearnes, Tillamook County, OR (Jeff Goddard, pers. commun.). 160. Janolus barbarensis (Cooper, 1863) The range of this species is extended south to Puerta Vallarta, Mexico (Hermosillo-Gonzalez 2003). 161. Janolus fuscus (O’ Donoghue, 1924) The range of this species is extended north to Homer, Alaska (Goddard and Foster 2002). It has also been documented from Sado island, Sea of Japan and Onagawa and Hokkaido, Pacific Ocean side in Japan (Nishina Masayoshi, pers. commun.). Suborder Aeolidacea (= Aeolidina) 163. Chlamylla sp. 1 {now: Flabellina sp. and/or Flabellina intermedia (Bergh 1899)]| Currently, there is some confusion concerning whether we have one or two new species of Chlamylla in the 1991 edition of Pacific Coast Nudibranchs. On review, the species in the photo- graph probably belongs to the genus Flabellina (S. Millen, pers. commun.). The specimen shown in the photograph as species #163 of the 1991 edition was collected in the Strait of George, British Columbia, was 9 mm in length, and was found on a solitary Tubularia sp. This specimen is considered to be an undescribed Flabellina (S. Millen, pers. commun.). The rhinophores can be extended the length of the body, although they are contracted in the 1991 photo. The ground color is transparent grey with pale peach suffused on the dorsum. Lanceolate cerata are 46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 2 | continuous and irregularly arranged on notal flange. Ceratal cores are peach-pink with white © cnidosacs, inside a clear sheath. The rhinophores, and long, cylindrical oral tentacles are dusted with opaque white at the ends. The other specimens referred to in the 1991 edition are nearly identical to this photo but have been collected in deep water trawl nets off the coast Oregon by Goddard (1990), and in Alaska. These specimens are yellow in color and are considered to be Flabellina intermedia (Bergh 1899) (Alan Kuzirian, pers. commun.). 165. Flabellina fusca (O’ Donoghue, 1921) [now: Flabellina trophina (Bergh, 1894)] Flabellina fusca (O’Donoghue, 1921) is a junior synomyn to Flabellina trophina (Bergh, 1894) (Roginskaya 1990). Flabellina trophina is often confused with Flabellina verrucosa, both species having reddish ceratal cores. The primary distinguishing characters are that F. triopina has a pointed snout, which is often upturned, and the cerata are continuous on a flange rather than in clusters inserted direct- ly on the notum. Plate 4C shows a dark color morph of the species from Alaska. In a message to the Sea Slug Forum from Irina Roginskaya, it appears that this species has a wide diet feeding on other aeolid nudibranchs as well as polychaete worms. 166. Flabellina iodinea (Cooper, 1862) The range of this species is extended to San Benitos Islands, Baja California, Mexico (Hermosillo-Gonzalez 2003). 167. Flabellina marcusorum Gosliner and Kuzirian, 1990 This species is known from the Pacific and Atlantic Oceans. Its range along this coast has been extended south to the Gulf of Chiriqui, Panama and to Islas Galapagos, Ecuador (Gosliner 1994), and along the Atlantic Coast, it ranges north to Venezuela and Curacao in the Caribbean and to Santa Marta Bay, Columbia, and to Brazil (P. Montoya, pers. commun.; Humann and DeLoach 2002; Sea Slug Forum). 169. Flabellina salmonacea (Couthouy, 1838) [now: Flabellina japonica (Volodchenko, 1941)] According to Sandra Millen (pers. commun.), Pacific Coast specimens differ from the Atlantic Coast sibling species, F: salmonacea, by their yellow cerata and penial morphology; they are actu- ally Flabellina japonica. Thus, the range should be changed to Sea of Japan and British Columbia, Canada. The species was named for the Japan Sea. 172. Flabellina sp. 1 It has been suggested that this species might be a Cumanotus due to its wide body and basal- ly joined rhinophores (S. Millen, pers. commun.). The photo in the 1991 edition of Pacific Coast Nudibranchs is of a specimen collected at Mission Bay, California. The reference to the British Columbia specimens was likely to a similar looking Cumanotus. This species has a unique swimming style, where the cerata move in unison, forward and back- wards in a rowing manner (Neil McDaniel, pers. commun.). BEHRENS: PACIFIC COAST NUDIBRANCHS AT 173. Flabellina sp. 2 [now: Flabellina vansyoci Gosliner, 1994] This colorful aeolid from the southern portion of this coast has been recently described by Terry Gosliner (Gosliner 1994), and given the name Flabellina vansyoci; the name honors California Academy of Sciences’ biologist Robert Van Syoc. The radular formula for the species is 36 x 1.1.1. The rachidian tooth bears 5—6 triangular denticles as shown here (Fig. 15). The margin teeth bear 15—20 denticles. The size of the species ranges from 15—30mm. Originally, the species was documented to occur at Magdelena Bay, Baja California and Islas Ladrones, Panama. Bertsch et al. (2000) extended the range northward to Punta Eugenia, Baja California. The species has also been observed further south, at La Paz, Baja California Sur (Angulo-Campillo 2002) and Puerto Vallarta (Hermosillo-Gonzalez 2003), Mexico, and along the west coast of Costa Rica (INBIO 2003). Ficure 15. Flabellina vansyoci Gosliner, 1994. Radular teeth. 174. Flabellina sp. 3 [now: Flabellina amabilis Hirano and Kuzirian, 1991] This species is Flabellina amabilis from northern Sea of Japan (Sandra Millen, pers. com- mun.). Its radula is triseriate, with a formula of 13-17 x 1.1.1. The rachidian teeth have 4—8 denti- cles and the lateral teeth 5—8 denticles. 175. Babakina festiva (Roller, 1972) Redfern (2001) documents the presense of Babakina festiva in the tropical Atlantic, in the Bahamas. 178. Eubranchus olivaceus (O’ Donoghue, 1921) [now: Eubranchus rupium (Moller, 1842)] Just and Edwards (1985) and Martynov (1998) indicated that the northeast Pacific Eubranchus olivaceus is a synonym of the north Atlantic Eubranchus rupium. The range of this species on this coast is thus extended to Amalga Harbor, north of Juneau, Alaska. It has been observed feeding on the hydroid species Obelia (Rita O’Clair, pers. commun.). More recently, the range has been extended to Homer, Alaska (Goddard and Foster 2002). 181. Eubranchus steinbecki Behrens, 1984 The range of this species is extended to La Paz, Baja California Sur, Mexico (Angulo- Campillo 2002). 184. Catriona rickettsi Behrens, 1984 Previously known only from San Francisco Bay and La Jolla, California, Jeff Goddard reports large numbers of this species were found on large patches of Tubularia sp. on the ocean side of the south jetty, Umpqua River, Douglas County, Oregon, during February 1994, 1996 and 1997. Also, Bertsch et al. (2000) extended the range south to Punta Eugenia, Baja California, Mexico. 48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 [Note concerning Cuthona: Martynov (2002) recommends a number of taxonomic changes for Pacific coast Cuthona returning several species to the genus Trinchesia. I cannot support these recommendations and do not follow them here. ] 186. Cuthona albocrusta (MacFarland, 1996) The range of this species is extended north to Cordova Marina, Prince Williams Sound, Alaska, and south to Bird Rock, La Jolla, California (Goddard 2000a), Bahia Tortugas (Bertsch et al. 2000) and La Paz, Baja California Sur, Mexico (Angulo-Campillo 2002) 187. Cuthona cocoachroma Williams and Gosliner, 1979 The northern limit of this species is extended from Oregon to Kayostla Beach, Olympic Peninsula, Washington (Goddard et al. 1997). 190. Cuthona flavovulta (MacFarland, 1996) The northern limit of this species is extended from Oregon to Kayostla Beach, Olympic Peninsula, Washington (Goddard et al. 1997). 191. Cuthona fulgens (MacFarland, 1996) The northern limit of this species is extended from Oregon to Kayostla Beach, Olympic Peninsula, Washington (Goddard et al. 1997). 193. Cuthona lagunae (O’ Donoghue, 1926) The range of this species is extended north to Humbug Mountain, Curry County, Oregon (Jeff Goddard, pers. commun.), and south to Bahia Tortugas, Baja California (Bertsch et al. 2000). 194. Cuthona perca (Marcus, 1958) Perrone (1995) reports the occurrence of this species in the Mediterranean, and he discusses taxonomic variation between several of the populations of this global species. 195. Cuthona phoenix Gosliner, 1981 This species has now been reported from Costa Rica (INBIO 2003). 197. Cuthona pustulata (Alder and Hancock, 1854) The range of this species is extended northward to Homer, Alaska (Goddard and Foster 2002) where it was found feeding on the hydroid, Sarsia sp. 198 Cuthona rolleri Behrens and Gosliner, 1988 A recent deepwater study conducted in the Santa Maria Basin in Central California collected a specimen of C. rolleri at a depth of 575 m. This is the first subtidal collection of this species, which previously was known only from intertidal collections, and it extends the range south to Point Sal, San Luis Obispo County, California (Gosliner 1996). i) j — BEHRENS: PACIFIC COAST NUDIBRANCHS 49 200. Cuthona viridis (Forbes, 1830) The range of this species is extended south to Bremerton, Washington (Andy Lamb, pers. com- mun.). Roginskaya (2000) reports that in summer C. viridis 1s the most common nudibranch in the intertidal zone of the Murman coast of the Barents Sea. Roginskaya also gives a detailed account of this species’ reproductive biology and development. 203. Fiona pinnata (Eschscholtz 1831) Schrédl (2003) reports this species from the southern hemisphere along the coast of Chile. 204. Anetarca armata Gosliner, 1991 Since the description of the species in 1991, it has been determined that it occurs on both coasts of Baja California. Farmer (1990) describes studies conducted by Jim Lance in Bahia de los Angeles, Gulf of California, Mexico that report a fascinating association of this nudibranch 1.e., where it lives, feeds, and lays its eggs, on a symbiotic hydroid species (Plate 4D), which occurs solely on the shelled gastropod, Decipifus californicus (Plate 4E). Reported also from Puerto Vallarta, Mexico (Hermosillo-Gonzalez 2003) and Costa Rica (INBIO 2003). 206. Facelina stearnsi Cockerell, 1901 Specimens have been recently collected at a depth of 110 ft. in La Jolla Canyon, San Diego, California (Peter Brueggeman and George Spalding, pers. commun.). The range of the species is extended southward to La Paz, Baja California where numerous specimens have been observed on an offshore shipwreck (Bob Sinclair, pers. commun., and Angulo-Campillo 2002). 208. Hermosita sangria Gosliner and Behrens, 1986 This species has been reported from Costa Rica (INBIO, 2003). 209. Noumeaella rubrofasciata Gosliner, 1991 The range is extended to Puerto Vallarta (Hermosillo-Gonzales 2003) and Guanacaste, Costa Rica (Peter Ajtai, pers. commun.). 210. Phidiana hiltoni (O’ Donoghue, 1927) The range of this species is extended northward to Duxbury Reef, Marin County, California (T. Gosliner, pers. commun.). 212. Aeolidia papillosa (Linnaeus, 1761) Goddard and Schickel (2000) extend the geographic range of this cosmopolitan species along the Pacific Coast south to Punta Rosarito, Baja California. Schrédl (1997) reports a variation of Aeolidia papillosa, Aeolidia papillosa var. serotina from Bahia de Coliuma, Chile and in the Falkland Islands and Argentina (Schrédl 2003). 213. Aeolidiella chromosoma (Cockerell and Eliot, 1905) Along the West Coast, the range of this species is extended to Costa Rica (INBIO 2003); in 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 the eastern Pacific it has recently been reported from the Izu Peninsula and Sagami Bay, Japan (Jun Imanoto and Nishina Masayoshi, pers. commun.). 214. Aeolidiella indica Bergh, 1888 [now: Anteaeolidiella indica (Bergh, 1888)] Aeolidiella indica 1s placed in a new genus, Anteaeolidiella. Miller (2001) reviewed the aeoid nudibranchs of New Zealand. His review included the report of A. indica from that region. Because of internal anatomical features found in A. indica, which separated it from other species in Aeolidiella, he created a new genus Anteaeolidiella for it. The range for this species should include the Galapagos Islands, Ecuador (Hickman and Finet 1999): 216. Cerberilla mosslandica McDonald and Nybakken, 1975 Jeff Goddard (pers. commun.) reports a single specimen matching the description of C. moss- landica from 65 m of water, 6.4 km west of Bandon State Beach, Oregon. 217. Cerberilla sp. 1 [now: Cerberilla cf. pungoarena Collier and Farmer, 1964] This species has become common in the Channel Islands. (Plate 4F) (Kathy deWet, pers. com- mun.). This speciesis considered to be Cerberilla pungoarena Collier and Farmer, 1964, from Puerto Refugio, Isla Angel de la Guardia, Gulf of California, Mexico. REFERENCES ABBOTT, R.T. 1974. American Seashells: the Marine Mollusca of the Atlantic and Pacific Coasts of North America, 2nd ed. Van Nostrand Reinhold Co., New York. 663pp., 24pls. ANDERSON, E.S. 1971. The Association of the Nudibranch Rostanga pulchra MacFarland 1905 with the Sponges Ophilitaspongia pennata, Esperiopsis originalis, and Plocamia karykina. Ph.D. Dissertation. University of California, Santa Cruz, Santa Cruz, California. ANGULO-CAMPILLO, OrSO. 2002. New distributional records of Opisthobranch molluscs from the Gulf of California, Mexico. The Festivus 34(10):117—121. ANGULO-CAMPILLO, ORSO, J. ARREOLA ROBLES, AND HANS BERTSCH. 2001. Registro de talla récord para Pleurobranchus areolatum (Morch, 1863) (Opisthobranchia: Pleurobranchidae) en el Golfo de California. Océanides 16(1):91—92. BEHRENS, DAvip W. 1980. Pacific Coast Nudibranchs: A Guide to the Opisthobranchs of the Northeastern Pacific. Sea Challengers, Los Osos, California. 112pp. BEHRENS, David W. 1991. Pacific Coast Nudibranchs: A Guide to the Opisthobranchs Alaska to Baja California, 2nd ed.. Sea Challengers. Monterey, California. 107pp. BEHRENS, DAvip W., and Robert Henderson. 1981. Two new cryptobranch dorid nudibranchs from California. The Veliger 24:!70-128. BEHRENS, Davip W. AND ANGEL VALDES. 2001. The identity of Doris (s.l.) species MacFarland, 1966 (Mollusca, Nud.branchia, Discodorididae): A persistent mystery from California waters. Proceedings of the California Academy of Sciences 52(15):183-193. BERGH, RUDOLPH. 1878. Malacologische Untersuchungen. Pages 603-645 in C. Semper, ed., Reisem im Archipel der Philippinen, Theil 2 Heft 14. Kreidel, Wiesbaden, Germany. BERGH, RUDOLPH. 1894. Die Opisthobranchien. Reports on the dredging operations off the West Coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer “Albatross,” during 1891, Lieut. Commander Z.L. Tanner, U.S.N. commanding. Bulletin of the Museum of Comparative Zoology, Harvard 25(10):125-235. BERTSCH, HANS. 2002. The natural history of Doriopsilla gemela Gosliner, Schaefer, and Millen, 1999 | BEHRENS: PACIFIC COAST NUDIBRANCHS 51 (Opisthobranchia: Nudibranchia) at Bahia de los Angeles, Baja California, Mexico. Annual Report, Western Society of Malacologists 33:7-8. BERTSCH, HANS., ORGO ANGULO CAMPILLO, AND JOSE Luis. ARREOLA. 2000. New distributional records of opisthobranchs from the Punta Eugenia region of the Baja California peninsula, a record based on 1997-1998 CONABIO-sponsored expeditions. The Festivus 32(7):99-104. BRODIE, GILIANNE D., RICHARD C. WILLAN, AND JOHN D. COLLINS. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia: Dendrodorididae) in the Indo-Pacific Region. Journal of Molluscan Studies 63:407-423. CADIEN, DONALD. 1998. Good things/ Small package. SCAMIT Newsletter 17(3):4—S. CaMACHO-GaRCiA, YOLANDA E., AND ANGEL VALDES. 2003. Caryophyllidia-bearing dorid nudibranchs (Mollusca: Nudibranchia: Doridina) from Costa Rica. Proceedings of the California Academy of Sciences 54(4):65-79. CARLTON, J.T. 1979. History, Biogeography and Ecology of the Introduced Marine and Estuarine Inverte- brates of the Pacific Coast of North America. Ph.D. Dissertation. Unversity of California, Davis. 904pp. Cura, F-S, AND M. SKEEL. 1973. The effect of food consumption on growth, fecundity and mortality in a sacoglossan opithobranch, Olea hansineensis. The Veliger 16:153-158. FARMER, WESLEY M. 1978. Tambja and Roboastra (Mollusca: Opisthobranchia) from the Gulf of California and the Galapagos Islands. The Veliger 20(4):375-—385. FARMER, WESLEY M. 1990a. A strange occurrence at Bahia de Los Angeles: Egg laying on Nassarius and Decipifus. The Festivus 22(8):85-—95. FARMER, WESLEY M. 1990b. Platydoris macfarlandi, Hanna 1951 from Bahia de Sebastian Viscano and Isla Navidad, Baja California, Mexico. The Veliger 26(4):48. FISCHER, MARIA ANGELICA, AND JESUS ORTEA. 1996. New records of the genus Ancula Lovén, 1846 (Nudibranchia: Goniodorididae) on the American Pacific Coast. The Veliger 39(1):90-92. GEIGER, HEATHER, AND ALAN R. HOLyoak. 1996. Preference of adults of the dorid nudibranchs Archidoris montereyensis (Cooper, 1862), Diaulula sandiegensis (Cooper, 1862) and Triopha catalinae (Cooper, 1863) for shaded over lighted conditions. The Veliger 39(1):95-97. GODDARD, J.H.R. 1978. Observations on the opisthobranch mollusks of Punta Gorda, California, with notes on the distribution and biology of Crimora conejo. The Veliger 29:267-273. GODDARD, J.H.R. 1996. Lecithotrophic development in Doto amyra (Nudibranch: Dendronotacea), with a review of developmental modes in the genus. The Veliger 39(1):43-—54. GopDARD, J.H.R. 2000a. Northern and southern range extensions of the aeolid nudibranch, Cuthona albocrus- ta. Opisthobranch Newsletter 26(2):9. GODDARD, J.H.R. 2000b. Sponge prey of Chromodoris macfarlandi. Opisthobranch Newsletter 26(2):9. GODDARD, J.H.R, and N.R. Foster. 2002. Range extensions of sacoglossan and nudibranch mollusks (Gastropoda: Opisthobranchia) to Alaska. The Veliger 45(4):331-336. GoppDarD, J.H.R, and E. Schickel. 2000. Range extension of six opisthobranchs to Punta Rosarito, Baja California. Opisthobranch Newsletter 26(5):21—22. GODDARD, J.H.R, T.A. Wayne and K.R. Wayne. 1997. Opisthobranch mollusks and the pulmonate limpet Trimusculus reticulatus (Sowerby, 1835) from the outer Washington coast. The Veliger 40(4):292—297. GOSLINER, TERRENCE M. 1979. A review of the systematics of Cylichnella Gabb (Opisthobranchia: Scaphandridae). Nautilus 93:85-92. GOSLINER, TERRENCE M. 1991. The Opisthobranch gastropod fauna of the Galapagos Islands. Chpt. 13 in Matthew James, ed., Galapagos Marine Invertebrates. Plenum, New York. 474 pp. GOSLINER, TERRENCE M. 1994. New records of Flabellinidae (Opisthobranchia: Aeolidacea) from the tropical Americas, with descriptions of two new species. Proceedings of the California Academy of Sciences 48(9):171-183. GOSLINER, TERRENCE M. 1995. Introduction and spread of Philine auriformis (Gastropoda: Opisthobranchia) from New Zealand to San Francisco Bay and Bodega Harbor. Marine Biology 122:249-255. GOSLINER, TERRENCE M. 1996. The Opisthobranchia. In P.H. Scott, J.A. Blake, and A.W. Lissner, eds., Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel.,. vol. 9. The Mollusca, Part 2. Santa Barbara Museum of Natural History, Santa Barbara, California. SY PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 GOSLINER, TERRENCE M. 2004. Phylogenetic systematics of Okenia, Sakishimaia, Hopkinsiella and Hopkinsia (Nudibranchia: Goniodorididae) with descriptions of new species from the Tropical Indo-Pacific. Proceedings of the California Academy of Sciences 55(5):125—161, 29 figs. GOSLINER, TERRENCE M., AND DaAviID W. BEHRENS. 1996. Two new species of nudibranch mollusks from the Gulf of Farallones and Cordell Bank National Marine Sanctuaries, central California. The Veliger 39(4):348-353. GOSLINER, TERRENCE M., AND A.M. KUuZIRIAN. 1990. Two new species of Flabellinidae (Opisthobranchia: Aeolidacea) from Baja California. Proceedings of the California Academy of Sciences 47:1-15. GOSLINER, TERRENCE M., MARIA C. SCHAEFER, AND SANDRA V. MILLEN. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodorididae) from the Pacific coast of North Ameica, including a comparison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42(3):201-210. HERMOSILLO-GONZALEZ, ALI. 2003. New distributional records (Mollusca: Opisthobranchia) for Bahia de Banderas, Mexico (Eastern Pacific). The Festivus 35(3):21-28. HERNANDEZ C., MARCO ANTONIO. 1992. Inventario Preliminario de Moluscas Marinas de la Costa de El Salvador. Trabajo de investigacion presentado para secretaria ejecutiva del medio ambiente SEMA. 172 PPp- HICKMAN, CLEVELAND P., AND YVES FINET. 1999. A Fieldguide to the Marine Mollusks of Galapagos. An illus- trated Guidebook to the Common Intertidal and Shallow-water Snails, Bivalves and Chitons of the Galapagos Islands. Sugar Spring Press, Lexington, Virginia. 150 pp. HUMANN, P. AND N. DELOACH. 2002. Reef Creature Identification — Florida, Caribbean Bahamas, 2nd ed. New World Publications: Jacksonville, Florida. 420 pp. INBIO. 2003. Sea Slugs of the Pacific coast of Costa Rica. www.inbio.ac.cr/papers/babosasmarinas/ingels/ index.html. JAECKLE, W.B.1983. Range extensions for two California cuthoniids (Gastropoda: Nudibranchia) from Santa Cruz Island. Opisthobranch Newsletter 15(11—12):53. JENSEN, KATHE. 2001. Aplysiopsis oliviae from British Columbia. Sea Slug Forum. http://www.seaslugforum/aplyoliv.htm#m2521. 7 November 2001. Just H., AND M. EpMunDs. 1985. North Atlantic nudibranchs (Mollusca) seen by Henning Lemche. Ophelia (Suppl.) 2:1—150. KAISER, KIRSTIE L,. AND CLAYTON W. Bryce. 2001. The recent molluscan marine fauna of Isla de Malpelo, Colombia. The Festivus (Occasional Paper 1). 147 pp. Krua, P.J. 1998. Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutch- es in a population of the sacoglossan Alderia modesta. Marine Biology 132(3):483-494. LONHART, STEVE I., AND JEFF W. TUPEN. 2001. New range records of 12 marine invertebrates: the role of El Nino and other mechanisms in southern and central California. Bulletin of the Southern California Academy of Sciences 100(3):238—248. MACFARLAND, FRANK Mace. 1905. A preliminary account of the Dorididae of Monterey Bay, California. Proceedings of the Biological Society of Washington 18:35—54. Marcus, E. 1961. Opisthobranch mollusks of California. The Veliger 3(suppl. 1):1-85. McDonaLp, G.R., AND J.W. NYBAKKEN. 1997. A list of worldwide food habits of nudibranchs. University of California, Santa Cruz web site — http://veliger.org/nudibranch_food.html. McLEAN, JAMES, AND GENE COAN. 1996. Marine mollusks of Rocas Alijo. Jn Robert W. Schmieder, ed., Rocas Alijos: Scientific Results from Cordell Expeditions. Kluwar Academic Publications, Boston. 481 pp. Martynoy, A.V. 1994. Materials for the revision of nudibranchiate molluscs of the family Corambidae (Gastropoda, Opisthobranchia) taxonomy. Communication 1. Zoologichesky Zhurnal 73(10):3-15. (In Russian.) Marrynoy, A.V. 1997. Opisthobranch mollusks of coastal waters of Commander Islands with notes on their fauna in the Far-East Seas of Russia. Pages 230-241 in A.V. Rzhavsky, ed., Donnaya Fauna I Flora Schelifa Komandorskikh Ostrovoy. DalOnauka, Vladivostok, Russia. (In Russian.) Martynov, A.V. 1998. Opisthobranch mollusks (Gastropoda: Opisthobranchia) of the family Eubranchidae: Taxonomy and two new species from the Sea of Japan. Zoologicheskii Zhurnal 77:763—777. (In Russian.) Marrtynov, A.V. 2002. Two new species of the genus Trinchesia Ihering, 1879 from Peter the Great Bay, it BEHRENS: PACIFIC COAST NUDIBRANCHS 53 Japan Sea (Nudibranchia, Tergipedidae), with notes on the taxonomy of the family. Ruthenia 12(1):45—54. (in Russian.) Martynov, A.V,. AND O.N. BARANETS. 2002. A revision of the genus Colga Bergh (Opisthobranchia, Polycer- idae), with description of a new species from the North Pacific. Ruthenica 12(1):23—43. (In Russian.) MIKKELSEN, P.M. 1996. The evolutionary relationaships of Cephalaspidea s./. (Gastropoda: Opisthobranchia): a phylogenetic analysis. Malacologia 37(2):375—-442. MILLEN, SANDRA V., AND HANS BERTSCH. 2002. Three new species of dorid nudibranchs from southern California, USA and the Baja California peninsula, Mexico. The Veliger 43(4):354—366. MILLEN, SANDRA V., AND JAMES NYBAKKEN. 1991. A new species of Corambe (Nudibranchia: Doridoidea) from the northeastern Pacific. Journal of Molluscan Studies 57(4):209-215. Miter, M.C. 2001. Aeolid nudibranchs (Gastropoda: Opisthobranchia) of the family Aeolidiidae from New Zealand waters. Journal of Natural History 35:629-662. Morton, B., AND CHIU S.T., 1990. The diet, prey size and consumption of Philine orientalis (Opisthobranchia: Philinidae) in Hong Kong. Journal of Molluscan Studies 56:289-299. Murray, JAMES ALAN. 1966. Neural Correlates of Orientation to Water-flow in Tritonia diomedea. Ph.D. Dissertation (42526). University of Washington, Seattle. ODHNER, N.H.J. 1937. Coryphella islandica n. sp., a new nudibranchiate mollusk from Iceland. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 101:253—257. OKUTANI. T., ed. 2000. Marine Mollusks of Japan. Tokai University Press. 320 pp. OrTEA, J.. AND E.M. LLeRA. 1981. Un nuevo doérido (Mollusca: Nudibranchia) de La Isla Isabel, Nayarit, Mexico. /berus 1:47-52. Percy, W.G., J. FISHER, R. BRODEUR, AND S. JOHNSON. 1985. Effects of the 1983 El Nifio on coastal nekton off Oregon and Washington. Pages 188—204 in W.S. Wooster and D.L. Fluharty, eds., El Nifio North: Nino Effects of the Eastern Subarctic Pacific Ocean. Washington Sea Grant Program. University of Washington, Seattle. 312 pp. PERRONE, ANTONIO S. 1995. Una specie di nudibranchi del genere Cuthona Alder and Hancock, 1855, nuova per il Mediterraneo: Cuthona perca (Marcus, 1958) (Opisthobranchia: Nudibranchia). Bollettino. Malacologico 31(1-4):28-36. (In Italian) PicTON, B. E., AND C.C. Morrow. 1994. A Field Guide to the Nudibranchs of the British Isles. Tmmel Publishing Ltd., London, UK. POORMAN, FoRREST L., AND LEROY H. POORMAN. 1988. A report of the molluscan species in the San Carlos rectangle, Sonora, Mexico, collected by Forrest L. and Leroy H. Poorman from December 1953 to December 1983. The Festivus 20(6):47-63. REDFERN. C. 2001. Bahamian Seashells: a Thousand Species from Abaco, Bahamas. Bahamian seashells.com, Inc., Boca Raton, Florida. Rivest, B.R. 1984. Copulation by hyperdermic injection in the nudibranchs Palio zosterae and P. dubia (Gastropoda, Opisthobranchia). Biological Bulletin 167(3):534-554. RoGINsKAYA, I.S. 1972. Calycidoris gentheri (Gastropoda, Nudibranchia). Taxonomy and distribution. Zoologicheskii Zhurnal 51:913—918. (In Russian.) ROGINSKAYA, LS. 1990. Data of feeding, distribution and synonymy of Coryphella trophina (Bergh, 1894) (Nudibrancia: Gastropoda). Pages 47-57, 151-152, and 156 in A.P. Kuznetsov, ed., Feeding and Bioener- getics of Marine Bottom Invertebrates. Academy of Sciences of the U.S.S.R., P.P. Shirshov Institute of Oceanology, Moscow. (In Russian.) RoaInsKAyA, I.S. 2000. Notes on Trinchesia viridis (Forbes, 1840) (Nudibranchia, Tergipedidae) and its veligers. Opisthobranch Newsletter 26(3):13—16. RUDMAN, W.B. 1995. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-Pacific: further species from New Caledonia and the Noumea romeri color group. Molluscan Research 16:1-43. SCAMIT. 2001. A taxonomic listing of soft bottom macro- and megainvertebrates from infaunal and epiben- thic monitoring programs in the Southern California Bight. Edition 4. Southern California Association of Marine Invertebrate Taxonomists (SCAMIT). SCHRODL, MICHAEL. 1997. Range extensions of Magellanic nudibranchs (Opisthobranchia) into the Peruvian faunal province. The Veliger 40(1):38—42. 54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 2 ! i) \ } U | SCHRODL, MICHAEL. 2003. Sea Slugs of Southern South America. Conchbooks, Hackenheim, Germany. 165 Pp: SCHRODL, MICHAEL, AND SANDRA MILLEN. 2001. Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidae: Chromodorididae). Journal of Natural History 35:1143-1171. SHRAKE, J. 1991. Occurrence of a rather unusual opisthobranch found off southern California. Opisthobranch | Newsletter 23(5):18. SKOGLUND, CAROL. 2002. Panamic Province molluscan literature. Additions and changes from 1971 through 2001. III. Gastropoda. The Festivus 33(Suppl). 296 pp. SOWELL, R.R. 1949. Taxonomy and Ecology of the Nudibranchiate Mollusca of the Coos Bay, Oregon Region. M.S. Thesis. Oregon State University, Corvallis. 54 pp. SWENNEN, C., AND R. DEKER. 1995. Corambe batava Kerbert, 1886 (Gastropoda: Opisthobranchia), an immi- grant in the Netherlands, with a revision of the family Corambidae. Journal of Molluscan Studies 61: 97-107. THOMPSON, T.E., AND G.H. BROWN. 1984. Biology of Opisthobranch Molluscs, vol. 2. The Ray Society, London. 229 pp. TROWBRIDGE, CYNTHIA D. 2002. Northeastern Pacific sacoglossan opisthobranchs: natural history, review, bib- liography and prospectus. The Veliger 45(1):1—24. VALDES, ANGEL. 2002. A phyllogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society 136:535—636. VALDES, ANGEL, AND DAVID W. BEHRENS. 1998. A new species of Doriopsilla (Mollusca, Nudibranchia, Dendrondorididae) from the Pacific coast of North America. Proceedings of the California Academy of Sciences 50(13):307—3 14. VALDES, ANGEL, AND HANS BertTscH. 2000. Redescription and range extension of Bathydoris aioca Marcus and Marcus 1962 (Nudibranchia: Gnathodoridoidea). The Veliger 43(2):172-178. VALDES, ANGEL, AND PHLLIPPE BOUCHET. 1998. A blind abyssal Corambidae (Mollusca, Nudibranchia) from the Norwegian Sea, with a re-evaluation of the systematics of the family. Sarsia 83:15—20. VALDES, ANGEL, AND TERRENCE M. GosLINER. 2001. Systematics and phylogeny of the caryophyllidia-bear- ing dorids (Mollusca, Nudibranchia), with the description of a new genus and four new species from Indo- Pacific deep waters. Zoological Journal of the Linnean Society 133:103-198. VALDES, ANGEL, 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. VALLES, YVONNE, ANGEL VALDES, AND JESUS ORTEA. 2000. On the phanerobranch dorids of Angola (Mollusca, Nudibranchia): a crossroads of temperate and tropical species. Zoosystema 22(1):15-31. WatTSON, R.B. 1886. Report on the Scaphopoda and Gastropoda collected by H.M.S. Challenger during the years 1873-1876. Scientific Results of the Voyage of the H.M.S. Challenger, vol. 15. Zoology, pt. 2. 680 PPp- Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3, pp. 55-88, 24 figs., 1 table. April 22, 2004 The Panamic Gorgonian Genus Pacifigorgia (Octocorallia: Gorgoniidae) in the Galapagos Archipelago, with Descriptions of Three New Species Gary C. Williams! and Odalisca Breedy? ! Department of Invertebrate Zoology and Geology, California Academy of Sciences Golden Gate Park, San Francisco, California 94118, U.S.A. Email: gwilliams @ calacademy.org; 2 Centro de Investigaci6én en Ciencias del Mar y Limnologia (CIMAR), Universidad de Costa Rica 2060, San José, Costa Rica The eastern Pacific gorgonian genus Pacifigorgia Bayer, 1951, is here described from the northwestern fringe of the Galapagos Archipelago. The two small islands of Darwin and Wolf are relatively isolated from the rest of the archipelago. Two of the new taxa are apparently endemic to these outer islands, as previous subtidal surveys of the other Galapagos Islands, as well as other regions of the Panamic Biogeographic Province, have not encountered conspecific material. The third new species is apparently restricted to central islands of the archipelago. The previously described P. darwini (Hickson, 1928) is the most common and widespread species and is found throughout the archipelago, except for the outlying islands of Darwin and Wolf. A taxonomic assessment of Hickson’s Gorgonia darwinii var. douglasii is also included. The genus is restricted to the eastern Pacific (Gulf of California to Peru), with the exception of Pacifigorgia elegans, endemic to the tropical western Atlantic (Trinidad to Brazil). The addition of the new taxa described here makes twenty-one species belonging to the genus that are considered valid. The marine fauna of oceanic islands have been subjects of great scientific interest chiefly because of their biogeographic relevance. Endemism, dispersal patterns, and other aspects of evo- lutionary biology might be better understood through comparisons to mainland faunas (Quammen 1996). The main oceanic islands and archipelagos situated in the tropical eastern Pacific are Isla del Coco (Costa Rica); Isla Malpelo (Columbia); Islas Revillagigedo (Mexico), Ile Clipperton (France); and Islas Galapagos (Ecuador). Even though numerous expeditions to explore the flora and fauna of these islands have been made, the octocorallian fauna remains as one of the least known faunal elements. The literature pertaining to the Galapagos octocoral fauna is depauperate. The main sources here include Hickson (1928), Bayer (1978), and Breedy and Guzman (2003). Sources relevant to the genus Pacifigorgia are Bayer (1951, 1953, 1956), Breedy (2001), and Breedy and Guzman (2002, 2003). Hickson (1921, 1928) and Deichmann (1941) provided the first records of shallow water octocorals of the Galapagos Archipelago. Hickson (1921) described Cavernularia darwinii (Pennatulacea) from Isla Cristobal (Chatham Island), and then seven years later, he described two new species of Pacifigorgia (under the generic name Gorgonia) (Hickson, 1928). Deichmann (1941) described the species Muricea galapagensis (Plexauridae) from Isla Isabela (Albemarle 38) 56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 Island), which was pulled up on an anchor chain from 53 fathoms (97 m depth). The validity of this species has to be corroborated. Aside from Hickson’s two species and the recently described Pacifigorgia curta Breedy and Guzman, 2003, from Isla del Coco, there are not published records from other oceanic islands of the tropical eastern Pacific. Two presently undetermined species of Pacifigorgia are known from Islas Revillagigedo. These are represented by two dried preserved specimens, CAS 097905 and CAS 097906. The only published records of the genus Pacifigorgia from the Galapagos, appeared in Hickson’s publication “The Gorgonacea of Panama Bay.” He described two taxa: Pacifigorgia dar- winii (Hickson, 1928), and Pacifigorgia darwinii var. douglasii (Hickson 1928). Pacifigorgia dar- winii was revised and redescribed by Breedy and Guzman (2002), whereas P. darwinii var. dou- glasii, was originally figured only from sclerites, which is all that is known to exist from the type specimen (Breedy and Guzman 2002). In this paper, we monograph all the known taxa from the Galapagos Archipelago, which includes the descriptions of three new species, based on the study of recently discovered taxa, and material found in several collections. The two previously described taxa are also included in the present paper, making a total of five species of the genus Pacifigorgia presently known to inhabit the shallow coastal waters of the Galapagos Archipelago. Four of these are considered valid, whereas one is here considered to be of dubious status. The three new species of sea fans (gorgonian corals) were discovered during SCUBA opera- tions conducted during the 1994 California Academy of Sciences marine expedition to the Galapagos Islands, Ecuador. McCosker (1994) briefly describes this expedition. The genus Pacifigorgia was described by Bayer (1951:94) for net-like reticulated species with sclerites as spindles and capstans (scaphoids absent), geographically confined to the Pacific coast of tropical America (with one exception from the tropical western Atlantic). The following genera are known to comprise the shallow-water gorgonian fauna (<50 m) of the Galapagos Archipelago. These five genera represent approximately eleven species in the two families Plexauridae and Gorgoniidae: Adelogorgia (1 species), Eugorgia (1 species), Leptogorgia (1 species), Muricea (ca. 4 species), and Pacifigorgia (4 valid species, plus | dubious species) (pers. observ. and Cleve Hickman — pers. commun.). METHODS Specimens were collected by the senior author (GCW) from several islands in the Galapagos Archipelago, between 2°N to 1.5°S Latitude, and 89°W to 92°W Longitude, during the California Academy of Sciences marine expedition to the Galapagos Islands in May of 1994. All material was collected by SCUBA in shallow water (<50 m depth), and preserved in 70% ethanol. The speci- mens for study came from this recent collection as well as from expeditions by C. Hickman and P. Humann (1993-2001). Other material came from the collections of the California Academy of Sciences, San Francisco; the Charles Darwin Research Station, Galapagos; the voucher collection of the Museum of Comparative Zoology of Harvard University, Boston, made available to us by Ardis Johnston; and type specimens and microscope slides from The Natural History Museum, London, through the kindness of Sheila Halsey. Most of this material was collected by hand using SCUBA or skin diving. Most of the specimens are preserved in 70-75% ethanol and were original- ly fixed in ethanol. A few of the specimens were preserved dried. All collection sites are indicated on the maps (Figs. 5, 9, 16, 23), along with a range comparison map (Fig. 24). Sclerites were isolated using sodium hypochlorite (household bleach); see the website: WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 57 hitp:/Avww.calacademy.org/research/izg/OctoResearchTech.him (Octocoral Research Center or Octocoral Home Page — Research Techniques), and Breedy and Guzman (2002:784) for details. Also of relevance here are Bayer (1961:23—24), and Fabricius and Alderslade (2001:38). The subject for Figure 1A was photographed in a small glass aquarium from freshly collected material, using a Pentax 35 mm camera, with a 50 mm macro lens and flash unit. Figure 1C is a scanprint made directly from the holotype, using a Umax Astdra 1200S flatbed scanner. Micrographs for figures 1B and 2 were made using a Nikon Coolpix 990 digital camera, a Nikon SMZ-10 dissecting microscope, and an Olympus CH-2 compound microscope. Scanning electron micrographs were made using a Leo 1400 Series or a N-2360 Hitachi scanning electron micro- scope. Sclerite drawings were made using an Olympus CH-2 compound microscope with an attached drawing tube. Digital images and plates of photographs, micrographs, and scanning elec- tron micrographs were made using Adobe Photoshop software. Abbreviations used in the text are BM or NHM (The Natural History Museum, London; formerly British Museum — Natural History); CAS (California Academy of Sciences, San Francisco); CDRS (Charles Darwin Research Station, Puerto Ayora, Isla Santa Cruz, Galapagos Islands); MCZ (Museum of Comparative Zoology, Harvard University, Boston). Specimens examined in this study are deposited in the CAS octocoral collection (Department of Invertebrate Zoology and Geology), the CDRS, and the MCZ. Type specimens were obtained by loan from the BM. Previous records of some species have been acquired from literature sources (Hickson, 1928; Stiasny, 1941, 1943; Breedy and Guzman, 2002), as well as from personal obser- vations and museum specimens. Terminology used in this paper conforms to that of Bayer, Grasshoff and Verseveldt (1983), and Breedy and Guzman (2002). SYSTEMATIC ACCOUNT Family Gorgoniidae Lamouroux, 1812 Pacifigorgia Bayer, 1951 Rhipidigorgia (partial) Valenciennes, 1855:13. Milne Edwards and Haime, 1857:173. Horn, 1860:233. Rhipidogorgia (partial) Verrill, 1864:32. Duchassaing and Michelotti, 1864:20. Verrill, 1869:424. Litigorgia (partial) + Eugorgia (partial) Verrill, 1868:414. Leptogorgia (partial) Verrill, 1869:420. Verrill, 1870:548. Gorgonia Bielschowsky, 1918:32. Kiikenthal, 1924:338. Bielschowsky, 1929:141. Stiasny, 1941:268. Stiasny, 1943:74. Pacifigorgia Bayer, 1951:94. 1953:103. 1956:212. Breedy, 2001:182. Breedy and Guzman, 2002:791. 2003:3. DIAGNOsIs.— Sea fans composed of reticulated networks by anastomosis (as in the western Atlantic genus Gorgonia), but with sclerites similar to those of the genus Leptogorgia from the eastern Pacific and Atlantic Ocean. The coenenchymal sclerites are girdled spindles and radiates, whereas scaphoids are absent. Color permanent, incorporated in the sclerites. TYPE SPECIES.— Gorgonia stenobrochis Valenciennes, 1846; subsequent designation by Bayer, 1951:94. DIVERSITY AND DISTRIBUTION (see Fig. 5, inset).— Twenty species from the Eastern Pacific (Baja California to Chile, Cocos Islands, Revillagigedo Islands and Galapagos Islands); one species in the western Atlantic (Trinidad and Venezuela to Brazil). 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 Pacifigorgia dampieri Williams and Breedy, sp. nov. (Figs. 1-5, 24) MATERIAL EXAMINED.— HOoLotyPeE: CAS 097040, Ecuador, Galapagos Islands, Isla Darwin, 15-21 m depth, (Station G8), 13 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral I’. One whole wet specimen, preserved in 75% ethanol. PARATYPE: CAS 097021, Ecuador, Galapagos Islands, Isla Wolf, 9-15 m depth, (Station G5), 12 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral II’. One whole wet specimen, preserved in 75% ethanol. OTHER MATERIAL: CAS 097022, Ecuador, Galapagos Islands, Isla Wolf, 12-18 m depth, (Station G4), 12 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral IT’. One whole wet specimen, preserved in 75% ethanol. CAS 097023, Ecuador, Galapagos Islands, Isla Wolf, 12-18 m depth, (Station G4), 12 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral II’. One whole wet specimen, preserved in 75% ethanol. CAS 097025, Ecuador, Galapagos Islands, Isla Wolf, 9-15 m depth, (Station G5), 12 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral I”. One whole wet specimen, preserved in 75% ethanol. CAS 097025, Ecuador, Galapagos Islands, Isla Wolf, 9-15 m depth (Station G5), 12 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral IP’. One whole wet specimen, preserved in 75% ethanol. CAS 147483, Ecuador, Galapagos Islands, Isla Darwin, 15—21 m depth, (Station G8), 13 May 1994, collected by GC. Williams with SCUBA, aboard “Mistral IP’. One whole wet specimen, preserved in 75% ethanol. CDRS 02-39, Ecuador, Galapagos Islands, Isla Wolf, depth not recorded, 19 May 2001, collected by C. Hickman. GENERAL DESCRIPTION.— Fans are relatively stiff, planar or comprised of a main fan with sev- eral secondary fans in different planes, which radiate outward from the central region of the main fan. Secondary fans may radiate perpendicularly from the main fan. Colonies are wider than high, often irregularly-shaped, up to 200 mm in height and 270 mm in width. Multiple fans are intercon- nected by relatively thick branches (up to 8 mm in width) that diverge upwards from the holdfast. In some cases, fans with separate holdfasts anastomose to produce a maze-like appearance. The fusion between fans may take place in the coenenchyme of the main stem or main branches. Colonies often have strong, encrusting, orange-colored holdfasts. In smaller colonies, the holdfast may spread out conspicuously over the substratum. Networks are intricate, regular, and of open meshes. The meshes are often more-or-less square, but are also variable: circular, rounded-rectangular, elliptical or triangular meshes 2—5 mm long by 1-3 mm wide; elongated and trapezoidal meshes 4-12 mm long by 0.7—6.0 mm wide. Colonies average approximately six meshes/cm?. Mesh branches are thin, up to 1.0 mm in diameter. Prominent, rounded midribs , up to 7 mm in width, extend up into the fans, or reach a short dis- tance upwards from the base of the fans. They are of a distinct orange color, with distinct longitu- dinal grooves of yellow and orange sclerites, very apparent in dried specimens. The terminal branchlets are short, 2-5 mm in length. Free twigs emanating from the interior of the fans are short (up to 2 or 3 mm long) and scarce. The mound-like protuberances resulting from the retracted polyps are small (mostly < | mm long), oval in shape with slit-like apertures, slightly raised, and red-orange at the bases with yellowish apices. These mounds are disposed in more-or-less distinct, usually four longitudinal rows along the branches. On a particular face of the fan, the mounds are arranged biserially, alternately so on narrow branches, alternate to opposite on thicker branches. Polyps are milky white with very thin points-like arrangements of sclerites. The coenenchymal sclerites are mostly girdled spindles with blunt, rounded ends, but a dom- inance of spindles with acute ends is also observed in some specimens (e.g., CAS 097025). The spindles vary in length from 0.08 mm -0.16 mm and are up to 0.04 mm in width. They have 4-6 WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO >) whorls of tubercles, with both ends acute or blunt, or with one end acute and the other blunt. Compact, relatively wide radiates or capstans (0.40—0.60 mm long), and relatively elongated eight radiates (0.60 mm—0.80 mm long and up to 0.04 mm wide), are also common in all the specimens. These characteristically have two whorls of tubercles and warty ends. The coenenchymal sclerites are entirely red or yellow, whereas some are bicolored (red and yellow). Anthocodial sclerites are elongated yellowish rods (up to 0.11 mm in length and 0.02 mm in width), with lobed or scalloped margins, and with short tubercles at the ends. Color in life is brick-red to rust-orange, often with orange main branches. The mounds formed by retracted polyps are mostly brick-red to dull orange. The color is conserved in alcohol-preserved specimens or is slightly paler in dried specimens. DESCRIPTION OF HOLOTYPE.— Growth form and size (Fig. 1A—B): The holotype measures 77 mm high by 163 mm wide, and is comprised of a single planar fan. Secondary fans are absent. The main stem is very short above the holdfast (<5 mm long). A strong main trunk-like branch is absent. Instead, two thickened branches (each approximately 4 mm in width) emanate from the basal stem, which form a more-or-less V-shape, and subtend the fan. The free terminal branches are 2-10 mm long. Networks are composed of relatively large, open meshes of variable shape. The meshes are mostly quadrilateral , trapezoidal, or deltoid in shape, whereas some are elongated. Meshes vary from 2—8 mm long and 2—3 mm wide; mostly 6—8 meshes/cm2. Mesh branches are narrow, most- ly 1.0-1.5 mm wide. Polyps (Fig. 1B): The polyps are all retracted and form mound-like, rounded protuberances in longitudinal rows along the branches. The mounds are approximately 1.0 mm long, usually oval in shape with slit-like apertures. Sclerites (Figs. 1C—D, 2-4): The coenenchymal sclerites are girdled spindles, some with blunt to rounded ends and others with acute tips, mostly 0.08—0.14 mm in length. Eight radiates are also present, mostly 0.05—0.07 mm long (Figs. 1C—D, 2-4). The terminal branches have a predomi- nance of spindles (Fig. 2A—E, G—J, M), whereas the holdfast region has mainly radiates (Fig. 3). Some of these are ornately sculptured (Fig. 3J-K, O), but others are more sparsely ornamented (Fig. 3A, E-F, M). The anthocodial sclerites are bisquit-shaped rods, 0.03—0.04 mm long (Fig. 2K, N-P), or elongated rods with more or less scalloped margins, up to 0.06 mm long (Fig. 2F, L). Sclerites are red, yellow, or bicolored (Figs. 1C—D). The various kinds of sclerites are shown at the same scale in Fig. 4. Color (Fig. 1A—B): The coenenchyme is brick-red to rust-orange throughout, with brick-red to dull orange mounds formed by the retracted polyps. The sclerites are mostly deep-red or bright yel- low, whereas some are bicolored red and yellow. Wet preserved specimens appear deep red, dried specimens pale orange. ETYMOLOGyY.— This species is named for William Dampier, English explorer, naturalist, and buccaneer, who visited the Galapagos Archipelago between 31 May and 12 June of 1684, and described various aspects of the flora and fauna, over a century and a half before Charles Darwin’s visit of 1835 (Dampier, 1927:75-83). DIsTRIBUTION.— Apparently restricted to outlying Darwin and Wolf Islands, in the extreme northwestern fringe of the Galapagos Archipelago; 9-21 m in depth (Figs. 5, 22). REMARKS.— All colonies examined, including the holotype, have numerous pinkish white ophiuroids attached to the them (Fig. 1A). These brittle stars measure approximately 10-15 mm in length and presumably represent a single species, which is unidentified at present. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 60 Volume 55, No. 3 gia dampieri sp. nov. A. Whole colony, wet-preserved; scale bar = 30mm. B. Detail of FIGURE 1. Holotype of Pacifig« branching pattern; nenchymal sclerites: scale bar = 0.10 mm. D. Micrograph of FF coe scale bar = 0.08 mm. scale bar = 6 mm. C. Micrograph « ones; d re cluding bicol« in sclerites coenenchymal WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 61 FIGURE 2. Holotype of Pacifigorgia dampieri sp. nov. Scanning electron micrographs of coenenchymal and anthocodi- al sclerites from the outermost branch tips. A. 0.06 mm. B. 0.11 mm. C. 0.08 mm. D. 0.09 mm. E. 0.05 mm. F. 0.06 mm. G. 0.14 mm. H. 0.06 mm. I. Scale bar = 0.06 mm. J. 0.05 mm. K. 0.03 mm. L. 0.06 mm. M. 0.10 mm. N. 0.04 mm. O. 0.03 mm. P. 0.04 mm. Anthocodial sclerites: K, N, O, P; all others are coenenchymal sclerites. 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 3. Holotype of Pacifigorgia dampieri sp. nov. Scanning electron micrographs of sclerites from the holdfast. A. 0.06 mm. B. 0.07 mm. C. 0.05 mm. D. 0.05 mm. E. 0.06 mm. F. 0.06 mm. G. 0.07 mm. H. 0.07 mm. I. 0.07 mm. J. 0.07 mm. K. 0.06 mm. L. 0.07 mm. M. 0.07 mm. N. 0.06 mm. O. 0.06 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 63 4. Pacifigorgia dampieri sp. nov. (CASIZ 097022). Scanning electron micrographs of sclerites shown at the e bar = 0.40 mm. 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 e Culpepper Darwin Pinta |) Abingdon flarchena Genovesa <> Bindloe =} Tower N James San Salvador ‘ Santiago a Bartolome _-Seymour Roca Redonda _ devis § Baltra ; Bebicg ™ > Plaza air Cruz a= Fernandina ~—— Narborough east} ° Isabela Pinzon ° es Albemarle — = = Duncan poanta Fe a San Cristobal Chatham Santa =~ indefatigable SS eee re 25 100 km Charles | = Floreana , “» Hood Santa Maria Espanola FIGURE 5. Pacifigorgia dampieri sp. nov. Map of geographic distribution; a = collecting stations. Inset shows geo- graphic range of the genus Pacifigorgia (in black). Pacifigorgia darwinii (Hickson, 1928) (Figs. 6-9, 24) Gorgonia darwinii Hickson, 1928: 390-392. Stiasny, 1941:268-270. Pacifigorgia darwinii: Breedy and Guzman, 2002:808-8 13. MATERIAL EXAMINED.— LECTOTYPE: BM 1930.6.17.10, Ecuador, Galapagos Islands, Isla Isabela, Tagus Cove, depth not recorded, 1923-1924, collected by C. Crossland. PARALECTOTYPES: BM 1936.8.30.1, 1961.2.6.148, 1961.2.6.150, same data as lectotype. BM 1936.8.29.1 (one of the original syntypes now designated as “Type” in the Darwin collection but more properly one of the paralectotype series), Ecuador, Galapagos Islands (specific island not recorded), depth not record- ed, 1835, collected by C. Darwin (Beagle Collection No. 1306, 21 November 1901). OTHER MATERIAL: CAS 097019, Ecuador, Galapagos Islands, Isla San Cristobal, Leon Dormido, 15—26 m depth (Station G2), 10 May 1994, collected by GC. Williams with SCUBA aboard “Mistral I”, one whole specimen and one portion of another colony wet preserved in 75% ethanol. CAS 097031, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, 3-18 m depth (Station G9), 14 May 1994, collected by GC. Williams with SCUBA aboard “Mistral IT”, two whole wet specimens preserved in 75% ethanol. CAS 097032, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, 3-18 m depth (Station G9), 14 May 1994, collected by GC. Williams with SCUBA aboard “Mistral IT”, four whole wet specimens, preserved in 75% ethanol. CAS 097035, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, <18 m depth (Station G10), 14 May 1994, collected by GC. Williams with SCUBA aboard “Mistral II’, one whole wet specimen, preserved in 75% ethanol. CAS 097039, Ecuador, Galapagos Islands, north shore of Isla Fernandina, <2] m depth (Station G13), 15 May 1994, collected by GC. Williams with SCUBA aboard “Mistral II”, one whole wet specimen preserved in 75% ethanol. CAS 097054, Ecuador, WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 65 Galapagos Islands, Isla Isabela, Punta Vincente Roca, <18 m depth (Station G10), 14 May 1994, collected by GC. Williams with SCUBA aboard “Mistral IT”, twelve whole wet specimens, pre- served in 75% ethanol. CAS 097117, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, 3-18 m depth (Station G9), 14 May 1994, collected by GC. Williams with SCUBA aboard “Mistral II”, one whole wet specimen, preserved in 75% ethanol. CAS 097901, Ecuador, Galapagos Islands, Isla San Cristobal, 21 m depth, June 1993, depth not recorded, collected by P. Humann with SCUBA, one whole dried colony. CAS 105031, Ecuador, Galapagos Islands, Isla Floreana, Devil’s Crown, 6 m depth, date not recorded, collected by P. Humann with SCUBA, one portion of a colony originally fixed in 10% formalin and wet preserved in 75% ethanol. CAS 147486, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, 9 m depth, 19 June 2001, collected by C. Hickman with SCUBA, one portion of a colony wet preserved in 75% ethanol. CAS 147487, Ecuador, Galapagos Islands, Isla Fernandina, Punta Espinosa, 21 m depth, 21 June 2001, collected by C. Hickman with SCUBA, one portion of a colony wet preserved in 75% ethanol. CDRS 03-86 and 03-77, Ecuador, Galapagos Islands, Islas Los Hermancs, 9 m depth, 18 January 2003, collected by C. Hickman. CDRS 01-95, Ecuador, Galapagos Islands, Isla Pinzon, 9 m depth, 22 June 2001, collected by C. Hickman. CDRS 1994, Ecuador, Galapagos Islands, Isla Isabela, Punta Vincente Roca, 18 m depth, 3 May 1994, collected by C. Hickman. CDRS 1991, Ecuador, Galapagos Islands, Isla Fernandina, Punta Espinosa, depth not recorded, 24 June 1991, collected by C. Hickman. GENERAL DESCRIPTION.— The lectotype was designated, examined, and described in detail by Breedy and Guzman (2002: 808). Also included in their description was a specimen studied here (CAS 097035). We, therefore, provide only a brief descriptive account. The colonies examined are up to 250 mm, dark purple, with yellow or white oval rings, and have open and evenly reticulated branching. Most colonies are composed of 2—3 parallel fans. The mesh branches are up to 1.5 mm in diameter. The network is composed of polygonal, oblong, or sometimes round meshes, up to 13 mm in length and 3 mm in width (approximately 7 meshes/cm2). The stem is short, not exceeding 5 mm in height, without distinct midribs. The free terminal branchlets are blunt and up to 10 mm in length. The oval mounds resulting from the retracted polyps are generally flat, and form yellow, cream, or white rings around the apertures of the polyps. These protuberances are mostly arranged in two longitudinal rows along most of the branches, and in multiple rows (4 or 5) on the thicker branches. The polyps are white with yellowish to pale pink sclerites arranged in points. The coenenchymal sclerites are mostly purplish red, whereas some are yellow and a few are bicolored. They are blunt spindles or capstans up to 0.09 mm long by 0.05 mm wide, and spindles with acute ends to 0.10 long and 0.04 mm wide. The anthocodial sclerites are flattened pale yellow rods up to 0.12 long and 0.02 mm wide, with scalloped or lobed margins. The specimen figured (CAS 097035) is a flat, kidney-shaped fan with reticulations even and open throughout, 185 mm wide by 122 mm high (Fig. 5A). The network is composed of mostly rounded rectangular meshes (2—5 mm in diameter) to irregularly-shaped elongated ones (8—15 mm long by 14 mm wide) (Fig. 5B). The basal stem is very short, 5 mm in length. The retracted polyps form low rounded mounds in two opposite longitudinal rows along each face of the colony. The coenenchyme is dark reddish-purple, whereas the polyps mounds are mostly yellow (Fig. 5). Sclerites are mostly reddish purple (Fig. 5C), some are yellow, and a few are bicolored (red and yellow). Coenenchymal sclerites are mostly elongated spindles, some with tapering or distinctly pointed ends, 0.09-0.11 long (Figs. 5C, 6E—-O), and mostly blunt eight radiates, 0.06—0.08 mm in length (Fig. 7). Some radiates lack ornate sculpture (Fig. 6A—D). Anthocodial sclerites are flattened rod-like sclerites with scalloped margins, up to 0.12 mm long (Fig. 6K). DISTRIBUTION.— Southern islands of the Galépagos Archipelago (south of the Equator), 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES — Volume 55, No. 3 PDT FIGURE 6. Non-type specimen of Pacifigorgia darwinii (CAS 097035) A. Whole colony, wet preserved; scale bar = 48 mm. B. Detail of branching pattern; scale bar = 13 mm. C. Micrograph of coenenchymal sclerites; scale bar = 0.15 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 67 FiGurE 7. Non-type specimen of Pacifigorgia darwinii (CAS 097035). Scanning electron micrographs of sclerites from the tips of the ultimate branches. A. 0.06 mm. B. 0.04 mm. C. 0.06 mm. D. 0.07 mm. E. 0.09 mm. F. 0.09 mm. G. 0.05 mm. H. 0.06 mm. J. 0.10 mm. J. 0.10 mm. K. 0.12 mm. L. 0.10 mm. M. 0.10 mm. N. 0.10 mm. O. 0.11 mm. 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 8. Non-type specimen of Pacifigorgia darwinii (CAS 097035). Scanning electron micrographs of sclerites from the holdfast. A. 0.06 mm. B. 0.06 mm. C. 0.06 mm. D. 0.08 mm. E. 0.07 mm. F. 0.06 mm. G. 0.07 mm. H. 0.07 mm. I. 0.06 mm. J. 0.06 mm. K. 0.07 mm. L. 0.07 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 69 e Culpepper Darwin Abingdon Marchena eenOvoed om, Bindloe ©) Tower Roca Redonda ag N James San Salvador Santiago Bartolome _-seymour Baltra Fernandina _ @ _ Plaza Narborough gs Barrington ree San Cristobal Chatham Floreana P QQ Hood Santa Maria Espanola FiGuRE 9. Pacifigorgia darwini. Map of geographic distribution; A = collecting stations; * = location of Galapagos Islands. including Isabela, Fernandina, Los Hermanos, Floreana, Gardner off Floreana or Gardner off Espanola (not specified), and San Cristobal; 3-26 m depth (Fig. 8). The presently known distribu- tion forms a crescent-shaped arc from east to west in the southern part of the archipelago (Fig. 24). REMARKS.— This species is the most common one of the genus around the islands. It is clear- ly distinct from the other taxa. We found some variation in the sclerites — there is a dominance of spindles with acute ends in some samples. This is the same situation with the other species found in Galapagos, which herein we describe. Also, the mound-like protuberances resulting from the retracted polyps varies from yellow to cream or white — San Cristobal (yellow, cream, or white); Isabela (cream or yellow); Fernandina (yellow); Floreana (white). Pacifigorgia douglasii (Hickson, 1928) (Figs. 10-11, 24) Gorgonia darwinii var. douglasii Hickson, 1928:392-393. Pacifigorgia douglasii (Hickson, 1928). Breedy and Guzman, 2002:813-815. PREFATORY NOTE.— Breedy and Guzman (2002:813) considered Hickson’s Gorgonia darwi- ni var. douglasii as a member of the genus Pacifigorgia, and raised the variation to specific status as a new combination. They believed the differences between Pacifigorgia darwinii and Hickson’s taxon to be significant enough to justify raising the latter to species level. They designated the material cited below as lectotype. They also provided a description of the surviving material. MATERIAL EXAMINED.— LECTOTYPE: BM 1961.2.6.161, 162, two microscope slides of scle- rites only, Ecuador, Galapagos Islands, Isla Onslow, 8 m in depth, 1923-24, original material col- lected by Major Douglas. DESCRIPTION.— See Breedy and Guzman (2002:813-—815) for a detailed description and eval- uation. The sclerites are figured in the present paper (Fig. 10). 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 DISTRIBUTION.— Reported from one locality, Onslow Island (off the north shore of Floreana), Galapagos Archipelago, 8 m depth, attached to a fishing line (Fig. 11). REMARKS.— We have not found a specimen of the colony that matches the lectotype sclerite preparation. These colonies appear to be missing and presumably all that remains of material rep- resenting this taxon are two microscope slides of sclerites. We recently found in the NHM collection, a non-type specimen lot (BM 30.6.17.11) labeled as Gorgonia darwinii var. douglasii, containing three fragments of what we consider to be Pacifigorgia darwinii. One of the fragments represents the variety of P. darwinii with white rings, whereas the other two fragments are more representative of the typical form with yellow to cream WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 71 @ Culpepper Darwin Pinta » Abinadon WMarchena Genovesa Bindl @® Tower Roca Redonda eo N James San Salvador Santiago Bartolome oe Seymour , edervis Baltra = Rabida 2 Plaza Fernandina @ | mie anta Cruz Narborough « x Indefatigable pele rt » Pin20n Santa Fe Barrington Soe 3°" San Cristobal A Chatham = Charles & Floreana > Hood Santa Maria Espanola Ficure 11. Pacifigorgia douglasii. Map of geographic distribution; A = collecting stations; * = location of Galapagos Islands. polyps mounds. Even though the morphology of the three fragments agrees in many respects to Hickson’s description of Gorgonia darwinii var. douglasii, the sclerites are more consistent with those of P. darwinii. They differ in form and size from the sclerites on the lectotype slides (see the assessment of Breedy and Guzman 2002). After examining many specimens, we conclude that the fragments can all be aligned with P. darwinii. There is not sufficient reason to separate the white- ringed variety from the typical form. It should therefore be viewed within the range of variability for the species. Because of the depauperate nature of the type material, and the lack of any other specimen that matches Hickson’s description, it is not possible to validate the taxonomic status of Pacifigorgia douglasii, and we must therefore consider this to be a dubious taxon. In addition, northern Floreana Island is also a collecting station for typical specimens of Pacifigorgia darwinii (Figs. 9, 24). Pacifigorgia rubripunctata Williams and Breedy, sp. nov. (Figs. 12-16, 24) MATERIAL EXAMINED.— HoLotyPe: CAS 167958, Ecuador, Galapagos Islands, Isla Santa Cruz, Academy Bay, 3-6 m depth, 1964 (day and month not recorded), collected by Andre and Jacqueline De Roy. One whole wet specimen preserved in 75% ethanol. PARATYPE: CAS 147485, Ecuador, Galapagos Islands, Isla Rabida, 6 m depth, (Station 01-37), 17 June 2001, collected by Cleave Hickman. One whole specimen preserved in 75% ethanol. OTHER MATERIAL: CAS 167959, same data as holotype, two dried fragments of colonies. CAS 053266, same data as holotype, one whole dried colony. CAS 101762, Ecuador, Galapagos Islands, Isla Espafiola, Punta Sudrez, 7.6 m depth, 4 March 1977, collected by John McCosker and party. Two whole wet specimens, originally preserved in formalin, later transferred to 75% ethanol. CAS 096904, Ecuador, Galdpagos Islands, Isla Santiago, Sullivan Bay near Isla Bartolomé, depth not PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 3 | 30 mm. B. colony, wet-preserved; scale bar of branching pattern; scale bar = 20 mm. 1 rubripunctata sp. nov. A. Whole 0.10 mm. C. Detail bar scale WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO W3 recorded, 21 January 1938, collector not identified, one fragment of a colony wet preserved in 75% ethanol. CAS 147484, Ecuador, Galapagos Islands, Isla Rabida, 6 m depth, (Station 01-32), 17 June 2001, collected by Cleave Hickman. CAS 105033, Ecuador, Galapagos Islands, Nameless Islet off west coast of Isla Santa Cruz, 20 m depth, date not recorded, collected by Paul Humann, one whole specimen wet preserved in 75% ethanol. MCZ 36260 and MCZ 36262, Ecuador, Galapagos Islands, Isla Santa Cruz, Punta Astrada, 8-10 m depth, 1977, collected by S. Taylor. GENERAL DESCRIPTION.— Colonies wider than high, up to 120 mm 9in height, and 150 mm in width, composed of a single primary fan and 2—3 smaller secondary fans. Secondary fans stick out from different parts of the main fan, and radiate in right angles, hence, they grow perpendicularly to the main fan. Colonies seem to have encrusting holdfasts. Color when dry is a light violet with red spots, which are sparsely or closely disposed, and bright reddish-orange polyp mounds. Wet preserved specimens are yellowish to white, spotted with red and purple, and polyp mounds red- dish-orange, rarely yellowish. Networks are regular and comprised of angular meshes. The meshes are up to 15 mm in length and 3 mm in width (density averages approximately 6 meshes/cm2). Mesh branches up to 1.5 mm in diameter. No distinct midribs crossing the fans are observed, only thick short branches (up to 10 mm width) at the base. These extend for a short distance to one side of the fan, parallel to the sub- stratum. Short stems raise the colony a little from the substratum. Terminal branchlets are short, up to 2 mm in length. Free twigs are very scarce. Polyp mounds are slightly raised, arranged in pairs in longitudinal rows, on the external part of the branches, and separated by branches of dark pur- ple coenenchyme in the middle of the lines of polyps. Polyps are white with flattened anthocodial sclerites arranged in weak points. Coenenchymal sclerites (Fig. 15) are mostly spindles with blunt ends, and radiates (capstans) with elongated ends, but a dominance of spindles with acute ends is also observed (e.g. CAS 105033). The sclerites are red, orange, and colorless with different predominance of colors, but some are bicolored. Spindles are long (up to 0.15 mm in length, and 0.04 mm in width) and with 4-8 whorls of tubercles, with both ends acute or blunt, or with one acute and the other blunt. Radiates (capstans) are wide (up to 0.09 mm in length, and 0.05 mm in width) with two whorls of tubercles and warty ends. A few crosses with acute or round ends (up to 0.09 long by 0.06 mm wide) are also present. Anthocodial sclerites are pale yellow rods (up to 0.10 mm in length, and 0.02 mm in width) with smooth or lobed margins (Fig. 15 shows all sclerites at the same scale). DESCRIPTION OF HOLOTYPE.— Growth form and size (Fig. 12A, C). The holotype measures 120 mm in height by 94 mm in width. It is comprised of two similarly sized parallel fans that are joined by a common basal main stem. Only one of these fans can be seen in Fig. 12A. The main stem is flattened and measures 2 mm by 7 mm in transverse section. The holdfast portion is miss- ing since the specimen was cut just above the holdfast to separate it from the substratum. The sur- viving portion of the main stem is approximately 40 mm in length and gives rise to an intricate and uniform network without conspicuous secondary branches. The networks are regular and com- prised of meshes of variable shape. The meshes in the upper portion of one of the fans are mostly more or less square and arranged in longitudinal rows. Meshes of the upper portion of the other parallel fan vary from narrow and elongated to square or rectangular, and are not disposed in dis- tinct rows. Meshes vary in size from 2 mm minimum width to 15 mm maximum length. The mesh branches are mostly 1.0-1.5 mm in width. The density of meshes varies from approximately 5—9 meshes/cm?. The free tips of terminal branches are up to 5 mm in length. Polyps (Fig. 12C): The polyp mounds formed by the retracted polyps are only slightly raised, often irregularly-shaped (deltoid to trapezoidal) as well as circular, with slit-like apertures. The mounds are distributed in four longitudinal rows (two rows on each face of a particular fan) along 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 13. Holotype of Pacifigorgia rubripunctata sp. nov. Scanning electron micrographs of sclerites from the outer- most branch tips. A. 0.08 mm. B. 0.09 mm. C. 0.06 mm. D 0.06 mm. E. 0.06 mm. F. 0.08 mm. G 0.06 mm. H. 0.05 mm. I. 0.09 mm. J. 0.07 mm. K. 0.05 mm. L. 0.04 mm. M. 0.04 mm. N. 0.06 mm. O. 0.07 mm. P. 0.06 mm. Q. 0.08 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO US FIGURE 14. Holotype of Pacifigorgia rubripunctata sp. nov. Scanning electron micrographs of sclerites from the hold- fast. A. 0.08 mm. B. 0.06 mm. C. 0.08 mm. D. 0.09 mm. E. 0.08 mm. F. 0.07 mm. G. 0.07 mm. H. 0.06 mm. I. 0.07 mm. J. 0.08 mm. K. 0.09 mm. L. 0.06 mm. M. 0.06 mm. N. 0.07 mm. O. 0.08 mm. P. 0.07 mm. Q. 0.08 mm. R. 0.07 mm. S. 0.08 mm. T. 0.07 mm. U. 0.08 mm. V. 0.09 mm. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES ~ Volume 55, No.3 | FiGurE 15. Pacifigorgia rubripunctata sp. nov. (CASIZ 147484). Scanning electron micrographs of sclerites shown at the same scale; scale bar = 0.40 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO Ud e Culpepper Darwin Pinta § Abingdon” Marchena eenovceca Bindloe m/s N Roca Redonda aD James San Salvador Santiago Dy, Bartolome _« Seymour t 4 Baltra Fernandina } Narborough ahh anta Cru fe page retigebe n goanta Fe arrington F 5 3°" San Cristobal Chatham Charles ABP ‘a Floreana " A®) Hood Santa Maria Espanola Ficure 16. Pacifigorgia rubripunctata sp. noy. Map of geographic distribution; A = collecting stations; * = location of Galapagos Islands. the mesh branches. Polyps have weak points sparsely disposed anthocodial sclerites that are elon- gated and flattened. Sclerites (Figs. 12B, 13-15): Sclerites are red-purple, orange, or bicolored (orange at one end and colorless at the opposite end) (Fig. 12B). Coenenchymal sclerites are robust spindles or more elongated spindles with acute ends (0.08—0.11 mm long by 0.03—0.05 mm wide), and eight radiates (capstans) (0.04—0.08 mm long by 0.03—0.04 mm wide) (Figs. 13—14). A few cross-like sclerites are also present (approximately 0.08 mm long by 0.05 mm wide). Anthocodial sclerites are flat- tened rods with undulating margins (mostly < 0.10 mm long). Color (Fig. 12A, C): The coenenchyme throughout is a deep purplish red. The polyp mounds are orange, and contrast sharply with the surrounding coenenchyme. The polyps are white. ETYMOLOGyY.— The specific epithet of this new species is derived from the Latin rubra (red) and punctatus (spotted); in reference to the fine red-orange spots along the branches of most spec- imens, which are created by the retracted polyps. DISTRIBUTION (Figs. 16, 24).— The range of this species is apparently restricted to a line extending from northwest to southeast, running through the center of the Galapagos Archipelago (Fig. 24): known from Islas Santiago, Bartolomé, Rabida, Santa Cruz, and Espanola (Fig. 16). VARIATION.— A considerable amount of color variability is evident in this species. The coenenchyme varies from deep red or dark reddish purple to greyish-white, or yellow. The polyp mounds can vary from red to orange in most material, or pale yellow in one specimen (CAS 167959). The colors between the coenenchyme and the polyp mounds contrast sharply in all spec- imens. 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No.3 — Pacifigorgia symbiotica Williams and Breedy, sp. nov. (Figs. 17-24) MATERIAL EXAMINED.— HOoLotyPeE: CAS 097118, Ecuador, Galapagos Islands, Isla Darwin, Arch, 12-18 m depth, (Station G6), 13 May 1994, collected by Gary C. Williams with SCUBA, aboard “Mistral IT’. One whole wet specimen preserved in 75% ethanol. PARATYPE: CAS 097115, same data as holotype. One whole wet specimen preserved in 75% ethanol. OTHER MATERIAL: CAS 097041, same data as holotype. One whole wet specimen preserved in 75% ethanol. CAS 097948, Ecuador, Galapagos Islands, Isla Wolf, depth not recorded, 15 December 1898, collector not recorded, two dried fragments of colonies. CAS 097902, Ecuador, Galapagos Islands, Isla Wolf, 14 m depth, June 1993, collected by Paul Humann, one dried frag- ment of a colony. ; GENERAL DESCRIPTION.— Fans are stiff, planar or with smaller secondary fans. The main fans are wider than high (71—285 mm in width by 65-122 mm in height). Colonies have holdfasts that are strong and spreading (12—60 mm in maximum dimension). The main stem and primary branch- es are somewhat flattened and relatively wide (4-10 mm wide). This characteristic, along with the somewhat thickened mesh branches, gives the colonies a relatively rigid aspect. The networks are intricate with open meshes. The meshes are highly variable in shape and size. They are often rectangular or deltoid, but very narrow and elongated ones as well as small cir- cular ones are also evident. Meshes vary from 2—5 mm in width by 2—12 mm in length. Colonies usually have 6-8 meshes/cm2. Mesh branches are relatively thick (1.5—2.5 mm in width). The thickened mesh branches make the meshes appear relatively small. The free terminal branchlets at the distal margin of the colonies are 2—8 mm in length, thick and blunt or truncated at the tips. The mound-like protuberances produced by the retracted polyps are relatively robust, hemispherical or mammiform, |—2 mm in diameter, with ovoid or slit-like apertures. They cover most of the surface area of the mesh branches. The coenenchymal sclerites are of two kinds. In the holdfast and main stem region, there is a predominance of relatively elongated eight radiates (0.06—0.08 mm long) with a relatively wide separation between the whorls of tubercles. In the mesh branches and the distal extremities of the colonies, the sclerites are girdled spindles, many of which have strongly attenuated ends (0.08—0.12 mm long); as well as compact, ovoid eight radiates (0.03—0.08 mm long). Some of the smaller radi- ates lack ornate tuberculation. The anthocodial sclerites are flattened, elongated rods with more or less scalloped or undulating margins (0.05—0.13 mm long). These are arranged in eight weakly dis- posed points. A crown is not apparent. The color of the sclerites is mostly either dark red or bright yellow. However, some sclerites are bicolored — yellow and red. The superficial coenenchyme is deep reddish purple, whereas the polyp mounds are golden yellow to lemon yellow or orange-yellow. The polyps are white, with eight weak points of mostly yellow rod-like sclerites. DESCRIPTION OF HOLOTYPE.— Growth form and size (Fig. 17A, C; 18). The holotype meas- ures 98 mm high by 138 mm wide, composed of a single planar fan. An incipient secondary fan arises from near the distal terminus of the colony, and is parallel to it (22 mm high by 44 mm wide). The origin of this secondary fan is approximately 8 mm below the distal terminus of the main fan. The holdfast is spreading (23 mm wide by 28 mm long). The main stem is very short and flattened (approximately 6 mm long by 7 mm wide by 3 mm thick). This main stem gives rise to four pri- mary branches, 12—18 mm long. An intricate network of mesh branches emanates from the distal regions of the primary branches. Networks are comprised of relatively small meshes with thick- ened mesh branches; approximately 6-8 meshes/cm2. Meshes are highly variable and irregularly WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 79 FIGURE 17. Pacifigorgia symbiotica sp. nov. A. Living coral with two epizoic mimics, a palaemonid shrimp (left arrow), and an ovulid gastropod (right arrow); scale bar = 10 mm. B. Three coenenchymal sclerites from the holotype showing vari- ation in color; scale bar = 0.8 mm. C. Wet-preserved holotype; scale bar = 45 mm. 80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 18. Pacifigorgia symbiotica sp. nov. A-D. Micrographs of colony external morphology from the preserved holo- type - branching, reticulation, and retracted polyps. Scale bars = 1 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 81 FiGure 19. Pacifigorgia symbiotica sp. nov. A. Coenenchymal sclerites. B. Polyp sclerites. Scale bars = 0.05 mm. 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 20. Pacifigorgia symbiotica sp. noy. Scanning electron micrographs of sclerites. A-C. Polyp sclerites. A. 0.14 mm. B. 0.12 mm. C. 0.10 mm. D-N. Coenenchymal sclerites. D. 0.06 mm. E. 0.06 mm. F. 0.06 mm. G. 0.05 mm. H. 0.06 mm. I. 0.06 mm. J. 0.09 mm. K. 0.08 mm. L. 0.10 mm. M. 0.07 mm. N. 0.08 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 83 FiGURE 21. Pacifigorgia symbiotica sp. nov. Scanning electron micrographs of coenenchymal sclerites from the branch tips. A. 0.07 mm. B. 0.08 mm. C. 0.07 mm. D. 0.07 mm. E. 0.06 mm. F. 0.06 mm. G. 0.03 mm. H. 0.05 mm. [. 0.12 mm. J. 0.06 mm. K. 0.11 mm. L. 0.06 mm. 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 3 FIGURE 22. Pacifigorgia symbiotica sp. nov. Scanning electron micrographs of coenenchymal sclerites from the hold- fast. A. 0.06 mm. B. 0.08 mm. C. 0.07 mm. D. 0.06 mm. E. 0.07 mm. F. 0.07 mm. G. 0.06 mm. H. 0.07 mm. I. 0.07 mm. J. 0.08 mm. K. 0.06 mm. L. 0.07 mm. WILLIAMS AND BREEDY: PACIFIGORGIA IN THE GALAPAGOS ARCHIPELAGO 85 e Culpepper Darwin Pinta Abingdon — Genovesa Marchena a se «a, Bindloe ad Roca Redonda ED N James San Salvador Santiago Bartolome _-seymour Fernandina Narborough Santa Cru 5 z3 Indefatigable ° Pinzon Pp Tot ounean genta, Fe 7 San Cristobal =e Chatham Charles |) * Floreana-

= & Si ry MY ¥ ELAR GAEL: of abaxial foliar surface, x ca. 3; D. portion of inflorescence rachis showing bracts and bracteoles, x ca. 12; E. smooth appressed hairs of abaxial leaf surface, x ca. 20; F. bracteole, x ca. 12; G. petal, x ca. 7; H. stamens, profile view (left) and dorsal view (right), x ca. 12; I. berry, x ca. 4; J. seeds, x 32. (A— F, I and J from Almeda et al. 6437; B from Folsom & d FIGURE 2. Clidemia coloradensis Almeda. A. habit, x 2; B. representative leaf (abaxial surface), x 2; C. enlargement Collins 1814; G and H from Croat 37253.) ALMEDA: NEOTROPICAL GENUS CLIDEMIA 95 anthers 1.5—2 x 0.5 mm, linear-oblong, yellow with a dorsally inclined apical pore; connective nei- ther prolonged below the thecae nor appendaged. Ovary completely inferior, 5-locular, apex glabrous. Style 5—5.5 mm long, + straight, glabrous; stigma capitate. Berry 5-8 x 6-7 mm when dry, turning pink then blue-purple when ripe. Seeds 0.5 mm long, + triangular in profile view and + obovoid in dorsal view, testa tuberculate. PHENOLOGY.— Collected in flower during July and August, in fruit from January through April. DISTRIBUTION.— Locally common in primary cloud forests from southeastern Costa Rica to western Panama at 700-1700 m. PARATYPES.— Costa Rica: Cartago: Turrialba. Moravia. Cerro Tigre. Siguiendo por la que- brada innominada, bajando hasta Lago Ayil, 09°53’40”N, 83°22’55”W, 28 Mar. 1996, Kuss 49 (CAS, CR). PANAMA. Bocas del Toro: Edwin Fabrega Dam and Reserve in Fortuna along the Continental Divide Trail, 12.9 km N of Sitio de Presa (Dam site) offices above the Dam, 08°45’04’N, 82°15’04’W, 7 Feb 1996, Almeda et al. 7538 (CAS, PMA). Bocas del Toro/Chiriqui border: windswept cloud forest on slopes and valleys of the Cerro Colorado region, 27 Jan. 1989, Almeda et al. 6437 (CAS, MO, PMA, US). Cerro Colorado, 11.2 km along ridge road from main road to Escopeta, 16 Aug. 1977, Folsom 4879 (MO). Chiriqui: Cerro Colorado, along road to cop- per mine, 34.1 km beyond bridge over Rio San Felix near town of San Felix, 15 July 1976, Croat 37253 (CAS); Cerro Colorado top, Bocas Road, 17—18 Feb. 1977, Folsom & Collins 1814 (CAS); between Pinolz and Divide on Chiriqui side. Caldera Trail, mostly mossy forest above the oak for- est, 19 Apr. 1961, Kirkbride & Duke 860A (MO); between Quebrada Hondo and divide on Caldera Trail, 20 Apr. 1968, Kirkbride & Duke 954 (MO, 2 sheets). DisCussIoN.— The slender or little-branched habit of Clidemia coloradensis and its deep for- est habitat are typical of many formicarial species of this genus in the Mesoamerican region. Clidemia coloradensis, however, never appears to produce formicaria (ant domatia) of any kind. This species appears to flower during the rainy season which may account for the fact that few good flowering collections have been available for study. Clidemia coloradensis is readily recognized by a distinctive combination of characters. The indument on upper internodes, the inflorescence axis, and elevated veins on abaxial foliar surfaces consists of smooth appressed hairs (1-3 mm long) intermixed with densely matted woolly hairs. The inflorescence is a long-pedunculate modified cluster of dichasia with ultimate units that often appear umbelliform. The calyx teeth are shorter than or only slightly exceeding the calyx lobes (Fig. 21), the anther pores are dorsally inclined (Fig. 2H), the stigma is capitate, and the seed coat is tuberculate (Fig. 2J). The apiculate bracteoles that are concave on the adaxial side are also dis- tinctive (Fig. 2F). Because of its distinctiveness, no close relatives in any part of the American trop- ics are evident at this time. ETYMOLOGY.—The specific epithet is derived from Cerro Colorado in western Panama where most of the fertile material of this species has been collected. Clidemia davidsei Almeda, sp. nov. (Fig. 3) TyprE.— Costa Rica: Puntarenas: Cordillera de Talamanca; area around Rio Canasta, 9.5 air- line km NW of Agua Caliente, between Cerro Frantzius and Cerro Pittier; elev. 1500-1600 m, 9°02’N, 82°59’W, 6 Sep. 1984, G Davidse et al. 28374 (holotype: CAS!, isotypes: BM, CR, MO, PMA). Ramuli teretes sicut petioli foliorum venae primarieae subtus inflorescentia hypanthiaque 96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 dense vel modice setosi (pilis 0.5—1 mm longis pro parte glanduliferis) et sparse cum pilis stellatis obsiti. Lamina 6.5—16.5 x 3—8.5 cm ovata vel elliptico-ovata apice acuminata basi rotundata vel obtusa, 5—7-plinervata. Inflorescentiae pauciflorae in foliorum superiorum axillis oppositis; flores 5-meri, bracteolis 0.5—1.5 x 0.25—0.5 mm; calycis tubus 0—0.25 mm, lobis 1.5—2 x 0.5 mm subu- latis, dentibus exterioribus obsoletis. Stamina isomorphica glabra, antherarum thecae 1 x 0.4 mm oblongae poro dorsaliter inclinato; connectivum nec prolongatum nec appendiculatum. Ovarium 5- loculare et omnino inferum apicem versus sparsiuscule glandulis vel paullulo stellatis armatum. Lax shrub 1—3 m tall, the terete upper internodes, petioles, elevated primary veins of abaxial foliar surfaces, inflorescence, and hypanthia copiously to moderately covered with smooth (gland- tipped in part) spreading hairs 0.5-1 mm long with a ground layer of sessile stellate hairs. Leaves of a pair equal to somewhat unequal in size; petioles 1.3—5 (—6.2) cm long; blades 6.5—13.3 (—16.5) x 3-6.5(—8.5) cm, chartaceous and brittle when dry, ovate to elliptic-ovate, apex acuminate, base rounded to obtuse, margin entire to inconspicuously denticulate, 5—7-plinerved with the innermost pair of primary veins diverging from the median vein 3—7 mm above the blade base, the adaxial surface moderately covered with spreading smooth hairs and some glandular hairs 0.5—1 mm long intermixed with a scattering of stellate hairs on the actual surface and a concentration of stellate hairs on the impressed primaries toward the blade base, the secondary and higher order veins of the abaxial surface beset with smooth spreading hairs 0.5 mm long, the understory consisting of a very sparse scattering of stellulate hairs or the understory indument lacking. Inflorescence 1.5—2 cm long, axillary dichasia branched from the base or short-pedunculate and commonly paired at a node; bracteoles 0.5—1.5 x 0.25—0.50 mm, narrowly elliptic to oblong, apiculate and tipped with a short hair (otherwise glabrous). Pedicels (at anthesis) 0.5 mm long. Hypanthia 1—2 x 1.5 mm (at anthesis), campanulate to suburceolate. Calyx tube 0.25 mm long or obsolete, the calyx lobes 1.5—2 x 0.5 mm, subulate, beset with smooth spreading hairs like the hypanthium, calyx teeth lacking. Petals 5 in number, 2.54 x 0.5—1.5 mm, glabrous, white, + oblong. Stamens 10, isomorphic, fila- ments 1.5 mm long, glabrous; anthers | x 0.4 mm, oblong, yellow with a dorsally inclined apical pore; connective thickened dorsally but neither prolonged below the thecae nor appendaged. Ovary 5-locular, completely inferior, apex sparsely and deciduously glandular or stellulate-puberulent, elevated into a low truncate cone at anthesis but becoming + flat or rounded on maturing berries. Style 3 mm long, glabrous; stigma truncate to capitellate. Berry 4-5 x 4-5 mm when dry, blue- black or purple-black when ripe. Seeds 0.5 mm long, obovoid, testa + smooth. PHENOLOGY.— Collected in flower during January, March, July and August, and in October; in fruit from August through October, January and March. DISTRIBUTION.— Local and uncommon in cloud forests, river and lake margins, and forest clearings in southern Costa Rica with one outlying population in north-central Nicaragura at 1100-1750 m. The elevation on a single collection (Williams et al. 28572) is given as 3200 m. I Suspect this report may be erroneous and should be regarded with suspicion until this species is rec- ollected at that elevation again. PARATYPES.— NICARAGUA: Jinotega: Las Alturas de Kilambé, 13°37’N, 85°40’W, 26 Mar. 1981, Moreno & Sandino 7568 (CAS). Costa Rica: Puntarenas: Zona Protectora Las Tablas, slopes of Cerro Chai along Rio Bellavista drainage on trail to Cerro Echandi, 15 Feb. 1991, Almeda et al. 6637 (CAS, CR, MO); Finca Las Alturas, NW of the Lecheria, 17 Feb. 1991, Almeda et al. 6706 (CAS, CR); above coffee fincas along Rio Coto Brus, near Coton, 23 km N of La Union on the Panama border, 9 Aug. 1974, Croat 26604 (CAS, US); foothills of the Cordillera de Talamanca, FiGuRE 3. Clidemia davidsei Almeda. A. habit, x 2; B. representative leaf (adaxial surface), x 34; C. representative leaf (abaxial surface), x */4 ; D. flower (at anthesis), x ca. 10; E. petal, x ca. 11; F. stamens, %4 dorsal view (left) and ventral view (right); G. berry, x 5; H. enlargement of berry surface, x ca. 9; I. seeds, x 30. (A —I from Schnell 1046.) 97 ALMEDA: NEOTROPICAL GENUS CLIDEMIA 98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 lower montane forest along the Rio Bellavista, NW of Las Alturas, ca. 8°57’N, 82°51’W, 30 Aug. 1983, Davidse 24318 (CAS, CR, MO); cloud forest above Wilson’s finca, 6 km S of San Vito de Java, 14 Aug. 1967, Raven 21763 (MO); cloud forest above Wilson’s finca, 6 km S of San Vito de Java, 17 Aug. 1967 Raven 21843 (CR, DUKE, MO); vicinity of Finca Las Cruces, San Vito de © Java, 22-25 Aug. 1968, Schnell 1046 (US); just S of San Vito de Java, W of Cruces for ca. 3 km | along the road running along the Fila Cruces, 19 May 1984, Taylor 3456 (DUKE); cloud forest near lake on ridge SW of Las Cruces, 21 July 1977, Webster 21985 (CAS, DAV); cloud forest on ridge, 5 km S of San Vito, 11 May 1971, Wilbur 14456 (DUKE). San José: on Panamerican highway N of San Isidro del General, 15 Oct. 1965, Schnell 175 (US); Cordillera de Talamanca, about 25 km N of San Isidro del General along Panamerican highway, 29 Jan. 1965, Williams et al. 28572 (US). DISCUSSION.— This species, which was first collected in 1965, is easily recognized by its indu- ment of spreading hairs (gland-tipped in part) with a ground layer of stellate hairs (Fig. 3H), axil- lary dichasia that are commonly paired at a node (Fig. 3A), absence of exterior calyx teeth (Fig. 3G), inferior ovary, and smooth seed coat (Fig. 31). Collections of C. davidsei examined for this study have been variously identified as C. dentata D. Don, C. costaricensis Cogn. & Gleason, and C. petiolaris (Schltdl. & Cham.) Schltdl. ex Triana. It is most like C. dentata in general aspect and foliar size. In C. dentata the inflorescence is pseudolateral, the calyx consists of a truncate rim that is obscured and exceeded in length by subulate exterior teeth, and the adaxial surface of the torus (vascular ring at the apex of the hypanthium) is adorned with 10 fimbriate scales that are absent in C. davidsei. The ovary of C. dentata is 5-locular like that of C. davidsei but it is '4-inferior (vs. completely inferior) and its seed coat is rugulate or granulate (vs. essentially smooth). Both C. costaricensis and C. petiolaris differ most notably from C. davidsei in having pseudolateral inflo- rescences that are never paired at a node and a calyx that is clearly differentiated into lobes and exterior teeth. Costa Rican material of C. davidsei, which was all collected in the southwestern area of the country, is morphologically homogeneous. The single cited collection from Nicaragua, however, was puzzling and not placed in the recent treatment that I prepared for Flora de Nicaragua (Almeda 2001). It has slightly longer eglandular cauline trichomes (to 2 mm), longer petioles (to 6.2 cm), and larger leaf blades (16.5 x 8—8.5 cm) than the Costa Rica populations. In all other details it appears to be a good match for C. davidsei and is tentatively assigned to this species. Its status will require further consideration when better material comes to light. ETYMOLOGY.— This species is named for Gerrit Davidse (b. 1942), collector of the type, authority on the systematics of Poaceae, and a founding editor of the ambitious Flora Mesoamericana project. Clidemia evanescens Almeda, sp. nov. (Fig. 4) TyPE.— PANAMA. Coclé; forested slopes above El Copé along abandoned road leading to the Continental Divide, elev. 700-850 m, 25 Feb. 1988, F) Almeda et al. 5933 (holotype: CAS!; iso- types: AAU!, INB!, MO!, MEXU!, NY!, PMA!, US!). Ramuli teretes sicut petioli foliorum venae primariae subtus inflorescentia hypanthiaque dense pilis asperis induti. Lamina 9-27 x 3.4—-14.5 cm elliptica vel elliptico-ovata apice gradatim acumi- nata vel caudato-acuminata basi obtusa vel rotundata, 3—5-nervata. Inflorescentia primum termi- nalis demum lateralis pauciramosa vel in foliorum superiorum axillis; flores 4-meri, bracteolis 1.5—3 x 1-2 mm, calycis lobis interioribus 2—2.5 x 1.5 mm triangularibus, dentibus exterioribus 0.5—1.5 mm eminentibus. Stamina isomorphica glabra, antherarum thecae 1.5 x 0.5 mm oblongae poro dorsaliter inclinato; connectivum vix (0.1 mm) prolongatum, dente dorso-basali 0.5 mm glan- ALMEDA: NEOTROPICAL GENUS CLIDEMIA 99 dulis 0.1 mm longis marginato. Ovarium 4-loculare et omnino superiorum apice dense glandulis inconspicuis induto. Shrub 0.54 m tall with the terete upper internodes, petioles, elevated primary and secondary veins on abaxial foliar surfaces, inflorescence rachis, bracteoles, hypanthia, and calyx lobes mod- erately to copiously covered with asperous-headed hairs. Leaves of a pair + equal to somewhat unequal in length; petioles 1.54.7 cm long; blades 9—27 x 3.4—14.5 cm, subcoriaceous when dry, elliptic to elliptic-ovate, apex gradually acuminate to caudate-acuminate, base obtuse to rounded, margin essentially entire, 3—5-nerved, the secondary and higher order veins moderately to copious- ly beset with minute glands. Inflorescence 1—2.5 cm long, typically an axillary cluster of dichasia with branching initiated at the node or terminal with both lateral buds at a node elongating and overtopping the inflorescence; bracts and bracteoles 1.5—3 mm long and 1—2 mm wide at the base, triangular. Flowers 4-merous on pedicels 0.5—1 mm long that are obscured by subtending bracts and bracteoles. Hypanthia (at anthesis) subcylindric to narrowly campanulate. Calyx tube obsolete, the calyx lobes 2—2.5 x 1.5 mm, triangular, typically reflexed, calyx teeth 0.5—1.5 mm long, subu- late. Petals 1.5—2 x 0.5—-1 mm, glabrous, white, oblong, conspicuously reflexed at anthesis and early caducous. Stamens isomorphic; filaments 1.5 mm long, glabrous; anthers 1.5 x 0.5 mm, yel- low or white, linear-oblong, + truncate to somewhat emarginate at the apex with a dorsally inclined pore; connective prolonged dorso-basally into a deflexed gland-edged appendage 0.5 mm long. Ovary 4-locular, completely superior but tightly enveloped by the hypanthium, apex consisting of a shallow bowl-like depression with a slightly raised perimeter, copiously but inconspicuously glandular-puberulent. Style 6 mm long, glabrous; stigma punctiform to truncate. Berry 6 x 6 mm, bright orange at maturity, prominently 8-costate. Seeds 0.5 mm long, + triangular in outline with an irregularly rugose testa and a foot-like projection at the narrow end. PHENOLOGY.— Flowering and fruiting intermittently throughout the year. DISTRIBUTION.— Primary and secondary rain forests and cloud forests from north-central Costa Rica disjunctly to western Panama, and southwestern Colombia from 850—1900 m. PARATYPES.— Costa Rica: Alajuela: Reserva Biol6gica Monteverde, Rio Pefias Blancas. Finca de Jestis Rojas, 10°18’N, 84°44’W, 4 Dec. 1989, Bello 1540 (CAS, CR); Upala, Bijagua, El Pilén, subiendo por la margen derecha del Rio El Roble hasta al Volcan Tenorio, 10°41’N, 85°00’20’W, 9 July 1988, Herrera et al. 2037 (CAS, CR). PANAMA: Coclé: Alto Calvario above El Copé, ca. 6 km N of El Copé, Atlantic slope, along trail through forest W off old lumber trail lead- ing to Las Ricas, Limon and San Juan, 8°39’N, 80°36’W, 23 June 1988, Croat 68793 (CAS); El Copé, Division continental arriba de Barrig6n y el aserradero viejo. Camino de caballo a Coclesito, 27 Apr. 1992, Pena et al. 469 (CAS, PMA). Comarca de San Blas: Cerro Brewster, 9°18’N, 79°16'W, 21 Apr. 1985, de Nevers et al. 5452 (CAS); Cerro Habit, vicinity of peak, 78°49’W, 9°23’N, 19 Dec. 1980, Sytsma et al. 2694 (CAS). Chiriqui: Edwin Fabrega Dam and Reserve in Fortuna. Quebrada Arena, 8°45’N, 82°05’W, 15 Jan. 1989, Almeda et al. 6217 (CAS); Gualaca- Chiriqui Grande, 4.8 mi. beyond IRHE facilities at the Dam, 4 mi. N of bridge over Bayano Lake, 8°46’N, 82°16’W, 23 Sept. 1987, Croat 68018 (CAS, MO). Veraguas: along trail to summit of Cerro Tute about 2 mile above the Escuela Agricultura Alto Piedra near Santa Fé, 29 Jan. 1989, Almeda et al. 6490 (CAS, MO, PMA). CoLomsiA: Narifio: Reserva Natural La Planada, munici- pio de Ricuarte, 18 Nov. 1993, Restrepo 776 (US); La Planada Biological Reserve, ca. 7 km S of Chucunez, along trail to Pialapi beyond Quebrada La Calledita, 01°10’N, 77°55’W, 8 Aug. 1990, Luteyn & Sylva 13943 (CAS, NY). Discussion.— Among the 4-merous species of Clidemia that lack formicaria, C. evanescens is a standout because of its densely packed indument of roughened or asperous-headed hairs, con- spicuously 8-ribbed hypanthia (Fig. 4H), small oblong petals (Fig. 4D, F), deflexed gland-edged | | Volume 55, No. 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 100 ij MILE] a a wr 8 ALMEDA: NEOTROPICAL GENUS CLIDEMIA 101 dorso-basal staminal appendages (Fig. 4G), completely superior ovary, and berries that are orange (vs. blue-purple) at maturity. The inflorescence position of this species is variable. It is typically an axillary cluster of dichasia with branching initiated at the vegetative node but at times it can be ter- minal with both lateral buds at a vegetative node elongating and overtopping the inflorescence. Among congeners, C. evanescens, is most like C. densiflora (Standl.) Gleason and it seems likely that these two species are closely related. The latter also has 4-merous flowers, linear-oblong petals, anther connectives that are dorso-basally prolonged into a glandular appendage, and berries that are orange at maturity. In C. densiflora, the flowers are sessile (vs. pedicellate) and mostly fas- ciculate, the merely roughened hypanthial hairs are spreading but not tightly packed to conceal the actual surface as they are in C. evanescens. In addition to these differences, C. densiflora has modally smaller leaf blades [7—14(—18) x 2.5—5.5(—9) cm vs. 9-27 x 3.4—15 cm] and shorter peti- oles (0.3—0.8 cm vs. 1.54.7 cm). ETYMOLOGY.— The specific epithet for this species is from the Latin word evanescens, mean- ing quickly disappearing or fading, in reference to the petals that fall away from the flowers quick- ly at anthesis. Clidemia folsomii Almeda, sp. nov. (Fig. 5) TyPE.— PANAMA: Veraguas: along trail to summit of Cerro Tute about 2 mile above the Escuela Agricultura Alto Piedra near Santa Fé, 29 Feb. 1989, F. Almeda et al. 6488 (holotype: CAS!; isotypes: AAU!, CAS!, INB!, MEXU!, MO!, PMA!). Ramuli teretes sicut petioli laminae supra et subtus inflorescentia hypanthiaque pilis laevibus plerumque 1—2.5 mm longis pro parte glanduliferis modice obsiti et pilis stellatis modice vel sparse intermixtis. Lamina plerumque ad petioli apicem vesiculifera; vesicae 3.5—7.5 cm longae oblongae; Jamina 11—26.5 x 7.5—21 cm cordata vel ovata apice caudato-acuminata 7—9-nervata vel 7—9-plin- ervata. Inflorescentia in foliorum superiorum axillis multiflorae; flores 5-meri bracteolis 0.5—1 x 0.25 mm; calycis tubus 1 mm altus, lobis interioribus 1.5 x 1.5 mm triangularibus, dentibus exte- rioribus subulatis setosis 1.5 mm eminentibus. Stamina isomorphica glabra, antherarum thecae 1—1.5 x 0.5 mm oblongae vel subulatae poro dorsaliter inclinato; connectivo non prolongato basi dorsaliter appendice hebeti ca. 0.2 mm longo descendenti armatum. Ovarium 5-loculare et omnino inferum apice modice glanduloso-puberulo. Little-branched subshrub 1—2 m tall with terete internodes. Uppermost internodes, petioles, adaxial foliar surfaces, elevated primary veins on abaxial foliar surfaces, inflorescence rachis, bracts, bracteoles, pedicels, hypanthia, and calyx teeth moderately covered with smooth (mostly gland-tipped) spreading hairs 1—2.5 mm long with an understory of sessile or stipitate-stellate hairs. Leaves of a pair somewhat unequal in size, otherwise identical in all details and consistently bear- ing paired elongate formicaria 3.5—7.5 cm long extending from the blade base onto the upper por- tions of the petiole. Petioles 4-10 cm long, blades 11—26.5 x 7.5—21 cm, membranaceous and brit- tle when dry, cordate varying to ovate, apex caudate-acuminate, base broadly cordate, margin den- ticulate to crenulate (at least distally) varying to subentire, 7—9-nerved or -plinerved with the inner- most primary pair diverging up to 2 cm above the blade base, the secondary and higher order veins moderately covered with + sessile or subsessile stellate hairs. Inflorescence 6—13 cm long, axillary, laxly branched paniculiform dichasia typically paired at a node; bracteoles 0.5—1 mm long and 0.25 FiGure 4. Clidemia evanescens Almeda. A. habit, x ca. /2: B. representative leaf (abaxial surface), x 4; C. enlargement of abaxial foliar surface (midvein and innermost pair of primary veins), x ca. 2; D. portion of inflorescence (simple dicha- sium with flower at anthesis), x ca. 5: E. bracteole, x ca. 18; F. petal (adaxial surface), x ca. 14; G stamens, dorsal view (left) and profile view (right), x ca. 10; H. berry, x 4; I. seeds, x ca. 48. (A —I from Almeda et al. 5933.) | 102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 mm wide at the middle, linear-oblong. Flowers 5-merous, subsessile or on pedicels to 1 mm long. | Hypanthia (at anthesis) urceolate. Calyx tube ca. 1 mm long, the calyx lobes 1.5 x 1.5 mm, broad- _ ly triangular, collectively spreading and bowl-like; calyx teeth 1.5 mm long, subulate, exceeding and obscuring the calyx lobes. Petals 34.5 x 1.5-2.5 mm, glabrous, translucent white, pink, or | white tinged with pink or purple, narrowly obovate to oblong. Stamens isomorphic; filaments 2 mm long, glabrous; anthers 1—1.5 x 0.5 mm, white, + oblong to + subulate in ventral view, emarginate at the apex with a dorsally inclined pore; connective thickened dorsally and prolonged dorso-basal- ly into a blunt spur ca. 0.2 mm long. Ovary 5-locular, completely inferior; apex elevated into a low + truncate dome surrounding the stylar scar, minutely glandular-puberulent like the adaxial vascu- lar ring of the hypanthium. Style 5-6 mm long, glabrous; stigma punctiform to truncate. Berry 6 x 6 mm, blue at maturity. Seeds 0.5 mm long, obovoid to + triangular in outline with a verruculose to rugulate testa. PHENOLOGY.— Flowering and fruiting from November through June. DISTRIBUTION.— Cloud forests, elfin forests, and stream banks in western Panama from 800 to 1300 m. PARATYPES.— PANAMA: Chiriqui: vicinity of Fortuna Dam along trail from highway near Forestry Nursery down to Rio Hornito, 8°45’N, 82°15’W, 27 Apr. 1988, McPherson 12510 (CAS); distrito Boquete, Fortuna Dam site, 7 Feb. 1985, van der Werff & van Hardeveld 6607 (CAS). Coclé: Cerro Gaital, east slope and ridges leading to the summit, 8°40’N, 80°07’W, 24 Feb. 1988, Almeda et al. 5899 (AAU, CAS, INB, MEXU, MO, PMA); Alto Calvario, cloud forest above sawmill on continental divide, 5.2 miles above El Copé, 6 Dec. 1979, Croat 49152 (CAS): Alto Calvario above El Copé, ca. 6 km N of El Copé, along trail which leads down to Las Ricas, Limon and San Juan, 22 June 1988, Croat 68742 (CAS); Continental Divide above El Copé, 8°38’N, 80°39’W, 27 Nov. 1985, de Nevers et al. 6407 (CAS); Cerro Gaital, elfin forest, 8°37’N, 80°6’W, 26 Nov. 1984, de Nevers & Charnley 4388 (CAS); mountains N of El Valle de Anton, 28 May 1967, Lewis et al. 1771 (MO); El Copé, Divisi6n continental arriba de Barrig6n y el aserradero viejo, camino de caballo a Coclesito, 27 Apr. 1992, Pena et al. 467 (PMA); hills just N of El Valle, 14 Aug. 1971, Wiehler & Dressler 71287 (US-2 sheets). Panama: Cerro Campana near the FSU field station trail up the mountain, 9 Aug. 1970, Kennedy & Luteyn 435 (CAS, DUKE, US). Veraguas: trail to Reserva Biolo6gica Serrania de Tute and the summit of Cerro Tute about 0.7 km beyond the Escuela Agricola Rio Piedra just outside Santa Fé, 18 Feb. 1996, Almeda et al. 7614 (CAS, PMA); 6.4 km outside Santa Fé on the road that passes the agriculture school, 5 May 1977, Folsom 2955 (CAS); Cerro Tute, along ridge trail towards summit, 08°30/N, 81°07’W, 21 Mar. 1987, McPherson 10743 (CAS). DIscussiIoNn.— Clidemia folsomii is one of nearly 30 taxa (see Appendix I) in the genus Clidemia that produce formicaria or domatia to shelter ants. In Clidemia, these formicaria consist of paired inflated pouch-like outgrowths borne at the branch nodes, at the base of the leaf blade, on the petioles, or extending along a portion of the petiole and leaf blade. The ants that inhabit these domatia presumably defend the plants from herbivore attack but the exact nature of the symbiotic relationship has not been studied for the majority of Clidemia species (Benson 1985). Ants collect- ed in the formicaria of one population of this species (Almeda 5899) were identified as Solenopsis sp. (subgenus Diplorhoptrum) by P.S. Ward. According to Ward, the ants collected in these doma- tia were probably nesting opportunistically; there are a number of generalist, arboreal ant species in this subgenus. The size, shape, and placement of formicaria typically provide diagnostic features of taxonom- ic importance. The formicaria of C. folsomii, for example are elongate, produced at the very base of the leaf blade, and extend onto the distal half of the petiole (Fig. SA, B). This together with the — os ALMEDA: NEOTROPICAL GENUS CLIDEMIA 103 bbb ha te i: x UY WR o cal aN : Wye Ryne’ sree aA ay a road ee Oost bef Z Fy Ficure 5. Clidemia folsomii Almeda. A. habit, x 4; B. representative leaf (abaxial surface), x ca. 's; C. flower bud, x ca. 5: D. petal, x 12; E. stamens, profile view (left), dorsal view (middle), and ventral view (right), x ca. 12; F. berry, x 6, G. seeds, x 36. (A—-F from Almeda et al. 6488; G from Almeda et al. 5899.) 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 4 | large cordate leaf blades (Fig. 5B), axillary, paired, and laxly branched dichasia at each node (Fig. — 5A), indument of spreading (mostly gland-tipped) hairs with an understory of stellate hairs, and © well-developed calyx teeth (Fig. 5F) make this species unmistakable among congeners. ! Morphologically, Clidemia folsomii does not appear to be particularly close to any of the other formicarial species in the genus. It is superficially most like C. plumosa (Desr.) DC., a non-formi- carial species of Jamaica, Hispaniola and St. Thomas. Both species have elongate petioles, large cordate or ovate leaf blades, paired laxly branched dichasia at the upper nodes, a copious indument of spreading hairs, and elongate setose calyx teeth. In C. plumosa, the conspicuous pubescence consists of hairs that are eglandular (vs. mostly gland-tipped), the understory indument consists of small glands that look granular (vs. stellate hairs), and the flowers are 4-merous (vs. 5-merous). ETYMOLOGY.— This species is named for James P. Folsom (b. 1950), student of neotropical Orchidaceae and veteran collector of Panamanian plants who first collected fertile material of this species in 1977. Clidemia inopinata Almeda, sp. nov. (Fig. 6) TypE.— Mexico: Chiapas: Municipio La Trinitaria, Ejido Cuauhtemoc, Km 25, 20 Aug. 1984, Méndez Ton 7868 (holotype: CAS!, isotype: MEXU!). Ramuli rotundato-quadrangulati sicut petioli inflorescentiaque pilis laevibus erectis vel reflex- is 2-8 mm longis. Lamina 7—24.5 x 4.2—13.5 cm ovata vel ovato-oblonga apice acuminata basi obtusa vel rotundata, supra modice longo-setosa pilis gracilibus laevibus 1-4 mm longis, subtus in nervis primariis et secundartis dense longo-setosa in nervulis superficieque modice vel sparse glan- dulosa, 5—7-nervata. Inflorescentia primum terminalis demum lateralis; flores 5-meri, bracteolis 3.5—8 x 2-3 mm. Calycis lobi interiores 0.75—1 x 0.75—1 mm suborbiculares, dentibus exterioribus 3 mm longis subulatis vel oblongis apice setiferis. Stamina isomorphica glabra, antherarum thecae 2.5 x 0.5 mm oblongae poro dorsaliter inclinato; connectivum nec prolongatum nec appendicula- tum. Ovarium 5-loculare et 34 inferum apice conico sparsissime glanduloso-puberulo. Shrub 1—2 m tall. Internodes rounded-quadrate; uppermost branches, petioles, and inflores- cence axes copiously hirsute with spreading to somewhat reflexed smooth hairs 2-8 mm long. Leaves of a pair + equal to unequal in size; petioles 1.1—-8 cm long; blades 7—24.5 x 4.2—13.5 cm, ovate to ovate-oblong, apex acuminate, base obtuse to broadly rounded, margin denticulate to ser- rulate and ciliate, 5—7-nerved, uniformly setose adaxially with smooth spreading hairs 14 mm long, the abaxial surface also copiously beset with spreading smooth hairs 1-3 mm long on the ele- vated primaries and higher order veins but only the primary and secondary veins sparingly under- lain with minute deciduous glands. Inflorescence 4-9 cm long, a pseudolateral modified dichasi- um openly and divaricately branched from the base; bracts and bracteoles 3.5—8 x 2—3 mm, persist- ent, lanceolate to elliptic-lanceolate, glabrous on both surfaces with hirsute-ciliate margins. Pedicels 1-2 mm at anthesis lengthening to 34 mm on the infructescence. Hypanthia oblong to subcylindric, 4 mm long from the base to the torus (vascular ring), copiously covered with smooth spreading hairs 3—8 mm long underlain with a sparse cover of minute and inconspicuous glands. Calyx tube 0.25 mm long, the calyx lobes 0.75—1 x 0.75 mm, + semicircular, glabrous and entire to subentire; calyx teeth 5 in number, 3 mm long or 4.5—6 mm long including the terminal hair, lin- ear-oblong to subulate, beset with an indument like the hypanthium. Petals 5 in number, 5—7 x 2.5 mm, glabrous, white, oblong to oblong-obovate. Stamens 10, isomorphic; filaments glabrous, 2 mm long; anthers 2.5 x 0.5 mm, white, linear-oblong, + emarginate at the apex with a dorsally inclined pore; connective conspicuously thickened dorsally and barely prolonged below the anther ALMEDA: NEOTROPICAL GENUS CLIDEMIA 105 sacs and unappendaged. Ovary %*4-inferior, 5-locular, apex elevated into a low truncate + crateri- form cone with one or few spreading hairs and/or few minute glandular hairs. Style 5 mm long, straight, glabrous; stigma capitellate when receptive. Berry 6-8 x 4-5 mm, red to purple when mature. Seeds ca. 0.4 mm long, ovoid, the testa uniformly asperulate. PHENOLOGY.— Flowering collections have been made from December through July; fruiting collections from April through January. DISTRIBUTION.— Understory shrub of primary and secondary forests, forest margins, clear- ings, and disturbed sites near rivers from southeastern Mexico (Veracruz) disjunctly south through Guatemala to Honduras from 180-1230 m. PARATYPES.— Mexico: Chiapas: municipio of La Trinitaria, 10 km east northeast of Dos Lagos above Santa Elena, 19 Dec. 1980, Breedlove 48532 (CAS); 15 km east-northeast of Dos Lagos above Santa Elena, 29 Dec. 1981, Breedlove 56636 (CAS); municipio La Trinitaria, 10 km east-northeast of Dos Lagos above Santa Elena, 19 Jan. 1982, Breedlove & Almeda 57505 (CAS); municipio La Trinitaria, km 25, camino Lagos de Montebello—Acayal, 3 May 1985, Espejo et al. 1660 (MEXU). Oaxaca: municipio Santa Marfa Chimalapa, San Antonio Nuevo Paraiso, 13 May 1995, Torres 669 (CAS); municipio Sta. Maria Chimalapa, ca. 15 km ESE de Sta. Maria, 16°52’N, 94°34’30’W, 28 May 1987, Hernandez 2520 (US); “Rio Uluapan”, 8 km al E de San Bartolomé Ayautla, carretera Huautla—Jalapa de Diaz, Distr. Teotitlan, 20 July 1982, Torres et al. 855 (CAS, MEXU); municipio Matias Romero, 7.2 km al O de Esmeralda, en la terraceria La Laguna—Sarabia, luego 2.6 km al S por camino, 9 Mar. 1982, Wendt et al. 3640 (CAS, US). Tabasco: municipio Huimanguillo, camino del ejido villa de Guadalupe a la torrey microondas, 6 Novy. 1994, Guadarrama et al. 4132 (MEXU). Veracruz: municipio Catemaco, lado NE de Lago Catemaco en Cerros al E de Coyama, 26 Oct. 1971, Beaman 5180 (CAS, MEXU); Bastonal—Sierra Santa Marta road, ca. 14 km E of Lago Catemaco, 29 May 1981, Gentry et al. 32431 (CAS, MO); 11 km S de Tebanca, Camino a Bastonal, 26 Apr. 1982, /barra 96 (CAS); municipio Hidalgotitlan, campamento La Laguna, along Rio Las Cuevas and nearby pastures, 17°16’N, 94°31’W, 6 Mar. 1984, Taylor 390 (CAS, F, XAL); municipio Hidalgotitlan, desde el Poblado 6 al sur por la brecha y la vereda al horcajo oriental del Rio Cuevas, 17°15’N, 94°30’W, 17 July 1980, Wendt et al. 2603 (CAS, US). GUATEMALA. Alta Verapaz: Chapultepec Farm, km 62 of Coban—Sebol Road, 25 May 1964, Contreras 4801 (CAS, LL); 7 miles up road to Oxec along road which turns off Highway 7E between Tucurt and E] Estor ca. 6 km NE of Panzos, 20 July 1977, Croat 41649 (CAS, MO); 2 km al S de Jolomylix, Teleman, Panzos, Sierra de las Minas, 20 July 1988, Martinez et al. 22958 (CAS, MEXU). Izabal: Montafias del Mico, 7-8 km W of Santo Tomas de Castilla on road to microwave tower, 19 Aug. 1988, Stevens et al. 25598 (MEXU). Honpuras. Atlantida: southern boundary of Lancetilla Valley (Near Tela), on ridge separating Lancetilla watershed from that to the SW of San Francisco, halfway to Cerro Pefia Blanca, 15°41’30’N, 87°28’30’W, 9 Nov. 1988, MacDougal et al. 3416 (CAS, MO). Comayagua: margins of Lake Yojoa near Pito Solo, 9 Aug. 1948, Williams & Molina 14622 (MO). Yoro: along road from San José de Texiguat to Campo Nuevo in Cordillera Nombre de Dios, ca. 28 km SW of Tela—La Ceiba hwy. (#13), ca. 15°29’N, 87°27 W°, 27 Apr. 2000, Daniel & Araque 9496 (CAS, EAP); slopes above E part of San José in the Rio Lean Valley, between Rio Texiguat and Rio Guan Guan, slopes of Cordillera de Nombre de Dios, 6 Nov. 1988, MacDougal et al. 3283 (CAS, MO). DIsCcussION.— Diagnostic characters of C. inopinata include the openly and divaricately branched inflorescence (Fig. 6A), the conspicuous and persistent bracteoles (Fig. 6C), the deeply pigmented (blackish) hypanthia that are copiously covered with smooth spreading hairs (Fig. 6F) underlain with a sparse cover of minute inconspicuous glands, the white anther thecae, and the uni- formly asperulate seed coat (Fig. 6G). Clidemia fulva Gleason has an aspect that is very reminis- 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 FIGURE 6. Clidemia inopinata Almeda. A. habit, x ‘4; B. representative leaf (abaxial surface), x 2; C. bracteole, x 6; D. petal (adaxial surface), x 8; E. stamens, profile view (left) and ventral view (right), x ca. 7; F. berry, x ca. 9; I. seeds, x ca. 56. (A from Breedlove & Almeda 57505; B from the holotype; D, E and G from Breedlove 56636; C and F from Breedlove 48832.) ALMEDA: NEOTROPICAL GENUS CLIDEMIA 107 cent of C. inopinata and the latter has typically been identified as the former. Both species share similar inflorescence architecture with persistent bracts and bracteoles, a copious indument of smooth spreading hairs, elongate calyx teeth, and white petals and anther thecae. In C. fulva, how- ever, the leaf base is often unequal (vs. never unequal), pedicels on the infructescence are 4-8 mm long (vs. 34 mm), hypanthial hairs are 2.5—4 mm long (vs. 3-8 mm), the seeds are 0.75 mm long (vs. 0.4 mm), and the seed coat is smooth (vs. uniformly asperulate). Clidemia fulva also has a more limited geographic and elevational distribution than C. inopinata. It occurs in Chiapas, Mexico, and in a limited region of adjacent Guatemala (Alta Verapaz, Izabal, and Peten) at elevations of 200-500 m. ETyMOoLOoGy.— The epithet for this species is derived from the Latin word inopinatus, unex- pected, alluding to the fact that it has long gone unrecognized under the name of a closely related species. Clidemia lanuginosa Almeda, sp. nov. (Fig. 7) TypE.— PANAMA: Bocas del Toro: Fortuna Dam area along continental divide trail bordering Chirigui Province, 08°45’04’N, 82°15’04’W, elev. 1200-1300 m, 10 Mar. 1988, F: Almeda et al. 6070 (holotype: CAS!; isotypes: INB!, MO!, PMA'!). Ramuli teretes sicut petioli inflorescentiaque modice pilis laevibus vel usque ad 3—5 mm obsi- ti et dense persistenterque lanuginosa pilis gracillimis laxis intermixtis. Lamina 8.5—22 x 5—12.5 cm ovata vel ovato-elliptica apice acuminata basi rotundata supra modice setosae pilis laevibus 3—6 mm longis, subtus in venis setosa et modice setulosa pilis caduco-glanduliferis, 7—9-plinervata; formicaria ca. 2—5.5 cm longa in laminarum basibus omnino immersa. Inflorescentia primum ter- minalis demum lateralis; flores 4 (-5)-meri, bracteolis 1-2 x 0.25 mm. Calycis lobi interiores 0.25 x 0.5—0.75 mm suborbiculares, dentibus exterioribus 0.5-4 mm longis subulatis apice setiferis. Stamina isomorphica glabra, antherarum thecae 1.75—2 x 0.2 mm subulatae poro dorsaliter incli- nato; connectivo nec prolongatum nec appendiculatum. Ovarium (3—) 4 (—5)-loculare et omnino inferum apice glabro. Little-branched subshrub 0.5—2 m tall, the terete upper internodes, petioles, and inflorescence rachis moderately covered with smooth spreading hairs 3—5 mm long with a dense understory of deciduous appressed somewhat woolly (often gland-tipped and crisped) hairs. Leaves of a pair equal to unequal in size, otherwise identical in all details and consistently bearing paired elongate inflated formicaria 2—5.5 cm long on the adaxial base of the blade; petioles 0.5—4 cm long; blades 8.5—22 «x 5-12.5 cm, membranaceous and brittle when dry, ovate to ovate-elliptic, apex acuminate, base broadly rounded, margin subentire to denticulate, 7—9—plinerved, the paired primary veins arising at successive points above the blade base with the inner-uppermost primaries diverging from the median vein 24.5 cm above the blade base, the adaxial surface moderately and uniform- ly covered with smooth spreading hairs 3-6 mm long, the primary and higher order veins on the abaxial surface beset with smooth spreading hairs 2-5 mm long and an understory of minute decid- uous glandular hairs. Inflorescence 2-8 cm long, a paniculiform dichasium that is initially termi- nal but then becomes pseudolateral with growth and elongation of lateral branches; bracteoles 1—2 mm long (including terminal hair) and 0.25 mm wide, subulate to oblong-subulate, sparingly beset with minute glandular hairs on the abaxial surface. Flowers 4 (—5)-merous on pedicels 0.5—1 mm long. Hypanthia (at anthesis) cylindric to cylindric-campanulate, sparsely covered with smooth spreading hairs 1-3 mm long and an understory of appressed glandular hairs. Calyx tube obsolete, the calyx lobes 0.25 x 0.5-0.75 mm, semicircular to rounded-triangular, tardily deciduous, hyaline, | 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4° deciduously glandular-puberulent abaxially and glandular-ciliolate at the margins; calyx teeth | 0.5—4 mm (excluding apical hair), subulate, sparingly beset with smooth spreading hairs 2 mm long | and an understory of minute glandular hairs. Petals 2-4 x 1 mm, glabrous, white or pink, oblong | to oblong-obovate. Stamens isomorphic; filaments 1.75—2 mm long, glabrous; anthers 1.75—2 x 0.2 © mm, yellow, subulate with a dorsally inclined pore; connective thickened dorsally but not pro- longed below the thecae and unappendaged. Ovary (3—) 4 (—5)-locular, completely inferior; apex elevated into a short cone (0.25 mm) and vaguely lobulate collar at anthesis that disappears on mature fruits. Style 4 mm long, glabrous; stigma punctiform to truncate. Berry 2.5—3 x 2.5-4 mm, blue at maturity. Seeds 0.5 mm long, triangular, the convex face somewhat angular with a densely tuberculate testa. PHENOLOGY.— Flowering material has been collected from February through November; fruiting specimens have been collected in March, June, July, September, and November. DISTRIBUTION.— Local and uncommon, often in deep shade of cloud forests and rain forests from southeastern Costa Rica to Panama (Bocas del Toro east to Darién) at 300-1450 m. PARATYPES.— CosTA RicA: Limon: Cant6n de Talamanca, R.I. Talamanca, Cordillera de Talamanca, Alto Urén, 9°27’00"N, 82°59°30’W, 24 Sep. 1994, Gallardo & Lépiz 281, (INB); Cant6n de Talamanca, R.I. Talamanca, Cordillera de Talamanca, Alto Urén, 9°21’35’N, 82°59’45’”W, 13 Sep. 1992, Gallardo 88 (INB); Reserva Indigena Talamanca, Sukut, desembo- cadura del Rio Sukut en el Rio Urén, camino al sureste, hacia Purisqui, 9°23’30’N, 82°58’00’W, 7 July 1989, Hammel et al. 17571 (CAS, CR, INB, MO). PANAMA: Bocas del Toro: above Chiriqui Grande on a side road about 10 miles below the Continental Divide about 2 '/ miles east on that road, 8°55’N, 82°10’W, 19 Jan. 1989, Almeda et al 6334 (CAS, PMA): road from Fortuna Dam to Chiriqui Grande, 10 miles from Continental Divide, 8°51’N, 82°10’W, 2 Aug. 1984, Churchill 5933 (CAS): Oleoducto Road, near Continental Divide. Fortuna Dam area, 8°48’N, 82°12’W, 5 Feb. 1984, Churchill et al. 4494 (CAS): después de la Divisi6n Continental, en la carretera que va de Fortuna a Chiriqui Grande, 27 Aug. 1993, Correa et al. 9554 (PMA); along road between Fortuna Dam and Chiriqui Grande, 7.3 mi. N of bridge over Fortuna Dam, 3.2 mi. N of Continental Divide, 8°45’N, 82°15’W, 10 Mar. 1985, Croat & Grayum 60220 (CAS); road from Fortuna Dam to Chiriqui Grande, 3.1 mi. N of Continental Divide, 8°50’N, 82°15’W, 10 Mar. 1985, McPherson 6770 (CAS); Edwin Fabrega Dam and Reserve in Fortuna. Trail to the hydrological station on the Rio Hornito between Quebrada Nelson and Quebrada Mono, 7 Feb. 1996, Almeda et al. 7521 (CAS, CR, MO, PMA); along road between Gualaca and Fortuna Dam, at 8.3 mi. NW of Los Planes de Hornito, 9 Apr. 1980, Antonio 4150 (CAS); Fortuna, camino de Quebrada Bonita. Hacia el este del Rio Chiriqui, 7 Apr. 1987, Valdespino et al. 563 (CAS, PMA); Fortuna, plots 5—8 de Jaime Cavelier, frente al vivero del IRHE, 26 Apr. 1988, Valdespino et al. 846 (CAS, PMA). Chiriqui: S. O. del campamento de Fortuna (Hornito), sitio de presa subiendo hasta la finca Pittit, bordeando el bosque y regresando por el lado S del campamento, 14 Aug. 1976, Correa et al. 2424 (US); Fortuna Dam area, around creek to . of road drawing into Rio Hornito S of reservoir, 8°43/N, 82°14’W, 14 June 1984, Churchill 5375 (CAS); Fortuna Dam area, on Kaolin hill, just N of reser- voir, 31 July 1984, D’Arcy et al. 15902 (US). Darién: trocha Rio Frio hacia la cima mas alla de Rancho Tuque (3 horas de camino), 22 Sep. 1989, Aranda et al. 949 (CAS, PMA). Veraguas: Distrito de Sante Fé, Serrania de Tute, 8°33’N, 81°07’W, 5 July 1996, Aranda et al. 2653 (CAS, PMA); in forest below summit of Cerro Arizona (Cerro Tute), above Santa Fé, 22 Apr. 1980, Hammel & Kress 8569 (CAS); ridge east of Cerro Tute, NW of Santa Fé, 25 Oct. 1975, Dressler 5203 (US); about 20 km NW of Santa Fé near the Continental Divide, 29 Mar. 1975, Kennedy & Dressler 3417 (US); trail to Cerro Tute, above Escuela Agricola Alto de Piedra, just W of Santa Fé, 8°32’N, 81°07’W, 5 June 1982, Knapp and Dressler 5417 (CAS). ALMEDA: NEOTROPICAL GENUS CLIDEMIA 109 SvvXs NY SY \ AAA iS) yyy .\ Ni Ficure 7. Clidemia lanuginosa Almeda. A. habit, x ca. /%2; B. representative leaf (adaxial surface), x '; C. representa- tive leaf (abaxial surface), x '4; D. portion of inflorescence rachis, x 9; E. bracteoles, x 17; F. petal (adaxial surface), x 14; G. stamens, profile view (left) and dorsal view (right), x 11; H. berry, x ca. 9; I. seeds, ca. 56. (A — G from Almeda et al. 6070; H from Churchill 5375; 1 from D’Arcy et al. 15902.) 110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No.4 — DIscussIOn.— Clidemia lanuginosa is readily recognized by its 7—9-plinerved leaf blades with paired primary veins arising at successive points above the blade base (Fig. 7C), didymous formicaria (Fig. 7B) restricted to the adaxial base of the blade, and oblong, glandular-puberulent bracteoles that are tipped with a prominent apical hair (Fig. 7E). The indument on upper intern- odes, petioles, and the inflorescence is also distinctive. It consists of smooth spreading hairs 3—5 mm long with a dense understory of deciduous appressed woolly (often gland-tipped and some- what crisped) hairs. Clidemia lanuginosa is variable in characters that are typically constant in many other con- geners. For example, the calyx lobes are persistent on some collections but tardily deciduous on others. Floral merosity and ovary locule number are also variable. The isomerous flowers of this species are typically 4-merous with a 4-locular ovary but the flowers on inflorescences of individ- ual plants in populations from both Costa Rica and Panama can be prevailingly 4-merous with some that are 5-merous, and the ovary can vary from 3- to 5-locular. None of this variation appears to correlate with other vegetative or reproductive features. Several collections from Cerro Tute, Panama, stand out because they have flowers that are consistently 5-merous with shorter calyx lobes and formicaria that occupy the blade base and about half of the distal portion of the petiole. These anomalous collections may represent a distinctive taxon; they are here excluded from the circumscription of C. Januginosa pending additional study. ETYMOLOGY.—The specific epithet is derived from the Latin word, lanuginosus, woolly, to emphasize the copious understory of woolly hairs on upper internodes, petioles, and the inflores- cence rachis. Clidemia pectinata Almeda, sp. nov. (Fig. 8) TyPE.— PANAMA. Bocas del Toro: along oleoducto road between continental divide and Chiriqui Grande, 21 miles S of Chiriqui Grande, 8°50—55’N, 82°9-15’W, elev. 850 m, 3 May 1985, Hammel 13748 (holotype: MO!). Ramuli teretes sicut folia supra hypanthiaque glabri. Petioli 7-14.3 cm longi dense setosi pilis laevibus 4-9 mm longis. Lamina 20—20.5 x 10—13 cm ovato-elliptica apice breviter hebeti-acumi- nata vel hebeti-acuto basi rotundata, subtus sparse glanduloso-setulosa alioqui glabra, 7-plinerva- ta. Inflorescentia primum terminalis demum lateralis. Pedunculus ad 18 cm longus cum pedicellis modice pilis glanduliferis inconspicuis obsitus; flores 5-meri; bracteolae 1-2 mm longae. Calycis tubus ca. 0.1 mm longus, lobis interioribus 0.5 x 1 mm oblatis, dentibus exterioribus 0.5 mm longis triangularibus. Stamina isomorphica glabra, antherarum thecae 2 x 0.25 mm oblongae poro dor- saliter inclinato; connectivum nec prolongatum nec appendiculatum. Ovarium 5-loculare % infer- um apice glabro. Shrub to 1.5 m tall with terete glabrous internodes, the nodes of uppermost branches and veg- etative buds deciduously setose with smooth spreading hairs 2-6 mm long. Leaves of a pair essen- tially equal in size; petioles 7—14.3 cm long, glabrous on the abaxial surface but densely setose on the adaxial surface with smooth spreading hairs 4-9 mm long that appear pectinate (one-sided) and directed adaxially when dry; blades chartaceous and brittle when dry, 20—20.5 x 10-13 cm, entire but coarsely ciliate, ovate-elliptic, apex short-acuminate to acute, base broadly rounded, 7-plin- erved, glabrous adaxially, the abaxial surface appearing glabrous but actually minutely and some- what deciduously glandular-papillose. Inflorescence a pseudolateral, pendulous and long-peduncu- late (to 18 cm long) corymbiform-umbelliform cluster of dichasia (34 cm long including second- ary peduncles); the primary peduncle deciduously setose with smooth hairs 0.75—2 cm long; the 111 ALMEDA: NEOTROPICAL GENUS CLIDEMIA oe ZX\\\ ne ante et VA 9% ae FiGURE 8. Clidemia pectinata Almeda. A. habit, ca. 2; B. representative leaf (abaxial surface), x '4; C. petal (adaxial le view (left) and ventral view (right), x 12; E. berry, x 4; F. seeds, x ca. 34. (A— D from the holotype; E and F from Correa et al. 9855.) surface). x 11; D. stamens, profi 112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 4 — secondary peduncles, pedicels, and very young hypanthia sparsely to moderately beset with minute — inconspicuous glands; bracts and bracteoles 1-2 mm long (including apical hair) and 0.25-0.5 mm — wide, subulate, essentially glabrous. Flowers 5-merous on pedicels 1-2 mm long. Hypanthia (at anthesis) narrowly campanulate, beset with minute glandular hairs when young but essentially glabrous at maturity. Calyx tube ca. 0.1 mm long, the calyx lobes 0.5 x 1 mm, oblate, hyaline and glabrous; calyx teeth 0.5 mm long, triangular. Petals 3 x 2 mm, obovate, glabrous. Stamens isomor- phic; filaments 1.5 mm long, glabrous, abruptly constricted distally just below the thecae; anthers 2 mm long, ca. 0.25 mm wide, oblong, truncate apically with a dorsally inclined pore; connective thickened dorsally but neither prolonged nor appendaged below the thecae. Ovary 5-locular, %4- inferior, apex glabrous and smooth around the stylar scar. Style 3 mm long, straight and glabrous; stigma truncate. Berry 4x5 mm, reddish-purple and conspicuously lobed at maturity. Seeds 0.5—0.75 mm long, bluntly triangular and angular with a densely tuberculate testa. PHENOLOGY.— Only two collections of the species are known; the May collection is in mature bud, the August collection is in mature fruit. DISTRIBUTION.— Evidently local and rare in rain forests along stream banks on the Caribbean versant of western Panama in the region extending from Fortuna to Chiriqui Grande at 600-850 m. PARATYPE.— PANAMA: Bocas del Toro: después de la Division Continental en la carretera que va de Fortuna a Chiriqui Grande, 27 Aug. 1993, Correa et al. 9855 (CAS, PMA, SCZ). DISCUSSION.— Clidemia pectinata 1s perhaps the rarest and surely one of the most distinctive species of Clidemia. The combination of glabrous internodes (Fig. 8A), adaxial leaf blades, and mature hypanthia (Fig. 8E), pectinately pubescent petioles (Fig. 8B), and pendent long-peduncu- late inflorescences (Fig. 8A) are unknown in any other species of Clidemia. The limited material available for study has precluded an assessment of variation and I was unable to relocate even ster- ile individuals of this species during field work in Panama in early 1996. Measurements of floral parts were taken on large but unexpanded flower buds of the holotype so even the size of most flo- ral organs are likely to be larger when material is collected with expanded flowers. ETYMOLOGY.— The epithet for this species is derived from the Latin word, pectinatus, 1.e., with narrow close-set divisions like a comb. This refers to the posture and density of the petiolar hairs that are one-sided and directed adaxially when dry. Clidemia quinquenervia (Mill.) Almeda, comb. nov. BASIONYM: Melastoma quinquenervium Mill., Gard. Dict. ed. 8, sp. 15. 1768. Octopleura quinquenervia (Mill.) Triana, Trans. Linn. Soc. London 28:145. 1871. Ossaea quinquenervia (Mill.) Cogn., DC. Monogr. Phan. 7:1064. 1891. Type: Exact locality not legible, Herb. Miller s.n. (holotype: BM!). Clidemia ? decurrens Beurl., Bid. Portobellos Flora. Kongl. Vetensk, Acad. Handl. 40:127. 1854. syn. nov. TyPE: Panama. Porto Bello, in silvis ad viam versus Panama, Apr. 1826, Billberg 280 (holotype: S, photo: MO!). Ossaea diversifolia (Bonpl.) Cogn., Bull. Acad. Roy. Sci. Belgique Ser. 3, 14:968. 1887. Basionym: Melastoma diversifolium Humb. & Bonpl., Melast. 138, t. 59. 1816. Type: Colombia. Rio Magdalena, Bonpland 1617 (holotype: P!; isotypes: P! - 3 sheets, US!). For additional synonymy see Wurdack (1971:128). Shrub 0.5—2 (—6) m, the terete upper internodes and branchlets, petioles, primary abaxial leaf veins, and inflorescence axes covered with a dense indument of stipitate or subsessile clavate asperous-headed hairs. Leaves of a pair somewhat unequal in size, blades 10—24 x 5—15 cm, broad- ly elliptic to ovate, 5—7 (—9)-plinerved, abruptly narrowed at the base and narrowly decurrent along the petiole, apex obtuse, acute, or short-acuminate, the margins ciliolate-denticulate, moderately to sparsely setulose or glabrate adaxially, sparsely and deciduously stellulate-furfuraceous and ALMEDA: NEOTROPICAL GENUS CLIDEMIA 113 resinous-glandular on the secondary, higher order veins and blade surface abaxially. Inflorescence 2—4 cm long, a modified dichasium mostly divaricately branched from the base; flowers 5-merous on pedicels 0.25—0.5 mm long; the bracteoles mostly 0.5 mm long, subulate, persistent. Hypanthium bluntly 10-costate, deciduously resinous-glandular, occasionally with a sparse admix- ture of spreading simple hairs. Calyx vaguely undulate, ca. 0.25 mm long but concealed by the setulose exterior teeth that are 0.5—1 mm long. Petals 0.75—1.25 x 0.75—1 mm, obovate-oblong, densely papillose-furfuraceous on both surfaces with a projecting infra-apical, abaxial tooth. Filaments 1—1.5 mm long, anthers 1.5—2 mm long, yellow with a truncate to ventrally inclined api- cal pore; connective prolonged dorso-basally into a gland-edged spur ca. 0.25 mm long. Ovary 5- locular, completely inferior, apex somewhat depressed, minutely and deciduously resinous-glandu- lar puberulent; berry 4-5 x 2.5 mm when dry, compressed, initially white to pink-purple but turn- ing blue-black. Seeds 0.5 mm long, the testa smooth. PHENOLOGY.— Flowering and fruiting throughout the year. DISTRIBUTION.— Uncommon to locally common in gallery forests, semideciduous forests, rain forest margins, river or stream banks, pasture margins, and disturbed sites, often in deep or par- tial shade from Honduras south through southern Central America to Colombia, Venezuela, and Ecuador at 0O—700(—1400) m. REPRESENTATIVE SPECIMENS EXAMINED.— HONDURAS: Gracias a Dios: Ahuas Bila, 200 km SO de Puerto Lempira, orilla del Rio Wanki, Coco 0 Segovia, 5—13 May 1985, Nelson & Cruz 9345 (CAS). NICARAGUA: Jinotega: Macizos de Penas Blancas, 5 May 1976, Neill 249 (CAS); Zelaya: Sector de Negro Was, entre El Empalme y Rosita, ca. 13°45’ N, 84°25’W, 5 Sep. 1984, Ortiz 2123 (CAS). Costa Rica: Cartago: Atirro, 1894 (w/out exact date), Donnell Smith 4775 (US). Heredia: La Selva, near Puerto Viejo, 21 May 1972, Opler 826 (US). Limon. Ca. 5.5 km W of Limon off road to Siquirres, 27 July 1977, Almeda et al. 3261 (CAS, CR); ca. 7 km SSW of Home Creek on the road to Bribri, 26 July 1977, Almeda et al. 3256 (CAS, CR). Puntarenas: Cant6n de Golfito, P. N. Corcovado, Peninsula de Osa, Estaci6n Sirena, Sendero a Rio Claro, 14 Oct. 1993, Aguilar 2529 (CAS, INB, MO); disturbed sites above Golfito along the road to the television tower, 16 July 1977, Almeda et al. 3094 (CAS, CR); Reserva Absoluta Cabo Blanco, Estaci6n Cabo Blanco, 9°35/N, 85°06’W, 4 Nov. 1991, Chavarria 305 (CAS, INB, MO); Punta Catedral, ca. 7 km SE of Quepos, 9°22.5’N, 84°09’W, 20 Aug. 1985, Grayum & Sleeper 5896 (CAS, CR, MO); Nicoya Peninsula, canyons and ridges toward Punta Georgia, Punta Blanca trail, 23 Aug. 1995, Sanders et al. 17706 (CAS, UCR); Reserva Bioldgica Carara, Sendero Quebrada Bonita-Bijagual, 9°46’10’N, 84°33’50’W, 8 Dec. 1989, Zuniga & Jiménez 18 (CAS, CR, INB). San José: Zona Protectora El Rodeo Ciudad Coldén, 9°54’N, 84°16’W, Varela 241 (CAS, INB, MO). PANAMA: Bocas del Toro: Lincoln Creek, 28 Jan. 1921, Carleton 87 (US); Changuinola to 5 miles S at junction of Rio Changuinola and Terebe, 17-19 Dec. 1966, Lewis et al. 833 (F, MO, PMA). Canal Area: Madden Forest (Parque Nacional Soberania), Las Cruces Trail, 3.6 mi N of Gamboa road turn-off, 23 July 1994, Croat & Zhu 77060 (CAS, MO, PMA); Barro Colorado Island, Bat Cave, 18 Sep. 1982, Schmalzel & Schupp 955 (CAS, MO). Chiriqui: vicinity of Puerto Armuelles, 28-31 July 1940, Woodson & Schery 855 (MO). Coclé: forest on hills above road 18 km past Sardinilla on way to Nombre de Dios, 2 Aug. 1974, Croat 26096 (MO). Colén: north of Diamante, ridge NW of aban- doned mine on Quebrada de la Mina, 9°24’N, 79°35’W, 11 Jan. 1984, Churchill & de Nevers 4216 (CAS, MO, PMA); 1-2 km from the Portobelo Highway up the Rio Guanche, 9°30/N, 79°40’W, 17 Feb. 1982, Knapp 3641 (CAS). Comarca de San Blas: Aligandi area along trail from ocean to waterfall on river, 7 Oct. 1978, Hammel & D’Arcy 4989 (CAS, MO, PMA). Darién: de la estacién de Cruce Mono hasta la quebrada Ipelisa (limite del parque), 28 Aug. 1990, Aranda et al. 1517 (CAS, PMA); trail between Boca de Cupe and Rio Piedras on the trail to Paca and Cana, 27 June 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES | Volume 55, No. 4 1959, Stern et al. 733 (US). Los Santos/Veraguas border: mountains of S Azuero Peninsula near — proposed route of road from El Cortezo to Arenas, 0-5 km SW of El Cortezo, 29 Oct. 1978, Hammel 5461 (CAS, MO, PMA). Panama: area around Torti Arriba, 31 Aug. 1977, Folsom 5153 — (CAS, MO, PMA); hills S of Guacuco, 8 km E of Ipeti, 78°20’W, 8°55’N, 18 Sep. 1982, Hamilton © & D’Arcy 1353 (CAS). Veraguas: Dto. De Montijo, Isla de Coiba, campamento de La Salina, subi- — da hacia Rio Negro, 21 Oct. 1995, Martin & Ballesteros MB1345 (CAS, PMA). COLOMBIA: Antioquia: Quebrada Isaias, east of Turbo, 8 July 1946, Haught 4946 (US). Bolivar: Boca Verde, Rio Sinu, 13-14 Feb. 1918, Pennell 4227 (US). Choc6é: Rio Ciego, 17 June 1967, Duke 13290 (US); Corregimiento San Francisco, Vereda Coquital, Sitio “El Paramo, Quebrada Zardi, 8°23’N, 77°07’ W, 22 May 1989, Betancur et al. 119] (US). Cundinamarca: San Antonio, 14 June 1948, Haught 6241 (CAS). Magdalena: Parque Nacional Tayrona, along mule trail from Pueblito to Calabazo, ca. 11°19’N, 73°58’W, 26 Oct. 1972, Kirkbride 2565 (US). Tolima: Mariquita, 10 Nov. 1938, Haught 2429 (DS). VENEZUELA: Portuguesa: Distr. Guanare, 17 km NW of Tucupido by road, N slope of fila Las Palmas, 69°57’W, 9°O1’N, 30 Oct. 1982, Davidse et al. 21455 (CAS, MO, US). Zulia: Dto. Mara, cuenca del Rio Guasare, alrededores del Destacamento Guasare No. 1 (La Yolanda), 10°52’10”N, 72°29’30’W, 9 Nov. 1982, Bunting et al. 11977 (CAS). Ecuapor: Napo: Rio Napo between Coca (Puerto Francisco de Orellana) and Armenia Vieja, 15 Nov. 1974, Harling & Andersson 11986 (US). Pastaza: Rio Pastaza, between Destacamento Chiriboga and Apachi Entza, ca. 76°55’—77°8’W, 2°20-32’S, 24 July 1980, @llgaard et al. 35173 (CAS). DISCUSSION.— It is surprising that the generic placement of this species has not been adjusted until now since it was first proposed as a new species in 1768. It was described as Clidemia ? decurrens Beurl. in 1854 (see citation above) but students of neotropical Melastomataceae subse- quently relegated Beurling’s name to the synonymy of Octopleura quinquenervia or Ossaea quin- quenervia without comment (Triana 1871; Cogniaux 1891; Wurdack 1980). Gleason (1939) makes no mention of it in his regional revision of Clidemia. Among congeners, C. guinquenervia is closely allied to the rare Costa Rican endemic, C. coro- nata Gleason. Both species share a similar indument of asperous-headed hairs, densely papillose- furfuraceous petals, a completely inferior 5-locular ovary, and a smooth seed coat. Important dis- tinctions between these species can be found in readily observed vegetative characters. In C. coro- nata the leaves are 5—7-nerved (vs. 5—9-plinerved), the blades are ciliate and + entire (vs. ciliolate- denticulate) and the blade base is not decurrent on the petiole (vs. conspicuously so). Although both species also have anther connectives that are prolonged dorso-basally into appendages, the appendages on C. coronata are rounded and entire whereas those on C. quinquenervia are spur-like and gland-edged. Clidemia rodriguezii Almeda, sp. nov. (Fig. 9) Type.— Costa Rica: Cartago: Canto6n de Paraiso, Cuenca del Reventazon. Orosi. Entrada a Tausito, aprox. 3 km después del cruce hacia la Estaci6n Tapanti, 9°46’20”N, 83°47'/10”W, 1400- 1600 m, 15 Apr. 1998, A. Rodriguez et al. 3239 (holotype: INB!; isotypes: CAS!, CR!, MO!, NY!, US!). Ramuli quadrangulati; internodi alatis demum carinatis, nodis tumidus et carinatis. Petioli 6.3—12 cm long; lamina 6—23 x 11.7—16.2 cm ovata vel subcordata apice acuminata basi rotunda- ta, 7—9-nervata, nervi subtus in axillis domatiis instructi. Inflorescentia lateralis in ramulis infra folia plerumque oriunda 1—1.5 cm longa multiflora; flores 4-meri, bracteolis 1.5—2 x 0.25—0.5 mm. Calycis tubus | mm longus, lobis interioribus 0.5 x 1 mm ovato-triangularibus, dentibus exteri- oribus 2 x | mm oblongis. Stamina isomorphica glabra, antherarum thecae 2 x 0.25 mm subulatae | ALMEDA: NEOTROPICAL GENUS CLIDEMIA itd) poro ventraliter inclinato; connectivum nec prolongatum nec appendiculatum. Ovarium 4-loculare et omnino inferum glabrum. Shrub 0.4—1.5 m tall, the glabrous cauline internodes thick and quadrate, distinctly alate when young becoming carinate with age with opposing faces 0.8—1.2 cm wide, the nodes bearing promi- nent expanded (0.25—0.75 mm) interpetiolar ridges. Very young nodes, vegetative buds, and young petioles (adaxial surface) copiously setose with smooth early deciduous hairs 0.5—1 mm long. Leaves of a pair equal to somewhat unequal, otherwise identical, bearing two pairs of vesicular domatia (24 mm long) on the abaxial blade base where the primary veins diverge from one anoth- er; petioles 6.3—12 cm long; blades, 16—23 x 11.7—16.2 cm, chartaceous and brittle when dry, ovate to subcordate, apex acuminate, base broadly rounded, margin ciliate-serrulate, 7—9-nerved, glabrous on the adaxial surface at maturity or with remnant tufts of smooth hairs along basal por- tions of the impressed primary veins, essentially glabrous abaxially. Inflorescence 1—1.5 cm long, a congested cluster of axillary dichasia, typically paired at each node below the leaves; rachis nodes persistently setose with flexuous hairs 0.5—1 mm long, the internodes glabrate and the pedicels copiously beset with minute stellulate or branched hairs; bracteoles 1.5—2 x 0.25—0.5 mm, oblong io narrowly triangular, glabrous except for a short terminal hair. Flowers 4-merous on pedicels 1.5—2 mm long. Hypanthia (at anthesis) suburceolate with a cylindric distal neck about 1 mm long, copiously stellulate-puberulent with an occasional scattering of smooth spreading flexuous hairs. Calyx tube 1 mm long, the calyx lobes 0.5 x 1 mm, ovate-triangular, fleshy, glabrous, erect and concealed by the calyx teeth; calyx teeth 2 x 1 mm, oblong, glabrous, widely spreading at anthesis and in fruit. Petals 2 x 0.75 mm, glabrous, white, oblong. Stamens isomorphic; filaments 2.5 mm long, glabrous; anthers 2 mm long, 0.25 mm wide, white, subulate with a ventrally inclined pore; connective somewhat thickened dorsally but neither prolonged nor appendaged below the thecae. Ovary 4-locular, completely inferior, apex glabrous and smooth. Style 5 mm long, glabrous; stig- ma punctiform. Berry 3—5 x 3—5 mm, purple at maturity. Seeds 0.5 mm long, ovoid to obovoid with a vaguely rugose testa. PHENOLOGY.— Flowering material has been collected in April and August; fruiting specimens in March, April, and August. DISTRIBUTION.— Rare and local in central and southeastern Costa Rica where it is known from the lower northern slopes of the Fila Matama, vicinity of Tapanti National Park, and the Parrita Valley north of Quepos at 700-1600 m. PARATYPES.— Costa Rica: Limon: Almirante. Fila divisoria entre la Cuenca superior del Rio Xichiari y la cuenca superior del Rio Boyei, 09°45’50’N, 83°19’45”W, 12 Aug. 1995, Herrera 8446 (CAS, CR). San José: Canton de Tarrazu. Faja Costefia del Valle de Parrita en el camino a Quepos. Fila San Isidro, luego del cruce a San Isidro, 09°33’25”N, 84°04’48”W, 26 Mar. 1995, Morales & Urefia 3767 (CAS, INB). Discussion.— This little-collected species is characterized by a combination of many unusu- al characters. The uppermost internodes are quadrate, winged, and quite thick with opposing faces 0.8—1.2 cm wide (Fig. 9A). The nodes have prominent expanded interpetiolar ridges (Fig. 9A). The 4-merous flowers are borne in axillary congested dichasia that are typically paired at a node (Fig. 9A). The hypanthia are constricted distally into a conspicous neck (Fig. 9F). The exterior calyx teeth are foliaceous, adnate to and conceal the actual calyx lobes (Fig. 9C). Two pairs of vesicular domatia are produced at the abaxial blade base where the primary veins diverge from one another (Fig. 9B) but these are unusually small when compared to other domatia-bearing species in the genus. In leaf shape and inflorescence architecture, C. rodriguezii is similar to and may be most close- ly related to C. discolor (Triana) Cogn., another species with 4-merous flowers and a 4-locular 116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 FicurE 9. Clidemia rodriguezii Almeda. A. naturally defoliated branch with infructescences, x ca. 1; B. representative leaf (abaxial surface), x 2; C. berry, x ca. 7; D. petal, x 17; E. stamen (profile view), x ca. 17; F. immature berry (longitu- dinal section), x ca. 10; G seeds, x ca. 32. (A— G from Rodriguez et al. 3239.) ALMEDA: NEOTROPICAL GENUS CLIDEMIA 117 ovary that is completely inferior. The latter has slender unwinged upper internodes that are round- ed or rounded-quadrate and lack prominent interpetiolar ridges. The leaf blades of C. discolor also lack vesicular domatia at the abaxial base and the internodes of the inflorescence rachis are fari- nose-furfuraceous with an admixture of clavate glands (vs. glabrate to minutely stellulate). Clidemia rodriguezii is also reminiscent of C. epiphytica (Triana) Cogn. Both species have 4- merous flowers, a 4-locular completely inferior ovary, and foliaceous calyx teeth that are adnate to and obscure the calyx lobes. The latter differs most notably in being a climbing vine with distinct- ly dimorphic leaf blades at each node that lack vesicular domatia. ETYMOLOGY.— This species is named for Alexander Rodriguez G. (b. 1972), student of Costa Rican Asteraceae and collector of the type series. His collections were accompanied by excellent pickled material of flowers and fruits that facilitated the study and illustration of this species. Clidemia tenebrosa Almeda, sp. nov. (Fig. 10) TyPE.— PANAMA: Coclé: forested slopes above El Copé along abandoned road leading to the Continental Divide, 8°38’N, 80°38’W, 700-850 m, 24 Jan. 1989, F: Almeda et al. 6392 (holotype: CAS!; isotypes: MO!, PMA!). Ramuli primum obtuse sulcato-quadrangulati demum teretes sicut petioli folia inflorescentia hypanthiaque pilis laevibus (pro parte glanduliferis) dense induti. Lamina 22-44.5 x 10-16.5 cm elliptica vel ovato-elliptica apice gradatimque acuminato basi late acuta vel obtusa, subtus in venis primariis secundariisque sparse pilis glanduliferis induti et sparse vel sparsissime stellato-puberu- la, 7—9-plinervata; petioli liberi 2.5—5.5 cm longi; formicaria 4-7 cm longa ad petiolorum apices evoluta et in laminarum bases immersa. Inflorescentiae multiflorae in foliorum superiorum axillis oppositis; flores 5-meri, bracteolis 1.5—2 x 0.25 mm; calycis tubus ca 0.5 mm, lobis interioribus 0.75—1 x 1—1.5 mm triangularibus, dentibus exterioribus 4-5 mm longis subulatis. Stamina isomor- phica glabra, antherarum thecae 1.5 x 0.5 mm oblongae vel subulatae poro paulo dorsaliter incli- nato; connectivum nec prolongatum nec appendiculatum. Ovarium 5-loculare et omnino inferum apice modice glanduloso-puberulo. Monopodial or sparingly branched shrub 1-3 m tall with rounded-quadrate internodes. Upper internodes, petioles, both leaf surfaces, inflorescence rachis, pedicels, hypanthia and calyx teeth copiously covered with smooth (gland-tipped in part) spreading hairs 1.5—3 mm long. Leaves of a pair essentially equal in size and consistently bearing paired elongate formicaria 4-7 cm long extending from the adaxial blade base onto the distal portions of the petiole; petioles 3-8 cm long: blades 2244.5 x 10-16.5 cm, chartaceous, flexuous when fresh and very brittle when dry, elliptic to ovate-elliptic, apex gradually acuminate, base rounded to obtuse, margin inconspicuously den- ticulate, 7—-9-plinerved with the innermost pair of primary veins diverging from the median vein 2-3 cm above the blade base, the primary and higher order veins sparsely beset with a ground layer of minute glands sometimes sparingly intermixed with minute stellate hairs. Inflorescence 2—3.5 cm long, axillary congested dichasia that are paired at a node, the rachis, pedicels and hypanthia sparsely to moderately beset with a ground cover of tardily deciduous glands and stellulate hairs; bracteoles 1.5-2 x 0.25 mm (including the apical hair), subulate, inconspicuous and obscured by the surrounding indument of spreading hairs. Flowers 5-merous on pedicels 0.5—1 mm long. Hypanthia (at anthesis) subcylindric. Calyx tube ca. 0.5 mm long, the calyx lobes 0.75—1 x 1-1.5 mm, triangular, glandular-puberulent adaxially and along the margins; calyx teeth 4-5 mm long, subulate, greatly exceeding and obscuring abaxial surface of the calyx lobes. Petals 4-5 x 2—-3.5 mm, glabrous, translucent white, obovate to oblong. Stamens isomorphic; filaments 2.5—3 mm eee eee eae ea a Se aS ie ashes <= 3 Z LV a) k ah eo S 2 ‘S Pas | f SUN AR cea cciee it CRS se conn , ° ; Shs Dn i S Z = S 22) Lh O > = 1) a < S < Z 5 : S fn ae - i, S YN O Z la ra ro S © 22 ae 118 B XK Ca . ‘A; C. immature x 12: EF. seeds, x ca. 38. (A and B from Knapp 3734; C from Almeda et al. 6392: D and E from Croat 67552: F from Almeda et al. 7657.) 5) > ight) FiGurE 10. Clidemia tenebrosa Almeda. A. habit, x 's; B. representative leaf (adaxial surface) berry, x 5; D. petal (adaxial surface), x 8; E. stamens, ventral view (left) and profile view (ri ALMEDA: NEOTROPICAL GENUS CLIDEMIA 119 long, glabrous; anthers 1.5 mm long, 0.5 mm wide, white, linear-oblong to subulate, truncate at the apex with a somewhat dorsally inclined pore; connective somewhat thickened dorsally but neither prolonged nor appendaged below the thecae. Ovary 5-locular, completely inferior, apex somewhat crateriform at anthesis, minutely glandular puberulent like the adaxial vascular ring of the hypan- thium. Style glabrous, 6 mm long; stigma capitate. Berry 6 x 5 mm, blue at maturity. Seeds 0.5 mm long, obovoid with a smooth to vaguely rugulate testa. PHENOLOGY.— Flowering material has been collected from January through March, July, September, and December; fruiting collections have been gathered in January, February, July, September, and December. DISTRIBUTION.— Cloud forests of west-central Panama at 200-1100 m, typically in deep shade. PARATYPES.— PANAMA: Coclé: forested slopes above El Copé along abandoned road leading to the Continental Divide, 25 Feb. 1988, Almeda et al. 5935 (CAS); about 7—10.5 km beyond El Copé in Omar Torrijos National Park along end of rocky trail to Rio Blanco and Lim6én beyond Alto Calvario, 21 Feb. 1996, Almeda et al. 7657 (CAS, MO, PMA); vicinity of La Mesa, N of El Valle de Anton, along N slope of Cerro Gaital, 8°37’N, 80°08’W, 14 Jul. 1987, Croat 67286 (CAS); Alto Calvario along summit of Continental Divide, 5.5 mi. N of El Copé, 8°39/N, 80°36’W, 13 Sep. 1987, Croat 67552 (CAS); area between Cano Blanco del Norte, Cano Sucio and Chorro del Rio Tife, 3 Feb. 1983, Davidse & Hamilton 23514 (CAS); Atlantic drainage near Continental Divide at sawmill above El Copé, 7 Mar. 1982, Hammel & Trainer 11314 (CAS); trail from Rio Blanco del Norte to Cafio Sucio, 8°44’N, 81°40’W, 21 Feb. 1982, Knapp 3734 (CAS); between Cafio Sucio and waterfall at base of Cerro Tife, ca. 4 hr. hike, 8°38’N, 80°36’W, 13 Dec. 1980, Sytsma et al. 2557 (CAS). Panama: sendero al Cerro Campana, 8°40’N, 79°55’W, 23 Sep. 1993, Correa & Montenegro 10036 (PMA); Cerro Campana near the FSU field station trail up the mountain, 9 Aug. 1970, Kennedy & Luteyn 435 (CAS, US); Altos de Campana, sendero del Tigre, 11 July 1998, Montenegro & Galdames 1979 (CAS). DIscUssION.— Clidemia tenebrosa is like C. collina, C. folsomii, and C. myrmecina in having 5-merous flowers and formicaria positioned at the base of the blade but extending on to a portion of the petiole. Among these species, C. tenebrosa most closely resembles C. collina. Both species have plinerved leaf blades with at least the innermost pair of primary veins diverging from the mid- vein well above the blade base on the abaxial surface and a short inflorescence axis that is obscured by the dense indument. In C. collina, however, the leaf blades are prevailingly ovate varying to ovate-elliptic and mostly 9.5—21 cm long (vs. prevailingly elliptic varying to ovate-elliptic and 2244.5 cm long), the inflorescence is pseudolateral but not paired at a node (vs. axillary and paired at a node), the understory indument on hypanthia is lacking or sparingly covered with minute deciduous glands (vs. a sparse to moderate cover of minute stellulate hairs and a scattering of minute glands), the anther thecae are yellow (vs. white), and the seed coat is densely tubercu- late (vs. smooth to vaguely rugulate). The flowers and fruits of C. tenebrosa and other congeners with congested copiously hairy inflorescences are difficult to study once they are pressed and dried. This coupled with the highly fugacious nature of floral parts and brittleness of the inflorescence rachis typically make for specimens that are incomplete. To facilitate the preparation of specimens that are adequate for study, collectors should make every effort to preserve reproductive structures in a liquid medium in the field. ETYMOLOGY.— The epithet for this species is derived from the Latin word, tenebrosus, dark, gloomy, or of shaded places, alluding to its shaded forest habitat. 120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 Clidemia ventricosa Almeda, sp. nov. (Fig. 11) foe tne Rint eee eres TypE.— PANAMA: Comarca de San Blas: El Llano—Carti road, 19.1 km from Interamerican ~ Hwy., 9°19’N, 78°55’W, elev. 350 m, 5 Mar. 1985, G de Nevers et al. 4963 (holotype: CAS!; iso- | types: CAS!, INB, KUNA, MEXU, MO, PMA). Ramuli teretes sicut foliorum venae primariae subtus inflorescentia hypanthiaque pilis lae- vibus plerumque 3—7 cm longis induti. Lamina 11—23.5 x 4.1—8.5 cm ovato-elliptica vel ovata apice attenuata vel acuta basi rotundata, 5—7-nervata vel 5—7-plinervata; formicaria 1-2 cm longa ad petiolorum apices evoluta et raro in laminarum bases immersus. Inflorescentia primum termi- nalis demum lateralis; flores 5-meri, bracteolis 3.5-4.5 x 0.25—0.5 mm; calycis tubus | mm, lobis interioribus 0.5—0.75 x 1 mm triangularibus vel orbicularibus, dentibus exterioribus 2.5—3.5 mm longis subulatis. Stamina isomorphica glabra, antherarum thecae 3—4.2 x 0.5 mm subulatae poro dorsaliter inclinato; connectivum nec prolongatum nec appendiculatum. Ovarium 5-loculare et 2/3 inferum apice modice glandulosum-puberulo. Sparingly branched shrub 0.5—1 m tall. The terete upper internodes, abaxial surface of petioles, and elevated primary veins on abaxial foliar surfaces copiously covered with appressed smooth hairs 3—7 cm long. Leaves of a pair equal to somewhat unequal in size; petioles 14 cm long with bulbous paired formicaria 1-2 cm long borne distally and sometimes barely extending onto the blade, the adaxial surface of the petioles and formicaria setose with spreading hairs 0.6—1.7 cm long; blades 11—23.5 x 4.1—8.5 cm, chartaceous, brittle when dry, oblong-ovate to elliptic-ovate, apex attenuate to acute, base rounded, margin entire, 5—7-nerved or if plinerved then the innermost pair of primary veins diverging from the median vein 0.5 cm above the blade base, the higher order veins and actual surface inconspicuously beset with scattered minute glands or nearly glabrous. Inflorescence 1.5—2 cm long, a pseudolateral congested modified dichasium, the rachis, hypanthia, and calyx teeth moderately setose with widely spreading hairs 0.4—1 cm long underlain by a scat- tering of very minute inconspicuous glands; bracts and bracteoles 3.5—4.5 x 0.25—0.5 mm (includ- ing the apical hair), subulate to oblong-subulate, inconspicuous and obscured by the surrounding setose indument. Flowers 5-merous on minutely glandular-puberulent pedicels 1-2 mm long. Hypanthia suburceolate at anthesis, prolonged distally into a cylindric neck 1 mm long above the ovary apex and below the vascular ring. Calyx tube 1 mm long, the calyx lobes 0.5—0.75 x 1 mm, rounded-triangular to semicircular, glandular-puberulent along the margins; calyx teeth 2.5—3.5 mm long (excluding the apical hair), greatly exceeding and somewhat obscuring abaxial surfaces of the calyx lobes, oblong to subulate. Petals 8 x 2.5 mm, glabrous, pink or reportedly yellowish with a tinge of purple at the tip (Nee 10474), oblong to narrowly obovate. Stamens isomorphic; fil- aments 3—3.5 mm long, glabrous, abruptly constricted at the junction with the thecae; anthers 34.2 x 0.5 mm, yellow, subulate with a dorsally inclined pore; connective thickened and somewhat ridged dorso-basally but neither prolonged nor appendaged below the thecae. Ovary 5-locular, 2/3- inferior, apex prolonged into a deciduously glandular truncate cone 1 mm long at anthesis. Style 9-9.5 mm long, glabrous, straight; stigma subcapitate to truncate. Berry 6-7 x 6-8 mm when dry, purple at maturity. Seeds 0.5 mm long, bluntly triangular in outline with a minutely asperulate testa. PHENOLOGY.— The single known flowering collection was made in March; fruiting speci- mens have been collected from August through October. DISTRIBUTION.— Local and uncommon in low rainforests of north-central Panama, often in deep shade along creeks and in the vicinity of streams at 300-400 m. PARATYPES.— PANAMA: Comarca de San Blas: El Llano—Carti road, 18 km from Interamerican Hwy., 9°19’N, 78°55’W, 7 Sept. 1984, de Nevers 3877 (CAS); Nusagandi, El Llano- IDA ALMEDA: NEOTROPICAL GENUS CLIDEMIA ; C. representa- ‘hs ; E. petal (adaxial surface), x 5; F. stamens, B. representative leaf (adaxial surface), x ': tive leaf (abaxial surface), x '4; D. flower (just prior to anthesis), FiGure 11. Clidemia ventricosa Almeda. A. habit, * ca. profile view (left) and dorsal view (right), x 6; G. young fruiting hypanthium, x 2; H. seeds, x ca. 26. (A—G from de Nevers et al. 4963; H from Folsom et al. 6182.) <2) 122. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 i Carti road, along a creek on the Atlantic slope, 9°19’N, 78°15’W, 10 Aug. 1984, de Nevers & : Gonzdles 3673 (CAS); El Llano—Carti road, headwaters of Atlantic drainage, 9°19’N, 78°55’ W, 29 Aug. 1984, de Nevers & Gonzdles 3784 (CAS); El Llano—Carti road, 9°19’N, 78°55’W, 1 June ~ 1985, de Nevers & Herrera 5821 (CAS). Panama: road from El] Llano to Carti, 12.4 km N of — Panamerican Hwy., 31 Oct. 1977, Folsom et al. 6182 (CAS); wet forest along El Llano—Carti road, | 12 km N of Panamerican Hwy., 11 Mar. 1974, Nee 10474 (MO, US). DISCUSSION.— Clidemia ventricosa can be recognized by its glabrous adaxial foliar surfaces, mostly petiolar formicaria (Fig. 11B) that sometimes barely extend on to the blade base, smooth appressed hairs on the upper internodes, abaxial surface of the petioles, and elevated primary veins on abaxial foliar surfaces (Fig. 11C), pseudolateral congested dichasia (Fig. 11A), and widely spreading hairs on the inflorescence rachis, hypanthium, and calyx teeth (Fig. 11C). Among the Central American species of Clidemia with 5-merous flowers, only C. taurina and some popula- tions of C. pubescens have petiolar formicaria like C. ventricosa. Both of these species are easily distinguished from C. ventricosa by their spreading or retrorse hairs on upper internodes and spreading hairs on elevated primary veins on the abaxial foliar surfaces. ETYMOLOGY.— The epithet for this species is derived from the Latin word, ventricosa, swollen, in reference to the conspicuously bulbous ant domatia at the distal end of each petiole. ACKNOWLEDGMENTS Fieldwork for this study was funded, in part, by U.S. National Science Foundation Grants DEB 76-83040, DEB 78—25620, BSR 86-14880 (Flora Mesoamericana), the G. Lindsay Field Research Fund of the California Academy of Sciences, and two anonymous private donors. A number of organizations provided logistical support during field studies in Mexico, Guatemala, Costa Rica and Panama over many years. I am especially grateful to the Museo Nacional de Costa Rica, the Missouri Botanical Garden, the Organization for Tropical Studies, the Instituto Nacional de Biodiversidad, Costa Rica, the Smithsonian Tropical Research Institute, the University of Mexico, and the University of Panama. I also thank a number of colleagues who contributed to this study directly or indirectly in diverse ways: B. Anderson, D. E. Breedlove, W. Burger, M. Correa, T. Croat, T. Daniel, G. Davidse, G. de Nevers, C. Galdames, M. Grayum, B. Hammel, G. McPherson, J. Solomon, M. Sousa, D. Stevens, G. Umana Dodero, I. Valdespino, P. S. Ward, R. L. Wilbur, N. Zamora, and the late J. J. Wurdack. The line drawings were prepared by J. Speckels (Figure 1); H. Pazdirkova (Figures 8, 9 and 11); S. Myers (Figure 5); C. Sudekum (Figures 2 and 7); E. del Valle (Figures 3 and 4); H. Kho (Figure 6); and M. Stalcup (Figure 10). I am also appreciative of the many courtesies received from curators and staffs of the following herbaria for either loans, gifts of specimens, or assistance during study visits: BM, BR, C, CAS, CR, DAV, DS, DUKE, F, G INB, K, LL, M, MA, MEXU, MICH, MO, NY, P, PMA, SCZ, TEFH, TEX, UC, UCR, US, USJ, and WIS. LITERATURE CITED ALLEN, P.H. 1956. The Rain Forests of Golfo Dulce. University of Florida Press, Gainesville. (Reissued in 1977 by Stanford University Press, Stanford, California). ALMEDA, F. 2000. New Costa Rican and Panamanian species of Miconia (Melastomataceae: Miconieae). Proceedings of the California Academy of Sciences 52(4):33—54. ALMEDA, F. 2001. Melastomataceae. Pages 1339-1419 in W. D. Stevens, C. Ulloa Ulloa, A. Pool, and O.M. Montiel, eds., Flora de Nicaragua. Monographs in Systematic Botany from the Missouri Botanical Garden., 85(II). ALMEDA: NEOTROPICAL GENUS CLIDEMIA 123 BENSON, W.W. 1985. Amazon ant-plants. Pages 239-266 in GT. Prance and T.E. Lovejoy, eds., Amazonia. Pergamon Press, Oxford, England. CoGNiAux, A. 1888. Melastomaceae (Clidemia). In C.F. P. von Martius, Flora Brasiliensis 14(4):468—511. CoGniAux, A. 1891. Mélastomacées. Jn A. and C. de Candolle, eds., Monographiae phanerogamarum 7: 1-1256. G Masson, Paris. GLEASON, H.A. 1939. The genus Clidemia in Mexico and Central America. Brittonia 3:97-140. Jupp. W.S. 1989. Taxonomic Studies in the Miconieae (Melastomataceae). II. Cladistic Analysis of Axillary- flowered taxa. Annals of the Missouri Botanical Garden 76:476—-495. TRIANA, J. 1871. Les Mélastomacées. Transactions of the Linnean Society, London 28:1-188. Wurpback, J.J. 1971. Certamen Melastomataceis XVI. Phytologia 21(2):115—130. Wurpback, J.J. 1980. Melastomataceae. Jn G. Harling and B. Sparre, eds,. Flora of Ecuador 13:1-406. 124 DAO © SVATON OOS IP ONOS) Guo Ie (ey SG OVO) OO PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 4 Appendix I Species of Clidemia known to produce pouch-like ant domatia (formicaria) on some or all leaves, petioles, or branchlets nodes, and their geographic distributions. acostae Wurdack — Ecuador ayangannensis Wurdack — Guyana allardii Wurdack var. allardii — Colombia, Ecuador, Peru allardii Wurdack var. maranonensis Wurdack — Ecuador, Peru, Brazil ciliata D. Don var. elata (Pittier) Uribe — Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Peru ciliata D. Don var. testiculata (Triana) Uribe — Venezuela, Colombia collina Gleason — Panama crenulata Gleason — Belize, Guatemala, Honduras, Costa Rica, Nicaragua, Colombia, Ecuador foliosa Gleason — Peru folsomii Almeda — Panama heterophylla (Desr.) Gleason — Colombia, Ecuador, Peru heptamera Wurdack — Venezuela, Guyana inobsepta Wurdack — Colombia, Ecuador juruensis (Pilger) Gleason — Venezuela, Brazil, Peru killipti Gleason — Colombia lanuginosa Almeda — Costa Rica, Panama myrmecina Gleason — Panama, Colombia neblinae Wurdack — Venezuela pilosa D. Don — Venezuela, Colombia, Ecuador, Peru pubescens Gleason — Costa Rica, Panama rodriguezii Almeda — Costa Rica (NB. The vesicular domatia of this species may be acarodomatia.) setosa (Triana) Gleason — Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama spectabilis Gleason — Nicaragua, Costa Rica, Panama sprucei Gleason — Ecuador, Peru taurina Gleason — Costa Rica, Panama tenebrosa Almeda — Panama tococoidea (DC.) Gleason — Venezuela, Brazil, Colombia ventricosa Almeda — Panama Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5, pp. 125-161, 29 figs., 1 table April 22, 2004 Phylogenetic Systematics of Okenia, Sakishimaia, Hopkinsiella and Hopkinsia (Nudibranchia: Goniodorididae) with Descriptions of New Species from the Tropical Indo-Pacific Terrence M. Gosliner Department of Invertebrate Zoology and Geology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 U.S.A.; Email: tgosliner@ calacademy.org. The anatomy of eight species of Okenia, four species traditionally placed in Hopkinsia and one species of Sakishimaia was examined. The descriptions of previ- ously described species, Okenia pellucida, O. japonica, Hopkinsia rosacea, H. plana, H. pilosa, H. nakamotoensis and Sakishimaia kondoi were expanded from prior mor- phological studies. Six new species of Okenia are described from the Indo-Pacific tropics. Okenia brunneomaculata is described from specimens collected from Indonesia and the Philippines. Okenia purpureolineata is known only from Okinawa in the Ryukyu Islands. Okenia kendi is described from specimens from Indonesia and the Philippines. Okenia virginiae is described from South Africa and is also known from Oman. Okenia lambat is described from the Philippines. Okenia liklik is described based on specimens from Papua New Guinea and the Philippines. Preliminary phylogenetic studies reveal several important aspects about the rela- tionships of these taxa. Collectively, these taxa appear to form a monophyletic clade. Okenia mediterranea is the most basal taxon in the present analysis and is the sis- ter group to all of the other taxa. Hopkinsia, Sakishimaia and Hopkinsiella are nest- ed within Okenia in the analysis presented. The characteristics employed to distin- guish Sakishimaia and Hopkinsiella from Hopkinsia are not distinctly different or represent autapomorphies or symplesiomorphies. Hopkinsia, when Sakishimaia and Hopkinsiella are included, constitutes a monophyletic taxon. However, maintenance of Hopkinsia as a distinct taxon renders Okenia paraphyletic. On this basis, Hopkinsia, Hopkinsiella and Sakishimaia are considered as synonyms of Okenia to preserve the monophyly of this oldest taxon. The Indo-Pacific Goniodorididae consist of species that have been placed in the genera Ancula Loven, 1846; Goniodoris Forbes and Goodsir, 1839; Goniodoridella Pruvot-Fol, 1933; Hopkinsia MacFarland, 1905; Hopkinsiella Baba, 1938; Murphydoris Sigurdsson, 1991; Okenia Menke, 1830; Sakishimaia Hamatani, 2001; Spahria Risbec, 1928 and Trapania Pruvot-Fol, 1931. These taxa have been sporadically studied for the last 100 years with additional species described in a wide variety of faunistic and systematic treatments. Recently, Rudman’s (1987) monographic study of Indo-Pacific species of Trapania has significantly added to our knowledge of Indo-Pacific Goniodorididae. Few taxonomic studies have focused on the distinctions between the genera Okenia, Hopkinsiella, Hopkinsia and Sakishimaia. Bouchet and Ortea (1983) revised the system- atic distinctions between the former three genera and concluded that Okenia and Hopkinsia repre- sent distinct genera, whereas Hopkinsiella is regarded as junior synonym of Hopkinsia. Most recently, Hamatani (2001) described two new species from Japan, a species of Hopkinsia and a member of a new genus, Sakishimaia. 125 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 Recent investigations of tropical coral reef ecosystems throughout the Indo-Pacific tropics from South Africa, the Philippines, Indonesia, Papua New Guinea, the Marshall Islands, and the Hawaiian Islands have yielded specimens of six new species of Okenia. Description of these taxa is the focus of this investigation. The discovery of additional specimens Okenia pellucida Burn, 1967; O. japonica Baba, 1949; Hopkinsia pilosa Bouchet and Ortea, 1983; Hopkinsia nakamotoen- — sis Hamatani, 2001 and Sakishimaia kondoi Hamatani, 2001 has permitted the amplification of — these species. Anatomical examination of these taxa and preliminary phylogenetic analyses under- taken here require systematic revision of several goniodoridid taxa. All specimens examined here are housed in the Department of Invertebrate Zoology and Geology of the California Academy of Sciences (CASIZ) and the South African Museum (SAM). SPECIES DESCRIPTIONS Okenia Menke, 1830 Type species: Okenia elegans (Leuckart, 1828), by monotypy. Idalia Leuckart, 1828 (type species, /dalia elegans (Leuckart, 1828), by original designation). Idaliella Bergh, 1881 (type species, /daliella elegans (Leuckart, 1828), by original designation). Idalina Norman, 1890 (type species, /dalina elegans (Leuckart, 1828), by original designation). Cargoa Vogel and Schultz, 1970 (type species Cargoa cupella Vogel and Schultz, 1970, by original designa- tion). Hopkinsia MacFarland, 1905 (type species: Hopkinsia rosacea MacFarland, 1905, by monotypy). SYN. NOV. Hopkinsiella Baba, 1938, (type species, H. hiroi Baba, 1938, by original designation). SYN. NOV. Sakishimaia Hamatani, 2001, (type species, S$. kondoi Hamatani, 2001, by original designation). SYN. NOV. Okenia pellucida Burn, 1967 (Figs. 1F, 2-3) Okenia pellucida Burn, 1967:52, pls. X—XI, figs. 1-3. MATERIAL EXAMINED.— CASIZ 078645, seven specimens, Maalena Bay, Maui, Hawaiian Islands, 6 m depth, 4 June 1991, P. Fiene Severns. CASIZ 078493, 26 specimens, | dissected, Marang River Mouth, 16 km s. of Kuala Terengganu, Malay Peninsula, Malaysia, intertidal zone, 26 July, 1991, T.M. Gosliner. CASIZ 168016, two specimens, Sand Island, Palmyra Atoll, Oceania, 10 m depth, 27 May 2002, T.M. Gosliner. DISTRIBUTION.— Australia (Burn 1967), United Arab Emirates (Behrens 2001), Hawaii (Gosliner, Johnson, and Bertsch 1986), Japan (Kurihara 1999) Palmyra Atoll and Malaysia (both from present study). NATURAL History.— Okenia pellucida, together with its elongate egg masses, 1s found com- monly on large masses of the ctenostomatous bryozoan, Zoobotryon sp. EXTERNAL MORPHOLOGY.— The body is elongate (7-20 mm in length) with numerous elon- gate, rounded papillae situated along the notal margin and on the remainder of the notum. The body (Fig. 1F) is translucent white with numerous thin brown lines that form an interconnecting network on the notum. The rhinophores also contain patches of brown pigment. The rhinophores are elon- gate with 11 well—spaced lamellae. The gill consists of 8 bipinnate branches. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular with a rounded buccal pump directed dorsally. Numerous small, elongate, pyriform oral glands are present at the opening of the buccal mass into the mouth. The radular sac is short and extends ventrally from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ventral to the buc- GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 127 Ficure 1. Living animals. A. Okenia kendi sp. nov., Sepok Point, Luzon, Philippines. B. Okenia lambat sp. nov. (CASIZ 084283), Devil’s Point, Luzon, Philippines. C. Okenia virginiae sp. nov. (SAMA 35558), Vetchies Pier, Durban, South Africa. D. Okenia purpureolineata sp. nov., Seragaki Tombs, Okinawa, Japan, photo by R.F. Bolland. E. Okenia brunneomaculata sp. nov., Luzon, Philippines. F. Okenia pellucida Burn, 1967 (CASIZ 071398), Hawaii. G. Okenia liklik Sp. noy., Barracuda Point, Madang, Papua New Guinea. All photos by T.M. Gosliner except as otherwise noted. 128 cal pump. A rounded, lobate salivary gland is present on either side of the buccal mass ante- rior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregu- lar polygonal plates. The cuticle expands as it enters the buccal pump. The radular formula is 16 x 1.1.0.1.1. in one specimen examined. The inner lateral teeth (Fig. 2) are wide basally with an elongate acute bifid cusp. The masticatory margin of the inner lateral bears 17 triangular denticles that increase in size in the direction of the outer margin. The outer laterals are small and reduced (Fig. 2) with two short, blunt cusps along the outer edge. REPRODUCTIVE SYSTEM (Fig. 3).— The preampullary duct is thin and elongate and expands into the elongate, sausage—shaped ampulla. The ampulla divides into a short oviduct that enters the female gland mass and a more elongate vas deferens. The proximal por- tion of the vas deferens is prostatic but relative- ly narrow. It curves and continues as an elon- gate loop that eventually widens slightly into the highly convoluted muscular, ejaculatory portion. The penial bulb is wide and conical. From it protrudes a rounded, fleshy penial papilla. The vagina is short and wide at its exit adjacent to the penis. More distally it narrows and continues as a thin duct that enters the base of the large, pyriform bursa copulatrix. From the base of the bursa is a second duct that joins with the smaller, more rounded receptaculum seminis. Near the base of the receptaculum the uterine duct separates from the duct joining the receptaculum and bursa and enters the female gland mass. The female gland mass consists of three portions, the albumen, membrane and mucous glands. The mucous gland is the largest of the nidamental glands. DISCUSSION.— Okenia pellucida is imme- diately identifiable by its translucent white body color with a network of interconnecting thin brown lines. The present material agrees closely with that originally described by Burn PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 FIGURE 2. Okenia pellucida Burn, 078493). Scanning electron micrograph of radular teeth. FiGureE 3. Okenia pellucida Burn, 1967 (CASIZ 078493). Reproductive system, am = ampulla, be = bursa copulatrix, fem = female gland mass, p = penis, rs = receptaculum sem- inis, V = vagina, vd = vas deferens; scale = 0.6 mm. (1967), with several exceptions. Burn did not describe the presence of oral glands surrounding the anterior end of the buccal mass. The original description included specimens with as many as 28 1967 (CASIZ ) GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 129 rows of radular teeth whereas the present material had only 16 rows. Burn also described the inner lateral teeth as having a smooth masticatory margin. The present material from Malaysia and spec- imens previously reported from the Hawaiian Islands (Gosliner, Johnson, and Bertsch 1986) have a series of triangular denticles along the masticatory margin. Despite these differences, the consis- tency in external anatomy, color pattern and reproductive anatomy suggest that all the material identified previously and from this study are conspecific. Recent records of this species support its wide ranging distribution from the United Arab Emirates to the Hawaiian Islands and Palmyra Atoll. This wide distribution of Okenia pellucida is not only facilitated by dispersal of plank- totrophic larvae, but also by its association with the ctenostomatous bryozoan, Zoobotryon sp., which forms large masses carried by oceanic currents. Okenia brunneomaculata Gosliner, sp. nov. (Figs. LE, 4-5) TYPE MATERIAL.— Ho.oryPe: CASIZ 168019, 20 m. depth, Ligpo Island, Calumpan Penin- sula, Batangas Province, Luzon, Philippines, 1-2 m depth, 24 March 1993, T.M. Gosliner. PARATYPES: CASIZ 085876, 4 specimens, 2 dissected, 20 m. depth, Ligpo Island, Calumpan Penin- sula, Batangas Province, Luzon, Philippines, 1-2 m depth, 24 March 1993, T.M. Gosliner. CASIZ 083853, 1 specimen, 20m. depth, Sepok Point, Maricaban Island, Batangas Province, Luzon, Philippines, 1—2 m depth, 26 February 1992, T.M. Gosliner. CASIZ 168017, three specimens, Tul- amben, Bali, Indonesia, 21 October 2001, M.D. Miller. CASIZ 168018, two specimens, Tulamben, Bali, Indonesia, 21 October 2001, M.D. Miller. ETYMOLOGY.— The specific name, Okenia brunneomaculata, comes from the scattered brown spots found on the dorsal surface of the notum. DISTRIBUTION.— This species is known only from Luzon Island, Philippines and Bali, Indonesia (present study). NATURAL History.— This species is found on ctenostomatous bryozoans on the undersides of the large foliose sponge colonies. EXTERNAL MORPHOLOGY.— The living animals (Fig. 1E) are small, 4-6 mm in length. The preserved specimens are 2—3 mm. long. The body is moderately short and relatively high. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papil- lae. The body color is uniformly translucent white. There is a lateral band of chocolate brown and irregular spots of the same color scattered over the notum. There are 5—6 pairs of elongate notal papillae along the sides of the body. The two anteriormost pairs of papillae are situated in front of the rhinophores and are anteriorly directed. Four to seven elongate papillae are scattered mid-dor- sally anterior to the gill. A single mid-dorsal papilla is present posterior to the gill. The gill consists of 3-5 unipinnate branches. The rhinophores are elongate with 10 well—spaced lamellae. The ante- rior end of the foot contains two elongate triangular lobes that are united medially. The genital aper- ture is situated on the right side of the body approximately a third of the length of the body poste- rior to the head. The foot is narrow and widens anteriorly The oral tentacles (Fig. 4A) are elongate and extend well beyond the margin of the foot. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular with a rounded buccal pump directed posteriorly. There are no obvious oral glands around the mouth. The radular sac is short and extends ventrally from the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth but no discrete jaw elements are visible. The cuticle expands as it enters the buccal pump. The radular formula is 14 x 1.1.0.1.1. in one specimen examined. The inner lateral teeth (Fig. 5A) are broad basally with an acutely pointed cusp and 9-11 elongate denticles along the masticatory 130 margin. The outer laterals are slightly smaller and reduced (Fig. 5B) with bifid cusp with two elongate projections along the outer side. REPRODUCTIVE SYSTEM (Fig. 4B).— The preampullary duct is short and thin and expands into sausage-shaped ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic portion of the vas defer- ens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of two folded segments. The distal end of the prostatic segment narrows into a short, curved ejaculatory portion that terminates in a narrow penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is narrow and elongate and connects directly to the large, spherical bursa copulatrix. At the base the bursa, the receptaculum duct emerges and par- allels the vagina for some distance prior to join- ing the large, pyriform receptaculum seminis. The uterine duct emerges from near the base of the receptaculum and enters the female gland mass. DISCUSSION.— Several other species of Okenia have a whitish body color with brown markings: O. angelensis Lance, 1966; O. zoobotryon (Smallwood, 1910); O. impexa Marcus, 1957; O. mija Burn, 1967, O. pelluci- da Burn, 1967. All of these other species have much shorter notal papillae than does O. brun- neomaculata. In O. brunneomaculata, the notal papillae are as long as the width of the body. The dorsal brown spots and bands of O. brun- neomaculata are much larger than those found PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 FIGURE 4. Okenia brunneomaculata sp. nov. (CASIZ 085876). A. Ventral surface of head; scale = 0.5 mm. B. Reproductive system, am = ampulla, be = bursa copulatrix, p = penis, pr = prostate, rs = receptaculum seminis; scale = 0.2mm. — FiGuRE 5. Okenia brunneomaculata sp. noy. (CASIZ 085876). Scanning electron micrographs of radular teeth. A. Inner lateral teeth. B. Outer lateral teeth. Scale = 2um. in the remaining species. The internal anatomy of O. mija remains unknown. Of the remaining species, O. zoobotryon and O. angelensis have outer lateral teeth that are short and rounded with bifid cusp (Marcus 1957; Valdés and Ortea 1995; Lance 1966). Okenia brunneomaculata and O. impexa have elongate outer laterals with a bifid cusp. In O. impexa, the outer lateral tooth has an additional basal denticle that is not present in O. brunneomaculata. Okenia impexa also has a sin- gle mid-dorsal papilla, whereas O. brunneomaculata has numerous dorsal papillae. : a i i i GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 131 Okenia japonica Baba, 1949 (Figs. 6-7) Okenia (Okenia) japonica Baba, 1949:45, fig. 45, pl. 15, fig. 52, Rudman and Darvell, 1990:39, pl. 2a. MATERIAL EXAMINED.— CASIZ 070216, 1 specimen, 170 ft. [52 m] deep, Seragaki Beach, Okinawa, 22 April 1989, R.F. Bolland. CASIZ 099075, 1 specimen, 190 ft [58 m] deep, Seragaki Tombs, Okinawa, 12 March 1994, R.F. Bolland. CASIZ 105350, 2 specimens, 180 ft [55 m] deep, Seragaki Beach, Okinawa, 24 March 1994, R.F. Bolland. CASIZ 115378, 1 specimen, 190 ft [58 m|] deep, Seragaki, Okinawa, 30 April 1989, R.F. Bolland. CASIZ 115382, 2 specimens dissected, 195 ft [59 m] deep, Seragaki, Okinawa, 11 April 1997, R.F. Bolland. RFB 2135B 180 ft [55 m] deep, Seragaki, Okinawa, 30 April 1989, R.F. Bolland. DISTRIBUTION.—This species is known from Sagami Bay and Ku, Japan (Baba 1949) and Hong Kong (Rudman and Darvell 1990) and is here recorded from Okinawa in the Ryukyu Islands. NATURAL HIsToORY.—Okenia japonica was collected on erect arborescent bryozoans in 50-60 m depth (R. Bolland, pers. commun.). EXTERNAL MORPHOLOGY.—The living ani- mals are 10-13 mm in length. The body is mod- erately elongate and relatively high. There is well—developed, distinct notal border. The foot extends posteriorly and is devoid of notal papil- lae. The body color is uniformly translucent white with a dense, uniform covering of opaque white. There are 6—7 pairs of elongate notal papillae along the sides of the body. The two anteriormost papillae are situated in front of the rhinophores and are anteriorly directed. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of 4-5 unipinnate branches. The anteriormost branch is frequently bifid at the base. The rhinophores are elongate with 7-8 congested lamellae. The anterior end of the foot contains two elongate triangular lobes that are united medially. The genital aperture is situated on the right side of the body approxi- mately a third of the length of the body posterior to the head. The foot is narrow and is wider ante- riorly. The oral tentacles (Fig. 6A) are broadly triangular and extend beyond the lateral margins of the foot. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 6B) with a rounded buccal pump directed dorsally. Numerous, large, FIGURE 6. Okenia japonica Baba, 1949 (CASIZ115382). A. ventral view of head, scale = 1 mm. B. Buccal mass, bp = buccal pump, e = esophagus, g = oral glands, r = radula sac, sg = salivary gland; scale = 0.25 mm. C. Reproductive sys- tem, am = ampulla, be = bursa copulatrix, p = penis, pr = prostate, rs = receptaculum seminis; scale = 0.2 mm. 12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 elongate, pyriform oral glands are present at the re ae opening of the buccal mass into the mouth. The y isl < fo) radular sac is short and extends ventrally from _ pees the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the open- ing of the mouth. It contains irregular polygo- nal plates. The cuticle expands as it enters the buccal pump. The radular formula is 20x 1.1.0.1.1. The inner lateral teeth (Fig. 7) are wide basally with an elongate acute cusp. The masticatory margin of the inner lateral bears about 19-22 triangular denticles that increase in size in the direction of the outer margin. The outer laterals are small and reduced (Fig. 7) with a short cusp along the outer edge. aT PERT (CaN ETO Go eae FIGURE ile Okenia japonica Baba, 1949 (CASIZ 115382). ete) Scanning electron micrograph of radular teeth; scale = 20 of the central nervous system are highly con- um. centrated and surround the esophagus, at the posterior end of the buccal mass. The cerebral and pleural ganglia are largely fused. A sessile eye is present at the base of either cerebral ganglion. The pedal ganglia are smaller than the cerebro- pleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM (Fig. 6C).— The preampullary duct is short and thin and expands into an ovoid ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albu- men gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of a single, undivided portion. The distal end of the prostatic segment narrows into an elongate, straight ejaculatory portion that terminates near the base of the wide, elongate penis. There is no distinct penial papilla. Inside the distal end of the penis are penial spines. The vagina exits adjacent to the penis. It is wide and elongate basally and narrows more proximally. The narrow portion con- nects directly to the spherical bursa copulatrix and the smaller, pyriform receptaculum semunis. Near the middle of the narrow portion of the vagina, the uterine duct separates and enters the albu- men gland. DISCUSSION.— This species has been documented only once since its original description (Baba 1949; Rudman and Darvell 1990). The presence of a yellowish white body with fine opaque white markings is consistent between the original description and the present material. The num- ber of notal appendages, the presence of a single medial papilla and a basally bifid anterior gill branch are also consistent between the original and present material. The denticulation and shape of the radular teeth are also very similar. The only difference noted is that Baba described 5—8 mm animals with 32-35 row of radular teeth whereas the present material (10-13 mm) has only 20 rows. The large oral glands were not noted by Baba nor was the reproductive anatomy described. jy ai GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 133 There is little doubt that the present material from deep water in Okinawa represents the same species originally described by Baba. Okenia purpureolineata Gosliner, sp. nov. (Figs. 1D, 8-9) TYPE MATERIAL.— Ho.otyPe: CASIZ 168020, 160 ft deep, Seragaki Beach, Okinawa, 2 June 1989, R.F. Bolland. PARATYPES: CASIZ 099097, 1 specimen, 190 ft [58 m] deep, Seragaki Tombs, Okinawa 12 March 1994, R.F. Bolland. CASIZ 099085, 1 specimen, 175 ft [53 m] deep, Seragaki Tombs, Okinawa 18 February 1994, R.F. Bolland. CASIZ 070256, 1 specimen, dissected, 160 ft [49 m] deep, Seragaki Beach, Okinawa, 2 June 1989,, R.F. Bolland. CASIZ 070217, | specimen, 180 ft. [55 m] deep, Seragaki Beach, Okinawa, 30 April 1989, R.F. Bolland. ETYMOLOGyY.— The species name, Okenia purpureolineata, is derived from the presence of longitudinal purple lines on the dorsal surface of the body. DISTRIBUTION.— This species is known only from Okinawa in the Ryukyu Islands (present study). NATURAL History.— Okenia purpureolineata was collected on erect arborescent bryozoans in 49-60 m depth (R. Bolland, pers. commun.). EXTERNAL MORPHOLOGY.— The living animals (Fig. 1D) are 10-12 mm in length. The body is moderately elongate and relatively high. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papillae. The body color is uniformly translucent purple. There is an irregular network of darker purple lines on the dorsal surface of the notum. Darker purple pigment is also present on the gill lamellae. The rhinophores are dark purple basal- ly and red throughout most of their length. There are seven pairs of elongate notal papillae along the sides of the body. The two anteriormost pairs of papillae are situated in front of the rhinophores and are anteriorly directed. Two medial papillae are present mid-dorsally anterior to the gill. The gill consists of 4-5 unipinnate branches. The rhinophores are elongate with 18-21 congested lamellae. The anterior end of the foot contains two elongate triangular lobes that are united medially. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. The foot is narrow and is wider anteriorly. The oral tentacles are broadly triangular and extend beyond the lateral margins of the foot. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 8A) with a rounded buccal pump directed posterodorsally. Numerous large, elongate, pyriform oral glands are present at the opening of the buccal mass into the mouth. The radular sac is short and FIGURE 8. Okenia purpureolineata sp. nov. (CASIZ 070256). A. Buccal mass, bp = buccal pump, e = esopha- gus, g = oral glands, r = radula sac, sg = salivary gland; scale = 0.25 mm. B. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, pr = prostate, rs = receptaculum seminis, v = vagina; scale = 0.35 mm. 134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 extends ventrally from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ventral to the buc- cal pump. A rounded, lobate salivary gland is present on either side of the buccal mass ante- rior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregu- lar rounded plates. The cuticle expands as it enters the buccal pump. The radular formula is 28 x 1.1.0.1.1. The inner lateral teeth (Fig. 9A) are wide basally with an elongate acute cusp. The masticatory margin of the inner lateral bears 9-12 coarse, triangular denticles. The outer laterals are small and reduced (Fig. 9B) and quadrangular. CENTRAL NERVOUS SYSTEM.— The ganglia of the central nervous system are highly con- centrated and surround the esophagus, at the posterior end of the buccal mass. The cerebral and pleural ganglia are largely fused. A sessile eye 1s present at the base of either cerebral gan- glion. The pedal ganglia are smaller than the cerebropleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM (Fig. 8B).— The preampullary duct is short and thin and oe expands into a large sausage—shaped ampulla. ms i The ampulla divides into a short, narrow ak i oviduct and the wider prostatic portion of the FIGURE 9. Okenia purpureolineata sp. nov. (CASIZ vas deferens. The oviduct enters the lobate 070256). Scanning electron micrographs of radular teeth. A. female gland mass that consists of a small albu- View of half row of radular teeth; scale = 19 pm: BOute) = lateral teeth; scale = 12 um. men gland, a lobed membrane gland and a larg- er, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The pro- static portion of the vas deferens is wide and bilobed. The distal end of the prostatic segment nar- rows into an elongate, convoluted ejaculatory portion that terminates at the top of the widest por- tion of the elongate penis. There is no distinct penial papilla. Inside the distal end of the penis are penial spines. The vagina exits adjacent to the penis. It is narrow and elongate throughout its length. The proximal end of the vagina enters the spherical bursa copulatrix. At the base of the bursa anoth- er duct connects the large, pyriform receptaculum seminis. At the base of the receptaculum, the short uterine duct separates and enters the albumen gland. DIscUSSION.— Okenia purpureolineata is similar to O. japonica in its external body shape. Both species occur sympatrically in relatively deep water from Okinawa. Although O. japonica is uniformly white in color, O. purpureolineata is purple with darker purple lines on the notum and red rhinophores. Okenia purpureolineata has two mid-dorsal papillae but only one is present in O. japonica. Both species have large oral glands surrounding the anterior end of the buccal mass, but GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 135 there are more numerous glands in O. purpureolineata, that are smaller in size. The inner lateral teeth of O. purpureolineata have 9-12 large denticles along the masticatory border whereas there are 19—22 finer denticles on the inner laterals of O. japonica. In the reproductive system of O. pur- pureolineata, the receptaculum seminis and bursa copulatrix are well separated from each other whereas in O. japonica they are immediately adjacent to each other. Okenia kendi Gosliner, sp. nov. (Figs. 1A, 10-11) Okenia sp. 2 Gosliner, Behrens and Williams, 1996:171, fig. 605. TYPE MATERIAL.— Ho.otyPe: CASIZ 110451, 1 specimen, Bethlehem, Caban Island, Batangas Province, Luzon, Philippines, 10 m depth, 24 April 1997, T.M. Gosliner. PARATYPES: CASIZ 085880, 2 specimens, Babbit, Calumpan Peninsula, Batangas Province, Luzon, Philippines 25 March 1993, T.M. Gosliner. CASIZ 085892, 2 specimens, Layalayag, Maricaban Island, Batangas Province, Luzon, Philippines, 22 March 1993, T.M. Gosliner. CASIZ 085877, 2 speci- mens, Sepok, Maricaban Island, Batangas Province, Luzon, Philippines, 10 m depth, 23 March 1993, T.M. Gosliner. CASIZ 085883, 2 specimens dissected, Sepok, Maricaban Island, Batangas Province, Luzon, Philippines, 10 m depth, 26 March 1993, T.M. Gosliner. CASIZ 097397, 1 spec- imen, Devil’s Point, Maricaban Island, Batangas Province, Luzon, Philippines, 10 m depth, 13 March 1994, T.M. Gosliner. CASIZ 105681, 1 specimen, Sepok, Maricaban Island, Batangas Province, Luzon, Philippines, 10 m depth, 26 February 1995, T.M. Gosliner. CASIZ 106593, 1 specimen, Bethlehem, Caban Island, Batangas Province, Luzon, Philippines, 15 m depth, 19 April 1996, T.M. Gosliner. CASIZ 093923, 2 specimens, Tandarusa Wreck and Pulau Kecil, Lembeh Strait, Manado, Sulawesi, Indonesia, 5-6 November 1993, P. Fiene Severns. ETYMOLOGy.— The specific epithet kendi is the Filipino word for candy, in reference to its appearance to a sweet confection. DISTRIBUTION.— This species is known from the Luzon Island, Philippines (Gosliner, Behrens, and Williams 1996) and Sulawesi, Indonesia (present study). NATURAL History.— Okenia kendi is found on the underside of the large brown leafy sponge Phyllospongia lamellosa (Esper, 1799), on shallow reefs in 5—15 meters of water, where it feeds upon encrusting bryozoans (Gosliner, Behrens, and Williams, 1996). EXTERNAL MORPHOLOGY.— The living animals (Fig. 1A) attain 30 mm in length. Preserved specimens range in size from 15—25 mm. The body is wide and dorso—ventrally flattened. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papil- lae. The general body color is translucent yellowish white. There is a broad mid-dorsal band of maroon pigment that runs from anterior to the rhinophores to the posterior end of the foot. The band divides around the mid-dorsal papillae and the two bifurcations rejoin each other posterior to the gills. The rhinophores and gill also have maroon pigment but also possess opaque white markings along their lengths. The medial and marginal papillae are maroon at the base and possess purple pigment more distally. There is a lateral line of maroon present on either side of the body between the notum and the foot. There are eight pairs of elongate notal papillae along the sides of the body. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of three unip- innate branches that are bifid or trifid at the base. The rhinophores are elongate with 31 congested lamellae. The anterior end of the foot contains two elongate rounded lobes that are united medial- ly. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. The foot is broad and is divided anteriorly. A groove separates the foot from the mouth and oral tentacles (Fig. 10A). 136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 er : x i FicureE 11. Okenia kendi sp. nov. (CASIZ 085883). Scanning electron micrograph of radular teeth; scale = 23 um. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 10B) with a rounded ; ; bem buccal pump directed posteriorly. There are a FIGURE 10. Okenia kendi sp. nov. (CSIZ 085883). A. ventral view of head; scale = 0.5 mm. B. Buccal mass, bp few elongate oral glands around the mouth. The = buccal pump, e = esophagus, g = oral glands, r=radula radular sac is short and extends ventrally from sac, sg = salivary gland, scale = 0.5 mm. C. Reproductive the buccal mass. The esophagus is thin and system, am = ampulla, be = bursa copulatrix, fgm = female elongate and inserts into the buccal mass imme- eae sae est AV tae Sealers diately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle sur- rounds the lips at the opening of the mouth. It contains irregular rounded plates. The cuticle expands as it enters the buccal pump. The radular formula is 26 x 1.1.0.1.1. The inner lateral teeth (Fig. 11) are broad basally with an acutely pointed cusp and 35-38 elongate denticles along the masticatory margin. The outer laterals are smaller and reduced (Fig. 11) with an elongate cusp along their outer side. CENTRAL NERVOUS SYSTEM.— The ganglia of the central nervous system are highly concen- trated and surround the esophagus, immediately posterior to the buccal mass. The cerebral and pleural ganglia are largely fused. A sessile eye is present at the base of either cerebral ganglion. The pedal ganglia are smaller than the cerebropleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM (Fig. 10C).—The preampullary duct is short and thin and expands into an elongate ampulla. The ampulla divides into a short, narrow oviduct and the wider prostat- ic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamen- GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 137 tal glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of two folded portions. The distal end of the prostatic segment narrows into an elon- gate, convoluted ejaculatory portion that terminates in a wide, elongate penis. There is no distinct penial papilla. Inside the distal end of the penis are penial spines. The vagina exits adjacent to the penis. It is wide and short and connects directly to the spherical bursa copulatrix. Near the base of the vagina, the uterine duct separates and enters the albumen gland. From the base of the bursa cop- ulatrix, a narrow duct connects the bursa with the large, pyriform receptaculum seminis. DISCUSSION.— Two described species of Indo-Pacific species of Okenia are similar in exter- nal appearance to O. kendi. Okenia plebeia (Bergh, 1902) and Okenia barnardi Baba, 1937, have a series elongate appendages along the notal margins and possess a single elongate mid-dorsal papilla. The color pattern of living specimens of O. plebeia is unknown. The radula of O. plebeia has a similar formula to that of O. kendi. with 27—28 rows of teeth. The inner lateral teeth of O. plebeia have approximately 30 denticles along the masticatory margin whereas O. kendi has 35-38 denticles. The outer lateral teeth of O. plebeia are rounded but those of O. kendi have a sharply pointed apical extension. The reproductive system of O. plebeia has a short prostate with a muscu- lar portion that is four times the length of the penis. In O. kendi, the prostate is long and the mus- cular portion of the vas deferens is only about twice the length of the penis. Okenia barnardi has a markedly different color pattern to that of O. kendi. It is light brown with a darker brown band near the notal margin that is punctuated by opaque white spots. An opaque white line is present dorsally on the posterior end of the foot. In contrast, O. kendi has a pattern of maroon blotches located on the notum. The anatomy of O. barnardi is largely unde- scribed. The radula was depicted in the original description (Baba 1937). The outer lateral teeth of O. barnardi have a rounded apex whereas there is a pronounced apical denticle on the outer later- als of O. kendi. Okenia virginiae Gosliner, sp. nov. (Figs. 1C, 12-13) Okenia sp. Gosliner 1987: 92, fig. 159. Okenia barnardi Debelius, 1996:196, upper photo, misidentification, not O. barnardi Baba, 1937. TYPE MATERIAL.— HoLotyPe: SAM A35558, one specimen, dissected, Vetchies Pier, Durban, South Africa, 28 April 1982, T.M. Gosliner. ETYMOLOGY.— Okenia virginiae is named for my mother, Virginia Malaney Gosliner, in recognition of her nurturing my earliest interest in natural history. DISTRIBUTION.— This species is known from the Natal, South Africa (Gosliner 1987) and Oman (Debelius 1996). NATURAL History.— Okenia virginiae was found on the underside of a small rock in 2 meters of water. The specimen when dissected was found to be parasitized by three trematodes. EXTERNAL MORPHOLOGY.— The living animal (Fig. 1C) was 10 mm in length. It is 8 mm long in the preserved state. The body is wide and dorsoventrally flattened. There is well-developed, dis- tinct notal border. The foot extends posteriorly and is devoid of notal papillae. The general body color is translucent white. There is a broad mid-dorsal band of brown pigment that runs from ante- rior to the rhinophores to the posterior end of the foot. It is interrupted by an irregular network of white pigment. The notal papillae, rhinophores and gill have irregular maroon bands along their length. There are 8 pairs of elongate notal papillae along the sides of the body. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of three unipinnate branches that are bifid or trifid at the base. The rhinophores are elongate with 21 congested lamellae. The 138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 FIGURE 12. Okenia virginiae sp. nov. (SAMA 35558). A. ventral view of head scale = 0.5 mm. B. Buccal mass, bp = buccal pump, e = esophagus, g = oral glands, r = radula sac, sg = salivary gland; scale = 0.1 mm. C. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina; scale = 0.2 mm. anterior end of the foot contains two elongate rounded lobes that are united medially. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. The foot is broad and is divided anteriorly. A groove separates the foot from the mouth and elongate, triangular oral tentacles (Fig. 12A). DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 12B) with a rounded buc- cal pump directed posteriorly. There are a few elongate oral glands around the mouth. The radular sac is short and extends ventrally from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junction of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregular polygonal plates with a denticulate margin (Fig. 13B). The cuticle expands as it enters the buccal pump. The radular formula is 24 x 1.1.0.1.1. The inner lateral teeth (Fig. 13A) are broad basally with an acutely pointed cusp and 22-33 small, irregular denticles along the masticatory margin. The outer laterals are smaller and reduced (Fig. 13A). They have a short cusp along their outer side. CENTRAL NERVOUS SYSTEM.— The ganglia of the central nervous system are highly concen- trated and surround the esophagus, immediately posterior to the buccal mass. The cerebral and pleural ganglia are largely fused. A sessile eye is present at the base of either cerebral ganglion. The pedal ganglia are smaller than the cerebropleural ganglia and are separated by a short commissure. GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 139 Paired buccal ganglia are situated ventral to the BUN esophagus. » REPRODUCTIVE SYSTEM (Fig. 12C).— The preampullary duct is short and thin and expands into an elongate ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of two folded portions. The distal end of the prostatic segment narrows into an elongate, convoluted ejaculatory portion that terminates in a short, wide penis. There is no distinct penial papilla. Inside the distal end of the penis are penial spines. The vagina exits adjacent to the penis. It is wide and short and connects directly to the spherical bursa copulatrix. Near the base of the vagina, the uterine duct separates and enters the albumen gland. Distal from the base of the bursa copulatrix, a narrow duct connects the bursa with the large, pyriform receptaculum seminis. Pint se Discussion.— This species has been FicurE 13. Okenia virginiae sp. nov. (SAMA 35558). recently erroneously identified as Okenia Scanning electron micrographs. A. Half-row of radular barnardi (Debelius, 1996:196, upper photo). — teeth; scale = 100um. B. Jaw rodlets; scale = 6 um. Okenia virginiae differs from O. barnardi in its color pattern by having dark brown pigment located in the center of the notum rather than submar- ginally and in having a network of white lines rather than opaque white spots. The outer lateral teeth are broad basally with a rounded apex in both O. barnardi and O. virginiae, in contrast to O. kendi, where the outer lateral teeth each have a pointed cusp. The reproductive anatomy of O. barnardi has not been described. Okenia virginiae has a reproductive system with a short vagina with the uterine duct entering the vagina near its base. This arrangement has not been described for any other member of the genus. A Okenia lambat Gosliner, sp. nov. (Figs. 1B, 14-15) TYPE MATERIAL.— Holotype: CASIZ 084283, 1 specimen, dissected, Devil’s Pt., Maricaban Island, Batangas Province, Luzon, Philippines, 10 m depth, 23 February 1992, T.M. Gosliner. ETYMOLOGyY.— The specific epithet Jambat is derived from the Filipino word for net, refer- ring to the network of white markings that is visible through the brown notal pigment. DISTRIBUTION.— This species is known from the Luzon Island, Philippines (present study). NATURAL History.— Okenia lambat has been found under a small stone in 10 meters of water. 140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 Ficure 14. Okenia lambat sp. nov. (CASIZ 084283). A. ventral view of head, scale = 0.5 mm. B. Buccal mass, bp = buccal pump, e = esophagus, r = radula sac, sg = salivary gland: 0.2 mm. C. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis; scale = 0.1 mm. EXTERNAL MORPHOLOGY.— The living animal (Fig. 1B) was 7 mm in length. Preserved, the specimen is 6 mm long. The body is wide and dorsoventrally flattened. There is a well-developed, distinct notal border. The foot extends posteriorly and is devoid of notal papillae. The general body color is opaque yellowish white. There is a broad mid-dorsal band of brown pigment that runs from anterior to the rhinophores to the posterior end of the foot. It is interrupted by an irregular network of white pigment. The notal papillae, rhinophores and gill have irregular maroon spots along their length. There are 8 pairs of elongate notal papillae along the sides of the body. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of three elongate, unipinnate branches. The two lateral branches are undivided and the anterior one is bifid to the base. The rhinophores are elongate with 18 congested lamellae. The anterior end of the foot contains two elongate rounded lobes that are united medially. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. The foot is broad and is divided anteriorly. A groove separates the foot from the mouth and oral tentacles (Fig. 14A). DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 14B) with a massive rounded buccal pump directed posteriorly. There are a few elongate oral glands around the mouth. The radular sac is relatively short and extends posteroventrally from the buccal mass. The esoph- agus is thin and elongate and inserts into the buccal mass immediately ventral to the buccal pump. A rounded, lobate salivary gland is present on either side of the buccal mass anterior to the junc- tion of the esophagus with the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregular rounded rods. The cuticle expands as it enters the buccal pump. The radular formula is 18 x 1.1.0.1.1. The inner lateral teeth (Fig. 15A) are broad basally with an acute- ly pointed cusp and 18-19 denticles along the masticatory margin. The outer laterals are smaller and reduced (Fig. 15B). They have an elongate cusp along their upper margin of the outer side and a rounded lobe basally. REPRODUCTIVE SYSTEM (Fig. 14C).— The preampullary duct is short and thin and expands into a wide saccate ampulla. The ampulla divides into a short, narrow oviduct and the wider pro- static portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nida- mental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens 1s rel- atively narrow and consists of two folded portions. The distal end of the prostatic segment narrows into an elongate, curved ejaculatory portion that terminates in a short, wide penis. There is no dis- tinct penial papilla. Inside the distal end of the penis are penial spines. The vagina exits adjacent to the penis. It is wide and short and connects directly to the spherical bursa copulatrix. Near the base GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 141 of the bursa, the uterine duct separates from the vagina and enters the albumen gland. Also from the base of the bursa copulatrix, an addi- tional narrow duct connects the bursa with the small, rounded receptaculum seminis. DiIscussiION.— Okenia lambat is very sim- ilar in its external appearance to O. virginiae. Both species have a series of lateral papillae with a single medial one and have dark brown dorsomedial pigment interrupted by a network of white lines. The general body color of O. virginiae is translucent white to cream whereas that of O. lambat is opaque yellowish white. Internally there are other differences that dis- tinguish these taxa. In O. lambat, the mastica- tory margin-bearing denticles extend virtually to the apex of the primary cusp, in O. virginiae whereas the denticles terminate well short of the apex. The outer lateral teeth of O. lambat each bear an extended, acutely—pointed cusp, but in O. virginiae the cusp is much shorter and more rounded. Despite the fact that both O. lambat and O. virginiae are unique in having a much-shortened vagina, there are differences in the reproductive anatomy of the two species. in O. lambat the ejaculatory portion of the vas deferens is relatively short and is of uniform diameter throughout its length. In contrast, the elongate ejaculatory vas deferens of O. virgini- ae consists of a thin distal portion that expands into a wider portion that enters the penial bulb. = In O. lambat, the uterine duct joins the vagina FIGURE sy, Okenia lambat sp. nov. (CASIZ 084283). atthe base ofthe receptaculum seminis where- _pmne en micah of mal ah Vi as in O. virginiae it enters the genital atrium at tooth: scale = 8.6um. the base of the vagina. eee SS Okenia liklik Gosliner, sp. nov. (Figs. 1G, 16-17) TYPE MATERIAL.— Ho.otype: CASIZ 168021, one specimen, Barracuda Pt., Tab Island, Madang Lagoon, Madang, Papua New Guinea, 7 February 1988, R.C. Willan. PARATYPES: CASIZ 168022, one specimen, Barracuda Pt., Tab Island, Madang Lagoon, Madang, Papua New Guinea, 7 February 1988, R.C. Willan. CASIZ 168023, one specimen, dissected, Bus Stop, Balayan Bay, Batangas, Luzon Island, Philippines, 10 m depth, 9 May 2001, A. Valdés. Etymology.— The specific epithet /iklik is the Papuan pidgin word for small, in reference to the tiny body size of the mature specimens of this species. DISTRIBUTION.— This species has been found from Papua New Guinea and Luzon Island in the Philippines (present study). 142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 EXTERNAL MORPHOLOGY.— The living animals (Fig. 1G) are small, 3-4 mm in length. The body is elongate and relatively high. There is a well-developed, continuous notal border. Immediately ventral to the notum is a series of small subdermal mantle glands situated in the pos- | 2 Re RS terior half of the body (Fig. 16A). The foot extends posteriorly and is devoid of notal papillae. The — body color is uniformly purplish pink. There is mid-dorsal band of chocolate brown that is inter- rupted by connected or three interrupted, irregular opaque white markings. The gill is uniformly — opaque white. All but the posterior pair of papillae are tipped with an orange subapical marking © and, in some cases, a chocolate brown apex. The posterior pair and the single medial papilla are purple with chocolate brown, but lack orange pigment. There are 8—9 pairs of elongate notal papil- lae along the sides of the body. The two anteriormost pairs of papillae are situated in front of the rhinophores and are anteriorly directed. A single mid-dorsal papilla is present anterior to the gill. It has the form of a crenulate crest rather than a more typical digitiform papilla. The gill consists of 3 unipinnate branches. The rhinophores are elongate with 17-18 congested lamellae. The anterior end of the foot contains two elongate triangular lobes that are united medially. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to — the head. The foot is narrow and is divided anteriorly. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 16B) with a large round- ed, elongate buccal pump directed posteriorly. There are a several elongate oral glands around the FiGurE 16 (left). Okenia liklik sp. nov. (CASIS 168023). A. Lateral view of preserved specimen, a =anus, bm = buccal mass, g = oral glands, mg = mantle glands; scale = 0.2 mm. B. Buccal mass, bp = buccal pump, e = esophagus, r = radula sac; scale = 0.1 mm. C. Reproductive system, am = ampulla, be = bursa copulatrix, p = penis, pr = prostate, rs = receptac- ulum seminis, v = vagina; scale = 0.1 mm. FiGurE 17 (right). Okenia liklik sp. nov. (CASIS 168023). Scanning electron micrograph of radular teeth. A. Inner lat- eral teeth; scale = 3um. B. Inner and outer lateral teeth; scale = 3m. GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 143 mouth. The radular sac is relatively short and extends posteroventrally from the buccal mass. A labial cuticle surrounds the lips at the opening of the mouth. No distinct rodlets are evident around the opening of the buccal cuticle. The radular formula is 12 x 1.1.0.1.1. The inner lateral teeth (Fig. 17A, B) are broad basally with an acutely pointed cusp and 24—26 fine denticles along the masti- catory margin. The outer laterals are smaller and reduced (Fig. 17B) with a single acute cusp. REPRODUCTIVE SYSTEM (Fig. 16C).— The preampullary duct is short and expands into a wide saccate ampulla. The ampulla divides into a short, narrow oviduct and the wider prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albu- men gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic segment of the vas deferens is relatively nar- row and consists of two folded portions. The distal end of the prostatic segment narrows into an elongate, curved ejaculatory portion that terminates in a short, wide penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is wide and short and connects directly to the small, triangular bursa copulatrix. Near the base of the vagina, the short uterine duct separates from the vagina and enters the albumen gland. Also from the middle of the vaginal duct, an addi- tional narrow duct connects the bursa with the large, rounded receptaculum seminis. Discussion.— Okenia liklik has a distinctive purplish pink body color with orange, brown and opaque white markings. No other described species has been described as having glands situated near the posterior margin of the mantle. As in some other species of Okenia, such as O. japonica and O. purpureolineata, the body is elongate and narrow and large oral glands are present sur- rounding the mouth. The relatively large, unifid outer lateral teeth are more elongate than those found in other described species. Okenia rosacea (MacFarland, 1905) (Figs. 18-19) Hopkinsia rosacea MacFarland, 1905:53; 1906:149, pl. 31, figs. 24,25, pl. 21, figs. 97-103. Okenia rosacea (MacFarland, 1905) comb. nov. MATERIAL EXAMINED.— CASIZ 072075, specimens, one dissected, Great Tide Pool, Pacific Grove, California, 1 June 1973, Gary McDonald. DISTRIBUTION.— This species was originally described from Monterey Bay, California (MacFarland 1905) has been found from Coos Bay, Oregon to Isla San Martin, Baja California (Behrens 1991). NATURAL History.— This species feeds upon the cheilostomatous bry- ozoan, Eurystomella sp. (Behrens 1991). DIGESTIVE SYSTEM.— The buccal mass is large and the radular sac is elon- gate, extending well behind the posterior end of the muscular mass. The radular teeth of Okenia rosacea are distinctive. The inner laterals (Fig. 18) are elongate with a recurved apex forming a distinctive hook. The outer laterals are reduced to a small vestigial plate. REPRODUCTIVE SYSTEM (Fig. 19).— The preampullary duct is short and expands into a wide saccate ampulla. The ampulla divides into a short, narrow oviduct and the much wider prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental FiGuRE 18. Okenia rosacea (MacFarland, 1905). (CASIZ 072075) Scanning electron micrograph of inner lateral tooth; scale = 200 um. 144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 glands exit ventrally to the vagina and penis. The prostatic segment of the vas deferens is rel- atively wide and consists of two folded por- tions. The distal end of the prostatic segment narrows into an elongate, convoluted ejaculato- ry portion that terminates in a short, wide penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is narrow and elongate and connects directly to the large, sac- cate bursa copulatrix. Near the distal end of the vagina, the long uterine duct separates from the vagina and enters the albumen gland. Also from the distal end of the vaginal duct, an additional narrow duct connects to the small, saccate receptaculum seminis. Penial hooks were not observed. DISCUSSION.— Okenia rosacea is similar to O. hiroi (Baba, 1938) in its external anatomy and uniform pink coloration. The two species FIGURE 19. Okenia rosacea (MacFarland, 1905). differ markedly in their radular anatomy: (CASIZ 072075). Reproductive system, am = ampulla, be = whereas O. rosacea has elongate, curved inner >¥"S4 copulatrix, fgm = female gland mass, p = penis, pr = 3 aoe : prostate, rs = receptaculum seminis; scale = 1.0 mm. lateral teeth, those of O. hiroi are denticulate and hamate in shape. Only O. rosacea and O. nakamotoensis have elongate inner lateral radular teeth that terminate in a sharply curved apex. Those of O. nakamotoensis are even thinner and more elongate than those found in O. rosacea. Subsequent phylogenetic analysis shows that Hopkinsia is nested within Okenia and is therefore regarded as a synonym of the older taxon, Okenia. Okenia plana Baba, 1960 (Figs. 20B, 21-22) Okenia plana Baba, 1960:80, pl.7, figs 2a—d). Hopkinsia plana (Baba, 1960), Bouchet and Ortea, 1983:231; Rudman and Darvell, 1990:37, pl. If. MATERIAL EXAMINED.— CASIZ 001026, six specimens, two dissected. Cooley’s Landing, East Palo Alto, San Francisco Bay, California, 2 February 1977, D. Chivers. CASIZ 101373, five specimens, two dissected, Berkeley Fishing Pier, San Francisco Bay, California, 6 March 1963, L. Andrews. CASIZ 083837, 1 specimen, dissected, Seafari, Calumpan Peninsula, Batangas Province, Luzon, Philippines, 1-2 m depth, 17 February 1992, T.M. Gosliner. DISTRIBUTION.— This species was originally described from Japan (Baba, 1960) and later recorded from Hong Kong (Rudman and Darvell 1990), California (Steinberg 1963; Behrens 1991), New Zealand and Australia (Rudman 1998). The present specimen extends the range to the Philippines. The specimens from California represents a human introduction (Behrens 1991), NATURAL History.— In the Philippines this species was found under coral rubble in about 6 meters of water. Specimens introduced into San Francisco Bay have been observed on the encrust- ing cheilostomatous bryozoan, Membranipora sp., growing on arborescent algae (present study). EXTERNAL MORPHOLOGY.— The living animals (Fig. 20B) attain 10-15 mm in length. Preserved specimens were 4-9 mm long. The body is broad and ovoid. There is no distinct notal border. The foot is short and does not extend beyond the oval of the general body. The body is uni- GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS a / FiGuRE 20. Living animals. A. Okenia pilosa (Bouchet and Ortea), 1983 (CASIZ 109784), Barracuda Point, Madang, Papua New Guinea. B. Okenia plana Baba, 1960 (CASIZ 083837), Anilao, Luzon, Philippines. C. Okenia kondoi (Hamatani, 2001), Balicasag Island, Bohol, Philippines. D. Okenia nakamotoensis (Hamatani, 2001), Balicasag Island, Bohol, Philippines, photo by G. Williams. All photos by T.M. Gosliner except as otherwise noted. 146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 formly translucent yellowish white with the exception of small brown spots scattered over the surface of the notum and occasional opaque white spots. The notal papillae, rhinophores and gill all possess larger brown and yellow blotches. There are 5 pairs of elongate papillae scattered on either side of the dorsal surface of the body. A single medial papillae is present mid-dorsally anterior to the gill. The gill con- sists of 9-11 unipinnate branches. The rhinophores are relatively short with 12-15 congested lamellae. The anterior end of the foot is deeply indented anteriorly forming a v-shaped edge that encloses the mouth (Fig. 21A). The oral tentacles are indistinct but are separated from the foot by a groove. The geni- tal aperture is situated on the right side of the dorsal surface of the body approximately a third of the length of the body posterior to the head. DIGESTIVE SYSTEM.— The buccal mass is B thick and muscular with a rounded buccal pump situated anteriorly. The radular sac is rel- atively short. The esophagus is thin and elon- gate and inserts into the buccal mass immedi- ately dorsal and anterior to the entrance of the FIGURE 21. Okenia plana Baba, 1960. A. Ventral view of preserved specimen, (CASIZ 083837); scale = 0.6 mm. B. Reproductive system, (CASIZ 001026), am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina; scale = 0.5 mm. radular sac. A rounded, lobate salivary gland is present on either side of the buccal mass at the point where the esophagus enters the mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregular rounded plates. The cuticle expands as it enters the buccal pump. The radular formula is 21—29 x 1.1.0.1.1. in two specimens examined. The inner lateral teeth (Fig. 22) are thin and elongate and with a straight apex. They are devoid of denticles. The outer laterals are small and reduced with two pointed denticles along the outer margin, occasionally with 2—3 smaller denticles along the edge. REPRODUCTIVE SYSTEM (Fig. 21B).— The preampullary duct is short and expands into a wide saccate ampulla. The ampulla divides into a short, narrow oviduct and the narrow vas deferens that expands into a massive, lobate prostatic portion. The oviduct enters the highly folded female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. More distally the lobate pro- static segment of the vas deferens narrows somewhat into a second glandular section that is rela- tively wide and consists of two folded sections. The distal end of the prostatic segment narrows © into an elongate, thin, curved ejaculatory portion that terminates in a narrow penis. There is no dis- tinct penial papilla. The thin vaginal duct exits adjacent to the penis. It is elongate and narrow and connects directly to the large, spherical bursa copulatrix. Near the base of the bursa, another elon- | gate duct enters the small pyriform receptaculum seminis. Also from the base of the receptaculum, ~ the short uterine duct enters the large female gland mass. Discussion.— Okenia plana lacks the pink color of O. rosacea and O. hiroi and is uniformly ~ | ‘ ‘ GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 147 - b> | ae FIGURE 22. Okenia plana (Baba, 1960). Scanning electron micrographs of radular teeth. A. Half row of radular teeth of specimen from San Francisco Bay (CASIZ 001026); scale = 10um. B. Half row of radular teeth of specimen from the Philippines (CASIZ 083837); scale = 10um yellowish white with black spots and opaque white markings on the notum and notal papillae. Its color pattern is similar to that found in O. pilosa Bouchet and Ortea, 1983. The two species differ in the arrangement of the notal papillae and in their radular anatomy. In O. plana there are almost always five pairs of marginal notal papillae and a single mid-dorsal papilla, whereas in O. pilosa there are 35-40 scattered papillae over the surface of the notum. In O. plana the inner lateral teeth are short and evenly arched, whereas in O. pilosa the primary cusp is far more elongate. In O. plana, the outer lateral tooth has two primary denticles with occasional 2—3 smaller denticles. In O. pilosa the outer lateral teeth have up to 14 denticles along their outer margin. There is little anatomical difference in the Californian and Philippine specimens examined here. The primary difference is in the color pattern. The specimens from California have larger, darker brown spots that are more regularly distributed over the surface of the notum and notal papillae. There is little difference in the shape and denticulation of the radular teeth. The reproduc- tive anatomy of this species has not previously been described. The most distinctive features of the reproductive anatomy are the lobed prostate, the extremely narrow ejaculatory portion of the vas deferens and the thin vagina. Okenia pilosa (Bouchet and Ortea, 1983) (Figs. 20A, 23-24) Hopkinsia pilosa Bouchet and Ortea, 1983:227, figs 1-7; Rudman and Darvell, 1990:36, pl. le. Okenia pilosa (Bouchet and Ortea, 1983) comb. nov. MATERIAL EXAMINED.— CASIZ 109784, 10 m depth, Barracuda Point, Tab Island, Madang 148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 Lagoon, Madang, Papua New Guinea, 4 November 1996, T. M. Gosliner. DISTRIBUTION.— This species was originally described from New Caledonia (Bouchet and Ortea 1983) and later recorded from Hong Kong (Rudman and Darvell 1990). The present speci- men extends the range to Papua New Guinea. NATURAL History.— This species is found under coral rubble in about 6 meters of water. EXTERNAL MORPHOLOGY.— The living animals (Fig. 20A) reach 13 mm in length. The pre- served specimen was 11 mm long. The body is broad and ovoid. There is no distinct notal border. The foot is short and does not extend beyond the oval of the general body. The body is uniformly translucent white with the exceptions of a network of opaque white and yellow pigment on the notum and rhinophores. The gill rachises are also translucent yellow. Scattered brown spots are present on the notum. There are 35-40 elongate papillae scattered over the dorsal surface of the body. A single medial papilla is present mid-dorsally and is situated immediately anterior to the gill. The gill consists of 12 bipinnate branches. The rhinophores are relatively short with 7-8 congest- ed lamellae. The anterior end of the foot is deeply indented anteriorly forming a v-shaped edge that A B FIGURE 23 Okenia pilosa (Bouchet and Ortea, 1983) (CASIZ 109784). A. Ventral view of preserved specimen; scale = 1.0 mm. B. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, Vv = vagina; scale = 0.4 mm. encloses the mouth (Fig. 23A). The oral tentacles are indistinct but are separated from the foot by a groove. The genital aperture is situated on the right side of the dorsal surface of the body approx- imately a third of the length of the body posterior to the head. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular with a rounded buccal pump sit- uated anteriorly. The radular sac is relatively short. The esophagus is thin and elongate and inserts into the buccal mass immediately dorsal and anterior to the entrance of the radular sac. A rounded, lobate salivary gland is present on either side of the buccal mass at the point where the esophagus enters the mass. A labial cuticle surrounds the lips at the opening of the mouth. It contains irregu- lar rounded plates. The cuticle expands as it enters the buccal pump. The radular formula is 19 x 1.1.0.1.1. The inner lateral teeth (Fig. 24) are thin and elongate and with a straight apex. They are devoid of denticles. The outer laterals are small and reduced with an approximately 14 denticles along the masticatory margin. REPRODUCTIVE SYSTEM (Fig. 23B).— The preampullary duct is short and expands into a wide saccate ampulla. The ampulla divides into a short, narrow oviduct and the wide prostatic portion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albu- men gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic segment of the vas deferens is relatively wide and consists of two convoluted, folded portions. The distal end of the prostatic segment narrows GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 149 into an elongate, curved ejaculatory portion that terminates in a short, wide penis. There is no distinct penial papilla. The vagina exits adjacent to the penis. It is elongate and narrow and connects directly to the small, spherical bursa copulatrix. Near the base of the bursa, a short duct enters the pyriform receptaculum seminis. Also from the base of the receptacu- lum, the short uterine duct enters the large female gland mass. Discussion.— As in Okenia plana, a translucent body color with mottled brown spots and opaque white pigment makes the liv- ing animals of O. pilosa exceedingly cryptic when encountered on their host bryozoans. The anatomical distinctions between these two taxa are reviewed in the discussion following O. plana. In their original description of H. pilosa, Bouchet and Ortea (1983) revised the systemat- ics of taxa included in Hopkinsia. They trans- ferred Okenia plana Baba, 1960 and ee Ce Ni, a Me ee Hopkinsiella, with its monotypic species, H. FicuRE 24. Okenia pilosa (Bouchet and Ortea), 1983 hiroi, to Hopkinsia. They noted that radular dif- (CASIZ 109784). Scanning electron micrograph of radular Ree teeth; scale = 10um. ferences between Hopkinsia rosacea and Hopkinsiella hiroi were of specific value rather than generic value. They noted that the primary fea- ture that distinguishes Hopkinsia from Okenia is the absence of a distinct pallial ridge in the for- mer. Bouchet and Ortea stated that members of the two genera feed on different prey with species of Hopkinsia feeding on Bryozoa whereas species of Okenia feed on tunicates. However, it is evi- dent from many studies, including the present work, that several species of Okenia are also special- ists on bryozoans rather than tunicates (Burn 1967; Rudman and Darvell 1990; present study). Subsequent phylogenetic analysis supports the inclusion of all of these taxa within a single mono- phyletic taxon, Okenia. ‘ee Okenia nakamotoensis (Hamatani, 2001) (Figs. 20D, 25-26) Okenia sp. 1 Gosliner, Behrens and Williams, 1996: 171, fig. 604. Hopkinsia nakamotoensis Hamatani, 2001:151, fig. 1 A-E. Okenia nakamotoensis (Hamatani, 2001) comb. nov. MATERIAL EXAMINED.— CASIZ 096238, 5 specimens, 2 dissected, Balacasag Island, Bohol, Philippines, 29 m. depth, 21 March 1994, T. M. Gosliner. CASIZ 096244, 2 specimens, Balacasag Island, Bohol, Philippines, 34 m. depth, 22 March 1994, T. M. Gosliner. CASIZ 117028, 1 speci- men, 20 m depth, R. Buoy Pinnacle, Kwajalein Atoll, Marshall Islands, 28 March 1992, S. Johnson. CASIZ 120736, 3 specimens, Cement Ship Pinnacle, Enewetak Atoll, Marshall Islands, 21 August 1983, S. Johnson. CASIZ 120732, 4 specimens, 1 dissected, Enewetak Atoll, Marshall Islands, 5 June 1983, S. Johnson. CASIZ 120733, 5 specimens,15 m depth, R. Buoy Pinnacle, Kwajalein PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 150 Atoll, Marshall Islands, 18 June 1982, S. Johnson. CASIZ 120734, 5 specimens, R. Buoy Pinnacle, Kwajalein Atoll, Marshall Islands, 22 June 1982, S. Johnson. CASIZ 120735, 7 specimens, 1-20 m depth, R. Buoy Pinnacle, Kwajalein Atoll, Marshall Islands, J. Johnson. CASIZ 093922, 5 specimens, un- named reef near Koon Island, East Banda Sea, Indonesia, 17-30 m depth, 14 November 1993, P. Fiene Severns. DISTRIBUTION.— This species is known from the Kuroshima Island, Japan (Hamatani 2001) Indonesia, Bohol and Cebu Islands (Philippines) and Enewetak and Kwajalein Atolls (Marshall Islands) (Gosliner, Behrens, and Williams 1996; present study). NATURAL HISTORY.— This species is found on outer reef faces in 10-30 meters of water, where it feeds upon the bright red bry- ozoan, Tropidozoum cellariforme Harmer, 1957 (Gosliner, Behrens, and Williams 1996). EXTERNAL MORPHOLOGY.— The living animals (Fig. 20D) attain 20 mm in length. Preserved specimens range in size from 3—13 mm. The body is elongate and wide. There is FIGURE 25. Okenia nakamotoensis (Hamatani, 2001) (CASIZ 096238). A. Buccal mass, bp = buccal pump, e = esophagus, sg = salivary gland, r = radula sac; scale = 1.0 mm. B. Reproductive system, am = ampulla, be = bursa cop- ulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina; scale = 0.5 mm. no distinct notal border. The foot extends pos- teriorly and is devoid of notal papillae. The body is uniformly red with the exceptions of the gill rachises and the base of the notal papillae, which are pinkish white. The gill lamellae and apices of the notal papillae are the same color as the remainder of the body. There are five pairs of elongate notal papillae along the sides of the body. Two of these are situated anterior to the rhinophores. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of 34 unipinnate branches. The rhinophores are elongate with 13—20 congested lamellae. The anterior end of the foot is rounded and forms a projection anterior to the mouth. The oral tentacles are fused to form an oral veil. The genital aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular (Fig. 25A) with a rounded buc- cal pump situated anteriorly. The radular sac is elongate and is approximately equal in length to the rest of the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass imme- diately dorsal and anterior to the entrance of the radular sac. A rounded, lobate salivary gland is present on either side of the buccal mass at the point where the esophagus enters the mass. A labi- al cuticle surrounds the lips at the opening of the mouth. It contains irregular rounded plates. The cuticle expands as it enters the buccal pump. The radular formula is 16 x 1.1.0.1.1. The inner lat- eral teeth (Fig. 26A) are extremely thin and elongate and with a hooked apex (Fig. 26B). They are devoid of denticles. Some of the older teeth are eroded and have a simple rounded or irregularly fractured apex. The outer laterals are small and reduced (Fig. 26C) with an anteriorly directed cusp. CENTRAL NERVOUS SYSTEM.— The ganglia of the central nervous system are highly concen- trated and surround the esophagus. The cerebral and pleural ganglia are largely fused. A sessile eye GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS Itsy FIGURE 26. Okenia nakamotoensis (Hamatani, 2001) (CASIZ 096238). Scanning electron micrographs. A. Inner lateral teeth, scale = 150 um. B. Apices of inner lateral teeth; scale = 34 um. C. Outer lateral tooth; scale = 20 um. is present at the base of either cerebral ganglion The pedal ganglia are smaller than the cerebro- pleural ganglia and are separated by a short commissure. Paired buccal ganglia are situated ventral to the esophagus. REPRODUCTIVE SYSTEM (Fig. 25B).— The preampullary duct is short and thin and expands into an ovoid ampulla. The ampulla divides into a short, narrow oviduct and the wide prostatic por- tion of the vas deferens. The oviduct enters the lobate female gland mass that consists of a small albumen gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventrally to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of approximately three coils. The distal end of the prostatic segment narrows into a short, straight ejaculatory portion that terminates in an indistinct, but slightly wider penis. There is no dis- 52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 tinct penial papilla. Inside the distal end of the penis are penial spines. The thin elongate vagina exits adjacent to the penis. It connects to the large, spheroid bursa copulatrix. From the base of the bursa a short duct connects to the small, serial receptaculum seminis. From the proximal end of the receptaculum the uterine duct joins with the female gland mass. DISCUSSION.— The original description of O. nakamotoensis (Hamatani, 2001) includes illus- trations of the external anatomy, the buccal mass and the radular teeth. All of these features, as orig- inally described, are consistent with the present material from the Marshall Islands and the Philippines. The shape of the inner lateral teeth closely resembles that described for the typé species of the genus Hopkinsia, O. rosacea. In both taxa, the inner laterals are thin and elongate with a hook—shaped apex, although the teeth are thinner and more elongate in O. nakamotoensis than in O. rosacea. Okenia nakamotoensis differs externally from O. rosacea. It is bright red with white pigment situated basally on the notal appendages and red apices whereas O. rosacea is uni- formly pink in color. Also, O. rosacea has more numerous randomly distributed notal appendages while O. nakamotoensis has five pairs of lateral appendages and a single mid-dorsal appendage anterior to the gill. The reproductive systems of the two species are similar in configuration with several notable differences. Although the ampulla of O. rosacea is larger than that found in O. nakamotoensis, the prostatic portion of the vas deferens of O. nakamotoensis is more highly con- voluted, consisting of several loops, than that of O. rosacea. Okenia kondoi (Hamatani, 2001) (Figs. 20C, 27-28) Sakishimaia kondoi Hamatani, 2001:153, Fig 2a-e. Okenia kondoi (Hamatani, 2001) comb. nov. MATERIAL EXAMINED.— CASIZ 158272, 5 specimens, Balacasag Island, Bohol, Philippines, 29 m. depth, 21 March 1994, T.M. Gosliner. CASIZ 106551, 7 specimens, 1 dissected, Agus Point, Mactan Island, Cebu, Philippines, 20 m. depth, 27 April 1996, M. Miller and T.M. Gosliner. CASIZ 158273, 4 specimens, CASIZ 120732, 4 specimens, Enewetak Atoll, Marshall Islands, 5 June 1983, S. Johnson. DISTRIBUTION.— This species is known from the Kuroshima Islands, Japan (Hamatani, 2001), Bohol and Cebu Islands (Philippines) and Enewetak and Kwajalein Atolls (Marshall Islands) (Gosliner, Behrens, and Williams 1996; present study). NATURAL History.— This species is found together with O. nakamotoensis on outer reef faces in 10-30 meters of water, where it feeds upon the bright red bryozoan, Tropidozoum cellari- forme Harmer, 1957. EXTERNAL MORPHOLOGY.— The living animals (Fig. 20C) attain 20 mm in length. Preserved specimens range in size from 3—13 mm. The body is elongate and narrow. There is no distinct notal border. The foot extends posteriorly and is devoid of notal papillae. The body is uniformly red with the exceptions of the gill rachises and the base of the notal papillae, which are pinkish white. The gill lamellae and apices of the notal papillae are the same color as the remainder of the body. There are four pairs of elongate notal papillae along the sides of the body. Two of these are situated ante- rior to the rhinophores and are more thin and elongate than the more lateral ones. A single medial papilla is present mid-dorsally anterior to the gill. The gill consists of 3—5S unipinnate branches. The rhinophores are elongate with 10—20 congested lamellae. The anterior end of the foot expands into two rounded lobes that are united as a small hood that surrounds the mouth (Fig. 27A). The geni- tal aperture is situated on the right side of the body approximately a third of the length of the body posterior to the head. GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 153 FIGURE 27 (left). Okenia kondoi (Hamatani, 2001) (CASIZ 106551). A. ventral view of head; scale = 0.2 mm. B. Reproductive system, am = ampulla, be = bursa copulatrix, fgm = female gland mass, p = penis, pr = prostate, rs = recep- taculum seminis, v = vagina; scale = 0.3 mm. FIGURE 28 (right). Okenia kondoi (Hamatani, 2001) (CASIZ 106551). Scanning electron micrographs. A. Radular teeth; scale = 35 um. B. Penial armature; scale = 100 um. DIGESTIVE SYSTEM.— The buccal mass is thick and muscular with a rounded buccal pump sit- uated anteriorly. The radular sac is short and protrudes only slightly from the buccal mass. The esophagus is thin and elongate and inserts into the buccal mass immediately dorsal and anterior to the entrance of the radular sac. A rounded, lobate salivary gland is present on either side of the buc- cal mass at the point where the esophagus enters the mass. Lobate oral glands are present at the anterior end of the buccal mass around the mouth. The radular formula is 25 x 1.1.0.1.1. The inner lateral teeth (Fig. 28A) are truncate with a broad base and an elongate masticatory margin. The tooth contains 16-17 elongate denticles. The outer laterals are small and reduced to elongate, fan-shaped, quadrangular plates. REPRODUCTIVE SYSTEM (Fig. 27B).—The preampullary duct is short and thin and expands into an elongate, cylindrical ampulla. The ampulla is curved and divides into an elongate, narrow oviduct and the wide prostatic portion of the vas deferens. The oviduct enters the rounded, elon- gate receptaculum seminis. The lobate female gland mass consists of a small albumen. gland, a lobed membrane gland and a larger, smooth mucous gland. These nidamental glands exit ventral- ly to the vagina and penis. The prostatic portion of the vas deferens is wide and consists of two folded adjacent portions. The distal end of the prostatic segment narrows into a long, curved ejac- ulatory portion that terminates in a distinct conical penis. Protruding from the penial bulb is a cutic- ular portion bearing rows of chitinous hooks (Fig. 28B). The thick short base of the massive bursa 154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 copulatrix functions as the vagina. Near its base is a short duct that bifurcates and joins the albu- men gland and the elongate receptaculum seminis. DIscussion.— Hamatani (2001) described Sakishimaia as a distinct genus based on the pres- ence of a cylindrical slender body, absence of a pallial margin and velum and the absence of sec- ond (outer) lateral teeth. Scanning electron micrographs of the radula of O. kondoi (Fig. 28A) clear- ly show the presence of outer lateral teeth. The only characteristic unique to O. kondoi is the more body shape that is more elongate than species of Hopkinsia. This feature is shown to represent an autapomorphy for this taxon. The phylogeny presented here indicates that O. kondoi is nested with- in Hopkinsia, which in turn is nested within Okenia. Consideration of Sakishimaia as a distinct genus renders Hopkinsia paraphyletic. Similarly maintaining Hopkinsia makes Okenia paraphylet- ic. Thus, I consider it more prudent to consider Sakishimaia as a synonym of Okenia to maintain the monophyly of this taxon. PHYLOGENETICS To date, no phylogenetic analysis has been undertaken for members of the genera Okenia, Hopkinsia, Hopkinsiella and Sakishimaia. In the absence of phylogenetic studies there has been no objective test of the monophyly of these taxa. Recent works including these taxa (Bouchet and Ortea 1983; Hamatani 2001) have based systematic revisions exclusively on traditional rather than phylogenetic criteria. It is therefore important to conduct a preliminary phylogenetic analysis of members of these taxa. To facilitate such an analysis, morphological and anatomical studies were undertaken. Most data used to construct the matrix for phylogenetic analysis are derived from the morphological studies presented here. In addition to the species specifically discussed here, speci- mens of Okenia angelensis Lance, 1966 were examined (CASIZ 072351, 4 specimens, one dissect- ed, Morro Bay, California, May-June 1981, David Behrens). Morphological description for other species were taken from the literature to represent morphological variation and geographical breadth. Specifically, anatomical details for the following taxa were derived from the following morphological studies: Diaphorodoris luteocincta (Schmekel and Portmann, 1982), Goniodoris nodosa (Thompson and Brown, 1984), Okenia impexa (Marcus, 1958; Schmekel and Portmann 1982), O. zoobotryon (Marcus, 1957, as O. evelinae; Valdés and Ortea 1995), O. quadricornis (Valdés and Ortea, 1995), O. elegans (Valdés and Ortea, 1995), O. aspersa (Schmekel and Portmann, 1982), O. mediterranea (Valdés and Ortea, 1995) and O. hiroi (Baba, 1938). To determine the phylogenetic relationships of species of Okenia, Hopkinsia, Hopkinsiella and Sakishimaia, twenty morphological characters and one biological character were examined for 23 taxa, including Diaphorodoris luteocincta and Goniodoris nodosa as outgroup taxa. These taxa were chosen primarily because their anatomy is reasonably well described. These taxa also repre- sent an excellent sample of the morphological variation that is known to exist in these taxa and also represent global geographical coverage of the known distribution of the Goniodorididae. All char- acters were treated as unordered. The following characters were considered: 1. Body shape: In the outgroup taxa and in many species of Okenia and Hopkinsia the body is short and ovoid (OQ). In some species of Okenia and Sakishimaia kondoi the body is narrow and elongate (1). Mantle between notum and foot: In the outgroup and species of Okenia, the lateral margins of the body provide a surface distinct from the notum and foot (0). In these taxa, the genital opening is situated on this lateral surface of the right side of the body. In species of Hopkinsia and Sakishimaia, there is no distinct lateral surface and the notum slopes to the foot (1). In these taxa, the genital opening is on or near the lat- eral margin of the notum. Notal ridge: In the outgroup taxa the notal edge is demarcated by a distinct thickening of tissue around its to eS) GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 35) entire margin (0), whereas all of the ingroup taxa lack any thickening around the mantle margin (1). 4. Mid-dorsal crest: In the outgroup taxa and Okenia mediterranea, there is a distinct mid-dorsal longitudi- nal crest that extends between the rhinophores and gill (0). In the remaining taxa, no thickening of tissue is present (1). Caudal crest: The outgroup taxa, O. mediterranea and O. quadricornis have a caudal crest extending along the middle of posterior portion of the foot (0). This feature is absent in the remaining taxa studied here (1). 6. Oral tentacle shape: In Diaphorodoris luteocincta and several species of Okenia, the oral tentacles are rounded in shape (0). In other species, including Goniodoris nodosa and most species of Okenia, the oral tentacles are triangular and tentacular (1), whereas in species of Hopkinsia and Sakishimaia, the oral ten- tacles are fused with the mouth region to form an oral veil (2). 7. Buccal pump: In most species examined here, including the outgroup taxa and most of the ingroup taxa, the buccal pump portion of the buccal mass is moderately expanded (0), whereas in Okenia virginiae and O. lambat it is greatly expanded into a massive structure (1). 8. Radular sac: In the outgroup and vast majority of the ingroup taxa, the radular sac is much shorter than the rest of the buccal mass (0) whereas in Hopkinsia rosacea and H. nakamotoensis it is almost as long or longer than the buccal mass (1). 9. Marginal papillae: In the outgroup taxa, the margin of the notum may be undulate or irregular, but distinct lateral papillae are absent (0). In all of the ingroup taxa, elongate lateral papillae extend outward from the sides of the body (1). 10. Mid-dorsal papillae: In the ingroup taxa there are numerous conical papillae evenly distributed over the surface of the notum. This arrangement is also found in some species of Okenia and Hopkinsia pilosa. Mid-dorsal papillae are absent in O. aspersa and O. quadricornis (1). Okenia purpureolineata is unique in having two mid-dorsal papillae anterior to the gill (2). In many species of Okenia, Hopkinsia and Sakishimaia, there is a single mid-dorsal papillae anterior to the gill (3). 11. Rhinophoral lamellae: In the outgroup taxa and the vast majority of ingroup taxa, the rhinophores are char- acterized by having crowded lamellae along most of their length (0). In Okenia angelensis, O. impexa, O. pellucida, O. zoobotryon and O. brunneomaculata the rhinophoral lamellae are well separated from each other (1). 12. Inner lateral tooth shape: In the outgroup and most ingroup taxa the inner lateral teeth have a broad shape with a thick pointed cusp (0). In Hopkinsia rosacea and H. nakamotoensis the inner lateral teeth are high- ly modified and much elongated (1). 13. Inner lateral tooth denticulation: The ingroup taxa, Hopkinsia hiroi, and many species of Okenia the mas- ticatory margin of the inner lateral tooth bears numerous fine teeth (0). Some species of Okenia and Sakishimaia (O. impexa, O. pellucida, O. zoobotryon, O. brunneomaculata, S. kondoi) have inner lateral teeth with fewer, coarser denticles (1). In Hopkinsia rosacea, H. pilosa, H. plana and H. nakamotoensis the inner lateral teeth entirely lack denticles (2). 14. Inner lateral tooth apex: In the outgroup taxa and the vast majority of ingroup taxa the apex of the inner lateral tooth is gently curved with an acute apex (0). In Hopkinsia rosacea and H. nakamotoensis, the apex of the inner laterals are sharply curved with a rounded apex (1). 15. Outer lateral tooth: The shape of the outer lateral teeth varies considerably in the taxa considered. In the outgroup and most of the ingroup taxa the outer laterals have a single apex (0). In Okenia impexa and O. brunneomaculata the outer lateral teeth have a deeply divided tooth with two sharp projections (1). The outer lateral tooth of Hopkinsia rosacea and H. nakamotoensis is greatly reduced in size to a tiny plate (2). The outer lateral tooth of Sakishimaia kondoi is flat and quadrangular in shape (3). The outer lateral tooth of Hopkinsia plana and H. pilosa is multidenticulate (4). 16. Oral glands: In the outgroup taxa and the vast majority of ingroup taxa studied here, there are minute oral glands at the anterior base of the buccal mass, adjacent to the mouth (0). Large oral glands are present in Okenia japonica, O. liklik and O. purpureolineata (1). The development of oral glands remains unknown for Okenia aspersa, O. zoobotryon and Hopkinsia hiroi (?). 17.Ampulla length: The ampulla provides the hermaphroditic duct from the ovotestis to the genital aperture. It also serves as an endogenous sperm storage organ. In the outgroup taxa and many ingroup taxa, the ampulla is elongate (0). In Okenia japonica, O. virginiae, O. impexa, O. brunneomaculata, O. lambat, O. tn 156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 purpureolineata, H. pilosa, H. plana, H. rosacea, H. nakamotoensis and Sakishimaia kondoi, it is short (1). The form of the ampulla is not known for O. aspersa or O. quadricornis (?). 18. Receptaculum seminis: In the outgroup and the vast majority of ingroup taxa the diameter of the receptac- ulum seminis is about the same size as the bursa copulatrix (0) whereas in species of Hopkinsia and Sakishimaia kondoi, the diameter of the receptaculum seminis is much smaller than the bursa copulatrix (1). The form of the receptaculum seminis is not known in Okenia quadricornis, O. aspersa, O. zoobotry- on or Hopkinsia hiroi (?). 19. Uterine duct: In the outgroup taxa and the majority of ingroup taxa the uterine duct is proximal to the ~ receptaculum seminis (0). In O. japonica, O. virginiae, O. lambat, O. kendi and O. liklik, the uterine duct emerges from the vagina (1). The uterine duct position is unknown for Okenia quadricornis, O. aspersa or Hopkinsia hiroi (?). 20.In the outgroup taxa and the vast majority of the ingroup taxa, the vagina is long (0). In Okenia virginiae, O. lambat and O. kendi, the vagina is short (1). The length of the vagina is unknown for Okenia quadri- cornis, O. aspersa, O. zoobotryon or Hopkinsia hiroi (?). 21.Members of the taxa studied either feed on bryozoans (0) or tunicates (1). Food associations are unknown for Okenia liklik (?). In order to determine phylogenetic relationships, the following data were entered into a data matrix using MacClade 3.01 (Table 1). This analysis was performed using PAUP 3.1.1, for the data matrix. All characters were treated as un—ordered and unweighted. A Heuristic search using step- wise addition from 1000 random starting trees was undertaken using the DELTRAN option, result- ing in 200 trees with a length of 41, a consistency index of 0.683 and retention index of 0.824. The strict consensus of these trees is shown in Figure 29. Bremer support analysis was conducted to test the robustness of the tree. DISCUSSION The preliminary phylogeny presented here is moderately well supported, given the number of characters. Bremer support values are included in Figure 29. Most nodes have a Bremer support value of 1. The two most basal nodes have a Bremer support value of at least 2. Additionally, the clade that contains Hopkinsia rosacea and H. nakamotoensis also has a value of at least 2. Further analysis was not possible, owing to the large number of trees that were produced (more than 60,000) after adding trees with two more steps. Adding a third step to the tree lengths far exceed- ed the computing ability of the computer dedicated to phylogenetic analysis. The greater robust- ness of these basal clades is especially important for the subsequent systematic conclusions that are discussed. The phylogenetic hypothesis presented in Figure 29 suggests several important aspects regard- ing the phylogeny of Okenia, Hopkinsia, Hopkinsiella and Sakishimaia. These taxa collectively form a monophyletic taxon characterized by two synapomorphies (presence of marginal papillae and absence of a notal ridge). Okenia mediterranea is the sister taxon to all the remaining mem- bers of the ingroup. Several large monophyletic clades are formed by species traditionally includ- ed in Okenia. This first clade contains the European Atlantic taxa, O. elegans, O. aspersa and O. quadricornis and is the sister group to all the remaining taxa. This clade is supported by an ecolog- ical synapomorphy, that all these taxa feed on tunicates rather than bryozoans. It is also interesting to note that Valdés and Ortea (1995) considered O. aspersa and O. quadricornis as the same species. This issue needs to be more fully studied with comparative material from Atlantic and Mediterranean waters. Members of this clade have short, rounded oral tentacles, as in the outgroup taxon Diaphorodoris luteocincta. Okenia leachi (Alder and Hancock, 1854) may also be a mem- ber of this clade. It also has rounded oral tentacles and feeds upon tunicates, but its anatomy remains largely unknown. 157) GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS Pe a ee eee gO Op Om Sp Oa ee Oar |0 ae (Rex ara o (cor CRs at Eee | PRPPPP PPP Pr Perr Perr rp aac Ch OR OE Ch ROR i [Osa deo [ Beem aes) Gor arse are o | 0 0 er 0 0 2 OH PPT Pee ee eee eye CCC i (ee 9 ak) Peon [Om Ole Oa yor | OF son 0 [xorg os Soe pe peor me | 0 | [ee eee is eee (RO or | 0, om Romi ON Lm ae | Oe One| O [Oo hate ope Toteyepe | ete 0/0] 0 oe por pepepe piss L 1/0] O [Qaroy kog One Ore Omi Om sO [Oa On 0 Spepr peer ep o OFS F ORs OR RCS iCie Oke | Oe Os Om | BOR OF OS RO SRO, SOs ROG OME OS | SO.s|cOm ROR pee} ibe 900J ndwe! jes {09es oeuul JOU! 0 D z a al ° p poqg 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 The sister group of these three species contain three distinct clades. The first of these, support- ed by two apomorphies (tentacular oral tentacles, sparse rhinophoral lamellae) contains O. ange- lensis, O. zoobotryon, O. pelllucida and the sister species, O. impexa and O. brunneomaculata. Okenia impexa and O. brunneomaculata share two synapomorphies, coarse denticles on the inner lateral tooth and a bifid outer lateral tooth. Additionally, all of these taxa have a translucent white body color with varied brown markings. They also all appear to feed upon ctenostomatous bry- ozoans. It appears that O. mija Burn, 1967 may also be a member of this clade. Its internal anato- my remains largely unknown, but it has a similar color pattern, feeds on ctenostomatous bryozoans and has sparse rhinophoral lamellae. Collectively members of this clade inhabit a wide geograph- ical area and include subtropical and tropical Atlantic, Eastern Pacific and Indo-Pacific taxa. Understanding the detailed phylogeny of this clade may also shed some additional light on the his- torical biogeography of this taxon. The second clade is supported by three synapomorphies (absence of a distinct lateral surface of the body, oral tentacles fused with the mouth region to form an oral veil and the presence of a short ampulla). Sakishimaia kondoi is nested within the species traditionally placed within Hopkinsia. Hopkinsiella hiroi is basal within this clade, but lacks any autapomorphies. Hopkinsia rosacea and H. nakamotoensis are sister species, a relationship that is supported by five synapo- morphies. These taxa are all Pacific in their distributions, from the Indo-Pacific tropics and sub- tropics and temperate Japan and California. The third clade is restricted to the Indo-Pacific tropics and adjacent subtropical waters of Japan. It is supported by three synapomorphies, the presence of tentacular oral tentacles, the pres- ence of a single mid-dorsal papillae and the presence of short ampulla. This clade is divided into two sister clades. One of these, contains Okenia liklik, O. purpureolineata and O. japonica. These three species have an elongate body and large oral glands. These taxa are restricted to the margins of the western Pacific. The other subclade contains O. kendi, and the sister species, O. lambat and O. virginiae. This clade is supported by the presence of a short vaginal duct and a uterine duct that emerges from the vagina. Okenia barnardi Baba, 1937, also appears to be a member of this clade. This is suggested by its body shape, arrangement of lateral papillae and the presence of a single medial papillae anterior to the gill. The anatomy of this species remains largely undescribed. Several systematic conclusions are evident from this phylogenetic analysis. On the basis of this preliminary phylogeny, Sakishimaia must be considered to be a junior synonym of Hopkinsia, since it is nested within Hopkinsia and maintaining it would render Hopkinsia paraphyletic. The monophyly of the ingroup is supported when only single outgroup taxa are included (either Diaphorodoris luteocincta or Goniodoris nodosa) and the other taxon is included in the ingroup. The monophyly of Hopkinsia (including Sakishimaia kondoi) is well supported. However, Okenia is paraphyletic when Hopkinsia is maintained, as the latter taxon is nested well within Okenia. One possible systematic solution is to create a series of subgeneric names in order to main- tain Hopkinsia. This does not seem to add any information not already contained in the phylogeny and adds unnecessary names to the nomenclature. A second solution is to include all of the ingroup taxa within a single monophyletic taxon, Okenia. In this case, Hopkinsia, Hopkinsiella and Sakishimaia would be junior synonyms of Okenia. It is with regret that I feel compelled to adopt this latter solution. Hopkinsia is a much beloved name that honors the founding of the Hopkins Marine Station of Stanford University and was erect- ed by Frank Mace MacFarland, former Stanford Professor of Biology and President of the California Academy of Sciences. Nevertheless, it is the most simple solution that preserves mono- phyly and does not create a nomenclature that is unnecessarily complicated. Therefore, Hopkinsia, Hopkinsiella and Sakishimaia are here considered as junior synonyms of Okenia. GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS PaUPLIE PEW BIUeIO suehaje eIuaxO esiedse BIUSYO siusooupenb elueyO Ipuay BUS} JEQUIE| B1UEXO eeiuiBia eIUexO eyeeuljeindind eiueyO IPH BlUeHO eoaiuodel e1uexO JOPUOH BIEWIYSEXES sisueojoweyeu eisuydoyH} eBeoesol BISUIydoH eup|d eisurdoy esojid eisundoy louly Byersupjdoy uofy0go0z BIUexO epionjed eiuexO eyeinoewosuUTG BIUaYO exedwi BIUexO sisugjaGue e1uexO Bsopou siOpoiuos 2y9uID08}N| suOpoloUdeig 10 17 Ww) 19 19 10 10 13 13 14 13 13 17 13 10 19 18 eS 17 i 21 4} a4 312+ FIGURE 29. Strict consensus tree of preliminary phylogeny. Apomorphies are indicted by character numbers included in the analysis. Characters which exhibit reversal are underlined. Bremer support values for each node are indicated as large numerals on the right side of each node. Bremer values not indicated on figure are 1. 159 160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 5 At least 12 additional described species of Okenia have not been fully studied for the anatom- ical characters included in this study. At least four undescribed species of Okenia are known to occur in the eastern Pacific tropics and two undescribed species are known from Australian waters. These taxa should be examined and incorporated into subsequent phylogenetic studies. Additional phylogenetic studies of the remaining goniodoridid and onchidoridid genera are also to necessary to produce a broader understanding of the phylogeny and systematics of these poorly understood dorid nudibranchs. ACKNOWLEDGMENTS This research has been supported by a grant from the National Science Foundation, PEET Program (DEB-9978155). Fieldwork was supported by the California Academy of Sciences, the Christensen Fund and the Christensen Research Institute and the South African Museum. Kit Stewart also provided financial support for some fieldwork. Specimens were also collected by sev- eral colleagues including Jerry Allen, Dave Behrens, Bob Bolland, Yolanda Camacho, Shireen Fahey, Pauline Fiene Severns, Mike Ghiselin, Rebecca Johnson, Scott Johnson, Gary McDonald, Mike Miller, Angel Valdés, Yvonne Vallés and Gary Williams. Their assistance and support is greatly appreciated. LITERATURE CITED Basa, K. 1937. Opisthobranchia of Japan (II). Journal of the Department of Agriculture, Kyushu Imperial University 5:289-344. Basa, K. 1938. Opisthobranchia of Kii, Middle Japan. Journal of the Department of Agriculture, Kyushu Imperial University 6(1):1-19. Basa, K. 1949. Opisthobranchia of Sagami Bay. wanami Shoten, Tokyo, Japan. 211pp. BaBA, K. 1960. The genera Okenia, Goniodoridiella and Goniodoris from Japan (Nudibranchia- Goniodorididae). Publications of the Seto Marine Biological Laboratory 8(1):79-83. BEHRENS, D. 1991 Pacific Coast Nudibranchs. Sea Challengers: Monterey. 107 pp., 217 plates. BEHRENS, D. 2001. http://slugsite.us/bow/nudwk275.htm BeERGH, R. 1902. The Danish Expedition to Siam 1899-1900. I. Gasteropoda Opisthobranchiata. Det Kongelige Danske Videnskabernes Selskabs Skrifter. 6 Raekke. Naturvidenskabelig og Matematisk Afdeling 12(2):153-218. BoucuHET, P., AND J. ORTEA. 1983. A new Hopkinsia feeding on Bryozoa in the South Pacific (Mollusca: Opisthobranchia). Venus 42(3):227—233. Burn, R.F. 1967. Descriptions of two new species of Okenia (Nudibranchia, Doridacea) from south-eastern Australia. Proceedings of the Royal Zoological Society of New South Wales 1965/1966:52-57. DeBELIUS, H. 1996. Nudibranchs and Sea snails. Indo-Pacific Field Guide. 1KAN Unterwasserarchivy, Frankfurt,Germany. 321 pp., 1000 unnumbered figs. GosLiner, T.M. 1987. Nudibranchs of Southern Africa. Sea Challengers, Monterey, California. 136 pp., 268 pls. GOsLINER, T.M., D. BEHRENS, AND G WILLAMS. 1996. Coral Reef Animals of the Indo-Pacific. Sea Challengers, Monterey, California. 314 pp., 1103 pls. GOSLINER, T.M., S. JOHNSON, AND H. BERTSCH. 1986. Additions to the opisthobranch gastropod fauna of the Hawaiian Islands. Western Society of Malacologists, Annual Report 18:14-17. HAmataNl, I. 2001. Two new species of Goniodorididae (Opisthobranchia; Nudibranchia) with a new genus from Kuroshima Island, Okinawa, Japan. Venus 60(3):151—156. KurRIHARA, T. 1999. Sea Slug Forum: hittp:/Avww.seaslugforum.net/display.cfmbase=okenpell&id=1399 (first record of Okenia pellucida from Japan). GOSLINER: TROPICAL INDO-PACIFIC GONIODORIDIDAE NUDIBRANCHS 161 Lance, J. 1966. New distributional records of some northeastern Pacific Opisthobranchiata (Mollusca: Gastropoda) with descriptions of two species. The Veliger 9(1):69-81. MACFARLAND, F.M. 1905. A preliminary account of the Dorididae of Monterey Bay, California. Proceedings of the Biological Society of Washington 18:35-54. MACFARLAND, F.M. 1906. Opisthobranchiate Mollusca from Monterey Bay, California and vicinity. Bulletin of the Bureau of Fisheries 25:109-151. Marcus, E. 1957. On Opisthobranchia from Brazil (2). Zoological Journal of the Linnean Society 43(292): 390-486. RuUDMAN, W.B.1987. The genus Trapania (Nudibranchia, Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53(2):189-212. RuDMAN, W.B. 1998. http:/Avww.seaslugforum.net/hopkplan.htm. RUDMAN, W.B., AND DARVELL, B.W. 1990. Opisthobranch molluscs of Hong Kong. Part 1. Goniodorididae, Onchidorididae, Triophidae, Gymnodorididae, Chromodorididae, (Nudibranchia). Asian Marine Biology 7:31-79. SCHMEKEL, L., AND A. PORTMANN. 1982. Opisthobranchia des Mittlemeeres, Nudibranchia und Saccoglossa. Faune flora del Golfo di Napoli. Monografia della Stazione Zoologica di Napoli, no. 40, 410 pp., 36 pls. Springer-Verlag, Heidelberg, Germany. Steinberg, J. 1963. Notes on the opisthobranchs of the West Coast of North America II. Further nomenclat- ural changes in the Order Nudibranchia. The Veliger 6(2):63-67. THOMPSON, T., AND G. BRown. 1984. Biology of Opisthobranch Mollusks, vol. 2. Ray Society, London, UK. 229 pp., 41 pls. VaLpEs, A., AND J. ORTEA. 1995. Revised taxonomy of some species of the genus Okenia, Menke, 1830 (Mol- lusca: Nudibrancia) from the Atlantic Ocean, with the description of a new species. The Veliger 38(3): 223-234. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 6, pp. 162-168, 3 figs., 1 table April 22, 2004 Eptatretus lakeside sp. nov., a New Species of Five-Gilled Hagfish (Myxinidae) from the Galapagos Islands Michael M. Mincarone! and John E. McCosker? ! Museu Oceanografico do Vale do Itajai, Univali, CP 360, Itajai, SC, Brazil, 88302-202. Email: mincarone @ be.univali.br; 2 California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599. Email: jmccosker@ calacademy.org. We describe Eptatretus lakeside sp. nov. from a deepwater (762 m) specimen trapped off Fernandina Island, Galapagos Islands. The new species differs from all known Eptatretus in having: five pairs of gill pouches; 3-cusp multicusps in anterior and posterior rows; 6 unicusps in each anterior and posterior row; 36 total cusps; 19 tail pores; 88 total pores; palatine tooth triangular; and body coloration pinkish-orange. A key to the species of Galapagos hagfishes is provided. The myxinoid fauna of the Galapagos Archipelago was unknown until 1995 when the California Academy of Sciences (CAS) / Harbor Branch Oceanographic Institute (HBOI) expedi- tion made numerous deepwater collections using the submersible Johnson Sea-Link. During that cruise, three species of Eptatretus, E. grouseri, E. mccoskeri, and E. wisneri were trapped and all were described as new (McMillan 1999). A subsequent expedition in 1998 involving CAS, HBOI, and the National Museum of Natural History (USNM), again using the submersible Johnson Sea- Link, captured additional specimens of hagfishes, including a single remarkable specimen of Eptatretus, which 1s described herein as new. Hagfishes are rare in the eastern tropical Pacific but abundant in the temperate and subtemper- ate eastern Pacific. In fact, the extensive deepwater collections off Central America and the Galapagos made by the U.S. Fish Commission Steamer Albatross in 1891 (Garman 1899) resulted in but one hagfish specimen. It was captured over a rocky bottom in 730 fathoms (1335 m) in the southern end of the Gulf of Panama and described as Myxine circifrons Garman, 1899. In contrast to the Albatross, the successful capture of hagfishes by the recent expeditions can be explained by the difficulty that the Albatross expedition had in trawling over unusually rocky terrain, and by the undeniable benefits provided by manned submersibles in exploring and collecting in complex deepwater habitats. Currently, thcve are 30 recognized species of Eptatretus, 12 of which are known from the east- ern Pacific (McMillan and Wisner 1984; Wisner and McMillan 1988, 1990; McMillan 1999). Undoubtedly more species remain to be discovered. MATERIALS AND METHODS The type of the new species as well as the holotypes of all other hagfishes from the Galapagos Islands are deposited in the Department of Ichthyology of the California Academy of Sciences (CAS), San Francisco. Other type specimens examined are deposited in the Scripps Institution of Oceanography (SIO), La Jolla, and National Museum of Natural History (USNM), Washington, — Dic 162 MINCARONE AND MCCOSKER: NEW GALAPAGOS HAGFISH 163 Methods of measuring and counting follow those of Fernholm and Hubbs (1981) and McMillan and Wisner (1984). The names of anatomical structures follow Wisner and McMillan (1995) and Mok (2001). Length of the specimens (in mm) is given as total length (TL), the distance from the front of the rostrum to the end of the caudal finfold. All other measurements are given in percentage of TL. Counts of gill pouches (GP), gill apertures (GA), and cusps are taken for both sides, whereas slime pore counts are from the left side. Measurements and counts are given in Table 1 and compared with data of all other Eptatretus species previously known from the Galapagos Islands. We provide drawings from the anterior and posterior sets of cusps, including the palatine tooth, which are not usually useful characters to identify hagfish species, but in this case they aid in distinguishing species. We provide a key to the Eptatretus species from the Galapagos Islands, which was modified from McMillan (1999). Eptatretus lakeside Mincarone and McCosker, sp. nov. (Figs. 1, 3; Table 1) MATERIAL EXAMINED.— Holotype: CAS 201880 (field number JM-155), an immature female, 275 mm TL, from off Cabo Douglas, NW Fernandina Island, Galapagos Islands, 00°17’30’S, 91°39’36’W, 762 m depth, collected on 17 July 1998 by David Pawson and Godfrey Merlen, using a baited metallic minnow trap, deployed while aboard the submersible R/V Johnson Sea-Link (JSL Dive 3101). DIAGNOsIs.— Eptatretus lakeside can be distinguished from its congeners by a combination of the followings characters: five pairs of gill pouches; 3-cusp multicusps in anterior and posterior rows; 6 unicusps in each anterior and posterior rows; 36 total cusps; 19 tail pores; 88 total pores; palatine tooth triangular; and body coloration pinkish-orange. DESCRIPTION OF THE HOLOTYPE Body subcylindrical and slender, slightly deeper than wide at prebranchial, branchial, and trunk regions and strongly compressed at tail. Rostrum blunt- ly rounded; nasopharyngeal duct cylindrical, tube-like, slightly projecting. One pair of conspicu- ous nasal-sinus papillae on the inner dorsal surface of the nasal sinus. Three pairs of barbels on the head, the first two pairs about equal in size and adjacent to the nasopharyngeal duct; the third pair is immediately adjacent to the oral cavity. Ventral finfold (VFF) conspicuous, 2 mm high, begin- ning 25 mm behind the last gill aperture and extending backward to the cloaca. Caudal finfold quite thin and rounded, extending around tail to dorsal surface, ending about over cloaca. Total length 275 mm; prebranchial length 68 mm; branchial length 17 mm; trunk length 145 mm; tail length 50 mm; body width at prebranchial region 14 mm; body depth at mid-trunk includ- ing VFF 20 mm; body depth excluding VFF 18 mm; body depth over cloaca 15 mm; tail depth 17 mm. Three-cusp multicusps in anterior and posterior rows of cusps; 6 unicusps in each anterior and posterior row; total cusps 36. Cusps long, slender, and pointed; palatine tooth triangular (Fig. 3). A segmentally arranged row of slime pores on each side, extending from beyond head to behind cloa- ca. Prebranchial pores 15; branchial pores 4; trunk pores 50; tail pores 19; total pores 88. Four branchial pores are intercalated with five gill apertures on both sides. No slime pore associated with the pharyngocutaneous duct. Five pairs of gill pouches corresponding to five pairs of gill apertures. Last branchial duct confluent with the pharyngocutaneous duct on the left side. First pair of gill pouches lies posterior to end of dental muscle; ventral aorta branches at the second gill pouch. Body color in alcohol pinkish; barbels and face the same color as body; eyespots present but inconspicuous; ventral finfold pale; caudal finfold with a narrow pale margin. In life, the specimen was pinkish-orange. DisTRIBUTION.— Known only from the Galapagos Islands. 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 6 FiGureE 1. Dorsal view of preserved holotype of Epratretus lakeside (CAS 201880; 275 mm TL). ETYMOLOGY.— Named /akeside, a noun in apposition. We take great pleasure in honoring the Lakeside Foundation of California, which has generously supported the work of the senior author and many other foreign scholars. Key to Galapagos Species of Eptatretus la. Five or six gill pouches and. apertures on each side. ..... <2... 2.225 a5 ohne 2 lib: Eight gill pouches and apertures on each side. ::..2... 22.25 +25.24%-2 eo Roe 3 2a. Body coloration pinkish-orange; 3/3 multicusp pattern; 36 total cusps; 88 total pores Uae rN Ie ies eps hw Gls eu teen als chayel ens eae eae Eptatretus lakeside, new species 2b. Body coloration dark brown; 3/2 multicusp pattern; 44-48 total cusps; 71—79 total pores RRS a fete wei: AA ae hs oes Dele Ged ee eins Eptatretus grouseri McMillan, 1999 3a. Prebranchial length 24-26 % of TL; 3/3 multicusp pattern; 48-51 total cusps; 14-15 pre- branchialypores Ari5hi02e. seis Se Om. Pee Bee Eptatretus mccoskeri McMillan, 1999 3b. Prebranchial length 19-23 % of TL; 3/2 multicusp pattern; 44 total cusps; 9 prebranchial pores By Se. Oe) ed 8) gs 2 os ns to een ee eile meal Eptatretus wisneri McMillan, 1999 COMPARISON.— Three five-gilled species of Eptatretus were previously known: E. grouseri McMillan, 1999 from the Galapagos Islands, E. profundus (Barnard, 1923) from South Africa, and E. eos Fernholm, 1991 from the Tasman Sea. All have a 3/2 multicusp pattern, whereas E. lakeside has 3/3. Eptatretus lakeside also differs from E. grouseri in the following characters, respectively: MINCARONE AND MCCOSKER: NEW GALAPAGOS HAGFISH 165 FIGURE 2. Left lateral view of preserved Eptatretus grouseri (CAS 201882; 420 mm TL). number of unicusps on each row (6 vs. 8-10); total cusps (36 vs. 44-48); tail pores (88 vs. 71-79); shape of palatine tooth (triangular and depressed vs. conic [Fig. 3]); and its body coloration (pink- ish-orange [Fig. 1] vs. dark brown [Fig. 2]). Also, the teeth of E. lakeside are more slender and more elongate than those of FE. grouseri (Fig. 3). Eptatretus lakeside differs from E. profundus in the following characters, respectively (based on the redescription of the holotype of E. profundus bo FiGuRE 3. Dentition of (A) Eptatretus lakeside (CAS 201880, 275 mm TL), and (B) Eptatretus grouseri (CAS 201882, 420 mm TL). 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 6 TABLE |. Measurements and counts of Eptatretus from the Galapagos Islands. E. lakeside E. grouseri E. mccoskeri — E. wisneri Holotype Holotype Non-types (2) Holotype — Holotype Total length TL (mm) Dies 370 315-420 310 355 Measurements in % of TL Prebranchial length 4.7 20.3 21.0-22.2 25.8 18.9 Branchial length 6.2 8.1 6.3-6.5 10.0 PAI Trunk length 50.9 57.0 54.0-55.7 48.4 Sil Tail length 18.2 14.6 16.9-17.5 18S 77 Body width 5.0 a) 5.4-6.3 TES 6.5 Body depth Inc. ventral finfold 2 6.8 7.9-9.0 oh 8.0 Exc. ventral finfold 6.4 6.8 7.9-8.8 8.2 Vall Over cloaca 5.4 Dyd 6.4-6.5 kes 6.5 Tail 6.0 6.8 7.6-7.9 8.9 8.3 Counts Cusps Multicusps 3/3 3/2 3/2 3/3 3/2 Anterior unicusps* 6+6 9+9 9-10 10+10 9+9, Posterior unicusps* 6+6 8+8 9-9 9+10 9+9 Total cusps 36 44 46-48 51 46 Slime pores, left side Prebranchial 15 12 11-12 13 9 Branchial 4 + 5-5 7 7 Trunk 50 46 42-48 43 43 Tail 19 15 13-14 10 14 Total pores 88 77 71-79 73 15 Gill apertures* 5+5 5+5 5-6 8+8 8+8 Gill pouches* 54+5 54+5 5-6 8+8 8+8 * Left + right count. by Strahan 1975): number of unicusps on each row (6 vs. 8); tail pores (19 vs. 15); and its body coloration (pinkish-orange vs. dark brown). Despite the multicusp condition, E. eos shares some characters with E. lakeside: both have a pinkish body coloration; a long tail (about 18% of TL); an elongated tube-like nostril (longer in E. eos); and the same number of unicusps (six). However, E. lakeside differs from E. eos in the number of prebranchial pores (15 vs. 26), trunk pores (50 vs. 75-77), tail pores (19 vs. 26-27), and total pores (88 vs. 128-130) (based on the original descrip- tion of the holotype of E. eos by Fernholm 1991). The other two Eptatretus species from the Galapagos Islands, E. mccoskeri and E. wisneri, can be easily separated from E. lakeside by their having eight pairs of gill pouches and a brownish-black body coloration. The triangular shape of MINCARONE AND MCCOSKER: NEW GALAPAGOS HAGFISH 167 the palatine tooth is probably an autapomorphy of E. lakeside (Fig. 3A). Nemamyxine kreffti McMillan & Wisner, 1982, is the only other hagfish that has a similar palatine tooth, but its tooth is more depressed and has a rounded point (Mincarone 2001). In most hagfish species the palatine tooth has a conical shape like a bird’s claw (Fig. 3B). The Galapagos species of Eptatretus, albeit based on a very limited sample size, appear to be stratified by depth. The only known E. mccoskeri were trapped at 215 m on a seamount SE of San Cristobal Island on the eastern edge of the archipelago. The other species were captured off Fernandina Island (F), along the western edge of the archipelago, and from Seymour (S) Island (= James) in the center of the archipelago. They were trapped at the following depths: E. wisneri, 512-563 m (FP); FE. grouseri, 648-722 m (F and S); and E. lakeside, 762 m (F). REMARKS.— The location of capture, Cabo Douglas, is located along the NW corner of Fernandina Island and drops steeply into deep water (more than 1000 m depth at a distance less than 2 km from shore). The specimen was collected using a galvanized metal minnow trap baited with fish flesh and set from the submersible at 0845 and retrieved approximately one hour later. The bottom was nearly flat with a slight downward slope (<10°) and covered with fine gray sedi- ment and occasional lava boulders (~ 0.5 m in diameter) in the vicinity of other large lava reefs. The temperature at depth was 8°C and there was a current of ~0.3 knot. The habitat is further described by Iwamoto and McCosker (2001) who described a new macrourid, Coryphaenoides gypsochilus, from that site. Also observed at and near that locality were several pelagic holothuri- ans (Pelagothuria nanatrix Ludwig), hagfishes (Eptatretus grouseri McMillan and E. wisneri McMillan), catsharks (Apristurus spp.), combtooth dogfish (Centroscyllium nigrum Garman), chi- maeras (Hydrolagus spp.), witch-eels (Facciolella equatorialis (Gilbert)), viperfish (Chauliodus sloani Bloch and Schneider), grenadier (Nezumia loricata loricata (Garman)), cardinalfish (Epigonus merleni McCosker and Long), batfishes (Dibranchus erinaceus (Garman)), ateleopo- dids (Guentherus altivelis (Os6rio)), bythitids (Diplacanthopoma jordani (Garman)), and an unde- scribed scorpionfish (Phenacoscorpius sp.). COMPARATIVE MATERIAL— Eptatretus grouseri: CAS 86428, holotype, 370 mm TL, Punta Espinosa, Fernandina Island, Galapagos Islands, 00°14’36’S, 91°26’36”W, 722 m; SIO 97-77, paratype, 138 mm TL, taken with holotype; CAS 201882, 2 specimens, 315-420 mm TL, Seymour Island, Galapagos Islands, 00°21’42”S, 90°15’00”’W, 648 m, 25 July 1998, Johnson Sea-Link (JSL Dive 3113). Eptatretus mccoskeri: CAS 86431, holotype, 310 mm TL, San Cristobal Island, Galapagos Islands, 01°06’19”S, 89°06’56”W, 215 m; SIO 97-75, paratype, 290 mm TL, taken with holotype; USNM 344905, paratype, 284 mm TL, taken with holotype. Eptatretus wisneri: CAS 86429, holotype, 355 mm TL, Cabo Hammond, Fernandina Island, Galapagos Islands, 00°27’56’S, 91°37'33”W, 563 m; SIO 97-76, paratype, 316 mm TL, Galapagos Islands, 00°17’30’S, 91°38’54”W, 512 m. ACKNOWLEDGEMENTS We thank: Eliecer Cruz and Mario Piu of the Galapagos National Park for permission to per- form research in the GNP; Robert Bensted-Smith and the staff of the CDRS for assistance in the Galapagos; the Harbor Branch Oceanographic Institute and its sub pilot Don Libertore; Carole Baldwin, Godfrey Merlen, and David Pawson, colleagues while in Galapagos; the technicians, col- lection managers and curators of the California Academy of Sciences, Scripps Institution of Oceanography, and the National Museum of Natural History for assistance with specimens; Rafael A. Brandi for preparation of the figures; the IMAX Corporation, Al Giddings, David Clark, and Mandalay Media Arts for their assistance; Jules M.R. Soto for support of the senior author; the 168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 6 Lakeside Foundation, and its President Paul L. Davies, Jr., for supporting the work of the senior author at the California Academy of Sciences; Tomio Iwamoto for his critical review of this man- uscript; and William N. Eschmeyer for his advice and assistance with this project. LITERATURE CITED FERNHOLM, B. 1991. Eptatretus eos: a new species of hagfish (Myxinidae) from the Tasman Sea. Japanese Journal of Ichthyology 38(2):115-118. FERNHOLM, B., AND C.L. Husss. 1981. Western Atlantic hagfishes of the genus Eptatretus (Myxinidae) with description of two new species. Fishery Bulletin 79(1):69-83. GARMAN, S. 1899. The Fishes. Jn Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer “Albatross” during 1891, Lieut.-Commander Z.L. Tanner, U.S.N. Commanding. XXVI. Memoirs of the Harvard Museum of Comparative Zoology 24:1-431. Iwamoto, T., AND J.E. McCosker. 1991. Notes on Galapagos grenadiers (Pisces, Gadiformes, Macrouridae), with the description of a new species of Coryphaenoides. Revista Biologial Tropical 49(supl. 1):21—27. McMILLAN, C.B. 1999. Three new species of hagfish (Myxinidae, Eptatretus) from the Galapagos Islands. Fishery Bulletin 97(1):110-117. McMILLAN, C.B., AND R.L. WISNER. 1984. Three new species of seven-gilled hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean. Proceedings of the California Academy of Sciences 43(16):246-267. MINCARONE, M.M. 2001. Further description of the hagfish Nemamyxine kreffti McMillan & Wisner, 1982 (Agnatha, Myxinidae). Mare Magnum 1(1):19-22. Mok, H.-K. 2001. Nasal-sinus papillae of hagfishes and their taxonomic implications. Zoological Studies 40(4):355—364. STRAHAN, R. 1975. Eptatretus longipinnis, n. sp., a new hagfish (family Eptatretidae) from South Australia, with a key to the 5—7 gilled Eptatretidae. The Australian Zoologist 18(3):137-148. WISNER, R.L., AND C.B. MCMILLAN. 1988. A new species of hagfish, genus Eptatretus (Cyclostomata, Myxinidae), from the Pacific Ocean near Valparaiso, Chile, with new data on E. bischoffii and E. poly- trema. Transactions of the San Diego Society of Natural History 21(14):227-244. WISNER, R.L., AND C.B. MCMILLAN. 1990. Three new species of hagfishes, genus Eptatretus (Cyclostomata, Myxinidae), from the Pacific coast of North America, with new data on E. deani and E. stoutii. Fishery Bulletin 88(4):787—-804. WISNER, R.L., AND C.B. MCMILLAN. 1995. Review of new world hagfishes of the genus Myxine (Agnatha, Myxinidae) with descriptions of nine new species. Fishery Bulletin 93(3):530-550. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 7, pp. 169-173, 2 figs., 2 tables April 22, 2004 A New Species of Finless Snake Eel (Anguilliformes: Ophichthidae) from Ascension Island, with Comments on Ichthyapus acutirostris John E. McCosker California Academy of Sciences, Golden Gate Park, San Francisco, California 94118; Email: jmccosker@ calacademy.org Ichthyapus insularis, new species, is described from the intertidal of Ascension Island. It differs from its widespread Atlantic congener, I. ophioneus, in its vertebral number and tail length, and from all other Jchthyapus in its vertebral number, body proportions, and preopercular pore condition. The identity of I. acutirostris Brisout de Barneville (1847), the generic type, is aligned with specimens from Japan. In preparing the ophichthid section of the Fishes of the Western North Atlantic, Eugenia Bohlke, Jim Bohlke, and I examined a myriad of finless ophichthids (McCosker et al. 1989) and discovered that one species, Ichthyapus ophioneus, was widely distributed from Bermuda to Brazil, and that its only Atlantic congener was an undescribed species from Ascension Island. In prepara- tion for the forthcoming ophichthid chapter of FAO’s Fishes of the Eastern Central Atlantic, I now describe that new species. In preparing this description, I believe that I have helped to solve the long-lasting conundrum, “what is the identity of Ichthyapus acutirostris Brisout de Barneville (1847)?” MATERIALS AND METHODS Type specimens of the new species are deposited in the Department of Ichthyology of the California Academy of Sciences (CAS), San Francisco. Measurements are straight-line (point to point) and made with dial calipers and recorded to the nearest 0.1 mm. Body length comprises head and trunk lengths; head length is measured from the snout tip to the posterodorsal margin of the gill opening; trunk length is taken from the end of the head to mid-anus. Vertebral counts (which include the hypural) are taken from radiographs. Preanal vertebrae are counted until mid-anus. Institutional abbreviations follow the Standard Symbolic Codes for Institutional Research Collections in Herpetology and Ichthyology (Leviton et al. 1985). Ichthyapus insularis McCosker, sp. nov. (Figs. 1-2; Tables 1-2) MATERIAL EXAMINED.— HoLotyPe: USNM 214480, an immature male, 427 mm TL, from Southwest Bay, Ascension Island, in 0-1 m, collected by S. Olson on 12 July 1970. PARATYPES: USNM 375444, 8(227-423 mm TL), collected with the holotype. CAS 218803, 387 mm TL, col- lected with the holotype. ANSP 158903, 2 (166-244 mm TL), from southwest side of English Bay, Ascension Island (07°53’48”S, 14°23’06”W), collected among rocks and sand by W.D. Anderson on 16 July 1980. 169 170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 7 DIAGNOsIs.— An elongate, finless species of sphagebranchin (sensu McCosker 1977) ophichthine with: tail 57-60% and head 9—11% of total length; 4 preopercular pores and 5 pores in supratemporal canal; teeth conical, uniserial on jaws and vomer; body mostly colorless in preser- vative; and mean vertebral formula -45-123.5, total vertebrae 121-126. COUNTS AND MEASUREMENTS OF HOLOTYPE (IN MM).— Total length 427; head 46.8; trunk 130.2; tail 250; body depth at gill openings 11.5; body width at gill openings 11.1; body depth at anus 9.8; body width at anus 9.6; snout 6.5; tip of snout to rictus 15.0; eye diameter 1.2; interor- bital distance 2.8; gill opening length 7.3; isthmus width 2.2. Vertebral formula -44-121. 120 later- al-line pores, 8 in branchial region, 46 pores before the anus. DESCRIPTION OF THE HOLOTYPE.— Body elongate, nearly cylindrical throughout, snout and tail tip sharply pointed, depth at gill openings 37 in TL. Branchial basket notably wider and deep- er than body. Head and trunk 2.4 in TL; head 9.1 in TL, 2.8 in trunk. Snout sharply pointed, its underside flattened and bisected by a groove nearly to anterior nostrils. Lower jaw short, included, upper and lower lips meet when mouth is closed. Mouth moderately elongate. Rictus well behind rear margin of eye. Eye minute, 12.1 in upper jaw and 39 in head, its center above middle of upper jaw. Anterior nostril non-tubular, a convoluted opening on underside of snout, about midway between tip of snout and tip of lower jaw when mouth is closed. Posterior nostril with an irregular rim, Opening inside of mouth. Branchial open- ings low, ventral, with a median fold forming a pouch; branchial region expanded, creating a bulbous region in posterior half of head. Head pores (Fig. 1) conspicuous. Single median interorbital and temporal pores. Supraorbital pores | + 4, infraorbital pores 5 + FiGuRE 1. Lateral view of the holotype of Ichthyapus 2, supratemporal pores 5, lower jaw pores 4, — insularis (USNM 214480, 427 mm TL). Arrows indicate preopercular pores 4. (The head pores of the 11 locations of the median interorbital and the temporal pores. paratypes are nearly identical to those of the holotype, save 2 with 5 rather than 4 right lower jaw pores and | with 6 rather than 5 Supratemporal pores.) 120 lateral-line pores, 8 before gill opening, 46 before anus. Teeth (Fig. 2) uniserial, small, conical and sharp. Intermaxillary with a chevron of 11 teeth, followed by a short gap and a linear row of 13 vomerine teeth. Jaw teeth nearly sube- qual, small and close-set, 24-28 teeth in upper jaw and 22—26 teeth in lower jaw. Body mostly colorless in isopropanol. Faint brown pigment smudges surround the second and third supraorbital pores and over- laying the nape and cheeks; a faint brown pos- teriorly-directed “v” arises behind posterior mid-orbit and extends to the interorbital pore. S1ZE.— The largest specimen examined is ¥ 07 dean ARES EN spec- SUES 2). Pinson of the holotype of chefs insu- laris (USNM 214480, 427 mm TL) (upper jaw, left; lower imens appears (examined radiographically, but jaw, right). The lips have been slightly retracted to expose the not by dissection) to be a mature female. teeth and posterior nostrils. MCCOSKER: NEW SPECIES OF FINLESS SNAKE EEL ETYMOLOGy.— From the Latin insularis, of an island, in reference to its apparently endemic location. DISTRIBUTION.— Known only 171 TABLE |. Counts and proportions (in thousandths) of the holo- type and 11 paratypes of Ichthyapus insularis. TL= total length. HL= head length. from Ascension Island. é pee SMe | Mean bar kencee = REMARKS.— Inasmuch as species TL (mm) = 166-427 of Ichthyapus lack fins and, particular- HL/TL 99 90-110 ly in preservative, display little or no Head and trunk/TL 415 401-426 differences in coloration, the most use- —_Tajj/TL, 585 401-426 ful ase . Sea species Depth at gill opening/TL 5 20-29 seems to be the preopercular pore (POP) condition, ee aera pepe aT ae ane and body/tail proportions. Those char- snout ae ee acters show limited variation as EYe/HL 25 19-29 demonstrated by an examination of 77 _ Interorbital/HL 65 Saad specimens of J. ophioneus that had a_ Gill Opening/HL 138 113-166 range of total vertebrae from 125-139, — Isthmus/HL 43 32-61 and the POP condition of 75 specimens __ Preanal vertebrae 45 44-46 was 4 and only two had 3 pores Total vertebrae 123.5 121-126 (McCosker et al. 1989). On that basis, Ichthyapus insularis appears to be most similar to 1. ophioneus (Evermann and Marsh), a wide- spread western Atlantic species from Bermuda, Bahamas, Florida, the Greater Antilles, and St. Helena Island. The two, however, differ in their vertebral number (121—126 vs. 125-139) and the comparative length of head and trunk to total length (.40-43 vs. .36—40). Specimens of the new species were compared to and separable from /. ophioneus from St. Helena Island. And, although not visible on all specimens, the faint dorsal head coloration of J. insularis is lacking in all speci- mens examined of /. ophioneus. The new species can be differentiated from its Pacific and Indian Ocean congeners on the basis of characters listed in Table 2. The only recent treatment of Ascension Island shorefishes is that of Lubbock (1980). In it he mentions but did not examine an Ascension Island specimen of “Sphagebranchus ophioneus found washed up on the beach by the Shackleton-Rowett expedition.” I have not examined that specimen either, but I believe that it will prove to be /. insularis. Lubbock listed 71 shorefish species at Ascension and found 11 to be endemic. Recognizing that /. insularis is indeed endemic, the degree of endemism of his ichthyofaunal list is thereby ele- vated to 12 species or 17%. More recent collecting at Ascension and the reidentification of many museum specimens (for example, two additional ophichthids, Callechelys bilinearis and Phaenomonas longissima, are now known from Ascension) will undoubtedly modify Lubbock’s analysis. COMPARATIVE MATERIAL EXAMINED.— 1062 specimens of Ichthyapus ophioneus, 50-479 mm TL, from 86 lots from Bermuda, Bahamas, Florida, West Indies, Cuba, Puerto Rico, Lesser Antilles, Panama, Belize, Venezuela, Brazil and St. Helena, including the holotypes of Sphagebranchus ophioneus (USNM 49526) and Sphagebranchus conklini (FMNH 57731). Those specimens are listed in McCosker et al. (1989:325). On the Identity of Ichthyapus acutirostris The provenance of the type specimen of Ichthyapus, I. acutirostris Brisout de Barneville (1847), is unknown in that the type specimen (MNHN 2119, two were described, but only one 172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 7 TABLE 2. Characteristics of the species of Jchthyapus. Abbreviations are: 1, this study; 2, McCosker and Castle 1986: 3, Blache and Bauchot 1972; 4, Hatooka et al. 1995; 5, Randall and McCosker 1975; 6, McCosker et al. 1989; 7, McCosker 2002; POP = preopercular pore condition; body is the proportion of head and trunk (in hundredths); * = holotype. POP Vertebrae Body/TL Source I. acuticeps 4 101-108 47-50 1D I. acutirostris* 3 133 40 1,3 I. Japan 3 129-133 42-44 4 I. Easter Island 3 130-134 42-45 5 I. insularis 4 121-126 - 41-43 1 I. ophioneus 4 125-129 3540 6 I. selachops 4 137-144 38-40 i ee eae 4 120-130 30-41 1 ee 3 or4 120-124 39-43 1 (Hawaiian archipelago) remains) lacks locality information other than “Haute-Mer, Océan équatorial.” The specimen was collected by “M. Rang” (Monsieur Paul-Charles-Alexandre-Léonard Rang, 1793-1843), a French malacologist who collected primarily in the tropical Atlantic but also visited India, Madagascar (Bauchot et al. 1990:121), and Japan (P. Béarez, in litt., 7 Jan. 2004). There is no indication the he traveled to the South Pacific. Since its description, /chthyapus acutirostris has been ascribed to specimens from St. Helena Island (Cadenat and Marchal 1963) and from Baja California, Mexico (Blache and Bauchot 1972). The type of /. acutirostris has 133 vertebrae and three preopercular pores (POP). It thereby does not agree with any Atlantic species. Blache and Bauchot compared the type specimen to a speci- men of [chthyapus from Baja California (MNHN 1893-58) and suggested that the two were con- specific. I have examined both specimens and find the latter to be Ichthyapus selachops (Jordan and Gilbert 1882), a widespread eastern Pacific species known from Baja California to Ecuador, which has 137-144 vertebrae (n = 17) and four POP. (Table 2 lists the POP and vertebral condition of all known Jchthyapus.) The only species with three POP are /. acutirostris, some specimens of Hawaiian J. vulturis (the relationship of Hawaiian /. vulturis to those across the Indo-Pacific requires further analysis), and specimens of Jchthyapus from Easter Island (identified as /. vulturis by Randall and McCosker 1975) and Japan (identified as /. vulturis by Hatooka et al. 1995). Vertebral numbers of all Hawaiian /. vulturis are lower than those of /. acutirostris, and on that basis I presume that their identity would reside with specimens now known either from Easter Island or Japan. In that the collector never visited the south Pacific (few if any collections had been made at or near Easter Island before the 20th century) but had traveled to Japan (aboard the Levant and after 1836), I sug- gest that the remaining holotype of /. acutirostris is of the same species as the two specimens iden- tified by Hatooka et al. (1995) from Japan. A comparison of the proportions, pore patterns, and den- tition as seen in the illustration of the Japanese specimen (Hatooka et al. 1995, fig. 2) with the illus- trations of the holotype of /. acutirostris (Blache and Bauchot 1972, figs. 14-15) demonstrates that MCCOSKER: NEW SPECIES OF FINLESS SNAKE EEL 173 they are identical. Jchthyapus vulturis is known from Japan. I have examined a specimen from Ogasawara Islands (BPBM 35175, 99 mm TL) that has 120 vertebrae and four POP. The identity of the Easter Island specimens remains undetermined. And finally, although species of I[chthyapus live in shallow water and are easily captured with rotenone and occasionally captured by dredge sampling, it is possible that additional unknown taxa remain that could be identified with /. acu- tirostris. ACKNOWLEDGMENTS I particularly thank the late Eugenia BOhlke (ANSP) for her contributions to our work with the FWNA ophichthid eels. Thanks are also due: Marie Louise Bauchot and Phillipe Béarez of MNHN for their assistance with /. acutirostris; the curatorial staffs of AMNH, ANSP, BMNH, BPBM, CAS, FMNH, MCZ, SIO, UMML, and USNM for their assistance with specimens; William Eschmeyer (CAS) for advice concerning nomenclature; Mysi Hoang (CAS) for assistance with the figures; and David Smith (USNM) and Tomio Iwamoto (CAS) for reviewing portions of this man- uscript. LITERATURE CITED Baucuot, M. L., J. DAGET, AND R. BAucHoT. 1990. L’ichtyologie en France au début du XIXéme siécle. Vhistoire naturelle des poissons du Cuvier et Valenciennes. Bulletin du Muséum national d Histoire naturelle, 4e sér., 12(1):3-142. BLACHE, J., AND M. L. BAucHoT. 1972. Contribution a la connaissance des Poissons Anguilliformes de la cOte occidentale d’ Afrique. Bulletin de l’IFAN, sér. A, 34(3):692-773. BRISOUT DE BARNEVILLE. 1847. Note sur un nouveau genre d’Anguilliformes. Revue Zoologique 1847:219-220. CADENAT, J., AND E. MARCHAL. 1963. Résultats des campagnes océanographiques de la Reine-Pokou aux iles Sainte-Héléne et Ascension. Poissons. Bulletin de l’IFAN, sér. A, 25(4):1235-1315. HatTooKa, K., T. YOSHINO, AND A. ONO. 1995. A rare ophichthid eel, Ichthyapus vulturis, from Izu Peninsula, Japan (Pisces: Ophichthidae). Bulletin of the Osaka Museum of Natural History 1995(49):19-22. Leviton, A.E., R.H. Grpss, JR., E. HEAL, AND C.E. DAwson. 1985. Standards in herpetology and ichthyolo- gy: part I. Standard symbolic codes for institutional resources collections in herpetology and ichthyology. Copeia 1985:802—832. LupBBock, R. 1980. The shore fishes of Ascension Island. Journal of Fish Biology 17:283-303. McCosker, J.E. 1977. The osteology, classification, and relationships of the eel family Ophichthidae. Proceedings of the California Academy of Sciences ser. 4, 41(1):1-123. McCosker, J.E. 2002. Notes on Hawaiian snake eels (Pisces: Ophichthidae), with comments on Ophichthus bonaparti. Pacific Science 56(1):23—34. McCosker, J.E., E.B. BOHLKE AND J.E. BOHLKE. 1989. Family Ophichthidae. Pages 254-412 in Fishes of the Western North Atlantic, Part Nine, Vol. One: Orders Anguilliformes and Saccopharyngiformes. Sears Foundation for Marine Research, Yale University, New Haven, Connecticut. McCosker, J.E., AND P.H.J. CasTLe. 1986. Family Ophichthidae. Pages 176-186 in M. Smith and P. Heemstra, eds., The Sea Fishes of Southern Africa. Macmillan Publ., Johannesburg, South Africa. RANDALL, J.E., AND J.E. McCosker. 1975. The eels of Easter Island with a description of a new moray. Los Angeles County Museum of Natural History Contributions in Science No. 264. 32 pp. Copyright © 2004 by the California Academy of Sciences San Francisco, California, U.S.A. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 55, No. 8, pp. 174-183, 3 figs. April 22, 2004 A Synopsis of Justicia Section Mesoamericanae (Acanthaceae) Thomas F. Daniel Department of Botany, California Academy of Sciences Golden Gate Park, San Francisco, CA 94118; Email: tdaniel@ calacademy.org Four morphologically and palynologically similar species of Justicia from southern Mexico and Central America are treated as a new section of the genus, Justicia sect. Mesoamericanae. Justicia calliantha (Honduras), J. nicaraguensis (Nicaragua), J. sulfurea (Guatemala and El Salvador), and J. torresii (Mexico) share a similar inflo- rescence structure, floral form, and pollen type but differ from one another in ves- ture, size and color of the corolla, and calyx length. With estimates of 600 species worldwide, Justicia is the largest genus of Acanthaceae and the largest genus of the family in the Mesoamerican region. At least 97 species have been reported from this region between the Isthmus of Tehuantepec and the eastern border of Panama (Daniel 1995, 1997, 2001, in press, unpublished). Phylogenetic relationships among species of Justicia and related genera were discussed by McDade et al. (2000), and additional sampling of taxa and molec- ular markers in the Justicieae are underway. In an infrageneric study of Justicia, based on a worldwide subset of species, Graham (1988) recognized 16 sections and seven subsections. Additional sections in the genus have been recog- nized by Hansen (1987), Hilsenbeck (1990), Ensermu Kelbessa (1990), Immelman (1992), and Daniel (2003). The descriptive activity at the infrageneric level subsequent to Graham’s (1988) account attests both to the importance of her treatment and to the morphological diversity in the genus that remained (and still remains) to be placed into a taxonomic framework. Numerous American species cannot be accomodated in any of the sections that have been previously recog- nized (e.g., see Daniel 1999, 2002; Daniel and Wasshausen 1990) and Graham (1988) treated numerous species as “peripheral” to her infrageneric taxa. Herewith, a group of morphologically similar and predominately Mesoamerican species of Justicia that cannot be accomodated in exist- ing infrageneric taxa of that genus is accorded sectional status. Justicia section Mesoamericanae T.F. Daniel, sec. nov. TYPE. Justicia sulfurea (Donn. Sm.) D.N. Gibson. Herbae perennes vel frutices; inflorescentia in axillis foliorum, pedunculata, brevispicata vel breviracemosa vel fasciculata, floribus 1—5, sessilibus vel brevipedicellatis; calyx 4-lobus, lobis aequalibus vel subaequalibus; corolla flava, aurantiaca, vel rubra, tubo ampliato gradatim apicem versus, fauce indistincta; stamina thecis basi ecalcaratis; pollinis granae 2-aperturatae; capsula 15-33 mm longa, pubescens trichomatibus glandulosis; semina tuberculata. Perennial herbs to shrubs. Young stems multistriate with greenish striations. Inflorescence of pedunculate short-spikes, racemes, or subcapitate clusters of flowers from axils of leaves, spikes or clusters solitary or paired at nodes. Flowers sessile to short-pedicellate, 1-5 per inflorescence; calyx 4-lobed, lobes equal to subequal in size; corolla yellow, orange, or red, tube gradually expanded distally, lacking a distinct throat, + equal to or longer than lips; thecae lacking basal 174 ; DANIEL: JUSTICIA SECTION MESOAMERICANAE 175 appendages; pollen 2-aperturate, apertures flanked on each side by 2(—3) rows of + circular insu- lae, insulae and interapertural regions reticulate. Capsules 15—33 mm long, pubescent with glandu- lar (and often eglandular as well) trichomes; seeds 4, sublenticular, covered with + conical tuber- cles. Section Mesoamericanae consists of 4 species that occur from southern Mexico (lat. 17°51’N in Oaxaca) to northern Nicaragua (lat. 13°45’N in Atlantico Norte). Each of the species is relative- ly restricted in distribution and their distributions are not known to overlap. The species occur in a diversity of biotic communities. Macromorphological and palynological characters of the section are shown in Figures | and 2, respectively. The section is distinguished by the combination of axillary, pedunculate, and few-flowered inflorescences; more or less equally 4-lobed calyces; relatively large corollas with the tube gradu- ally expanded distally; diaperturate pollen; and glandular pubescent capsules. The greenish (turn- ing brown with age) seeds bearing prominent tubercles may also be characteristic of the section, but seeds remain unknown for two of the species. The inflorescence is a few-flowered spike or raceme of alternate, sessile, single-flowered dichasia subtended by two bracteoles in the axil of a bract. The flowers are sessile (spikes) to short- pedicellate (racemes). The fertile portion of the spikes is often reduced with the rachis between bract-bearing nodes very short or nearly absent. When the rachis is not evident, the spikes appear as pedunculate clusters of flowers and the distinction between bracts and bracteoles is then diffi- cult to discern. The entire inflorescence is sometimes partially or entirely enclosed by revolue leaves. Diaperturate pollen in Justicia 1s secondarily bilateral in symmetry and somewhat more com- plicated to describe that radially symmetric pollen (see Daniel 1998). Pollen in section Mesoamericanae has a P:E in apertural view of 1.404 to 2.021 and a P:E in interapertural view of 3.711. Because the ratio of the longer equatorial axis to the shorter equatorial axis is 1.90 (based on grains of J. torresii only), the pollen can be characterized as globose-oblong (Walker and Doyle 1975). Pollen among the four species of section Mesoamericanae is remarkably uniform in sculp- tural characteristics (Fig. 2). Using Graham’s (1988) key to and descriptions of sections of Justicia, plants treated here would appear most similar to species in section Drejerella (Lindau) V. Graham (i.e., with inflores- cences short-pedunculate and consisting of fewer than seven flowers and with red to yellow flow- ers 24 to 63 mm long). Species of section Mesoamericanae differ from those in section Drejerella by having a 4 (vs. 5)-parted calyx, somewhat longer inflorescence peduncles (0.5—2 vs. less than 1 cm long), and 2 (vs. 3)-aperturate pollen. Pollen of species in section Mesoamericanae (Figure 2) conforms to that described and figured by Graham (1988) as her type 7. Such pollen is encountered in at least four of Graham’s sections. One of these, section Chaetothylax (Nees) V. Graham, con- tains some species with 4-parted calyces. However, species in that section differ by having highly modified anthers (i.e., thecae oblique, unequal in size, and the lower with a basal appendage) and white, pink, blue, or wine-red corollas. The other sections with similar pollen (Orthotactus (Nees) V. Graham, Plagiacanthus (Nees) V. Graham, and Simonisia (Nees) V. Graham) all have 5-parted calyces and differ in various other characters as well. Key to Species of Justicia sect. Mesoamericanae la. Corolla red, externally pubescent with glandular and eglandular trichomes; Honduras. Oe by ROC RE NOTE REE A ORE Coc PREM TD TR OT PRN a SS CON a cer ee. Oh Pics Guewe OBI Phare 1. J. calliantha 1b. Corolla yellow or orange, externally pubescent with eglandular trichomes only. ......... A, 176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF ai Volume 55, No. 8 <= — DANIEL: JUSTICIA SECTION MESOAMERICANAE WD) 2a. Cauline trichomes mostly antrorse to antrorsely appressed; calyx 11-14 mm long; thecae BP Ses IMT ONS NI CKICO ser. Megan arts eles are erin Dar amin ta oes tc SEM NOt od ARLE yd 4. J. torresii 2b. Cauline trichomes mostly retrorse; calyx 5-10 mm long; thecae 1.5—2.5 mm long; Central ANETEITIODs 9 15 928 ONS PEE OE ah CHER MCP EE CLS ORS RMRSECIMNGRT , acMRDReeMace Re nerPT tat peli gees tae oa oa 3 3a. Corolla yellow, (17—) 22—29 mm long, lobes of lower lip 1.7—2.5 mm long; Guatemala and E] SELVEGIOE